LIBRARY UNIVERSITY OF CALIFORNIA DAVIS Digitized by the Internet Archive in 2007 with funding from IVIicrosoft Corporation http://www.archive.org/details/animalparasitesoOOfantrich THE ANIMAL PARASITES OF MAN BY H. B. FANTHAM, M.A.Cantab, D.Sc.Lond. Lecturer on Parasitology, Liverpool School of Tropical Medicine ; iSectionctl Editor in Protozoology^ " Tropical Diseases Bulletin,^' Londofi, atc.^ •"''', J. W. W. STEPHENS, M.D.Cantab., D.P.H. Sir Alfred [o7ies Professor of Tropical Medicine^ Liverpool University^ etc. AND F. V. THEOBALD, M.A.Cantab., F.E.S., Hon. F.R.H.S. Professor of Agricultural Zoology, London University ; Vice- Principal and Zoologist of the South-eastern Agricultural College ; Mary Kingsley Medallist ; Grande Medaille Geoffroy St. Hilaire, Soc. Nat. d'' Accliui. de France^ etc. PARTLY ADAPTED FROM Di. Max Braun's *' Die Tierischen Parasiten des Menschen " (4th Edition, 1908) and an Appendix by Dr. Otto Seifert. NEW YORK WILLIAM WOOD AND COMPANY MCMXX. LIBRARY .UNIVERSITY OF CALIFORNL\ DAVT.S N PREFACE 111 PREFACE. The English edition of Braun's '' Die Tierischen Parasiten des Menschen," produced in 1906, being out of print, the pubhshers decided to issue another edition based on the translation of Brain's fourth German edition, which appeared in 1908, to which had been added an appendix, by Dr. Otto Seifert on Treatment, etc/ • ^ .'. When the work was considered with a view to A hevv edition, it was found that a vast amount of new matter had to be incorporated, numerous alterations essential for bringing it up to date were necessi- tated, and many omissions were inevitable. The result is that parts ol the book have been rewritten, and, apart from early historical references, the work of Braun has disappeared. This is more particularly the case with the Protozoa section of the present work. The numerous addi- tions, due to the great output of scientific literature and other delays in publication, have led to the book being somewhat less homogeneous than we desired, and have necessitated the use of appendices to allow of the presentation of new facts only recently ascertained. Many new illustrations have been added or substituted for older, less detailed ones. Some of these new figures w^ere drawn specially for this book* The first section, on the Protozoa, has been wTitten by Dr. Fantham, there being little of the original text left except parts of the historical portions, and thus the section on Protozoa must be considered as new. The second section, on Worms (except the Acanthocephala, Gordiidae and Hirudinea), has been remodelled by Professor Stephens to such an extent that this, too, must not be looked upon as a translation of Braun's book. With regard to the Arthropoda, much remains as in the last English edition, but some new matter added by Braun in his fourth German edition is included, and much new matter by Mr. Theobald has been incorporated. As regards the Appendix by Dr. Seifert, the first section has been remodelled, but the sections on the Helminthes and the Arthropoda are practically translations of the original. The authors desire to express their thanks to Miss A. Porter, D.Sc, J. P. Sharpies, Esq., B.A., M.R.C.S., and H. F. Carter, Esq., F.E.S., for valuable help. They also wish to thank the authors, editors, and IV THE ANIMAL PARASITES OF MAN publishers of several manuals and journals for their courtesy in allowing the reproduction of certain of their illustrations. In this connection mention must be made more particularly of Professor Castellani, Dr. Chalmers, Professor Doflein, Dr. Leiper, the late Professor M inch in. Professor Nuttall, Dr. Wenyon, Mr. Edw. Arnold, Messrs. Bailliere, Tindall and Cox, Messrs. Black, Messrs. Cassell, Dr. Gustav Fischer, Messrs. Heinemann, the Cambridge University Press, the Editors of the Annals of Tropical Medicine and Parasitology, the Editors of the Journal of Experimental Medicine, and the Editor of the Tropical Diseases Bnlletin. H. B. F. J. W. W. S. December, J915. F. V. T. CONTENTS CONTENTS, PAGE PREFACE ... ... ... ... ... ... ... ... ... Hi ERRATA ^ xxxii ON PARASITES IN GENERAL i Occasional and Permanfent Parasitism ... ... ... ... ... i Entozoa, Endoparasites, Helminthes, Turbellaria ... ... ... ... 2 Hermaphroditism ... ... ... ... ... ... .. 4 Fertility ot Parasites ... ... ... ... ... ... ... 5 Transmigrations ... ... ... ... ... ... ... ... 5 Commensals, Mutualists ... ... ... ... ... ... ... 6 Incidental and Pseudo-parasites ... ... ... .. ... ... 6 The Influence of Parasites on the Host ... ... ... ... ... 8 Origin of Parasites ... ... ... ... ... ... ... 10 Derivation of Parasites ... ... ... ... ... ... ... 19 Change of Host ... ... ... ... ... ... ... ... 20 Literature ... ... ... ... ... ... ... ... 22 THE ANIMAL PARASITES OF MAN 25 A. Protozoa ... ... ... ... ... ... ... 25 Classification of the Protozoa. Class I. Sarcodina ... Order. Amcebina Foraminifera Heliozoa . . , Radiolaria Class II. Mastigophora III. Sporozoa Sub-class I. Telosporidia Order. Gregarinida Coccidiidea Hamosporidia Sub-class 2. Neosporidia Order. Myxosporidia Microsporidia Sarcosporidia Haplosporidia Class IV. Infusoria V. SUCTORIA 27 27 27 27 28 28 28 28 28 28 28 28 28 28 28 29 29 29 VI THE ANIMAL PARASITES OF MAN The Animal Parasites of Man — contd, ProtOZOSi— con ^d. PAGE Class I. Sarcodina, Biltschli, 1882 ... ... ... ... ... 29 Order. Amcebina, Ehrenberg ... ... ... ... ... ... 29 A. Human Intestinal Amoeb£e ... ... ... ... ... 29 Entamceha coli, Losch, 1875, emend. Schaudinn, 1903 .. 32 Entamoeba histolytica, Schaudinn, 1 903 ... ... ... 34 Enta/nceba te(rage7ta, WiQxecV, 1907... ... ... 38 Noc's Entamoeba, 1909 ... ... ... . . 41 Entamceba buccalis, Prowazek, 1904 ... ... ... 43 Entatnceba undulans, QsL^itWaxviy i()0^ ... ... ... 43 Entamoeba kartulisi, Doflein, 1901 ... ... ... 44 A7no;ba gingivalis, A. buccalis, A. dentalis ... ... 44 Genus. Paramceba, Schaudinn, 1896 ... ... ... 44 Paramceba (Craigia) hominis, Craig, 1906 ... ... 45 B. Amoebae from other Organs ... ... ... ... .. 45 Entamoeba pult?ionalis, Artault, 1898 ... ... ... 45 Amceba urogeititalis , Baelz, 1883 ... ... ... 45 Aviceba miurai, Ijima, 1898 ... ... ... ... 46 Appendix: " Rbizopods in Poliomyelitis Acuta" ... ... ... 46 Order. Foraminifera, d'Orbigny ... ... ... ... ... 47 Sub-order. Monothalamia (Tc-taceous Amoebse) ... ... ... ... 47 Genus. Chlamydophrys, Cienkowski, 1876 ... ... ... 47 Chlamydophrys enchelys, Ehrenberg ... ... ... 47 Leydenia geinmipara, Schaudinn, 1896 ... ... .. 49 Class II. Mastigophora, Diesing ... ... ... ... ... 50 Sub-class. Flagellata, Cohn emend. Biitschli ... ... ... ... 50 Order. Polymastigina, Blochmann ... ... ... ... ... 52 Genus. Trichomonas, Donne, 1837 ... ... ... 52 Trichomonas vaginalis, Donne ... ... ... ... 52 Trichomonas intestinalis, R. Leuckart, 1 879 = Trichomonas hominis, Davaine, 1854 ... ... ... ... 54 Genus. Tetramitus, Perty, 1852 ... ... ... ... 57 letratnitus mesnili,^Qnyox\y 1910 ... ... ... 57 Genus. Lamblia, R. Blanchard, 1888 ... ... ... 57 Lamblia intestinalis, Lambl, 1 8 59 ... ... ... 57 Order. Protomonadina, Blochmann ... ... ... ... ... 60 Family. Cercomonadida, Yieni evaerxA. 'RviischW ... ... ... 61 Genus. Cercomonas, Dujardin emend. Biitschli ... ... 61 Cercomonas hominiSf'Da.vdXnG, 1 854 ... ... ... 61 Monas pyophila, R. Blanchard, 1895 •■• ••• ••• ^2 Family. Bodonida, Biitschli ... ... ... ... ... 63 Genus. Prowazekia, Hartmann and Chagas, 19 10 ... ... 63 Prowazekia urinaria, Hassall, 1859 ... ... ... 63 Prowazekia asiatica, C2i5\.Q\\2iX\i Siwd OmXmex?,, 1910 ... 65 Prowazekia favajiensis,Y\n, igi2 ... ... ... 66 Prowazekia crttzi, Hartmann and Chagas, igio ... ... 66 Prowazekia weinbergi, MsLthis and heger, igio ... ... 66 Prowazekia parva, Nagler, 1910 ... ... ... 66 Family, Trypanoso?nidcB, Doflein ... ... ... ... 66 Genus. Trypanosoma, Gruby, 1843 ••• ••• ••• ^7 Historical ... ... .. ... ... 67 General ... ... ... ... ... ... 69 Morphology ... ... ... ... ... 70 Trypanosoma gainbiense, Dutton, 1902 ... ... ... 72 Trypanoso?na nigeriense, Macfie, 1913 ... ... ... 76 CONTENTS Vii The Animal Parasites of Man — contd. Protozoa.— con^d. page Trypanosoma rhodesiense, Stephens and P'antham, 19 lo ... 76 General Note on Trypanosomes with Posterior Nuclei 83 Trypanosoma crt4zi, C\\2ig2i%, 1 909 ... ... ... 83 Trypanosotna lewisi, Kent, 1 88 1 ... ... ... 88 Trypanosoma brucei, Plimmer and Bradford, 1899... ... 93 Trypanosoma evansi^ Steel, 1885 ... ... ... 95 Trypanosoma equinum^V ogQs, igoi ... ... ... 96 Trypanosofna equiperdtwi^ Doflein, 1901 ... ... ... 97 Trypanosoma theileri^ Bruce, 1902 ... ... ... 98 Trypanosoma hippicum^ Darling, 1910 ... ... ... 98 Endotrypamim schaudinni, Mesnil and Brimont, 1908 ... 99 Trypanosoma boy let, Lafont, 1912 ... ... ... 99 Monomorphic Trypanosomes ... ... ... ... 99 Trypanosoma vivax, 7.\Qvi\2iX\v\, 1905 ... ... ... 99 Trypanosoma caprce, Kleine, 1910 ... ... ... lOO Trypanosoma con^olenscy Broden, 1904 ... ... ... 100 Trypanosoma simice,'Qx\xcCi 1912 ... ... ... lOO Trypanosoma uniforme., Bruce, 1910 ... ... ... loi General Note on Development of Trypanosomes in Glossina 10 1 Adaptation of Trypanosomes ... ... ... lOl Genus. Herpetomonas, Saville Kent, 1881 ... ... ... 102 Genus. Crithidia, Leger, 1902, emend. Patton, 1908... ... 104 Genus. Leishmania, Ross, 1903 ... ... ... ... 104 Zm/^wa«2'a «?9'j) ... ... ••• 167 /•/ai^w^^/Mw 7^/zV/Mw, Sergent, 1907 (in birds) ... >.. 170 Cultivation of Malarial Parasites .. ... ... 170 Differential Characters of the Human Malarial Parasites 171 P'amily. Piroplasmidce, Franca, 1909 ... ... .•• ... 172 Genus. Babesia, Starcovici, 1893 ... ... ... ... 174 Genus. Ty^^zV^rza, Bettencourt, Fran9a and Borges, 1907 ... 178 Theileria parva, Theiler, 1903 ... ... ... ... 178 Iheileria mutans, Theiler, 1907 \ ... ... ... 1 80 Genus. Anaplasj/ia, Theiler, 1910 ... ... ... ... 180 Genus. Paraplasma, Seidelin, 1911 ... ... ••• 180 Sub-class. Neosporidia, Schaudinn ... ... ... ... ... 181 Order. Myxosporidia, Biitschli ... ... ... ... ... 181 Order. Microsporidia, Balbiani ... ... ... ... ... 184 Order. Actinomyxidia, Stole. ... ... ... ... ... ... 187 Order. Sarcosporidia, Balbiani ... ... ... ... ... 187 Sarcospoiidia oh?,et\tA xn Moxi ... .. ... 193 Order. Haplosporidia, C2i\x\\txy zxxd Me^mXy i2><)() ... ... ... 194 Rhinosporidium kinealyi, Minchin and Fantham, 1905 ... 195 Class IV. Infusoria, Ledermiiller, 1763 ... ... ... ... ... 198 Genus, Balantidium, Claparede et Lachmann ... ... 200 Balantidium coli, Malmsten, 1857 ... ... ... 200 Balantidium minutum, Schaudinn, 1899 ... ... 204 Genup. Nyctotherus, Leidy, 1849 ••• ••• •• ••• 204 Nyclotherus faba, Schaudinn, 1899 ... ... ... 205 Nyctotherus giganieus, P. Krause, 1906 ... ... ... 205 \Nyctotherus\africanus,QzsX&\\2.x\\,\iyd^ ... ... ... 206 The Chlamydozoa ... ... ... ... ... ... ... 207 Protozoa Incert^ Sedis ... ... ... ... ... ... 210 Sergenfella ^ominis, Brumpt, igio ... ... ... 210 B. Platyhelminthes (or Flat Worms) ... ... ... ... ... 211 Classification of the Platyhelminthes. Class I. Turbellaria (or Eddy Worms) ... ... ... ... 212 Order I. Rhabdoccelida ... ... ... ... ... ... 212 2. Tticladida ... ... ... ... ... ... 212 3. Polycladida ... ... ... ... ... ... 212 Class II. Trematoda (Sucking Worms) ... .. ... ... 212 III. Cestoda (Tapeworms) ... ... ... ... ... 212 Class II. Trematoda, Rud. ... ... ... ... ... ... 212 Development of the Trematodes ... ... ... ... . . 222 Biology ... ... ... ... ... ... ... ... 229 CONTENTS IX The Animal Parasites of Man — contd. Platyhelminthes— t6... ... 236 Family. Fasciolidce, Railliet, 1895 ••• ••• ••• •• ^37 Sub-family. FasciolincB, Odhner, 1910 ... ... ... ... 237 * Genus. Fasciola, L., 1758... ... ... ... ... 237 Fasciola hepatica,\u., Vji^Z ... ... ... ... 237 Halzoun... ... ... ... ... ... 242 Fasciola giganti'ca, Cobbold, 1856 ... ... ... 244 Sub-family. Fasciolopsitice, Odhner, 1910 ... ... ... 245 Genus. Fasciolopsis, Looss, 1898 ... ... ... ... 245 Fasciolopsis buski, Lank., 1857 ... ... ... ... 245 Fasciolopsis rathouisi, Ward, 1903 ... ... ... 246 Fasciolopsis goddardi. Ward, 1910 ... ... .. 247 Fasciolopsis fiilleborni, Rodenwaldt, 1909 ... .. 247 ^ Family. Troglotremidce, Odhner, 1914 ... ... ... ... 249 Genus. Paragoninius, BxeiMn, 1899... ... ... ... 249 Faragonitnus ringeri^ Cobb., 1880 ... ... ... 249 Family. Opisihorchiidcs, Braun, 1 90 1 ... ... ... ... 252 Sub-family. Opislhorchiince, Looss, 1899 ... ... ... 252 Genus. Opisthorchis, R. Blanch., 1845 ••• ••• "• ^52 Opisthorchis felineus, Riv., 1885 ... ... ... 252 Genus. Paropistkorchis, Stephens, 1912 ... ... ... 255 Paropisthorchis caninus, ^2ixV.Gry 1^12 ... ... ... 255 Genus. Amphimerus, Barker, 191 2 (?) ... ... ••• 257 Amphimerus fiovejra, Barker, 1912 (?) ... ... ••■ 258 Genus. Clonorchis, Looss, 1907 ... ... ... ... 258 X THE ANIxMAL PARASITES OF MAN The Animal Parasites of Man — contd. Platyhelminthes— r Echinostoma malayantirn, Leiper, 1911 ... Sub-family. Himasthlince, Odhner, 1910 Genus. Artyfechinostornutn, Clayton-Lane, 1915 Artyfechinostomutn siifrartyfex, Clayton-Lane, 191 5 Family. Schistosomidce, Looss, 1899 ... ... Genus. Schistosoma^ Weinl.. 1858 ... Schistosoma hivmatobitwi^ Bilharz, 1852 ... Schistosoma mansoni, Sambon, 1907 Schistosoma ;aponicu?f?,\s.2i\.s\M2idia, 1904... Class IIL Cestoda, Rud., 1808 ... Anatomy OF THE Cestoda ... Development OF THE Tapeworms Biology Classification of the Cestoda of Man. Order. Pseudophyllidea, Carus, 1863 Family. Dibothriocephalida, Liihe, 1902 Sub-family. DibothriocephalincB, Liihe, 1899 Order. Cyclophyllidea, v. Beneden Family. DipylidiidcB, Liihe, 1910 ... Hymenolepididce, Railliet and Henry, 1909 ... Davaineidi?, Fuhrmann, 1907 Sub-family. Davaineincey Braun, 1900 ... Family. Tc^niidce, Ludwig, 1886 ... The Cestodes OF Man Famiy. Dibothriocephalidce Sub-family. Dibothriocephalince Genus. Dibothriocephalus, Liihe, 1899 Dibothriocephalus latus, L., 1748 Dibothriocephalus cordatus, R. Lkt,, 1863 Dibothriocephalus parvus, Stephens, 1908 Genus. Diplo^onoporus, Lonnbrjj., 1892 Diplogonoporus grandis, R. Blanch., 1894 Sparganuni, Diesing, 1854 Sparganum mansoni, Cobb., 1883 Sparganum proliferum, Ijima, 1905 CONTENTS XI The Animal Parasites of yiAn—contd. Platyhelminthes — confd. Family. Dipylidiidce, Liihe, 1 910 Genus. Dipylidium, R. Lkt., 1863... Dipylidium canhuun, L. 1 758 ... Family. ^;/wg«^/^/za?'/fl('^, Railliet and Henry, 1909 Genus. Hymenolepis, Weinland, 1858 Hymenolepis nana, v. Sieb., 1852 Hytnenokpis diminuta, Rud., 18 19 Hymenolepis lanceolata, Bloch, 1782 Family. Davaineidce, Fuhrmann, 1907 ... Sub-family. Davaineina;, Braun, 1900 ... Genus. Davainea, R. Blanch., 189 1 Davainea madagascariensis, Davaine, 1869 Davainea (?) asiaiica, v. Linst., 1901 Family. Tceniid(C, Ludwig, 1886 Genus. Tanm, L., 1758 ... Tania solniin^'L,., p. p.^ I'jd'j ... Cysticercus acanthotrias, Weinland, 1858... Tli'wza ^r^///««'z, Stephens, 1908 Tcenia marginata, Batsch., 1786 Tienia serrata, Goeze, 1782 Tcenia crassicollis, Rud., 1810 ... Tc^nta saginala, GoezQ, i'j2>2 Ticnia africana, \. Linst., 1900... Tcenia conftisa, Ward, 1896 Tcenia echinococcHs, v. Sieb., 1853 Structure and Development of Ecchinococcus (Hydatid) Echinococctts mtiltilocularis (Alveolar Colloid) Serum Diagnosis of Echinococcus ... C. Nemathelmintljes Class. Nematoda Anatomy of the Nematodes Development of the Nematodes Classification of the Nematoda. Family. Anguillulidic, Gervais and van Beneden, 1859 Angiostomidcd, Braun, 1895 Gnathostoviidce Dractmculidce, Leiper, 19 12 Filar iidce, Claus, 1885 TrichinellidLe, Stiles and Crane, 1 9 10 Dioctophyinidcc Strongylida, Cobbold, 1864 Physalopteridce Ascaridce, Cobbold, 1864 Oxyuridce The Nematodes observed in Man ... Family. Anguillulidcc Genus. Rhabdiiis, Dujardin, 1845 ••• Rhabditis pellio, Schneider, 1866 Rhabditis niellyi, Blanchard, 1885 Rhabditis, sp. Genus. Anguillula, Ehrenberg, 1826 Anguillula aceli, Miiller, 1783 ... xu THE ANIMAL PARASITES OF MAN The Animal Parasites of yiA^—contd. Nemathelminthes— 3imt\s,i^o2> ... ... ... 438 Sub-family. Ancylostoinimc, Raillet, 1909I ... ... ... 438 Group. CEsophagostomecc, Railliet and Henry, 1909 ... ... ... 439 Genus. Ternidens, Railliet, 1909 ... ... ... ... 439 Ternidens deminutus, Railliet and Henry, 1905 ... ... 440 Genus. (Esophagostofiiuni, Molin, 1861 ... ... ... 441 CEsophagostoinuiii bniniptiy Railliet and Henry, 190'; ... 441 CEsophagostofiiHtn stephanostonmm var. thomasi, Railliet and Henry, 1909 ... ... ... ... ... 443 (Esophagostovium apiostonnini, Willach, 1891 ... ... 444 Group. AncylostomecE, Railliet and Henry, 1909 ... ... ... 445 Genus. Ancylostoma, Dubini, 1843, emend. Looss, 1905 ... 445 Ancylosioma duodenale,T)\x\nn\, 1843 ... ... ... 445 Ancy loslovi a ceylanicuvi, \aOQ%%, 19 ii ... ... ... 456 Ancylostotna brazilietise^Qoxn&z ^^Y^ixxz, \<^\o ... ... 456 Group. .5/^«i7i'/tfz//$g ... ... ... 47^ Echinorhynchus moniliformis, V>x&xii%^x, \^\.<^ ... ... 478 E. Gordiidae ... ... ... ... •. •• ••• ••• 479 F. Hirudinea s. Discophora (Leech) ... ... ... ... ... 480 Family. 6';/aMo^fl'^///^^ (Leeches with Jaws) ... ... ... 481 Genus. Hirudo, L., 1758 ... ... ... ... ... 481 Hirudo medicinalis,L..y lT^2i ... ... ... ... 481 Hirudo troctina, ]o)M\%\.or\, 1816 ... ... ... 482 Genus. Limnatis, Moq.-Tandon, 1826 ... ... ... 482 Limnatis nilolica, Savigny, 1820 ... ... ... 482 Genus. Hicmadipsa, Tennent, 1 86 1... ... ... ... 482 Family. Khynclwbdellidce (Leeches with Rostrum) ... ... 482 Genus. Htetnentaria, de Filippi, 1849 ... ... ... 482 Hamentaria officinalis y dQ¥\\\^^\ ... ... ... 482 Genus. Placobdella, R. Blanchard ... ... ... ... 482 Placobdella calenigera, M.o(\.-l!2indoTi .. ... ... 482 G. Arthropoda (Jointed-limbed Animals) ... ... ... ... ... 483 ^. Arachnoidea (Spiders, Mites, etc.) ... ... ... ... ... 483 Order. Acarina (Mites) ... ... ... ... ... ... 484 Family. Trombidiidcc {K\xx\xi\x\^M\\.^%) ... ... ... ... 485 Genus. Ti ombidiian, 'L.dXx^'iWe. {^cxidi Lepttis) ... ... ... 485 Leplus atitumnalis, ShsiViy 1790... ... ... ... 485 Trombidium tlalsahuate^ Lemaire, 1867 ... ... ... 486 Akamiishi ox Kedani ... ... ... ... ... 487 Family. 7"oxi, i%^o ... ... ... 489 Genus. Neph7-ophages ... ... ... .. ... 490 Nephrophages sanguinariusy Miyake and Scriba, 1893 ... 490 Family. Eupodidce ... ... ... ... ... .. 491 Genus. Tydeus, Koch ... ... ... ... ... 491 7 ydeus mo/eslus, Mon\ez, iSSg ... ... ... ... 491 Family. Gainasidcc (Coleopterous or Insect Mites) ... ... 491 Genus. Dermanyssus, Duges ... ... ... ... 492 Derjnatiyssus gallince, di^ Gqqx, 1778 ... ... .. 492 Dertnanyssns hirundinis, Hermann, 1804 ,,. ... 492 Genus. Holothyrus ... ... ... ... ... 493 Holothyrus coccinella, Gervais, 1842 ... ... ... 493 CONTENTS XV The Animal Parasites of Man — contd. Arthropoda- .\ Laverania^TYi^oh^X^ Genus. Celliay Theobald ... Genus. Neocellia, Theobald Genus. Kertesziuy Theobald Genus. Manguinhosia, Ciuz Genus. Chagasia, Cruz Genus. Calveriina, Ludlow Genus. Birbnella, Theobald Sub-family. Megarhinina Genus. Alegar hinus, Robineau Desvoidy Genus. Toxorliynchites, Theobald ... Sub-family. Culicince ... Genus. Mucidus^ Theobald Genus. Psorophora^ Robineau Desvoidy Genus. Janthinosoma, Arribalzaga ... Genus. Stegomyia^ Theobald Stegomyia fascial a, Fabricius (Yellow Fever Mosquito) Stegoviyia scutellaris. Walker Genus. Theobaldia, Neveu-Lemaire Theobaldinella^ Blanchard Theobaldia antmlata, M.G.\gQn Genus. Culex, Linnaeus Genus. Melanoconion^ Theobald Genus. Grabhamia, Theobald Genus. Psetidotceniorhynchus, Theobald ; Tccniorhynchus^ Theobald, non-Arribalzaga Genus. Tceniorhynchus^ Arribalzaga ; Mansonia, Blanchard ; Panopliiesy Theobald Genus. Chrysoconops, Goeldi Other Nematocera Family. Simulidce Family. ChirononiidcE (Midges) Sub-family. CeratopogonincE Family. Psychodidce (Owl Midges) Brachyera (Flies) Family. Phoridcs AphiochcEta ferruginea, Brun Phora ruJipeSy 'M.^\g. ... Family. Sepsidcz Piophila casei,\,. Family. ^S^r/^zflfe (Hover and Drone Flies) Family. Drosophilidce Drosophila melanogastery Br. ... ... ... Family. Muscidce Teichomyzafusca, Macq. Homalomyia canicularis^'Li., &{c. Honialomyia scalaris^ Fabr. Anthomyta desjardensii, Macq. ... ... ... Hydrotaa meteor ica^ L. Cyrtonetira siabulans ... PAGE 568 568 568 568 568 569 569 569 569 569 570 570 570 570 570 S70 571 571 571 571 571 574 575 575 575 575 575 576 576 576 577 577 577 577 579 580 581 582 582 583 583 583 583 583 584 584 584 584 584 585 585 585 585 CONTENTS XIX The Animal Parasites of Man — contd. Arthropoda— contd. page yl/wi-^ro! ^/^w3 714 714 715 715 729 733 733 • • 733 .. 733 .. 734 734 734 735 736 736 737 377 737 various 737 739 739 740 741 742 742 742 742 744 745 745 747 748 749 XXI 1 THE ANIMAL PARASITES OF MAN APPENDIX ON TREMATODA AND NEMATODA ... 753 Trematoda ... 753 Artyfechinostomum sufrartyfex ... ... 753 Metagoniimis ( Yokogawd) yokogawai ... 753 Opisthorchis sp. ... 753 Schistosome cercarim ... 753 Distomata cercariiF - 753 Group. Ferrocercous cercarice ... • •• 753 Family. Schistosoviida •• 753 Cercaria bilharzia, Leiper, 1915 .. 754 Cercaria dz^/iarzie//a, heiper, igis ... 754 Schistosoma mansotii, Sambon, 1907 • 754 Nematoda ••■ 754 Ancylostomiasis ... 754 Ground Itch ... •• 754 Ascaris lumbricoides ... ... 754 Filariasis ■•• 755 Onchocerca volvtilus 755 Sirongyloides stercoralis ••• 755 BIBLIOGRAPHY ... .... .. ... 756 INDEX .. 836 LIST OF ILLUSTRATIONS XXllI LIST OF ILLUSTRATIONS. PACK Fig. I Amcebacoli. (After Loesch) ... ... ... ... ... ... 29 2 Encysted intestinal amoebae. (After Grassi) ... ... ... ... 31 3 Eniama'ba coli,\\^e-cyc\Q. (After Castellani and Chalmers) ... ... 32 4 Enta7?ia'ba CO li) so-c&Wtdi 2m\.og2iTc\y. (From Minchin) ,.. ... ... 34 5 Entama'ba histolytica {tetrageita ioxm). (After Hartmann) ... ... 35 6 Entamceba histolytica, ingestion of red blood corpuscles. (After Hartmann) 35 7 Entamoeba histolytica, section through infected intestinal ulcer. (After Harris) 36 8 Entamceba histolytica {tetragena), trophozoite and nuclei. (After Hartmann) 38 9 Entamceba histolytica [tetragena), cysts. (After Hartmann) ... ... 39- 10 Entamceba buccalis. (After Leyden and Lowenthal) ... ... ... 43 11 Entamceba kartuli si. (After Kartulis) ... ... ... ... ... 44 12 Amoeba miurai. (After Ijima) ... ... ... ... ... 46 13 Chlamydophrys enchelys. (After Cienkowski) ... ... ... ... 48 14 Chlamydophrys enchelys, Qncy?,\.Qdi. (After Cienkowski) ... ... ... 49 15 Leydenia gemmipara, Schaudinn ... ... ... ... ... 50 16 Trichomonas vaginalis. (Ater Kunstler) ... ... ... ... 53 17 Trichomonas intestinalis. (After Grassi) ... ... .. ... 54 18 Trichomonas intestinalis. (Original, Fantham) ... ... ... 55 19 Lamblia intestinalis. (After Wenyon, from Minchin) ... ... ... 58 20 Lajjiblia intestinalis. (After Grassi and Schewiakoff) ... ... ... 59- 21 Cercomonas hominis. (After Davaine) ... ... ... ... 61 22 Cercoinonas hominis, ixoxu zxi ^c!t{vc\ozo(:.CK!i% cy%\.. (After Lambl) ... ... 61 23 Monas pyophila. (After Grimm) ... ... ... ... ... 62 24 Prowazekia urinaria. (After Sinton) ... ... .,. ... 64 25 Prowazekia urinaria, excystation. (After Sinton) - ... ... ... 65 26 Trypanosotna brucei vci d\\\%\on. (After Laveran and Mesnil) ... ... 70 27 Trypanosoma lewisi, rosettes. (After Laveran and Mesnil) ... ... 71 28 Trypanosoma gambiense. (After Dutton) ... ... ... ... 73 29 Trypanosoma ga7nbiense,d&\Q\o^mtr\i\nvQx[ehi2^.e\\ost. (Original, Fantham) 73. 30 Trypanosoma gavibiense,^Qv^\o^vc\Qx\X.\x\ Glossina palpalis. (After Robertson) 75 31 Trypanosoma rhodesiense. (After Stephens and Fantham) ... ... 77 32 Chart showing daily counts of number of Trypanosomes per cubic millimetre of peripheral blood from a case of Rhodesian sleeping sickness. (After Ross and Thomson) ... ... ... ... ... ... 79 33 Trypanosofna crtizi, schizogony. (After Chagas, from Castellani and Chalmers) ... ... ... ... ,.. ... ... 84 34 Trypanosoma cruzi in muscle. (After Vianna, from Castellani and Chalmers) 85 35 Trypanosoma cruzi, development in Triatoma megista. (After Chagas, from Castellani and Chalmers) ... ... ... ... .. 86 36 Trypanosoma cruzi, forms found in salivary glands of Triatoma. (After Chagas, from Castellani and Chalmers) ... ... ... ... 87 37 Trypanosoma lewisi, from rat's blood. (After Minchin) ... ... ... 89 38 Trypanosoma lewisi, {xovci %'(.ovc\2i.c\v oix2X-'dit2,. (After Minchin) ... ... 91 39 Trypanosoma lewisi, ixom.XQ.c\.\xv(i oi xdX-'^Q^. (After Minchin) ... ... 92 40 Trypaiwsoma brucei. (After Laveran and Mesnil) ... ... ... 94 ^ 41 Trypanosoma evansi. (Original, Fantham) ... ... ... ... 9^ xxiv THE ANIMAL PARASITES OF MAN PAGE Fig. 42 Trypanosoma eqttinum. (After Laveran and Mesnil) ... ... ... 96 43 Trypanosoma equiperdum {(^\\g\v\-d\,Yzx\\\\9.vc{) ... ... ... ... 97 44 Trypanosoma theileri. (After Laveran and Mesnil) ... ... ... 98 45 Trypanosoma vivax. (Original, Fanlham) ... ... ... ... lOO 46 Trypanosoma congoknse. (Original, Fanthain) ... ... ... ... lOO 47 Trypanosoma uniforme. (Original, Fantham) ... ... ... ... lOO 48 Trypanosoma rotatoriiitn. (After Laveran and Mesnil) ... ... ... lOl 49 Herpetomonas, Crithidia, Trypanosoma. (After Porter) ... ... ... lo.^ 50 Leishmania donovani. (After Christophers, Patton, Leishman ; from Cas- tellani and Chalmers) ... ... ... ... ... ... 106 51 Toxoplasma gondii. (After Laveran and MaruUaz, from 7><7/. Z)w. ^«//(i?/««) 1 13 52 Toxoplasma pyrogenes. (After Ca.ste\\a.ni, from Tr op. Dis. Bulletin) ... 1 13 53 Spirochcetabalbianii. (After Fantham and Porter) ... ... ... 1 14 54 Spirochceta duttoni. (After Fantham) ... ... ... ... ... 1 17 55 SpirochcEta duttoni and its coccoid bodies in the tick. (After Fantham) ... 118 56 Treponema pallidum. (After Bell, from Ca«;tellani and Chalmers)... .. 124 57 Treponema pallidum, 2i\iip2iX2L\.\x% iox c\x\t\w2ii\on o{. (After Noguchi) ... 125 58 Treponema pertenue. (After Castellani and Chalmers) ... ... ... 127 59 Monocystis agilis. (After Stein) ... ... ... ... ... 130 60 Cr^^arma /tfw^a, stages of growth of trophozoite ... ... ... 130 61 Xyphorhynchus Jirvms. (After Leger) ... ... ... ... 13 1 62 Gregarina munieri. (After Schewiakoff) ... ... ... ... 131 63 Monocystis agilis, si^oxQ?,. (After Biitschli) ... ... ... ... 132 64 Gregarines, conjugation and spore formation. (After Calkins and Siedlecki, modified) ... ... ... ... ... ... ... 133 65 Stylorhynchus oblongatus. cy?,X. 2LX\di gz^meies. (After Leger) ... ... 133 66 Gregarines, various spores. (After Leger) ... ... ... ... 134 67 Eimeria [Coccidium) schuhergi, life-cycle diagram of. (After Schaudinn) ... 139 68 Eimeria avium in gut epithelium of grouse chick. (After Fantham) .. 143 69 ^zw^^/d! az^/«w, life-cycle, diagram of. (After Fantham) ... ... 144 70 iS'zw^r/dt j/?oxQ. (After Doflein) ... Chloromyxum leydigi. (After Thelohan) Myxobolus ffeifferi, spore formation. (After Keysselitz, from Minchin) Nosema apis. (After Fantham and Porter) Nosema bombycis from silkworm. (After Balbiani) Nosema bombycis ^ spores. (After Thelohan) Hexactinomyxon psam/noryctis, spore. (After Stole) • Sarcocystis miescheriana in muscle of pig. (After Kiihn) "... Sarcocystis miescheriana, mature trophozoite Sarcocystis tenella in section, as seen in oesophagus of sheep Sarcocystis teneUa^yowxi^ Uo'^YiOzoxit.. (After Bertram) ... Sarcocystis miescheriana, end portion of trophozoite. (After Bertram) Sarcocystis blanchardi from ox. (From Wasielewski, after van Eecke) Sarcocystis tenella. (After Laveran and Mesnil) Haplosporidium heterocirri. (After CauUery and Mesnil) Haplosporidian spores. (After Caullery and Mesnil) Rhinosporidium kinealyi, portion of ripe cyst. (After Minchin and Fantham) Balantidium coli. (After Leuckart) ... Balantidium coli, free and encysted. (After Casagrandi and Barbagallo Balantidium minutum. (After Schaudinn) Nyctotherus faha. (After Schaudinn) ... Nyctotherus giganteus. (After Krause) Nyctotherus africantcs. (After Castellani) Trachoma bodies in conjunctival cells. (Original, Fantham) Half of a transverse section through Fasciola hepatica, L. Harmostomtcm leptostomum, Olss. Median section through the anterior part of Fasciola hepatica Polystomum integerrimum. (After Zeller) Allocreadiur/i isoporum,'Loosa. (After Looss) ... Terminal flame cell of the excretory system. (Stephens).,. Diagram of female genitalia. (Stephens) Diagram of male and part of female genitalia. (Stephens) Osnm o{ Fasciola hepatica, 1^. Mira.cid\\im of Fasciola he/>atica. (After Leuckart) A group of cercariae of Echinostoma sp. ... > Development of Fasciola hepatica, L. (After Leuckart)... Young redia oi Fasciola hepatica. (From Leuckart) Older redia. of jDistoma echinatum Cercana. of Fasciola hepatica. (After Leuckart) Encysted cercaria of Fasciola hepatica. (After Leuckart) Watsonius watsoni. (After Shipley) ... Watsonius watsoni : ventral projection composed from a series of sections. (After Stiles and Goldberger) Gastrodiscus hominis. (After Leuckart) Fasciola hepatica, L. Fasciola hepatica, showing the gut and its branches Fasciola hepatica, \,. (After Glaus) ... Fasciola hepatica : egg from liver of sheep. (After Thomas) Limnceus truncatulus,M\\\\. (From Leuckart)... Yowng Fasciola hepatica. (From Leuckart) XXvi THE ANIMAL PARASITES OF MAN PAGE Fig, 145 Fasciola gigantica. (After Looss) ... ... ••• ••• ••• 243 146 Fasciolopsis buski, \.2iXi\i. (After Odhner) ... ... ••• ••• 245 147 Fasciolopsis raihouisi, VoU. (After Claus) ... ... ••• ••• 246 148 Fasciolopsis fiillebo7-ni. (After Fulleborn) ... ... ••• ... 248 149 Paragonimus ringeri, Cohh. (After Katsurada) ... ... .•• 250 150 Paragonimus ringeri, Cohh. (After Kubo) ... ... ... ••■ 250 150A Paragonimus westermanii, Ktxh. (After Leuckart) ... ... ... 25a 15 ' "Eggoi Paragonimus ringeri, Cohh. (After Katsurada) ... ... ... 251 ^52 "Egg o{ Opisthoj'c his felineus ... ... ... .•• ••• ••• 253 153 Opisthorchis felineus. (After Stiles and Hassall) ...' ... ... 253 154 Opisthorchis pseudofelineus. (After Stiles) ... ... .-• ... 254 155 Parapisthorchis caninus. (After Stephens) ... ... ••• ... 256 156 Amphiinerus nove}'ca,Bxz.Mn. (After McConnell) ... ... .. 257 157 Metorchis conjunctus. (After Cobbold) ... ... ■•• .•• 258 158 Clonorc his sinensis. (After Looss) ... ... ... .•• ... 259 1$} OwSioi Clonorchis sinensis. (After Looss) ... ... ... ... 259 160 CJonorchis endemicus. (After Looss) ... ... ... . . .•• 260 161 Clonorchis endemicus : tggs. (After Looss) ... ... ... ... 260 162 Metorchis truncalus ... ... ... ... •.■ ••. ••• 262 163 Heterophyes heterophyes. (After Looss) ... ... ... ... 263 164 MetagonijHUs yokogawai. (After Leiper) ... ... ... ... 264 165 Dicrocceliu7n dendriticum ... ... ... .•• ... ... 265 166 Eggs of Dicroccelium dendriticum ... ... ... ... ... 266 167 M\xsLC\d\2i.o{ Dicroccelium dendriticum. (After Leuckart) ... ... 266 168 Echinostoma ilocanum. (After Brumpt) ... ... ... ... 268 169 Echinostoma ilocanum. (After Leiper) ... ... ... ... 268 170 Echinostoma malayanumyL.Q\^ex. (After Leiper) ... ... ... 269 171 Schistosoma hismatobium. (After Looss) ... ... ... ... 270 172 Transverse section through a pair of Schistosoma hcematobiutn in copula. (After Leuckart) ... ... ... ... ..• ... ... 271 173 Anterior end of the male Schistosoma hcematobiuvt. (After Looss) ... 271 174 Schistosoma hcematobium. (After Leuckart) ... ... ... ... 276 175 Schistosoma hcematobiu7Jt, owxvci oL (After Looss) ... ... ... 277 176 Schistosoma Japonicum. (After Katsurada) ... ... ... ... 278 177 Schistosojna japonicum. (After Katsurada) ... ... ... ... 279 178 Schistosojna japonicum. (After Looss) ... ... ... ... 279 \%o\ Schistosoma japonicum ixoxciAog. (After Katsurada) ... ... ... 280 181 J 182 Schistosoma japonicum. (After Catto) ... ... ... ... 281 183 Schistosoma japonicum. (After Katsurada) ... ... ... ... 282 184 Schematic representation of a small part of a transverse section of Ligula sp. (After Blochmann) ... ... ... ... ... ... 287 185 Half of a transverse section through a proglottis of Tcenia crassicollis ... 288 186 Dipylidium (anintim. (After Benham) ... .. ... ... 289 187 Longitudinal section of the head and neck of Tcenia crassicollis ... ... 290 188 Tcenia ccenurus. (After Niemisec) ... ... ... ... ... 291 189 Yo\xx\g Acanthobothrium coronatnm. (After Pintner) ... ... ... 292 190 Scolex of a cysticercoid from Arion sp. (After Pintner) ... ... ... 292 191 Proglottis of 7>«z'a .yo^/irzocepka/us /attts ... ... 311 206 Fairly mature proglottis of Z)z<^c//zrz£>^^//^a/?^j /a/M^ ... ... ... 311 207 Dibothriocephahis lahis, (After Benham and Schauinsland) ... ... 312 208 VXerocercoid of Dibothriocephalus latus ... ... ... ... 313 209 A piece of the body wall of the Burbot, Z^/d! zv//^aw ... ... ... 313 210 CephaWc end of Dibothriocephalus cordatHs. (After Leuckart) ... ... 315 211 Diplogonoporus grandisy\X\^ey 1899. (After Ijiraa and Kurimoto) ... 317 212 Diplogonoporus grandis. (After Ijima and Kurimoto) ... ... ... 317 213 Cephalic end of ^^^fl^-^awww ;//a«j-^«/, Cobb. (After Leuckart) ... ... 318 214 Sparganuin mansoni . (After Ijima and Murata) ... ... ... 318 215 Sparganum pro lifer. (After Ijima) ... ... ... ... •••319 216 Spai'ganum prolifefitm. (After Stiles) ... ... ... ... 319 217 Dipyliditim caninuiu. (After Diamarr) ... ... ... ... 320 218 Dipylidium caninum. (After Benham and Moniez) ... ... ... 320 219 Dipylidium caninum : central portion of a proglottis. (After Neumann and Railliet) ... 220 Dipylidium caninum : development of embryo. (After Benham, Grassi, and Rovelli) ... 221 Larva (cysticercoid) of Dipylidium caninum. (After Grassi and Rovelli) 222 Hymenolepis nana, v. Sieb. (After Leuckart) ... 223 Hymenolepis nana: head. (After Mertens) 224 Hymenolepis nana : an egg. (After Grassi) 225 Longitudinal section through the intestinal villus of a rat. (After Grassi and Rovelli) ... 226 Hymenolepis nana {murina) : cross-section of proglottis from a rat. (After v. Linstow) 227 Hymenolepis nana : longitudinal section of an embryo. (After Grassi and Rovelli) ... 228 Hytnenolepis di^ninuta. (After Zschokke) 229 Hymenolepis diminuta. (After Grassi) 230 Hymenolepis diminuta. (After Bizzozero) 231 Hymenolepis diminuta. (Stephens, after NicoU and Minchin) 232 Hymenolepis lanceolata. (After Krabbe) 233 Hymenolepis lanceolata. (After Wolffhiigel) 234 ScoXeyi of Davainea madagascariensis. (After Blanchard) 235 Two fairly mature proglottids of 7<2;/za WzV/w ... 236 WeaA of Tcenia solium 237 Large and small hooks of Tcenia solium. (After Leuckart) 238 TcEtiia soliiwi. (After Leuckart) 239 Two mature proglottids of 7}2«/a Wzz/w 240 Large and small booklets of Tctnia marginaia. (After Leuckart) ... 241 yfa.twxe segment of TiEnia saginata 242 Cephalic end of Tcenia saginata 243 Tania saginata. (After Leuckart) 244 A piece of the muscle of the ox, with three specimens of Cysticercus bovis (After Ostertag) ... 245 "^.a-twresegrnent of Tcenia africana. (After v. Linstow) ... 246 Proglottis of Tenia africana. (After v. Linstow) 247 Head of Tcenia africana. (After v. Linstow) ... XXVlll THE ANIMAL PARASITES OF MAN PAGE Fig. 248 Tania confiisa. (After Guyer) ... ... ... ... ... 344 249 Tania confusa. (After Ward) ... ... ... ... ... 344 250 Tcenia echinococcus ... ... ... ... ... ... ... 345 251 Echinococcus veterinorw)i. (After Leuckart) ... ... ... ... 347 ^ r Diagrams of mode of formation of brood capsule and scolices (Stephens) ... 348 253 Section through an invaginated echinococcus scolex. (After Deve) ... 350 254 A piece of the wall of an Echinococcus veterinoriivi stretched out and seen from the internal surface ... ... ... ... ... ••• 35° 255 Echinococcus hominis in the liver. (After Ostertag, from Thomas) ,.. 351 256 Section through an echinococcus scolex in process of vesicular metamorphosis. (After Deve) ... ... ... ... ... ... ... 35^ ^^' \ Diagram of transformation of a scolex into a daughter cyst. (Stephens) ... 352 257A ) 258 Hooklets of echinococcus. (After Leuckart) ... .. ... ... 355 259 Echinococcus multilocularis \n\\\^\w^x o{\\i^ o\. (After Ostertag) ... 357 260 Diagram of a transverse section of .(^j^flr^V /MW(5rzV^?V/(fj. (After Brandes) ... 362 261 Anlerior end of an Ascaris megahce/>ha/a. (After Nassonow) ... ... 362 262 Transverse section through Ascaris lumbricoides at the level of the oesophagus behind the nerve ring. (After Goldschmidt) ... ... ... .■• 3^4 263 Schematic representation of the nervous system of a male Ascaris megalo- cephala. (After Brandes) ... ... ... ... ... ... 3^5 264 Diagram of female genitalia ... .. ... ... ... ... 3^8 264A Diagram of male genitalia of a strongylid ... ... ... ... 368 265 Transverse section through the ovarian tube of Belascaris cati of the cat ... 369 266 lA'sXo. Q{\!nex\\2}o^\\\z ioxvn oi Angiostonmin nigrovenosuiii ... ... 370 267 Transverse section through the posterior extremity of the body of Ascaris lumbricoides {maXe) ... ... ... ... ... ••• 37^ 268 Hind end of a male Ascaris lumbricoides cut across at the level of the dilator cells of the gut. (After Goldschmidt) ... ... ... ... 371 269 A piece of the trunk muscle of the pig with encapsuled embryonic Trichinge... 373 270 Strongyloides siercoralis, itmSiU. (After Looss) ... ... ... 380 271 Strongyloides siercoralis, mzXe. (After Looss) ... ... ... ... 380 272 Strongyloides stercoralis,iexfX2\e. (After Loo.ss)... ... ... ... 382 273 Strongyloides stercoralis. (After Looss) ... ... ... ... 3^2 274 Strongyloides stercoralis. (After Looss) ... ... ... ... 3^3 275 Gnathostoma siamense. (After Levinsen) ... ... ... ... 385 276 Gmnedi worm {Dracuncul us medinensis). (After Leuckart) ... ... 387 277 Anterior extremity of Guinea worm. (After Leuckart) ... ... ... 387 278 Dracunculus medinensis. (After Claus) ... ... ... ..'. 3^7 279 Transverse section of female Guinea worm. (After Leuckart) ... ... 388 280 Cyclo/>s virescens, (emsde ... ... ... ... ... ... 3^9 281 Filaria bancrofti. (After Leiper) ... ... ... ... ... 39^^ 282 Mf. bancrofti in thick film, dried and stained with hsematoxylin. (After Fiilleborn) ... ... ... ... ... ... ... 397 283 Schematic drawings of the anatomy of Ml. loa and Mf. bancrofti. (After Fiilleborn) ... ... ... ... ... ... ... 399 284 F. demarquayi. (After Leiper) ... ... ... ... ... 4^3 285 Mf. demarquayi \n thick film, dried and stained with haematoxylin. (After Fiilleborn) ... ... ... ... ... ... ... 4^4 286 Filaria {Tj conjunctivie. (After Addario) ... ... .. ... 405 287 Filaria (?) conJunclivcE. (After Grassi) ... ... ... ... 405 288 Setaria equina. (After Railliet) ... ... ... ... ... 408 289 Setaria equina : anterior end. (After Railliet) ... ... ... ... 408 290 Loa loa : the anterior end of the male. (After R. Blanchard) ... ... 410 291 Zca /^a ; anterior portion of the female. (After Looss) ... ... ... 410 LIST OF ILLUSTRATIONS XXIX PAGE Fig. 292 Loa loa in situ. (After Flilleborn and Rodenwaldt) ... ... ... 410 293 Loa loa : male and female. (After Looss) ... ... ... ... 410 294 Loa loa: the hind end of a male and of a female. (After Looss) ... ... 411 295 Loa loa : lateral view of tail of male showing papillae. (After Lane and Leiper) 411 296 Loa loa. (After Leiper) ... ... ... ... ... ... 411 297 i^. /m : in thick film, dried and stained with hsematoxylin. (After Fiilleborn) 413 298 Acanthocheilonema Persians. (After Leiper) ... ... ... ... 414 299 Mf. ferstans. (After Flilleborn) ... ... ... ... ... 415 300 Dirofilaria magalhdesi. (After v. Linstow) ... ... ... ... 417 301 Trichuris trichiura ... ... ... ... ... ... ... 420 302 Trichinella spiralis. (After Claus) ... ... ... ... ... 422 303 Isolated muscular fibre of a rat, invaded by Trichinella. (After Hertwig- Graham)... ... ... ... ... ... ... ... 425 304 Calcified Trichinella in the muscular system of a pig. (After Ostertag) ... 426 305 Various phases of the calcification of Trichinella of the muscles ... ... 426 306 Diociophyme gigas. (After Railliet) ... ... ... ... ... 432 307 'E'ggs oi Dioctophy77te gigas. (After Railliet) ... ... ... ... 432 308 Metastrongylus apri. (Stephens) ... ... ... ... ... 433 I Irichostrongyhis instabilis. (After Looss) ... ... ... ... 434 ^ I Trichostrongylus probolurus. (After Looss) ... ... ... ... 435 3 3 \ Trichostrongylus vitrinus. (After Looss) ... ... ... ... 436 3H i 3^5 \ Hcemonchus contortus. (After Ransom) ... ... ... ... 437 316 [ 317 Mecistocirrus fordi. (After Stephens) ... ... ... ... ... 439 318 Ternidens deminuius. (After Railliet and Henry) ... ... .. 440 \CEsophagostoviumstephanostomiim\2Ji.thomasi. (After Thomas) ... ... 442 r CEsophagostomum stephanostomnm var. thomasi. (After Thomas)... ... 444 ^^■^ i Ancylosloma duode7iale, male a.nd (emsile. (After Looss) ... ... 446 324 ) 325 Ancylostoma duodenale, showing ventral teeth. (After Looss) ... ... 447 326 Ancylostoma duodenale : diagrammatic representation of excretory system. (After a drawing by Looss) ... ... ... ... ... 448 327 Ancylostoma duodenale. (After Railliet) ... ... ... ... 449 328 Ancylostoma duodenale : bursa of male. (After Looss) ... ... ... 450 329 Ancylostoma duodenale : eggs in different stages of development. (After Looss) ... ... ... ... ... ... ... ... 451 330 Ancylostoma duodenale : larva. (After Leichtenstern) ... ... ... 452 331 Ancylostoma dtwdejiale. (After Looss) ... ... ... ... 453 332 Ancylostoma ceylanicum. (After Looss) ... ... ... ... 456 333 Ancylostoma hraziliense. (After Gomez de Faria) ... ... ... 456 334 Necator americamis. (After Looss) ... ... ... ... ... 457 335 Necator ame7-icanus : lateral view. (After Looss) ... ... ... 458 336 Necator americanus : bursa of male. (After Looss) ... ... ... 458 337 Syngamus kingi : anterior end of male. (After Leiper) ... ... ... 460 338 Syngamus kingi: anterior end of female. (After Leiper) ... ... 460 339 'B\xx?>2i oi Syngamus tj-ackealis. (Stephens) ... ... ... ... 461 340 Physaloptera mordens, luti^^eXy igoy. (After Leiper) ... ... ... 462 341 Ascaris lumbricoides. (From Claus) ... ... ... ... ... 4^3 342 0\\xxi\ o{ Ascaris lumbricoides... ... ... ... ... ... 4^3 34^ Owwm o[ Toxasca7is limbata ... ... ... ... ... ... 466 344 Transverse section through the head pait of Belasca7-is cati from the cat. (After Leuckart) ... ... ... ... ... ••■ ... 466 Fig. XXX THE ANIMAL PARASITES OF MAN PAGE ^^ \ Male a.nd iemale o( Ox^uris verf/ii'cu/aris ... ... ... ... 4^8 347 Oxytiris verviicularis : egg freshly deposited ... ... ... •■• 4^8 348 0:rj/wrzV z;(?r/;^/V«/am ; egg twelve hours after deposition ... ... ... 468 ^^^KT^ienxaXe oi Echinorhynchus augustatus ... ... ... ... 47^ 348B Anterior portion of the female apparatus of Echinorhynchus acus. (After Wagener) ... ... ... ... ... ... ... 47^ 348c Yj^^oi Echinorhynchus gigas. (After Leuckart) ... ... ... 477 348D The internal organs of the leech. (After Kennel) ... ... ... 480 348E Hirudo tnedicinalis, (After Claus) ... ... ... ... ... 481 349 Leptns atitwiinaHs. (After Gudden) ... ... ... ... ... 485 350 Leptus auhimnalis. (After Trouessart) ... ... ... ... 485 351 The kedani mite. (After Tanaka) ... ... ... ... ... 487 352 Tetratiychus telarius \2i.x. rzisseolns,\\.oc\\. (After Artault) ... ... 488 353 Pediculoides ventricostis. (After Laboulbene and Megnin) ... ... 489 354 Nephrophages sangutnai'his, male, ventral surface. (After Miyake and Scriba) 490 355 Nephrophages sangtiinarius : female, dorsal aspect. (After Miyake and Scriba) ... ... ... ... ... ... ... ... 490 356 Tydeits inolestus. (After Moniez) ... ... ... ... ... 491 357 Dermatiyssus gallincB. (After Berlese) ... ... ... ... 492 358 Dermanyssus hirtindinis. (After Delafond) ... ... ... ... 492 359 Ixodes ricinuSy \\\z\e. (After Pagenstecher) ... ... ... ... 498 360 Yt.vi\z\eoi Ixodes ricinus. (After Pagenstecher) , ... ... ... 498 361 Argas rejlextis. (After Pagenstecher) ... ... ... ... ... 506 362 Argas per sictis. (After Megnin) ... ... ... ... ... 507 363 Tyroglyphus farince : male. (After Berlese) ... ... ... ... 512 364 Tyroglyphus longior, Gqiw. (After Fum. and Robin) ... ... ... 512 365 Rhizoglyphus parasiticus : male and female. (After Dalgetty) ... ... 514 366 Histiogaster [entomophagus ?) spermaticus. (After E. Trouessart) ... ... 515 367 Sarcoptes scabiei. (After Fiirstenberg)... ... ... ... ... 518 368 Sarcoptes scabiei : male, ventral aspect. (After Fiirstenberg) ... ... 519 369 Sarcoptes minor var. cati. (After Raiiliet) ... ... ... ... 521 370 Demodex folliculorum oi the diOg. (After Megnin) ... ... ... 522 371 Lingiiatula rhinaria : female ... ... ... ... ... 524 372 \jax\z.oi Linguatula rhinaria {Pentastoma denticttlatum). (After Leuckart) 524 373 linguaiula rhinaria. (After M. Koch) ... ... ... ... 525 374 MowXh-y^zxX.^ o^ Pedictdus vestimenti. (After Denny) ... ... ... 533 375 Ovum of the head louse ... ... ... ... ... ... 533 376 Head louse, male ... ... ... ... ... ... ... 533 377 Pediculus vestimentiy B\xxn\. : adult female ... ... ... ... 533 378 Phthirius inguinalis, Leach... ... ... ... ... ... 534 379 Head of the bed bug from the ventral surface ... ... ... ... 535 380 Derniatophilus penetrans : young female. (After Moniez) ... ... 544 381 Derniatophilus penetrans : older female. (After Moniez) ... ... 544 382 Pulex irrilans ... ... ... ... ... ... ... 546 383 Larva of flea. (After Raiiliet) ... ... ... ... ... 546 384 Fulex serraticeps ... ... ... ... ... ,,, ... 546 385 Head of a male and of a female Anopheles. (After Giles) ... ... 549 386 Head of a male and of a female Culex. (After Giles) ... ... ... 549 387 yio^^\\i-\>2iX\% oi Anopheles claviger . (After Grassi) ... ... ... 550 388 Anopheles fjiaculipennis. (Afier Nuttall and Shipley) ... ... ... 550 389 Longitudinal section of an Anopheles, showing alimentary canal. (After Grassi) ... ... ... ... ... ... ... ... 551 390 Anopheles tNaculipennis,yie\gen. (After Grassi) ... ... ,.. 552 391 'La.wa.oi Anopheles maculipenniSfYahx. (After Grassi) ... ... ... 553 392 Larva of Culex. (After Grassi) ... ... ... ... ... 553 LIST OF ILLUSTRATIONS XXxi PAGE F^G. 393 Fupz o( j4nop/ieks macu/ipenniSy Me\g. (Afler Grass!) ... ... ... 554 394 Heads of Culex and Anopheles. (After Daniels) ... ... ... 556 395 Eggs of Culex, of Anopheles, of Stegomyia, of Taeniorhynchus, and of Psorophora ... ... ... ... ... ... ... 557 396 Diagram showing the structure of a typical mosquito. (Theobald) ... 558 397 Types of scales, head and scutellar ornamentation, forms of clypeus. (Theobald, etc., etc.) ... ... ... ... ... ... 559 398 Neuration of wing. Explanation of wing veins and cells. (Theobald) ... 560 399 Wmg of ^nophe/es piacti/ifenmsy Meigea ... ... ... ... 566 4CXD Wing of a Culex ... ... ... ... ... ... ... 575 401 Wing of Simulium ... ... ... ... ... ... ... 579 402 Wing of Chironomus ... ... ... ... ... ... 579 403 A Ceratopogon, or midge ... ... ... ... ... ... 580 404 An owl midge, Phlebotonnis sp. (From Giles's "Gnats or Mosquitots ") ... 581 405 \jax\^o{ Homaloniyia canicular is ... ... ... ... ... 585 406 'L3lx\2& oi Calliphora vomitoria ... ... ... ... ... 585 407 Larva of Chrysomyia niacellaria. (After Conil) ... ... ... 585 408 The screw-worm fly (Chjysomyia macellaria) ... ... ... ... 587 409 Ochromyia larva on the skin of man, South Africa. (After Blanchard) ... 590 410 Head end of •• larva of Natal." (After Gedoelst) ... ... ... 591 411 Lund's larva. (After Gedoelst) ... ... ... ... ... 593 412 Dermaiobia noxialis, GoxxdiOi ... ... ... ... ... ... 597 413 IjSiXVd^oi Derniatobia cyaniventris. (After Blanchard) ... ... ... 597 414 'L.z.xvz.of Derviatobia cyani7fenttis. (After Blanchard) ... ... ... 597 415 The ox gad fly (7a/5a«z^f <5(7t^2««j, Linn.) ... ... ... ... 601 416 'Y\\t.\yi\vc\'^ {Hceinatopota phivialis.,\Jvaxi.) ... ... ... ... 602 417 Head of Glossina longipalpis. (Afier Griinberg) ... ... ... 604 418 Antenna of Glossina pal iidipes, male. (After Austen) ... ... .. 604 419 Glossina palpalis 2ind Y)\xil)yi\nxii. (After Brumpt) ... ... ... 607 420 The tsetse-fly {Glossina morsitans^ West wood) ... ... ... ... 608 421 The stinging fly {Stomoxys calcitransy Linn.) ,.. ... ... ... 609 422 Trichomonas irom Q.2tz\xm 2ixv^ gyxX oi X2X. (Original, Fantham) ... ... 735 We regret to have taken without permission from the *' Transactions of The Society of Tropical Medicine and Hygiene," London, the following diagrams : — Pages Figures 268 No. 169 269 „ 170 391 „ 281 411 „ 295 and 296 414 „ 298 460 „ 337 and 338 and tender our regret to the Society in question for havint^ done so. ^ XXX THE ANIMAL PARASITES OF MAN PAGE ^ Y Male and {emale of Oxyuri's ver/nuularis ... ... ... ■•• 4^8 347 Oxyurts vermicularis : egg freshly deposited ... ... ... ••• 468 348 Oxyuris vermicularis : egg twelve hours after deposition ... ... ... 468 T,^2>A T'\\Qvaz\& oi Echinorhynchus atigtistatus ... ... ... ... 47^ 348B Anterior portion of the female apparatus of Echinorhynchiis acus. (After Wagener) ... ... ... ... ... ... ••• 47^ 348c 'Egg oi Echinorhynchus gigas. (After Leuckart) ... ... ... 477 348D The internal organs of the leech. (Afier Kennel) ... ... ... 480 348E Hirudo inedicinalis. (After Claus) ... ... ... . . ... 481 349 Leptiis atiHiiunaHs. (After Gudden) ... ... ... ... ... 485 350 Lepttis autuimiaJis. (After Trouessart) ... ... ... ... 485 351 The kedani mite. (After Tanaka) ... ... ... ... ... 487 352 Tetrajiychus telarius va.x. rzisseolus,\\.och. (After Artault) ... ... 488 353 Pediadoides ventricosiis. (After Laboulbene and Megnin) ... ... 489 354 Nephrophages sangtdnarius, male, ventral surface. (After Miyake and Scriba) 490 355 Nephrophages sanguiriarius : female, dorsal aspect. (After Miyake and Scriba) ... ... ... ... ... ... ... ... 49° 356 Tydeiis molestiis. (After Moniez) ... ... ... ... ... 491 357 Dermauyssus gallincB. (After Berlese) ... ... ... ... 492 358 Dermanyssus hirundinis. (After Delafond) ... ... ... ... 492 359 Ixodes ricinus, m2L\e. (After Pagenstecher) ... ... ... ... 498 360 Femsde oi Ixodes riciftus. (After Pagenstecher) . ... ... ... 498 361 Argas tejiextis. (After Pagenstecher) ... ... ... ... ... 506 362 Argas per sicus. (After Megnin) ... ... ... ... ... 507 363 Tyroglyphus farina : male. (After Berlese) ... ... ... ... 512 364 Tyroglyphtis longior, Getv. (After Fum. and Robin) ... ... ... 512 365 Rhizoglyphus parasiticus : male and female. (After Dalgetty) ... ... 514 366 Histiogasier {entomophagus }) spermaticus. (After E. Trouessart) ... ... 515 367 Sarcoptes scabiei. (After Fiirslenberg)... ... ... ... ... 518 368 Sarcoptes scabiei : male, ventral aspect. (After Furstenberg) ... ... 519 369 Sarcoptes minor sax. cati. (After Raiiliet) ... ... ... ... 521 LIST OF ILLUSTRATIONS XXXI PAGE ^^G- 393 Vn^pdi oi Anopheles macuIipenniSf'M.ti^. (After Grassi) ... ... ... 554 394 Heads of Culex and Anopheles. (After Daniels) ... ... ... 556 395 Eggs of Culex, of Anopheles, of Stegomyia, of Taeniorhynchus, and of Psorophora ... ... ... ... ... ... ... 557 396 Diagram showing the structure of a typical mosquito. (Theobald) ... 558 397 Types of scales, head and scutellar ornamentation, forms of clypeus. (Theobald, etc., etc.) ... ... ... ... ... ... 559 398 Neuration of wing. Explanation of wing veins and cells. (Theobald) ... 560 399 'W'mgoi Anopheles niaailifemns,MQ\gen ... ... ... ... 566 400 Wing of a Culex ... ... ... ... ... ... ... 575 401 Wing of Simulium ... ... ... ... ... ... ... 579 402 Wing of Chironomus ... ... ... ... ... ... 579 403 A Ceratopogon, or midge ... ... ... ... ... ... 580 404 An owl midge, Phlebotomns sp. (From Giles's "Gnats or Mosquitots ") ... 581 405 \j3LX\z.oi Honialojnyia canicularis ... ... ... ... ... 585 406 \jZXV2& oi Calliphora vomitoria ... ... ... ... ... 585 407 Larva of Chrysomyia viacellaria. (After Conil) ... ... ... 585 408 The screw-worm fiy {Chrysofnyia macellana) ... ... ... ... 587 409 Ochromyia larva on the skin of man, South Africa. (After Blanchard) ... 590 410 Head end of ** larva of Natal." (After Gedoelst) ... ... ... 591 411 Lund's larva. (After Gedoelst) ... ... ... ... ... 593 412 Der77iatobia noxialis, Goudot ... ... ... ... ... ... 597 413 'L^rwz. o{ Dermatohia cyaniventris, (After Blanchard) ... ... ... 597 414 l^dirvQ. oi Derfnatobia cyani7ientr is. (After Blanchard) ... ... ... 597 415 The o\ g2id ^y {Tabamt^ bovimts^ Uinn.) ... ... ... ... 601 416 The hr\m^ {HiBniatopoia plnvialiSfUinn.) ... ... ... ... 602 417 Hedid oi Glossina longipalpis. (After Grunberg) ... ... ... 604 418 Antenna of Glossina pallidipes^ male. (After Austen) ... ... .. 604 419 Glossina palpalis zxv^ }^\x^xr\wvcv. (After Brumpt) ... ... ... 607 420 The tsetse-fly {Glossina inorsitans. West wood) ... ... ... ... 608 421 The stinging fly (Stofnoxys calcitransy Linn.) ... ... ... ... 609 422 7>/V^tfw<7«aj- from caecum and gut of rat. (Original, Fantham) ... ... 735 423 Chilomastix {Tetramitus) mesnili. (Original, Fantham) ... ... 736 XXXll THE ANIMAL PARASITES OF MAN ERRATA. P. 31, line 6 from bottom: dekte "human," as Leidy really worked with Endamceba blaticBy parasitic in the gut of the cockroach. ^ P. 43, line 12 from bottom : for "John's " read "Johns." P. 44, line 13 from hotiora: for ^"^ Amceba buccalis, Sternberg," read ^^ Amoeba buccalisy Steinberg." P. 46, line 13 from top : for " breath " read ** breadth." P. 53, In footnote *, line 6 from bottom : insert "see" before Arch. f. Protistenk. P. 75 : To paragraph regarding development of the parasite in the fly's salivary glands, add that the crithidial phase takes two to five days. P. Ill, line 8 from top : the date of Sangiorgi should be 191 1. P. 142, line 7 from top : insert " Genus." before Eimeria. P. 252, Insert heading "Family. Opisthorchiidae, Braun, 1901," above "Sub-family. Opisthorchiinae, Looss, 1899." P. 351, description of fig. 255, line 3 : for " Thoma " read " Thomas." P. 471, line 15 from bottom : for " alcohol 100 parts " read "alcohol 100 c.c. " P. 472, line II from bottom : for " Or (2) 10 per cent, formalin^^^ read " Or (2) fix in hot 10 per cent, formalin.''^ P. 493, line 21 from top : for " Conoy " read " Couvy." P. 589, line 2 from top : for " carnosa" read " carnaria." P. 620, line 15 from top : for " fo " read " of." P. 622, line 12 from bottom : delete comma after quantity. P. 626, line 6 from bottom : delete comma after Mackie (1915). P. 638 : insert title "TREMATODES" above that of " Fascioliasis. ' ' P. 709, line 9 from bottom : omit second Pediculus capitis. P. 748, line 8 from top : for " cytologica " read " cy tological. " P. 753, line 4 from bottom : for ** Fercocercous " read ** Furcocercous." P. 755, line 7 : for " Oncocerca " read " Onchocerca." ON PARASITES IN GENERAL, By the term PARASITES is understood living organisms which, for the purpose of procuring food, take up their abode, temporarily or permanently, on or within other Hving organisms. There are both phints and animals (Phytoparasites and Zooparasites) which lead a parasitic life in or upon other plants and other animals. Phytoparasites are not included in the following descriptions of the forms of parasitism, but a very large number of animal parasites (zooparasites) are described. The number of the latter, as a rule, is very much underrated. How great a number of animal parasites exists may be gathered from the fact that all classes of animals are subject to them. Some of the larger groups, such as Sporozoa, Cestoda, Trematoda and Acaiithocephalaj consist entirely of parasitic species, and parasitism even occurs among the vertebrates {Myxine). It therefore follows that the characteristics of parasites lie, not hi their structure, but in the manner of their existence. Parasitism itself occurs in various ways and degrees. According to R. Leuckart, we should distinguish between OCCASIONAL (temporary) and PERMANiiNT (stationary) PARASITISM. Occasional parasites, such as the flea {Pnlex irritans), the bed-bug {Cimex lechUarius) , the leech (Hinido niedicinalis), and others, only seek their " host " to obtain nourishment and find shelter while thus occupied. Without being bound to the host, they usually abandon the latter soon after the attainment of their object {Cimex, Hirudo), or they may remain on the body of their host throughout their entire development from the hatching of the ^gg {Pediculiis). It follows from this mode of living that the occasional parasites become sometimes distinguishable from their free-living relatives, though only to a slight extent. It is, there- fore, seldom difficult to determine the systematic position of temporary parasites from their structure. In consequence of their mode of life, all these temporary parasites live on the external surface of the body of their host, though more rarely they take up their abode in cavities easily accessible from the exterior, such as the mouth, nose and gills. They are therefore frequently called Epizoa or Ectoparasites ; but these designations 2 THE ANJMAL PARASITES OF MAN do not cover only the temporary parasites, because numerous epizoa (as for instance the louse) are parasitic during their entire life. In contradistinction to these temporary parasites, the permanent parasites obtain shelter as well as food from their host for a long period, sometimes during the entire course of their life. They do not seek their host only when requiring nourishment, but always remain with it, thus acquiring substantial protection. The permanent para- sites, as a rule, live within the internal organs, preferably in those which are easily accessible from the exterior, such as the intestine, with its appendages. Nevertheless, permanent parasites are also found in separate organs and systems, such as the muscular and vascular systems, hollow bones and brain, while some live on the outer skin. Here again, the terms Entozoa and Endoparasites do not include all stationary parasites ; to the latter, for instance, the lice belong, which pass all their life on the surface of the body of their host, where they find shelter and food and go through their entire development. The ectoparasitic trematodes, numerous insects, Crustacea, and other animals live in the same manner. All *^ Helminthes," however, belong to the group of permanent parasites. This term is now applied to designate certain lowly w^orms which lead a parasitic life (intestinal worms) ; but they are not all so termed. For instance, the few parasitic Turbellaria are never classed with the helminthes, although closely related to them. The turbellarians, in fact, belong to a group of animals of which only a few members are parasitic, whereas the helminthes comprise those groups of worms of which all species (Cestoda, Treinatoda, Acantho- .cephala)f or at least the majority of species {Nematoda), are parasitic. Formerly the Linguatulidse {Petitastoma) were classed with the helminthes because their existence is also endoparasitic, and because the shape of their body exhibits a great similarity to that of the true helminthes. vSince the study of the development of the Linguatulidae (P. J. van Beneden, 1848, and R. Leuckart, 1858) has demonstrated that they are really degenerate arachnoids, they have been separated from the helminthes. It is hardly necessary to emphasize the fact that the helminthes or intestinal worms do not represent a systematic group of animals, but only a biological one, and that the helminthes can only be dis- cussed in the same sense as land and water animals are mentioned, i.e.y without conveying the idea of a classification in such a grouping. It is true that formerly this was universally done, but very soon the error of such a classification was recognized. Still, until the middle of last century, the helminthes were regarded as a systematic group, although C. E. V. Baer (1827) and F. S. Leuckart (1827) strenuously -opposed this view. Under the active leadership of J. A. E. Goeze, PERMANENT PARASITISM 3 J. G. H. Zeder, J. G. Bremser, K. A. Rudolphi and F. Dujardin, the knowledge of the hehninthes (hehninthology) developed into a special stud}', but unfortunately it lost all connection with zoology. It required the intervention of Carl Vogt to disestablish the helminthes as one class of animals, by uniting. the various groups with those of the free-living animals most closely related to them {Platyhelminthes, N ematlielinintlies). Permanent parasitism in the course of time has caused animals adopting this mode of life to undergo considerable, some- times even striking, bodily changes, permanent ectoparasites having as yet undergone least, alteration. The latter sometimes bear so unmistakably the likeness to the group to which they belong, that even a superficial knowledge of their structure and appearance often suffices for the recognition of their systematic position. For instance, though the louse, like many decidedly temporary parasites, has lost its wings — a characteristic of insects — in consequence of parasitism, yet nobody would deny its insect nature ; such also occurs in other temporary parasites {Ciniex, Pidex), On the other hand, the changes in a number of permanent ectoparasites (such as parasitic Crustacea) are far more considerable, and correspond with those that have occurred in permanent endoparasites. These alterations depend partly on retrogression and partly on the acquisition of new peculiarities. In the former case, the change consists in the loss of those organs which have become useless in a permanent parasitic condition of existence, such as wings in the louse, and the articulated extremities seen in the larval stage of parasitic Crustacea. The loss of these organs goes hand in hand with the cohesion of segments of the body that were originally separate, and alterations in the muscular and nervous systems. In the same manner another means of locomotion is lost — the ciliated coat — which is possessed by many permanent parasites during their larval period. To all appearances, this character is not secondary and recently acquired, but represents a primary character inherited from free-living progenitors, and still transmitted to the altered descendants, because of its use during the larval stage (e.g., the larvae of a great many Trematodes, the oncospheres of some Cestodes). Amongst the retrogressions, the loss of the organs of sense may be mentioned, particularly the eyes, which are still present, not only in the nearest free-living forms but also in the free-living larvae of true parasites. It is only quite exceptionally that the eyes are subsequently retained, as a rule they are lost. Lastly, in a great many cases the digestive system also disappears, as in parasitic Crustacea, in a few nematodes and trematodes, in all cestodes and Acanthocephala. There remain at most the rudiments of the muscles of the fore -gut, but these are adapted to entirely different uses. 4 THE ANIMAL PARASITES OF MAN The new characters which permanent parasites may acquire are, first of all, the remarkably manifold clasping and clinging organs, which are seldom (as in parasitic Crustacea) directly joined on to already existing structures. In those instances in which organs for the conveyance of food are retained, these likewise frequently undergo transformation, in consequence of the altered food and manner of feeding. Such alterations consist, for instance, in the transformation of a masticating mouth apparatus into the piercing and sucking organs of parasitic insects. Hermaphroditism (as m Trematodes, Cestodes, and a few Nema- todes) is a further peculiarity of many permanent parasites ; moreover, the association in couples that occurs, especially in trematodes, may lead to complete cohesion and, exceptionally, also to re-separation of the sexes. In many cases the females only are parasitic, while the males live a free life, or there may be in addition the so-called complementary males. Occasionally the male alone is parasitic, and in that case lives within the female of the same species, which may live free, like certain Gephyrea {Bonellia) ; or the female also may be parasitic, as Trichosoma crassicaudum, which lives in the bladder of the sewer rat {Miis decumanns). We have numerous proofs that demonstrate how considerably the original features of many parasites have become changed. We need only draw attention to the aforementioned Linguatulidae, also to many of the parasitic Crustacea belonging to various orders. In all of these a knowledge of the larval stages — in which there is no alteration, or at most only a slight degree of change — serves to determine their systematic position, i.e.^ the nearest conditions of relationship. The most remarkable changes are observed in those groups that contain only a few parasitic members, the majority leading a free life. A striking instance is afforded by a snail, the well-known Ento- concha mirabilts, Miiller. This mollusc consists merely of an elongated sac living in a Holothurian {Synapta digitata). It possesses none of the characteristics of either the Gastropoda or any molluscs, and in its interior there is nothing to be observed but the organs of generation and the embryos. Nevertheless, the Entoconcha is decidedly a parasitic snail, as is clearly proved by its larvae, but it is a snail which, in consequence of parasitism, has lost all the characteristics of molluscs in its mature condition, but still exhibits them in the early stages of development. Certain nematodes show very clearly to what devious courses parasitism may lead. The Atractonema gibbosnin, the life-history of which has been described by R. Leuckart, and which lives in the larvae and pupae of a dipterous insect (Cecidoniyia), exhibits, in its FERTILITY OF PARASITES 5 early stage, the ordinary characteristics of other threadworms. A few weeks later — the males having died off immediately after copu- lation — the females are transformed into spindle-shaped bodies, the mouth and anus of which are closed. They carry with them an irregu- larly shaped appendage, in which the segmenting ova are situated, and in which the further conditions of life of the Atradotiema are accomplished. A minute examination has demonstrated that this appendage is the prolapsed and enlarged vagina of the animal which has become merely a supplementary attachment. The conditions present in the Sphcendaria, the nematoid nature of which was long undiscovered, are still more remarkable. It was only when Siebold proved that typical nematodes were hatched from their eggs that their nature was recognized. The nematodes thus produced have not the slightest resemblance to the parent. The researches of Lubbock, A. Schneider, and more particularly of R. Leuckart, have shown that what we call Sphcerularia homhi is not an animal but merely an organ — the vagina — of a nematode worm. This vagina at first grows, sac-like, from the body of the tiny nema- tode ; it gradually assumes enormous dimensions (2 cm. in length) ; it contains the sexual organs and parts of the intestine. The remain- ing portion of the actual animal then becomes small and shrivelled ; it may be easily overlooked, being but an appendage to the vagina with its independent existence, and it finally disappears altogether. The GREAT FERTILITY of parasites is another of their peculiarities, though this may be also the case to a certain degree with some of the free-living animals, the progeny of which are likewise exposed to enormous destruction. More remarkable, however, is the fact that the young of the endoparasites only very exceptionally grow to maturity by the side of their parents. Sooner or later they leave the organ inhabited by tlie parents, frequently reach the open, and after a shorter or longer period of free existence seek new hosts. During their free period, moreover, a considerable growth may be attained, or metamorphosis may take place, or even multiplication. In the exceptional cases in which the young remain within the same host, they nevertheless usually quit the organ inhabited by the parents. They likewise rarely attain maturity within the host inhabited by the parents, but only, as in other cases, after having gained access to fresh hosts. These transmigrations play a very important rdle in the natural history of the internal parasites, but they frequently conceal the cycle of development, for sometimes there are INTERMEDIATE GENERATIONS, which themselves invade intermediate hosts. Even when there are no intermediate generations, the system OF intermediate HOSTS is frequently maintained by the endoparasites. 6 THE ANIMAL PARASITES OK MAN According to the kind of food ingested by parasites, it has recently become usual to separate the true parasites from those animals that feed on the superfluity of the food of the host, or on products which are no longer necessary to him, and to call the latter MESSMATES or COMMENSALS. As examples, the Ricinidae are thus designated, because, like actual lice, they dwell among the fur of mammals or the plumage of birds. They do not, however, suck blood, for which their mouth apparatus is unsuited, but subsist on useless epidermic scales. These epizoa, according to J. P. van Beneden, are, to a certain extent, useful to their hosts by removing deciduous materials which under certain circumstances might become harmful to them.^ This investigator, who has contributed so greatly to our knowledge of parasites, assigns the Ricines to the MUTUALISTS, under which term he comprises animals of various species which live in common, and confer certain benefits on one another. The mutualists are usually intimately connected in a mutually advan- tageous association known as '^ symbiosis."^ Incidental and Pseudo Parasites. — In many cases the parasites are confined to certain hosts, and may therefore be designated as specific to such hosts. Thus, hitherto, Tcenia solinm and Tcenia saginata in their adult condition have only been found in man ; Tcenia crassi- collis only in the cat ; Brandesia (Distoma) turgida and Halipegus {Distoma) ovocaudatus only in Rana esculenta, and so forth. In many other cases, however, certain species of parasites are common to several, and sometimes many, species of hosts ; Dipylidium caninnm is found in the domestic cat as well as in the dog; Fasciola hepatica is found in a large number of herbivorous mammals (nineteen species), Diplodiscus (Amphistomunt) suhclavatus in numerous urodele and ecaudate am- phibia, Holostomnm variahile in about twenty-four species of birds, and so on. In these cases the hosts are almost invariably closely related, belonging, as a rule, to the same family or order, or at any rate to the same class. Trichinella spiralis^ which is found in man, and in the pig, bear, rat, mouse, cat, fox, badger, polecat and marten, and is capable of being artificially cultivated in the dog, rabbit, sheep, horse, in other mammals, and even in birds, is one of the most striking exceptions. Some parasites are so strictly confined to one species of host that, even when artificially introduced into animals very closely related ^ According to Sambon, the Ricinidae are by no means advantageous to their hosts. These Hemipterous parasites give rise to an intolerable itching which may cause loss of rest, emacia- tion, and sometimes even death. Birds suffering from phthiriasis of the Ricines are usually in bad health. - For further information on these conditions, see "Die Schmarotzer des Thierreichs," by P. J. van Beneden, Leipzig, 1876 ; and *' Die Symbiose," by O. Hertwig. INCIDENTAL AND PSEUDO-PARASITES 7 to their normal host, they do not thrive, but sooner or later, often very quickly, die off, and very rarely establish themselves. For example, repeated attempts have been made to rear the adult Tcenia soli mil in the dog, or to rear Cysticercus cellulosce in the ox, or the Cyst icerc lis of Tcenia saginata in the pig, but they have always proved unsuccessful. Only exceptionally has it been possible to transfer Ccenunis cerebralis, the larval stage of a tapeworm (Tcenia ccenurus) of the dog from the brain of the sheep to that of the domestic goat. On the other hand, in the case of the Trichinellae transference to a different host is easily accomplished. Under natural conditions, it is not uncommon for certain kinds of specific parasites to occur occasionally in unusual hosts. Their relationship to the latter is that of incidental parasites. Thus Echinorhynchiis gigas, a specific parasite of the pig, is only an incidental parasite of man ; Fasciola hepatica and Dicroccelium lanceatnm are specific to numerous kinds of mammals, but may be found incidentally in man. On the other hand, Dibothriocephalus latuSj a specific parasite of man, may occasionally take up its abode in the dog, cat and fox. As a rule, all those parasites of man that are only rarely met with, notwithstanding that human beings are constantly being observed and examined by medical men, are termed incidental parasites of man. In many cases we are acquainted with the normal or specific host of these parasites. Thus we know the specific host of Balantidium colt, Eimeria stiedcBf Fasciola hepatica, Dipylidiuin caninuin, etc. ; in others the host is as yet unknown. In the latter case the question partly relates to such forms as have been so deficiently described that their recognition is impossible, partly to parasites of man in various regions of the earth, the Helminthes and parasites of which are totally unknown or only slightly known, or finally to early developmental stages that are difficult to identify. Animals that usually live free, and exception- ally become parasitic, may likewise be called incidental parasites. In this category are included a few Angiiillulidce that have been observed in man ; also Leptodera appendicidata, which usually lives free, but may occasionrJly become parasitic in black slugs {Arion einpiricoriun) : when parasitic it attains a larger size, and produces far more eggs than when living a free life. In order to avoid errors, the term " incidental parasites " should be confined to true parasites which, besides living in their normal host, may also live in other hosts. Leuckart speaks of facultative parasitism in such forms as Leptodera. L. Oerley^ succeeded in artificially causing Leptodera (Rhabditis) pellio to assume facultative parasitism by introducing these * Oerley, L., *' Der Rhabditiden und ihre medizinische Bedeutung," Berlin, 1886, p. 65. 8 THE ANIMAL PARASITEvS OF MAN worms into the vagina of mice, where the parasites remained aHve and multipHed. Lcptodera pellio dies in the intestines of mammals and man ; it remains alive in frogs, but always escapes into the open with the faeces. Recently the incidental parasites of man have also been called "Pseudo-parasites" or " Pseudo-helminthes." Formerly, how- ever, these terms were applied not only to living organisms that do not and cannot live parasitically, and that only exceptionally and incidentally get into man, but also to any foreign bodies, portions of animals and plants, or even pathological formations that left the human system through the natural channels, and the true nature of which was misunderstood. Frequently these bodies were described as living or dead parasites and labelled with scientific names, as if they w^ere true parasites. A study of these errors, which formerly occurred very frequently, w^ould be as interesting as it would be instructive. It is better not to use the expression pseudo-parasites for incidental parasites, but to keep to the original meaning, for it is not at all certain that pseudo-parasites are not described, even nowadays. The Influence of Parasites on the Host. — In a great many cases, we are not in a position to state anything regarding any marked influence exercised by the parasite on the organism, and on the con- ditions of life, of the host. Most animals and many persons exhibit few signs of such influence, an exception being infestation with helminthes and certain other parasites w^hich produce eosinophilia in the blood. As a general rule, the parasite, which is always smaller and weaker than its host, does not attempt to endanger the life of the latter, as simultaneously its own existence would be threatened. The parasite, of course, robs its host, but usually in a scanty and sparing manner, and the injuries it inflicts can hardly be taken into account. There are, however, numerous cases^ in which the situation of the parasites or the nature of their food, added to their number and movements, may cause more or less injury, and even threaten the life of the host. It stands to reason that a Cysticercus cellulosce situated in the skin is of but slight importance, whereas one that has penetrated the eye or the brain must give rise to serious disorders. A cuticular or intestinal parasite is, as a rule, less harmful than a blood parasite. A helminth, such as an Ascaris lunibricoides or a tapeworm, that feeds on the residues of foodstuffs w^ithin the intestine, will hardly ' Liihe, M., " Ueber d. Fix. d. Helm. a. d. Darmwand ihrer Wirthe u. die dadurch verursachten path-anat. Veranderungen d. Wirthsdarmes," Trans, of IVth Intern. Zool. Cong., Berlin, 1901 ; Mingazzini, P., '* Ric. sul var. modo di fiss. delle tenie alia par. int. e sul loro assorbimento," Ric. Lab. Anat. Roma e altri Lab. bioL, vol. x, 1904; Shipley, A. E., and E. G. Fearnsides, "The Effects of Metazoan Parasites on their Hosts," /ottrn, Econ. Biol., 1906, i, 2. THE INFLUENCE OF PARASITES 9 afifect its host by depriving it of this material. The case is different when the parasites are very numerous, especially when the heavily infested host happens to be a young individual needing all it ingests for its own requirements, and therefore unable to sustain the drain of numerous intruders in the intestine. Disturbances also set in more rapidly when the intestinal helminthes are blood-suckers, the injury to the host resulting from the kind of food taken by the parasite. Generally, the disorders caused by loss of chyle are insignificant when compared with those induced by the growth and agglomera- tion of the helminthes. The latter may cause chiefly obstructions of small vessels or symptoms of pressure in affected or contiguous organs, with all those complications which may arise secondarily, or they may even lead to the complete obliteration of the organ invaded. Of course the symptoms will vary according to the nature of the organ attacked. In consequence also of the movements of the parasites, disorders are set up that may tend to serious pathological changes of the affected organs. The collective migrations, undertaken chiefly by the embryos of certain parasites (as in trichinosis, acute cestode tuberculosis), are still more harmful, as are also the unusual migrations •of other parasites, which, incidentally, may lead to the formation of so-called worm abscesses or to abnormal communications (fistulae) between organs that are contiguous but possess no direct connection. Recently, several authors have called attention to the fact that the helminthes produce substances that are toxic to their host ; and II le effects of such poisons explain the pathology of helminthiasis far more satisfactorily than the theory of reflex action. In a number of cases these toxic materials (leucomaines) have been isolated and their effects on living organisms demonstrated by actual experiments. It also appears that the absorption of materials formed by the decomposition of dead helminthes may likewise cause toxic effects. However, our knowledge of these conditions is as yet in its initial stage. ^ Nearly all the symptoms caused directly or indirectly by parasites are of such a nature that the presence of the parasites cannot be diagnosed with any certainty, or only very rarely. The most that can be done is to deduce the presence of parasites by the exclusion of other causes. Fortunately, however, there are sufficient means 1 Moursson et Schlagdenhaufifen, "Nouv. rech. clin. et phys. sur quelq. liquides organ.,*' €. R. Acad. Set., Paris, 1882, p. 791 ; Debove, '* Del'intox. hydat.,"^«//. et Mint. Soc. m'ed. des Hopit., 1888 ; Linstow, v., " Ueb. d. Giftgehalt d. helm.," Internat. Monatsschr. f. Anat. u. Phys., xiii, 1896; Peiper, **Z. Symptomatol. der thier. Paras.," Deutsche med. Wochenschr., 1897, No. 40; Mingazzini, P., *' Ric. sul veleno d. elm. int.," Rass. intern, d. med. modern. Ann., 1901, ii, No. 6; Vaullegeard, A., " Etud. exp. et crit. sur Taction d. helm.," Btill. Soc. Linn, de Normandie, 1901, 5, Ser. T, vii, p. 84, and others. lO THE ANIMAL PARASITES OF MAN by which we may confirm the diagnosis in a great many cases. Such means consist not only in a minute examination of the patient by palpation, percussion and local inspection, but also in the micro- scopical examination of the natural secretions and excretions of the body, such as sputum, nasal mucus, urine and fpeces. Though such examinations may entail loss of time, they are necessary in the interest of the patient. It appears, moreover, that quackery, which has gained considerable ground even in the treatment of the helminthic diseases of man, can thus be considerably limited. Origin of Parasites} — In former times, when the only correct views that existed related to the origin of the higher animals, the mode of multiplication of parasites as well as of other lowly animals was ascribed to spontaneous generation {geueratio ceqnivoca), and this opinion prevailed throughout the middle ages. The writers on natural science merely devoted their time to the interpretation of the views of the old authors, and perpetuated the opinions of the ancients on questions, which, even in those days, could have been correctly explained merely by observation. It was only w^hen observations were again recommenced, and the microscope was invented, that the idea of spontaneous generation became limited. Not only did the microscope reveal the organs of generation or their products (eggs) in numerous animals, but Redi succeeded in proving that the so-called Helcophagi (flesh maggots) are only the progeny of flies, and never appear in the flesh of slaughtered animals when fully developed flies are prevented from approaching and depositing their eggs on it. Swammerdam likewise knew that the *^ worms" living in the caterpillars of butterflies were the larvae of other insects (ichneumon flies) which had laid their eggs in their bodies ; he also discovered the ova of lice. The tw^o authors men- tioned were, however, unwilling to see that the experience they had gained regarding insects applied to the helminthes. Leeuwenhoek also vehemently opposed the theory of a spontaneous generation, maintaining that, on a basis of common-sense, eggs, or at all events germs, must exist, even though they could not be seen. The use of the microscope also revealed a large number of very small organisms in the water and moist soil, some of which un- doubtedly resembled helminthes. Considering the wide dissemination of these minute organisms, it was natural to conjecture that after their almost unavoidable introduction into the human system they should grow into helminthes (Boerhave, Hoffmann). Linnaeus went even further, for he traced the descent of the liver-fluke of sheep from ' Die Geschichte der *' Klinisch wichtigen Parasiten," behandelt H. Vierordt im **Handb. d. Gesch. d. Med. hrsg." v. M. Neuburger u. J. Pagel, Bd. ii, 1903. ORIGIN OF PARASITES II a free-living planaria {Dendroccelum lacteuui), the Oxyuris verniicnlaris from free-living nematodes, and the Tcenia lata (i.e., Dibothriocephalus latus) from a tapeworm {Schistocephalits solidns) found free in the water. Linnaeus' statements met with general approval. However, we must bear in mind that at that time the number of helminthes known was very small, and many of the forms that we have long ago learned to differentiate as specific were then regarded as belonging to one species. Linnaeus' statements were partly supported by similar discoveries by other investigators, such as Unzer, and partly also by the discovery of eggs in many helminthes. It was believed that the eggs hatched in the outside world gave rise to free-living creatures, and that these, after their introduction into the intestine, were trans- formed into helminthes. By means of these eggs the old investigators tried to explain the hereditary transmission of the intestinal worms, which was universally believed until the commencement of the last century. Some authors went so far as to regard the intestinal worms as congenital or inherited ; they maintained the possibility of direct transmission, as in suckling, and denied that the eggs reaching the external world had anything to do with the propagation of the parasites. The more minute comparison between the supposed free-living stages of the helminthes and their adult forms, and the impossibility of finding corresponding free forms for the ever-increasing number of parasitic species, revealed the improbability of Linnaeus' statements (O. Fr. Miiller). It was the latter author also who recognized the origin of the tapeworms {Schistocephalits, Ligtila) found free in the water. They originate from fishes which they quit spontaneously. However, in spite of the fact that van Doeveren and Pallas correctly recognized the significance of the eggs in the trans- mission of intestinal worms, these statements remained disregarded, as did Abildgaard's observation, experimentally confirmed, that the (immature) cestodes from the abdominal cavity of sticklebacks became mature in the intestines of aquatic birds. Moreover, at the end of the eighteenth and the commencement of the nineteenth centuries, after helminthology had been raised to a special branch of study by the successful results of the investigations of numerous authors (Goeze, Bloch, Pallas, Miiller, Batsch, Rudolphi, Bremser), many of whom experienced a ^' divine joy " in searching the intestines of animals for helminthes, some authors reverted to generatio a^quivoca, without, however, entirely denying the existence of organs of genera- tion and eggs. The fact that a few nematodes bore living progeny — a fact of which Goeze w^as already aware — had no influence on the erroneous opinion, as in such cases it was considered that the young continued to develop beside the old forms. There were also 12 THE ANIMAL PARASITES OF MAN many helminthes known that never developed sexual organs and never produced eggs, and which therefore were referred to generatio cequivoca. People were convinced that the intestinal mucous mem- brane or an intestinal villus could transform itself into a worm, either in a general morbid condition of the body, or in pathological changes of a more local character. The appearance of helminthes was even regarded as useful and as a means for the expulsion of injurious matter. These views, firmly rooted and supported by such eminent authorities as Rudolphi and Bremser, could not easily be overthrown. First, a change took place in the knowledge of the trematodes. In 1773, O. Fr. Miiller discovered Cercaricv living free in water. He regarded them as independent creatures and gave them the name that is still used at the present time. Nitzsch, who also minutely studied these organisms and who recognized the resemblance of the anterior part of their bodies to a Fasciola, did not, however, arrive at a correct conclusion. He regarded the combination rather as that of a Fasciola with a Vibrio, for which he mistook the charac- teristic tail of the cercaria. He also noticed the encystment (trans- formation into the *'pupa") on foreign bodies of many species of these animals, but was of opinion that this process signified only the termination of Hfe. Considerable attention was attracted to the matter when Bojanus first published a paper entitled ''A Short Note on Cercaria and their Place of Origin." He pointed out that the cercariae creep out of the ''royal yellow^ worms," which occur in freshwater snails {LuiuKxa, Pakidina)y and are probably generated in these worms. Oken, in whose journal, his (1818, p. 729), Bojanus published his discovery, remarks in an annotation, " One might lay a wager that these Cercariae are the embryos of Distomes." Soon after (1827), C. E. V. Baer was able to confirm Bojanus' hypothesis that the cercariae as a '' heterogeneous brood " originated from spores in para- sitic tubes in snails (germinating tubes). Moreover, Mehlis [his, 183 1, p. 190) not only discovered the opercula of the ova of Distoma, but likewise saw the infusorian-like embryo emerge from the eggs of Typhloccelmn {Monostomum) flaviim and Cathcemasia {Distoma) hians. A few years later (1835) v. Siebold observed the embryos (miracidia) of the Cyclocoelnm {Monostomum) mutabile, and discovered in their interior a cylindrical body that behaved like an independent being (" necessary parasite"), and was so similar in appearance to the ''royal yellow worms " (Bojanus) that Siebold considered the origin of the latter from the embryos of trematodes as, at all events, possible. Meanwhile, V. Nordmann of Helsingfors had in 1832 seen the miracidia of flukes provided with eyes swimming in water; v. Siebold (1835) had observed ORIGIN OF PARASITES 13 the embryos, or oncospheres, of tapeworms furnished with six booklets in the so-called eggs of the Taenia;- while Creplin (1837) had discovered the 'Mnfusorial" young of the Diphyllobothrium {Bothrio- cephahis) ditrcninui, and conjectured that similar embryos were to be found in other cestodes with operculated eggs. At all events, the fact was established that the progeny of the helminthes appeared in various forms and was partly free living. The researches of Eschricht (1841) were likewise of influence, as they elucidated the structure of the Bothriocephali, and proved that the encysted and sexless helminthes were merely immature stages. J. I. Steenstrup (1842) was, however, the first to furnish explana- tions for the numerous isolated and uncomprehended discoveries. Commencing with the remarkable development of the Coelenterata, he established the fact that the Helminthes, especially the endopara- sitic trematodes, multiply by means of alternating and differently formed generations. Just as the polyp originating from the Qgg of a medusa represents a generation of medusae, so does the germinal tube ('^ royal yellow w^orm ") originating from the ciliated embryo of a Distoma, etc., represent the cercaria. These w^ere consequently regarded as the progeny of trematodes, and Steenstrup, guided by his observations, conjectured that the cercaria, whose entrance into the snails he had observed accompanied by the simultaneous loss of the propelling tail, finally penetrated into other animals, in which they became flukes. Part of this hypothetical cycle of development was erroneous, and in other particulars positive observation was lacking, but the path pursued was in the right direction. Immediately after the appear- ance of Steenstrup's celebrated work, v. Siebold expressed his opinion that the encapsuled flukes certainly had to travel, i.e.y to be trans- mitted with their bearers into other hosts, before becoming mature. This view was experimentally confirmed by de Filippi, La Valette St. George (1855), as w^ell as by Pagenstecher (1857), while the meta- morphosis of the ciliated embryo of Distoma into a germinal tube was first seen by G. Wagener (1857) in Gorgodera (Distoma) cygnoides of frogs. All that we have subsequently learned from the works of numerous investigators about the development of endoparasitic trema- todes has certainly increased our knowledge in various directions, and, apart from the deviating development of the Holostomidce has, as a whole, confirmed the briefly sketched cycle of development. Steenstrup's work on the cestodes did not attract the same atten- tion as his work on trematodes. Steenstrup always insisted on the '^ nurse" nature of the cysticerci and other bladder-worms. Abildgaard (1790), as well as Creplin (1829 and 1839), had already furnished the information that certain sexless cestodes (Schistocephalns and 14 THE ANIMAL PARASITES OF MAN Ligtila) from the abdomen of fishes only become mature after their transference to the intestine of aquatic birds. These passive migrations were confirmed in an entire series of other cestodes, particularly by v. Siebold (1844, 1848, 1850) and E. J. van Beneden (1849), not by actual experiment^ but by undoubted observation. It was correctly believed that the ova or oncospheres penetrate into certain intermediate hosts, in which they develop into unseg- mented larvae. Here they remain until, with their host, they are swallowed by some predacious animal. They then reach the intestine, being freed from the surrounding membranes through the process of digestion, and settle themselves there to form the adult chain of proglottides. Though some few scientists, such as P. J. van Beneden and Em. Blanchard, deduced from these observations that the bladder-worms (Cysticerci), which had hitherto been regarded as a separate class of helminthes, were only larval Taeniae, this correct view was not at first universally accepted. The foundation was too slight, and van Beneden was of opinion that the Cysticerci were not necessary, but only appeared incidentally. V. Siebold was a strenuous opponent to this theory, notwith- standing his experiences on the change of hosts of the Tetra- rhynchus. Together with Dujardin (1850) he conjectured that the Taeniae underwent a deviating cycle of development. He w^as of opinion that the six-hooked oncospheres left the lintestine, in which the older generation lived, and were scattered about with the faeces, and finally re-entered per os (i.e., with water and food) a host similar to the one they had left, in the intestine of which they w^ere directly transformed into tapew^orms. A change of host such as occurred in other cestodes was not supposed to take place (the history of the cestodes was at this time not entirely estab- lished). As the oncospheres of the Taenia are enveloped in one calcareous or several softer coverings which they cannot leave actively, and as, in consequence of this condition, innumerable oncospheres cannot penetrate into an animal, and others cannot reach the proper animal, v. Siebold conceded, at least for the latter, the possibility of a further development. But this was only supposed to occur because they had either invaded wrong hosts, or, having reached the right hosts, had penetrated organs unsuit- able to their development, and had thus gone astray in their travels, and had become hydropically degenerated taeniae. This was v. Siebold's explanation of bladder-worms. Naturally, v. Siebold himself conjectured that a recovery of the diseased tapeworm might occur, in a few exceptional cases, after transmission into the correct host, as, for instance, in the CysiicerciLS fasciolaris of mice, the host of which is the domestic cat, and in which there is a seemingly normally ORIGIN OF PARASITES 15 developed piece of tapeworm situated between the caudal vesicle and the cysticercus head. Guided by correct views, F. Kiichenmeister undertook in Zittau the task of confirming the metamorphosis of Cysticercus pisiformis of hares and rabbits, into tapeworms in the intestine of the dog by means of feeding experiments. The first reports on the subject, published in 1851, were not likely to meet with universal approval, because Kiichenmeister first diagnosed the actual tapeworm he had been rearing as Tcenia crassiceps, afterwards as Tcenia serrata, and finally as Tcenia pisiformis n. sp. However, in any case, Kiichenmeister, by means of the reintroduction of experimental investigation, rendered a great service to helminthology. The publication of Kuchenmeister's works induced v. Siebold to undertake similar experiments (1852 and 1853), which were partly published by his pupil Lewald in 1852. But the positive results obtained hardly changed Siebold's opinion, for although he no longer considered the bladder-worms as hydropically degenerated tapeworms, he still regarded them as taeniae that had strayed. The change of opinion was partly due to an important work of the Prague zoologist, v. Stein (1853). He was able to examine the development of a small bladder-worm in the larvae of the well-known meal-worm {Tenebrio molitor) and to demonstrate that, as Goeze had already proved in the case of Cysticercus fasciolaris of mice, first the caudal vesicle is formed and then the scolex, whereas Siebold believed that in bladder-worms the posterior end of the scolex was formed first, and that this posterior end underwent a secondary hydropic degeneration. In opposition to v. Siebold, Kiichenmeister successfully proved the necessity of the bladder- worm stage by rearing tapeworms in dogs from the Cysticercus tenuicollis of domestic mammals and from the Ccenurus cerebralis of sheep. He, and simultaneously several other investigators independently, succeeded, with material provided by Kiichenmeister, in rearing the Ccenurus cerebralis in sheep from the oncospheres of the Tcenia ccenurus of the dog (1854). R. Leuckart obtained similar results in mice by feeding them with the mature proglottides of the Tcenia crassicollis of cats (1854). Kiichenmeister also repeatedly reared the Taenia solium of man from the Cysticercus cellulosce of pigs (1855), and from the embryos of this parasite P. ]. van Beneden succeeded in obtaining the same Cysticercus in the pig (1854). As Kiichenmeister distinguished the Tcenia mediocanellalaj known to Goeze as Tcenia saginata, amongst the large taeniae of man (1851), so it was not long before R. Leuckart ■(1862) succeeded in rearmg the cysticercus of the bookless tapeworm in the ox. It is particularly to this last-named investigator that helminthology is indebted more than to any other author. He l6 THE ANIMAL PARASITES OF MAN followed the gradual metamorphosis from oncospheres to cystic worms in all its details. In view of all the researches that were made, and which are too numerous to mention individually, the idea that bladder-worms are abnormal or only incidental forms had to be abandoned. Every- thing pointed to the fact that in all cestodes the development is divided between two kinds of animals ; in one — the host, the adult tapeworm is found ; while in the other, the intermediate host, we find some form or other of an intermediate stage (cysticercus in the broadest sense). The practical application of this knowledge is self-evident. If no infected pork or beef is ingested, no tapeworm can be acquired, and also the rearing of cysticerci in the human body is prevented by avoidmg the introduction of the eggs of tapeworms. Though these results were definitely proved by numerous researches, yet they have been repeatedly challenged, notably by J. Knoch (1862) in Petrograd, who, on the basis of experiments, sought to confirm a direct development without an intermediate host and ciliated stage, at all events as regards Dibothriocephalus laUis. However, the repeated communications of this author met with but little favour from competent persons, partly because the experiments were conducted very carelessly, and partly because their repetition on dog and man (R. Leuekart) had no results (1863). It was only in 1883 that Braun was able to prove that the developmental cycle of Dibothriocephalus latus is similar to that of other Cestodes. The results obtained in other places by Parona, Grassi, Ijima and Zschokke render any discussion of Kiichenmeister's conclusions unnecessary.^ Long after Knoch, a French author, P. Megnin, also pleaded for the direct development of some cestodes, and especially some taeniae. He (1879) also sought to prove a genetic connection between the bookless and armed tape- worms of mammals, but the arguments he adduced, so far as they rest on observations, can be easily refuted or attributed to mis- interpretation. Only one of these arguments is correct, namely, that the number of the species of taeniae with which we are acquainted is far larger than that of the corresponding cystic forms ; but this disparity alone cannot be taken as a proof of direct development. It can only be said that our knowledge in this respect is deficient. As a matter of fact, we have during recent years become acquainted with a large number of cystic forms, hitherto unknown, belonging to taeniae which have long been familiar. It must also be borne in mind that no man in his lifetime can complete an examination for bladder-worms of the large number of insects, for instance, which ' Refer to the collected literature under Dibothriocephahis latus, and the reply to Klichen- meister by Braun ('* Ueber den Zwischenwirt des breit. Bandw." Wurzb. : Stuber, 1886). DEVELOPMENT OP^ PARASITES \*J may destroy an entire generation of an insectivorous species of bird within a small district. Naturally it does not follow that direct development in the cestodes is altogether lacking. The researches of Grassi (1889) have furnished an example in Hyiiienolepis {iceiiia) imirina, which shows that development may sometimes take place without an intermediate host, notwithstanding the retention of the cystic stage. It was found that the oncospheres of this species, introduced into rats of a certain age, after a time grow into tapeworms without leaving the intestine, but not directly, for they bore into the intestinal wall, where they pass the cystic stage, the cysts afterwards falling into the intestinal lumen, where they develop into tapeworms. The recent experiments of NicoU (1911) show that the larval stages of Hyinenolepis uiurina also occur in the rat-flea, Cera tophyllits fasciatus. Important observations were soon made on the remaining groups of helminthes. The discussion on the origin of parasites soon became confined to the helminthes. Amongst the Nematoda, it had long been known that encapsuled forms existed that had at first been regarded as independent species, but very soon they were pronounced to be immature forms, in consequence of their lack of sexual organs. Though Dujardin and also v. Siebold regarded them as ''strayed" animals, v. Stein (1853) very promptly demon- strated that the progeny of the nematodes were destined to travel by discovering a perforating organ in the larval nematodes of the mealworm. This was first experimentally confirmed (i860) by R. Leuckart, R. Virchow and Zenker, all of whom succeeded not only in bringing to maturity the muscle Trichinae (known since 1830) in the intestine of the animals experimented upon, but were likewise able to follow the migrations of the progeny. Of course, the encapsulating brood remained in the same organism, and in this respect deviated from the broods of other helminthes which escape into the outer world and find their way into other animals, but the encapsuled nematodes could no longer be regarded as the result of straying. Subsequently, R. Leuckart worked out, more or less completely, the history of the development of numerous nematodes, or pointed out the way in which further investigations should be made. It has been found that in nematodes far more frequently than in other helminthes, the typical course of develop- ment is subject partly to curtailment and partly to complications, which sometimes considerably increase the difficulties of investigation and have hitherto prevented the attainment of a definite conclusion, though the way to it is now clear. In a similar manner the works of R. Leuckart have cleared up the development of the Acanthocephala and Lingttatulida. Of l8 THE ANIMAL PARASITES OF MAN course, mucli still remains to be done. So far, we do not even know all the helminthes of man and of the domestic animals in all their phases of life, and still less is known of those of other animals. We are indebted to the discoveries of the last fifty years for the knowledge arrived at, though compara- tively few names are connected with it. The gross framework is revealed, but the gaps have only been filled up here and there. However, we may trustfully leave the completion of the whole to the future, without fear that any essential alterations will take place. The deductions to be drawn are as follows : That the helminthes like the ectoparasites multiply by sexual processes, that the entire course of development of the helminthes is rarely or never gone through in the same host as is the case with several ectoparasites, that the progeny at an earlier or later stage of development, as eggs, embryos, or larvae, quit the host inhabited by the older generation, and almost always attain the outer world : only in Trichinella does the develop- ment take place directly in the definite host. Where the eggs have not yet developed they go through the embryonic evolution in the outer world. The young larvae are transmitted, either still enclosed within the Qgg or embryonic covering, to the intermediate host or more rarely they are transferred straight to the final host. In other cases they may hatch out from their envelopes, and after a longer or shorter period of free life, during which they may partake of food and grow% they, as before, penetrate, usually in an active way, into an intermediate host, or at once invade the final host. Exceptionally {e.g.f Rhabdonema), during the free life there may be a propagation of the parasitic generation, and in this case only the succeeding genera- tion again becomes parasitic, and then at once reaches its final host. The young forms which have invaded the final host become mature in the latter, or after a longer or shorter period of parasitism again wander forth (as the CEstridae, Ichneumonidae, etc.), and reach the adult stage in the outer world. The young stages, during which the parasites undergo metamorphoses or are even capable of producing one or several intermediate generations, are passed in the intermediate hosts until, as a rule, they are passively carried into the final host and there complete their cycle of development by the formation of the organs of generation. This mode of development, the spending of life in two different kinds of animals (intermediate and final host), is typical of the helminthes. This is manifested in the Acanthocephala, the Cestoda, the majority of the endoparasitic Trematoda, a number of the Nematoda, and the Linguatulidae. There are now and then exceptions, however, in which, for instance, the host and intermediate host change order {Trichinella y Hymenolepis miirina). DERIVATION OF PARASITES I9 Parasites are hardly ever inherited amongst animals.^ According to a few statements, however, Trlcliinella and Cceniirus are supposed to be transmissible from the infected mother to the foetus. Other- wise most animals acquire their parasites, especially the Entozoa, from without, the parasites penetrating either actively, as in animals living in the water, or passively with food and drink. A particular pre- disposition to worms is not more likely than a spontaneous origin of parasites. Derivation of Parasites. — Doubt now no longer exists as to the derivation of the temporary and of many of the stationary ecto- parasites from free-living forms. This conclusion is founded on the circumstance that not only are there numerous intermediate degrees in the manner of living and feeding between predacious and parasitic animals, but that there is more or less uniformity in their structure. The differences that exist are easily explained as consequences of altered conditions of life. The case is more difficult in regard to groups that are exclusively parasitic (Cestoda, Treniatoda, Acantho- cephala, Liiiguatiilidce, and Sporozoa), or groups that are chiefly parasitic {Nematoda), because in these cases the gulf that divides these forms from free-living animals is wider. It is true that we know that the nearest relatives of the Lingtiatuiidoi are found amongst the Arachnoidea, and indeed in the Acarlna ; that, moreover, the structure and development of the Sporozoa refers them to the Protozoa^ and allows some of them to be regarded as the descendants of the lowest Rhlzopoda, We know that the Trematoda, and through these the Cestoda, are closely related to the Turbellaria, from which they may be traced. The Neinatoda, and still more the Acanthocephala, stand apart. This is less evident, how^ever, in the Nematoda, for there are numerous free-living members of these from which it is possible that the parasitic species may be descended. Indeed, this seems more than probable if such examples as Leptodera, Rliabdonema and Strongyloides are taken into consideration, as well as the conditions of life of free-living nematodes. These mostly, if not exclusively, spend their lives in places where decomposing organic substances are present in quantities ; some species attain maturity only in such localities, and there propagate very rapidly. Should the favourable conditions for feeding be changed, the animals seek out other localities, or they remain in the larval form for some time until more favourable conditions set in. It is comprehensible that such forms are very likely to adopt a parasitic manner of life which at first is facultative {Leptodera, Angnillula), but may be regarded as the tran- ' However, in the Protozoa there are examples of hereditary transmission of parasites, e.g., in the case of Babesia {Piroplasma) bovis and Babesia cants in their invertebrate hosts (ticks) ; in Crithidia melophagia and Crithzdia hyalommce ; and in the case of SpirocJueta diittoni in its invertebrate host (a tick). 20 THE ANIMAL PARASITES OF MAN sition to true parasitism. The great advantages attached to a parasitic Hfe consist not only in protection, but also in the supply of suitable food, and consequently in the easier and greater production of eggs, and thus fully account for the gradual passage of facultative parasitism into true parasitism. In many forms the young stages live free for some time {Strongylidce), in others, as is the case in Rhabdoiieina, parasitic and free-living generations alternate ; in others, again, the free period is limited to the agg stage or entirely suppressed. Though it is possible thus to connect the parasitic with the free- living nematodes, by taking their manner of life into account, this matter presents greater difticulties in regard to other helminthes. It is true that the segmented Cestoda may be connected with and traced from the less known and interesting single-jointed Cestoda {Ainphilina, Archigetes, Caryophyllc^us, Gyrocotyle). Trematodes are all parasites, with the exception of one group, Temnocephaiidiv, several genera and species of which live on the surface of the bodies of Crustacea and turtles of tropical and sub-tropical freshwaters. Teriinocephalid^^ nre, nevertheless, predacious. They feed on Infusoria, the larvae of small insects and Crustacea. So far as is known they do not nourish them- selves on part of the host. They belong to the group of commensals, or more correctly, to that of the space parasites, which simply dwell with their host and do not even take a portion of the superfluity of its food. However, space parasitism may still be regarded as the first stage of commensalism, which is again to be regarded as a sort of transition to true parasitism. It is possible that parasitism came about in this way in the trematodes, in which connection we must first consider the turbellaria- like ancestors of the trematodes. Much can be said in favour of such a genetic relationship between turbellaria and trematodes, and .hardly anything against it. It should also be remembered that amongst the few parasitic turbellaria there are some that possess clinging discs or suctorial pores, and these are only differentiated from ectoparasitic trematodes by the possession of a ciliated integu- ment, which is found only in the larval stages of the latter. The Acanthocephala occupy an isolated position. Most authors certainly regard them as related to the nematodes ; in any case, the connection is not a close one, and the far-reaching alterations which must have occurred prevent a clear view. Perhaps the free original forms of Acanthocephala are no longer in existence, but that such must have existed is a foregone conclusion. An explanation of the change of host so frequent in parasites is more difficult than that of their descent. R. Leuckart is of opinion that the present intermediate hosts, which belong principally to the lower animals, were the original hosts of the parasites, and fostered DERIVATION OF PARASITES 21 both their larval and adult stages. It was only in course of time that the original hosts sank to the position of intermediate hosts, the cause for this alteration being that the development of parasites, especially of the helminthes, through further development and differentiation extended over a larger number of stages. The earlier stages remained in their original hosts, but the later stages sought out other hosts (higher animals). To prove this, Leuckart points out that the mature stages of the helminthes, with but few exceptions, occur only in the vertebrates which appeared later in the development of the animal kingdom, while the great majority of intestinal worms of the lower animals only represent young stages, which require transmission into a vertebrate animal before they can become mature. The few helminthes that attain maturity in the lower animals (Aspidogaster, Archigetes) are therefore regarded by Leuckart as primitive forms, and he compares them with the developmental stages of helminthes, Aspidogaster with rediae, Archigetes with cystic ^rcoids. He classes the nematodes that become mature in the invertebrates with Angiiil- Inlidce, i.e., with saprophagous nematodes from which the parasitic species descend. Leuckart therefore regards the change of hosts as secondary, so does Sabatier. The latter, however, adduces other reasons for this (lack of clinging organs and the necessity to develop them in an intermediary stage) ; but in this connection he only considers the Cestoda. In opposition to Leuckart, R. Moniez, however, is con- vinced that the migrations of the helminthes, as well as the system of intermediate hosts, represent the original order of things. Moniez tiaces all Entozoa from saprophytes, but only a few of these were able to settle directly in the intestine and there continue their development. These are forms that at the present day still lack an intermediate host, such as Trichocephalus, Ascaris, and Oxyurls. In most other cases the embryos, however, consisted of such sapro- phytes as were, in other respects, suitable to become parasites, but were incapable of resisting the mechanical and chemical influences of the intestinal contents. They were therefore obliged to leave the mtestine at once, and accomplished this by penetrating the intestinal walls and burrowing in the tissues of their carriers. In this position, assisted by the favourable conditions of nutrition, they could attain a relatively high degree of development. Mechanical reasons pre- vented a return to the intestines, where the eggs could be deposited. Most of them doubtless died off as parasites, as also their young stages do at present when they penetrate WTong hosts. Some of them, nevertheless, passively reached the intestine of beasts of prey. Many were destroyed in the process of mastication ; for a small part, however, there was the chance of reaching the intestine of a beast 22 THE ANIMAL PARASITES OF MAN of prey undamaged, and there, having become larger and more capable of resistance, maturity was attained. By means of this in- cidental coincidence of various favourable circumstances, these pro- cesses, according to Moniez, have been established by heredity and Iiave become normal. This is not the place to express an opinion either for or against the various hypotheses advanced, but the existence of these diametri- cally opposed views alone will show the great difficulty of the question. Independently, however, it appears more natural to come to the conclusion that parasitism, as well as change of hosts, were gradual transitions. As a conclusion to this introductory chapter, a list of some of the most important works on the parasitology of man and animals is appended. LITERATURE. GOEZE, J. A. E. Versuch einer Naturgeschichte der Eingeweidewiirmer thieri- scher Korper. Blankenburg, 1782. 4to, 471 pp., with 44 plates. Zeder, J. G. H. Erster Nachtrag zur Naturgeschichte der Eingeweidewiirmer. von J. A. E. Goeze. Leipzig, 1800. 4to, with 6 tables. RUDOLPHI, C. A. Entozoorum sive vermium intestinalium historia naturalis. I, Amstelod., 1808; ii, i8og. 8vo, with 18 plates. RUDOLPHi, C. A. Entozoorum synopsis. Berol., 1819. 8vo, with 3 plates. Bremser, J. G. Ueber lebende Wiirmer im lebenden Menschen. Wien, 1819. 8vo, with 4 plates. Bremser, J. G. Icones helminthum, systema Rudolphii entozoologicum illus- trantes. Viennae, 1824. Fol. (Paris, 1837). DUJARDIN, F. Histoire naturelle des helminthes ou vers intestinaux. Paris, 1845. 8vo, with 12 plates. DiESING, C. M. Systema helminthum. 2 vols. Vindobonnae, 1850, 185 1. 8vo. Supplements by the same author : Revision der Myzhelminthen (Report of the Session of the Imp. Acad, of Science. Wien, xxxii, 1858) ; with addendum (ibid., xxxv, 1859); Revision der Cephalocotyleen (ibid., xlix, 1864, and xlviii, 1864) ; Revision der Nematoden (ibid., xlii, 1861) ; Supple- ments (ibid., xliii, 1862). Beneden, p. J. VAN. Memoire sur les Vers intestinaux. Paris, 1858. 4to, with 12 plates. KtJCHENMElSTER, F. Die in und an dem Korper des lebenden Menschen vorkom- menden Parasiten. Leipzig, 1855. 8vo, with 14 plates. Leuckart, R. Die menschlichen Parasiten und die von ihnen herriihrenden Krankheiten. I, Leipzig, 1863; II, Leipzig, 1876. 8vo. COBBOLD, T. Sp. Entozoa ; an Introduction to the Study of Helminthology. London, 1864. 8vo. Supplement, London, 1869. Davaine, C. Traite des entozoaires et des maladies vermineuses de I'homme et des animaux domestiques. 2nd edit. Paris, 1877. 8vo. LiNSTOW, O. V. Compendium der Helminthologie, ein Verzeichniss der bekannten Helminthen, die frei oder in thierischen Korpern leben, geordnet nach ihren Wohnthieren, unter Angabe der Organe, in dencn sie gefunden sind, und mit Beifiigung der Litteraturquellen. Hanov., 1878. 8vo. Supplement, including the years 1878-1888, Hanov., 1888. COBBOLD, T. Sp. Parasites; a Treatise on the Entozoa of Man and Animals, including some Account of the Entozoa. London, 1879. 8vo. Leuckart, R. Die Parasiten des Menschen und die von ihnen herriihrenden Krankheiten. 2nd edit. Leipzig, 1879-1886. The Protozoa, Cestodes, Trematodes and Hirudinea have hitherto appeared (continued by Brandes). BtJTSCHLi, O. Protozoa in Bronn's Klass. u. Ordn. d. Thierreichs. Vol. i, Leipzig, 1 880-1889. 8vo, with 79 plates. LITERATURE 23 Braun, M. Trematodes in Bronn's Klass. u. Ordn. d. Thierreichs. Vol. iv, i, Leipzig, 1879-1893. 8vo, with 33 tables. (The first thirteen sheets, com- prising the history of the worms up to 1830, were compiled by H. Pagenstecher.) ZlJRN, F. A. Die thierischen Parasiten auf und in dem Korper unserer Haus- saugethiere, sowie die durch erstere veranlassten Krankheiten, deren Behandlung und Verhiitung. 2nd edit. Weimar, 1882. 8vo, with 4 plates. COBBOLD, T. Sp. Human Parasites; a Manual of Reference to all the Known Species of Entozoa and Ectozoa. London, 1882. 8vo. KUCHENMEISTER, F., and F. A. ZURN. Die Parasiten des Menschen. 2nd edit. Leipzig, 1888., 8vo, with 15 plates. Blanchard, R. Traite de zoologie medicale. I, Paris, 1889; II, i8go, Svo. Neumann, L. G. Traite des maladies parasitaires non microbiennes des animaux domestiques. 2nd edit. Paris, 1892. Svo. English edit., trans- lated by G. Fleming. • 2nd edit., revised by J. Macqueen. 1905. London : Bailliere, Tindall and Cox. Looss, A. Schmarotzerthum in der Thierwelt. Leipzig, 1892. Svo. Railliet, a. Traite de zoologie medicale et agricole. 2nd edit. I, Paris, 1S95. Svo. ParoN!\, C. L'elmintologia italiana da' suoi primi tempi all' anno 1890. Genova, 1894. Svo. Braun, M. Cestoda in Bronn's Klass. u. Ordn. d. Thierreichs. Vol. iv, 2, Leipzig, 1S94-1900. Svo, with 24 plates. MOSLER, F., and E. Peiper. Thier Parasit. (Spec. Path. u. Ther. v. H. Nothnagel. Vol. vi.) Wien, 1894. Svo, with 124 illustrations. Laveran^ a., et R. Blanchard. Les hematozoaires de I'homme et des anim, Paris, 1895. i2mo, with 30 figs. Sluiter, C. R. De dierl, paras, v. d. mensch en van onze huisdier. Haag, 1895. Svo. Blanchard, R. Malad. parasit., paras, animaux, paras, veget. a I'exclus. des bacter. (Traite de pathol. gen. de Ch. Bouchard, vol. ii.) Paris, 1S95. Svo, with 70 figs. HUBER, J. Ch. Bibliographie der klin. Helminthol. Miinchen, 1S95. Svo. With Supplement, 1898, and continued as Bibl. d. klin. Entomol. Miinchen, 1S99-1900. MONIEZ, R. Traite de parasitol. anim. et veget. appl. a la medecine. Paris, 1896. Svo, with 116 figs. Weichselbaum. Parasitologie (Weil's Handb. d. Hyg.). Jena, 1S9S. Svo, with 78 illustrations, Kraemer, a. Die thierischen Schmarotzer des Auges (Grafe and Samische's Handb. d. ges Augenheilk.). Leipzig, 1S99. Svo, with 16 illustrations. Cholodkowsky, N. a. Icones helm, hominis. St. Petersburg, 1898-99. Fol. (atlas with 15 plates). Perroncito, E. I parassiti dell' uomo e degli animali utili e le piii comuni malattie da essi prodotti. Ila ed. Milano 1902. 8°. con 276 fig. e 25 tav. Stiles, Ch. W. and A. Hassall. Index Catalogue of Medicine and Veterinary Zoology. AVashington, 1902 (U.S. Dept. of Agric, Bur. of Anim. Ind., Bull. No. 39). Neveu-Lemaire, M. Precis de parasitologie humaine, parasites vegetaux et animaux. 4 e edit. Paris, 1911. Hofer, B. Handbuch der Fischkrankheiten. Miinchen, 1904. 8°. 18 Taf. 222 Abb, GuiART, J., and L. Grimbert. Precis de Diagnostic chimique, microscopique et parasitologique. Paris, 1906. With 500 figs. OSTERTAG, R. Handbuch der Fleischbeschau. V. Aufl. mit 265 Abb. Stutt- gart, 1904. Stiles, Ch. W. The International Code of Zoological Nomenclature as applied to Medicine (Hygienic Lab., Bull. No. 24, Washington, 1905)- Stiles^ C. W., and Hassall, A. Trematoda and Trematode Diseases. (Index Catalogue of Med. and Vet. Zoology.) Hygienic Lab., Bull. No. ZT^ Washington, 1908. 24 THE ANIMAL PARASITES OF MAN Stiles, C. \\., and Hassall, A. Cestoda and Cestodaria. Hygienic Lab., BulL No. 85, Washington, 1Q12. Laloy, L. Parasitisme et mutualisme dans la nature. Paris, igo6. 8vo, 284 pp., 82 figs. Theobald, F. V. A Monograph of the Culicidae of the World. 5 vols, and plates. iQoi-igio. London : Brit. Museum, Nat. Hist. James^ S. p., and LlSTON^ W. G. The Anopheline Mosquitoes of India. 2nd edit. igii. Calcutta : Thacker, Spink and Co. Howard, L. O., Dyar^ H. G., and Knab, F. The Mosquitoes of North and Central America and the West Indies. 2 vols. 1Q12. Washington : Carnegie Institution. Austen, E. E. African Blood-sucking Flies, iqoq. London : Brit. Museum, Nat. History. Austen^ E. E. A Handbook of Tsetse-flies, igii. London: Brit. Museum, Nat. History. Castellani, a., and Chalmers^ A. J. Manual of Tropical Medicine. 2nd edit. 1,747 pp. igi3. London : Bailliere, Tindall and Cox. KOLLE and Wassermann. Handbuch der pathogenen mikroorganismen. Jena : Gustav Fischer. MiNCHIN, E. A. An Introduction to the Study of the Protozoa. igi2. London : Arnold. Laveran, a., et Mesnil, F. Trypanosomes et Trypanosomiases. 2nd edit. igi2. Paris : Masson and Co. DOFLEIN^ F. Lchrbuch der Protozoenkunde. 3rd edit. igii. Jena: Gustav Fischer. NuTTALL^ G. H. F., Warburton, C, Cooper, W. F., and Robinson^ L. E. Ticks — a Monograph of the Ixodoidea. Pt. I (igo8). Pt. II. (igii). University Press, Cambridge, England. Brumpt, E. Precis de Parasitologie. 2nd edit. igi3. Paris : Masson and Co. Patton, W. S., and Cragg, F. W. A Text-book of Medical Entomology. igi3. Christian Literature Society of India : London, Madras, and Calcutta. JOURNALS. For current researches the following, among others, should be consulted : — Annals of Troi)ical Medicine and Parasitology^ Liverpool. Annales de VInstitut Pasteur, Paris, Archives de Parasitologie, Paris. Archives de Zoologie Exferimentale et Generate, Paris. ArchiiJ fiir Protistenhunde, Jena. Archiv fiir Schiffs- itnd Tro-pen-Hygiene, Leipzig. Bulletin of Entomological Research, London. Bulletin de VInstitut Pasteur^ Paris. Bulletin de la Societe de Pathologic Exotique, Paris. Bulletins of the Bureau of Animal Industry^ Washington. Centralhlatt fiir Bakteriologie und Parasitenkunde, Jena. Com ft. Rend. Acad. Sci., Paris. Comft. Rend. Soc. Biol., Paris. Indian Journal of Medical Research^ Calcutta. Journal of Ex-perimental Medicine, New York. Journal of Medical Research^ Boston. Memorias do Instituto Osivaldo Cruz, Rio de Janeiro. Parasitology, Cambridge. Proceedings of the Royal Society, London. Quarterly J ournal of Microscopical Science, London. Review of Applied Entomology, London. Tropical Diseases Bulletin (London : Tropical Diseases Bureau). Zeitschrift fiir hifehtionshranhheiten, Berlin. THE ANIMAL PARASITES OF MAN Man is one of those organisms in or on which a whole host of parasites find conditions suitable for their existence: Protozoa, Platyhelminihes, Nematoda, Acanthocephala, Hirudinea, and a large number of Arthropoda (Arachnida as well as Insects) all include members which are parasites of man. These animals either live on the external surface of the body or within the intestine and its appendages. Other organs and systems are not quite free from foreign organisms — we are acquainted with parasites in the skeletal system, in the circulatory system, in the brain, in the muscles, in the excretory and genital organs, and even in the organs of sense. It is possible, and perhaps might be advantageous, to arrange and describe the parasites of man according to the situations in which they are found (parasites of the skin, intestinal parasites, etc.). Their description in the various stages of develop- ment would, however, be disturbed when, as is generally the case, the different stages are passed in different organs, and a work which treats more fully of the natural history of the parasites than of the local disorders to which they give rise would suffer thereby. It is, therefore, preferable to describe the parasites of man in their systematic order, and to mention their different situations in man in describing each species. A. PROTOZOA, . BY H. B. FANTHAM, M.A., D.Sc. All those animal organisms which throughout their entire life never rise above the unicellular stage, or merely form simple, loose colonies of similar unicellular animals, are grouped under the term Protozoa (Goldfuss, 1820), as the simplest types of animal life. All the vital functions of these, the lowest forms of animals, are carried out by their body substance, the protoplasm (sarcode). Often particular parts possess special functions, but the limits of a cell are never over-stepped thereby. These special parts of the cell are called " cell-organs " ; recently they have been termed " organella?." The living protoplasm has the appearance of a finely granular, viscid substance which, as a -rule, when not surrounded by dense investing membranes or skeletons, exhibits a distinct kind of movement, which has been termed amoeboid. According to the species, processes of different forms and varying numbers called pseudopodia are protruded and withdrawn, and with their assistance these tiny organisms glide along — it might almost be said f^ow along— over the surface. In most Protozoa two layers of cytoplasm may be recognised, and distinguished by their appearance and structure, namely, the superficially situated, viscid, and quite hyaline ectosarc or ectoplasm, and the more fluid and always granular endosarc or endoplasm, which is entirely enveloped by the ectoplasm. The two layers have different functions ; the movements originate from the ectoplasm, which also undoubtedly fulfils the functions of breathing, introduction of food and excretion. The endoplasm, which in some forms (Radiolaria) is separated from the ectoplasm by a membrane, undertakes the digestion 26 THE ANIMAL PARASITES OF MAN of the food. To this distribution of functions between the various layers ot cyto- plasm is due the development of particular cellular organs, such as the appearance of cilia, flagella, suctorial tubules (in the Suctoria) and the myophan striations, which are contractile parts of the ectoplasm in Infusoria and Gregarines. In many cases (Flagellata, Ciliata), an area is differentiated for the ingestion of food (oral part, cytostome) to which there is often added a straight or curved opening (cytopharynx), through which the food reaches the endoplasm. The indigestible residue is either cast off through the oral part or excreted by a special anal part (cytopyge). In rare cases, structures sensitive to light, the so-called pigment or eye spots are developed, e.g.^ Eugle7ta. In the case of Infusoria the endoplasm circulates slowly, and agglomerations of fluids (food vacuoles) sometimes appear around each bolus of food ; in these vacuoles the food is digested under the action of certain materials (ferments). Even in the lowliest Protozoa fluids to be excreted are, as a rule, gathered into one, or, more rarely, several contractile vacuoles, which regularly discharge their contents. This action, however, is to a certain extent governed by the temperature of the surrounding medium. In some Infusoria a tube-like channel in the cytoplasm is joined to the contractile vacuole which usually occupies a certain position ; this forms a sort of excretory duct, and there are also supply- canals leading to these organellae. Very frequently various substances are deposited in the endoplasm, such as fatty granules, drops of oil, pigment granules, bubbles of gas or crystals. More solid skeletal substances are secreted in or on the ectoplasm. To the latter belong the cuticle of the Sporozoa and Infusoria, the chalky shells containing one or several chambers of the Foraminifera, the siliceous and very ornamental framework of the Radiolaria, and the chitinous coat of many Flagellata, Infusoria, etc. Some forms make use of foreign bodies found in their surroundings, such as grains of sand, to construct their protective coverings. The food often consists of small animal or vegetable organisms and of organic waste ; it is usually introduced in toto into the endoplasm. On the other hand, the Suctoria extract nourishment from their prey by means of their tentacles. Many parasitic species also ingest solid food, others feed by endosmosis. In all cases one nucleus at least is present. It is true that the existence of non- nucleated Protozoa, the so-called Mofiera, is still insisted upon, but some of these have already proved to be nucleated, and the presence of nuclei in the others will no- doubt be established. Very often the number of nuclei increases considerably, but these multinucleate stages are always preceded by uninucleate stages. In the Infusoria, in addition to the larger or principal nucleus (macronucleus) there is usually a smaller reproductive nucleus (micronucleus). This dualism of the nuclear apparatus is considered by some to be general, and usually to appear first at the onset of reproduction. The form and structure of the nucleus vary greatly in different species. There are elongate, kidney-shaped, or even branched nuclei as well as spherical or oval ones. In addition to vesicular nuclei with a distinct karyosome and incidentally also with a nuclear membrane, homogeneous and more solid formations are frequently encountered. The nuclei are always differentiated from the protoplasm by their reactions, particularly in regard to certain stains. In many Protozoa an extra-nuclear mass, sometimes compact, sometimes diffuse,, arises from or near the nucleus. This mass, whose staining reactions resemble those of the nucleus, is termed the chromidial apparatus. On the dualistic hypothesis, two varieties of chromidia occur, one originating from the vegetative nucleus (macro- nucleus), being chromidia in the restricted sense, the other derived from the reproductive or micronucleus being termed sporetia. Chromidia consist of altered (? katabolic) nuclear material. CLASSIFICATION OF THE PROTOZOA 27 The nucleus plays the same part in the life of the single celled organisms as it does in the cells of the Metazoa and Metaphyta. It appears to influence in a certain manner all, or at least most, of the processes of life, such as motility, regeneration, growth, and generally also digestion. Its principal influence, however, is exercised in the propagation of the cells, as this is always brought about by the nucleus. The PROPAGATION of the Protozoa is effected either by division or by means of direct budding. In division, which is preceded by direct or indirect (mitotic) division of the nucleus, the body separates into two, several, or even a great many segments. In this process the entire substance of the body is involved, or a small residual fragment may be left, which does not undergo further division and finally perishes. In the budding method of multiplication a large number of buds are formed, either on the surface or in the interior of the organism. Where divisions or buddings follow one another rapidly, without the segments separating immediately after their production, numerous forms develop, which are often unlike the parental forms, and these are termed swarm spores or spores. Divisions imperfectly accomplished lead to the formation of protozoal colonies. Sometimes encystment^ takes place previous to division. Frequently, also, sexual processes appear, such as the union of two similar (isogamous) or dissimilar (anisogamous) individuals. In the latter case sexual dimorphism occurs, with the formation of males (microgametes) and of females (macrogametes). The union may be permanent (copulation), the process being comparable with the fertilisation of the ovum by a spermatozoon. On the other hand, attachment may be transient (con- jugation) when, after the exchange of portions of the nucleus, the couple separate, to multiply independently of each other. Sometimes there is an ALTERNATION OF GENERATIONS, as there may be several methods of propagation combined in the same species, either direct multiplication, conjugation, or copulation being practised ; the different generations may thus, in certain cases, be unlike morphologically. Protozoa inhabit salt water as well as fresh water ; they are also found on land in very damp places, and invade animals as parasites. Classification of the Protozoa. Class I. — Sarcodina {Rkizopoda). Protozoa, the body substance of which forms pseudopodia ; many of them are capable of developing chitinous, chalky, or siliceous coverings or skeletal structures, which, however, permit the protrusion of the pseudopodia either over the entire periphery or at certain points. They possess one nucleus or several. Order i. — Amoebina (Lobosa) naked or with a simple shell, sometimes formed of a foreign substance ; the pseudopodia may be lobose or finger-shaped ; there may be a contractile vacuole ; generally only one nucleus. They live in fresh or salt water, in the soil, and also parasitically. Order 2. — Foraininifera (Reticulata). Mostly provided with a calcareous shell, usually consisting of several chambers, and allowing the protrusion of the pseudopodia either at the periphery or only at the opening. The pseudopodia are filamentous and frequently anastomosed ; there is no contractile vacuole ; there are usually several nuclei. Mostly marine. Order '^i. — Heliozoa. Naked, or with a chitinous or simple radial siliceous skeleton ; the pseudopodia are filamentous, and are frequently supported ' Independently of propagation, many protozoa protect themselves from death by encyst- ment when the water in which they are living dries up ; in this condition the wind may carry them over wide tracts of land. 28 THE ANIMAL PARASITES OF MAN by firmer axes, which exhibit no tendency to anastomosis ; there is a contractile vacuole ; one or several nuclei. Live in fresh water. Order ^.—Radiolaria. The body has radially-disposed filamentous pseudo- podia, and the nucleus is hidden in the central capsule ; there is almost always a siliceous framework, consisting of pieces arranged radially, tangentially, or lattice-like ; there is no contractile vacuole, but fluid- containing hydrostatic vacuoles are present in the peripheral protoplasm. Marine. Class II. — Mastigophora {Flagellata). Protozoa with one or several long flagella used for locomotion and for acquiring food ; in stationary forms their only function is to take in food. Cvtostome and contractile vacuole mav be present. Maybe either naked or provided with protective coverings; one or more nuclei. They live either in fresh or salt water, or may be parasitic. This class is again divided into several sub-classes and orders, of which only the Euflag>:llata, with the Protomonadina and Polymastigoda are of interest here. Class III. — Sporozoa. Protozoa that only live parasitically in the cells, tissues, or organs of other animals. They ingest liquid food by osmosis ; the surface of the body is covered with an ectoplasmic layer, or cuticle ; they have no cilia in the adult state, but may form pseudopodia. Flagella occur, but only on the male propagating individuals. There may be one or numerous nuclei, but no contractile vacuole. Propagation by means of spores, mostly provided with sporocysts, is characteristic. Sub-class I. — Telosporidia. These are usually of constant form, rarely amoeboid ; they are uninucleate in the mature state; they live within host cells in the first stage. Spore-formation occurs at the end of the life-cycle. Order i. — Gregari?iida. Body of a constant, usually elongate form, surrounded by a cuticle, In the early stage they lead an intracellular existence; in the mature stage they live within the intestine or body cavity of inverte- brate animals, especially the Arthropoda, and, like intestinal parasites, are provided with clinging organs. Copulation usually isogamous ; the spores have coats (chlamydospores) and usually contain several minute germs (sporozoites). Order 2. — Coccidiidea. Body of uniform spherical or oval shape : they lead an intracellular life, but are not freely motile in cavities of the body. Fertilization is anisogamous ; the spores have coals or shells (sporocysts), and usually contain several sporozoites. Exhibit alternation of generations. Order 3. — Hcemosporidia. Parasites of the blood corpuscles of vertebrate animals ; they exhibit amoeboid movement; fertilization is anisogamous; many present alternation of generations and hosts ; spores naked. Sub-class 2. — Neosporidia. They are multinucleate when adult, and the form of the body varies exceedingly (often amoeboid) ; spore-formation com- mences before the completion of growth. Order i. — Myxosporidia. The spores have valvular coats, with or without caudal appendages, with two, rarely four, polar capsules. They live free in such organs as the gall or urinary bladder, but are chiefly found in connective tissue. They occur especially in fishes. Order 2. — Microsporidia. Spores with coats or sporocysts ; no caudal appendage, with one polar capsule. They usually live in the tissues of Arthropoda. Order 3. — Sarcosporidia. Elongate parasites of the muscular fibres of amniotic vertebrates, on rare occasions they occur also in the connective tissue ; the spores, which are kidney or sickle-shaped, are naked and apparently have no obvious polar capsule. SARCODINA 29 Order ^. — Haplosporidia. Simple organisms, forming simple spores; they occur in Rotifers, Polychaetes, Fish and Man. Class IV. — Infusoria {Ciliata). The body is generally uniform in shape, with cilia and contractile vacuole, frequently also with cytostome ; usually has macro- and micro-nucleus ; live free in water and also parasitically. The "orders Holotricha., Heterotricha, Olicrotricha^ Hypotricha and Peritricha are classified according to the arrangement of the cilia. Class V. — Suctoria. Bodies with suctorial tubes, contractile vacuoles, macro- and micro-nucleus, no cytostome. They generally invade aquatic animals as cavity parasites, yet also attack plants ; early stage ciliated. Live sometimes as parasites on Infusoria. [The Suctoria are frequently regarded as a sub-class of the Infusoria.] The Protozoa and Protophyta are sometimes united under the term Protista (Haeckel, i866). The Spirochaetes are Protists (see pp. 1 14 — 128). Class I. SARCODINA, Butschli, 1882. Order. Amoebina, Ehrenberg. A. Human Intestinal Amoebse. The first record of the occurrence of amceba-like organisms in the human intes- tine, that is, in intestinal evacuations, was that of Lambl (1859) ; nevertheless, the case was not quite conclusive, as the occurrence of testaceous amoebae of fresh water {Arcella, Difftiigid) was also reported. In 1870 Lewis found amoebae associated with disorders of the large intestine in patients in Calcutta. A year later Cunningham reported from the same locality that he had observed on eighteen occasions, in one hundred examinations of dejecta from cholera patients, colourless bodies with amoeboid Fig. I. — Avuxba co/i, Lcisch, in the intestinal mucus. (After Losch.) movements, which became encysted and multiplied by fission. The daughter forms were said to be capable of dividing again, but they might also remain in contact. Contractile vacuoles were not noticed. The same bodies were observed also in simple diarrhoea (twenty-eight cases out of one hundred.) The case reported by Losch in 1875 attracted more attention. It was that of a peasant, aged 24, who came from the province of Archangel. He was admitted 30 THE ANIMAL PARASITES OF MAN into Eichwald's clinic at Petrograd with symptoms of dysentery. In the dis- charges containing blood and pus, Losch found amcebae in large numbers. When at rest these amoebae measured from 20 /i to 35 ft ; in a state of movement their length might extend up to 60 /i (fig. i). The pseudopodia appeared only singly, and, since they were hyaline (ectoplasmic), were thus distinguished from the markedly granular endoplasm that enclosed a spherical nucleus of from 5 /u to 7 ;u in diameter. One or more non-contractile vacuoles were present. Quinine enemata had the effect of making the amoebae disappear from the fasces and thus causing the diarrhoea to abate. Four months after admission the patient died from the results of intercurrent pneumonia. At the autopsy ulceration of the large intestine was found, especially in the lower parts. Losch connected the amoebae with the ulcerations by experiments made on four dogs by injecting them with recently passed stools {per OS et anum). Eight days after the last injection numerous amcebae were found in the faeces of one of these dogs ; eighteen days after the injection the animal was killed. The mucosa of the rectum was inflamed, covered with blood-stained mucus and ulcerated in three places. Numbers of amoebae were found both in the pus of the ulcers and in the mucus. The three other dogs remained healthy. From these observations Losch concluded that the species of amoeba described by him as Amccba colt could not be regarded as the primary cause of the disease, but that it was certainly capable of increasing a lesion of the large intestine already present, or at least of preventing its healing. B. Grassi (1879) found in the stools of healthy as well as in those of diarrhoeic patients from various localities in Northern Italy, amoebae similar to those discovered by Losch. As this was of frequent occurrence, the pathogenicity could not be definitely established. Normand, formerly naval surgeon at Hong-Kong, observed numerous amoebae in the dejecta of two patients suffering from colitis. Many further investigations, which cannot be quoted in detail, showed not only that intestinal amoebce were widely distributed in man, but indicated with greater certainty their role as agents of dysentery. The Commission sent out by the German Government in the year 1883 to investigate cholera in India and Egypt — whose members discovered the cholera bacillus — also collected information with regard to dysentery. In five cases of dysentery examined posi mortem at Alexandria, with the exception of one case in which ulceration of the colon had already cicatrized or was approaching cicatrization, R. Koch found amoebae as well as bacteria in sections from the base of the ulcers, although such had previously escaped notice in examina- tion of the dejecta. Encouraged by these results, Kartulis (1885), who had discovered amceba-like bodies in the stools of patients suffering from intestinal complaints at Alexandria, continued his investigations. The results, obtained from more than 500 cases, gave rise to the theory that typical dysentery vviis caused by amoebae as were also the liver-abscesses that often accompany it. Kartulis supported his theory not only by the regular occurrence of amoebae in the stools of dysenteric patients and their absence in other diseases, and by the occurrence of the parasites in ulcers of the large intestine and in the pus from liver-abscesses, but also by experiments which he performed on cats. These were infected by injection /^r a«//;// of stool material rich in amoebae from subjects of dysentery. The infection took place also when amoeba-containing, but bacteria-free, pus from liver-abscesses was used. It has been objected that the infection of man with Amceba coli^ as the dysenteric amoebae were then generally designated, does not take place j2^.?r attu7n hnx. per os. This difficulty, however, diminished in proportion as the encysted states of amoebae (fig. 2), long known in the case of other Protozoa, became understood. The infection of man (Calandruccio, 1890) and of cats (Quincke and Koos) succeeded solely when material containing such stages was used. Amoebae introduced into the intestine multiply thereby fission (Harris, 1894). However, this theory, to which various other authors SARCODINA 21 gave support on the grounds of their own observations, encountered opposition. Thus it was estabbshed that amoebas were not found in patients in every place where dysentery was endemic, or else they were much rarer than was expected. Further amoebae were present in the most varied kinds of intestinal diseases, both of infective and non -infective characters. Also they were present in quite healthy persons. Moreover, for various reasons, infection experiments on animals failed to supply proof, and finally a bacterium was discovered (Shiga, 1898) to be the excitant of one form of dysentery. Agglutination attested the specific part played by this organism, as it was produced by the blood serum of a person suffering from or recovered from dysentery, but not by the serum of one who was uninfected. Bacillary dysentery consequently was a distinct entity. The final step to be taken was to decide whether there was a specific amoebic enteritis (amoebic dysentery or amoebiasis, according to Musgrave). Fig. 2. — Encysted intestinal amoebae showing nuclear muliiplication. (Afier B. Grassi.) This question should decidedly be regarded from the positive point of view. It is intimately connected with another, namely, whether there are not several species of intestinal amoebae. The possibility of this had already been recognized. In addition to the Amceba coli Losch, R. Blanchard distinguished yet another, Amoeba intesiinalis^ and designated thereby the large amoebae described in the first com- munication made by Kartulis ; later on he stated the distinction between the species. Councilman and Lafleur ^ (1891) considered the amceba of dysentery to h& Amceba coli Losch and so re-named the species Amoeba dysentericd. Kruse and Pasquale (1893) employed the same nomenclature, but retained the old name Amoeba coli Losch for the non-infectious species. Quincke and Roos (1893) set forth three species: a smaller species (25 /x) finely granular, pathogenic for men and cats {Ajnosba coli Losch) ; a larger species (40 /*) coarsely granular, pathogenic for men but not for cats {A. coli mitis) ; and a similar species non-pathogenic either for man or cat {A. intestini vulgaris), Celli and Fiocca (1894-6) went still further, they distinguished : (i) Amoeba lobosa vdiueiy gutlula {=A.guttula Duj), variety oblonga (=A. oblonga Schm.) and variety coli (= A. coli Losch). (2) Amoeba spinosa n. sp. occurring in the vagina as well as in the intestine of human patients suffering from diarrhoea and dysentery. (3) Amoeba diaphana n. sp. found in the human intestine in cases of dysentery. (4) Amoeba vermicularis Weisse, present in the vagina and in dysentery ; and (5) Amoeba reticularis n. sp. in dysentery. Shiga distinguished two species; a larger pathogenic species with a somewhat active movement, and a small harmless species with a somewhat sluggish movement. Bowman mentions two varieties, Strong and Musgrave (1900) two species— the patho- genic Amceba dysenteries and the non-pathogenic Amoeba coli ; Jager (1902) and Jiirgens (1902) mention at least two species. In the following year (1903) a work by Schaudinn was published which marked a real advance. This, in conjunction with the establishing of a special genus {^Endamoeba or Entainceba) for human intestinal amoebae first by Leidy^ and then by Casagrandi and Barbagallo,^ for the time cleared up the confused nomenclature, the old name Amoeba coli being retained for the ' '• Amcebic Dysentery," Johns Hopkins Hosp. Repts., ii, pp. 395-548, 7 plates. - "On Amceba blattae,'' Proc. Acad. Nat. Set., Philadelphia (1879), xxxi, p. 204. 3 '■'' Entamceba hominis s. A?nceba coli (Losch). Annali d'lgiene speriment. (1897), vii, p. 103. See also further remarks on p. 34. 32 THE ANIMAL PARASITES OF MAN harmless intestinal amoebae of man, whereas the pathogenic species was desij^inated Entama'ba histolytica. The history of more recent work is incorporated in the accounts of the entamoebas given below. Entamceba coll, Losch, 1875, emend. Schaudinn, 1903. Syn. : Amceba coli, Losch, 1875. EntauKsba /wniinis^ Casagr. et Barbag. 1897. The amoeboid trophozoite, according to Losch, measures 26 yu, to 30 fjL and upwards; according to Grassi 8 /i to 22 /x ; according to Schuberg 12 /x to 26 /x. A separation of the body substance into ectoplasm and endoplasm is only perceived during movement. The pseudopodia, which are generally only protruded singly, are broad Fig. 3. — Entamceha coli : life-cycle, a — e, stages in binary fission ; A — Z>, schizogony, wiih formation of eight merozoites ; 2 — 10, cyst formation cr sporogony, with formation of eight nucleate cysts. (After Castellani and Chalmers). and rounded at the end (lobopodia) and are hyaline, while the remainder of the body is granular. The ectoplasm is less refractile than the rest of the cytoplasm; it also stains less intensely (fig. i), and is best seen on protrusion of a pseudopodium. Red blood corpuscles are larely, if ever, found ingested in the cytoplasm. The nucleus is vesicular, and is spherical when inactive, measuring 5 /A to yz-t, with a thick nuclear membrane. In the centre of the ENTAMCEBA CO LI 33 nucleus is a chromatinic body or karyosome or sometimes several small nuclear bodies formed of plastin and chromatin ; the remaining chromatin is arranged on the achromatic network in the form of fine granules, especially thickly deposited on the nuclear membrane. Entamoeba coll lives as a commensal in the upper portion of the large intestine, where the faeces still possess a pulpy . consistency. With their concentration and change in reaction lower in the bowel, the parasites either die or else if they are at a suitable stage of development form resistant cysts. These cysts (fig. 2) can be found in great abundance in normal faeces, as Grassi first observed. Slight laxantia or intestinal diseases of any kind producing increased peristalsis, however, show amoebae even in the unencysted condition, provided that the person harbours intestinal amoebae generally. The intensity of infection varies according to the locality; thus Schaudinn found that 50 per cent, of the persons examined w^re infected with harmless amoebae in East Prussia, 20 per cent, in Berlin and about 66 per cent, on the Austrian littoral. The life-history (fig. 3) of the parasite exhibits two phases : (a) asexual multiplication in the intestine, either by binary fission or by schizogony with formation of eight merozoites, and (b) sporogony leading to the production of eight-nucleate cysts. Infection results from ingestion of cysts. Only cysts with eight nuclei are infective. The diameter of such cysts is about 15 /^ to 20 fM. There are varying accounts of the details of the life-cycle of Entamosba colt in its different stages. Thus, regarding schizogony or multiple fission it was formerly stated that the nucleus of the parent amoeba divided into eight portions, which after dissolution of the nuclear membrane, passed outwards into the cytoplasm, which segregated around each. Eight merozoites were thus produced. More recently the process of schizogony has been considered to consist in the repeated division of the nucleus into two, four, and finally eight nuclei (fig. 3, A — d), and the formation of eight merozoites or amoebulas. The process of encystment is initiated by the extrusion of all liquid and foreign bodies from the protoplasm, which assumes a spherical form (fig. 4, a). The rounded uninucleate amoeba then secretes a soft gelatinous coat, which finally diffisrentiates into a double contoured cyst wall in older cysts. According to Casagrandi and Barbagallo, the size of the cyst varies from 8 ju to 30 /*, and averages about 15 /*. According to Schaudinn (1903) the cytological changes during cyst formation are as follows. The nucleus of a rounded uninucleate form divides into two (fig. 4, B). Each of these nuclei fragments into chromidia (fig. 4, C), some of which are absorbed, while others reunite so that the cell becomes binucleate again. Each of these nuclei, by a twice repeated division, produces three nuclei (fig. 4, D), the smaller two of which degenerate and were regarded as reduction nuclei. There is a clear zone or vacuole in the middle of the cyst during these maturation processes, dividing the cyst into two halves. After the nuclear reduction the clear space disappears, and each nucleus (termed by some a gamete nucleus) divides into two pronuclei (fig. 4, e). The pronuclei of the pairs were said by Schaudinn to differ slightly. Copulation occurs between pairs of unlike pronuclei, and is an example of autogamy (fig. 4, F). When complete, each of the fusion nuclei (synkarya) divides twice, giving 3 34 THE ANIMAL PARASITES OF MAN rise first to four and finally to eight nuclei. Eight amcebulae are thus formed within the cyst. According to Hartmann and Whitmore (191 OS however, autogamy does not occur within the cysts of E. colt. They consider that eight small amoebulas are formed (fig. 3, 2—10) which escape from the cyst and then conjugate in pairs (fig. 3, 10 — 12), afterwards growing into a new generation of trophozoites. Only some 10 to 20 per cent, of the cysts evacuated with the fasces undergo the full course of development, the majority perish previously. In old dry faeces, only cysts with eight nuclei are found, and it is these alone that cause the infection. Entamceba ivilliamsi, E. biitschlii, E. hartmanni and E. poleki (Prowazek) are probably only varieties of E. colt. Fig. 4. — So-called autogamy of ^«/a?/za'/^a ^(7//. A, rounded amoeba; B, nucleus dividing ; c, the two daughter-nuclei giving off chromidia ; d, each nucleus has formed two reduction nuclei ; e, cyst membrane formed, and gamete nuclei are dividing ; f, cyst with two synkarya. The principal feature distinguishing Entamoeba coll from E, histolytica is the formation of eight-nucleate cysts by the former aS' contrasted with the tetra-nucleate cysts of the latter. The cyst-wall of E. coll is thicker than that of E. histolytica (tettagena). Further, E. coli does not usually ingest red blood corpuscles, nor are " chromidial blocks" present inside its cyst (see p. 40). According to Chatton and Lalung-Bonnaire'^ (191 2) the entamcebae of vertebrates should be placed in a separate genus Loschia, as they differ in their life-history from E. blatta% the type species of Entamoeba. Leidy (1879), however, named the genus Eudanioeba, but further researches are necessary on biological variation among these organisms. Entamoeba histolytica, Schaudinn, 1903. Syn. : Amoeba coli, autt. p. p. Ainosba dysenfericp, autt. p. p. The average size of the amoeboid trophozoite is 25 fji, to 30 fi. In fccces diluted with salt solution the amoebae swell to 40 ^ and more. There is sometimes separation of the body substance into a strongly refractile vitreous ectoplasm and a corneous endoplasm, pronounced ' Archiv f. Protistenktinde^ xxiv, p. 182. - Bull. Soc. Path. Exotique^ v, p. 135. ENTAMCEBA HISTOLYTICA 35 even in repose, although the former is not equally thick at all parts of the periphery. In the endoplasm generally there are numerous foreign bodies (bacteria, epithelial cells, colourless and red blood corpuscles (fig. 6), and occasionally living flagellates of the intestine). The nucleus is 4 /z to 6 /a in diameter, and may be difficult to recognize because it is sometimes weakly refractile and poor in chromatin. Its shape is slightly variable ; it is usually excentric, sometimes wholly peripheral at the limit of the two parts of the body. Vacuoles are not present in quite fresh specimens, but appear later. In the study of E. Jiisfolytica, the morphological characters of the trophozoite or vegetative stage of the organism formerly separated as E. tetragena (figs. 5, 6, Sa) must be considered (see p. 38). Fig. ^.—Entamceba histolyiica [^tetragena form), showing three successive changes of form due to movement. X IICX). (After Hartmann.) The history of the development ot these species, which give rise to amoebic enteritis as distinguished from bacillary dysentery, was formerly not so well known .as that of E. coli. Upon being introduced into cats {per anum) dysenteric amoebae provoke symptoms similar to those in man. In the latter, besides metastatic liver abscesses, abscesses of the lungs, and, according lo Kartulis, cerebral abscesses are occasionally produced. Marchoux (1899) states that when the disease has lasted for some time liver abscesses are produced in cats also. In the large intestine of infected cats the amoebae creep over the epithelium, and here and there they force the epithelial cells apart, as well as removing them or pushing them in front of them ; the amoebae thus insert themselves into the narrowest fissures. They penetrate also into the glands through the epithelium, and thence into the connective tissue of the mucosa. Intestinal and glandular epithelia perish under the influence of these parasites : the cells are pushed aside, fall to pieces or are absorbed by the amoebae, mucosa the amoebae migrate further, and often accumulate above the muscles. Finally they rupture this and force their way into the submucosa. In cats. Fig. 6. — Entamaba histolytica which has ingested many red blood corpuscles. x iioo. (After Hart- mann.) In the connective tissue of the 36 THE ANIMAL PARASITES OF MAN apparently, the penetration is not so great as in men, according to Kruse and Pasquale. During their migration the parasites also gain access to the lymph- follicles of the wall of the intestine, which become swollen and commence to suppurate ; follicular abscesses arise and after their rupture follicular ulcers. The diseased patches in the mucosa are markedly hyperaemic and numerous hiemor- rhages are set up. Roos and Harris state that the amoebae also oenetrate into the blood-vessels (fig. 7) and this explains the occurrence of metastatic abscesses.' The whole submucosa is severely swollen at the diseased spot and undergoes small- celled infiltration in the neighbourhood of the colonies of amoebae. From these findings Jiirgens (1902) draws the conclusion^ which is followed here, that the amoebae Fig. 7.— Section through wall of large intestine (of a man) close under an ulcer caused by Entamoeba histolytica. A, amoebse that have penetrated partly in blood-vessels (Bv), partly ill tissue of submucosa to the muscularis. Magnified. (After Harris.) are causative agents of the enteritis of cats, which disease is well defined, both pathologically and anatomically. Subsequent researches confirm the experience of earlier authors ; great precautions were taken to exclude errors, hence, as with Gross and Harris, no exception can be taken to their results. The inoculation material was derived from soldiers who suffered from amoebic enteritis in China and who were admitted into the garrison hospital at Berlin. In order to be ' Lung abscesses generally arise by the bursting of a liver abscess through the diaphragm into the right lower lobe of the lung, sometimes also through conveyance of amoebcE by means of the blood-stream (Banting). ^ These findings were confirmed by Schaudinn by means of investigations on cats and men. Cf. also Alfred Gross, Marchoux, P. G. Woolley, W. E. Musgrave, H. F. Harris and others. ENTAMOEBA HISTOLYTICA yj independent of the patients themselves, transmission experiments from cat to cat were performed, after the first experiments on cats yielded positive results. This was also effected by rectal feeding as employed by earlier workers. Such appeared necessary in order to prevent the evacuation of the inoculation material /^r «««w, as well as to avoid the employment of morphia and ether narcosis. Forty-six cats were used for the experiments. Ten cats received tested stools containing motile amoebie from soldiers suffering from amoebic enteritis contracted in China. Sixteen other cats received stools from cats infected by inoculation. All the animals sickened and suffered from the disease. Five cats received dejecta from human amoebic enteritis in which, however, no motile amcebae were present. Thirteen cats received stools from soldiers who suffered from bacillary dysentery. None of the latter cats took the complaint and none showed changes in the large intestine upon sectioning. The injection of various bacteria, obtained from a stool of amoebic enteritis pathogenic to cats, remained without result in both the cats employed for this experiment. Lastly, two cats, which had been kept with those artificially infected, were taken ill spontaneously and suffered from the disease. In the opinion of Harris, who ascertained the harmless nature of bacteria derived from the intestinal flora containing dysenteric amoebae, young dogs are capable of being infected. Within the large intestine an active increase of Entamoeba histolytica must occur. Nevertheless, Jiirgens did not definitely find changes that might be interpreted in this sense. Schaudinn (1903) observed division and gemmation in vivo. Both processes, in which the nucleus divides by amitosis, can only be distinguished by the fact that the daughter individuals are similar in binary fission but dissimilar in gemmation, whether they make their appearance singly or in greater numVjers. Schizogony, resulting in the formation of eight individuals, which is so characteristic for Ent- amoeba coli, was not observed. (But schizogony, into four merozoites, is now known to occur. Gemmation processes are apparently degenerative.) Resistant stages, which serve for transmission to other hosts, are according to Schaudinn' first formed when the diseased portions commence to heal, or more accurately, the recovery commences when the vegetative increase of the amoebae in the intestine discontinues. The so-called spores of E. histolytica were distinguished very definitely from those of E. coli; they were said to consist of spheres of only 3 to 7 /i in diameter, which were surrounded by a double membrane, at first colourless, but becoming a light brownish yellow colour after a few hours, and possessing a protoplas- mic content containing chromidia. They were said to arise by fragments of chromatin passing outwards from the nucleus of the amoeba into the surrounding cytoplasm (fig. 9, a) and undergoing so marked an increase that finally the whole cytoplasm became filled with chromidia. The remainder of the nucleus underwent degenera- tion and became extruded. On the surface of the cytoplasm there then arose small protuberances containing chromidia. These processes had been observed in the living organisms. They gradually divided and separated from membranes which later became yellow. The remainder of the amoeba perished. Craig^ had also seen phases of this process of development. It must be remarked that, according to recent researches, these processes of exogenous sporulation are degenerative in character (see p. 41). The small spores may be fungi. The " sporulation " processes are only men- tioned here as a warning. They are now only of historic interest. By means of an experiment made on a cat, Schaudinn ascertained that ingestion of permanent cysts, which resist desiccation, is the cause of the infection. The animal took food containing dry faeces with amoeba cysts ; these fasces came from a patient suffering from amoebic ' Arb. a. d. kaiserl. Gesundheitsamtey xix, pp. 547-576. 2 " Life cycle of Amoeba coli in Human Body," American Medicine, 1904, vii, p. 299 ; viii, p. 185. 58 THE ANIMAL PARASITES OF MAN enteritis in China. On the evening of the third day the cat evacuated blood-stained mucous faeces which contained large numbers of typical Entamceba histolytica. On the fourth day after the infection the animal experimented upon died, and the large intestine showed the changes previously stated. E histolytica also is found in the large intestine. This was originally shown to be the case by Kartulis, and the fact has recently been confirmed from many quarters. It is also present in the metastatic abscesses of which it is the cause {cf. among other authors, Rogers, Brit. Med. Journ., 1902, ii, No. 2,177, P- 844 ; and 1903, i, No. 2,214, P- 1315)- It should lastly be pointed out in this connection that mixed infections also take place. For instance, in addition to E. histolytica^ E. coli, and, under certain circum- stances, flagellates may be found together. In the same way E. coli may come under observation even in bacillary dysentery. On the other hand, Schaudinn stated that in cases of dysentery endemic in Istria, Entamoeba coli, if it had hitherto been present, disappeared, to return again after recovery from the illness. ^■ «s Fig. 8. — Entama-ba histoiy.i.u. a, trophozoite {tetragena type) containing red blood corpuscles, x 1,300; h and c, two isolated nuclei showing different appearances of karyosome, centriole and nuclear membrane, x 2,600. (After Hartmann.) (Entamceha tetragenUy Viereck, 1907.) This amoeba must now be considered to be a part of the hfe- cycle of Entamoeba histolytica, in fact a very important part of that cycle, especially in its tetranucleate cystic stages. This organism, the so-called Entaniccba tetragena, may occur in the human intestine in cases of amoebic dysentery, especially in mild or chronic cases. It was discovered by Viereck in 1907 in patients suffering from dysentery contracted in Africa. Soon after- wards an independent description was published by Hartmann, who called the ainoeba E. africana. It was also studied by Bensen and Werner. Recently (1912-13) much work has been published on this amoeba by Darling and others ; in this way its relationship to Schaudinn's E. histolytica has been made known. In general morphology it somewhat resembles Entamceha coli, and ENTAMOEBA HISTOLYTICA (TETRAGENA) 39 its discoverer at first mistook it for a variety of that species. Accord- ing to Hartmann, a distinct ectoplasm is only clearly visible when a pseudopodium is protruded (fig. 5). The granular endoplasm may contain ingested red blood corpuscles (fig. 6). The large, round nucleus is visible in the fresh state (fig. 8, a). So-called chromidial masses (? crystalloidal substances) may occur in the cytoplasm. Some investigators, as Hartmann,^ lay stress on the internal structure of the nucleus (fig. 8, b, c), best seen in preparations fixed wet and stained with iron-haematoxylin. The nucleus is limited by a well- marked nuclear membrane, on the inside of which granules or nodules of chromatin may occur. There is a karyosome, which, in success- FiG. 9. — Entamceba histolytica {tetragena form). «, emission of chromatin from nucleus ; b, nuclear division ; t to 4"5yLfc. A con- tractile vacuole is not visible. The pseudopodium is broad. It was discovered in the mouths of persons with dental caries at Rovigno and also at Trieste, being most easily found in dense masses of leucocytes, also among leptothrix and spirochaste clusters. It can be easily dis- tinguished from leucocytes by more intense staining with neutral red. Multiplication proceeds by fission. Transmission may take place through the small spherical cysts. This species (fig. 10) has since been observed in Berlin, and is also occasionally found in carcinoma of various regions of the oral cavity. (Leyden and Lowenthal, 1905). ■■^^ Fig. 10. — Entamoeba buccalis^ Prow, a-d, the same specimen observed during five minutes. X 1,000. e, amoeba fixed and stained with iron-hsematoxylin. x 1,500. (After Leyden and Lowenthal.) Entamoeba buccalis, Prow., is said to be allied to a protozoon which A. Tietze has found either encysted or free in the lumen of the orifice of the parotid gland of an infant aged 4 months. The gland had undergone pathological change, and had therefore been extirpated. The organisms, which were roundish and three to four times the size of the normal epithelial cells of the gland, were without a membrane and possessed a nucleus in which the chromatic substance appeared to be contained in a karyosome. Bass and John's^ (Feb. 191 5) and Smith, Middleton and Barrett (1914) state that E. buccalis is the cause of pyorrhoea alveolaris. Entamoeba undulans^ Aide Castellani, 1905. Under this name a protozoon is described which A. Castellani found in addition to Entamceba histolytica and Trichojtionas intestinalis in the fasces of an European planter living in Ceylon, who had suffered from amoebic enteritis and liver abscess. The shape of the body was roundish or oval, 25 /t to 30 /t in the greatest diameter. It was without a flagellum, but with an undulating membrane, and capable of pro- truding a long pseudopodium from different parts of its body at short intervals. The nucleus could not always be recognized in life ; it was, however, always demonstrable ' Joiirn. Amer. Med. Assoc, Ixiv, p. 553. 44 THE ANIMAL PARASITES OF MAN by staining. One or two contractile vacuoles were present. The protoplasm was finely granular, showing no differentiation into ecto- and endo-plasm. According to Braun, in spite of the author declaring himself expressly against the flagellate nature of the parasite, such a nature may be assumed to be tolerably certain in view of the description and illustration. It is now considered that Entamcsba undulans is a portion of a flagellate, namely, T?'ichomonas. Entamoeba kartulisi, Doflein, 1901. Doflein gave this name to amoebae, from 30 /^ to 38 /x in diameter, which KartuHs(i893) found on examining the pus of an abscess in the right lower jaw of an Arab, aged 43, and in a portion of bone that had been extracted. The movements of the amoebae (fig. 11) were more active than those of ^' dysenteric amoebae." Their coarsely granular cytoplasm contained blood and pus corpuscles, and a nucleus was generally ; only recognizable after staining. Vacuoles I ^/' were not seen with certainty. Flexner re- ^ ported upon a similar case, and Kartulis published five additional cases. As in these &rDofl.'; ftotThTpt'oTan cases dental caries was present the infection abscess in the lower jaw, show- is likely to have proceeded from the oral mfnt^'^{AftIr'S-?uliso'"°''^" cavity as a result of the carious teeth. Craig^ (191 1) considers that this parasite is probably identical with Entamoeha histolytica. In the literature the following species have been reported as occurring in the oral cavity of man : — Amoeba gingivalis, Gros, 1849. [? identical with Eniamoeha bticcalis.~[ Amceba buccalis, Sternberg, 1862. Amoeba dentalis, Grassi, 1879. Far too little, however, is known concerning these to regard them as definite species, that is, independent organisms ; Grassi thinks it even possible there may have been a confusion in their case with salivary corpuscles. If they really are amcebse they are all of them probably identical with E7itamceba buccalis. Genus Paramoeba, Schaudinn, 1896. Schaudinn established the genus Paramcsba for a marine rhizopod which multi- plied by division, became encysted at the end of its vegetative life and then segmented into swarm bodies with two flagella. These multiplied by longitudinal fission, and finally passed into the condition of Amoebae. Whether the human parasite described by C. F. Craig (1906) as ' "The Parasitic Amoebse of Man," Lippincott, Philadelphia. AMCEBA UROGENITALIS 45 Paramoeba hominis. belonged to this genus was for a time uncertain. It is now placed in a new genus Craigia, Calkins, 1912, since it possesses only one flagellum.' In the amoebic stage it is 15 ^ to 25 ju in diameter ; ecto- and endo-plasm during rest are indistinguishable. The body substance is granular, with a spherical, sharply contoured nucleus and an accessory nuclear body. No vacuoles are present, but occasionally the endoplasm contains red blood corpuscles. The pseudopodia are hyaline, finger- or lobe-shaped, and are protruded either singly or in twos. Multiplication is by binary fission and by the formation of spherical cysts (15/ito 10 jx in diameter) in which occurs successive division of the nuclei, ultimately forming ten to twelve roundish bodies each of which soon develops a flagellum. The flagellate stages have similarly a spherical shape and attain a diameter of 10 /* to 15 fi. They also occasionally contain red blood corpuscles and pass either directly or after longitudinal division into the amoeboid phase. Craig found these Amoebae and the flagellate stage belonging to them in six patients in the military hospital at Manila (Philippine Islands), five of whom were suffering from simple diarrhoea whilst the sixth exhibited an amoebic enteritis and contained also Parai7taba hominis, with Entainoeba histolytica^ Schaudinn. In one of the other cases. Trichomonas intestinalis was present. B. Amoebae from other Organs. EntamcEba pulmonalis, Artault, 1898. Arlault^ discovered a few amoeboid forms with nucleus and vacuole in the contents of a lung cavity. In the fresh condition they were distinguishable from leucocytes by their remarkable capacity of light refraction. They were also much slower than the latter in staining with methylene blue or fuchsine. Their movements became more lively in a strong light. Water and other reagents killed them, and then, even when stained, they could not be distinguished from leucocytes. They have also been seen by Brumpt. R. Blanchard found amoebae which may belong here in the lungs of sheep. A. pulmonalis is perhaps the same as Entamoeba buccalis. Smith and Weidman^ (1910, 1914) described an entamoeba, E. mortinatalmm, from the lungs and other organs of infants in America. Amoeba urogenitalis, Baelz, 1883. This species was found in masses in the sanguineous urine as well as in the vagina of a patient in Japan, aged 23. Shortly before the death of the patient, which was caused by pulmonary tuberculosis, haematuria with severe tenesmus of the bladder had set in. The amoeba, which showed great motility, and had a diameter of about 50 fi when quiescent, exhibited a granular cytoplasm and a vesicular nucleus. Baelz is of opinion that these parasites were introduced into the vulva with the water used for washing the parts, and thence had ' See Craig (1913), Amer. Journ. Trap. Dis. and Prevent. Med.y i, p. 351. ' Arch, de Parasitologies i, p. 275. ^ A?fier. fourn. Trop. Dis. and Prevent. Med.y ii, p. 256. 46 THE ANIMAL PARASITES OF MAN penetrated into the bladder and vagina. Doflein places the organism in the genus Entamoeba, and it is perhaps identical with E. histolytica. Similar cases are also reported (1892-3) by other authors : Jiirgens, Kartulis, Posner^ and Wijnhoff. Jiirgens found small mucous cysts, filled with amoeboid bodies, in the bladder of an old woman suffering from chronic cystitis ; they were also found in the vagina. The amoeba observed by Kartulis in the sanguineous urine of a woman, aged 58, suffering from a tumour of the bladder, measured 12 /* to 20 |t, and exhibited slow movements by protruding short pseudopodia. The vacuoles and nucleus became visible only after staining with methylene blue. Posner's case related to a man, aged 37, who had hitherto been quite healthy and had never been out of Berlin. Suddenly, after a rigor, he passed urine tinged with blood. This contained, besides red and white blood corpuscles and hyaline and granular casts, large granular bodies (about 50 /* in length and 28 /i in breath), which slowly altered their shape, and contained red blood corpuscles in addition to other foreign matter. These bodies exhibited one or several nuclei and some vacuoles. From the course of the disease, which extended over a year, and during which similar attacks recurred, Posner came to the conclusion that the amoebae which had originally invaded the bladder had penetrated into the pelvis of the kidney, where they probably had settled in a cyst, and thence induced the repeated attacks. Wijnhoff observed four cases of amoeburia in Utrecht. Amoeba miurai, Ijima, 1898. Under this term the author describes protoplasmic bodies which Miura, in Tokyo, found in the serous fluid of a woman, aged 26, who had died from pleuritis and peritonitis endotheliomatosa. Two days before death these same forms had t^-^ — also appeared in the haemorrhagic faeces of the / */'""^\ patient. The bodies were usually spherical or tr | |fa fciri Bi ^ .i'-V \ ellipsoidal, and at one pole carried a small pro- \m|v', y:.<, J/ tuberance (fig. 12) beset with filamentous short ^^jjl^Lm^ "pseudopodia" (really a pseudopodium covered ^^^^ with cilia). Their size varied between 15 fx and 38 /t. The cytoplasm was finely granular, and TT.^ , y, L . . T- 1^0 difference was observable in the ecto- FlG. 12. — Amceba imtirai, Ij. , , , , 1 • X 500. a, fresh ; b, after treat- and endo-plasm, only the villous appendage ment with dilute acetic acid. was clearer. The cytoplasm contained vacuoles (After Ijima.) more or less numerous, none of which was contractile. After the addition of acetic acid one to three nuclei could be distinguished, 8 ^u to 15 /t in size. Actual movements were not observed. Taking everything into consideration, the independent nature of these bodies is, to say the least, doubtful, although it cannot be denied that they possess a certain similarity to the marine Amceba fluida, Grilber or Greeff, and to a few other species. (It is likely that cells present in serous exudation were mistaken for amoebae.) Appendix. ^* Rhizopods in Poliomyelitis acuta." In three cases of poliomyelitis acuta which were investigated by Ellermann, the spinal fluid obtained by puncture of the cord contained bodies, from 10 /a to 15 m in size, which had amoeboid movements and exhibited variously shaped pseudopodia in large numbers. After staining, a usually excentric nucleus, about i"5 /a in size, was demonstrated in them. CHLAMYDOPHRYS ENCHELYS 47 Order. Foraminifera, d'Orbigny. The order is divided by Max Schultze into Monothalamia and Polythalamia. Only a few of the former can be considered here. Sub-Order. Monothalamia. (Testaceous Amoebae). These forms occur frequently in fresh water, rarely in sea water. They possess a shell which is either pseudo-chitinous in character, or consists of foreign particles, or in a few cases is composed of siliceous lamellae. There is usually an orifice for the protrusion of pseudopodia. The only representative of the order of interest here is : — Genus. Chlamydophrys, Cienkowski, 1876. The genus is based on a form which A. Schneider carefully investigated and considered to be the Dijfflugia enchelys of Ehrenberg. L. Cienkowski redis- covered this same form and created for it the genus Chlamydophrys. We agree with this view, but not with the renaming of the organism (so common at the time). If the parasite in dung, Chlamydophrys stercorea Cienk: is identical with Difflugia enchelys of Ehrenberg, the old specific name should be retained. The genus is characterized by the possession of a hyaline, structure- less, slightly flexible shell which is ovoid or reniform. At the more pointed pole there is an orifice situated terminally or somewhat laterally^ serving for the emergence of the filiform pseudopodia (fig. 13, a). The protoplasm does not entirely fill the interior of the shell. An equatorial zone bearing excretory granules divides the shell internally into two almost equal portions. The anterior portion is rich in vacuoles and serves for the reception of nutriment and for digestion. The posterior part is vitreous, and contains the nucleus. One to three contractile vacuoles are situated in the equatorial zone. Chlamydophrys enchelys, Ehrbg. Syn. : Chlamydophrys stercorea^ L. Cienkowski. This species (fig. 13) is found in the faeces of various animals (cattle, rabbits, mice, and lizards), and also in quite fresh human faeces. According to Schaudinn, the parasite occurs so frequently in the human faeces that it must be considered of wide distribution. The species must traverse the intestine of man and animals during one stage of its life cycle, as Schaudinn showed by experiments on himself and on mice. He infected himself with cysts (fig. 14) by swallowing them, and evacuated the first Chlamydophrys as early as the following day. After the evacuation of numerous specimens on one of the following days the infection ceased. The nucleus of a living specimen is surrounded by a hyaline, strongly retractile chromidial mass, arranged in the form of a ring. Chromatin stains colour it darkly. Asexual inultipHcation (fig. 13, /?), which takes place in faeces, follows 48 THE ANIMAL PARASITES OF MAN a similar course to that of allied forms (e.g., Eiiglypha, Centropyxis). It commences by the cytoplasm issuing from the orifice of the shell and assuming the shape characteristic of the mother organism, but in a reverse position. The nucleus then divides by mitosis, when the daughter nuclei move apart from one another. The chromidial ring also divides into two portions by a process of dumb-bell like con- striction. The one daughter nucleus remains in the mother organism, the other moves towards the daughter individual, which then separates from the parent. Fig. 13. — Chlamydophrys ejichelys. a, free, motile form, showing nucleus, equatorial granules, vacuoles and pseudopodia ; b, dividing organism, x 760. (After Cienkowski.) In this species plasmogamic union of two or more individuals (up to twenty) is frequently observed. Such colonies may similarly divide, and in this way mon- strosities frequently arise. When drying of the fasces, or deficiency of food occurs, encystment takes place apparently spontaneously. The whole body, as stated by Cienkowski, issues from the shell, assumes a spherical shape (probably with discharge of water) and becomes surrounded with a thick membrane (fig. 14). After the addition of water and the escape of the encysted Chlamydophrys^ a new shell must be formed. Schaudinn, who has not given a more detailed description of the process of encyst- ment in this species, but refers to Cienkowski and to similar observations made on Centropyxis^ states of the latter that the encystment takes place within the shell. The sexual nmltiplication is accompanied by shedding of all the foreign bodies and of the degenerating nucleus. The protoplasm, now contracting into a sphere, remains behind in the shell with the chromidial mass. From the latter several new nuclei arise (sexual nuclei) often eight in number. The cytoplasmic sphere then segre- gates into as many spherical portions as there are nuclei present. When they have assumed an oval form, two flagella develop at one LEYDENIA GEMMIPARA 49 pole and the flagellispores swarm out of the shell/ The biflagellate swarm-spores, or gametes, copulate in pairs and apparently the indi- viduals of the pairs of gametes arise, from different mother organisms. The zygote secretes a thick covering which soon becomes brown and rough. These zygote cysts >- or resistant spores must now pass from the in- / \ testine of an animal in order to complete their ( I development. The escape of the cyst contents 1 ^ / does not always take place in the intestine ; 'v - -^=^^^^ often it does not occur until after def?ecaticn. ^-— ^ These shell-less individuals (amoebulae) soon ;>;jrj/j %//il/j7, ency'Sd''; become invested with a shell. But in the on the left the old capsule, alkaline intestinal contents, shell formation may kowskU ^^ ^^"" proceed even while the organism is in the in- testine, and multiplication may take place. Schaudinn's further communication was of special interest ; it was to the effect that Chlamydophrys was related to Leydenia gemmlpara, Schaudinn, 1896. In the fluid removed by puncture from two patients suffering from ascites in the first medical clinic in Berlin, cellular bodies with spontaneous movement were found, which Leyden and Schaudinn regard as distinct organisms. They remained alive without the use of the warm stage for four or live hours, the external temperature being 24° to 25° C. In a quiescent condition they were of a spherical or irregular polygonal form. Their surface was rarely smooth, being beset with protuberances and excrescences (fig. 15). The substance of the body was thickly permeated with light refractile granules with a yellowish shimmer. The hyaline ectoplasm was rarely seen distinctly. All sizes from 3 /^ to 36 yw, in diameter were observed. The movements were rather sluggish, the ectoplasm in the meantime appearing in the form of one or several lamellae, in which also strings of the granular endoplasm occurred, and frequently protruded over the border of the hyaline pseudopodia. The tendency for the joining of several individuals by means of their pseudopodia was so marked that associations ensued similar to those known in free-living Rhizopoda. The cytoplasm enclosed blood corpuscles as well as numerous vacuoles, one of which pulsated slowly about every quarter of an hour. A vesicular nucleus the diameter of which was about equal to one-fifth of the body was present. ' Schaudinn (1903), Arb. a. d. Kaiserl. Gesundh., xix, p. 547. 50 THE ANIMAL PARASITES OF MAN Multiplication took place by means of division andbudding (lig. 15, c), after previous direct division of the nucleus. The buds were supposed to divide repeatedly soon after their appearance, thus giving rise to minute forms of 3 /^. There was a suspicion in both cases that the ascites was associated with maHgnant neoplasms in the abdomen, and autopsy confirmed this view in one case. c X Fig. 15. — Leydenia gemmipara^ Schaud. a, in a quiescent condition, x looo ; ^, in the act of moving, X lOOO ; f, from a fixed preparation, showing a bud, X 1500. The parasite, which has seldom been observed, has been variously interpreted ; for example, it has been regarded merely as altered tissue cells. It is now known, from Schaudinn's researches, that Leydetiia geminipara is connected with abnormal conditions of Chlainydophrys, occasionally occurring as a commensal in the ascitic fluid. The form is produced when pathological conditions of the large intestine create an alkaline reaction of its whole contents. The formation of shells then often ceases, and these naked Chlaniydophrys are enabled to multiply atypically by division and gemmation. Such stages, which are no longer capable of a normal development, are the Leydenia, as Schaudinn has demonstrated. Class II. MASTIGOPHORA, Diesing. Sub-Class. FLAGELLATA, Cohn emend. Biitschli. During the motile part of their life the Flagellata possess one or more fiagella which serve for locomotion, and in many cases also for the capture of food. A few groups {Euglenoidi?2CEy Choanofla^ellata) have only one fiagellum, others two or several of about equal length {Isomasiigoda)^ or of various lengths {Monadina, Heteromastigoda^ Dinofiagellata). The long llagellum is the principal one ; the smaller ones on the same organism are accessory fiagella. The flagella directed MASTIGOPHORA 5 1 backwards, which occur in the Heteromastigoda and are used for clinging, are termed trailing flagella or tractella. At the base of the flagellum, which is almost always at the anterior end, a Choanoflagellate possesses a cytoplasmic funnel-shaped neck or collar. In the parasitic forms an undulating membrane is often present. The body of the Flagellata is usually small, generally elongate and of un- changeable form. It is frequently covered by a distinct cuticle, and, in certain groups, by a hard envelope, or it may be more or less loosely enveloped by a gelatinous or membranous covering. An ectoplasmic layer is thin and not always obvious. The granular cytoplasm contains a varying number of vacuoles, one of which may be contractile, and is generally situated near the area from which the flagella arise, that is, at the anterior extremity. The cytoplasm, moreover, contains the nucleus, which is nearly always single ; and in many species there are also yellow, brown, or green chromatophores of various shapes, such as occur in plants. Some species feed after the manner of green plants (holophytic), or of plants devoid of chlorophyll (saprophytic) ; others, again, ingest solid food, and for this purpose usually possess a cytostome ; the latter, however, in a few forms is not used for its original function, but is connected with the contractile vacuole. Many parasitic forms feed by endosmosis. A few species possess eye-spots with or without light- refracting bodies. Variation in the form of the nuclear apparatus occurs. One nucleus only, which may be compact or vesicular, is known in many species. This nucleus is situated either centrally or sometimes near the flagellar end of the body, but its position is subject to variation. The flagella may arise near the nucleus. Other structures, such as an axial filament and a rhizoplast, may be present. Some flagellates are binucleate, the two nuclei — which often differ in size and shape — being separated from each other. One of these nuclei is the principal, vegetative or trophic nucleus ; the other is an accessory nucleus, frequently termed the blepharoplast, flagellar or kinetic nucleus. One or more small basal granules are often present at or very near the origin of the flagella. Multiplication is by fission, usually longitudinal, which may occur in either the free or encysted forms. Division is initiated by that of the nucleus or nuclei (especially the kinetic nucleus). The basal granule divides also. Collars and chromatophores, if present, likewise separate into two. Variation in the method of doubling the original number of flagella occurs. In most organisms, especially uniflagellate forms, the flagellum splits lengthwise, after division of the basal granule, blepharoplast and nucleus. The daughter flagella may be of the same or different lengths and thicknesses. Other flagellates at division are said to produce new flagella in the neighbourhood of the original ones. The daughter organisms in such cases are provided with one or more parental flagella in addition to newly formed ones. It has been stated that in certain cases the parent flagellate retains all its flagella, while new ones arise ab initio in the cytoplasm of the daughter forms. Multiplication by longitudinal fission may be interrupted sooner or later by the production of gametes, which form zygotes, from which new generations of indi- viduals arise. In many flagellates gamete formation and sporogony are unknown, and asexual reproduction by fission alone prevails. Incomplete division results in the formation of colonies of individuals. These colonies must not be confused with the aggregation rosettes of flagellates found among the parasitic Mastigophora. The individuals of aggregation rosettes are capable of immediate separation from the rosette at will. A number of parasitic Flagellata produce non-flagellate stages which are very resistant to external conditions, the assumption of which forms serves to protect the organisms during their transference from one host to another. Such non-fla^ellate forms possess one or more nuclei, are usually of an oval or rounded contour, and 52 THE ANIMAL PARASITES OF MAN are capable of developing into the full flagellate on the return of more favourable conditions. These forms are often known as the post-flagellate stage of the organism. When ingested by a new host, the post-flagellate coat becomes more flexible, and the phase of the organism which now recommences growth is known as the pre-flagellate stage ; it gradually develops into the typical flagellate organism. Many Flagellata live free in fresh and salt water. They prefer stagnant water, rich in organic products of decomposition, such as puddles, swamps and pools. Those forms developing shells and colonies are, as a rule, adherent. A number of species are parasitic in man and animals, living mostly within the intestine or in the blood. It is usual to classify the Flagellata in four orders : Euflagellata, Dinoflas^ellata^ Choanoflagellata^ and Cysto flagellata^ of which only the Euflagellata are of mterest to us. This is a group comprising numerous species, for the further classification of which the number and position of the flagella are utilised. The Euflagellata observed in man belong to the Protomonadina as well as to the Polymastigina. The former possess either only one or two similar flagella, or one principal and one or two accessory flagella. The Polymastigina possess at least three flagella of equal size, or four to eight of unequal size, inserted at different points. An undulating membrane may be present in members of both groups. It must also be pointed out that unicellular organisms with one or several flagella are not always classified with flagellates, for such forms occur in Rhizopods as well as temporarily in the lower plants. In addition, the examination of the flagellates, especially the parasitic species, is very difficult on account of their diminutive size and great activity ; thus it happens that certain forms cannot with certainty be included in the group because their description is insufficient. Order. Polymastigina, Blochmann. The Polymastigina contains flagellates with three to eight flagella. Some of the Flagellata parasitic in man belong to the Polymastigina, and to two or three genera that are easily distinguishable. Genus. Trichomonas, Donne, 1837. The body is generally pyriform, the anterior part usually rounded, the posterior part pointed. There are at the anterior extremity three (? four) equally long flagella that are sometimes matted together. A blepharoplast (kinetic nucleus) and basal granule are present, together with a supporting structure known as an axial filament or axostyle. In addition there is an undulating membrane, bordered by a trailing flagellum, that commences at the anterior extremity and proceeds obliquely backwards. The nucleus, which is vesicular, is situated near the anterior extremity, and behind it are one or more vacuoles, none of which seems to be contractile. These flagellates are parasitic in vertebrate animals, and live chiefly in the intestine. Trichomonas vaginalis, Donne. The form of the body is very variable, and is elongate, fusiform or pear-shaped, also amoeboid. The length varies between 15/1, and 25 /i, and the breadth between 7 ytt and 12 /i. The posterior extremily is drawn out to a point and is about half the length of the remainder TRICHOMONAS VAGINALIS 53 of the body. The cuticle is very thin and the body substance finely granular. At the anterior extremity there are three — some say four ^ — flagella of equal length which are frequently united together, at least at the base, and are easily detached. There is an undulating membrane (fig. i6) which runs spirally across the body, arising from the place of insertion of the flagella, and terminating at the base of the caudal process. A cytostome seldom is recognizable in fresh specimens, but is apparently present. The nucleus is vesicular, elliptical and situated near the anterior extremity.^ Multiplication takes place by division (Marchand). Encysted forms are almost un- known. Trichomonas vaginalis lives in the vaginal mucus of vvomen of various ages, not in normal mucus, but in mucus of acid reaction. It is found in menstruating females as well as in females who have passed the menopause. It occurs in pregnant and non-pregnant women, even in very young girls, provided always that they have a vaginal catarrh with acid reaction of the secretion. Should the acid reaction change, as, for instance, during menstruation, the parasites disappear, as they do likewise on injection of any alkaline fluid into the vagina. A low temperature (below + 15° C.) is also fatal to the parasites. These flagellates can pass from the vagina through the urethra into the bladder, and produce severe catarrh, and are not easily removed. r. vaginalis appeared to be a parasite specific to the female organs and not transmissible to man. However, several observations have since been made that confirm the occurrence of this species in the urethra of the male. The infection apparently takes place through coitus when changes are present in the urethral mucous membrane. At any rate, three cases observed point to this circumstance. Attempts at experimental transmission to rabbits, guinea-pigs and dogs failed (Blochmann, Dock). So far, the manner in which women become infected is unknown. Fig. 16. — Trichomonas vaginalis, Donne. X 2,000 approx. (After Kunstler.) Four flagella are repre- sented, but usually only three are present. 1 To explain this discrepancy it is stated that the border of the undulating membrane can be detached in the form of an independent flagellum. But Parisi (1910) places such quaciriflagellate forms in the sub-genus Teiratrichomonas , Arch. f. Protistenk., xix, p. 232. ■^ According to Marchand, the nucleus is connected with a line, which becomes visible on addition of acetic acid, terminates at the posterior extremity, and does not correspond to the line of insertion of the undulating membrane. This formation probably is the same as the axostylein Trichomonas bairacho?'ttm, Perty. Blochmann (1884) also mentions two longitudinal rows of granules, which commence at the same place as the nucleus and converge posteriorly. 54 THE ANIMAL PARASITES OF MAN Trichomonas intestinalis, R. Leuckart, 1879= Trichomonas hominis, Davaine, 1854. Some authors believe that a second trichomonad inhabiting man, Trichomonas intestinalis, R. Lkt., is identical with Trichonionas vaginalis, Donne. Leuckart's species was based on the discoveries of Marchand (1875) and Zunker (1878), who stated that according to all appearances, and in their opinion, it was the same as Cercomonas intestinalis, Lambl, 1875 {nee 1859), which they found in the fasces of patients suffering from intestinal disorders. The organism is described by them as being pear- shaped and 10 yLt to 15 yLt in length and 3 /x to 4' yLt in breadth. The posterior extremity terminated in a point (fig. 17). Fig. 17. Trichomonas intestinalis, (After Grassi.) A row of twelve or more cilia was said to commence at the anterior end and extend Lkt. over the body. Leuckart stated that this parasite, placed by the two authors in the genus Cerconw?ias\ w^s a Trichomonas, and that they mistook the undulating membrane for cilia, and overlooked the flagella. Notwithstanding its striking similarity with T. vaginalis, it was said to be distin- guishable from that species by ditferences in the undulating membrane. Lambl's C. intestinalis^ (of 1875) which corresponds with C. hojnifiis, Davaine^ (1854), is regarded by Leuckart as a true Cercomonad (characterized by a flagellum and the absence of an undulating membrane, see p. 61), and is thus generically distinct from Trichomonas. The correctness of Leuckart's judgment in regard to Marchand-Zunker's flagellate was demonstrated by Grassi's researches, accounts of which were published soon after. In about 100 cases of bowel complaints in North Italy and Sicily, Grassi found Flagellata in the stools, which he first named Monocercomo7tas and Ci?ncr?io- monas, but later termed Trichomonas. However, in opposition to Leuckart, Grassi has also classified Uavaine's C. homi7iis {= C. intestinalis, Lambl, 1875) ^s Trichomonas, and most authors have followed his example. Hence arose the use of the name Trichomonas hominis. It was through Janowski (1896) that the former view was agam taken up. After a review of the literature, the occur- rence of Cercomonads in the intestine of human beings in addition to Tricho- monads was considered by the author to have been proved, and he added a description of the Trichomonads. According to this, all morphological distinction 1 Under the term Cercomonas intestinalis, Lambl in different years has described two entirely distinct Flagellata, namely, in 1859 ("Mikr. Unters. d. Darm- Excrete," Pnag. Vierteljahrsschr. f. prakt. Hlkde., Ixi, p. 51 ; and Lambl, A. d. Franz-Josephs- Kinder spitale in Prag, Prag, i860, i, p. 360), a form that at the present day is termed Lainblia intes- tinalis ; and in 1875 (^^^ the Russian Medical Report, No. 33), a species identical with Cercomonas hominis, Dav. ^Davaine, C. , *' Sur les anim. infus. trouv. dans les selles d. malad. atteints du cholera et d'autr. malad.," C. R. Soc. Biol,, 1854, ii, p. 129. TRICHOMONAS INTESTINALIS 55 between T. vaginalis^ Donne, and T. intestmalis^ LeuckaU, disappeared. On the other hand, it is worthy of note that the smaller size, the more pear-shaped form, and the longer flagella differentiate T. intestinalis (= T. hommis) from T. vaginalis} The easily deformed pear-shaped body has three free flagella anteriorly, and an undulating membrane with its flagellar bordc" terminating in a short free flagellum posteriorly (figs. 17, 18). The undulating membrane may coil itself spirally round the body. A supporting rod or axo- style projects as a posterior spine. It appears to begin near the nucleus and blepharoplast, which are situated near the more rounded, anterior end of the body. There may be a chromatoid basal supporting line along the body for the undulating membrane. Rows of chromatoid granules are sometimes situated along one side of the axostyle. A cytostome may sometimes be seen. In mice, Wenyon (1907) found these parasites to vary in length from 3 //, to 20 \i. They occur in the caecum and intestine of mice, where their internal structure seems more obvious than in man. The flagellates divide by longitudinal fission. T. intestinalis, R. Leuckart, appears to be capable of settling in all parts of the human intestine in which the contents have an alkaline reaction. Trichomonads have been cited as occurring in the oral cavity by depressior°^anter- Steinberg, Zunker, Rappin and Prowazek ; in the ioriy, undulating oesophagus by Cohnheim, and in the storpach by ^^ a^xos^y^l"! Strube, Cohnheim, Zabel, Hensen and Rosenfeld. x 2,500. Original. The normal situation seems to be the small intestine. The parasites then appear in the dejecta, especially in various intestine, diseases the course of which is connected with an increased peristalsis. They are also found in healthy persons, from whom they are obtained after the administration of laxatives. They have been regarded by some workers as commensals, which, however, have the power of accelerating 1 For the present the following should be regarded as synonymous : Protoryxomyces coprinarius^ Cunningham {Quart. Journ. Micr. ScL [2] 1880, xxi, p. 234), {Zeitschr. f. Biol., 1882, viii, p. 251). Monocercomorias hominis, Grassi, 1882. Cimcenomonas hominis^Qx2&%^_ 1882. Trichomonas hominis, Grassi, 1888. Cercomonas coli hominis^ May {DetUschei Archiv. f. klin. med., 1 89 1, xlix, p. 51). Monocercomonas kominis, Epstein [Frag. med. Wochenschr. 1893, ^^s. 38-40). Trichomonas confusa, Stiles {Zool. Anz., 1902, xxv, p. 689). Trichomonas elongata. Trichomonas elliptica, Cohnheim {Deutsche med. Wochenschr., 1903 xxix, Nos. 12-14). Trichomonas elotigata. Trichomonas caudata, Trichomonas Jlagellata, Steinberg (A7^ze/(?r Zeitschr. f. neiiere A/edicin, 1862). Trichomonas pulmonalis, A. Schmidt, {Miinch. med. Wochenschr., 1895, No. 51), and St. Artault {Arch, de parasit. 1898, i, p. 279). Fig. 18.— 7>/- chomonas intestin- alis from man, showing anterior 50 THE ANIMAL PARASITES OF MAN the onset of intestinal complaints, or at least of adding to them. They have been found in cases of carcinoma of the stomach, and in other diseases of that organ in which the acid reaction ceased. Naturally, whether all the reports relate to the same species of Trichomonas must remain undecided. Certain authors (Steinberg, Cohnheim, van Emden) accept several species. Provvazek speaks of a variety of T. intestinalis inhabiting the oral cavity. This was distinguished by a posterior process exceeding the length of the body fourfold, and by a somewhat unusual course of the undulating membrane. The food of this form, which was found in the whitish deposit present, especially in the cavities of carious teeth, consisted almost exclusively of micrococci. Schmidt and St. Artault named the Trichomonads found in pathological products {e.g.^ gangrene, putrid bronchitis, phthisis) of the lungs of man, as Trichomonas pulmonalis. Trichomonads have also been found by VVieting in lobular pneumonia in the lungs of pigs. It is still uncertain in what way the infection takes place. Experiments in the transmission of free trichomonads to mammals [per os)^ in which the same or allied species occur (guinea-pigs, rats, apes), have been without result. Probably encyst- ment is necessary. Such conditions are mentioned by May, KUnstler, Roos, Schurmayer, van Emden, Prowazek, Galli-Valerio and Schaudinn. According to Prowazek, intestinal trichomonads of rats become encysted for conjugation. In the cyst an accumulation of reserve food material occurs, causing distension. The nuclei of the conjugants each give off a reduction body and, after fusion, produce the nuclei for the daughter individuals. According to Schaudinn the intestinal tricho- monads lose their flagella before conjugation, become amoeboid and encyst in twos, the formation of a large agglomeration of reserve substance accompanying this. Galli-Valerio found double-contoured cysts in the faeces of trichomonad-infected guinea-pigs, after the faeces had been kept for a month in a damp chamber. When exposed to heat small flagellates escaped from them. Administration of such material containing cysts resulted in severe infection with trichomonads, and death of the experimental guinea-pigs followed. The cyst wall is clearly a protection against the deleterious acid reaction of the stomach contents. Alexeieff (1911) and Brumpt (1912) think that the trichomonad cysts of man are really fungi, while other workers also doubt encystment among trichomonads. Wenyon (1907) states that T. intestinalis in mice produces spherical contracted forms which escape from the body in the faeces. Air, water, and under certain circumstances even food may be regarded as vectors for the trichomonads. The occurrence of the organisms in the oral cavity, and still more so in the lungs, is in favour of the air being the transmitting agent. An observation made by Epstein supports the idea of water transmission. Multipli- cation of the trichomonads, once they have gained access to the body, is effected by longitudinal division commencing at the anterior end (Kiinstler). " Cercomonads " with several flagella and an undulating membrane, as well as trichomonads, have been observed by Ross in some cases of cutaneous ulcers. Mello-Leitao (1913)^ has described flagellate dysentery in children in Rio de Janeiro. He states that it is due to T. intestinalis and Lamhlia intestinalis either separately or together. Flagellate dysentery, he thinks, is benign and is the most frequent form of dysentery in infants. The flagellates are pathogenic to infants under three years of age. Escomel (1913)^ found 152 cases of dysentery in Peru due solely to Trichomonas. Such cases are probably widespread ' BriL Jow-n. Childretis Diseases, x, p. 60. ^ jguii. Soc. Path. Exot., vi, p. 120. TETRAMITUS MESNILI 57 Genus. Tetramitus, Perty, 1852. Tetramltus mesnili, Wenyon, 1910. Syn. : Macrostoma mesnili^ Chilomastix mesnili^ Fanapapea intestinalis. The genus Tetramitus differs from Trichomonas in possessing an undulating membrane inserted in a deep groove or cytostome. There are three anterior flagella. The pear-shaped organism measures 14 yu by 7 yLt, but smaller examples occur. T. mesnili occurs in the human intestine, having been described by Wenyon^ (1910) from a man from the Bahamas in the Seamen's Hospital, London. Its occurrence is widespread. Alexeieff considers that Macrostoma and Tetramitus are synonymous. The parasite is the same as Fanapapea intestinalis, Prowazek, 1911, from Samoa. Brumpt (191 2) found T. mesnili to be the causal agent of colitis in a Frenchwoman. Nattan-Larrier (1912) considers it of little pathological importance. GabeP (1914) described an interesting case of seasonal diarrhoea acquired in Tunis, in which a new Tetramitid was the causal agent. The organism was pear-shaped, without an undulating membrane, and measured 6*5 //, to 8 //, by 5 //. to 6 /x. The cytostome was large, and there was no skeletal support. Encystment occurred. Gabel named the organism Difdmns tiinensis and considered that it was pathogenic. Genus. Lamblla, R. Blanchard, 1888. Syn., Dimorphus, Grass), 1879, ^^<^ Haller, 1878; Me^astoma, Grassi, 1881, nee de Blainville. The body is pear-shaped, with a hollow on the under surface anteriorly. It has four pairs of tlagella directed backwards, of which three pairs lie on the borders of the hollow disc, and the fourth arises from the pointed posterior extremity. Lamblia intestinalis, Lambl, 1859. Syn.: Cercomonas intestinalis^ Lambl, 1859 {nee 1875); Hexamitus duodenalis, Davaine, 1875; Dimorphus inwris, Grassi, 1879; Megastoma enterieum^ Grassi, 1881 ; Megastoma intestinale, R. Blanch., 1886; Lamblia duodenalis^ Stiles, 1902. The organism is pear-shaped and bilaterally symmetrical. It is from 10//. to 21 /z long and 5 yw, to 12 yu, broad and possesses a thin cuticle. Anteriorly an oblique depression is present, which functions as a sucking disc (fig. 19, s). Its edges are raised above the general surface and are contractile. It corresponds to a peristome and acts as an adhesive organ (fig. 20, 6, c). No true cytostome is present. A double longitudinal ridge, representing axostyles, extends from the sucking disc to the tapering posterior extremity, which is prolonged as two flagella from 9 /x to 14 /^ long. Lamblia intestinalis possesses eight flagella (fig. 19). The first pair 1 Parasitology, iii, p. 210. '^ Arch. f. Protistenk., xxxiv, p. I. 58 THE ANIMAL PARASITES OF MAN of flagella, which cross one another, arise in a groove formed by the anterior edge of the sucking disc. Two pairs of flagella (lateral and median) are inserted on the posterior edge of the disc, while the posterior flagella occur at the tapering posterior extremity of the body. Basal granules are found at the bases of the flagella. The median flagella are most active in movement, the anterior and lateral flagella being less motile, as they are partially united to the body for part of their length. The nuclear apparatus is situated in the thin, anterior, hollowed part of the body. It is at first dumb-bell shaped, the " handle " of the dumb-bell being formed by a very slight connecting strand, which eventually separates, so that the flagellate becomes binucleate, and thus completes the general bisymmetry of the organism. There is a karyosome in each nucleus. Other bodies of unknown function, and possibly composed of chromatin, occur on or near the axostyles. . Fig. 19. —Lamblia intestinalis. A, ventral view ; B, side view ; N, one of the two nuclei ; ax.^ axostyles; fl'^,Jl'^,Jl^,fl^, the four pairs of flagella; s, sucker-like depressed area on the ventral surface; x, bodies of unknown function. (After Wenyon.) Division has not been observed in the flagellate stages of the Lamblia, but it occurs within the cysts. The resistant cysts (fig. 20, e) are oval and are surrounded by a fairly thick, hyaline cyst wall. They measure 10 //, to 15 /x by 7 //< to 9 /x, and may be tetranucleate. According to Schaudinn, the cysts arise from the conjugation of two individuals, and nuclear rearrangement occurs. LAMBLIA INTESTINALIS 59 L. intestinalis occurs in its flagellate stage in the duodenum and jejunum, and rarely as such in the other parts of the intestine. Normally it is found in the large intestine as cysts, which are voided with the faeces. The hosts of Lamblia include Mtis muscuhis, M. rattuSy M. decumanus, M. silvestris, Arvicola arvensis and A, amphihius the dog and cat, rabbit, sheep and man. Cysts voided with the faeces of infected animals reach plants or drinking water, and thence are transferred to man. The flagellate in these different hosts exhibits some variation in size and in the problematic chromatic bodies. Bensen has suggested the species L. intestinalis from man, L. ninris from the mouse and L. cunicnli from the rabbit. It is not certain whether these different species are necessary, as the variation may be due to differences of environment. Fig. 20. — Lamblia intestinalis. a, from the surface ; b, from the side ; c, on intestinal epithelium cells ; d, dead and ^ ^ "it ^^5^ '^^ ^ >; '^ <^ " inA^o.v ^ *■>!'*" ^^t eui-pjjii inAxoiv ■ — *— ^ ^ >i "o T'"' "* - — 3 ^ ^ ^s- »^ ':::»■ ;n 5 ^"~^*.- ■ 2; 5 o~> — ^ T^>' 'cl. 00 ■■= — '^"^ - =* S'd^ >o eurejS Al iJkxOlV * "^ > _ f ^to< ^ r. >o tM * W* ^ Q -^ „-— 1— »-"• *-'^P ~ ^ »U- ' '^C g "e * ' « o "* ^ """": ""■"— — — — Q, "■■"S ISW ^ "'^i^ : > S(2 2 --liL |ii:ii.::i::::::±: § -r - - .Ji: l< ^3 ^X Z^ 1^ ^ ^^ '^s. ^S ^ ^^ .:-=* ^c 1- ^ ^^ ':. c:i5 ^ ^ ^ ■ '^ ■*' ' ^ » S hfl § ^ "^^ i:* -i.S 5: ^^ t "^ g ig- ^ ^^ - --^^ ^%. SiJ ^ ^*lOH>-i.Q.ooixtovoT'cDO'-< HMH< ^ -««>-< -T •qi9LJU9aUPJ 'poo|g J.0 -Luujn;? J9cl 99UJ0C0U'pdfijx jx> gj^qtunyj -ajnguiaaLus^^ 8o THE ANIMAL PARASITES OF MAN parasite usually found in man and in animals sub-inoculated from cases of sleeping sickness in Nyasaland. It has since been found in German East Africa and Portuguese East Africa, while Ellacombe has described a case from North-western Rhodesia. (3) Serum Reactions. — Interesting experiments on this subject were performed during 1911 and 1912 by various French investigators. {a) Action of Immune Serum (Mesnil and Ringenbach)^ : (i) A goat was infected with T. rhodesiense. Twenty-two days later its serum mixed with T. rhodesiense was injected into a mouse. Result: Pro- tection. (2) The serum mixed with T. gamhiense was injected into a mouse. Result : Infection. (b) Action of Baboon Serum. — Contrary to T. gambiense, T. rhode- siense is very susceptible to human and baboon sera. Mesnil and Ringenbach^ showed that a dose of i c.c. of baboon (Papio anubis) serum cured mice infected with T. rhodesiense. In the same dose it acted very feebly on T. gambiense. (c) Action of Human Serum. — i c.c. of human serum cured T. rhodesiense mice in three out of four cases ; on T. gambiense mice there was no appreciable effect. Laveran and Nattan-Larrier^ have shown the same, namely, that human sera act on T. rhodesiense, but are quite without action on J. gambiense. (d) Trypanolytic Reactions. — Mesnil and Ringenbach* have also shown that the sera of animals (man, monkey and guinea-pig) infected with T. gambiense are trypanolytic for the homologous trypanosome, that is, T. gambiense, but have no action on the hetero- logous trypanosome, that is, T. rhodesiense. (4) Cross Immunity Experiments. — (a) Mesnil and Ringenbach^ immunized a monkey (Macacus rhesus) against T. gambiense. It was inoculated with T. rhodesiense on June 7, 191 1 ; on June 27 trypano- somes appeared, the infection being slight ; on July 4 it died. A control died in ten and a half days. (b) Laveran^ immunized a goat and mice against T. gambiense. When they had acquired a solid immunity, they were inoculated with T. rhodesiense. They became infected like the controls. (c) Laveran and Nattan-Larrier^ immunized a ram against T. brucei, it subsequently became infected with T. rhodesiense. {d) Laveran^ immunized a ram and a sheep against different strains of T. brucei. Inoculated with T. rhodesiense they both acquired acute infections and died. Conclusion : T, rhodesiense is not T. brucei. ' C.R. Soc.BioL, Ixxii, p. 58. ^ (j /^ Acad. Sc/., 153, p. 1,097. 8 C./^. Acad. Sci., 154, p. 18. 4 c.J^. Soc. Biol., Ixxi, p. 609. * C.R. Soc. Biol., Ixxi, p. 271. « Bull. Soc. Path. Exot., v, pp. 26, 241. ^ C.R. Acad. Sci., 154, p. 18. « Bull. Soc. Path. Exot., v, p. loi. TRYPANOSOMA KHODESIENSE 8 1 When the converse set of experiments is tried, namely, immuniz- ing an animal against T. rhodesiense, and then inoculating with T. gambiense, the difficulty immediately arises that it is impossible to immunize an animal against T. rhodesiense, owing to its virulence. But a partial and transitory immunity to T. rhodesiense can be obtained by treating the infected animal with drugs, such as arsenophenylglycin. The results, so far as Ihey go, seem to show that an animal immunized against T. rhodesiense is immune not only to T. rhodesiense, but also to T. ganibiense, a fact which, according to Mesnil and Leger, does not invalidate the specificity of T. rhodesiense, but tends to show that the two trypanosomes are closely related. (5) Mode of Transmission and Reservoir. — Kinghorn has shown that T. rhodesiense is transmitted by Glossina niorsitans in which it under- goes development. Kinghorn and Yorke^ found that about 16 per cent, of the wild game examined in Northern Rhodesia was naturally infected with T. rhodesiense. The wild game examined included water- buck, hartebeest, mpala, bushbuck and warthogs. One native dog near the Nyasaland border was found infected, but not domestic stock. Taute doubts whether T. rhodesiense really occurs in wild game. Approximately 3*5 per cent, of the tsetse flies fed on infected animals may become permanently infected with T. rhodesiense, and capable of infecting clean animals. Furthermore, a tsetse fly when once infective probably remains infective for the rest of its life. Kinghorn and Yorke, however, have shown that climatic condi- tions, namely, those of temperature, also affect the infectivity of the tsetse fly, as the ratio of flies capable of transmitting T. rhodesiense to those incapable of transmitting the virus is i : $34 ^^^ hot valley districts (e.g., Nawalia, Luangwa Valley, temperature 75° to 85° F.), while on elevated plateaux (e.g., Ngoa, on the Congo-Zambesi water- shed, temperature 60° to 70° F.) the ratio falls to i : 1312. Mechanical transmission by the tsetse fly does not occur, if a period of twenty-four hours has elapsed since the infecting meal. Developniental Cycle in the Fly. — The period which elapses between the infecting feed of the flies and the date on which they become infective varies from eleven to twenty-five days in the Luangwa Valley, according to Kinghorn and Yorke. Attempts carried out at laboratory temperature on the Congo-Zambesi plateau, during the cold season, to transmit T. rhodesiense by means of G. morsitans were always unsuccessful. The developmental cycle of the trypano- some in the fly is influenced by the temperature to which the flies are subjected (as stated above). The first portion of the develop- mental cycle proceeds at the lower temperatures (60° to 70° F.), but higher temperatures are necessary for the completion of the develop- ' Annals Trop. Med. and Parasitol., vii, p. 183. 82 _ THE ANIMAL PARASITES OF MAN ment of the trypanosome. Kinghorn and Yorke found that the trypanosomes may persist in the fly, at an incomplete stage of their development, for at least sixty days when the climatic conditions were unfavourable. The first portion of the developmental cycle of the trypanosome takes place in the gut of the fly. Invasion of the saHvary glands of the tsetse is secondary to that of the intestine, but is necessary for the infectivity of the fly. A relatively high mean temperature, 75° to 85" F., is essential for the passage of the trypanosomes into the salivary glands and the completion of their development therein. Kinghorn and Yorke^ state that the predominant type of trypano- some in the intestine of infected G. morsitans was a large broad form, quite different from that which is most common in the salivary glands. The trypanosome in the glands resembles the short form seen in the blood of the vertebrate host. The authors quoted state that both the intestinal and salivary gland forms of infective G. morsitans are virulent when inoculated into healthy animals. Bruce and colleagues^ have quite recently (June, 1914) published an account of their investigations of T. rhodesiense in G. morsitans in Nyasaland. (Incidentally it may be remarked that Bruce considers T. rhodesiense to be identical with a polymorphic strain of T. hnicei — see pp. 83, 94). The development of T. rhodesiense takes place in the alimentary canal and salivary glands, not in the proboscis, of the tsetse fly. In feeding experiments with laboratory bred flies, as well as with a few wdld flies, fed on infected dogs or monkeys, only 8 per cent, of the flies were found to be infected on dissection. Of such infected flies, however, only some allow of the complete develop- ment of the trypanosomes within them, in other words only about I per cent of the flies become infective. The length of time which elapses before a fly becomes infective varies from fourteen to thirty- one days, averaging twenty-three days, when kept at 84° F. (29° C). The dominant intestinal type of flagellate in the fly is that seen in the proventriculus, which contains many long, slender trypanosomes. These proventricular forms find their way to the salivary glands, wherein crithidial and encysted forms are seen. They change into "blood forms," which are short, stumpy trypanosomes and are infective. ''The infective type of trypanosome in the salivary glands — corresponding to the final stage of the cycle of development — is similar to the short and stumpy form found in the blood of the verte- brate host." The cycle is thus very similar to that of T, ganibiense in G. palpalis (fig. 30). Culture. — ]. G. Thomson (1912),^ and subsequently Thomson ' Annals Trap. Med. and Parasitol., vii, p. 281. ^ Froc. Koy. Soc, B, Ixxxvii, p. 516. ^Annals Trop. Med. and Parasitol., vi, pp. 103, 331. TRYPANOSOMA CRUZI 83 and Sinton, succeeded in cultivating T. rhodesiense in a modified Novy-MacNeal medium. The development obtained resembled that of the trypanosome in the intestine of Glossina, General Note on Trypanosomes with Posterior Nuclei. Posteriorly placed nuclei have been found to occur not only in T. rhodesiense by Stephens and Fantham (1910), but also in T. pecaiidi by Wenyon {191 2), in T. brucel by Blacklock (191 2), and in T. eqiii- perdiun by Yorke and Blacklock (191 2). Recently Stephens and Blacklock (1913)^ have shown that two trypanosomeS; different morphologically, have been confused under the name T. hriicei. One of these is polymorphic {i.e., it exhibits long and slender as well as short and stumpy forms) and came from Uganda, while the other is monomorphic and is the original Zululand strain described by Bruce from cattle suffering from ^^nagana." Bruce (1914) considers that morphological change has occurred in T. brucei in its passage through laboratory animals, and thus explains the diversity of views. The posterior nuclear forms described by Blacklock occurred in the Uganda strain of T. brucei. (See p. 95.) Similarly, a posterior nuclear form, T. equi, has been separated from T. eqniperdum. (See p. 98.) Again, Bruce and his colleagues on the Royal Society Commis- sion investigating sleeping sickness in Nyasaland, have stated (April, 1913) that ^'evidence is accumulating ih-^iZ, rhodesiense and T. brucei (Plimmer and Bradford) are identical." The exact identity of trypano- somes showing posterior nuclei is, then, far from settled, although Laveran by cross immunity tests has declared that T. brucei is distinct from T. rhodesiense. No one has yet seen posterior nuclei in T.gambiense. Trypanosonna cruzi, Chagas, 1909. Syn. : Schizotrvpanutn cruzi, Chagas, 1909. The trypanosome was discovered by Chagas^ in the intestine of the bug, Triatoma (Conorhinus) megista, in Brazil, and then in the blood of a small monkey bitten by the bug. A little later it was found in the blood of a child, aged two years, suffering from irregular fever, extreme anaemia and enlarged glands in the State of Minas Geraes, Brazil. Chagas found that he was able to infect many of the usual laboratory animals with the trypanosome, by allowing the bug to bite them. He was also able to culture the parasite on blood agar. Chagas found the Reduviid bug, Triatoma megista, in the houses of the poorer inhabitants of the Brazilian mining State, and that it attacked the people, more especially the children, at night, biting the face. On this account the insect is called *' barbeiro " by the ' Froc. Roy. Soc, B, Ixxxvi, p. 187. ^ Me7?i. Inst. Oswaldo Crtiz., i, p. 159. 84 THE ANIMAL PARASITES OF MAN inhabitants. The bite is somewhat painful. The disease has since been found in other parts of Brazil, e.g., Matta de Sao Joao in Bahia province, Goyaz, Matto Grosso and Sao Paulo provinces, as well as in Minas Geraes. Morphology. — The trypanosome has a large blepharoplast or kinetic nucleus. It is stated to occur both free and in the red blood corpuscles in the peripheral blood. It is about 20 /a long, on an average. Two forms of the parasite (fig. 33, 6, 7) are described in the human blood. In one free form there is a large egg-shaped blepharoplast and the posterior (aflagellar) end of the parasite is drawn out. The blepharoplast (kinetic nucleus) may have a chromatin appendage. The nucleus is oval or band-like, containing a karyosome. The flagellum, starting close to the blepharoplast or its appendage, has a free portion of variable length. The other free form in the blood has a more or less round, terminal blepharoplast, smaller than in the first form, without a chromatin appendage as a rule. The body of this second form is decidedly broader than that of the first mentioned. Fig. 33. — Jrypavosoma cruzi. Schizogony, i, merozoite in red blood corpuscle ; -?, parasite totally enclosed in red cell, no flagellum or undulating membrane; j-5, parasitts partially enclosed in red cell ; 6, 7, parasites in human blood ; 8-11, parasites in lungs of the monkey, Callithrix ; 12, jj, initial forms of schizogony ; i^, 75, schizogony in the lungs of Callithrix. (After Chagas.) The dimorphism has been interpreted sexually, the first mentioned forms being termed males, the second ones females. The correctness of this interpretation is very doubtful. No sign of longitudinal division was ever seen in the peripheral blood or in the internal organs. The '' endocorpuscular " forms may be completely or partially enclosed in the red cell or only attached thereto (fig. 33, i-S)' ^^ the beginning of infection the endocor- puscular forms are the more numerous. Some authorities, however, doubt these stages. Life-history in the Vertebrate Host. — Chagas found fluctuations in the number of the parasites in the peripheral blood. He believes the increase of the parasites to be periodic. TRYPANOSOMA CRUZI 85 The investigations of Chagas and of Hartmann have revealed two types of miiltiphcation which take place in the internal organs of the vertebrate host. (a) The first type — which possibly belongs to another organism, Pneiujiocystis carinil, see p. 90 — occurs in the capillaries of the lungs. Tlie flagellate parasite entering the lung capillaries loses its flagellum and undulating membrane. Its body becomes curved, and the two ends fuse, and so an oval mass is formed (fig. 33, 8-1 1). In some cases the blepharoplast disappears, in other cases it blends or fuses with the nucleus. The nucleus of the rounded parasite then divides into eight by successive divisions (fig. 33, 12-15). Next the body, which is surrounded by its own periplast, also divides, giving rise to eight tiny daughter individuals or merozoites (fig. 33, 75). The merozoites lie inside the periplast, which acts as a sort of " cyst wall." The merozoites are said to exhibit dimorphism, and Chagas has interpreted the dimorphism in terms of sex. The daughter forms, produced by the parent trypanosomes which kept their blepharoplasts, themselves have blepharoplasts as well as nuclei, and have been termed ^'males'' or '' microgametes." The merozoites, arising from parent trypanosomes which lost their blepharoplasts, have themselves only nuclei, and have been called ^'females" or ^^ macrogametes." In the case of the so-called ''female" forms the single nucleus divides into two unequal parts, of which the smaller becomes the blepharoplast, and a flagellum is formed later. The so-called '' males " possess early a rudiment of a flagellum. Both kinds of merozoites escape from the parent periplast wall, and enter red blood corpuscles. They grow into flagellates within the corpuscles, and then become free as adult trypanosomes in the blood-stream. (6) The second mode of multiplication is one of asexual reproduction (schizogony or agamogony). It was first described by Hartmann from hypertfophied endothelial cells of the lungs. It has since been found in the cardiac muscle, in the neuroglia of the central nervous system, and in striped muscle (fig. 34). fig. 34. — Trypanosoma In laboratory animals it has also been found ^':^^''- Transverse section ..... - . , ^ of a striated muscle con- m the testicle and suprarenal capsules. In taining rounded forms of these tissues the parasite is intracellular, ap- the parasite in the central ,, 1,1, • , , portion. X 1,000 approx. pearmg as a small rounded body with nucleus (After Vianna.) and blepharoplast, without flagellum or undu- lating membrane. In other words the parasite is Leislunania-Vike in the body tissues, and recalls the organism of kala-azar. Chagas considers this second mode of multiplication to be strictly asexual. By this means the number of parasites in the vertebrate 86 THE ANIMAL PARASITES OF MAN host is increased, and symptoms are produced. On the other hand the first mode of multiphcation, seen in the hing capillaries, is considered by Chagas to be a process of gametogony, in which sexual forms are differentiated. He finds that (i) the adult trypanosomes exhibit a dimorphism in human blood rarely seen in artificially infected guinea-pigs. In these guinea-pigs (infected from guinea-pigs) the so-called gametogony in the lungs is seldom seen. (2) The intermediate host, Triatoma (Coiiorliiiitis), becomes infective if fed directly on infected human blood, but very rarely so if fed on guinea- pigs. Chagas is led to believe that the occurrence of sexual forms constantly in the blood of man implies a greater resistance to infection on the part of man than on the part of guinea-pigs or other animals, assuming the general hypothesis that the formation of gametes represents a reaction of the Protozoon to unfavourable conditions. In human infection the number of parasites is always less than in laboratory animals, and their presence in the blood is transitory, lasting from fifteen to thirty days in acute cases. In many cases examination of the tissues at death has shown the presence of parasites in patients who did not exhibit them in the general circulation. Fig. 35. — Trypanosoma crtizi. Development in Triatoma inegista. i'6, forms found in the mid gut of Triatoma ; 7 flagellate forms found in ihe posterior part of the gut of Triatoma. (After Chagas.) Life History in the Invertebrate Host. — About six hours after the ingestion of infected blood by the bug {Triatoma inegista), the kinetic nucleus of the trypanosome moves toward? the nucleus, and the flagellum is usually lost' (fig. 35, 7-5). The parasite becomes rounded and Leishmania-Vike (fig. 35, j-5), and multiplies rapidly by division. After a time, multiplication having ceased, the rounded forms become pear-shaped and develop a flagellum at the more pointed end. Crithidial forms (fig. 35, 7) are thus produced and pass into the intes- tine, where they multiply and may be seen in about twenty-five hours after the ingestion of blood. The crithidial forms may also be found in the rectum and faeces. The last stage in the invertebrate is a small, trypanosome-Iike type, long and thin with a band-like nucleus and conspicuous kinetic nucleus. These parasites are found in the hind gut and in the body cavity. They find their way into the salivary glands, and are the forms (fig. 36) which are transmissible TRYPANOSOMA CRUZI 87 to a new vertebrate host. The development in the bug takes about eight days altogether, after which time the bugs are infective. There are thus three prin- cipal phases in the develop- ^^^x / ment of T. criizi in Triatoma (If iP ^ megista : (i) A multiplicative yS^^^..^----.,^^^^^ phase {Leishmania-Vike) in the x_.>'^''* stomach of the bug, (2) a . YiG.zd— Trypanosoma cmzi. Forms found .,,.,.,, .... , in the salivary glands of Triatoma megista. cnthidial phase, which is also (^fter Chagas.) multiplicative, in the hind-gut, and (3) a trypanosome phase, which is '^ propagative," and apparently passes through the wall of the alimentary canal into the body cavity and so into the salivary glands. Brumpt found that 7. cnizi could live in Ciniex lectnlarius, C. boueti, and Ornithodorus moiihata. The Cimex faeces may be infective. Blacklock found multiplication of the parasite in C. ledularius. Culture. — The trypanosome can be cultivated on Novy-MacNeal's blood agar, and the cultural forms resemble those described in the bug. Possible Reservoir. — Chagas thinks that probably the armadillo or " tatu " (Dasyptis novemcinctus) may be the reservoir of T. cruzi. He also thinks that Triatoma geniadata is a transmitter ; it lives in the burrows of the armadillo. Other carriers may be Triatoma infestans and T. sordid a. Clinical Features. — The trypanosomiasis of Brazil, produced by T. cruzi and spread by Triatoma spp. has received various names, such as oppilagao, canguary, parasitic thyroiditis, and coreotrypanosis. It is also known as the human trypanosomiasis of Brazil, South American trypanosomiasis, and Chagas' disease. Chagas^ reports two principal forms — acute and chronic. The acute infection is rare, and is characterized by increase in the volume of the thyroid gland, pyrexia, a sensation of crackling in the skin, enlarged lymphatic glands in the neck, axilla, etc., while the liver and spleen are increased in volume. Sclerosis of the thyroid gland is found at autopsy and fatty degeneration of the liver. During an attack of fever, trypanosomes are found in the blood. The acute form was only observed in children. In the chronic form Chagas reports several varieties : (a) A pseudo-myxcedematous form, occurring in most cases, especially up to the age of 15. There is hypertrophy of the thyroid gland or at least signs of hypothyroidism, general hypertrophy of glands, ^ Brazil Medico, Nov. 15, 1910. Longer account in Mem. Inst. Oswaldo Cruz, iii, pp. 219-275. See Sleep. Sick. Bull.y Nos. 35 and 40. 88 THE ANIMAL PARASITES OF MAN disturbance of heart rhythm, and nervous symptoms. (6) The myxoe- dematous form is characterized by similar symptoms, especially by considerable swelling of the thyroid body, and myxoedema of the subcutaneous cellular tissue ; sometimes there is a true pachydermic cachexia, (c) In the nervous form there are motor disturbances, aphasia, disturbances of intelligence or signs of infantilism, athetosis of the extremities and idiocy. There are also paralytic symptoms of bulbar origin, disturbances of mastication, phonation and deglutition, and in some cases convulsive attacks, (d) The cardiac form, charac- terized by disturbance of the heart rhythm. In all these forms the parasite is found at autopsy in the nervous substance, brain, bulb and heart. Vianna (1911)^ has studied the histopathology of the disease. Some of the chief points are : in the heart muscle destruction of the sarcoplasm, followed by interstitial myocarditis; in the central nervous system invasion of the neuroglia cells and inflammatory reaction ; in the suprarenal capsule invasion of medulla or cortex ; inflammatory reaction can also be seen in the kidneys, the hypophysis and thyroid gland. Recently Chagas states" that " schizotrypanosomiasis " has been found in a child 15 to 20 days old, and that Trypanosoma criizl has also been found in a foetus — the mother being infected with the trypanosome. The trypanosomiasis can, then, be transmitted hereditarily. Trypanosoma lewisi, Kent, 1881. The trypanosome has a nucleus somewhat displaced anteriorly, about one-third of the way from the anterior (flagellar) end of the body, a relatively straight edge to the undulating membrane, and a rod-shaped blepharoplast (fig. 37, A). It averages about 25 jju long and i'5 fju broad. Much attention has been devoted in recent years to the elucida- tion of the life history of the rat parasite, Trypanosoma lewisi. It is usually non-pathogenic to its host. It has been shown that the trypanosome can be transmitted from rat to rat by the rat-flea, Ceratophyllns fasciatus, and by Ctenocephalus canis (the so-called dog- flea). (See also p. 92). The flagellate may also persist, but doubtfully develop, in the rat-louse, Ha^matopinns spinnlosns. These researches may now be summarized. Life Cycle in the Vertebrate Host. — After infection of a rat, the trypanosomes usually appear in the animal's blood in five to seven days. This incubation period applies either to a natural or an artificial infection. The trypanosomes first observed in the rat's blood are diverse in form (fig. 37), being small, medium and large in size. This diversity is explained by the rapid multiplication taking place. A ' Mem. Inst. Oswaldo Cruz, iii, p. 276. ^ ^^^. Med. S. Paulo (1912), xv, p. 337. TRYPANOSOMA LEWISI 89 trypanosome may divide by equal longitudinal fission (fig. 37, C, D), but more commonly multiple fission occurs (fig. 37, G, H), and is unequal. Rosette forms are produced, in which the parent form can be recognized by its long flagellum (fig. 37, H) and attached to it are Fig. 37. — Trypanosoma lewisi, from rat's blood. A, ordinary form; B, small form; C, D, stages in equal binary fission ; e, elongate form {longocaudense type), resulting from division as seen in d ; f, unequal binary fission ; G, H, multiple fission into four and eight ; I, small form ; j, binary fission of small form ; K, division rosette. X 2,coo. (After Minchin and Thomson.) daughter individuals, smaller in size, from which flagella are growing. Minchin and |. D. Thomson (1912) find that the daughter forms may be set free sometimes with a crithidia-like facies (fig. 37, I), the blepharoplast being anterior but near to the nucleus. The daughter 90 THE ANIMAL PARASITES OF MAN forms, when set free, may themselves divide by binary or muUiple fission, in the latter case forming rosettes (fig. 37, K). Rosette forms were described by Moore, Breinl and H indie in 1908. Lingard, some years ago, described as a distinct species, T. longo- caudense, certain forms wnth markedly elongate posterior ends (fig. 37, E). According to Minchin, '' these forms appear to arise by binary fission" (fig. 37, D). These long drawn-out forms ''are of constant occurrence and very numerous at a certain stage of the multiplication period." It is about the eighth or tenth day after infection that the multiplication of T. lewisi is at its maximum in the rat's blood. About the twelfth or thirteenth day the trypanosomes seen in the blood appear uniform. According to Minchin (1912) ^ the rat ''gets rid of its infection entirely sooner or later, without having suffered, apparently, any marked inconvenience from it, and is then immune against a fresh infection with this species of trypano- some." There is, then, a cycle of development in the vertebrate host. Minchin notes that the records of the pathogenicity of T. leivisi in rats, causing their death, need further investigation. T. lewisi inoculated into dormice {Myoxus iiitela) and jerboas may become pathogenic thereto. Carini found cysts in the lungs of rats infected with T. lewisi. He thought the cysts were schizogonic stages of the trypanosome, comparable with those found in the lungs of animals sub-inoculated with T. criisi. Delanoe (1912)^ has found, however, that such cysts, containing eight vermicules, occurred in rats uninfected with T. lewisi. Delanoe concludes that the pneumocysts are independent of r. leivisi f and represent a new parasite, Pneuinocyslis carinii. The pneumocysts may be allied to the Coccidia, and must be considered when investigating the life-cycle of a trypanosome in a vertebrate host. Some of the stages of T. cruzi may possibly be of this nature. Life-cycle in the Invertebrate Host. — This occurs in fleas, and has been investigated in considerable detail by Minchin and Thomson in Ceratophyllns fasciatns, and by Noller in Ctenocephalns canis and Ctenopsylla ninscnli. When infected rat's blood is taken up by the flea, the parasites pass wMth the ingested blood direct to the mid-gut of the Siphonapteran. In the flea's stomach they multiply in a somewhat remarkable manner, namely, by penetration of the cells of the lining epithelium, and division inside the epithelial cells. Inside these lining cells the trypanosomes first grow to a large size and then form large spherical bodies, within which nuclear multiplication occurs (fig. 38, A — F). Any one of these large spherical bodies contains at first a number of nuclei, blepharoplasts and developing flagella, the original flagellum * ** Protozoa," p. 294. 2 c. R. Acad. Sci., civ, p. 658. TRYPANOSOMA LEWISI 91 still remaining attached for a time. The cytoplasm then divides into daughter trypanosomes which are contained within an envelope, formed by the periplast of the parent parasite. Inside the periplast envelope are a number of daughter trypanosomes " wriggling very actively ; the envelope becomes more and more tense, and finally bursts with explosive suddenness, setting free the flagellates, usually about eight in number, within the host-cell " (fig. 38, F). The daughter forms escaping from the host cell into the stomach lumen of the flea are fully formed, long trypanosomes. Fig. 38. — Trypanosoma lewisi. Developmental stages from stomach of rat flea. O, ordinary blood type ; A — F, stages occurring in gut-epithelium of flea, when the trypanosome becomes rounded and undergoes multiplication, forming in F eight daughter trypanosomes ; G, type of trypanosome resulting from such division which passes back to the rectum. X 2,000. (After Minchin.) The trypanosomes (fig. 38, G) pass into the flea's rectum. The next phase is a crithidial one. The parasites become pear-shaped, in which the blepharoplast (kinetic nucleus) has travelled anteriorly past the nucleus towards the fiagellum (fig. 39). The crithidial forms attach themselves to the wall of the rectum, and multiply by binary fission (fig. 39, D). A stock of parasites is thus formed which, according to Minchin and Thomson, ^^ persist for a long time in the flea — probably under favourable conditions, for the whole life of the insect " (fig. 39, A — I). From the crithidial forms of the rectum, according to Minchin, small infective trypanosomes arise by modification morphologically (fig. 39, ]— M). The flagellum grows longer and draws out more the anterior part of the body, the blepharoplast migrates posteriorly, behind the nucleus, and carries with it the flagellar origin. These trypanosomes are small, but broad and stumpy (fig. 39, N), and can 92 THE ANIMAL PARASITES OF MAN infect a rat. Minchin and Thomson formerly considered that the small, stumpy, infective trypanosomes pass forwards from the rectum into the stomach, and " appear to be regurgitated into the rat's blood when the flea feeds." However, the small infective trypanosomes were previously described by Swellengrebel and Strickland.^ They may be found in the flea's faeces. Noller (191 2)^ has found that the development of T. lewisi proceeds quite well in the dog flea (Cteno- cephalns canis) in Germany. Wenyon confirms this, and states that the human flea, Piilex irritans, and the Indian rat-flea, Xenopsylla cheopis, are also able to serve as true hosts for T. lewisi. Noller stated that rats were not infected with T. lewisi by infective fleas biting them, but by the rats licking up the faeces passed by the ^ Fig. 39. — Trypanosoma lewisi. Developmental stages from rectum of rat-flea. A, early rectal form; C, D, division of crithidial foim; E, group of crithidial forms; F — I, crithidial forms without free flagella, some becoming rounded ; J — M, transitional forms to tiypanosome type seen in N, which represents the final foim in the flea, x 2,000. (After Minchin.) fleas while feeding. This is not in agreement with Minchin and Thomson's earlier views of regurgitation, which, apparently, they have now abandoned.^ Wenyon (1912) confirms Noller's experi- ments. He took a dog flea, containing infective trypanosomes in its faeces, and allowed it to feed on a clean rat. The faeces of the flea, passed while feeding, were carefully *^ collected on a cover glass and taken up in culture fluid with a fine glass pipette." The contents of the pipette were discharged into the mouth of a second clean rat. Injury to the rat's mouth was carefully avoided. The first rat, on which the infective flea was fed, did not become infected, while the ' Parasitology, iii, p. 360. "^ Arch. f. Frotistenkunde, xxv, p. 386. ^ Report to Advis. Comm. Trop. Dis. Research Fund for 1913, p. 7-^. TRYPANOSOMA BRUCEI 93 second rat, in whose mouth infective flea faeces were placed, became infected in six days. When infective forms of T. lewisi have been developed within the ^iit of a rat flea, they may enter and infect the vertebrate host by ^ {a) being crushed and eaten by the rodent ; (6) the rat may lick its fur on which an infected flea has just passed infective excrement ; or (c) the rat may lick, and infect with flea excrement, the wound produced by the bite of the flea. The time taken for the full development of T. lewisi in the flea is about six days. The intracellular phase is at its height about the end of the first day ; the crithidial phase, in the flea's rectum, begins during the second day; the stumpy, infective trypanosomes are developed in the rectum about the end of the fifth day. Wenyon^ writes that, ''the fleas, when once infected with T. lewisi^ remain infected for long periods, for though many small infective trypanosomes are washed out of the gut at each feed, those that remain behind multiply to re-establish the infection of the hind gut. Further, the infection is still maintained even if the flea is nourished on a human being, so that fresh human blood does not appear to be destructive to the infective forms in the flea." The best method of controlling fleas during experiments is that due to Noller. He adopted the method of showmen who exhibit performing fleas, and secure them on very fine silver wire. Of fleas fed on an infected rat only about 20 per cent, become infective. About 80 per cent, are immune. If fleas are examined twenty-four hours after feeding, trypanosomes will be found in all, so that many of the parasites are destined to degenerate. It may be of interest to note that Gonder^ (1911) has shown that a strain of T. lewisi resistant to arsenophenylglycin loses its resistance after passage through the rat-louse, Hceniatopiniis splnulosus. These experiments suggest that physiological " acquired characters " may be lost by passage through an invertebrate host. Trypanosoma brucel, Plimmer and Bradford, 1899. Trypanosoma brucel was discovered by Sir D. Bruce in 1894 in cattle in Zululand and was named T. brucel by Plimmer and Bradford in 1899 in honour of its discoverer. This trypanosome is of con- siderable economic importance, as it is responsible for the fatal tsetse fly disease, or '' nagana/' in cattle, horses and dogs. The disease is widely distributed in Africa and is transmitted from host to host by the tsetse, Glossina morsitans, and other species of Glosslna, ' Nuttall, Parasitology, v, p. 275. "^ Report to Advis. Comm. Trop. Dis. Research Fund, October, 1912, p. 91. See also /ourn. Lond. Sch. Trop. Med., ii, p. 119. 3 Centralbl.f. Bakt., Orig., Ixi, p. 102. 94 THE ANIMAL PARASITES OF MAN The virus is maintained in nature in certain big game, such as wilde- beest, bushbuck and koodoo, which thus act as Uving reservoirs of disease from which the tsetse may become infected. These reservoir hosts are not injured, apparently, by the presence of the parasites. T. hriicei is rapidly fatal to the small laboratory animals, such as rats and mice. Horses, asses and dogs practically always succumb to its attacks, while a very small number of cattle recover from '^ nagana." The disease is characterized by fever, destruction of red blood corpuscles, severe emaciation and by an infiltration of coagu- lated lymph in the subcutaneous tissue of the neck, abdomen and extremities giving a swollen appearance thereto. The natural reservoirs in which T. hriicei has been long acclimatized are unaffected by the trypanosomes, while the newer hosts, such as imported cattle in Africa, are rapidly destroyed by their action. The general morphology and life history in the vertebrate host is that of a typical trypano- some (fig. 40). Its length is from 12 /t to 35 /a, its breadth from 1*5 /a to 4/x. Multiplication by longitudinal division proceeds in the peripheral blood (fig. 26), while latent, leishmaniform Fig. ^o.— Trypanosoma bodics are produced in the internal organs. iTv'etn a„d'^;„il.t''" Bruce and colleagues' have quite recently (June, 1914) described the development of a Zululand strain of T. hrucei in G. morsitans. The tsetse flies were bred out in Nyasaland. In vertebrate blood the brncei strain was polymorphic. The development was like that found for T. gambiense in G. palpalis (fig. 30), and by Bruce and colleagues for T. rhodesiense in G. morsitans in Nyasaland. Long trypanosomes were found in the proventriculus of the tsetse. Crithidial, rounded or encysted, and immature ^' blood forms " occurred in the salivary glands; and finally infective, stumpy, ''blood forms " were differen- tiated in the salivary glands. The period of development of T. hrucei in G. morsitans takes about three weeks, and then the fly becomes infective. Bruce believes that T. rhodesiense of Nyasaland and T. hrucei of Zululand are the same, their cycles of development in G. morsitans being " marvellously alike." (But see Laveran, p. 80.) T. hrucei has been cultivated with difficulty by Novy and MacNeal, using blood agar. The best treatment for nagana is arsenic in some form. It is probable that more than one trypanosome has been con- fused under the name T. hrucei, more especially as the occurrence of many species of trypanosomes in various animals in Africa was not ' Proc. Roy. Soc.^ Bj Ixxxvii, p. 526. TRYPANOSOMA EVANSI 95 suspected until comparatively recent times. It has been shown by Stephens and Blacklock (191 3) that the original Zululand strain of T. hnicei was monomorphic, while the organism sent from Uganda, and at the time believed by Bruce to be the same as the Zululand trypanosome, has been found to be polymorphic, with morpho- logical resemblances to T.rJiodesiense. Stephens and Blacklock^ have suggested the name T. iigandce for the polymorphic trypanosome, which, however, has marked resemblances with Trypanosoma pecaudi, and they are, perhaps, identical. T. pec audi was the name given by Laveran ^ in 1907 to the causal agent of '* baleri " in equines and sheep in the French Sudan. T. pecaudi, which is dimorphic, is widely distributed in Africa. An extremely small number of both T. pecaudi and T. ugaudce have been shown to possess posterior nuclei. T. pecaudi is transmitted by various species of Glossina, and is said to develop in the gut and proboscis of the fly. On the other hand, Bruce and colleagues (1914), examining a strain sent from Zululand in 1913, state that T. hrucei is polymorphic. Bruce (1914) suggests that passage through laboratory hosts has influenced and altered the morphology of the parasite. Trypanosoma evansi, Steel, 1885. Syn. : Spirochceta evansi^ Steel, 1885; Hcematomonas evansi, Crookshank, 1886; Tf'ichomonas evansi^ Crookshank, 1886. Trypanosoma evansi, first found by Evans in 1880, in India, is the causal agent of the disease known as ** surra." The malady affects more particularly horses, mules, camels and cattle in India and neigh- bouring countries, such as Burma and Indo-China. It occurs also in Java, the Philippines, Mauritius and North Africa. Elephants may be affected. A serious outbreak among cattle in Mauritius occurred in 1902, the disease being imported into the island. The symptoms are fever, emaciation, oedema, great muscular weakness and paralysis culminating in death. T. evansi varies from 18 //, to 34 //< in length and i'5 //- to 2 //, in breadth. It has a pointed posterior extremity, and, anteriorly, there is a free portion to the flagellum (fig. 41). It is possibly mono- morphic, but a few broad forms occur. The trypanosome multiplies by longitudinal fission in the blood. Rounded leishmaniform stages occur in the spleen of the vertebrate host, which stages Walker^ (1912) considers to be phases of schizogony. The parasite is transmitted in nature by various species of Tabanus and Stomoxys, though at present little is known of the life-history within these invertebrate hosts. ' Froc. Roy. Soc.^ B, Ixxxvi, p. 187. "^ C.R. Acad. Set., cxliv, p. 243. ■•* Philippine lourn. Sc. (Sect. B), vii, p. 53. 96 THE ANIMAL PARASITES OF MAN Fig. 41. — Trypanosoma evaiisi. x 2,000. (Original. From preparation by Fan- tham.) Dogs are said to contract the disease by feeding on animals dead of surra. A variety of T. evansl is the cause of '' mbori " in dromedaries in Africa (Sahara and Sudan). Another possible variety, or closely allied form, is T. soudaiiense, the causal agent of ''el debab " in camels and horses in North Africa, especially Algeria and Egypt. An extraordinary example of the possible infection of a human being with an animal trypanosom.e is recorded in the case of Pro- fessor Lanfranchi, of the Veterinary School, Parma. The Professor became infected with trypanosomes, although only nagana and surra were maintained in his laboratory, and he himself had never visited the tropics. He suffered from irregular attacks of fever and was cedematous, but his mind remained clear. The identification of the trypanosome from Lanfranchi's blood has been a matter of great difficulty. Apparently Mesnil and Blanchard (1914)^ consider the strain found in the patient is almost indistinguish- able in its reactions from T. gainbiense, though the parasite is mono- morphic. Lanfranchi considers that he was infected with T. evansi. Trypanosonna equinum, Voges, 1901. Syn. : Trypafwsoina ehnassiaiH^ Lignieres. Trypanosoma eqiiinum was found by Elmassian to be the cause of the fatal disease, '' mal de caderas," of horses and dogs, in South America (Paraguay, Argen- tine, Bolivia). The name refers to the fact that in the disease, as in other trypanosomiases, the hind quarters become paralysed. Cattle are refractory to inoculation. T. equinum is about 22 yu, to 24 /x long and about 1*5 fi broad (fig. 42). Although this trypanosome is very active, yet it is charac- terized by the blepharoplast (kinetic nucleus) being very minute or even absent, as the granule sometimes seen may be the basal granule of the fiagellum. Fig. 42. — Trypanosoma The mode of transmission of T. equinum ^qt^innm. x 2,000. (After , , -ji 1 1 J i • i TVT- Laveran and Mesnil.) is not known with absolute certainty. Migone has shown that the parasite causes a fatal disease in the large South * Bull. Soc. Path. Exot., vii, p. 196. TRYPANOSOMA EQUIPERDUM 97 American rodent, the capybara {Hydrochcenis capybara). This animal appears to be a reservoir of the parasite. Dogs may become infected by eating diseased capybaras, and it is suggested that the infection is spread from the dogs to horses by the agency of fleas. Some authorities consider that T. equimtin may be spread by various Tabanidce and by Stomoxys. Neiva (1913)^ doubts all these modes of transmission in Brazil, and suggests Chrysops or Trlatoma as vectors. Trypanosoma equiperdum, Doflein, 1901. Syn. : Trypanosoma rougeti, Laveran and Mesnil. The malady of horses known as " dourine " or '' mal du coit " is due to a trypanosome, T, equiperdtun, discovered by Rouget in 1894. "Dourine" — also known as ^'stallion disease" or '^covering disease" — is found among horses and asses in Europe, India, North Africa and North America. The trypanosome is transmitted by coitus, and so far as is known not by insect agency. The progress of the disease may be con- sidered under three periods. The period of oedema, when signs of oedema of the genitalia are seen. The oedema is generally painless and non-inflammatory. This period lasts about a month. It is succeeded by the period of eruption, which sets in about two months after infection. Circular oedematous areas (*' plaques "), often about the size of a two- fig. 43. — Trypamsowa shilling piece, appear under the skin of the sides equiperduvi. x 2oooapprox- , , . \ '^^ 1 ,, imately. (Original. From and hmd quarters, and also, at times, under the preparation by Famham.) skin of the neck, thighs and shoulders. The eruption is variable, but usually lasts about a week and leaves the animal in an enfeebled condition. Gland enlargement and swelling of the joints and synovia also may occur. The third period of the disease is described as that of ancemia and paralysis. The animal becomes very anaemic, emaciation is marked, superficial non-healing abscesses often form, and conjunctivitis and ulcerative keratitis can occur. Paralysis ensues, and in from two to eighteen months the animal dies. In the acute form of the disease the animal may die after the first period from acute paralysis. It is difficult to find the trypanosomes in naturally infected animals, and they are best obtained from the plaques of the eruption. Apparently the parasite occurs more in the lymph than in the blood. ' Brazil Medico^ xxvii, p. 366. 98 THE ANIMAL PARASITES OF MAN Ruminants are said to be refractory to this trypanosome. T. equiperdum is about 25 yu, to 28 /a in length on an average, but varies from 16 /x to 35 fjb. Its cytoplasm is relatively clear, and does not show chromatic granules (fig. 43). It is stated to be monomorphic. It has been shown recently by Blacklock and Yorke {1913)^ that there is another trypanosome giving rise to dourine in horses. This trypanosome is dimorphic (resembling T. pecandi and 7. iigandce), and is named T. eqni. Previously T. equiperdum and T. eqtii had been confused. Uhlenhuth, Hiibner and Worthe have demonstrated the presence of endotoxins in T. equiperdum. These endotoxins may be set free by trypanolysis. Trypanosoma theiteri, Bruce, 1902. This parasite, 60 /x to 70 //- long, and 4 /i to 5 /a broad, is dis- tinguished for its large size, though it is not so large as 7. ingens from Uganda oxen, whose length may be 72 //, to 122 fi, and breadth 7 /A to 10 fjL. The posterior end of T. theileri is drawn out. Small forms of the flag- ellate are known, 25 /x, to 53 /x in length. Probably other forms of the parasite have the nucleus posterior, and these Fig. 44. - Trypanosoma theileri. x 2,000. fl o rr^l 1 a f pc wpi-p f nrm pri v c;pn (After Laveran and Mesnil.) tlagcllates wei c toi meriy Sep- arated as r. transvaaliense (Laveran, 1902). Myoneme fibrils may be seen on its body. The pathogenicity of this organism is doubtful, it was formerly thought to be the causal agent of ^' gall-sickness " in cattle in South Africa. T. theileri also occurs in Togoland, German East Africa, and Trans- caucasia. Allied or identical parasites occur in cattle in India. Trypanosoma theileri, specific to cattle, is perhaps transmitted by the fly Hippobosca rufipes in South Africa, Trypanosoma hippicum, Darling, 1910. Trypanosoma hippicum causes the disease of mules known as '*murrina."2 It was found in mules imported to Panama from the United States. It can live in other equines. The parasite varies from 18 /x to 28 /A in length, and is from 1*5 ytt to 3 //. broad. Its undu- lating membrane is little folded. The trypanosome has a noticeable blepharoplast. It can penetrate mucous membranes, and it is thought that the trypanosome may be transmitted during coitus. It may also ^ Proc. Roy. Soc, B, Ixxxvii, p. 89. ^ Bull. Soc. Path. Exot., iii, p. 381. TRYPANOSOMA VIVAX 99 be spread meclianically by species of Miisca, Sarcophaga and Comp- somyia, sucking the wounds of infected animals and carrying over the trypanosomes to wounds on healthy ones. Endotrypanum schaudinni, Mesnil and Brimont, 1908. This organism was discovered in the blood of a sloth {Choloepus didactyliis), m South America (French Guiana).^ It possesses special interest, in that the best known form of the organism is endoglobular, inhabiting the erythrocytes of the sloth. A free trypanosome in the same animal was considered to be different from the endoglobular form, which w^as somewhat like a peg-top, and possessed a short fiagellum. Darling^ (November, 1914) has seen the organism in Panama. He describes free crithidial forms in shed blood, but not in the blood-stream of the sloth. Trypanosoma boylei, Lafont, 1912. This is a parasite of the Reduviid bug, Conorhiniis rubrofasciattis. The insect attacks man in Mauritius, Reunion and other places. Lafont infected rats and mice by intraperitoneal injection with the gut-contents of infected bugs. Trypanosomes appeared in the mice. Other flagellate types were assumed by the parasites in the bug. MoNOMORPHic Trypanosomes. A number of trypanosomes, characterized by relative uniformity in size and structure, may be considered under this heading. They occur in cattle, sheep, goats and horses in Africa, especially West Africa. Morphologically, they are characterized by the posterior (aflagellar) part of the body being swollen, while the anterior part narrows. The nucleus is central and situated at the commencement of the narrowing of the body. The blepharoplast is almost terminal, the undulating membrane is narrow and not markedly folded, so that the flagellar border lies close to or along the body. The fiagellum may or may not possess a free portion. Some recent workers have considered that T. briicel (Zululand strain) and T. evansl are also monomorphic, but they do not exhibit the general characteristics outlined above. T. hriicei and T. evansi have already been considered separately. The monomorphic trypanosomes, as defined above, include : — Trypanosoma vivax, Ziemann, 1905. This trypanosome^ occurs in cattle, sheep and goats, and was first found in the Cameroons. It is fatal to cattle. Equines are also affected. Antelopes are the possible reservoirs of the trypanosome. ' C. R. Soc. Biol.y Ixv, p. 581. 2 Joitrn. Ale.i. Research^ xxxi, p. 195. ^ See Bruce and colleagues (1910), Proc. Roy. Soc, B, Ixxxiii, p. 15. TOO THE ANIMAL PARASITES OF MAN It is probably transmitted by Glossina palpalis and other tsetse flies. Its movement is very active. It possesses a free flagellum (fig. 45) and it averages 23//, to 24 yit in length. T. cazalhoni (Laveran, 1906) — the causal agent of ''souma" in bovines and equines in the French Sudan — is probablv synonymous with T. vlvax. Trypanosoma caprae (Kleine, 1910) is allied, but is somewhat broader and more massive. It was found in goats in Tanganyika. Trypanosoma congolense, Bi oden, 1904. Probable synonyms. — Trypcmosoma dimorphon, Laveran and Mesnil, 1904 ; Trypa?iosofna namun, Laveran, 1905 ; Trypanosoma pecorum^ Bruce, 1910 ; Try- panosoma confusum, Montgomery, 1909. This trypanosome causes disease among horses {e.g., Gambia horse sickness), cattle, sheep, goats, pigs, and dogs. It is widely dis- tributed in Central Africa {e.g., Gambia, Congo, Uganda, Nyasaland), the strain probably being maintained naturally in big game. It is trans- mitted by various Glossince, and perhaps by Tabanus and Stomoxys. It is said to develop in the gut and proboscis of Glossina palpalis and G. morsitans. The trypanosome averages 13 /x to 14 yLt in length and has no free flagellum (fig. 46). It is about 2 fj, broad. Formerly T. tiaiium and T. pecorumwere said to differ in their pathogenicity, the former being said not to infect the smaller laboratory animals. Yorke and Blacklock (1913), however, consider that the virulence varies and that these trypanosomes are probably the same. Fig. 45. — Trypanosoma vivax. x2,oco. (Original. From preparation by Fan- tham.) Fig. 46. — Trypanosoma congolense. x 2,000. (Orig- inal. From preparation by Fantham.) Fig. 47. — Trypanosoma uni forme, x 2,000, (Orig- inab From preparation by Fantham.) The T. diinorphon originally obtained by Button and Todd (1903) in Gambian horse sickness has been shown to be a mixture of T. vivax and T, congolense. Trypanosoma simiae (7\ ignotum) is like T. congolense. It averages I7'5 /A long. It is virulent to monkeys and pigs. TRYPANOSOMA UNI FORME lOI Trypanosoma unlforme, Bruce, 1910. This trypanosome was found in oxen in Uganda/ It can be in- oculated to oxen, goats and sheep, but is refractory to dogs, rats and guinea-pigs. It has been found in antelopes. It resembles T. vivax, but is smaller (fig. 47), averaging 16 fju in length. A free flagellum is present. It is transmitted by Glossincv. Many other trypanosomes occur in mam- mals, while birds, reptiles, amphibia (fig. 48) and fish also harbour them. The discussion of these forms does not come within the scope of the present work. They are dealt with in Laveran and Mesnil's ''Trypanosomes et Trypanosomiases," 2nd edit., 19 12. General Note on Development of Trypanosomes in GLOSSINA. Yig.^^.— Trypanosoma rotatoriufn, from blood of Before concluding the account of trypano- a frog, x 1,400. (After .. , r ■ 1 11 1 ii J Laveran and Mesnll.) somes, it may be of mterest to remark that several African trypanosomes develop in various species of Glosshia, and are found in different parts of the alimentary tract and in the proboscis. Thus {a) T. vivax, T. uniforme and T. caprce develop in the fly's proboscis (labial cavity and hypopharynx) only ; {b) T. congolense, T, simicB and T. pecaudi develop first in the gut of the fly and then pass forward to its proboscis ; and (c) T. gambiense and T. rhodesiense develop first in the gut and later invade the salivary glands of the tsetse. The proboscis or the salivary glands in such cases are termed by Duke^ the anterior station of the trypanosome, wherein it completes its development. Adaptation of Trypanosomes. These flagellates may exhibit power of adaptation to changes of environment, such as those due to the administration of drugs, change of host, etc. A few examples of such mutations may be briefly considered : — (i) Blepharoplastless Trypanosomes. — T. briicei may become resistant to pyronin and oxazine. Accompanying this drug resistance is a change in morphology, namely, the loss of the blepharoplast (Werbitzki).^ A race or strain of blepharoplastless trypanosomes may be thus produced which retains its characteristic feature after as many as 130 passages (Laveran).^ Oxazine is the more powerful drug, and it acts directly on the blepharoplast. (Compare the natural blepha- roplastless character of T. equintun.) ' Froc. Roy. Soc, B; Ixxxiii, p. 176. 2 Repts. Sleeping Sickness Commission Roy. Soc. (1913), xiii, p. 82. 3 Centralbl. f. Bakt. (1910), Orig., liii, p. 303. * Bull. Soc. Path. Exot., iv, p. 233. 102 THE ANIMAL PARASITES OF MAN (2) Reference has been made on p. 93 to the experiments of Gonder, who showed that a strain of T. lewisi rendered resistant to arsenophenylglycin lost its resistance after passage through the rat louse. This is in marked contrast with the retention of drug resistance during passage b}^ inoculation from rat to rat. (3) T. lewisi from the blood of a rat when transferred to a snake seems largely to disappear, as very few flagellates are seen. When blood from the snake is inoculated into a clean rat, then trypano- somes reappear in the rat, but they are not all like those originally inoculated. It seems certain that, in such a case, changes in form and virulence of the trypanosome have occurred. Similar experiments were made with T. hriicei from rats to adders and other animals and back to rats. Changes in the form and virulence of T. hrucei occurred. These interesting experiments were performed by Wendelstadt and Fellmer.' Genus. Herpetomonas, Saville K'^nt, 1881. Herpetomonas is a generic name for certain flagellates possessing a vermiform or snake-like body, a nucleus placed approximately cen- trally, and a blepharoplast (kinetic nucleus) near the flagellar end. There is no undulating membrane (fig. 49, a). The organisms in- cluded in this genus certainly possess one flagellum, while according to Prowazek (1904) Herpetomonas miiscce-domesticcey the type species, possesses two flagella united by a membrane. Patton,^ Porter ^ and others affirm, however, that the biflagellate character of H. niiiscce- domesticcB (from the gut of the house-fly) is merely due to precocious division. The matter is further complicated by the generic name Leptomonas, given by Kent in 1881, to an uniflagellate organism found by Butschli in the intestine of the Nematode worm, Trilohus gracilis. This parasite, Leptomonas hiltschlii^ has not yet been completely studied. Until these controversial points relating to the identity or separation of Herpetomonas and Leptomonas have been satisfactorily settled, we may retain the better known name Herpetomonas for such uniflagellate, vermiform organisms. However, the name Leptomonas, having been used by Kent two pages earlier in his book {^' Manual of the Infusoria ") than Herpetomonas, would have priority if the two generic names were ultimately shown to be synonymous. A full discussion of these interesting and important flagellates hardly comes within the purview of the present work ; brief mention can only be given here to certain species. The Herpetomonads occur principally in the digestive tracts of insects, such as Diptera and Hemiptera. They are also known in the ' Zeitschr.f. Immunitatsforschung^ iv, p. 422 (1909), and v, p. 337 (1910). * Arch.f. Protist.^ xiii, p. i. ^ Parasitology^ ii, p. 367. HERPETOMONAS 103 guts of fleas and lice, but are not confined to blood-sucking insects. One example, H. deuocephali (Fantham, 191 2) ' occurs in the digestive tracts of dog fleas, Ctenocephalns canis, in England, France, Germany, Italy, India, Tunis, etc. It is a natural flagellate of the flea, and might easily be confused with stages of blood parasites in the gut of the dog flea. Dog fleas are stated by Basile to transmit canine kala-azar, which is believed to be the same as human infantile kala-azar. Confusion is further likely to arise since herpetomonads pass through pre-flagellate, flagellate and post-flagellate or encysted stages; pre- and post-flagellate stages being oval or rounded and Leishmania-like. The post-flagellate stages are shed in the faeces, and are the cross-infective stages by means of which new hosts are infected by the mouth. The possible presence of such natural flagellates must alw^ays be considered when experimenting with fleas, lice, mosquitoes, etc., as possible vectors of patho- genic flagellates like Leish- mania and Trypanosoma, H, pedicidi (Fantham, 191 2) occurs in human body lice.^ See further remarks on pp. 107, 112. Laveran and Franchini (1913-14)^ have recently succeeded in inoculating Herpetoinonas ctenocepJiali, from the gut of the dog flea, intraperitoneally into white mice, and produc- ing an experimental leish- maniasis in the mice. A dog was also infected. They have also succeeded in infecting mice with H. pattoni — a natural flagellate of the rat flea — by mixing in- fected rat fleas with the food of the mice, and by causing them to ingest infected faeces of rat fleas. Further, they have shown that infection with the herpetomonas occurs naturally by this method, that is, by the rodents eating the fleas and not by the insects inocu- lating the flagellates into the vertebrates when sucking blood. These experiments shed an interesting light on the probable origin of Leishmania and its cultural herpetomonad stage, which were very probably once parasitic flagellates in the gut of an insect. • Bull. Path. Exot., vi, p. 254. ^ Froc. Roy. Soc, B, Ixxxiv, p. 505. ^ C. R. Acad. Set., clvii, pp. 423, 744. Ibid., clviii, pp. 450, 770. Bull. Soc. Paih. Exot., vii, 605. Fig. 49. — a, Herpetomonas ; b, Crithidia Cy Trypanosoma. (After Porter.) 104 THE ANIMAL PARASITES OF MAN Fantham and Porter^ (1914-15) have shown that young mice may be inoculated or fed with Herpetomonas jaculum, from the gut of the Hemipteran, Nepa ciiierea (the so-called "water-scorpion"), with fatal results. The pathogenic effects are like those of kala-azar. They also showed that the post-flagellate stages of tlie herpetomonads seemed most capable of developing in the vertebrate. A herpetomonad, N. davidi^ has been found in the latex of species of the plant- genus Euphorbia in Mauritius, India, Portugal, etc. It is apparently transmitted to the plants by Hemiptera. The plants sometimes suffer from " flagellosis." Franchini (1913) ^ has described a new parasite, Hccnwcystozoon brasiliense, from the blood of a man who had lived in Brazil for many years. It possesses flagellate and rounded stages, and is closely allied to the herpetomonads. Genus. Crithidia, Leger, 1302, emend. Patton, 1908. Crithidia is the generic name of vermiform flagellates with a central nucleus, a blepharoplast or kinetic nucleus in the neighbour- hood of the principal nucleus, and a rudimentary undulating mem- brane bordered by a flagellum arising from a basal granule, which is the centrosome of the kinetic nucleus (fig. 49^). The anterior or flagellar end of the body is attenuated and fades off as the undulating membrane. Crithidia fasciculata, the type species, was found by Leger in the alimentary canal of Anopheles macidipennis. Crithidia occur in bugs, flies, fleas,' and ticks. Some of them are found in the body-fluid of the invertebrate host as well as in the gut. Others may be restricted to the body cavity or intestine respectively. C. melopliagia from the sheep-ked, Melophagns ovintis, and C. hyaloinmce from the haemocoelic fluid of the tick, Hyalotmna cegyptiutUj pass into the ovaries and eggs of their hosts, and the young keds or ticks are born infected. C. fasciciilata has been shown by Laveran and Franchini to be inoculable into white mice, producing a sort of experimental leish- maniasis therein. In one case cutaneous lesions were produced like those of Oriental sore. Crithidia are natural flagellates of Arthropoda, with their own pre-flagellate, flagellate and post- flagellate stages, and must not be confused with transitory crithidial stages of trypanosomes. Genus. Leishmania, Ross, 1903. With an oval body containing nucleus and blepharoplast (kinetic nu- cleus) but no flagellum. An intracellular parasite in the vertebrate host. Included in the genus Leishmania are three species, namely : — ' Froc. Camb. Philosoph. Soc, xviii, p. 39. "■ BtilL Soc. Path. Exot., vi, pp. 156, 333, 377. ^ See Porter, Parasitology, iv, p. 237. LEISHMANIA DONOVANI 105 (i) Leishmania donovaiii, Laveran and Mesnil, 1903, the parasite of Indian kala-azar, a generalized systemic disease, usually fatal, occurring in subjects of all ages. (2) Leislunania tropica, Wright, 1903, the parasite of Delhi boil, Oriental sore, Aleppo button — a localized, cutaneous disease, usually benign. (3) Leishmania infantujii, Nicolle, 1908, the parasite of infantile kala-azar, occurring in children (and a few adults) around the shores of the Mediterranean. The disease is perhaps a form of Indian kala-azar, and the parasite is probably identical with L. donovani. These diseases may be termed collectively leishmaniases. The morphology of the various species is practically identical. Leishmania donovani, Laveran and Mesnil, 1903. Syn. : Piroplasjna donovam\ Laveran and Mesnil. The parasite of Indian kala-azar was demonstrated in 1900 by Leishman from a post-mortem examination of a case of ^* Dum-Dum fever," but details were not published till May, 1903. In July, 1903, Donovan found similar bodies from cases in Madras. Rogers succeeded in cultivating the parasite in July, 1904.^ The original centre of the disease was probably Assam ; it occurs also in Madras, Ceylon, Burma, Indo-China, China and Syria. A variety of this leish- maniasis is found in the Sudan. The patient becomes emaciated, with a greatly enlarged spleen. There is anaemia and leucopenia. The parasite, commonly known as the Leishman-Donovan body^ is intracellular (fig. 50, 2, 5). It is found in the endothelial cells of the capillaries of the liver, spleen, bone-marrow, lymphatic glands and intestinal mucosa, and in the macrophages of the spleen and bone- marrow. Some host cells may contain many parasites. It is rather rare in the circulating blood, but may be found in the blood from the femoral, portal and hepatic veins. It does not occur in the red blood corpuscles as was formerly thought. The parasites liberated from the endothelial cells are taken up by the mononuclear and polymorphonuclear leucocytes. The Leishman-Donovan body is the resting stage of a flagellate. As found in man it is a small, oval organism, about 2*5 //, to 3*5//' in length by 2 yu, in breadth, and con- taining two chromatinic bodies, corresponding to the nucleus and kinetic nucleus (blepharoplast) of a flagellate. The latter element is the smaller and more deeply staining, and is usually placed at the periphery, transversely to the longer axis of the oval organism. ^ The literature up to 191 2, on kala-azar and other leishmaniases is reviewed in the Kala-azar Bulletin. Afterwards in the Tropical Diseases Bulletin. io6 THE ANIMAL PARASITES OF MAN There is sometimes a very short, slightly curved filament to be seen,, which may be a rhizoplast. Multiplication takes place by binary or multiple fission. The presence of the parasite used to be demonstrated by splenic or hepatic prmcture ; nowadays it can be demonstrated in. peripheral blood, e.g., of the finger, or by culture of infected blood. Fig. 50. — Leishmania donovani. /, 't lee lorms, each with nucleus and rod -shaped Mepharo- plast (after Christophers) ; 2, endothelial cell and leucocytes containing parasites (after Christophers) ; j, capillary in the liver showing endothelial cells containing parasites (after Christophers) ; ^, two parasites escaping from a leucocyte in the alimentary canal of the bug (after Palton) ; 5, further development in bug (after Patton) ; 6, young flagellate forms in bug (after Patton) ; 7-/7, culture forms (after Leishman) ; 7, 8^ 9, show development of flagellum. L. donovani can be cultivated in citrated splenic blood, under aerobic conditions, at 22° to 25° C. This was first accomplished by Rogers (1904). It is not so easily culturable as L. infantum on the Novy-MacNeal-Nicolle medium.^ L. donovani is inoculable with ' For the composition of this medium, see Appendix. LEISHMANIA TROPICA 107 some difficulty into experimental animals — in India, white rats, white mice, dogs and monkeys (Macaais spp.), have been inoculated. The Sudan variety, somewhat less virulent, is inoculable to monkeys. Row also produced a local lesion in Macaais slnicus by subcutaneous inocu- lation of L. dofiovani. Parasites taken from such a local lesion were found to be capable of producing a generalised infection in Macacus slnicus and white mice. In cultures the various species of Leishmania all grow into herpetomonad, uniflagellate organisms (fig. 50, 10), about 12 fi io 20 /j, in body length. On this account Rogers^ and Patton place the Leishman-Donovan body within the genus Herpetoinonas. The method of culture may be used in diagnosing leishmaniases. Kala-azar is very probably an insect- borne disease. Patton^ sus- pects the bed-bug to be the transmitter and finds (fig. 50, 4-6) that the Leishman-Donovan body can develop mto the flagellate stage in the digestive tract of the bed-bug. Feeding experiments are unsatisfactory, since there are very few cases in which the parasites occur in sufficient numbers in the peripheral blood to make the infection of the insect possible, or at any rate easy. In examining the alimentary tracts of insects for possible flagellate stages of Leishmania, it must be remem- bered that in many insects natural flagellate parasites, belonging to the genus Herpetontonas, m3.y occur therein ; such natural insect flagellates may be harmless, and have no connection with the life-cycle of L. donovani. Natural herpetomonads are known to occur in the alimentary tracts of flies, mosquitoes, sand-flies, fleas and lice, but not in bed-bugs. Further, if such flagellates are able to be inoculated into and live within vertebrate hosts, producing symptoms like those of leishmaniasis, the origin of kala-azar is indicated (see pp. 104, 112). Leishmania tropica, Wright, 1903. Syn. : Helcosoma tropicum, Wright, 1903 ; Z. wrighti, Nicolle, 1908; Ovoplasma orientale^ Marzinowsky and Bogrow. It is believed by some that the parasite was first described by Cunningham in 1885, and studied by Firth in 1891, being called by him Sporozoon furuncidosum. If these earlier studies were of the parasite, then its correct name is L. furunculosa^ Firth, 1891. The benign disease produced by this parasite has received many names, among the best known being Oriental sore. Tropical sore, Delhi boil and Aleppo button. These names, however, are not happy ones, as cutaneous leishmaniasis {e.g., on the ear) is now known to occur in the New World, for example in Mexico, Venezuela, Brazil and neighbouring States. However, it may be necessary to subdivide cutaneous leishmaniases later. ' Proc. Roy. Soc, B, Ixxvii, p. 284. 2 Sci. Mem, Govt. India, Nos. 27, 31 (1907-08). I08 THE ANIMAL PARASITES OF MAN In the Old World the disease occurs in India, Persia, Arabia and Transcaucasia. It is also known in Algeria, Northern Nigeria, Egypt, Sudan, Crete, Calabria, Sicily and Greece. The boils often occur on the face, and before ulceration the parasites may be found in the cells at the margin and floor of the " button." In searching for parasites the scab should be removed and scrapings made from the floor and edges. Where lesions occur atrophy of the epidermis takes place, and infiltration of mononuclear cells (e.g., plasma cells, lymphoid and endothelial cells) follows. The parasites are intracellular, being found inside mononuclear cells. In non-ulcerating sores, Cardamitis found some free parasites. Non- ulcerating forms are said to occur in the Sudan. In the Old World the sores are often limited to exposed surfaces of the body. Infection of mucous membranes (such as the lip, palate, buccal and nasal membranes) may occur, especially in South America, and are often known there as ^^ Espundia." Christopherson (1914) has recorded a case in Khartoum. Leishmania tropica is equally well cultivated on Novy-MacNeal- Nicolle medium or on citrated blood. The usual temperature for cultivation is 22° to 28° C, though Marzinowski claims to have cultivated the parasite at 37° C. L. tropica can be inoculated into monkeys and dogs, with the production of local lesions. Material from a human sore or flagellates from a culture may be thus successfully inoculated. Also infected material may be rubbed directly into a scarified surface. The incubation period is long, extending over several months. The duration of the disease may be from twelve to eighteen months. Recovery from one attack of tropical sore confers immunity, and the Jews in Bagdad inoculate their children with the disease on a part of the body which will be covered, and so secure immunity in adult life. The mode of transmission of L. tropica is unknown. Wenyon (1911)^ has found that the parasite develops into the flagellate stage in the digestive tract of Stegomyia fasciata in Bagdad. Patton (191 2)' has found similar development in the bed-bug in Cambay. The house-fly, Phlehotonins and Simulium have been suspected as trans- mitters in different parts of the world. An interesting announcement has been made recently (May, 1913), that Neligan has found that L. tropica occurs in dogs in Teheran, Persia, producing ulcers on the dogs' faces [cf. natural occurrence of L. infantiim in dogs — see p. no). Yakimoff and Schokhor (1914),^ have found the disease in dogs in Tashkent. Gonder* (191 3) has performed some interesting experiments ' Parasitology, iv, p. 387. ' Sci. Mem. Govt. India, No. 50. 3 Bull. Sue. Path. Exot., vii, p. i86-. * Arch. f. Schifs- u. Prop.. Hyg.^ xvii, p. 397. LEISHMANIA INFANTUM 109 showing the relation of infantile kala-azar to Oriental sore. Gonder infected mice with L. infantum and with L. tropica. He used culture material and injected intraperitoneally or intravenously. In each a general infection resulted, with enlargement of the liver and spleen. Later, however, mice injected with Oriental sore (North African variety) developed peripheral lesions on the feet, tail and head, and the lesions contained Leishuiania. No such peripheral lesions developed in the case of the mice infected with the kala-azar virus. Gonder suggested that Oriental sore, like kala-azar, is really a general infection overlooked in its earlier stages, and that it is in the later stages that peripheral lesions on the skin are developed. Row (1914)^ also obtained a general infection in a mouse by the injection of cultures of L. tropica from Oriental sore of Cambay. Leishmania Infantum, Nicolle, 1908.^ Infantile splenic anaemia has been long known in Italy. It also occurs in Algeria, Tunis, Tripoli, Syria, Greece, Turkey, Crete, Sicily, Malta,^ Spain and Portugal. This leishmaniasis is, then, distributed along the Mediterranean littoral; also in Russia. Cathoire (1904) in Tunis and Pianese (1905) in Italy were among the first to see the parasite. Nicolle then found the parasite in patients in Tunis, and further found spontaneous infection in dogs. The patients are usually children between the ages of 2 and 5 years. There are a few cases known in which the infantile type of leishmaniasis occurred in youths and adults of the ages of 17 to 19, while one patient in Calabria was 38 years old. The symptoms are like those of Indian kala-azar. Several Italian investigators and others consider that L. infantum is the same as L. donovani, and that the latter name should be used for the parasite of Mediterranean leishmaniasis. This view, as to the identity of L. donovani and L. infantum, seems coming into general favour. There are, however, differences between the Indian and infantile kala-azars, in addition to the ages of the patients affected, thus : {a) As regards cultures, it is found that L. infantum is readily grown on the Novy-MacNeal-Nicolle C' N.N.N.") medium (saline blood- agar), and that sub-cultures are easily obtained ; in citrated blood L. infantum grows with difficulty. The reverse is the case with regard to culture media for L. donovani, which grows with difficulty on the N.N.N, medium, but relatively easily in citrated splenic blood. (6) Considering inoculability into experimental animals, it is found that L. donovani is inoculated generally with some difficulty into white rats, * Bull. Soc. Path. Exot.^ vii, p. 272. ^ Arch. Inst. Pasteur Tunis, i, p. 26. ^ See Wenyon (1914), Trans. Soc. Trop. Med. attd Hyg., vii, p. 97; also Critien (191 1), Annals Trop. Med. and Parasitol., v, p. 37. no THE ANIMAL PARASITES OF MAN white mice and monkeys, and with greater difficulty into dogs, while L. infaninm can be inoculated into several experimental animals, especially into dogs and monkeys, with ease, (c) At present L. donovani is not known to occur spontaneously in animals, jDut L. infantiun is found naturally in dogs in the Mediterranean region, and the disease in dogs is often referred to as canine kala-azar. Kittens have occasionally been found infected. However, these differences must not be empha- sized too much. The material for cultivation is obtained from punctures of spleen, liver or bone-marrow of cases infected with L. infantum. It is not always easy, however, to infect from cultures, as the cultural flagellates inoculated into the body are often phagocytosed. Similarly, the material for animal inoculation is obtained from emulsions of infected spleen, liver or bone-marrow. Dogs and monkeys are easily inoculated with such material ; Nicolle inoculates into the liver or the peritoneal cavity. Mice, white rats, guinea-pigs and rabbits only show slight infections after such inoculations. Dogs infected experimentally with infantile leishmaniasis may show either acute or chronic symptoms. The acute course occurs more often in young dogs, and is usually fatal in three to five months. The chronic course is found more commonly in older dogs, and may last seventeen to eighteen months. In acute forms there is irregular fever, progressive wasting, diarrhoea occasionally, motor disturbances involving the hind quarters, and the animal dies in a comatose con- dition. In the chronic form the animal may appear well, except for loss of weight. The parasites may be found in the internal organs of these experimental dogs, but are not numerous in the peripheral blood except at times of high fever. Experimental monkeys live about three months. It may be interesting to record the number of dogs found to be infected naturally with leishmaniasis in various countries. In Tunis, Nicolle and Yakimoff found about 2 per cent, infected out of about 500 dogs examined. Sergent in Algiers found 9 infected out of 125 dogs examined. In Italy and Sicily, Basile found about 40 per cent, of the dogs to be infected out of 93 examined at Rome and Bordonaro. Cardamitis found 15 infected out of 184 examined in Athens. In Malta, Critien found 3 infected out of 30 dogs examined. Alvares found i infected dog out of 19 examined in Lisbon. Pringault has recently (December, 191 3) found an infected dog in Marseilles.^ Yakimoff and Schokhor found 24 per cent, infected out of 647 dogs examined in Turkestan. The distribution of the parasites in the body of the human patient is much the same as in the case of Indian kala-azar. Critien records Bull. Soc. Path. Exot.. p. 41 LEISHMANIA INFANTUM IH the finding of parasites in the mucous flakes of the stools of a three- year-old Maltese child/ Intestinal lesions rarely occur in infantile leishmaniasis. Etiology. — Infantile leishmaniasis is stated to be transmitted by fleas, especially dog fleas, Ctenocephalus cams (= Pulex serraticeps), and by Pulex irritans. Children living in contact with infected dogs may be bitten by infected dog fleas, and so contract the disease. Basile (1910-11) and Sangiorgi (1910) state that they found L. infantum parasites in the digestive tract of the dog flea. After searching they found infected dog fleas on the beds, mattresses, and pillows used by children suffering from the disease. Franchini (191 2) thinks that Anopheles maculipennis may be concerned in the transmission. Basile^ tried a number of experiments to show that infantile leishmaniasis is transmitted by fleas, thus : — (i) Fleas were taken from a healthy dog. They were placed in vessels containing infected spleen-pulp and allowed to feed thereon. The fleas were then killed and dissected, and portions of the gut- contents examined for parasites. The remainder of the gut was emulsified and injected into a young puppy, whose bone-marrow had been shown previously to be uninfected. Basile states that the puppy became infected. The parasites are said to increase in number in the flea's gut. (2) Two healthy pups, each a month old, and born in the labora- tory, were placed in a disinfected, flea-proof cage. A few days after, an infected dog was placed in the cage, so that fleas from the infected dog could pass on to the puppies. A month later the two pups became infected, parasites being found in them after liver puncture. A number of control puppies from the same litter remained uninfected and in good health. (3) Basile next used other laboratory-born puppies, a month old. Four of the litter were placed in a disinfected, flea-proof gauze cage in Rome. The cage was isolated from other dogs. Fleas obtained from an infected area in Sicily were placed in the cage. The puppies were examined by hepatic puncture, but w^ere found to be negative for two months. Then two of the puppies showed infection, and six days later the remaining two puppies were found to be infected, and all four died. They showed irregular temperatures, and were getting thin. Control puppies remained healthy. From these experiments Basile concludes that fleas transmit leishmaniasis. However, Basile did not exclude the possible occur- rence of natural herpetomonads in the gut of the fleas.^ Herpetomonas ' Quoted by Leishman (191 1) in his interesting review of Leishmaniasis, lourn. Foy. Army Med. Corps, xvii, p. 567, xviii, pp. I, 125. Also Quart, fourn. Med. v, pp. 109-152. "^ Numerous papers in Rendiconti R. Accad. dei Lincei (Rome), xix, xx (1910-II). 8 See Fantham, Brit. Med. foiirn.^ 1912, ii, p. 1 196. 112 THE ANIMAL PARASITES OF MAN ctenocephali is known to occur in the gut of Ctenocephalus canis A natural Herpetoinonas is also known in the gut of P^ilex irritans, as well as a Crithidla (C. piilicis, Porter). These natural flagellates of the fleas pass through non-flagellate stages, like the Leishman-Donovan body. In consequence Wenyon and Patton, among others, have criticized Basile's results. Further, other investigators, such as Wenyon and Da Silva (191 3), have repeated Basile's flea experiments and been unable to confirm them. In feeding and inoculation experiments the incubation period of the parasite may be long, and so it is necessary to wait a long time to see whether the parasite will develop. Iminnnity. — Nicolle has tried some experiments with L. infantum and L. tropica. He finds that in animals recovery from an attack of the former confers immunity against infection by the latter and vice-versa. Laveran^ records that a monkey having an immunity against L. infantum was also immune to L. donovani. As mentioned on p. 103, Laveran and Franchini (19 13), working in Paris, have succeeded in inoculating Herpetomonas ctenocephali, a natural flagellate in the gut of the flea, Ctenocephalus canis, into white mice. Leishmaniform stages of the flea flagellate were re- covered from the peritoneal exudate, blood and organs of the mice some weeks after inoculation. The parasites may also be conveyed by way of the digestive tract of the vertebrate. Similar experiments have succeeded with H. pattoni. These experiments go to show, together with those of Fantham and Porter with H. jaculum (seep. 104), that, in the words of the latter authors, *^ it may be expected that the various leishmaniases, occurring in different parts of the world, will prove to be insect-borne herpetomoniases." Genus. Histoplasma, Darling, 1906. Under the name Histoplasma capsulattim,^ Darling described small round or oval parasites, enclosed in a refractile capsule, and each con- taining a single nucleus. The bodies were found in cases of spleno- megaly in Panama. They occurred in the endothelial cells of the small blood-vessels of the liver, spleen, lungs, intestine and lymphatic glands, and also within the leucocytes. A few flagellates were stated to occur in the lungs. The parasite has usually been placed near Leish- mania, but recently Rocha-Lima has stated that Histoplasma is a yeast. Genus. Toxoplasma, Nicolle and Manceaux, 1908. The genus was created for crescentic, oval or reniform parasites, 2*5 ju to 6 /u by 2 Ai to 3 /i, possessing a single nucleus and multiplying by binary fission. They occur ^ Annales Inst. Pasteur (1914-15), xxviii, pp. 823, 885 ; xxix, pp. i, 71. "^ lotirn. Amer. Med. Assoc, xlvi, p. 1283 : /ourn. Exptl. Med. (1909), xi, p. 515. TOXOPLASMA 113 in mononuclear and polymorphonuclear cells in the blood, spleen, liver, peritoneum etc. (fig. 51). The parasites have been found in the gondi, dog, rabbit, mole, mouse, pigeon and other birds. Although various species names have been given to the parasites in these hosts, it seems probable, from cross infection experiments, that there is but one species with several physiological races. Splendore' (1913) has described a flagellate stage. Castellani (1913-14)^ has described similar parasites from a case of splenomegaly, with fever of long standing, in a Sinhalese boy. The Fig. 51. — Toxoplasma gondii, endocellular or free in the peritoneal exudate of infected mice. I, 2, mononuclear leucocytes containing toxoplasms. 3, polynuclear, containing parasites. 4, 5, 6, endothelial cells containing toxoplasms, agglomerated in 6. 7, agglomera- tion forms. 8-1 1, free forms. 12-13, division stages. X i,6do. (After Laveran and MaruUaz.) 4» ^^ ■V 9. ^^ ':i Fig. ^2.— Toxoplasma pyro^eius. I, body found in blood. 2-7, bodies found in spleen, [i is about the size of a red blood corpuscle, as drawn in the figures!. Magnification not stated. (After Castellani.) bodies were found in the spleen and more rarely in the blood (fig. 52). Castellani has named them Toxoplasma pyrogeiies. Further researches are needed. ^ Bull. Soc. Path. ExoL, vi, p. 31 J ■^ Journ. Trop. Med. and Hyg., xvii, p. 113. 114 THE ANIMAL PARASITES OF MAN THE SPIROCH.'^TES. The Spirochaetes are long, narrow, wavy, thread-like organisms, with a firm yet flexible outer covering or periplast. There is a diffuse nucleus internally in the form of bars or rodlets of chromatin distributed along the body. In some forms there is a membrane or crista present (fig. 53), which in the past was compared with the undulating membrane of a trypanosome, but the membrane of a spirochaete does not undulate. Progression is very rapid, corkscrew- like and undulatory movements occurring simultaneously. The genus Spirochceta was founded by Ehrenberg in 1833 for an organism which he discovered in stagnant water in Berlin. Ehrenberg named the organism Spirochceta plicatilis. According to Zuelzer (191 2) S. plicatilis does not possess a membrane or crista, but an axial filament. S. gigantea has been described by Warming from sea- water. Spirochaetes occur in the crystalline style and digestive tract of many bivalve molluscs. The first molluscan spirochaete to be studied was that of the oyster, named by Certes (1882) " Trypanosoma " balhianii (fig. 53). Similar spirochaetes, probably belonging to the same species, occur in various species of Tapes and in Pectcn (the scallop). S. halbianii has rounded ends (fig. 53). Other spirochaetes occur in fresh- water mussels {Anodonta spp). .S. anodonice, studied by Keysselitz (1906) and by Fantham (1907), has pointed ends. Gross (191 1) sug- gested the generic name Cristispira for mol- luscan spirochaetes, because they possess a well-marked membrane or "crista," which appears to be absent from S. plicatilis, accord- ing to Zuelzer's researches. Schaudinn in 1905 founded the genus Treponema for the parasite of syphilis {T. pallidum), discovered by him and by Hoffmann. According to Schaudinn the Treponemata have no membrane or crista. The pathogenic agent of yaws or framboesia, discovered by Castellani, is also placed in the genus Treponema, as T. pertenue. There remain the blood spirochaetes. It is somewhat disputed as to whether these organisms possess a membrane. The present writer considers that they have a slight membrane or crista. The name of Fig. 53. — SpirocJmta bal- hianii. a, basal granule or polar cap. /^, chromatin rod- lets, r, membrane ("'crista"). d, myonemes in membrane. ■{After Fantham and Porter.) THE spirochj5:tes 115 tlie genus in which to place the blood-inhabiting forms is somewhat uncertain and disputed. Various generic names given to them are Spirochceta, Treponema, Spiroschaudinriia (Sambon) and Borrelia (Swellengrebel). Included in this division are the causal agents of relapsing or recurrent fever. These Protists will be named, for description, Spirochaetes without prejudice as to the ultimate correct generic name. It is sometimes made a matter of argument as to whether the spirochcctes are Protozoa or Bacteria. Such arguments are somewhat unprofitable. Morphologically the spirochaetes are like the Bacteria in possessing a diffuse nucleus. They differ from Spirilhim, an undoubted bacterial genus, in being flexible and not possessing flagella. Molluscan spirochaetes, however, may appear to have flagella if their membrane becomes frayed or ruptured, when the myonemes therein (fig. 53), becoming separated, form apparent threads or flagella (Fantham, 1907-08).^ Again, the mode of division of spirochaetes has been used as a criterion of their bacterial or protozoal affinity. They have been stated to divide transversely, longitudinally, and by *' incurvation," or bending on themselves in the form of a U, "a form of transverse fission." The present writer believes that they divide both transversely and longitudinally, and that there is a periodicity in their mode of division at first longitudinal (when there are few spirochaetes in, say, the blood) and then transversely (w^hen spirochaetes are numerous in the blood).^ Some authors consider that longitudinal division is ex- plained by :" incurvation." The spirochaetes of relapsing fever show a remarkable periodic increase and decrease in numbers in the blood. They are transmitted bv ticks or by lice. They react to drugs {e.g., salvarsan or ^' 606 ") rather like trypanosomes, and — like Protozoa, but unlike Bacteria — they are cultivated with difficulty. These and other criteria have been used to endeavour to determine whether they are Protozoa or Bacteria. The present writer believes that they are intermediate in character, showing morphological affinities with the Bacteria and physiological and therapeutical affinities with the Protozoa. The group Spiro- chaetacea, as an appendix to the Protozoa, has been created for them by the present writer (Jan., 1908). Others have placed them in the Spirochaetoidea of the Bacteria or with the Spirillacea. Doflein (1909) called them Proflagellata. Further discussion is unnecessary, as they are undoubtedly Protista (see p. 29). There is no true conjugation, sex or encystment in spirochaetes, but morphological variation may occur.^ They may agglomerate. ' Qtiart. Journ. Microsc. Set., lii, p. I. ^ Froc. Roy. Soc, B, Ixxxi, p. 500. 3 Fantham, Parasitology, ii, p. 392. Il6 THE ANIMAL PARASITES OF MAN The Spirochaetes form an interesting chapter in the evokition of parasites. There are free Hving forms, parasitic forms in the guts of both vertebrates and invertebrates, and blood-inhabiting forms. These probably represent the order of evolution of parasitism. The blood-inhabiting forms are pathogenic to warm-blooded hosts. We must now consider the blood Spirochaetes and the Treponemata (organisms of syphilis and of yaws). THE SPIROCHETES OF THE BLOOD. There are at least two important human parasites included hereunder ; — (a) Spirochceta recurrentls (^S. obermeieri), (b) Spirochcvta diittoni. More is known of the life-cycle of Spirochceta dnttonij and it will be convenient to consider that first. Spirochaeta duttoni, Novy and Knapp, 1906. The specific name duttoni was also given, independently, to this parasite in 1906 by BreinI and Kinghorn. S. duttoni is the pathogenic agent of African tick fever in man, prevalent in the Congo State and other parts of Africa. The full- grown organism is about 16 //, to 24 /i, long, and has pointed ends. It is 0*25 yu, to 0*5 yLt broad. P. H. Ross and Nabarro were among the earliest to see a spirochaete in the blood of patients in Uganda. It is transmitted by the tick, Ornithodorus moxibata. In the blood of the patient some of the spirochaetes may show, after staining, lighter and darker portions (chromatin dots) and evidence of the possession of a very narrow membrane (fig. 54). The mode of division has already been discussed. Periodicity in the direction of division was first described by Fantham and Porter,^ (1909). Just before the crisis in African tick fever, BreinI has stated that S. duttoni becomes thinner in the spleen and bone-marrow and rolls up into skein-like forms, which are surrounded by a thin *^cyst" wall (probably the periplast). Such occur in apyrexial periods. Inside the cyst the spirochaete breaks up into granules. Balfour and Sambon have described somewhat similar rolled up forms, breaking into granules, inside the red blood cells of Sudanese fowls in the case of S. granulosa (possibly only a variety of S. gal- linarum). The intracorpuscular stage is not definitely established. The granule phase, however, is an essential one in the invertebrate transmitter (fig. 54c). In 1905,^ Button and Todd proved experimen- tally that 0. nioubata transmitted S. duttoni. They fed ticks, obtained * Proc. Roy, Soc, B, Ixxxi, p. 500. - Liverpool Sch. Trop. Med., Metnoir xsn'. Lancet, Nov. 30, 1907, p. 1523. SPIROCH^TA DUTTONl 117 from Congo native huts in which infected persons hved, on monkeys and the latter became infected. Button and Todd also found the offspring of infected ticks to be capable of transmitting the infection to experimental animals. They concluded that 0. mouhata was a true intermediate host. A little later in 1905, Koch stated that spirochaetes from the gut of the tick penetrated the gut wall and tissues and found their way into the eggs in the ovary. Koch figured tangled masses of spirochaetes as occurring in the tick eggs. He found ticks infective to the third generation. He thought that the infection was spread by the salivary fluid of the tick, in the act of biting. (This is now known to be incorrect.) Markham Carter (1907) corroborated Koch's work on the spirochaetes in the tick eggs, and they have been seen since by Kleine and Eckard (1913)- Sir William Leishman,^ in 1909-10, found that at ordinary temperatures the salivary glands of infected ticks (0. rnoiibata) were not themselves infective, and that the infection occurred by way of the ticks' excretion. The spirochaetes (contained in the ticks' excrement) found their way into the vertebrate host through the wound made by biting. While feeding, ticks pass large quantities of clear fluid from the coxal glands ; in this fluid an anticoagulin occurs. Some of the ticks also Fig. 54. — Spirochxta duttoni. a, blood pass thick, white Malpighian form showing slight membrane; <^, granules or ^ . [lu i- • + coccoid bodies clearly formed wiihin the organism; secretion, that is, excrement, ^^ beginning of extrusion of coccoid bodies in the towards the end of the feed. t'ck. (After Fantham.) LeishmaU; using experimental monkeys, showed that if infected ticks were interrupted while feeding, then no infection resulted in the monkeys. If, however, the ticks were allowed to finish their feed, and the Malpighian secre- tions were passed, then the experimental monkeys became infected. Fantham^ and Hindle^ (191 1)» independently, have repeated the experiments with mice. •% ' Journ. Roy. Army Med. Corps, xii, p. 123 ; Lancet (1910), clxxviii, p. ii. '2 Anncds Trop. Med. and Parasitol., v, p. 479. ' Parasitology, iv, p. 133. ii8 THE ANIMAL PARASITES OF MAN Coxal Leishman's methods and results may be summarized thus : SaHne emulsions of the organs of infected ticks were made, after the organs had been most carefully dissected out. The ticks were first kept for several days at certain constant temperatures, such as 24° to 25° C. or blood heat, 37° C. The saline emulsions of the organs were inoculated, separately, into experimental animals, and the results recorded : — At 24° C. At 37° C. Salivary glands Negative ... Positive Malpighian tubules Positive ... Positive Gut and contents Positive ... Positive Excrement Positive ... Positive Genital organs Positive ... Positive fluid is usually negative ; thick, white excrement from Malpighian tubes is posi- tive. When the ticks were incubated at 21° to 24° C. no spirochaetes, as such, were seen in the organs, except perhaps in the gut, where they often dis- appeared in a few days. When the ticks were pre- viously incubated at 35° to 37° C. for two to three days, spirochaetes, as such, re- appear in the gut, organs and haemoccelic fluid. The infection proceeds, from the salivary gland, but from the infective excrement, is, from the thick, white material voided by the tick while Fig. 55. — Spirochceta duttoni and its coccoid bodies in the tick {0. vioubata). — Mononuclear cells of the tick {O. moubata) containing [a) Spirochete breaking up into coccoid bodies ; {b) Similar tick-cell containing coccoid bodies or granules. Such mononuclear cells occur in various organs of ticks and in developing Malpighian tubules. (Original. From preparations by Fantham.) not that feeding, usually towards the end of the meal. This Malpighian excrement passes into the wound caused by the bite, being greatly aided by the clear and more limpid coxa fluid, which bathes the under surface of the tick's body, and mixes with and carries the infective excrement into the wound. Ticks remain infective for a long time. The spirochaetes in the gut of infected ticks divide by a process of multiple transverse fission into granules, which are composed of chromatin (fig. 54). These granules — sometimes known as coccoid bodies — are capable of multiplication. Leishman first found them in clumps inside the cells of the Malpighian tubules (cf. fig. 55). To summarize, when spirochaetes are ingested by a tick, some of them pass through the gut- wall into the haemoccelic (body) fluid. They then bore their way into the cells of various organs (fig. 55^7) SPIROCH^TA GALLINARUM II9 and break up into coccoid bodies. In this manner the granules find their way into the ovaries and ova, thus explaining how the young ticks are born infected. Inoculation of these chromatinic granules usually produces infection. Infective granules are also seen in the rudiments of the Malpighian tubules of embryo ticks. Bosanquet and Fantham (191 1), independently, have shown that molluscan spirochaetes also break up into similar granules or coccoid bodies. Gross has also demonstrated multiple transverse fission in molluscan forms. Marchoux and Couvy (19 13) and Wolbach (1914) consider the granules or coccoid bodies to be degeneration products. This is unlikely (see below). Schuberg and Manteufel have found that certain 0. moubata, perhaps 30 per cent, of the specimens of a given neighbourhood, may acquire a natural active immunity against infection with S. duttoni. S. duttoni, or a closely allied form (by some termed S. novyi), occurs in Colombia, and is spread by the tick Ornithodorus turicata. In Panama a similar spirochaete is probably spread by 0. talaje. Spirochseta gallinarum, Stephens and Christophers, 1905 (= Spirochaeta marchouxi, Nuttall, 1905). This Spirochaete, which occurs in fowls and is pathogenic, is transmitted by the tick Argas persicus. It is about 10 fi to 20 fj, long. There is a pathogenic spirochaete known to occur in geese, named by Sakharoff (1891) S. anserina, and found in Caucasia. This may be the same as S. gallinanim, in which case the name S. anserina will have priority. These organisms cause fever, diarrhoea, anaemia and death. The life history of the avian pathogenic spirochaetes has been studied by Balfour, by Hindle^ and by Fantham.^ It is essentially similar to that of S. duttoni. Marchoux and Couvy^ (1913) consider that the '^fragmentation of the chromatin " in spirochaetes is a process of degeneration. Working with A. persicus and S. gallinarum, they state that a large number of the spirochaetes ingested by the Argas almost immediately pass through the wall of the alimentary canal and appear in the haemo- ccelic fluid. Marchoux and Couvy consider that Leishman's granules may be found in the Malpighian tubules of various Arachnids. They found spirochaetes in the cephalic glands of infected Argas. They consider that spirochaetes remain as wavy spirochaetes within the tick, if they are to be infective, though the spirochaetes may become so thin as to be invisible ! The latter argument is obviously weak, and it was never asserted that all granules in the Malpighian tubules of infected ticks were derived from spirochaetes. With dark-ground ' Parasitology, iv, p. 463. ^ Annals Trop. Med. and Parasitol. {1911), v, p. 479. ^ Annales Inst. Pastenr, xxvii, pp. 450, 620. 120 THE ANIMAL PARASITES OP^ MAN illumination small, refractile spirochaetal granules may be seen to grow into spirochaetes. The granule phase of spirochaetes has recently been discussed by Fantham^ (1914)- Spirochaeta recurrentis, Lebert, 1874. Syn. : Spirochceta obermeieri^ Cohn, 1875. This organism was discovered by Obermeier (1873) in cases of relapsing fever in Berlin. Short forms 7 /x to 9 /^ long, and longer (probably adult) forms, 16 /^ to 19//-, are found in the blood. The width is 0-25 /x. Parasites 12 yu, or 13 /a long are often observed. The spirochaete is found in the blood during febrile attacks and relapses, but not during intervening periods. It can be inoculated into monkeys, rats and mice. It can live in the bed-bug, Cimex lectnlarms, and Nuttall has succeeded in transmitting S. recurrentis from mouse to mouse by the bites of the same bug. The French investigators Sergent and Foley (1908-9) in Algeria, and NicoUe, Blaizot and Conseil (191 2) in Tunis, have shown experimentally that S. recurrentis (var. berhera) is transmitted by lice. The latter workers also demonstrated the method of infection that commonly occurs, namely, by the scratching of the skin and crushing of lice containing spirochaetes on the excoriated surface of the body. Lice as transmitting agents for relapsing fever were indicated by Mackie^ in 1907. An epidemic among Indian school children furnished the materials.^ It was noted that out of 170 boys, 137 were infected, and the boys were very verminous. Among the girls, 35 out of 114 suffered, and few lice were found on them. Twenty- four per cent, of the lice taken from the boys contained spirochetes as compared with 3 per cent, of those from the girls. As the epidemic died out among the boys, the lice also became fewer> and an increase in the number of cases among the girls coincided with an increase in the number of lice. Spirochaetes were found in the gut, Malpighian tubules and genital organs of the lice. Mackie thought that infec- tion of the patients was brought about by the regurgitation of the spirochaetes when the lice fed, but proof of this was lacking. In 191 2, Nicolle, Blaizot and Conseil,* working in Tunis and using chiefly an Algerian strain of relapsing fever spirochaetes (some- times called S. berhera) j showed by direct experiments that infection by means of the bites of Pediculus vestinienti and P. capitis was untenable. As many as 4,707 infected lice were fed on one man, and 6,515 on another occasion were allowed to bite a man after they had fed on a monkey heavily infected with spirochaetes, yet no infection of the man followed. Examination of the lice showed that the spirochaetes left the gut soon after they were ingested, and ' Amials Trop. Med. and ParasitoL, viii, p. 471. "^ Brit. Med. Journ., Dec. 14, 1907, p. 1706. ' See also Nuttall, Herter Lecture on Spirochsetosis, Parasitology, v, p. 269. * C.R. Acad. Sci., cliv, p. 1636 ; civ, p. 481. SPIROCH^TA RECURRENTJS 121 passed into the body cavity, which swarmed with spirochaetes. The contents of the ah'mentary tract and the faeces of the Hce ahke were iininfective. The spirochaetes did not reappear in the gut till eight days after an infective feed, but some persisted as late as the nineteenth day when kept at 28° C. It was noted that the irritation due to the lice caused scratching, and that thereby lice became crushed on to the skin. An emulsion was made of two infected lice and rubbed on to the slightly excori- ated skin of one of the above workers. Infection followed five days later. A drop of emulsion placed on the conjunctiva of the human eye produced spirochaetosis after an incubation of seven days. The body contents of such lice, then, produce infection when they reach the blood by any excoriated or penetrable surface. The stages leading lip to infection in nature briefly are : The irritation due to the louse bites causes scratching, and the lice are crushed on to the skin. The slight abrasion is quite sufficient to permit the entry of the parasite. The louse bite alone is harmless. Infection by way of the eye is quite probable in Africa, remembering the constant trouble due to sand, dust, insects, etc., resulting in frequent touching of the eyes. The spirochaetes occur in the body fluid of the lice and can pass in it to the adjacent organs. Thus they probably find their way into the genital organs, and into the eggs of the lice. Eggs laid twenty to thirty days after the parent became infected have retained the infection, and the larvae issuing from such eggs must have contained some form of spirochaetes, for an emulsion of either the eggs or the larvae produced spirochaetosis when inoculated into monkeys. Further details regarding the spirochaetosis in the eggs of the lice and in the larvae are needed. Hereditary infection, however, has been demonstrated, but is not very common. Sergent and Foley (19 14) state that the spirochaete possesses a very small and virulent form which it assumes during apyrexial periods in man and during a period following an infecting meal in the louse. Nicolle and Blanc {1914) find that the organisms are infective in the louse just before they reappear as spirochaetes. Nicolle and Blaizot found that female lice were more susceptible to spirochaetes than males, four times as many females as males being infected. Tictin (1897) found S. recurrentis in bugs recently fed on patients, and infected a monkey with the fluids of crushed bugs. Karlinski (1902) found the spirochaetes in bed-bugs in infected houses. There is some other evidence to show that bugs may transmit the spirochaete in Nature. Further researches are needed regarding the relationship of bed-bugs and human spirochaetosis. Multiplication of S. recurrentis is by longitudinal and transverse division (including so-called '^incurvation"), and the organism forms small, ovoid bodies ('' coccoid " bodies) in the same way as S. diittoni. 122 THE ANIMAL PARASITES OF MAN S. recurrentis is the cause of European relapsing fever, and a number of possible varieties of it are associated with relapsing fevers in other parts of the world. Such spirochaetes only differ by bio- logical reactions, such as acquired immunity tests. They include : — 5. rossii^ the agent of East African relapsing fever ; S. novyiy the agent of North American relapsing fever; S. carteri, the agent of Indian relapsing fever; S. herhera, the agent of North African and Egyptian relapsing fever. Other Human Spirochetes are: — S. schaiidinni. This organism, according to Prowazek, is the agent of ulcus tropicum. It varies in length from lo/i to 20 /x. S. ahoriginalis has been found in cases of granuloma inguinale in British New Guinea and Western Australia. It also occurs in dogs, and may not be truly parasitic. S. vincenti. This spirochaete is 12 /a to 25 /i, in length, tapers at both ends and has few coils. It has been associated with angina vincenti. It often occurs in company with fusiform bacilli. S. hronchialis, found by Castellani in 1907 in cases of bronchitis in Ceylon. The parasites are delicate, but show morphological variation. This organism is important and has since been found in the West Indies, India, Philippine Islands and various parts of Africa, such as the Anglo-Egyptian Sudan, Uganda and West Africa. It has recently been the subject of research by Chalmers and O'Farrell, Taylor, and Fantham. S. phagedenis was found by Noguchi in a ten days old ulcerated swelling of the labium. The organism shows much variation in size, being 4/x to 30 //, in length. S* refringens (Schaudinn, 1905) occurs in association witli Treponema pallidum in syphilitic lesions, but is non-pathogenic. It is 20 /i, to 35 /A long and 0*5 jju to 075 fi broad, being larger than r. pallidum and more easily stained. Various spirochaetes have also been notified in vomits, chiefly in Australia ; others from the human intestinal tract, e.g., S. eurygyrata ; S. stenogyrata (Werner) ; S. hachaizce (Kowalski), in cholera motions; S, buccalis (Cohn, 1875) and S. detitiiim occurring in the human mouth and in carious teeth (S. dentium, Koch, 1877, being the smaller) ; S. acuminata and S. obtusa found by Castellani in open sores in cases of yaws. Animal spirochaetes of economic importance include : — S. anserinay highly pathogenic to geese. S. gallinariim ( = S. marchouxi) in fowls. (See p. 119.) S. theileri in cattle and S. ovina in sheep also occur in Africa ; their pathogenicity is not clear. S. laverani ( = S. miiris), occurring in the blood of and patho- CULTIVATION OF SPIROCHETES 1 23 genie to mice, is probably the smallest spirochsete from the blood, being only 3 /x to 6 fi long. Numerous spirochaetes have been recorded from the guts of various mammals, birds, fishes, amphibia and insects. Cultivation of Spirochetes. — Cultures of spirochaetes have been made with little success or with great difficulty until compara- tively recently, when Noguchi (191 2) devised a means whereby he has cultivated most of the pathogenic spirochaetes as well as some Treponemata. Noguchi has now cultivated S. duttoni, S. recurrentis, S. rossii, S. novyi and S. gallinariun from the blood ; S. phagedenis^ from human phagedaenic lesions ; S. refringens^ and spirochaetes from the teeth. His method is as follows : — A piece of fresh, sterile tissue, usually rabbit kidney, is placed in a sterile test-tube. A few drops of citrated blood from the heart of an infected animal, ^.^., rat or mouse, is added, and about 15 c.c. of sterile ascitic or hydrocoele fluid is poured quickly into the tube. Some of the tubes are covered with a layer of sterile paraffin oil, others are left uncovered. The tubes are incubated at 37° C. The best results are obtained if the blood is taken from an animal forty-eight to seventy- two hours after it has been inoculated, that is, before the spirochaetes reach their maximum multiplicative period in the blood. The presence of some oxygen seems indispensable for these blood spirochaetes, and they fail to develop in vacuo or in an atmosphere of hydrogen. For subcultures, 0*5 c.c. of a culture is added to the medium instead of citrated blood, and it is useful to add a little fresh, normal blood, either human or from an animal, such as a rat. Noguchi found that the events in cultures were :^ S. duttoni,^ maximum multiplication on the eighth to ninth day ; disintegration beginning on the tenth day, spirochaetes disappeared after about the fifteenth day. No diminution of virulence was found at the ninth day. S. rossii ( = S. kochi).^ Maximum development on the ninth day, after which the virulence diminishes. The incubation period is also prolonged. S. recurrentis^ (=S. obermeieri). Maximum growth on the seventh day. S. novyi. ^ — Maximum development on the seventh day. It is more difficult to grow than the preceding forms. All the above spirochaetes showed undoubted longitudinal division and transverse division was observed in part. S. gallinarum ^ can be cultivated as above, but transverse division ^ Jotirn. Exptl. Med., xvi, p. 261. ^ Journ. Exptl. Med., xv, p. 466. ^ Journ. Exptl. Med., xvi, p. 202. ■• Ibid., p. 205. ^ Ibid., p. 205. « Ibid., p. 20S. " Ibid., p. 620. 124 THE ANIMAL PARASITES OF MAN was usual here. Maximum growth occurred in the culture about the fifth day. Treponemata. The genus Treponciim (Schaudinn, 1905), includes minute, thread- like organisms, with spirally coiled bodies, the spirals being preformed or fixed. No membrane or crista is present, according to Schaudinn, though a slight one is said by Blanchard to be present in the organism of yaws. The ends of the organisms are tapering and pointed. Multiplication is by longitudinal and transverse division. The most important members of the genus are T. pallidum, the agent of syphilis, and T. perte/iiie, which is responsible for framboesia or yaws. Treponema pallidum, Schaudinn, 1905. Syn. : Spirochcrta pallida. Treponema pallidum was first described by Schaudinn and Hoff- mann in 1905 under the name of Spirochaita pallida. It has also been described under the names oiSpirouema pallida, Micro- sp iron em a pallida and Trypanosoma hits. Siegel in 1905 described an organ- ism which he called Cyto- rhyctes Inis and considered to be the agent of syphilis. Schaudinn reinvestigated Siegel's work and found T. pallidum, which he con- sidered to be the causal agent of the disease, and pronounced against Cyto- rhyctes Inis. It is probable now that both workers were correct, for Balfour (191 1) has seen the emis- sion of minute granules or " coccoid " bodies from T, pallidum and these granules probably correspond to the C. luis of Siegel. Recently E. H. Ross, having observed a spirochaete stage in the development of Kurloff bodies, thinks that T. pallidum is a stage in the life-history of a Lympho- cytozoon. MacDonagh has also described a complicated and some- what similar cycle, but these observations require further study and confirmation. T. pallidum varies from 4/x to 10 fjL in length, its average length being y fM, while its w4dth is usually about 0*25 /n. Longer individuals Fig. 56. — Trepo7teina pallidum. (After Bell, from Castellani and Chalmers.) TREPONEMA PALLIDUM 125 of i6yLt to 20 /x have been recorded. The body has from eight to ten spiral turns and forms a tapering process at each end (fig. 56). The organism is most difficult to stain, and its internal structure is Uttle known. It is possibly like that of Spirochceta diittoiil or S. halhianiiy as the '' granule shedding " observed by Balfour is strongly suggestive of the formation of resistant bodies by those spirochaetes. Hoffmann (19 1 2) has seen the formation of spores in T. pallidum. The Treponemata occur in the primary and secondary sores, but are difficult to find in the tertiary eruptions of syphilis. Noguchi and Moore (1913) and Mott' (19 1 3) have demonstrated T. pallidum in the brain in cases of general paralysis of the insane. Marie and Levaditi (19 14), however, consider that the treponeme found in the brain in such cases is different from T. pallidum. Cultivation of T. pallidum. — This has been accomplished successfully by No- guchi,^ using a modification of his method for spirochaete cultivation, for T. pallidum is much more difficult to grow than spirochaetes, being a strict anaerobe. The apparatus consists of two glass tubes, the upper being connected to the lower by a narrower tube passing through a rubber cork (fig. 57). Both tubes are carefully sterilized. A piece of fresh, sterile rabbit's kidney is placed in the lower tube, which \s filled with ascitic fluid, or ascitic fluid and bouillon mixture. The tube is in- oculated with syphilitic material and corked by inserting the upper tube. In the bottom of the upper tube a piece of sterile rabbit's kidney is placed and syphilitic material poured over it. A mixture of one part ascitic fluid and two parts of slightly alkaline agar is then poured over the tissue and allowed to solidify. When solid, a layer of sterile paraffin oil is poured on top of it, and the top plugged with cotton wool (fig. 57). The whole is then incubated at 37° C. for two or three weeks. The tissue removes traces of oxygen from the lower levels of the medium and also probably ' Brit. Med. [ourn.^ Nov. 15, 1913. p. 1,271. 2 /ourn. Exptl. Med., xv, p. 90; xvi, p. 211. 7iraf/tn V oil Solid ^•medium ^■Tissue nuid r- niQdium |«-Txs6ue Fig. 57. — Diagram of apparatus for cultivation of Treponema palli- dum by Noguchi's method. (After Noguchi.) 126 THE AxMIMAL PARASITES OF MAN provides a special form of nourishment. At first T. pallidum grows in the soHd medium, and then when the cultural conditions in the lower fluid portion become favourable, the organisms migrate thither and multiply abundantly. At first the culture is impure, but after several transferences a pure culture is obtained readily. The syphilitic material for culture is prepared by cutting off pieces of tissue from the lesions, washing in sterile salt solution containing i per cent, sodium citrate, and then emulsifying the tissue in a mortar with sodium citrate. Good cultures shaw rapid multiplication, w^hich is invariably by longitudinal division. In his various cultivation experiments Noguchi^ found morpho- logical and pathogenic variations in T. pallidiun. Three forms of the organism were found, namely, thicker, average and thinner types. The lesions caused in the testicle of the rabbit differ according to the variety inoculated, but more work is necessary on the subject. Noguchi^ has cultivated a separate organism, T. caUigyrum, from tlie surface of human genital or anal lesions, either syphilitic or non-syphilitic. It is apparently non-pathogenic, and is 6 //, to 14 /x long. Hata(i9i3)^ has modified the Noguchi technique for the cultivation of spirochaetes and treponemes, with a view to simplification and convenience. Hata substitutes normal horse serum for ascitic fluid and the " buffy coat " of the clot of horse blood in place of the small pieces of rabbit's kidney. It is unnecessary to place sterile paraffin on the surface of the medium. The horse serum is mixed with twdce its volume of physiological saline solution. The mixture is placed in tubes which are heated on a water-bath at 58° C, the temperature being raised gradually until it reaches 70° or 71° C. in three hours. The tubes are then heated at 71° C. for half an hour. After cooling, the contents will consist of an opaque semi-coagulated mass. This semi-coagulated serum and saline mixture may be substituted for Noguchi's ascitic fluid. The buff coagulum is cut into small pieces, about i c.c. in volumcx They must be forced with a sterile glass rod to the bottom of the semi- coagulated serum and saline mixture. The medium is inoculated wath a small quantity of infected blood and kept at 37° C. In the case of S. recurrentis, growth of spirochaetes is observed on the second day, reaching a maximum in five to seven days. The growth of the organisms proceeds rather more slowly, they live for a longer period and maintain their virulence better than in Noguchi's medium. ^ [ourn. Exptl. Afed., xv, p. 201. ^ /ourn. Exptl. Med.y xvii, p. 89. ^ Centralbl. f. Bakt., Orig., Ixxii, p. 107. TREPONEMA PERTENUE 1 27 Treponema pertenue, Caslellani, 1905. Syn. : Spirochceta pertenids ; S. pallidula^ Castellani, 19015. Castellani discovered the organism in 1905, in scrapings of yaws pustules. He first described it under the name of Spirochceta pertenuis. Treponema pertenue (fig. 58), though delicate and slender, shows great morphological variation both in length and thickness. It may be short, e.g., j /jl, but can attain 18//, to 20 /jl in length and may be even larger. In cultures made by Noguchi, thick, medium and thin forms were found, each giving rise to a different type of fram- boesial lesion when inoculated into the testicles of rabbits, thus suggesting the possibility of the occurrence of varieties of T. pertenue. The organism is difficult to stain, but occasionally deeper staining granules are found along its body. They may represent a diffuse nucleus. Granule formation similar to that of T. pallidum has been observed by Ranken, using dark-ground illumination. Many experiments have been made with a view to establishing the identity of the organism of yaws and also of differentiating between the causative agents of yaws and syphilis. Both monkeys and the human subject have been experimentally inoculated with yaws material and have developed the disease. In an early experiment, negroes were inoculated with the secretion from lesions . All r ii 11 11 Fig. 58. — freponema pertenue. of yaws. All of them developed the (After Castellani and Chalmers.) disease, nodules appearing, chiefly at the seat of inoculation, in from twelve to twenty days, followed by the usual eruption. Similar results were obtained with thirty-two Chinese prisoners, who were inoculated with yaws, twenty-eight becoming infected. A naturally infected yaws patient when inoculated with syphilis, contracted that infection, thus showing that yaws does not confer immunity to syphilis. This has also been observed naturally, when yaws patients have contracted syphilis. Experiments with monkeys have been successfully performed. The incubation period varies from sixteen to ninety-two days. Lesions appear first at the seat of inoculation, and in some monkeys the eruption is localized to this spot, though the infection is general, T. pertenue occurring in the spleen, lymphatics, etc. Monkeys inoculated with splenic blood of a yaws patient, and also sometimes with blood from the general circulation, have become infected. 128 THE ANIMAL PARASITES OF MAN Castellani and others have shown that monkeys successfully inoculated with syphilis do not become immune to yaws, and vice- versa. Craig and Ashburn, using the monkey Cynouiolgns philippincnsis, found these animals susceptible to yaws but not to syphilis. The ulcerated lesions of framboesia are rapidly invaded by numerous bacteria as well as by different spirochaetes, of which Castellani has described three distinct species. One is identical with Spirochceta refringens, Schaudinn, the other two are thin and delicate. One, S. ohtiisay has blunt ends ; the other S. acuminata, has pointed ends. T. pertenue is also present. The reasons for considering T. pertenue to be the specific cause of framboesia are : — (i) T. pertenue is the only organism present in non-ulcerated papules, in the spleen and in the lymphatics of yaws patients, or of monkeys artificially infected with the disease. By no method has any other organism been obtained. (2) Extract of framboesia material, free from all organisms other than T. pertenue, reproduces the disease if inoculated. (3) Extract of framboesia material deprived by filtration of T. pertenue is no longer infective on inoculation. The method of infection is contaminative, by direct contact. Women in Ceylon are frequently infected by their children. Any slight skin abrasion is sufficient to admit the parasite. In some cases, insects may carry the disease from person to person, and even in hospitals, when dressings are removed, it has been noticed that flies greedily suck the secretion from the ulcers. T. pertenue has been recovered from flies that have fed on yaws, and monkeys have contracted the disease when flies were placed and retained on them for a short time, after the insects had fed on yaws material. Cultivation. — T. pertenue has been cultivated by Noguchi, who finds three types of parasites in his cultures, as before mentioned. Its multiplication is by longitudinal division. Noguchi^ (191 2), has cultivated species of Treponema from the human mouth, e.g., T. ntacrodentium, T. niicrodeutluni and T. mucosnni, the latter from pyorrhea alveolaris. These parasites in the past may have been confused under the name Spirochceta dentium. Class III. SPOROZOA, Leuckart, 1879. The third group of the Protozoa consists entirely of parasitic organisms forming the class know^n as the Sporozoa or spore- producing animals. The members of this class are characterized ' foiirn. Exptl. Med., xv, p. 8l ; xvi, p. 194. TELOSPORIDIA 129 by possessing very great powers of multiplication, coupled with a capacity for producing forms that serve for the transference of the organisms to other hosts. These reproductive bodies, whether for increase of numbers within one host or for transmission to another host, are called spores. But, strictly, the term spore should be used only in the latter connection, when a protective or resistant coat known as a sporocyst envelops the body of the spore. The Sporozoa are widely distributed, occurring in various tissues and organs of Annelids, Molluscs, Arthropods, and Vertebrates. Their food, which is fluid, is absorbed osmotically. The life-cycle of a Sporozoon may be completed within one host or may be dis- tributed between two different hosts. The Sporozoa were divided by Schaudinn into two groups or sub-classes, called (i) the Telosporidia, and (2) the Neosporidia. The Telosporidia are Sporozoa in which the reproductive phase of the parasites is distinct from the growing or trophic phase, and follows after it. The Neosporidia include Sporozoa in which growth and spore-formation go on simultaneously. This classification is not final, for certain exceptions and difficulties are already known with regard to it. It is possible that the class Sporozoa is not a natural entity, but should be replaced by two classes of equal rank, corre- sponding in most respects with the Telosporidia and Neosporidia. The Telosporidia comprise the Gregarinida, the Coccidiidea, and the Haemosporidia. Doflein combines the two latter orders into one known as the Coccidiomorpha. The Neosporidia comprise the Myxosporidia, the Microsporldia, the Actinomyxidia, the Sarcosporidia, and the Haplosporidia. Doflein combines the first three orders into one, the Cnidosporidia. Sub-Class. TELOSPORIDIA, Schaudinn. Sporozoa in which the reproductive phases follow completion of growth. Order. Gregarinida, Aime Schneider emend. Doflein. Knowledge of the Gregarinida probably goes back as far as the year 1684, when Redi observed gregarines in the crab, Cancer pagiirus. Von Cavolini (1787) found them in Cancer depressus. The name Gregarina was created by L. Dufour (1828), who observed masses of these organisms in the gut of insects of different orders. Hammerschmidt (1838) and von Siebold found rich infestations in insects, while Dujardin (1835) and Henle described various genera from segmented worms. Henle (1835) also observed cysts containing "navicellae" in the sperm-sacs of segmented worms, and attention was drawn to his researches by the discovery by von Siebold (1839) of" pseudonavkrellse" in the gut of Sciara nitidicollis. Up to this time many workers considered the gregarines to be worms, but Kolliker (1845) investigated many of them and maintained their unicellular nature, while Stein's work (1848) showed the interrelation of the pseudonavicellas and the gregarines. 9 IjO THE ANIMAL PARASITES OF MAN The discovery of amoeboid germs in the pseudonavicellas by Lieberkiihn (1855) and the demonstration of myonemes further aided in the elucidation of their true systematic position. The entire process of conjugation, of which Du four had seen one phase, was followed by Giard under the microscope. From 1873 onwards Aime Schneider made important additions to the knowledge of the morphology, life-history, and systematic position of numerous gregarines. Butschli (1881) and L. Leger (1892) also contributed much work on the subject. The discoveries of Schaudinn with regard to the life-cycle of Coccidia gave a fresh stimulus to the study of the Gregarines, whereby the life-cycles of numerous forms and the phases thereof have been elucidated. Asexual multiplication is not common among the Gregarines, but is known to occur in the sub-order Schizogregarinea, formerly known as the Amoebosporidia. Although the Gregarinida are not known to be parasitic in man or other verte- brates, they are of great interest, inasmuch as they are among the earliest known Sporozoa, and therefore will be briefly described here. The Gregarines are usually elongate, somewhat flattened organisms (figs. 59, 60), whose bodies are enclosed in an elastic and often thick cuticle. The enclosed living substance shows a separation into ectoplasm and endoplasm, as is common among Protozoa. The cuticle is sometimes regarded as the outer portion or epicyte of the ectoplasm. A single, vesicular, spherical, or elliptical, large nucleus, with its chromatin concentrated to form a spherical karyosome, is present. The body of some gre- garines may be divided by ingrowing ectoplasmic partitions or septa, and are then said to be ^* septate" or "polycystid" (fig. 61). Other gregarines remain simple and non-septate. Fig. 59. — Monocystia agilis from seminal vesicles of Ltimbricus X 250. (After Stein.) .[(;dl abed e Fig. 60. — Gregaritia longa fiom larva of crane-fly {Tipula). a, in epithelial cell of host h, f, gradually leaving host-cell ; d, adhering to host-cell ; e, fully developed free trophozoite. and are termed 'Mnonocystid " (fig. 59). The monocystid gregarines occur especially in the body cavity of Chaetopoda and Insecta, more rarely in Echinodermata, in the parenchyma of Platyhelminthes, GREGARINIDA -. i^ also in the gut of Tunicata and Insecta (fig. 60) and in the seminal vesicles of Annelida. In the polycystid gregarines a single septum only is present as a rule, and thus the body presents two portions : (i) an anterior portion termed the protomerite ; (2) a posterior, larger portion, knOwn as the deuto- merite, which generally contains the nucleus. The protomerite is often modified anteriorly to form an organ of attachment, termed the epimerite (fig. 61), which is developed from the pointed rostrum of the sporozoite or primary infecting young gregarine. The structure of the epimerite may be complicated, being pro- vided with hooks, spines, knobs, and other appendages. An extension of the polycystid condition is seen in Tceniocystis mira Leger (from the dipteran larva, Ceratopogon solstitialis), whose body shows a number of partitions, giving the organism a superficial resemblance to a tapeworm. The ectoplasm of a gregarine exhibits three layers : (i) An epicyte (cuticle) externally of which the epimerite is composed ; (2) a sarcocyte which forms the septa if present ; (3) the deeper myocyte layer containing con- tractile elements in the form of fibrils or threads termed myonemes (fig. 62). The endoplasm is fluid and granular, con- taining many enclosures, which are of the nature of reserve food materials. They consist of fat drop- lets or of paraglycogen, and give the organisms an opaque appear- ance. Lithocystis contains crystals of calcium oxalate in its endoplasm. Many gregarines are capable of active movements, though they do not possess obvious locomotor organs: The movement is of a smooth, gliding character and two sug- gestions have been put forward to explain it. According to Schewiakoff, a gelatinous substance is secreted be- tween the layers of the ectoplasm. This is extruded posteriorly and thus the animal is pushed forward. On the other hand, Crawley consider^ that the movements are produced by contractions of the myonemes. Fig. 61. — Xyphorhynchus firmus with epimerite in intestinal epithelial cell of host. (After Leger.) (i E Fig. 62. — Gregarina munieri (from the beetle, Chrysomela hcenioptera). Sec- tion through surface layers. C«, cuticle ; E^ ectoplasm proper ; (7, gelatinous layer ; My, myonemes in myocyte layer. X 1500. (After Schewiakoff.) 132 THE ANIMAL PARASITES OF MAN sporocyst ; b, mature spore con- taining sporozoites ; ?i. Soc. Biol., Paris, 1858, v, 2 ; and Gaz. med. de Paris, 1858, p. 657 ; Leuckart, R., Die menschl. Paras., 1863, ist edition, i, pp. 49, 740. ^ Leuckart, R., Die menschl. Paras., 1863, 1st edition, i, p. 740.- ^ Leuckart, R., Die Pai-as. d. mensch., 1879, 2nd edition, p. 281. * Leuckart, R., ibid., p. 282. ^ Silcock, "A Case of Parasit. by Psorospermia," Trans. Path. Soc, London, 1890, xli, p. 320. * Pianese has confirmed the fact that Coccidia actually occur in the blood of the hepatic veins of infecteil rabbits. ' Die ei- ti. kiigelf. Psorosp. d. Wirbelt., 1870, p. 16. * Railliet and Lucet, " Obs. s. quelq. Cocc. intest.," C. R. Soc. Biol., Pari?, 1890, p. 660 ; Railliet, Trait. Zool. med. et agric, 2e ed., 1895, p. 140. ISOSPORA BIGEMINA 149 (c) Doubtful Cases. To these belong Virchow's case ' where, in the liver of an elderly woman, a thick walled tumour measuring 9 to 1 1 mm. was found. Among the contents of this tumour there were oval formations 56 fi long, surrounded by two membranes and enclosing a number of round substances. Virchow considered these foreign bodies to be eggs of pentastomes in various stages of development, others consider them to be Coccidia. The Coccidia which Podwyssotzki claims to have seen in the liver of a man, not only in the liver cells, but also in the nuclei, are also problematic.^ The parasite was called Caryophagus hominis. Again, other explanations can be given to an observation by Thomas, on the occurrence of Coccidiuin oviforme in a cerebral tumour of a woman aged 40. The growth was as large as a pea and surrounded by a bony substance.'^ Genus. Isospora, Aime Schneider, 1881. Syn. : Dipiospora^ Labbe, 1893. Belonging to the section Disporea^ that is, forming only two spores, each with four sporozoites. Isospora bigemina, Stiles, 1891. Syn.: '''' Cytospermiuiti villoru7ii i?itestinalium ca?tis et felis^'' Rivolta, 1874; " Coccidium Rivolta^'' Grass), 1882 ; Coccidium bigeininum, Stiles, 1891. This parasite lives in the intestinal villi of dogs, cats, and the polecat (Mustela putorins, L.). According to Stiles,'' the oocyst divides into two equal ellipsoidal portions or sporoblasts which become spores and then each forms four sporozoites. The oocysts of this species vary from 22 /x, to 40 yu, in length and from 19 fju to 28//. in breadth. Each spore is 10//, to 18 /^ long and contains four sporozoites. The parasites live and multiply, not only in the gut epithelium, but also in the connective tissue of the intestinal submucosa. Wasielewski has seen merozoites in the gut of the cat. Isospora bigemina (fig. 75) appears to occur also in man, for Virchow published a case which was communicated to him by Kjellberg, and attributed the illness to this parasite.^ Possibly also it would be more correct to ascribe the observation of Railliet and Lucet, which is mentioned under " Human Intestinal Coccidiosis," p. 148, to this species, as the Coccidia in that case were distinguished ' Arch, f. path. An.^ xviii, i860, p. 523. 2 Podwyssotzki, " Ueb. d. Bedeut. d. Coccid. in d. Path. Leber des Menschen," Centralbl. f. Bakt., vi, 1889, p. 41. 3 Thomas, J., "Case of Bone Formation in the Human Brain, due to the Presence of Coccidium oviforme,'' [ournal Boston Soc. Med. Sc, iii, 1899, p. 167; Centra/d/. /. Bakt. [ij xxviii, 1900, p. 882. * *' Notes on Paras.," No. 11, /ourn. of Comp. Med. and Vet. Sci., 1892, xiii, p. 517. * Arch, f path. An., i860, xviii, p. 527. ISO THE ANIMAL PARASITES OF MAN by their diminutive size (length 15 //,, breadth 10 /x). The case com- municated by Grunow may also possibly refer to hospora bigemina} Fig. 75. — hospora bigemina, Stiles (from the intestine of a dog), a, Piece of an intestinal villus beset with Isospora, slightly enlarged ; b, Isospora bigemina {15 fi in diameter), shortly before division ; c, divided ; d, each portion encysted forming two spores ; ^, four sporozoites in each part, on the left seen in optical section, together with a residual body— highly magnified. (After Stiles.) Roundish or oval structures of 6 /x to 13 yit in diameter occurred in the mucous membrane of the gut and in the faeces of a case of enteritis. Doubtful Species. In literature many other statements are found as to the occurrence of Coccidia-like organisms in different diseases of man. In some of the cases the parasites proved to be fungi. This was the case with the parasites of a severe skin disease of man, formerly called Coccidioides immitis and Coccidioides pyogenes. Other statements are founded on misapprehensions, or are still much disputed. If reference is here made to '■'' Eimeria hofnims" R. Blanchard, 1895, this is done on the authority of the investigator mentioned. The structures in question are nucleated spindle-shaped bodies of very different lengths (18 n. to 100 /*), which either occurred isolated or were enclosed in large globular or oval cysts, alone or with a larger tuberculated body ("residual body")- These formations were found by J. Kiinstler and A. Pitres in the pleural exudation removed from a man by tapping. The man was employed on the ships plying between Bordeaux and the Senegal River. Blanchard looks upon the fusiform bodies as merozoites and the cysts as schizonts of a Coccidium. On the other hand, Moniez declares the spindle bodies to be the ova and the supposed residual bodies to be " floating ovaries " of an Echinorhynchus. Severi's "monocystid Gregarines," which were taken from the lung tissue of a still-born child, are also quite problematical. No less doubtful are the bodies which Perroncito calls Cocciditwi jalinutn, and which he found in severe diseases of the intestine in human beings, pigs, and guinea- pigs ; Borini also reported another case. ' Grunow, "Ein Fall von Protozoen (Coccidien?) Erkrankung des Darmes," Arch. f. exper. Path, und Pharm., 1901, xlv, p. 262. H^MOSPORIDIA 151 Order. Haemosporidia, Danilewsky emend. Schaudinn. The Haemosporidia are a group of blood parasites, comprising forms differing greatly among themselves. Some of the forms need much further investigation. However, there are certain true Haemo- sporidia which present close affinities with the Coccidia, leading Doflein to use the term Coccidiomorpha for the two orders conjoined. The Haemosporidia present the following general characteristics : — (i) They are parasites of either red or white blood corpuscles of vertebrates during one period of their life-history. (2) They exhibit alternation of generations — asexual phases or schizogony alternating with sexual phases or sporogony — as do the Coccidia. (3) There is also an alternation of hosts in those cases which have so far been completely investigated. The schizogony occurs in the blood or internal organs of some vertebrate while the sporogony occurs in an invertebrate, such as a blood-sucking arthropod or leech. (4) Unlike the Coccidia, resistant spores in sporocysts are not generally produced, such protective phases in the life-cycle being unnecessary, as the Haemosporidia are contained within either the vertebrate or invertebrate host during the whole of their life. The Haemosporidia may be considered for convenience under five main types : — (i) The Plasinodiiitn or Hcemainceba type. This includes the malarial parasites of man and of birds. The asexual multiplicative or schizogonic phases occur inside red blood corpuscles and are amoeboid. They produce distinctive, darkish pigment termed melanin or haemozoin. Infected blood drawn and cooled on a slide may exhibit *' exflagellation " of the male gametocytes, i.e., the formation of filamentous male gametes. The invertebrate host is a mosquito. The malarial parasites of man are discussed at length on p. 155. Similar pigmented haemamoeboid parasites have been described in antelopes, dogs, and other mammals, and even reptiles. (2) The Halteriditun type. The trophozoite stage inside the red blood corpuscle is halter- shaped. Pigment is produced, especially near the ends of the organism. The parasites occur in the blood of birds. The invertebrate host of H. cohunhce of pigeons in Europe, Africa, Brazil and India, is a hippoboscid fly, belonging to the gtnviS Lynchia. Halteridium parasites are common in the blood of passerine birds, such as pigeons, finches, stone owls, Java sparrows, parrots, etc. The Halteridium embraces or grows around the nucleus of the host red cell without displacing the nucleus. Young forms and multipli- cative stages of H. columhce have been found in leucocytes in the lungs of the pigeon (fig. 76, 8-12). Male and female forms (gameto- cytes) are seen in the blood (fig. 76, ja, jb). The cytoplasm of the ^52 THE ANIMAL PARASITES OF MAN male gametocytes is pale-staining and the nucleus is elongate, while the cytoplasm of the females is darker and the nucleus is smaller and round. Formation of male gametes from male gametocytes (the so-called process of '* exflagellation ") may occur on a slide of drawn infected blood, also fertilization, and formation of the ookinete, as first seen by MacCallum. The correct generic name for Halteridia is, apparently, Hcumoproteiis. Wasielewski (191 3), working /2. Fig. 76 — Hcemoproteus {Halteriditim) colunibce. Life-cycle diagram : i, 2, stages in red blood corpuscle of bird; 3, 4, gametocytes (3a S , 3b ?); 5^, formation of microgametes ; 6, fertilization (in fly's gut); 7, ookinete; 8-12, stages in mononuclear leucocytes in lungs. (After Aragao.) on H, danilewskyi (var. falcoiiis), in kestrels, finds that the halter- idium may be pathogenic to nestlings. The cycle of H. noctucv described by Schaudinn (1904) lacks confirmation. The account of the life-cycle of H. colurnhce given by Aragao (1908) is illustrated in fig. 76. It agrees with the work of Sergent (1906-7) and Gonder (1915). Mrs. Adie (19 15) states that the cycle in Lynchia is like that of a Plasmodium. (3) The Leucocytozoon type. The trophozoites and gametocytes occur within mononuclear leucocytes and young red cells (erythro- H^MOSPORIDIA 153 blasts) in the blood of birds. Laveran and Franga consider that the Leucocytozoa occur in erythrocytes. The host cells are often greatly altered by the parasites, becoming hypertrophied and the ends usually drawn into horn-like processes (fig. 77), though some remain rounded. Leucocytozoa are limited to birds, and very rarely produce pigment. Male and female forms (gametocytes) are distinguishable in the blood (hg. 77), and the formation of male gametes (^^ex- fiagellation ") may occur in drawn blood. The Leucocytozoa were first seen by Danilewsky in 1884. They are usually oval or spherical. It is not easy sometimes to distinguish the altered host cell from the parasite, as the nucleus of the former is pushed to one side by the leucocytozoon. The cyto- plasm of the female parasite stains deeply, and the nucleus is rather small, containing a karyosome. In the male the cytoplasm stains lightly and the nucleus is larger, with a loose, granular structure. Many species of Leuco- cytozoa are recorded, but schizogony has only been described by Fantham (1910)^ in L. lovati in the spleen of the grouse {Lagoptis scoticus), and by Moldovan^ (191 3) ^^ L. ziejuanni in the internal organs of screech-owls. M. and A. Leger^ (19 14) propose to classify Leuco- cytozoa, provisionally, according as the host cells are fusiform or rounded. (4) The Hcemogregarina type. Included herein are many parasites of red blood corpuscles, with a few (the leucocytogregarines) parasitic in the white cells of certain mammals and a few birds. They are not amoeboid but gregarine-like, vermicular or sausage-shaped (figc 7cS). They do not produce pigment. They are widely distributed among the vertebrata, but are most numerous in cold-blooded vertebrates (fishes, amphibia and reptiles). The haemogregarines of aquatic hosts are transmitted by leeches, those of terrestrial hosts by arthropods. Fig. 77. — Leucocytozoon lovati. a, Male parasite (microgametocyte), within host cell, whose ends are drawn out ; b, female parasite (macrogametocyte) from blood of grouse. X 1,800. (After Fantham.) ' Annals Trop. Med. and Farasitol., iv, p. 255. 2 Centralbl.f. Bakt., Orig., Ixxi, p. 66. ^ Bull. Soc. Path. Exot., vii, p. 437. 54 THE ANIMAL PARASITES OF MAN The nucleus of haemogregarines is usually near the middle of the parasite, but may be situated nearer one end. The body of the parasite may be lodged in a capsule ('' cytocyst "). There is much variation in size and appearance among haemogregarines. Some are small {Laiikesterella) ; some attack the nucleus of the host cell {Karyo- lysiis) ; others have full grown vermicules larger than the containing host corpuscle, and so the haemogregarines bend on themselves in the form of U (fig. 7^, h). Schizogony often occurs in the internal organs of the host, sometimes in the circulating blood. The haemogregarines occurring in the white cells (mononuclears or polymorphonuclears) of mammals have been referred to a separate genus, Lencocytogregarina (Porter) or Hepatozoon (Miller). Such leucocytogregarines are known in the dog (fig. 79), rat, mouse, palm- squirrel, rabbit, cat, etc. Schizogony of these forms occurs in the internal organs, such as tfie liver, lung and bone-marrow of the hosts. Fig. 78. — Haemogregarines from lizards, a, H. schaudinni, var. africana, from Lacerta ocellata ; b, H. nohrei fiom Lacerta muralis ; c, H. marceaui in cytocyst, from Lacerta muralis. (After Fran9a.) They are apparently transmitted by ectoparasitic arthropods, such as ticks, mites, and lice. A few haemogregarines are known to be parasitic in the red blood corpuscles of mammals. Such are H. gerbilli in the Indian field rat,. Gerhilliis indicus ; H. balfoiiri (jaciili) in the jerboa, Jacuhis jacnlns, and a few species briefly described from marsupials. These parasites do not form pigment. Strict leucocytic gregarines have been described from a few birds by Aragao and by Todd. The sporogony of haemogregarines is only known in a few cases, and in those affinity with the Coccidia is exhibited. In fact, the Haemogregarines are now classified by some authors with the Coccidia. (5) The Babesia or Piroplasma type. These are small parasites of red blood corpuscles of mammals. They do not produce pigment. They are pear-shaped, round or amoeboid in Babesia, bacilliform H^MOSPOKIDIA 155 and oval in other forms referred to this group. Piroplasms are transmitted by ticks. These parasites are described at length on p. 172. Fig. 79. — Leucocytop-egarina canis. Life-cycle diagram. Constructed from drawings by- Christophers. (After Castellani and Chalmers.) Schizogony occurs in the bone-marrow. The parasite is transmitted from dog to dog by the tick, Rhipicephalus sanguineus, development in which, so far as known, is shown on the right. THE MALARIAL PARASITES OF MAN. Malaria, otherwise known as febris intermittens, chill-fever, ague, marsh fever, paludism, etc., is the name given to a disease of man, which begins with fever. It has been known since ancient times and is distributed over almost all the world, although very unevenly, but does not occur in waterless deserts and the Polar regions. In many places, especially in the civilized countries of Central Europe, the disease is extinct or occurs only sporadically, and large tracts of land have become free from malaria. The rhythmical course of the fever is characteristic. It begins apparently suddenly with chilliness or typical shivering, whilst the temperature of the body rises, the pulse becomes low and tense and the number of beats of the pulse increases considerably. After half to two hours the heat stage begins. The patient himself feels the rise of his temperature (shown by feeling of heat, dry tongue, headache, 156 THE ANIMAL PARASITES OF MAN thirst). The temperature may reach 41° C or more. At the same time there is sensitiveness in the region of the spleen and enlargement of that organ. After four to six hours an improvement takes place, and with profuse perspiration the body temperature falls rapidly, not often below normal. After the attack the patient feels languid, but otherwise well until certain prodromal symptoms (heaviness in the body, headache) which were not noticed at first, denote the approach of another attack of fever, which proceeds in the same way. The intervals between the attacks are of varying length which permit of a distinction in the kinds of fever. If the attacks intermit one day, occurring on the first, third and fifth days of the illness and always at the same time of day, it is termed febris tertiana ; if two days occur between fever days, it is called febris quartana. In the case of the fever recurring daily, later writers speak of typical febris quotidiana. But a quotidian fever may arise when two tertian fevers differing by about twenty-four hours exist at the same time (febris tertiana duplex). The patient has a daily attack, but the fever of the first, third and fifth days differs in some point (hour of occurrence, height of temperature, duration of cold or hot stage) from the fever of the second, fourth and sixth days. Similarly, two or three quartan fevers which differ by about twenty-four hours each may be observed together {febris quartana duplex or triplex) ; in the latter case the result is also a quotidian fever. Two kinds of tertian fever are dilfefentiated — a milder form occurring especially in the spring (spring tertian fever), and a more severe form appearing in the summer and autumn in warmer districts, especially in the tropics {suninier or autumn fever, febris cestivo-autumnaUSy febris tropica, febris perniciosa). The latter often becomes a quotidian fever. All the afore-mentioned infections are termed acute. They' are distinguished from the very different chronic malarial infection by the frequent occurrence of relapses, which finally lead to changes of some organs and particularly of the blood. The relapses are then generally marked by an irregular course of fever. The term masked malaria is used when any disturbance of the state of health of a periodic character shows itself and disappears after treatment with quinine.^ Generally it is a question of neuralgia. That intermittent fever was an infectious disease, although not one which was transmitted direct from man to man, had been assumed for a long time. Therefore it was natural, at a time when bacteriology was triumphing, to look for a living agent causmg infection in malaria, which search was, seemingly, successful (Klebs, Tomasi-Crudeli, 1879). Hence it was not surprising that the discovery of the 1 Quinine is still almost exclusively the remedy used in the treatment of malaria. It is prepared from the bark of the cinchona tree. This important remedy was introduced into Europe in 1640 from Ecuador by Juan del Vego, physician of ihe Countess del Cinchon. THE MALARIAL PARASITES OF MAN I 5/ real malarial parasites in November, 1880, by the military doctor A. Laveran^ in Constantine (Algeria), at first met with violent opposition, even after Richard (1882) had confirmed it and Marchiafava, Celli, Grassi and others, had further extended it. Not that the existence of structures found in the blood of malaria patients by Laveran and Richard was denied ; on the contrary, the investiga- tions of the opponents furnished many valuable discoveries, but the organisms were differently interpreted and considered to be degeneration products of red blood cor- puscles. Only when Marchiafava and Celli (1885) saw movements in the parasites, wiiich Laveran called Oscillaria malarice and later Hcematozoon malarice^ was their animal nature admitted and the parasites were r\7kx^^di Plasmodium malaricE. Shortly before this, Gerhardt (1884) had stated that the disease could be transmitted by the injection of the blood of a malarial patient to a healthy person. This supplied the starting point for further investigations, which were made not exclusively, but principally, by Italian investigators (Golgi, Marchiafava and Celli, Bignami and Bastianelli, Grassi and Feletti, Mannaberg, Romanowsky, Osier, Thayer and others). In 1885 Golgi described the asexual cycle in the blood, in the case of the quartan parasite. These investigations, after attention had been drawn by Danilewsky (1890) to the occurrence of similar endoglobular parasites in birds, were extended to the latter (Grassi and P'eletti, Celli and Sanfelice^Kruse, Labbe and others). The result was as follows : Malaria in man (and birds) is the result of peculiar parasites included in the Sporozoa by Metchnikoff, which parasites live in the erythro- cytes, grow in size and finally " sporulate," that is, separate into a number of " spores " which leave the erythrocytes and infect other blood corpuscles. Morphologically and biologically several species (and respectively several varieties) of malarial parasites may be distinguished, on which the different intermittent forms depend. Trans- mission of the blood of patients to healthy people produces a malarial affection which corresponds in character to the fever of the patient from whom the inoculation was made. The combined types of fever (tertiana duplex, quartana duplex or triplex) are explained by the fact that the patient harbours two or three groups of parasites which differ in their development by about twenty-four hours, whilst the irregular fevers depend on deviation from the typical course of development of the parasites. In addition to stages of the parasites which could easily be arranged in a developmental series concurrent with the course of the disease, other phases of the parasites also became known, such as spheres, crescents, polymitus forms, which seemed not to be included in the series and, therefore, were very differently interpreted. The decision reached at the beginning of the last decade of the last century, which found expression in comprehensive statements (Mannaberg, Ziemann and others), only concerned a part of the complete development of the malarial parasites. No one could with any degree of certainty demonstrate how man became infected, nor were there reliable hypotheses based on analogy with other parasites concerning the exit of the excitants of malaria from the infected person and their further behaviour. Numerous hypotheses had been advanced, but none was able to elucidate the ' The discovery of Laveran is in no way lessened by the fact that one investigator or another (according to Blanchard {^Arch. de Paras., vii, 1903, p. 152], P. F. H. Klencke in 1843) had seen, mentioned and depicted malarial parasites. {Neiie phys. Abhandl, auf. selbstdnd. Beob. gegr., Leipzig, 1843, p. 163, fig. 25). In 1847 Meckel had recognized that the dark colour of the organs in persons dead of malaria was due to pigment. Virchow in 1848 stated that this pigment occurred in blood cells. Kelsch in 1875 recognized the frequency of melaniferous leucocytes in the blood of malarial patients. Beauperthuy (1853) noticed that in Guadeloupe there was no malaria at altitudes where there were no '* insectes tipulaires," and suggested that the disease was inoculated by insects. 158 THE ANIMAL PARASITES OF MAN various observations made from time to time in dealing with malaria. One hypo- thesis only seemed to have a better foundation. Manson (1894), who knew from his own experience the part played by mosquitoes in the development of Filaria from the blood of man, applied this also to the malarial parasites living in the blood, whereby at least the way was indicated by which the Haemosporidia could leave man. The parasites were said finally to get into water through mosquitoes which had sucked the blood of malarial patients, and the germ spread thence to men who drank the water. In some cases the parasites were supposed to reach man by the inhaling of the dust of dried marshes. On the other hand, Bignami believed that the mosquitoes were infected in the open air by malarial parasites which occurred there in an unknown stage and the insects transmitted the germs to man when biting. R. Koch combined both hypotheses, without, however, producing positive proof. R. Ross, then (1897-8) an English military doctor in India, was the first to succeed in this. He had been encouraged by Manson to study the fate of malarial Plasmodia which had entered the intestine of mosquitoes with malaria-infected blood, especially in the case of the Plasmodium {Proteosojna) living in the blood of birds. He showed that the Proteo- soma penetrate the intestmal wall of the mosquitoes, grow and develop into large cysts which produce innumerable rod-like germs, which burst into the body cavity and penetrate the salivary glands. Ross allowed mosquitoes to suck the blood of birds affected by malaria, and some nine days later, let the infected mosquitoes which had been isolated suck healthy birds. After five to nine days Proteosoma were found to occur in the blood of the birds used. The Proteosoma and Halteridium of birds were also further investigated by MacCallum (1897-8), Koch and others, and important results followed. In any case Ross (1898) had clearly established the importance of mosquitoes in the spread of malaria among birds. It was now only a question of proving whether, and how far, mosquitoes were concerned with human malaria. Ross himself worked to this end. Here the experiments of Italian investigators (Bignami, Bastianelli, Grassi)^ were of importance. These investigators studied the fate of malarial parasites in man, produced malaria in men experimentally by the bites of infected mosquitoes, and established that only mosquitoes belonging to the genus Anopheles were concerned, and not species of Culex. These laiter are only able to transmit Proteoso7na to birds. It is true that Culex can ingest the human malarial parasites, but the latter do not develop in them. Development only occurs in species of Anopheles. In Anopheles (and similarly for Proteosoma in Culex) sexual repro- duction takes place ; crescents, spheres and polymitus forms are necessary stages of development in the mosquito. With these discoveries the campaign against malaria became more definite. It was directed partly against the transmitters, whose biology and life-cycle were more thoroughly investigated, instead of merely against the infection of the adult Anopheles. The latter do not, as was believed for some time, transmit the malarial germs to their offspring. They always infect themselves from human beings, whereby the relapses appearing in early summer, and the latent infection, especially of children of natives, play a principal part (Stephens and Christophers, Koch). Further, the crusade was directed against the infection of man by the bites of Anopheles. Important results have been obtained in these directions. Low and Sambon in 1900 lived in a mosquito-screened hut in a malarial part of the Roman Campagna for three of the most malarious months and did not contract the disease. In the same year Dr. P. T. Manson was infected with malaria by infected mosquitoes sent from Italy. The role of mosquitoes having been proved, it may be hoped that ultimately the eradication of malaria, or at least a considerable restriction of it, will be achieved. * Grassi, B. (1901), " Die Malaria," 250 pp., 8 plates. G- Fischer, Jena. DEVELOPMENT OF THE MALARIAL PARASITES OF MAN 1 59 It is of importance to record that, although malarial parasites occur in mammals (monkeys, bats, etc.) the human ones are not transmissible to mammals, not even to monkeys. The species, therefore, are specific to the different hosts (Dionisi, Kossel, Ziemann, Vassall). An important work dealing with the modern applications of the mosquito-malaria theory in all parts of the Tropics was published by Sir Ronald Ross in 191 1. It is entitled "The Prevention of Malaria" (John Murray, London, 21s.). DEVELOPMENT OF THE MALARIAL PARASITES OF MAN. The commencement of the developmental cycle and of the infection of man, is the sporozoites (fig. 80, /) which are passed into the blood of a person by the bite of an infected mosquito. Prior to this the para- sites collect in the excretory ducts of the salivary glands (fig. 80, 27) of the Anopheles. The sporozoites are elongate and spindle-shaped, 10 fx to 20 fi long and 1 //, to 2 //, broad, v^'ith an oval nucleus situated in the middle. They are able to glide, perform peristaltic contractions, or curve laterally. Schaudinn has studied the penetration of the red blood corpuscles (fig. 80, 2) by the sporozoites in the case of the living tertian parasite. The process takes forty to sixty minutes in drawn blood. After its entrance the parasite, which is now called a trophozoite, contracts, and becomes an active amoebula (fig. 80, 5). It develops a food vacuole and grows at the expense of the invaded blood corpuscle (fig. 80, 4), which is shown by the appearance of pigment granules (transformed haemoglobin) in it. When the maxi- mum size is attained, multiplication by schizogony (fig. 80, ^-y) begins with a division of the nucleus, which is followed by further divisions of the daughter nuclei, the number of which varies up to 16 or even 32, depending on the species of the parasite. Then the cytoplasm divides into as many portions as there are nuclei, the result being a structure suggestive of the spokes of a wheel or of a daisy, the centre of the resulting rosette being occupied by dark pigment. Finally, the parts separate from one another, leaving behind a residual body contain- ing the pigment, and the daughter forms issue into the blood plasma as merozoites (fig. 80, 7). They are actively amoeboid (fig. 80, 8) and soon begin to enter other blood corpuscles of their host, for the entry into which thirty to sixty minutes are necessary, according to Schaudinn's observations. ^ Here they behave like sporozoites which previously entered and ' It should be remembered that some authors (Laveran, Argutinsky, Panichi, Serra) argue against the intra-globular position of malarial parasites and state that they only adhere out- wardly to the red blood corpuscles. These views have recently been revived by Mary Rowley- Lawson, and she states that the malarial parasite is " extracellular throughout its life-cycle and migrates from red corpuscle to red corpuscle destroying each before it abandons it." {/ourn. Exper. Med.^ 1914, xix, p. 531.) 6 3 ff)"^^fi' "D^'^^'-wf 7 V Fig. 8o. — Life-cycle of the tertian parasite [Plasmodium vivax). Figs, i to 17, X 1,200; figs. 18 to 27, X 600. (After Llihe, based on figures by Schaudinn and Grassi.) i, sporozoite ; 2, entrance of the sporozoite into a red blood corpuscle ; 3, 4, growth of the parasite, now DEVELOPMENT OF THE MALARIAL PARASITES OF MAN l6l again produce merozoites. This process is repeated until the number of parasites is so large that, at the next migration of the merozoites, the body of the person infected reacts with an attack of fever/ which is repeated with the occurrence of the next or following generations. The growth and schizogony last different times, according to the species of the parasite, about forty-eight hours in the case of the parasite of febris tertiana or tropica, and seventy-two hours for the quartan parasite. The various intermittent forms produced by them depend on this specific difference in the malarial parasites. The schizogony can, however, only be repeated a certain number of times, supposing that the disease has not been checked prematurely by the administration of quinine, which is able to kill the parasites. It appears that after a number of attacks of fever the conditions of existence in man are unfavourable for the malarial parasites, and this brings about the production of other forms which have long been known, but also long misunderstood (spheres, crescents, polymitus). The merozoites in this case no longer grow into schizonts, or at least not all of them, but become sexual individuals called gametocytes (fig. 80, 9 — 12), which only start their further development when they have reached the intestine of Anopheles. This does not take place in every case, nor with all the gametocytes which exist in the blood of patients with intermittent fever. Of those parasites which remain in the human blood the male ones (microgametocytes) soon perish, the females (macrogametocytes) persist for some long time, and per- haps at last acquire the capacity of increasing by schizogony. They might thus form merozoites which behave in the body as if they had proceeded from ordinary schizonts (fig. 80, 13c — i^c). If their number increases sufficiently, in course of time the patient, who was apparently recovering, has a new series of fever attacks, or relapses, without there having been a new infection. This is the view of Schaudinn, who from researches of his own concluded that relapses were brought about by a sort of parthenogenetic reproduction of macrogametocytes. R. Ross, on the contrary, believes that in the sometimes called a trophozoite ; 5, 6, nuclear division in schizont ; 7, free merozoites ; 8, the merozoites which have developed making their way into blood corpuscles, (arrow pointing to the left) and increase by schizogony (3 — 7) ; after some duration of disease the sexual individuals appear ; ga — 12a, macrogametocytes ; gd — 12(5, microgametocytes, both still in the blood-vessels of man. If macrogametocytes (12a) do not get into the intestine of Anopheles they may perhaps increase parthenogenetically according to Schaudinn (12a; 13^ — 17^). The merozoites which have arisen (I7(:) become schizonts 3 — 7. The phases shown underneath th^ dotted line (13 — 17) proceed in the stomach of Anopheles. I'^^b and 14/^, formation of microgametes ; 13a and 14a, maturation of the macrogametes ; 15*^, microgamete ; 16, fertilization ; 17, ookinete; 18, ookinete in the walls of the stomach ; 19, penetration of the epithelium of the stomach ; 20 — 25, stages of sporogony on the outer surface of the intestinal wall ; 26, migration of the sporozoites to the salivary gland ; 27, salivary gland with sporozoites. * The incubation period, that is, the time between infection and the first attack of fever, is ten to fourteen days ; with severe infection fewer days (minimum 5 to 6) are needed. II l62 THE ANIMAL PARASITES OF MAN relatively healthy periods the number of parasites in the blood falls below that necessary to provoke febrile symptoms ; relapses then result merely from increase in the numbers of the parasites present in the individual. Ross's view is now generally accepted. If the gametocytes, which are globular, or in the pernicious or malignant tertian parasite crescentic (fig. 8i), gain access to the " I © (J © <7^ a h c d e Fig. 8i. — Stages of development of pernicious or malignant tertian parasites in the intestine of Anopheles maculipennis. (After Grassi.) a, macrogametocyte (crescent) still attached to human blood corpuscles; b, macrogametocyte (sphere) half an hour after ingestion by the mosquito ; Apiosojna^ Wandolleck, 1895 ' Piro- plasma, W. H. Patton, 1895 ; AmcEbosporidium^ Bonome, 1895. The organisms belonging to this genus are pyriform, round or amoeboid. The characteristic mode of division is as follows : Just before division the parasite becomes amoeboid and irregular in shape, (fig. 91, I — 5) with a compact nucleus. The latter gives off a nuclear bud. This nuclear bud divides into two by forking (fig. 91, 6, 7) The chromatin forks grow towards the surface of the body of the rounded parasite, and then two cytoplasmic buds grow out. The forking nuclear buds, which are Y-shaped, pass into the cytoplasmic outgrowths^ (fig. 91, S, 9). The buds gradually increase in size at • Arch. Inst. Bact. Camara Pestatta, iii, p. ii. '^ Parasitology, v (IQ12), p. 65. ^ Nuttall and Graham-Smith, Journ. Hyg., vii, p. 232. BABESIA 175 showing the usual mode of multiplication of Piroplasma canis in the circulating blood. 15 Fig. 91. — Babesia {Piroplasma) canis, life-cycle in stained preparations of infected blood of dog. (After Nuttall and Graham-Smith.) 176 THE ANIMAL PARASITES OF MAN the expense of the parent form until they become two pear-shaped parasites joined at their pointed ends. The connecting strand shrinks and the two daughter forms separate (fig. 91, 10 — 14). The pyriform parasites after having exhausted the blood corpuscle escape from it (fig. 91, 15), and seek out fresh host corpuscles, entering by the rounded, blunt end (fig. 91, i). It is the pyriform phase of the parasite which penetrates red blood corpuscles, not rounded forms, which die if set free. The pyriform parasite, however, becomes rounded (fig. 91, 2, 3), soon after its entry into a fresh host cell. This interesting mode of division by gemmation and chromatin forking has been made dia- gnostic of the genus Babesia by Nuttall.^ Rounded forms of Babesia divide by binary fission, and this direct method can also be adopted by the other forms of Babesia. The distribution of the chromatin in the pear-shaped Babesia, as seen in B. cams and B. bovis, is interesting. The main nuclear body consists of a karyosome surrounded by a clear area. There is also a loose (chromidial) mass of chromatin representing the remains of the chromatin forks seen during the formation of the parasite as a daughter form by gemmation. Occasionally there is a small dot or point, the so-called " blepharoplast" of Schaudinn and Liihe. This minute dot is not a flagellate blepharoplast, for there is no flagellate stage in the life-history of Babesia. These nuclear phenomena have been described by Nuttall and Graham- Smith and Christophers (1907)^ for B. cants, by Fantham (1907)^ for B. bovis, and by Thomson and Fantham (1913) from glucose-blood cultures of ^. cams. Babesia are tick borne, as was first shown by Smith and Kilborne (1893). The developmental cycle in the tickis in completely known. The best accounts are those of Christophers (1907)'* for B. cams and Koch (1906) for B. bovis, and these accounts are supplementary. The principal stages, so far as known, may be summarized thus : — (i) The piroplasms taken by the tick in feeding on blood pass into the tick's stomach. The pyriform parasites, which alone are capable of further development, are set free from the blood corpuscles. In about twelve to eighteen hours they become amoeboid, sending out long, stiff, slender, pointed pseudopodia. The nucleus of each parasite divides unequally into two. Similar forms have been obtained in cultures. These stellate forms may be gametes, and according to Koch fuse in pairs. (2) A spherical stage follows, possibly representing the zygote. This grows, and a uninucleate globular mass results. This form is found in large numbers on the third day, according to the observations of Koch. (3) A club-shaped organism is next formed. This may represent an ookinete stage. The club-shaped bodies are motile and gregarine-like, and are about four times the size of the blood forms. These club-shaped bodies and subsequent stages were described by Christophers in the development of B. cams in the dog-tick, Rhipicephalus sanguineus. ' " Piroplasmosis," Herter Lectures, Parasitology, vi, p, 302. "^ Sci. Mems. Govt. India, No. 29. 3 Quart. Journ. Microsc. Sci., li, p. 297. * Sci. Mems. Govt. India, No. 29. BABESIA 177 (4) The club-shaped bodies pass from the gut of the tick into the ovary, and so get into the ova. There they become globular, and later are found in the cells of the developing tick-embryo. The parasites are, then, transmitted hereditarily. Similar globular bodies are found in the tissue cells of the body of tick nymphs which have taken up piroplasms. The globular stage was called the "zygote" by Christophers, but it may correspond to the oocyst of Plasmodia. (5) The globular body divides into a number of "sporoblasts," which become scattered through the tissues of the larval or nymphal tick, as the case may be. (6) The sporoblasts themselves divide into a large number of sporozoites, which are small uninucleate bodies, somewhat resembling blood piroplasms. The sporo- zoites collect in the salivary glands of the tick. They are inoculated into the vertebrate when the tick next feeds. The chief species of Babesia and their pathogenic importance may be listed thus : — (i) Babesia bovis (Babes) produces infectious haemoglobinuria of cattle in Europe and North Africa. It is transmitted by Ixodes riciiiits. A similar parasite also occurs in deer. (2) Babesia bigeniina (Smith and Kilborne) produces Texas fever, tristeza, or red-water in cattle in North and South America, South Africa and Australia. It is transmitted by Bodphilns annulatiis in North America, by B. austral is in Australia, South America, and the Philippines, and by B. decoloratus in South Africa. The parasite is from 2 /a to 4 /x long, and from i'5 yu, to 2 yu, broad. Babesia bigemina may be the same parasite as B. bovis. (3) Babesia divergens (MacFadyean and Stockman) is a small parasite. It is found in cattle suffering from red-water in Norway, Germany, Russia, Hungary, Ireland, Finland, and France, and is transmitted by Ixodes ricinus. (4) Babesia canis (Plana and Galli-Valerio) gives rise to malignant jaundice or infectious icterus in dogs in Southern Europe, India, and other parts of Asia and North Africa, where it is transmitted by Rhipicephalus sanguineus. In Africa generally, especially South Africa, the disease is transmitted by Hceinaphysalis leacJii. Babesia canis varies from 07 /a to 5 //-, the size depending partly on the number of parasites w^ithin the corpuscle. It averages about 3 //,. It has been cultivated in Bass' medium (glucose and infected blood), see p. 172. In India Piroplasma gibsoni (Patton) infects hunt dogs and jackals. It is annular or oval in shape. (5) Babesia ovis (Babes) produces ^' Carceag," a disease of sheep in Roumania, the Balkan Peninsula, Italy, and Transcaucasia. It varies in size from i />«, to 3 //,. It is transmitted by Rhipicephalus bursa. The parasite has recently been recorded from Rhodesia. (6) Babesia caballi (Nuttall and Strickland) causes *' biliary fever" in equines. The parasite occurs in Russia, Roumania, and 12 iy8 THE PARASITES OF MAN Transcaucasia. It varies in size from j /jl to 2 fi. It is transmitted by Dermacentor reticulaius. It should be mentioned that Nuttallia equi also causes "piroplasmosis" in equines, with symptoms of hasmo^lobinuria and jaundice in Italy, Sardinia, many parts of Africa, Tianscaucasia, India, and Brazil. In Africa it is transmitted by Rhipicephalus evertsi. It has been shown experimentally that a horse recovered from Babesia caballi was susceptible to the inoculatit)n of Nuttallia equi blood. (7) Babesia pitheci (P. H. Ross) was found in a monkey, Cercopithecus sp., in Uganda. The pear-shaped forms measure 1*5 fi by 2*5 fjb. (8) Babesia inuris (Fantham)^ was found in w^hite rats. The pyriform parasites are 2 fi to ^ /jl long and i ft to i'5 fju broad; oval forms are 0-5 to 1*5 fi diameter. The usual symptoms of babesiasis (piroplasmosis) are high fever, loss of appetite, haemoglobin iiria, icterus, anaemia, paralysis, and death in about a week in acute cases. In chronic cases there is anaemia, and haemoglobinuria is less marked. When animals recover, there are still some piroplasms left in the blood. " Recovered " or *' salted" animals are not susceptible to reinfection, but ticks feeding on them acquire piroplasms, and are a source of danger to freshly imported animals. Treatment. — Trypan-blue is the best drug, as shown by Nut tall and Had wen* (1909). It should be administered intravenously in i to 1*5 per cent, aqueous solu- tion. A dose of 5 to 10 c.c. is curative for dogs, one of 100 to 150 c.c. for horses and cattle. Unfortunately, the tissues are coloured blue by the drug. The "salted" animals, after trypan-blue treatment, still harbour the parasites in their blood for years. Genus. Theileria, Bettencourt, Franqa and Borges, 1907. The organisms belonging to this genus are rod-like or bacilliform, and coccoid or round. The best known of the species of Theileria is T. parva^ the pathogenic agent of East Coast fever or Rhodesian fever in cattle in Africa. Theileria parva, Theiler, 1903. Syn. : Piroplasma farvum. In the blood corpuscles of infected cattle minute rod-like and oval parasites are seen. Some are comma siiaped and otht-rsare clubbed (fig. 92, 1-12). The rod-like forms measure i /^ to 3 /^ in length by 0*5 yu< in breadth; the oval forms are 0*7 //, to 1-5 /a in diameter. The intracorpuscular parasites are said by R. Gonder {1910) to be gametocytes, the rod-like forms being thought to be males, the oval forms to be females. Free parasites are practi- cally never seen in the blood. It is known that it is impossible to ' Quart, fourn. Microsc. Sci., 1, p. 493. 2 Parasitology, ii, p. 156. THEILERIA PARVA 179 produce the disease in a healthy animal by blood inoculation, but only by intraperitoneal transplantation of large pieces of infected spleen (Meyer). There may be as many as eight parasites in a cor- puscle. The chromatin is usually at one end of the organism. In some parasites the appearance of the chromatin suggests division, but such division, if it takes place, must be very slow, as it has not been actually seen in progress. The red blood corpuscles appear merely to act as vehicles for the parasites (Nuttall, Fantham, and Porter).^ 00© 14 16 Fig. 92. — Theileria parva. 1-12, intracorpuscular parasites, stained. (After Nuttall and Famham) ; 13-18, Koch's blue bodies, from stained spleen smear; 17-18, breaking up of Koch's body. (After Nuttall.) In the internal organs, especially the lymphatic glands, spleen and bone-marrow, are found multinucleate bodies known as Koch's blue bodies (fig. 92, 13-18). These are schizonts, according to Gonder.^ The actual Koch's blue bodies are said to be extracellular, but similar multinucleate bodies, schizonts, occur in lymphocytes. The schizonts divide and the merozoites resulting probably invade the red blood cor- puscles in the internal organs. Gonder considers that the sporozoites injected by the tick collect in the spleen and lymphatic glands, penetrate the lymphocytes and give rise to the schizonts. Gonder has studied the cycle of 7 . parva in the tick. He states that the gametocytes leave the host corpuscles and give rise to gametes, then conjugation occurs producing zygotes. The zygotes are then said to become active lo form ookinetes, and to enter the salivary glands of the tick. Multiplication is said to occur therein, producing a swarm of sporozoites. This work needs confirmation. T. parva is transmitted by Rhipicephalus appendiculatus^ R. simus, R. evertsi^ R. m/ens, and R. capensis. The parasites are not hereditarily transmitted in Rhipicephalus^ but when taken by the transmitter at one stage of its development the tick is infective in its next stage (t or Miescher's lube. a, sub- stance of muscle fibre ; b, envelope of saro'cysi ; <:, mus- cle nuclei ; r/, sports in cham- bers ; ^, ground subs'ai)ce. X 400. (From Wasielcwski, after van Eecke.) SARCOSPORIDIA 191 Laveran and Mesnil) contain an obliquely striated body (fig. 109) often homologized with the polar capsule, while the greater part of the spore is taken up by the nucleate sporozoite. Several authors state that they have also observed filamentous appendages (polar filaments) at one end of the spores, and have seen two kinds of spores in the same Sarcosporidium. Spores of various species of Sarcosporidia may contain metachromatic granules, often centrally placed (fig. 109). These granules may be metabolic or possibly may contain toxin (see below). The gymnospores of Sairocystis jiiuris, fr om the mouse, show active boring movements when kept in saline solution warmed to 35° or 37° C. S. inuris is very deadly to its host. From their structure the spores do not appear to have great powers of resistance to external conditions. They measure 1 2 yu, by 3 ytt to 4 />«- or less. Laveran and Mesnil (1899) isolated a toxin from S. tenella of the sheep and called it sarcocystin. This substance is especially pathogenic to experimental rabbits. The duration of life of the Sarcosporidia is a comparatively long one. The affected muscular fibres may remain intact and capable of performing their functions for a long time, but at last they perish, if the host lives long enough. Thus the Sarcosporidia of the muscles are then enveloped only by sarco- lemma, and finally, when this likewise dis- appears, they fall into the intra-muscular connective tissue. In many cases the Sarco- ^ ^ sporidia die off within their hosts, this, Fig. 109.— Spores of .Sar^^?- according to Bertram, being brought about ^otnTt' H- c^a.S; by a disintegration of the spores in the cen- b, stained, showing meta- tral chambers. In other cases the leucocytes ^f,^'''^'^ ^'^,^^;'. ""(Vt"; play a part in the destruction of the Sarco- Laveran and Mesnil.) sporidia, and sometimes it happens that lime salts are deposited in and around the vacant cylinders. In some places pigs, sheep, mice and rats are infected with sarcosporidiosis to a remarkable extent, in certain cases almost reach- ing 100 per cent. Young animals also are infected, and perhaps infection only takes place during youth. Although the natural mode of transmission of the Sarcosporidia remains to be determined, yet various experimental researches on the problem are of interest and importance. Theobald Smith (1901) found that mice could be experimentally infected with S. miiris by feeding them with the flesh of other infected mice. The incuba- tion period was a long one, namely forty to fifty days. Thus, on the forty-hfth day after feeding young Sarcosporidia were found, and seventy days after feeding spore formation began. Ripe spores were found two and a half to three months after the commencement of 192 THE ANIMAL PARASITES OF MAN these experiments. This mode of infection — a cannibahstic one — hardly seems likely to be the natural method for the infection of sheep and ruminants generally. Smith's researches have been confirmed. Negre' (1910) found that the faeces of mice fed on infected muscular tissue were infective to other mice when ingested by them. Negri*^ infected guinea-pigs with S. nuirls by feeding them on infected mouse flesh, and found that the parasite in guinea-pigs showed different characters from those exhibited by it in mice. Darling^ also succeeded in infecting guinea-pigs with S. mnrisy and Erdmann infected mice with S. tenella (from the sheep). According to Erdmann* (iqio) the Sarcosporidian spore germinates in the intestine of the host, which has recently ingested infected material. The spore liberates its contained toxin — sarcocystin — which acts upon the adjacent intestinal epithelium, whereby the latter is shed, and an amcebula creeps out of the spore. The amoebula is able to penetrate the denuded area and get directly into the lymph- spaces of the submucous coat of the intestine. The first period of development, lasting some twenty-eight to thirty days, is said to be passed in the lymph-spaces of the intestine. Later the amoebula reaches a muscle fibre. Writing in May, 1914, Erdmann^ records the appearance of small amoeboid and schizogony forms six days after infection of the host. Crawley^ (19 13) controverts some of these statements and considers that the Sarcosporidian spore, still sickle- shaped, bores its way into the epithelial cells of the intestine and comes to rest there. The spore then becomes round or elliptical, and peripheral masses of chromatin appear within it, suggesting schizogony. This happens about twelve hours after feeding, and in twenty-four hours the spores appear to have left the intestine. More recently (May, 1914), Crawley^ considers that there is sexual differentiation among the Sarcosporidian spores, a few hours after their ingestion by the host. Interesting discussions have occurred as to the site of the toxic sarcocystin within the spore. Metachromatic granules occur in the middle of the Sarcosporidian spore (fig. 109), and the toxin may be contained in these grains, as they disappear, according to Erdmann, before the amoebula penetrates the denuded intestinal wall. However, a polar capsule, containing a polar filament, may be present at one end of a Sarcosporidian spore. Laveran and Mesnil described a striated area at the more pointed end of the spore of S. tenella^ which area tliey consider to represent a polar capsule. Fantham^ (1913) ' C. R. Soc. Biol., Ixviii, p. 997. '^ Cefztralbl. f. Bakt., Orig., xlvii, p. 612 ; see also xlvii, p. 56 ; Iv, p. 373. ^Joitrn. Exptl, Med., xii, p. 19. ■» Sitz. Gesell. naturf. Freunde zti Berlin, p, y]-]. ^Proc. Soc. Exper. Biol, and Med., xi, p. 152. ^Science, xxxvii, p. 498. ^ Proc. Acad. Nat. Sci., Philadelphia, May, 1914, p. 432. ^ Proc. Cambr. Philosoph. Soc.y xvii, p. 221. SARCOSPORIDIA I93 found a vacuole-like, polar capsule area in the spores of S. colli from the African mouse-bird. The sarcocystin may be contained in the polar capsule. The nucleus of the spore is generally at the opposite, blunter end. Again, various authors have stated that Sarcosporidian spores may occur in the blood of the host at times. If so, then an intermediate host may be concerned in their transmission. Perrin suggested that Sarcosporidia might be spread by blow-flies and flesh-flies. The classification of the Sarcosporidia as proposed by R. Blanchard, which was based on their various habitats, can no longer hold, because the same species may occur in the muscles as well as in the connective tissue. For the present, the few species that are known maybe placed in one genus, Sarcocystis, Ray Lankester, 1882. The following species of Sarcocystis are of interest : — S. iniescheriana, Kiihn, 1865, in the pig. S. bertrami, Doflein, iQoi, in the horse. S. teiiella, Railliet, 1886, in sheep. S. tenella bubali in buffaloes in Ceylon and Egypt. S. blanchardi, Doflein, 1901, in cattle. S. niuris, Blanchard, 1885, in the mouse, to which it is lethal. S. Iiiieti, Blanchard, 1885, in the seal. S. coin, Fantham, 191 3, in the African mouse-bird, Coliiis eryt/ironieloii. Also various Sarcosporidia from antelopes, monkeys, opossum, birds, the gecko and wall-lizard are known. The spores of S, mnris, S. bertrami, S. tenella, and S. colli can multiply by longitudinal fission. Sarcosporidia observed in Man. (i) Lindemann ^ found on the valves and in the myocardium of a person who had died of dropsy certain brownish masses, 3 mm. in length and 1-5 mm. in breadth which he regarded as gregarines. If these were actually independent animal organisms it may be suggested that they were Sarcosporidia. Rivolta (1878) named the species S. liiidcjuanni. (2) Rosenberg^ found a cyst 5 m_m. in length and 2 mm. in breadth in a papillary muscle of the mitral valve of a woman, aged 40, who had died from pleuritis and endocarditis. The cyst contained no scolex nor booklets of taenia. Numerous small refracting bodies, round, oval or kidney-shaped, were found in a daughter cyst, as well ' ** Ueb. d. hyg. Bdtg. d. Gregarinen," Dtsche. Ztschr. f. Staatsarzneikunde, 1868, xxvi, p. 326. * " Ein Befund von Psorosp. in Herzmusk d. Menschen," Ztschr. f. Hygiene, 1892, xi, P- 435- ^3 194 THE ANIMAL PARASITES OF MAN as sickle-shaped bodies. The description hardly appears to indicate Sarcosporidia. (3) Kartulis^ observed Miescher's cylinders of various sizes in the liver (?) and in the muscular system, of a Sudanese who had succumbed to multiple abscesses of the liver and abdominal muscles. This may be considered as the first actual case of the occurrence of Sarcosporidia in man. Koch in 1887 described a case in Egypt. (4) The case reported by Baraban and St. Remy^ was at once demonstrated as certain. It related to a man who had been executed, and in the laryngeal muscles of whom Sarcosporidia were found ; the length of the parasites varied between 150 //, and 1,600 /a, their breadth between 77 /x and 168 ^. The affected muscular fibres were distended to four times their normal thickness. This species was described by Blanchard as *^ Miescheria " inuris, but according to Vuillemin, it was more probably Sarcocystis tenella of the sheep. (5) Vuillemin has also described a case of Sarcosporidia found in the muscles of a man who died from tubercle at Nancy. The author considered that the parasite corresponded to S. tenella. (6) Darling^ (1909) found Sarcosporidia in the biceps of a negro from Barbados. The Myxosporidia, Microsporidia, Actinomyxidia and possibly the Sarcosporidia may be included within the section Cnidosporidia (Doflein), since they possess spores containing polar capsules. Order. Haplosporidia, Caullery and Mesnil. The Haplosporidia are a group of organisms having both a simple structure and life-history. The simplicity may represent a primitive condition or may be due to degradation resultant on parasitism, and thus it is possible that the group is not a homogeneous one. The order Haplosporidia was created by Caullery and Mesnil in 1899, and includes parasites of rotiifers, annelids (fig. no), Crustacea, fish, pro- chordates and man. They may be present in the body cavity or alimentary tract, and can also occur in the septum nasi of man, in the nervous system of Cephalo- discus, and in tumours of fish. As the name implies, the spores of the Haplosporidia are simple, without polar capsules, and are uninucleate. In some genera, e.^., Haplosportdium, Urospori- dium (fig. Ill) there is a spore-coat or sporocyst which may be elongate or spiny. The developmental cycle of a Haplosporidian, such as Haplosporidium or Bertramia, begins with a small, uninucleate cell, often rounded, possessing a cell membrane that may be prolonged into processes. Growth takes place, coupled with an increase in the number of nuclei, so that a multinucleate trophozoite is produced. Later, this ' Kartulis, *'Ueb. pathog. Protoz. b. Menschen," Ztschr. f. Hyg. u. Inf., 1893, xiii, p. 1. ■Compare also Braun, M., Die Thier. Par. d. Mensch., 2nd Edit., Wrzbg., 1895, p. 92 ; Braun, M., "Z. York. d. Sarcosp., b. Menschen," Centralbl. f. Bakt. 1895. xviii, p. 13. '" Sur un cas de Tub. Psorosp. ob. chez rhomme," C. R. Soc. Biol., Paris, 1894 (x), I, p. 201. " Le Parasitisme d. Sarcosp. chez rhomme," Bibliogr. Anat. 1894, p. 79. ^ Arch. Internal Med., Ill, p. 183. HAPLOSPORIDIA 195 multinucleate trophozoite becomes segmented into a number of ovoid or spherical pansporoblasts, which give rise to few (one to four) spores. Such a spore, when set free, begins the life cycle over again. More recently (1905- 1907) two important organisms have been described and included in this group, namely, Neurosporidium cephalodiscP (Ridewood and Fantham) from the nervous system of the prochordate, Cephalodisciis nigrescens, and Rhi7iosporidiu7n kinealyi (or seeberi) from the septutn nasi of man. In the case , bolus of food. (After Casagrandi and Barbagallo.) pointed according to the degree of contraction (figs. 113, 114). The ecto- and endo-plasm are distinct, the latter is granular, containing drops of fat and mucus, granules of starch, bacteria, and occasionally BALANTIUIUM COLI 20I also red and white blood corpuscles. There are usually two con- tractile vacuoles, seldom more. The anus (cytopyge) opens at the posterior extremity. The macronucleus is bean- or kidney-shaped, rarely oval ; the micronucleus is spherical. Balaiitidium coll lives in the large intestine of man, in the rectum of the domestic pig, and has been found in monkeys. It propagates by transverse division, but conjugation and encystment are known to take place.^ Transmission to other hosts is eft'ected by the cysts of the parasite (fig. 114). Balaiitidinjn coll, first seen by Leeuwenhoek, was described by Malmsten in 1857 in a man aged 35 years, who had two years previously suffered from cholera, and since then had been subject to diarrhoea. The examination showed an ulcer in the rectum above the mid sphincter ani, in the sanguineous purulent secretion of which numerous Balantidia were swimming about. Although the r deer was made to heal, the diarrhoea did not cease and the stools contained numerous Balantidia, the number of which could only be decreased by extensive enemas of hydrochloric acid. The second case related to a woman who was suffering from severe colitis, and who died ten days after admission. The mal- odorous, watery evacuations contained innumerable Balantidia, in addition to pus, and at the autopsy the anterior portion of the large intestine was found to be infested with them. Subsequently this parasite has often been observed in human beings, and various cases have been recorded. These occurred in Russia, Scandinavia, Finland, Cochin China, Italy, Germany, Serbia, Sunda Islands, Philippine Islands, China, and in other parts of Asia and in America. Other cases were reported by Askanazy, Ehrnroth, Klimenko, Nagel, Koslowsky, Kossler, Waljeff, Strong and Musgrave, Glaessner^ and others. Sievers found B. coll very common in Fmland. In the majority of the cases described by Sievers from Finland, and in other cases from Central Europe, the patients suffered from obstinate intestinal catarrh, w^hich did not always cease even after the Balantidia had disappeared. On the other hand, Balantidia have occasionally still been found to persist, though in small numbers, after the catarrh has been cured. Some authors, nevertheless, do not • According to Gourvitch (" Bal. coli. Darmk. d. Menschen," /vnss. Arch. f. Path.^ klin. med. u. Bad., Petrograd, 1896), the conjugated Balantidia are supposed to fuse with each other and form oval cysts two or three times the size of the free organisms, and to divide into numerous globules within the cystic membrane ; the process, however, has hitherto not been confirmed. The supposed Balantidium cysts appeared in two patients who were simultaneously suffering from Dibothriocephahis latus, after the administration of anthelminthics. It therefore seems, according to the description, that in reality these forms were actually abnormally large, possibly swollen, young eggs of the tape-worm mentioned. • Centralbl.f. Rakt., Orig., xlvii, p. 351. 202 THE ANIMAL PARASITES OF MAN regard Balantidia as the primary cause of the various diseases of the large intestine, which often commence with the development of ulcers, but they consider that they may aggravate these diseases and render them obstinate. According to Solowjew, Askanazy, Klimenko and Strong and Musgrave, however, the parasites penetrate the intes- tinal wall, and give rise to ulcerations which may extend deeply into the submucosa, and even be found in the blood and lymphatic vessels of the intestinal wall. According to Stokvis, B. coli occurs also in the lung ; at all events this author states that he found one living and several dead param^cia (?) in the sputum of a soldier, returned from the Sunda Islands, who was suffering from a pulmonary abscess. Sievers has shown that B. coli might occur in persons not suffering from intestinal complaints, but E. L. Walker ^ (19^3) states that every person parasitised with B.. coli is liable sooner or later to develop balantidian dysentery. Since Leuckart confirmed the frequent presence of B. coli in the rectum of pigs, and corresponding observations were made in other countries, the pig is universally considered to be the means of the transmission of Balantidium to man. The encysted stages only serve for transmission, because, according to all observations, the free parasites have a very small power of resistance. They perish when the faeces have become cool ; they cannot live in ordinary, slimy, or salt water. As they are killed by acids even when much diluted, they cannot pass through the normal stomach alive except under the most unusual circumstances. The pigs, in whose intestines the Balantidium appears to cause little or no disturbance, evacuate numerous encysted Balantidia with the faeces, and their occasional transference to man brings about their colonization there, but perhaps only when a disease of the colon already exists. Experimental transmission of the free parasites to animals (per os or per anum) yielded negative results, even in the case of pigs. Casagrandi and Barbagallo (1896), however, had positive, as well as negative, results. They employed healthy young cats, or cats in which catarrhal entero-colitis had been artificially induced (which in other experiments is apt to cause the death of the animals experimented upon in about six or seven days), or finally cats that had dilatation of the rectum with alkaline reaction of the faeces. An attempt to infect three healthy cats by injecting human faeces containing Balantidium into the rectum proved negative, in so far as the faeces of the experimental animals had an acid reaction and contained no Balantidia, but at the autopsy performed eight days after infection a few^ encysted parasites were found in the mucus of the ileum. In the case of four cats ^ Philip. JL Sc, Sec. B, viii, p. 333. BALANTIDIUM COLI 203 suffering from entero-colitis, into which human faeces containing Balantidia were introduced per os, Balantidium cysts were found in the faeces three days after the last ingestion. Great numbers, more- over, w^ere found in the caecum and the posterior part of the small intestine at the autopsy of the animals, which died about eight days after the commencement of the experiments. Actual colonization, therefore, was not effected in either series of experiments. Free or encysted Balantidia of pigs were used for further experiments. The experiments proved negative when faeces containing cysts were injected into the rectum of healthy cats (three experiments), or cats (two) suffering from spontaneous intestinal catarrh, or when such material was introduced per os into three healthy cats. In the case of two cats with intestinal catarrh artificially produced, a small number of the active Balantidia injected into the rectum remained alive. Larger quantities of faeces containing encysted Balantidia were introduced into two other cats affected with the same complaint. These, certainly, did not appear in the faeces, but small numbers, free and alive, were found in the caecum. Similarly, encysted Balantidia were introduced into two cats with dilated rectum, and whose faeces had an alkaline reaction, in these cases no parasites appeared in the faeces, but three and five days later, when the two animals were examined, a very small number were discovered free in the large intestine. Klimenko did not succeed in infection experiments with B. coll on young dogs, whose intestines had been artificially affected by disease. More recent experiments by Brumpt have shown that young sucking pigs can be infected with Balantidium from infected monkeys {Macacns cynoniolgiis) and suffer heavily from the same, whereas the Balantidium of the pig is rarely harmful to its host. This and previous experiments may be thought to suggest that there are perhaps several pathogenic species, and also that harmless strains of Balan- tidium may occur. At the same time, it must be remembered that a large proportion of the cases recorded of Balantidian colitis occur among swineherds and butchers, that is, among people in frequent contact with pigs. Morphologically, there are practically no differ- ences between the Balantidia found in man, monkeys and pigs, and it is probable that one species only, under slightly different environ- mental conditions, may be responsible for the colitis observed. In any case, efficient prophylactic measures should be taken against balantidiasis in countries where it may occur, by confining the pigs and not allowing them to run in yards and dwellings. E. L. Walker (19 13) has given a good summary of work on balantidiasis. His own researches in the Philippines showed that monkeys could be infected by Balantidia both from pigs and men. Parasites may appear in the stools only at infrequent intervals. He 204 THE ANIMAL PARASITES OF MAN believes that the cihates are the prhiiary etiologic factor in the symptoms and lesions of balantidian dysentery. Behrenroth (1913) has given an interesting account of Balantidium coli and its pathogenic significance. Balantidium minutum, Schaudinn, 1899. The body is of oval form, with tlie anterior extremity pointed, and posterior extremity broad and rounded (fig. 115). The length is 20 fi to 32 /jL, and the breadth is 14 /jl to 20 /x. The peristome, which is fissure-like, extends to the centre of the body (fig. 115). The right lateral border of the peristome is beset with cilia the same length as those of the body, the left side terminates in a thin hyaline membrane that extends towards the back, and can pass over to the right side. A row of longer and stronger cilia (cirri) are on the left border of the peristome. The cuticle is refractile, the ectoplasm hyaline and the endoplasm granular, with numerous food vacuoles. A single contractile vacuole lies dor- sally and to one side at the posterior extremity. The macronucleus, which is always spherical, is central and is 6 /x to 7/x in diameter. The micronucleus, close in front of it, only measures i /a (fig. 115). The cysts are oval. These parasites were found in num- bers in the evacuations of a man aged 30, who was born in Germany and had repeatedly travelled between Hamburg and North America, where he made long stays. Tlie patient came to the Charite in Berlin to seek advice for constipation alter- nating WTth diarrhoea accompanied by abdommal pam. A second case (the parasite of which w^as described as Colpoda <:iiciilliis by Schulz) was observed in a patient in the same institution. As, in both cases, the parasites only appeared during the diarrhoea, and disappeared as soon as the faeces had assumed a normal consis- tency, or could only be demonstrated in a few encysted specimens, it may be assumed that the small intestine or the duodenum is their habitat. Fig. 115. — Balantidium minu- itim. P, peristome; N, nucleus; J/, micronucleus ; V, contractile vacuole. Food vacuoles are repre- sented in the endoplasm. (After Schaudinn.) Genus. Nyctotherus, Leidy, 1849. Flat, heterotrichous Infusoria, kidney- or bean-shaped. The peristome com- mences at the anterior pole of the body and extends along the concave side to the NYCTOTHERUS FAB A ^05 middle, where the oral aperture is situated. The cytopharynx is oblique and is more or less curved. The cytopyge is at the posterior extremity, where a single contractile vacuole is also situated. The macronucleus is almost in the centre of the parasite. The members of this genus live parasitically in the intestine of amphibia, insects and myriapods, and at least one species is found in man. Nyctotherus faba, Schaudinn, 1899. The body is bean-shaped, and a Httle flattened dorso-venlrally. It is 26 //, to 28 //, ^ong, 16 yL6 to 18 yLt bi'oad, and 10 //, to 12 /a thick (fig. 116). The peristome is on the right border and extends to the middle ; at the left there are large adoral cilia, the cilia on the right border not being larger than those on the body. The cyto- pharynx is short, slightly curved and turned backwards. The contractile vac- uole is large, spherical, situated at the posterior extremity, and its contents are voided through the anus at its left. The macronucleus is in the centre of the body; it is globular (6 /x to 7 //, in size), and contains four or five chromatin masses. ¥ig. 116.— JVyao/Aerus fada. r, rr^. . , ,. 1 , ., , . peristome; A^. nucleus ; ill/, micro- 1 he micronucleus lies close to it, and is nucleus; r, contractile vacuole. spherical or somewhat elongate measuring (After Schaudinn.) I /A to 1*5 w, (fig. 116). The cysts are oval. This species has hitherto only been seen once in the same patient in whom Balantiduim niinutum was discovered. Nyctotherus giganteus, P. Krause, 1906. Under the name Balaiifidiumgigaiiteiun n. sp., P. Krause described an Infusorian which was repeatedly observed with Trichoiuonas iutes- fiiialis in the alkaline evacuations of a typhoid patient in Breslau. The body is ovoid, narrower and rounded anteriorly and broader and stunted posteriorly. The peristome lies to one side ; the macro- nucleus is bean-shaped, the micronucleus small and globular ; one or two vacuoles are present. The anus is at the farther end. The organism is go /jl to 400 /^ long, 60 fi to 150 fju broad (fig. 117). After a prolonged stay outside the body, it becomes rounded and encystment occurs. In the thermostat the Infusoria remain alive at 37° C. for five weeks. The species, however, hardly belongs to Balautidiiuii, but to all appearances is a Nyctoihents and is distinguished from N. faha by the difference in size. >o6 THE ANIMr^L PAI^ASITES OF MAN [Nyctotherus] afrlcanus, Castellani, 1905. In the fa3ces of a native of Uganda who suffered froni sleeping sickness and diarrhoea and had in his intestine Ascaris liujibricoidcs, Trichoceplialus trichiiinis and Ancylostoiiui diiodenale, Castellani found a curiously shaped Infusorian, 40 /x to 50 /x long, and 35 //, to 40 />t broad, with spherical macro- and micronucleus and a contractile vacuole (tig. 118). He included the organism in the genus Nyctotherus, perhaps wrongly, or the parasite may have been deformed. After the patient's death the same parasite was found in the intestine and especially in the caecum. Fig. WJ.— Nycto- therus giganleiis. (Afier Krause.) Fig. i\%.— Nyctotherus africamis. (After Castel- lani.) G. Lindner, in Cassel, studied certain peritrichal Infusoria (stalkless Vorticella), and connected them, probably incorrectly, with the most varied diseases of man and domestic animals, even with Sarcosporidia of pigs. It may be mentioned that according to a communication by letter from Schaudinn, Vorticella may be found in freshly evacuated faeces, but always only after the administration of a water enema. In spile of this, several other investigators mention Vorticellas as intestinal parasites of man. The Chilodon dentafus (Ehrenberg) recorded in 1903 by J. Guiart as a parasite of man, which may be found in all infusions, can hardly have lived in the man from whose faeces it was cultivated, but may represent a chance admixture both in the faeces and the cultivations. C. uncifiahis was also found as a chance parasite of man by Manson and Sambon. According to Doflein^ (1911) certain Chilodon- like organisms have been found by Selenew in prostate secretions in gonorrhoea. Other species of the genus Chilodon are known, but only as ectoparasites {e.g., Chilodo?i cypriiii, Morofif, 1902, from the skin and gills of diseased carp). A number of other parasitic Ciliates are known, among which Ichthyophthirius jiiultifiliis, destructive to fish, is important. It lives in the skin and the layers immediately below it, forming small whitish pustules which may become confluent. ' Lehrhuch der Frotozoeukiinde, 3rd ed,, p. 963. CHLAMYDOZOA 20/ The pustules are most common on the head and fins, but occur also on the eyes and gills of the host. The young parasite, which is one of many formed in a cyst, is very small. At first it is free swimming, but soon attaches itself to the skin of a fish. It bores inwards and becomes surrounded by the irritated skin. There it attains a relatively large size, being 500 /a to 750 /^ and occasionally more in diameter. The body has a rounded terminal mouth, short cytopharynx and a number of minute contractile vacuoles. The macronucleus is large and horseshoe-shaped ; the small micronucleus is only seen in the very young animal. When full grown, the organism encysts and forces its way to the surface and bursts through, leaving a small, gaping wound behind. The cyst sinks to the bottom of the water, nuclear multiplication occurs and a number of young parasites are produced, which leave the cyst and either attack new hosts or else perish. Opalina ranai'um, parasitic in the rectum and urinary bladder of frogs and toads, shows great degradation and simplification due to parasitism, possessing no separate micronuclei, no cytostome, cytopharynx or cytopyge. It has many macronuclei, and is a large parasite. During summer and autumn nuclear multiplication followed by division of the body occurs, the process being repeated after the daughter forms have grown to the size of their parent. In spring, the Opalina divide rapidly, but do not grow much before dividing again. Finally, tiny forms, containing three to six nuclei, encyst and pass from the host with the fteces. As these latter are greedily devoured by tadpoles, the Opalina gain new hosts in which they develop. THE CHLAMYDOZOA. The name Chlamydozoa was proposed by Prowazek in 1907 for a number of minute, problematic organisms (fig. 119) believed to be the causal agents of certain diseases in man and animals, such as vaccinia and variola, trachoma, inclusion blenorrhoea in infants, molluscum contagiosum, and bird epithelioma contagiosum. Other diseases possibly due to Chlamydozoa ^ are hydrophobia, measles, scarlet fever, foot-and-mouth disease, the " Gelbsucht" disease of silk- worms, and perhaps even typhus (Prowazek, 191 3). The subject is difficult and controversial and can only be briefly discussed here. It is known that the viruses in all these diseases can pass through ordinary bacterial filters, that is, they belong to the group of '' filterable viruses." At such periods the organisms are extracellular or free. It is also known that in many of these cases the virus produces definite and characteristic reaction-products or cell-inclusions in the infected cells, during the intracellular phase of the life-history of the organism. As the organisms to be considered are problematic, it will be con- venient to summarize their history : — (i) Cell-inclusions, usually named after their discoverers, have been found in certain diseases, thus : In vaccinia Guarnieri's bodies, ' For a detailed account of the Chlamydozoa see Prowazek's Handbtich der Pathogenen Frotozoen, Bd. i (191 1- 12). Leipzig, J. A. Barth. 208 THE ANIMAL PARASITES OF MAN in scarlet fever Mallory's bodies, in hydrophobia Negri's bodies, in trachoma Prowazek's bodies occur. (2) At first these characteristic cell-inclusions were considered to be actual parasitic organisms causing the diseases in question. The bodies received zoological names and attempts were made to work out their supposed development cycles. The supposed parasites of vaccinia and variola were referred to a so-called genus Cytorydes, those of hydrophobia to Neuroryctes, of scarlet fever to Cyclasierhim, while those of molluscum contagiosum were referred to the Coccidia. Calkins in 1904 studied in detail the cell-inclusions of vaccinia and small-pox, calling them Cytorydes varlolce, Guarnieri. Calkins con- sidered that in the stratified cells of the epidermis they passed through two cycles, the one cytoplasmic, the other intranuclear. The first is the vaccinia cycle, the second the pathogenic (intranuclear) variola cycle. It is hardly necessary to follow all Calkins' stages here. Negri (1909) described a cycle for Nenrorydes hydrophobia. Calkins refers both Cytorydes variotcc and Neuroryctes hydrophohice to the Rhizopoda. Siegel (1905) described quite different organisms under the name Cvtorhydes. He listed several species : C. vacdnice of vaccinia and small-pox, C. scarlatlnce of scarlet fever, C. luis of syphilis (this is probably the granule stage of Treponema pallidiun), and C. aphtharuju of foot-and-mouth disease. (3) The afore-mentioned views were criticized, and the bodies were not considered to be living organisms but merely reaction products or cell-inclusions due to the effects of the virus on the host cells. Thus Guarnieri's bodies were stated to consist of extruded nucleolar or plastin material, having no developmental cycle. It was further asserted that infection could be produced by lymph in which Guarnieri's bodies had been destroyed. Similar assertions have been made regarding the Negri bodies, and others. The Cytorydes, Neurorydes, etc., are considered, according to these views, to be degeneration products of the nucleus or to be of a mucoid nature. (4) More recently a positive belief has gained ground that there are true parasitic organisms causing these diseases, and that the parasites are very minute, being termed Chlamydozoa by Prowazek and Strongyloplasmata by Lipschlitz. The Chlamydozoa are characterized by {a) their very minute size, smaller than any bacteria, so that they can pass through bacterial filters ; (6) they pass through intracellular stages, in the cytoplasm or the nucleus of the host cell, producing therein the reaction products or inclusions in the cell already recorded as characteristic or diagnostic of the diseases produced; (c) they pass through definite developmental cycles. Such a cycle consists essentially of growth CHLAMYDOZOA 209 and division. The mode of division of the Chlamydozoa resembles that of the centriole of a cell, by the formation of a dumb-bell-shaped figure. Two dots are observed connected by a fine line or strand which becomes drawn out and finally snaps across the middle. Prowazek and Aragao (1909) working on smallpox in Rio de Janeiro found that the chlamydozoal granules passed through a Berkefeld filter and that the filtrate was virulent. But if an "ultra-filter" were used, i.e., one coated with agar, then the granules were retained and the filtrate was no longer virulent. The surface of the ultra-filter was found to contain many granules. The Chlamydozoa are parasites of epiblastic tissues {e.g.y epidermal cells, nerve cells, conjunctival cells). The life-history of a Chlamydozoon (fig. 119), such as that of Fig. 119. — Chlamydozoa. Trachoma bodies in infected epithelial cells of the conjunctiva, (a) initial bodies (above) and cluster of elementary bodies (touching the nucleus) ; {b) cluster of granules surrounded by mantles, x 2,000 approx. (Original. From preparation by Fantham.) vaccinia, is, according to Prowazek, Hartmann and their school, as follows : — 1. The infection begins with elementary bodies or elementary corpuscles which live at first extracellularly. An elementary body is a minute speck of chromatin, apparently devoid of cytoplasm, which can pass through a bacterial filter. It can enter a host cell, but the entry is not a process of phagocytosis. 2. Inside the host cell the elementary body grows in size, and becomes an initial body (fig. 119, a), 3. A reaction on the part of the host cell results, for nucleolar, plastin substance is extruded from the cell-nucleus and surrounds the parasitic initial body. The latter is thus enveloped in a mantle (hence the name Chlamydozoa, from )(Xa/jLv<;, a mantle), and the characteristic cell-inclusion (Guarnieri's body, Negri's body, etc.) is produced. The nucleolar, mantle substance probably represents the ^* cytoplasm " of Cytoryctes, described by Calkins. 210 THE ANIMAL PARASITES OF MAN 4. The body next breaks up into a number of smaller bodies known as initial corpuscles. These, in their turn, divide by simple division (in the manner already described) into numerous elementary bodies (fig. 119). Thus, the life-cycle is completed. The Chlamydozoa are, then, the minute granules inside the body of the Cytoryctes variolce or the Neuroryctes hydrophobics, so that the whole body of the Cytoryctes or Neuroryctes corresponds to the mantle and parasite of the Chlamydozoon. The Cytoryctes group is said to cause destruction of the host cell. The Cytooikon group (e.g.y trachoma bodies) causes proliferation of the host cell. In September, 1913, Noguchi ^ described the cultivation of the parasite of rabies in an artificial medium, similar to that used by.., composed of the egg or ovarian cell, which is surrounded by numerous yolk cells or their products of disintegration. The two sexual openings usually lie close together, frequently in the fundus of a genital atrium ; they are rarely separated from one another. Shell glands also usually occur (p. 221). Reproduction is sexual, often, however, combined with asexual methods of propagation (segmentation, budding). The Platyhelminthes live partly free in fresh or salt water, exceptionally also on land. The greater part, however, live as parasites on or in animals. 2IO THE ANIMAL PARASITES OF MAN 4. The body next breaks up into a number of smaller bodies known as initial corpuscles. These, in their turn, divide by simple division (in the manner already described) into numerous elementary bodies (fig. 119). Thus, the life-cycle is completed. The Chlamydozoa are, then, the minute granules inside the body of the Cytoryctes variolce or the Neuroryctes hydropJiobice, so that the whole body of the Cytoryctes or Neuroryctes corresponds to the mantle and parasite of the Chlamydozoon. The Cytoryctes group is said to cause destruction of the host cell. The Cytooikon group {e.g.y trachoma bodies) causes proliferation of the host cell. In September, 1913, Noguchi ^ described the cultivation of the parasite of rabies in an artificial medium, similar to that used by,- parasitic forms. [In Platyhelminthes simple eye-spots frequently occur, and in a few an auditory vesicle.] z. . Blood-vessels and definite respiratory organs are lacking [except in Nemertinea\ ; the EXCRETORY APPARATUS (formerly termed water- vascular system) is typical of the entire class. It commences in the interstices of the parenchyma, with peculiar terminal cells (ciliated funnels;, which will be described later (p. 219), the capillary processes of which go on uniting into larger branches^ ^nd finally form two large collecting vessels, which, sometimes separately and. sometimes united, open to the exterior through one, two, or numerous pores. -^ Nearly all the Platyhelminthes are hermaphroditic, and in .nearly all there are, in addition to the ovaries producing ova, other glands attached to the female genital apparatus, namely, the vitellaria or yolk glands, which provide a substance termed yolk, which serves as nourishment for the embryo. The fully formed eggs have shells and are " compound," i.e.^ composed of the ^zg or ovarian cell, which is surrounded by numerous yolk cells or their products of disintegration. The two sexual openings usually lie close together, frequently in the fundus of a genital atrium ; they are rarely separated from one another. Shell glands also usually occur (p. 221). Reproduction is sexual, often, however, combined with asexual methods of propagation (segmentation, budding). The Platyhelminthes live partly free in fresh or salt water, exceptionally also on land. The greater part, however, live as parasites on or in animals. 212 the animal parasites of man Classification of the Platyhelminthes. Class /.— Turbellaria (or Eddy Worms). Flat worms for the most part, free living, and always covered with a ciliated epithelium. Order i. — Rhabdocoelida^ gut unbranched. Order 2. — Tricladida^ gut with three main branches. Order -^.—Polydadida, a central gut with lateral caeca. Development direct or through metamorphosis. They live in fresh and salt water or on land ; very seldom as parasites. Class //.— Trematoda (Sucking Worms'). [Usually known as Flukes. — F. V. T.] Flat worms, living as ecto- or endoparasites, that are only ciliated in the larval condition, and in their adult state are covered with a cuticle, the matrix cells of which lie in the parenchyma. They have either one, a few, or several suckers,- and frequently also possess chitinous fixation and adhesive organs. The intestine is single, but generally bifurcated, and not uncommonly there are transverse anasto- moses between the forks or diverticula on them. Excretory organs double, with two orifices at the anterior extremity or a single one at the posterior end. Development takes place by a metamorphosis or alternation of generations (p. 283). These worms are almost always hermaphroditic, with two or more female and one male sexual orifice. They live, almost without exception, as parasites on vertebrate animals, but the intermediate generations are passed in molluscs. Class III. — Cestoda (Tapeworms). Endoparasitic flat worms without an alimen- tary canal. The larval stages are rarely ciliated, but are usually provided with six spines ; the adult worm is covered with a cuticle, the matrix cells of which are em- bedded in the parenchyma. The body consists of a single segment (Cestodaria) or a chain of segments, in which case it consists of the scolex and the segments contain- ing the sexual organs (proglottides) (Cestodes s. str.). The scolex is provided with various adhesive and fixation organs, and there are calcareous corpuscles in the parenchyma. Excretory organs symmetrical, opening at the posterior end. These worms are always hermaphroditic, and then possess one or two female and one male sexual orifice. During development a larval intermediate stage ("measle") occurs and almost always in a different host to that in which the adult sexual worm lives. The adult stage is parasitic in vertebrate animals^ but the larval stage may occur in invertebrates. Class II. TREMATODA, Rud. These worms are usually leaf- or tongue-shaped, but also barrel- shaped or conical; they vary from o*i mm. to almost i m.^ in length ; most of them, however, are small (5 mm. to 15 mm.). The sur- face on which the orifice of the uterus and the male sexual opening are situated is termed the ventral surface ; the oral aperture, which also acts as anus, is always at the anterior end in the sub-order Prostomata (p. 230), but in the sub-order Gasterostomata it is ventral. ' This grouping goes back to the year 1800, and wai made by J. G. H. Zeder, a physician and helminthologist of Forchheim, who divided the helminths, which until 1851 were generally regarded as a special class of animals, into the groups of round, hook, sucker, tape and bladder worms, as which they are recognized up to the present time. In 1809, K. A. Rudolphi gave them the names Nematodes, Acanthocephali, Trematodes, Cestodes and Cystici. 2 A sucker or acetabulum (little cup) is a round, cup-shaped muscular organ, the muscles of which are sharply defined from those of the body. 2 Netnatobothrium filarina, van Bened., on the branchial chamber of the Tunny. TREMATODA 21 3 Suckers are always present and occur in varying numbers and positions at the anterior extremities as well as on the ventral surface, and occasionally on the lateral margin and on the dorsum ; the beginning of the intestine (mouth) is always surrounded by a sucker in the Prostomata. In or near the suckers there may be chitinous hooks, claws or claspers, or the surface of the body is more or less covered with spines, scales or prickles; in one genus (Rhopalias) there are projectile tentacles beset with spines on the sides of the anterior part of the body. The body of adult Trematodes is covered by a homogeneous layer of varying thickness, which either lies directly over the external layer (basement membrane) of the parenchyma, or over the muscles embedded in the parenchyma. This investing membrane (cuticle) arises from pear-shaped or spindle-shaped cells arranged singly or in groups (which lie between or internal to the diagonal muscles), and is connected with them by processes ; these cells one may regard as epithelial cells which have sunk down, or possibly as parenchymatous cells. An epithelium of one layer is also found on the body of young stages, but it disappears during growth, and only occasionally do its nuclei persist until adult life. In its place we then find the cuticle, which, moreover, extends into all the body openings more or less deeply. It is thus a debatable point whether the ^'investing layer " of flukes is a cuticle — that is, consists of modified epithelial cells — or whether it is a basement membrane, i.e., compressed and modified connective tissue cells ; in this latter case the true epidermis and cuticle have been cast off. In the former case the epidermal cells are the pear-shaped cells referred to above. According to recent authors it consists of two parts, an outer true cuticle and an inner basement membrane. There are also unicellular cuticular glands, lying isolated or in groups, which are termed cephalic, abdominal, or dorsal glands according to the position of their orifice. The PARENCHYMA is a connective substance, the structure of which is still a matter of dispute. It consists, according to some authors, of multipolar cells, the offshoots from which anastomose with each other so that a network, permeating the entire body and encom- passing all the organs, is produced. There exists also, as part of it, a homogeneous matrix, in the form of lamellae and trabeculae that border small cavities communicating with each other and filled with fluid. According to other authors, the parenchyma of the Trematodes con- sisted originally of cells, of which, however, only the cell membranes remain, while the protoplasm has been liquefied except for small residua around the nucleus. Between these cells an intercellular mass has appeared. By partial absorption of the walls, adjoining spaces unite. 214 THE ANIMAL PARASITES OF MAN and the originally flat cell walls become transformed into trabeculae. According to this vi^w the cavities filled with fluid are zw/ra-cellular, according to the former view /«/^y-cellular. Pigment cells occur only in a few species. The MUSCULAR SYSTEM of the Trematodes is composed of (i) a dermo-muscular tube, (2) the dorso-ventral or parenchymal muscles, (3) the suckers, and (4) the special muscles of certain organs. The dermo-muscular tube, which lies fairly close to the cuticle, consists of annular, diagonal, and longitudinal fibres which surround the entire body in one or several layers, and as a rule are more strongly developed on the ventral surface as well as in the anterior part of the body. The MUSCLES OF the parenchyma are found chiefly in the 7. I. A \ Md. F.v.s. C7t- F.v.s F.v.s Ex.v. Fig. 120. — Half of a transverse section through Fasciola hepatica, L. 25/1. Cm., Cuticle with scales; under the cuticle are circular muscles, and adjoining them the longitudinal and diagonal muscles; internal to the latter are the matrix cells of the cuticle; /., gut; the other similarly contoured cavities are gut diverticula that have been transversely or obliquely sectioned; i^.z^.^., vitellaria ; ^.^.z/., excretory vessels; T., testes; J/<;/. , median plane; the fibres passing from the ventral to the dorsal surface are the muscles of the parenchyma. The parenchyma itself is omitted. lateral parts of the body and pass through the parenchyma in a dorso-ventral direction ; their diverging brush-like ends are inserted on the inner surface of the cuticle (fig, 120). The suckers are specially differentiated parts of the dermo- muscular tube. Their concave inner surface is lined by the con- tinuation of the cuticle and their convex external surface is covered by a more dense tissue that frequently takes the form of a refractive membrane, thus separating them from the parenchymal muscles. The principal mass of the suckers consists of muscular fibres which run in three directions — equatorial, meridional and radial. The equatorial fibres correspond to the annular muscles, the meridional fibres to the longitudinal muscles, and the radial fibres to the muscles of the parenchyma; the radial fibres are always the most strongly TREMATODA 215 developed. The function of these muscles is evident from their position ; the meridional fibres flatten the suctorial disc and diminish the depth of its cavity, so that the internal surface may adhere to the object to be held ; if the equatorial fibres now contract, the sucker rises by elongating longitudinally, and its inner surface is drawn in by the contraction of the radial muscles. Thus the sucking disc becomes adherent. Usually also there is a sphincter Fig. 121. — Harmostoinum leptostomum, Olss., an immature specimen from Helix hortensis. Nervous system^ according to Bettendorf. A.s., ventral sucker; Cg.., cerebral ganglion; Ex.p., excretory pore ; G.p., genital pore; 6>.^., oral sucker; M.d., dorsal medullary nerve; M.I., lateral medullary nerve ; N.ph., pharyngeal neive ; M.v., ventral medullary nerve. Magnified. at the border of the suckers, which plays its part during the act of adhesion by constricting in a circular manner that part of the mucous membrane to which it is attached. The loosening of the fixed sucker is effected by relaxation chiefly of the radial fibres, by the contraction of the meridional fibres and certain bundles of muscles situated at the base and at the periphery of the suckers. The connective and elastic tissues between the muscles of the suckers probably also take part in the process. 2l6 THE ANIMAL PARASITES OF MAN Of the muscles of the organs which have developed from the parenchyma muscles we may briefly mention those bundles that are attached to certain parts of the genital apparatus, to the suckers, to the hooks and claws, and also, at all events in Fasciola hepatica, to the spines. The sheaths used for the projection of the tentacles of the RhopaliadcB are also muscular. The contractile elements consist of fibres of various lengths that are mostly parallel to one another, and frequently anastomose ; a cortical substance finely fibrillated can usually be distinguished from an internal homogeneous mass ; large nucleated cells of uniform size are always connected with them ; these have been variously interpreted, but have been proved to be myoblasts, one or more of their processes constituting the muscular fibres. The MOVEMENTS of the Trematodes consist in alterations of form and position of the body, as well as in creeping movements. In the NERVOUS system (fig. 121) can be distinguished a cerebral portion as well as strands (medullary strands) running from it^ and peripheral nerves. The cerebral portion always consists of two large ganglia situated in the anterior end of the body which pass dorsally over the oesophagus and are connected by means of a broad and thick commissure composed of fibres only. From each ganglion three nerves run anteriorly — the inner and dorsal nerve for supplying the anterior dorsal part of the body ; the median and ventral for the oral sucker; and the exterior and lateral likewise for the supply of the sucker. In a similar manner three strands run backwards from each ganglion — one dorsal, one lateral and one ventral. The dorsal and ventral strands become united and curve backwards ; the symmetrical lateral strands are connected by means of transverse commissures, the number of which vary according to the species. Such commis- sures also exist between the lateral and the two other strands on each side. There are ganglion cells along the entire course of the posterior cords, more particularly at the points of origin of the commissures. There also appears to be in addition a fourth anterior and posterior pair of nerves, the front pair for the oral sucker and the hind pair for the pharynx. The peripheral nerves, which spring from the posterior strands as well as from the commissures, either pass directly to the muscular fibres or to the sensory cells that are situated at the level of the subcuticular cells, or they reach these after the formation of a plexus situated immediately beneath the dermo-muscular layer ; the pro- cesses directed outwards terminate in small vesicles in the cuticle. As to other ORGANS OF SENSE, simple eyes, two or four in number, are known in several ectoparasitic species as well as in a TREMATODA 217- few free-living larval stages (Cercariae) of endoparasitic forms. In the adult stage, however, they usually undergo complete atrophy. The ALIMENTARY CANAL commences with an oral aperture, generally terminal or sub-terminal (ventral) at the anterior extremity, which leads into an oral cavity usually surrounded by a sucker ;. the oesophagus, of various lengths, is directed backwards and is generally surrounded by a muscular pharynx (fig. 122). In some cases there exists between the sucker and pharynx, pharyngeal pouches (praepharynx). Sooner or later the intestine divides into two lateral branches directed backwards, both of which end blindly (caeca) at the same level.' In many ectoparasites (Mono- genea [p. 222]) a connection exists between the genital glands and one of the intestinal branches (ductus vitello- intestinalis [fig. 123]). The oral cavity, pharyngeal pouches, pharynx, and oesophagus are lined with a continuation of the cuticle of the body ; the gut caeca are lined with tall cylindrical epithelium (fig. 120). The oesophagus and intestinal branches often have also one layer of annular and longitudinal muscles; the pharynx has essentially the structure of a sucker (fig. 122). The accessory organs of the ali- mentary canal consist of groups of unicellular salivary GLANDS that discharge into the oesophagus in front of or behind the pharynx, or even into the pharynx itself. Fio. 122. — Median section through the anterior part of Fasciola hepatica : the oral sucker, pharyngeal pouches, pharynx, oesophaeus, cuticle with spines, and the body parenchyma. ^ The following conditions represent deviations from this type: (i) In Gasterostomum the oral apeiture is situated in the middle of the ventral surface, and occasionally is even nearer to the posterior than to the anterior end. There is no proper oral sucker,, but the pharynx is thus termed. (2) A few genera, such as G aster ostomum^ Aspidogaster ^ Diplozoon, etc., have only one intestinal diverticulum, which is undoubtedly to be taken as representing the primitive condition, as it is also often found in the young stages of the Trematoda. (3) The branches of the intestines are curved and united behind (several Tristomidce and Monostomida), while in Polystomtwi integerrimum (in the bladder of frogs) there are several commissures betwee n the intestinal branches, and in the ^r-^zV/^- somidce the united intestinal branches proceed as one channel towards the posterior end. (4) The termination of the two intestinal branches is not always on a level ; they are therefore of different lengths. (5) When the oesophagus is very long the intestinal branches extend both forward and backward, so that the gut exhibits the form of an H. (6) In the broad and fiat species the gut-forks form diverticula mostly externally but also internally ; these again may branch (fig. 139). (7) In a few cases [Nematobothrium, Didymozoon) the intestine completely disappears up to the pharynx. 2l8 THE ANIMAL PARASITES OF MAN The food of the Trematodes consists of mucus, epithehal cells, the intestinal contents of the hosts, and often also of blood, and this not only in those species living in the vascular system, but also in species Oot, Ir. c.v v.d.e. Fig. 123. — Polystomum integerrimum, a -monogeneiic fluke from the urinary bladder of the frog, z., intestine; h., large hooks of the sucking disc ; h.k.^ smaller hooklets ; Lev., longitudinal vitelline ducts; 0., oral orifice ; OoL, ooiype ; ov., ovary ; s.p., suckers of the disc ; tr.c.v.y transverse vitelline ducts ; Ut.y uterus with ova; 7;., entrance to the vagina; v.d.e., vas deferens; v.d.i., ductus vitello-intestinalis ; the vitellaria and testes are not shown. Magnified. (After Zeller.) Fig. 124. — Allocreadium iso- porum, Looss. Excretory appa- ratus. Of the other organs, the oral sucker, pharynx, genital pore, ventral sucker, ovary and testes are shown ; the cylin- drical excretory bladder is in the posterior end. 38/1. (After Looss.) living as ectoparasites or in the intestine or biliary passages of their hosts. The final products of assimilation dissolved in the fluids of the body are distributed throughout the parenchyma and are thence TREMATODA 219 expelled by a definite tubular system (excretory apparatus, proto- nephridia, formerly also termed the water-vascular system). This system, which is distributed throughout the entire body (fig. 124), is symmetrically developed, and, in the ectoparasitic Trematodes, it opens, right and left, at the anterior end on the dorsal surface ; in all other flukes, however, it opens singly into the excretory pore (foramen caudale) at the centre of the posterior border ; in those cases, however, where a sucker is present at the posterior end, as in the Amphistomata, the excretory pore is situated on the dorsal surface close in front of the sucker. The EXCRETORY SYSTEM^ consists of several parts : (i) of the more or less numerous terminal "flame" cells or funnel cells (figs. 124, 125); (2) of the capillaries ending in them ; (3) of larger vessels receiving the capillaries ; and (4) of the excretory bladder. Terminal cells and capillaries may be compared to unicellular glands with long excretory ducts ; the cellular body (fig. 125) is comparatively large, stretched longitudinally, more rarely transversely, and provided with numerous processes, that are lost in the parenchyma ; within is a conical cavity (analogous to the secretory cavity of unicellular glands) which is continued directly into the structureless capillary ; at its blind end is a bunch of cilia projecting into the cavity, and which, during life, shows a flicker- ing motion (ciliary flame). The nucleus is situated in the protoplasm of the terminal cell at its blind end. The entire apparatus thus begins blindly — ix.j within the terminal cells, to which must be ascribed the capacity of taking up from the fluid that permeates the parenchyma the products which are first collected into their own cavities and thence excreted by means of the capillaries and vessels. The vessels possess definite walls, consisting of a membrane and a r^ucleated protoplasmic layer. They unite at many points on either side, and again pass into other canals (COLLECTING TUBES), which finally, travelling towards the posterior end, discharge into the excretory bladder (fig. 124). The form and size of the bladder vary much according to the different species, but it always possesses its own flattened epithelium, surrounded by circular and longitudinal muscles, the circular muscles Fig. 125.— Terminal flame cell of the excretory system. n., nucleus of cell ; c. , bundle ofcilia forming the "flame"; p., processes of cell extending into parenchyma; J., excre- tory capillary. (Stephens.) The following description relates in the main to the Distomata. 220 THE ANIMAL PARASITES OF MAN forming a sphincter around the opening. Frequently also the structure of the bladder extends to the tubules discharging into it, which therefore are not to be regarded as separate *' vessels," but rather as tubular diverticula of the bladder, directed anteriorly. In some few species the diverticula also branch and the branches anastomose, so that a network of tubules ensues which receives the vessels or capillaries. In such cases there are also ciliary tracts in the tubules. The contents of the entire appar- atus usually consist of a clear or some- times reddish fluid ; in some species there are larger or smaller granules, and occasionally also concretions occur. vag. -ut. J^ec. ut. ^ec. sent. Vit. R. Fig. 126. — Diagram of female genitalia. Ov.^ Fig. 127.— Diagram of male and part of female genitalia. tU., uterus; vag.^ ovary; ovd.^ oviduct; Z.r., Laurer's canal; vagina; 5 , opening of vagina ; ^.j., geni- Rec. sem.y receptaculum seminis ; Vit. R., vitel- larian reservoir ; t.v.d., transverse vitelline duct tal sinus ; g.p., genital pore; (^ , opening of ejaculatory duct or vas deferens ; c.s., Oo., ootype ; Sh. gl., shell gland; Rec. ut,, cirrus sac; f., cirrus ;/./., pars prostatica ; receptaculum uterinum ; ut., uterus. (The various parts are not to the same scale.) (Stephens.) s.v., seminal vesicle; e.J., ejaculatory duct or vas deferens ; v.e., vas efferens ; t., testis. (Stephens.) Sexual Organs. — Nearly all the Trematodes are hermaphrodites, and only a few (Schistosotnidce, Koellikeria) are sexually differen- tiated. The sexual organs usually lie in the " central field " limited by the gut caeca ; the vitellaria, on the other hand, are, as a rule, external to the gut caeca in the *' lateral fields." The male apparatus^ is composed of two variously formed testes (fig. 127) (globular, oval, indented, lobed, or ramified), which may lie side by side or one behind the other ; from each testicle a tube (vas efferens) originates ; sooner or later, both tubes as a rule unite to form the ejaculatory duct or vas deferens, which is frequently The following description relates mainly to the Distoniata. TREMATODA 221 enclosed in a muscular CIRRUS SAC, or more rarely passes directly into the genital pore. The cirrus, which is the thick muscular terminal portion of the vas deferens, can be everted and protruded from the cirrus sac and serves as an organ of copulation. The walls of the muscular portion of the tube (the cirrus) are attached to the walls of the cirrus sac, and hence when the sac contracts the cirrus cannot be protruded except by evagination of its lumen. Opening into the middle portion of the vas deferens, and as a rule enclosed in the cirrus sac, is found a mass of unicellular glands (prostate), the vesicula seminalis (which is likewise within, or may also be outside the sac) being the dilated first portion of the vas. The female genitalia (fig. 1 26) consist of an ovary, usually situated in front of the testes, the form of which varies according to the species, the usually double vitellaria, the ducts and a number of auxiliary organs ; the short oviduct directed towards the centre arises from the ovary, and is connected in the median line with the excretory duct of the vitelline glands. These grape-like glands possess longitudinal excretory ducts, which assume a transverse direction behind the ovary, unite together at the median line and form a single duct, often dilated into a vitelline receptacle, that unites with the oviduct. Near this point, moreover, there frequently opens a canal (Laurer's canal) which begins on the dorsal surface, and on the inner end of which a vesicle filled with sperm (receptaculum seminis) usually occurs (fig. 126). Moreover, there are also numerous radial unicellular glands (shell glands) at or beyond the point of junction of the oviduct, vitelline ducts and Laurer's canal. In this portion of the duct (ootype), which is usually dilated, the ovarian cells are fertilized, surrounded with yolk cells and shell material, and as ova with shells they pass into the uterus (a direct continua- tion of the oviduct), which, with its many convolutions, occupies a larger or smaller portion of the central field, and runs either direct to the genital pore or, forming convolutions, first runs posteriorly and then bends forward (descending and ascending limbs). In both cases the terminal part lies beside the cirrus pouch and discharges beside the male orifice either on the surface of the body or into a genital atrium. The terminal portion of the uterus, which is often of a particular structure, serves as a vagina (METRATERM). The cirrus sac may include (i) the genital atrium {i.e., the common sinus, into which the vas deferens and vagina may open), or (2) a variable extent of the vas from cirrus to seminal vesicle. Thus the latter may be outside the sac. In the absence of a sac, the genital sinus may be surrounded by a pseudo-sucker, as in Heterophyes (in some cases the ventral sucker itself, from its close proximity to the genital pore, serves as an accessory copulatory organ). In other cases 222 THE ANIMAL PARASITES OF MAN copulatory organs are formed by hooks projecting into the lumen of the terminal portion of the vas. The GENITAL PORE, which is the opening from the genital sinus on to the surface, is generally situated at or near to the median line on the ventral surface and in the anterior region of the body ; in most of the Distomata it is in front of the ventral sucker, in other cases, e.g.y in the Cryptocotylince, it is behind.^ The spermatozoa do not differ essentially in their structure from those of other animals ; the ovarian or Qgg cells are cells without integument and contain a large nucleus and a little protoplasm; the vitellaria also produce nucleated cells, in the plasm of which there are numerous yellow yolk granules ; the yolk cells detach themselves, like the ovarian cells, from the ovarium, and pass into the oviduct to surround each ovarian cell in the ootype. They disintegrate sooner or later in the completely formed Qgg and are utilized as food by the developing embryo. Development of the Trematodes. (i) Copulation. — Observation has demonstrated that the one or two vaginae occurring in the ectoparasitic Trematodes are utilized as female organs of copulation, and that the copulation is cross ; it is also known that Laurer's canal, which was formerly generally regarded as the vagina, has only quite exceptionally, if at all, served the digenetic Trematodes as such — it appears to be homologous with the canalis vitello-intestinalis of the Monogenea? — but the terminal portion of the uterus, termed the metraterm, is used for copulation. Cross-copulation occurs as well as auto-copulation and auto-fecunda- tion. The spermatozoa subsequently pass through the entire uterus, which is still quite short at the time the male organs are matured ; the maturation of which, as usually is the case in hermaphrodites, precedes ^ The typical position of the genitalia is subject to many deviations, which are of importance in the differentiation of the genera and families. The following are some few of these deviations : (i) The genital pore remains on the ventral surface, but is situated beside or behind the ventral sucker, or it becomes marginal, and is then found in front of or beside the oral sucker, or at a lateral edge, or, finally, in the centre of the posterior border ; the ducts also correspondingly alter their direction. (2) The ovary usually lies in front of the testes, not rarely, however, behind them or between them. (3) The three genital glands mostly lie together close in front of, or behind, the centre of the body ; they may be moved far back, and may incidentally become separated one from the other. {4) The vitellarium may be single, in which case it then may lie in the central field. (5) A few forms possess but one, others several or numerous testes. Amongst the ectoparasitic trematodes there are also species with but one testis ; but they mostly have several. As a rule, their uterus is short, but the ootype well developed. Special canals (vagina), single or double, are used for copula- tion, not the uterus. The vitelline ducts also communicate with the intestine through the canalis vitello-intestinalis (fig. 123). 2 Monogenea : Trematoda in which the anterior sucker, if present, is double. Development without an intermediate host. TREMATODES : DEVELOPMENT 223 that of the female organs. It is only later with the onset of egg forma- tion that the uterus is fully developed. Copulation, however, takes place also in the case of fully grown forms with completely developed uteri. (2) Formation of the Ova. — The ovarian cells arising from the ovary first become mature after their entry into the ootype by the formation of three polar bodies, fertilization then taking place. At the same time as the ovarian cell a number of yolk cells from the vitellarium and secretion, drop by drop, from the shell gland reach the ootype.^ The shell is then formed during the generally active contractions of the ootype w^alls and then passes on into the uterus. In the uterus of the endoparasitic trematodes the eggs accumulate more and more, often in large quantities, while in ectoparasitic species generally only Fig. 128. — Ovum of Fasciola hepatica, L., cut longitudinally. The lid has been lifted in the process. Within the egg are numerous yolk cells, and at the lid end there is the still unseg- mented ovum (dark). 240/1. Fig. 129. — Miracidium of Fasciola hepatica that has just hatched from the egg, with a dis- tinct cuticular ciliatedepithelium. Magnified. (From Leuckart.) one or some few eggs can be found. The completed ova are of various forms and sizes. They are mostly oval, at all events in the digenetic trematodes, and the yellowish or brown shell is provided with an opening at one pole which is closed by a watch-glass-shaped lid (operculum). Appendages (filaments) on the shell — at one or both poles — are uncommon, but are the rule in the ova of the Monogenea (ectoparasitic species). (3) Deposition of the Ova. — Soon after their formation, the Monogenea (ectoparasitic trematodes) deposit round the place of ' [Recent work {e.^., Goldschmidt, Zool. Anzeiger^ xxxiv, p. 482) has shown that the older views regarding the formation of the egg must be modified. In certain species, at any rate, the shell material is formed by the yellow droplets of the yolk glands and not by the so- called shell gland (Mehli's gland) secretion, which is clear and watery. The function of this secretion accordingly still requires explanation ; according to Looss it serves as a covering secretion for the egg-shell proper. It appears also that other granules, the yolk granules as distinct from the shell drop granules, are not always used up during the development of the embryo and hence do not function as yolk, so these also when they exist, and frequently they are wanting, must serve some other purpose, possibly that of imbibing water for the use, of the embryo.— J. W. W. S.] 224 THE ANIMAL PARASITES OF MAN their attachment on the skin or the gills or other organs of their hosts, eggs which attach themselves by means of their filaments. The embryonic development thus takes place outside the parent. This also holds good for the eggs of many endoparasitic species, although as a rule in these the eggs are always retained for a longer time in the uterus. Moreover, they usually here undergo a part or a whole of their development, and are eventually deposited in those organs in which the adult forms are parasitic, but this is not always the case, as the egg, e.g., of F. hepatica appears in bile (and faeces) quite unchanged. By the natural passages they eventually get out of the body, and in cases where such do not exist, as in the case of the blood-vessels, the eggs pass out by means of the kidneys. (4) The embryonic developinent, after irregular segmentation of the ovum into a number of blastomeres, leads to the formation of a solid blastosphere or morula, which is surrounded by a cellular investing membrane (yolk envelope), while the principal mass of the cells forms the embryo, which uses for its nourishment the yolk cells, which have in the meantime disintegrated {cf. footnote, p. 223). Usually, after the ova have reached water the embryos hatch out, leaving the yolk envelope in the egg-shell ; in other cases, however, the embryos only hatch out after having been subjected to the influence of the intestinal juices, that is to say, in the intestine of an intermediate host which has ingested with its food the ova that have escaped from the primary host. (5) The post-embryonic development of the Trematodes is accom- plished in various ways ; the process is the most simple in the ecto- parasitic species (Monogenea), the young of which should certainly be regarded as larvae, because they possess characteristics (cilia, simple gut, etc.) that are lacking in the adult worms, but which, nevertheless, pass into the adult state direct after a relatively simple metamorphosis. In the Holostomata,^ a group found chiefly in the intestine of aquatic birds, and which rarely occur in other vertebrates, the ova develop in water. The young are ciliated all over, and, after having entered an intermediate host (leeches, molluscs, arthropods, amphibians, fishes) living in the water, they undergo a metamor- phosis into a second larval stage ; they then encyst and await trans- mission into the final host, where they become adult Metastatic trematodes, i.e., trematodes without asexually produced generations (p. 229). In the remaining so-called digenetic trematodes (p. 230) one or two asexual generations interpose between the miracidium and terminal stage, so that quite a number of adult worms may originate from one egg. Usually the young, which are termed miracidia^ (fig. 129), • Holostomata : Prostomata with (in addition to the oral and ventral suckers) a third fixation apparatus, generally on a separate part of the body. ^ [Also known as ciliated embryos. — F. V. T.] TREMATODES : DEVELOPMENT 225 hatch in water, where they move with the aid of their ciha. Sooner or later they penetrate into an intermediate host, which is always a snail or a mussel, and while certain of their organs disappear, they grow into a gutless germinal tube (sporocyst, fig. 131). These are simple elongated sacs with a central body cavity. They may or may not have excretory tubules. In these, according to the species, the larval stages (CERCARi^) that will ultimately become adult worms are produced, or another intermediate generation is first formed, viz.^ that of the REDI^^ (figs. 132, 133), which are always provided with an intestine, and these then give rise to cercariae (figs. 130, 134). The cercariae, as a rule, leave their host and move about in the water with the assistance of their rudder-like tails. After a little time, how- ever, they usually again invade an aquatic animal (worms, molluscs, arthropods, fishes, amphibians), then they lose their tails and become Fig. 130. — A group of cercarioe of Echinostoma sp. (from fresh water). 25/1. encysted (fig. 135) ; here they wait until they attain, together with their host, the suitable terminal host, and in this new situation they establish themselves and reach maturity. Or, again, the cercariae may themselves encyst in water or on foreign bodies (plants) and wait until they are taken up directly by the terminal host, e.g.y sheep. Accordingly the following conditions are necessary for the com- pletion of the entire development : (i) The terminal host in which the adult stage lives ; (2) an intermediate host into which the miracidia penetrate and in which they become sporocysts ; (3) a second intermediate host in which the cercariae become encysted. In certain species, as in Fasciola hepatica, this second host is omitted, as the cercariae spontaneously encyst on plants, or again (in other species) encystment may occur within the first intermediate host, when, * [In Fasciola hepatica in the summer months the rediae give rise to daughter redise, which then give rise to cercariae. — J. W. W. S.] 15 220 THE ANIMAL PARASITES OF MAN ill fact, the cercarice (which in this case do not acquire an oar-like tail) do not swarm out of, but encyst themselves w^ithin their sporocysts. The development, moreover, may be further complicated by rediae appearing in addition to the sporocysts, though this occurs in the first intermediate host and not in a second one. Animals that harbour adult digenetic Trematodes thus become infected by ingesting encysted cercariae, which either occur (i) in certain animals (second intermediate hosts) on which they feed, or (2) in water, or (3) on plants, or finally (4) in the first intermediate host ; whereas animals harbouring encysted cercariae have been directly infected by the corresponding tailed stage, and animals harbouring germinal tubes (sporocysts or rediae) have been infected by the miracidia. Thus certain species of ducks and geese become infected with Echinostoma echinatiim by devouring certain water-snails {LimncBus, Fig. 131. — Development of Fasciola hepatica, L. a, the miiacidium in optical section showing cephalic lobe, X-shaped eye-spot resting on the cerebral ganglion, two germ balls ; below each of these a flame cell, and still lower germ cells lying in a caviiy (primitive body cavity). ^, young sporocyst with two eye-spots, and germ balls ; the cells lining the cavity are not shown. ^, older sporocyst with a young redia. Magnified. (After Leuckart.) Paludina) in which the encysted cercariae occur. Oxen become infected with Paraniphistomum cervi (= Aniphistomiun coniciim) by swallowing with water, cysts of this species which occur at the bottom of puddles and pits. Sheep are infected with Fasciola hepatica by eating grass to which the encysted cercariae of the liver-fluke are attached; our song-birds infect themselves or their young wnth Urogoninnts macro- stomus by tearing off pieces containing the corresponding sporocysts which are full of encysted cercariae from snails {Succinea amphibia), which act as the first intermediate hosts, and eating, or offering their young these pieces. (i) The MIRACIDIA of the digenetic Trematodes are compara- tively highly organized, and the mode of their formation from the TREMATODES : DEVELOPMENT 227 segmentation cells of the ovum is only imperfectly known. They have a cuticular epithelium (fig. 129) entirely or partly covered with cilia, beneath this a dermo-muscular tube composed of circular and longi- tudinal muscles ; also, a simple gut sac with an oesophagus, occa- sionally also with pharynx, salivary glands and boring spine, also a cerebral ganglion on which, in some species, there are eyes (fig. 131, <^). As to the excretory organs, they are represented by two symmetrically placed terminal flame cells, with excretory vessels opening separately ; there is a more or less ample (primary) body cavity between the parietes of the body and the gut ; from the cel- lular parietal lining of this cavity single cells (germ cells) become free (fig. 131, a, 6), and become rediae or cercariae. [The germ cells of the mira- cidium and the germ balls of the sporocyst arise, according to some observers, by further division of undifferentiated blastomeres ; ac- cording to others from the cells of the lining wall of its body cavity. It is from these free germ balls that the redia stage is developed. [In the germ ball or morula appears an invagination, giving rise to the cup-shaped gastrula stage. This elongates and forms the REDIA (fig. 131, c). [In the interior of the redia cells are budded off and develop into gastrulae, as in the case of the sporocyst. These become a fresh generation of rediae or give rise to the third stage (cercaria).] (2) The SPOROCYSTS, on the contrary, which are produced direct from the miracidia, are very simple, as all the organs of the latter disappear, even to the muscles and excretory organs, during or after penetration into the intermediate host, whereas the budded and still budding cells of the wall of the (primary) body cavity continue to develop rapidly and form germ balls. The sporocysts when fully developed have the appearance of tubes or fusiform bodies with rounded edge; they are frequently of a yellow colour. Their length Fig. 132. — Young redia of Fasciola he- patica^ with pharynx and intestine, with a circular ridge an- teriorly and a pair of processes poste- riorly and masses of cells (germ balls) in the interior. Mag- nified. (From Leuckart.) Fig. 133.— Older redia of Distoma echinatum, with ru- dimentary intestine i. ; cercariae, c. ; germ balls, b. ; and birth pore, g. Magnified. 228 THE ANIMAL PARASITES OF MAX rarely exceeds a few millimetres ; in some species their size increases exceedingly through proliferation, and they then occupy a large portion of the body of the intermediate host. (3) The REDiJE (figs. 132, 133), on the other hand, are more cylindrical and always have a simple intestine of varying length, provided with a pharynx ; they likewise possess, situated near the circular ridge, a '' birth pore " which serves for the exit of the cercariae originating within them. (4) The CERCARi^^ are veiy different; typi- cally they consist of the anterior body and the oar-like tail at the posterior end (fig. 134). The former, even to the genitalia, has the organiza- tion of the adult digenetic Trematodes, and thus allows the easy recognition of at least the characters of that large group to which the species in question belongs. On the other hand, however, there are also organs that are lacking in the adult form, such as, in many, the boring spine in the oral sucker, or the eyes situated on the cerebral ganglion ; moreover, also, cutaneous glands (fig. 134), the secretion of which forms the cyst membrane. The oar-like tail may be long or short (stumpy-tailed cer- caria) or entirely absent ; its free end may be partly split (furcate cercaria), or split to its base (bttcephalns) ; in various forms also the an- terior end of the tail is hollow, and has enclosed within it the anterior body, which is otherwise free. The size also of the cercaria belonging to the different species is very diverse ; in addition to forms swimming in the water that have the appearance of minute milky-white bodies, there are forms which measure as much as 6 mm. in length. Fig. 135.— Encysted cer- ° . 1 , , caria of FasaWa hepatica. The cncysted ccrcariae (fig. 135) are globular Magnified. (After Leuck- qj- oval, and are surrounded by a homogeneous art.) ^ ^ Fig. 134. — Cercaria of Fasciola hepatica ; the cuta- neous glands at the side of the anterior body. Mag- nified. (After Leuckart.)ij^ ^ The cercaria is the characteristic larval stage of the Trematodes, and corresponds to a cysticercus or cysticercoid, though there is the important difference that the cercaria has an enteric cavity. According to some observers the enteron is represented by the frontal sucker of some Cestodes, and by the rostellum of the majority of others. The sporocyst and redia are regarded as intercalated stages, viz., as cercarise exhibiting pedogenesis, i.e., development of young by a parthenogenetic process fr( m individuals [i.e.y cercaria) not yet adult. TREMATODES : BIOLOGY 229 membrane, which may be striated or contain granules. The tail is always cast off when encystment occurs, and organs peculiar to the cercaria stage (boring papilla, eyes) almost entirely disappear. On the other hand, the genitalia appear or become more or less highly developed, in extreme cases to such an extent that they become functional, and after autocopulation the creatures produce ova within the cysts. The cycle of development of the digenetic Trematodes has hitherto been generally explained as a typical alternation of generations, one sexual generation regularly alternating with one or two asexually reproducing generations. Recent authors, how^ever, regard the cells in the sporocysts from which rediae or eventually cercariae arise as parthenogenetically developing ova, and the sporocysts as well as the rediae as generations propagating parthenogenetically. In this case, however, it is an alternation of a sexual not with an asexual but with firstly a parthenogenetic generation (the sporocyst), the central cells of which are regarded as ova which develop parthenogenetically into the redia, and this the second parthenogenetic generation finally produces larvae (cercariae) capable of developing into the sexually mature form. Other authors, again, regard the development of the Digenea as only a complicated metamorphosis (p. 283), which is distributed over several generations before it is concluded. Biology. Endoparasitic Trematodes, as fully developed organisms, occur in vertebrate animals only, with very few exceptions; they inhabit almost all the organs (with the exception of the nervous and osseous systems and the male genitalia), but by preference the intestine in all its extent from the oral cavity to the anus; and, further, certain species or groups inhabit only quite restricted parts of the intestine. Besides in the intestine other species live in the liver, or in the bile-ducts, or in the gall-bladder; other accessory organs of the intestine, such as the pancreas, bursa Fabricii (of birds), are only infected by a few species. Many inhabit the lungs, or the air sacs in fowls, a few the trachea. Trematodes have also been known to occur in the urinary bladder, the urethra and the kidneys of all classes of vertebrates; they are also present in the vascular system of a few tortoises, birds and mammals ; in birds they even penetrate from the cloaca into the oviducts, and are occasionally found enclosed in the laid eggs ; one species is known to occur in the cavum tympani and in the Eustachian tube of a mammal (Dugong), another in the frontal sinus of the polecat ; several species infest the 230 THE ANIMAL PARASITES OF MAN conjunctival sac under the membrana nictitans of birds, one species even lives in cysts in the skin of song-birds. In an analogous manner the ectoparasitic Trematodes are not entirely confined to the surface of the body or the trachea of the lower vertebrate animals ; a few species appear exclusively in the urinafy bladder, in the oesophagus, and in the case of sharks in an accessory gland of the rectum. Trematodes live free and active within the organs attacked, though they may attach themselves by suction for a longer or shorter period ; in other cases, however, they bore more or less deeply into the intestinal wall with their anterior end, or lie in cysts of the intestinal wall which only communicate with the lumen through a small opening ; in those species living in the lungs of mammals the host likewise produces a cyst, which usually encloses two specimens; such association of a pair is also observed in other situations, and, though this is the rule in species sexually distinct, it is not entirely confined to these. As regards the AGE attained by endoparasitic Trematodes, there are but few reliable records, and these differ considerably ; the overwhelming majority of species certainly live about a year, or perhaps a little longer, but there are some whose term of life extends to several or many years. Trematodes are but rarely found encysted in the higher verte- brate animals ; the condition, however, is more frequent in amphi- bians, and especially in fishes, as well as in numerous invertebrate animals. Classification of the Trematodes of Man. The following classification, partly artificial, partly natural^ embraces only the flukes found in man : — Order. Digenea, v. Beneden, 1858. Anterior sucker single and median, present. Eggs few. The (specialized) terminal portion of the uterus serves as a vagina. Development indirect, i.e.^ an intermediate host is required. Sub-order. Prostomata, Odhner, 1905. Mouth surrounded by the anterior sucker. Group. Amphistomata, Rudolphi, 1801, ep., Nitzsch, 1819. Gut forked, two suckers, the posterior sucker (acetabulum) terminal or ventro- terminal behind the genitalia, or at most embraced by the vitellaria. Skin with no spines. Excretory bladder a simple sac opening dorsally near hind end. Testes in TREMATODES : CLASSIFICATION 23 1 front of ovary. Genital pore, median in anterior third of body. Thick flukes, ahnost circular in cross section. Family. Paramphistomidae, Fischoeder, 1901. Amphistomata : Body not divided into a conical anterior portion and disc-like caudal portion. Ventral pouch absent. Sub-family. Paramphistominae, Fisch., 1901. ParamphistomidiE : Oral sucker without evaginations. Not in man. Sub-family. Cladorchiinae, Fisch., 1901. Paramphistomidae : Oral sucker with evaginations ; testes, two, deeply cleft (fig- I37)' Genera: IVatsonius, Cladorckts, tic. Family. Gastrodlsciidae, Stiles and Goldberger, 1910. Amphistomata : With body divided into a conical cephalic and disc-like caudal portion (fig. 138). Posterior sucker ventro-terminal. Oral sucker with evaginations. Genera : Gastrodiscus and Homalogaster. Group. Distomata, Retzius, 1782. Gut forked, two suckers, the posterior sucker (acetabulum) ventral. It is always separated from the hind end by at least a part of the genitalia. Family. Fasciolidae, Railliet, 1895. Large flat forms, genital pore in front of ventral sucker, the latter powerful. Vitellarias of numerous follicles, united by branching vitellarian ducts, at the sides of the body meeting posteriorly and extending ventrally and dorsally. Cirrus and vagina without spines. No crown of strong spines around sucker. Testes much branched. Uterus not well developed. Excretory bladder much branched. Eggs large. Sub-family. Fasciolinae, Odhner, 1910. Large or median forms, gut much branched. Body has a shoulder separating head from body. Receptaculum seminis absent. Ovary branched, ventral sucker in anterior part of body. Genus : Fasciola. Sub-family. Fasciolopsinae, Odhner, 1910. Shoulder absent. Receptaculum seminis present. Ovary branched, gut takes a zig-zag course with kinks on it, ventral sucker in anterior part of body. Genus : Fasciolopsis. 2^.2 THE ANIMAL PARASITES OF MAN Family. Opisthorchiidae, Braiin, 1901, emend, auctor. Ovary in front of testes. Small to medium flukes, very transparent, tapering anteriorly. Vitellaria moderately developed not extending in front of sucker. Cirrus absent. Seminal vesicle a twisted tube free in parenchyma. Testes near hind end one behind the other, lobed or branched, but not dendritically. Excretory bladder Y-shaped, the two limbs short, the stem S-shaped passing between the testes. Receptaculum seminis well developed. Laurer's canal present. Uterine coils transverse, numerous. Eggs small. Sub-family. Opisthorchiinae, Looss, 1899, emend, auctor. OpisthorchiidcE in which the excretory pore is terminal. Excretory bladder long, dorsal to testes. Uterine coils not overlapping gut forks. Genera : Opisthorchis, Paropisthorchis, Clonorchis, Amphimerus, etc. Sub-family. Metorchiinae, Liihe, 1909. Opisthorchiidcr in which the excretory pore is ventral. Excretory bladder short, ventral to testes. Uterine coils partly overlapping gut forks and extend anteriorly beyond the sucker. Vitellaria compressed on the sides of the body. Genus : Metorchis. Family. Dicrocoeliidae, Odhner, 1910. Ovary behind testes. Testes behind the ventral sucker, between it and the ovary. Body thin and transparent. Cirrus sac encloses the pars prostatica and seminal vesicle. Skin smooth. Gut forks do not reach posterior end. Receptaculum seminis and Laurer's canal present. Vitellaria, moderate, lateral in mid-body slightly over- lapping the gut. Uterus with an ascending and descending branch and numerous transverse coils extending to hind end. Eggs dark brown, 25 ju to 60 m. Excretory bladder tubular in posterior third or half of body. Parasitic in bile-ducts of mammals and birds. Genus : Dicrocoeliuin, Family. Heterophyiidae, Odhner, 1914. Ovary in front of testes. Genital pore behind or on a level with ventral sucker. Genital pore surrounded by a pseudo-sucker {i.e., its muscle is not sharply separated from but blends with the body muscles). Cirrus sac absent, consequently vesicula seminalis and pars prostatica lie free. Vagina and ejaculatory duct unite into a common duct before opening. Small and very small forms. Body covered with scales. Genera : Heterophyes, Meiagonimus^ etc. Family. Troglotremidae, Odhner, 1914. More or less flattened Distomes of compact form, 2 to 13 mm. long. Ventral surface flat or somewhat hollowed, dorsal surface arched. Skin completely covered with pointed spines. Musculature weakly developed also in the suckers in those TREMATODES : CLASSIFICATION 233 forms that inhabit cysts. Gut with pharynx and a not very long oesophagus and caeca, which end more or less shortly before the hind end. Excretory bladder Y-shaped or tubular. Pars prostatica and seminal vesicle always distinct. Testes elongated, symmetrically placed in or behind the middle of the body. Ovary directly in front of the testes, right-sided, generally much lobed. Receptaculum seminis and Laurer's canal present. Vitellaria generally well developed, exclusively or for the most part confined to the dorsal surface^ leaving only a median band unoccupied. Uterus either very long, coiling here and there, or shorter and more convoluted. Eggs in first case small 17 ju to 25 ^u, in the second much larger 63 a* to 85 /x or even 120 /*(?) long. Parasitic in carnivora or birds, generally occurring in pairs in cyst-like cavities. Genera : Paragonhnus^ Pholeter^ Collyriclum, Troglotreina. Family. Echinostomidae, Looss, 1902. More or less elongated flukes^ small or very large, much flattened anteriorly, less so posteriorly, or even round. Suckers near one another, the anterior small and weak, the posterior large and powerful directed obliquely backwards. Stcrrounding the oral sucker dorsally and laterally but 7tot venirally is a fold or " collar '' bearing a row or rows of pointed spi?tes which are continued round laterally o?i to the ventral corners, the number being constant for each species, the corner spities large or specialized, skin anteriorly scaled or spiny. Alirne7ttary canal consists of a pharynx, epithelial '''• pseudo-ossophagus " and gut cceca reaching to posterior end. Testes behind one a?tother in hind body. Ovary on right side or median directly in front of the testes. Vitellaria lateral, usually extending to the hind end and not beyofid the ve?ttral sucker anteriorly. Genital pore fust in front of ventral sucker. Uterus i?i transverse loops. Genital sinus absent or present. Receptaculum seminis and Laurer's canal prese?it. Eggs thin shelled and large, bright yellow, 6$ /x to 120 /j. long. Excretory bladder ^ -shaped. Parasitic in gut of vertebrates, especially birds. Sub-family. Echinostominae, Looss, 1899. Cirrus sac usually reaching to centre of ventral sucker, but not beyond. Cirrus long, usually without spines, coiled when retracted. Sejninal vesicle tubular, twisted. On the head a ventral uniting ridge between the angles of the collar. Dorsal circlet of spines, siftgle or double, not interrupted unless the collar itself is dorsally divided. Genera : Echinostoma, etc. Sub-family. Himasthllnae, Odhner, 1910. Cirrus sac reaching far beyond ventral sucker. Cirrus armed with strong rose-thorn-shaped hooks. Vesicula seminalis tubular not coiled. Cervical collar not continued across ventral aspect. Spines on collar in one row. Body armed with fine needle-shaped spines. Family. Schistosomidae, Looss, 1899. Sexes separate. Genital pore behind the ventral sucker. Ventral sucker elevated above the surface. Pharynx absent. Gut forks reunite to form a single stem. In $ four or more testicular follicles. In ? a single ovary, just in front of the union of the gut forks. Vitellaria on either side of the united gut stem. 234 THE ANIMAL PARASITES OF MAN The Trematodes Observed in Man. Family. Paramphistomldae, Stiles and Goldberger, emend. 1910. Sub-family. Cladorchiinae, Fisch., 1901. Genus. Watsonius, Stiles and Goldberger, 1910. CladorchmcE.—^ody pyriform. Ventral pouch absent. Acetabulum ventral or (?) ventro-subterminal, very large, margins projecting, aperture small. Genital pore in front of bifurcation of gut, not surrounded by a sucker ; ductus herma- phroditicus apparently absent. Excretory pore at posterior end of excretory vesicle, behind Laurer's canal. Oral sucker with a pair of irregularly globular suctorial pouches ; oesophagus thickened distally ; caeca long, not wavy ; end in acetabular region. Male Or£-a;is.— Testes two lobed, smaller than acetabulum ; longitudinally, nearly or quite coinciding ; transversely they abut or slightly overlap ; preovarial in equatorial and caudal thirds. Pars musculosa not largely developed; cirrus pouch absent. Female Organs. — Ovary and shell gland post-testicular. Vitellaria extend from gut fork to slightly beyond gut ending ; uterus interccecal, partly post-testicular. Laurer's canal in front of excretory vesicle. Type Species. — Watso?tius watsoni, Conyngham, 1904. Watsonius watsoni, Stiles and Goldberger, 1910. Syn. : Ainphistojuuni -£v^/J•'h/itirn \jCkY /vcrM'htinr/i T rvricc iRnA Fasciola hepatica var. cBgyptiaca^ Looss, 1 896. This species is closely allied to Fasciola hepatica, but is distin- guished by its elongated body, short cephalic cone, almost parallel sides, larger ventral sucker, which is also closer to the oral sucker, and by its larger eggs. Length up to 75 mm., width up to 12 mm. Oral sucker i to 1*2 mm., ventral sucker up to 17 mm. in diameter. Eggs 150 />fc to 190 /A long by 75 //, to 90 //, broad. Habitat. — Bile-ducts of Camelopardalis giraffaj Bos taunts, Bos indicus, Bos bubalis^ Ovis aries and Capra hitcns. Distribution. — Africa. This species has once been observed in man by Gouvea, in Rio de Janeiro, in a French naval officer who became ill with fever, cough and slight blood-spitting. The lungs were normal except for a ' In the English translation of Kiichenmeister's work on Parasitology (London, 1857). The specimen is preserved in the Hunterian Museum, London, and is an adult liver fluke, measuring 18 mm. in length and 7 mm. in breadth. FASCIOLOPSIN^ 245 sharply circumscribed spot at the base of the left lung. Twenty days later during a fit of coughing the patient spat up a fluke 25 mm. long, characterized by its slender aspect and by the size of its ventral sucker, and its close proximity to the oral sucker. Considering the fact that Gouvea's patient had spent many weeks in July of the same year in Dakar (Senegambia), where according to Railliet Fasciola gigantica is common in slaughtered animals, and considering also the characters of the fluke, Railliet rightly assumes that one had to do with the African giant fluke and that the patient had infected himself in Dakar. Sub-family. Fasciolopsinae, Odhner, 1910. Genus. Fasciolopsis, Looss, 1898. Ventral sucker large, and elongated posteriorly injto a sac. Cirrus pouch long and cylindrical, its greatest length being occupied by the sinuous tubular seminal vesicle, on which exists a peculiar caecal appendage. Laurer's canal present. Fasciolopsis buski, Lank., 1857. Syn. : Distomuin buski, Lank., 1857 ; Dist. crassum, Cobbold, i860, nee v. Sieb., 1836. The length of the body varies ; it may measure 24 to 37 or even attain 70 mm.; the breadth is from 5*5 to 12 to 14 mm. In the pig the fresh parasites measure, smallest, 12 to 8 mm. ; largest, 35 to 16 mm. (Mathis and Leger). Skin without spines, but according to Heanly always present in man and pig specimens. The oral sucker measures 05 mm. in diameter; the ventral sucker is three to four times as large ; the pharynx is globular, 0*7 mm. in diameter ; the prepharynx is provided with a sphincter ; the intestinal caeca ex- tend to the posterior border with Fig. 146. — Fasciolopsis buski, Lank. F.j., ventral sucker ; C.p., cirrus pouch ; /., inles- tinal fork; 6'.2/., vitellaria ; 7'., testes; 0., ovary; Ms., sucker; Shg., shell gland; 6^/., uterus. Magnified. (After Odhner.) 246 THE ANIMAL PARASITES OF MAN two characteristic curves, one at the anterior border of the anterior testis, the other between the two testes. The genital pore is at the anterior border of the ventral sucker; the cylindrical cirrus pouch extends from behind the ventral sucker to half-way to the shell gland. The seminal vesicle extends forwards within the cirrus pouch as a convoluted tube. From its anterior portion is given off the caecal appendage, which has itself short lateral diver- ticula. It runs backwards, ending blindly about o*5 mm. from the posterior end of the cirrus sac. The seminal vesicle is continued as the pars prostatica (?) 0*5 mm. long, and this by the very short ejaculatory duct (13 /x), and finally by the fairly long cirrus, which is beset with very fine spines except at either extremity. The ovary and shell gland are situated at about the middle of the body with the testes behind them, and the uterus in front. The vitellaria extend from the ventral sucker to the posterior border. The eggs measure 120 fi to 130 /i in length and 77 fiioSo fi in breadth, and resemble those of Echinochasma sp. in dogs. The larval stages are said to occur in shrimps. Habitat. — Intestine of pig and man. Distribution. — In man : India, Siam, China, Assam, Sumatra. It is common in Cochin China (16 out of 133 Annamites, Noc), in Tonkin very rare. Dr. J. Bell has sent me [J. W. W. S.] human specimens from Hong Kong. In pigs : very common in South China (Heanly). Common in pigs in Hong Kong. Sixteen out of 248 pigs (i.e., 6 per cent.) infected in Hanoi. • Fig. 147. — Fasciolopsis rathouisi^ Poir. : the mouth at the top, and under it the genital pore and ven- tral sucker, behind which again is the uterus. The vitellaria are at the sides, and posteriorly in the central field the ramified testes ; the ovary is in front of the right testis. (After Claus.) Fasciolopsis rathouisi, Ward, 1903. Syn. : Distomum rathouisi^ Poirier, 1887. Fifteen to 19 mm. long by 8*5 to 10-5 mm. broad by about 3 mm. thick. Skin with spines (Leiper). Bluntly oval or elliptical with short cephalic cone which is absent in Fasciolopsis biiski. Oral sucker subterminal, 0*25 to o'29 mm. broad by 0*2 mm. in antero-posterior diameter. Distant from ventral sucker by about twice its diameter. Ventral sucker 1-32 to 1*38 mm. broad by 0*68 to 07 mm. in antero- posterior diameter. (Esophagus extremely short. Cirrus sac not conspicuous and straight as in Fasciolopsis buski, but is convoluted. Testes one behind the other (according to Poirier they lie beside one FASCIOLOPSIS FULLEBORNI 247 another), more compactly branched, broader and denser than in Fasciolopsis bnski. Ovary on right side, small, coarsely branched. Uterus in broad, closely grouped coils, packed with ova anterior to ovary. Vitellarian acini more numerous and somewhat differently distributed. Eggs 150/^ by 80 yit, thin shelled. [H. B. Ward, who has examined this species, and from whose account the above is mainly taken, considers that it is a good species, although the differences between it and Fasciolopsis buski are slight, while Odhner, who examined the original species, is of the opposite opinion. — J. W. W. S.] The parasite appears to cause diarrhoea, wasting and occasionally jaundice. Habitat. — Intestine of man. Distribution. — China, common in some parts (Goddard). Fasciolopsis goddardi. Ward, 1910. Twenty-one to 22 mm. long, 9 mm. broad. Skin with spines (Leiper). Uterus very closely coiled, most striking character is the large size of the vitelline acini. Imperfectly known. Distribution. — China (Shanghai). Fasciolopsis fulleborni, Rodenwaldt, 1909. The fully extended fluke is tongue-shaped, 50 by 14 mm. ; two contracted specimens measured 40 by 15 mm. and 30 by 16 mm. respectively. Skin without spines, with according to Leiper cephaHc cone not clearly defined. Oral sucker circular, 075 mm. in diameter, slightly larger than that of Fasciolopsis buski. Ventral sucker 2-6 mm. in diameter (that of Fasciolopsis buski i*6 to 2 mm.). Length 2*9 mm. (as in Fasciolopsis rathoulsl), the excess of length over breadth being due to the posterior elongated sac-like prolongation of the sucker. Prepharyngeal sphincter present. Pharynx 07 mm. in diameter. CEsophagus practically absent. Gut caeca similar to those of Fasciolopsis buski. Testes — regularly branched, separated by an incurving of the caeca, the anterior occupying a smaller area than the posterior. Ovary — very small, as in Fasciolopsis buski, on the right side. Shell Gland — almond-shaped, 2-3 by 1*2 mm. In Fasciolopsis buski it is round and smaller, i to i"5 mm. in diameter. Vltellarla — similar in distribution to those of Fasciolopsis buski, but the acini are strikingly small. Cirrus Sac — is the most characteristic feature of this species. It is a powerfully built, convoluted sac standing out clearly on the body. 248 THE ANIMAL PARASITES OF MAN t^* Oral sucker ■ Prepharynx Pharyny ':^*^ Vl'A7fra/ sucker Vagina Cut Cirrus sac (with seminal vesicle) ' Uterine coils - Receptaculum seminis " Shellgland and yolk receptacle "~ Ovary ---Gut r:- Testes l--;-' Excretory system y Vitellaria Excretory pore (Dorsal) Fig. i/\%.—Fasciolopsi5fulleborni, ventral aspect. {After Fiilleborn.) TROGLOTREMID^ 249 It is not a uniform, straight cylinder 0*25 to 0*33 mm. in diameter, as in Fasciolopsis hiiski, but even in fully extended flukes is typically convoluted. It is i mm. thick in the middle, but in other parts varies much from this. The posterior end of the cirrus sac is- at two-thirds or more of the distance from ventral sucker to shell gland. In the case of Fasciolopsis bnski the posterior end of the sac only extends half-way. Seminal Vesicle — has a peculiar convoluted, saccular and angular course, but the caecal appendage characteristic of the genus appears to be absent ! Excretory System. — The main stem gives off very regular transverse branches which are well seen posteriorly. Eg^s. — 100 //, by 73 //,. Thin shelled. Habitat. — Intestine. Mahommedan from Calcutta. [It is evident that a re-examination of fresh material is required before the validity of all these species can be accepted. — J. W. W. S.] Family. Troglotremidae, Odhner, 1914. ' Genus. Paragonimus, Braun, 1899. Body egg-shaped or somewhat elongated, generally more broadly rounded in front than behind. Covered all over with spear-shaped spines arrangedin groups. Gut caeca winding with dilatations or constrictions in parts. Ventral sucker in or in front of the middle of the body. Excretory bladder cylindrical, very long and broad, reaching in front to the bifurcation of the gut. The lateral excretory canals join the bladder only a little in front of the excretory pore. Genital pore median just behind the ventral sucker. Genital sinus duct-like. Cirrus sac absent. Male terminal organs very small. Ejaculatory duct present. Testes and ovary deeply lobed, the testes in or just behind the middle, the ovary somewhat laterally placed just behind the ventral sucker. Uterus forms a coil behind the ventral sucker. Eggs rather large, thin shelled, the ovarian cell still unsegmented on deposition. Receptaculum seminis, small. Parasitic in the lungs of mammals, enclosed in cyst-like cavities, generally in pairs. Type species. — P. westermajiii in the tiger. Paragonimus ringeri,' Cobb., 1880. Syn. : Distoma ringeri, Cobb., 1880; Distoma pulmonale., Baelz, 1883 ; Distoma pulmo7iis, Suga, 1883. The body is of a faint reddish-brown colour and plump oval shape. The ventral surface a little flattened; 7*5 to 12 mm. in length, 4 to 6 mm. in breadth, and 3-5 to 5 mm. thick (in man). The oral sucker (075 mm.) is subterminal; the ventral sucker (o*8 mm.) somewhat in front of the middle of the body. Pharynx spherical, 250 THE ANIMAL PARASITES OF MAN 0*3 mm. in diameter, or 0*4 by 0*3 mm. ; oesophagus, 0*02 mm. ; intestinal caeca convoluted, asymmetrical, the first part having the same structure as the oesophagus. The cuticle is covered with spines in groups ; the excretory pore opens at the posterior end rather on the ventral surface, the excre- tory ducts open into the elongated bladder at the hind end near the pore. Genital pore behind • the ventral sucker and median. Genital sinus 0*2 mm. long with thick wall, ejaculatory duct o'i3 mm., pars prostatica 0*2 mm., seminal vesicle duct-like of irregular outline. Behind the sucker the ovary on the left, and the closely packed uterine coil on the right (though amphitypy of these two organs is common) ; the two irregularly lobed testes lie side by side posteriorly. Vitellaria extensive, leaving only a median dorsal and ventral space Fig. 149. — Paragonimus ringeriy Cobb. : to the right, dorsal aspect ; to the left, ventral aspect. Natural size. (After Katsurada.) Fk;, 150. — Paragonimus ringeri, Cobb. : diagram of the internal organs. a, oeso- phagus ; 3, vitellaria (a portion only shown) ; f, common genital duct ; d, shell gland with oviduct, Laurer's canal and vitelline duct ; e, ovary; y, vitelline receptacle; g, excretory pore ; A, oral sucker ; i, pharynx ; k^ gut*; /, ventral sucker; w, uterine coils; «, vitellarian ducts ; (7, vas efferens ; /, testis. (After Kubo. ) Fig. 150A. — Paragonimus wesiermanii. Kerb. : seen from the ventral surface. Mouth, pharynx, intestinal caeca, at the sides of which the vitellaria are observed. The genital pore is behind the ventral sucker, and next to it, on the left, the ovaty ; on the right, the uterus ; the two testes posteriorly ; the excretory vessel in the middle. lo/l. (After Leuckart.) free. Seminal receptacle probably absent ; Laurer's canal present. The eggs are oval, brownish-yellow, fairly thin shelled, and measure onf-an average 81 '2 /a by 49*2 /x. The following species are also known : — P. westennaiiii, Kerb., 1878, in the tiger, and P. kellicoiti, Ward, 1908, in the pig, dog, and PARAGONIMUS RINGERI 251 cat (N. America). Ward and Hirsch give the following differences between the spines of the three forms : — P. ringeri. P. westermanii. P. kellicotti. Shape Distribution Chisel-shaped, mod- erately heavy. Circular rows, in Lancet-shaped, very slender. Circular rows. Chisel-shaped, heavy. Circular rows, groups. in groups. singly. Two other species, P. riidis, Diesing, 1850, in a Brazilian otter {Ltitra brasiliensis)f and P. compachis, Cobbold, 1859, in the Indian ichneumon, are but little known. Habitat. — Lungs, pleurae, and especially the bronchi of man and dog. The alleged occurrence (of eggs) in other organs may be due to confusion with those of Schistosoma japoniciim. Distribiition. — China, Korea, and especially in Japan, where, accord- ing to Katsurada, there are no districts that are entirely free from pul- monary flukes. The mountainous provinces of Okayama, Kumamoto, Nagano and Tokushima are the principal centres. Pathology. — The number present in the lung varies from two to twenty, about. Usually one cyst contains one worm, but in the dog each cyst contains two. The cysts admit the tip of the finger, and have a fibrous wall I mm. thick. They origin^ate partly from dilatation of bronchi and bronchioles. Others arise from the inflammatory reaction of lung tissue into which the worms have wandered. The worms and their eggs cause bronchitis and peribronchitis, catarrhal, haemorrhagic, or purulent, and areas of consolidation. Areas containing eggs in their centre resembling tubercle nodules are not uncommon, and extensive cirrhosis of the lung may be found. As a result of these changes, emphysema and bronchiectasis also occur. As to the development, only the following details are known : that the eggs, which before segmentation of the ovum reach the open in the sputum and through being swallowed also in the faeces, develop in water into a miracidium ciliated all over, which hatches and swims about freely. According to Manson this takes place in four to six weeks. Fig. 151. — Egg of Para- gonivius ringeri, Cobb. , from the sputum. Showing the ovarian cell and vitel- line cells and granules. 1,000/1. (After Katsurada.) 252 THE ANIMAL PARASITES OF MAN Sub-family. Opisthorchiinae, Looss, 1899. Genus. Opisthorchis, R. Bl?.nch., 1845. OpisthorchiinjE with lobed testes. Laurer's canal present. Parasitic in the bile- ducts of mammals and birds. Opisthorchis felineus, Riv., 1885. Syn. : Distoma co7ius, Gurlt, 1831 {tiec Creplin, 1825); Distoma lanceolatum^ V. Sieb., 1836, V. Tright, 1889 {?iec Mehlis, 1825 = Fasciolo lanceolata^Kud.^ 1803); Distoma sibiricum. Winogr., 1892 ; Distoma tenuicolle, Miihl., 1896. This parasite is yellowish-red in the fresh condition, and almost transparent. The body is fiat, with a conical neck at the level of the ventral sucker marked by a shallow constriction ; this, however, is only noticeable in fresh and somewhat contracted specimens. Posteriorly to the ventral sucker the lateral borders run fairly parallel ; the posterior end is either pointed or rounded off. The length and breadth vary according to the contraction, being usually 8 to 11 mm. by 1*5 to 2 mm. The suckers are about one-fifth to one-sixth of the length of the body distant from each other, and of about equal size (o'23 to 0*25 mm.). The oesophagus is hardly any longer than the pharynx, which lies close behind the oral sucker ; the intestinal caeca reach almost to the posterior border and are often tilled with blood. The excretory pore is at the posterior extremity, and the excretory bladder forks in front of the anterior testis. The testes in the posterior fourth of the body lie obliquely one behind the other ; the anterior one has four lobes, the posterior one five lobes ; the ovary is in the median line transversely, simple or slightly lobed ; behind it lies the large pear- or retort-shaped recepta- culum seminis and Laurer's canal. The uterus is in the median field. The vitellaria occupy the fairly broad lateral areas, in about the central third of the body, beginning behind the ventral sucker and terminating at about the level of the ovary ; the acini are small and arranged in groups of seven to eight, separated by interstices. The genital pore is close in front of the ventral sucker. The ^ggi^ are oval with sharply defined operculum at the pointed pole, 30 yit by 1 1 yit. This species, which is frequently confused with others, inhabits the gall- bladder and bile-ducts of the domestic cat especially ; but is also found in the dog, in the fox, and in the glutton {Gulo borealis). It has been observed in France, Holland, North Germany (being particularly frequent in East Prussia), in Russia, Scandinavia, Siberia, Japan, Tonkin, Hungary, and Italy. The North American form (from cats and Ca7iis latrans) is a distinct species {Opisthorchis pseudofelineus) . In man this species was first found by Winogradoff in Tomsk (nine cases), then by Kholodkowsky in a peasant from the neighbourhood OPISTHORCHIS FELINEUS 253 of Petrograd who had travelled a great deal in Siberia, and finally by Askanazy in five persons who were natives of the East Prussian district of Heydekrug. In Tomsk, Opisthorchis fellnetis is the most frequent parasite of man that comes under observation ^t post inorteni (6*45 per cent), whereas Tcviiia sagiuafa has only been found in 3*2 per cent., Echinococcif^ in 2*4 per cent., Ascaris himbricoides in i'6 per cent., and Oxyuris vennicidaris in 0*8 per cent, of the autopsies. In the district of Heydekrug, however, the species in question is also frequent, as in a few years five cases came to our knowledge (of which three were diagnosed by the discovery of the eggs in the faeces). In none of Winogradoff's nine cases had the death of the patient been caused direct by the parasites, yet more or less extensive changes in the liver were found in all of them; such as dilatation of the bile-ducts with inflammation and thickening of their walls, and foci of inflammation or atrophy in the liver substance ; icterus was present five times and atrophy of the liver an equal number of times ; ascites was observed three times, and in two cases, probably of recent date, the organ was enlarged. The num- ber of parasites found fluctuated between a few and several hundreds. In two of Askanazy's cases, which he examined more closely, carcinoma which had developed at the places most invaded by flukes was found at the posUmortem., so that perhaps there may be grounds for the connection which the author seeks to establish between cancer of the liver and the changes induced by the parasites ; these changes consist of numerous and even ramified '^^ r.s. Fig. 152.— Egg of Opis- thorchis feli- neus, Riv. 830/1. Fig. 153. — Opisthorchis felineus : from the cat. m., mouth; p.6., pharynx ; i., gut ; ^./., genital pore ; ac, ventral sucker ; ut., uterus ; v.g., vitellarium ; ov., ovary; s.g., shell gland; r.s., receptaculum seminis ; t. testes ; ex. /., excretory pore. (After Stiles and Hassall.) 254 THE ANIMAL PARASITES OF MAN m s.g. L.c. t. 7lt. v.g. v.df. r.S. / ex. c. ex. p. proliferations of the epithelium of the biliary duct into the connective tissue, which is likewise proliferated. The number of worms found in one case amounted to over lOO ; in a second case, in which the parasites had also invaded the pancreatic duct, their number was even larger. Winogradoff as well as Askanazy found isolated flukes in the intestine also. Unfortunately, nothing much is known of the history of the develop- ment of Opisthorchis felmeus ; we only know that when deposited the eggs already contain a ciliated mira- cidium, which, however, according to my experience, does not hatch out in water, but only after the entry of the eggs into the intestine of young Limriceus stagnalis ; no further de- velopment, however, occurs. Wino- gradoff states that he has seen the miracidia. hatch after the eggs had been kept in water for a month at 37° C. ; and has even observed free miracidia in the bile of man and of a dog respectively. Although the whole post-embryonal development of the cat fluke remains yet to be investigated, Askanazy by a series of experiments on cats and dogs has dis- covered the mode of infection. The intermediate hosts are fish, and mainly the ide, in this country called Tapar (Idus melanoiuSj H. and Kr.), and of subsidiary importance the Fig. 154. — Opisthorchis psiudofdineus : from the bile-duct of the cat (Iowa), w., oral sucker; p.h.^ pharyngeal bulb ; ^j., oesophagus ; i., intestine; va.^ vagina; g.p.m., male orifice; ac, ventral sucker; «/., uterus ; v.g., vitellarium ; s.g., shell gland ; v.dt.^ vitelline duct ; ov. , ovary ; r.s. , receptaculum seminis ; L.c, Laurer's canal; /., testis; ex.c, excretory bladder; ex.p., excretory pore. (After Stiles.) PAROPISTHORCHIS 255 roach (Leiiciscus rutiltis). Both species of fish as well as others are readily eaten raw by man on the Courland lagoon (Baltic). It is, more- over, significant that those persons whom Askanazy found infected with the cat fluke were also infected with Dihothriocephaliis latus, the intermediate host of which is also fish (Lota sp., Esox sp., Perca sp.). In one of his nine cases Winogradoff also saw a small fluke covered all over with spines, which he conjectured to be the young stage of Opisthorchis felineiis ; as, however, according to my experi- ence, this species, even in smaller specimens, is always without spines, the above hypothesis cannot be accepted. It is much more probable that one of the other species that also invade the liver of cats may accidentally be introduced into man; we know, in fact, that Metorchis albidtis, Braun, and Metorchis truncatus, Rud., are both covered with spines. As, however, the spines of the first-named species are rather apt to fall off, and also as it possesses a different shape (spatula-shaped), it may be assumed that probably Winogradoff had found Metorchis truncatiis, Rud., 18 19, in his patient. Genus. Paroplsthorchis, Stephens, 1912. Structure as in Opisthorchis, except that the ventral sucker and genital pore occur on the apex of a process or pedicle projecting from the anterior portion of the body. This process is about ^ mm. long, and is retractile. Paroplsthorchis caninus, Barker, 1912. Syn. : Distoma co?ijunctum^ Lewis and Cunningham, 1872 ; Opisthorchis noverca^ M. Braun, 1903 (^pro parte) ; Opisthorchis caninus^ Barker, 19 12 (?). Length varies from 275 to 575 mm. in preserved specimens, average 3*6 to 5'2 mm. Body uniformly spinose, though as a rule spines are not present on the pedicle. Body slightly concavo-convex, the con- cavity being ventral. Oral sucker 0*28 mm. Pharynx 0*224 by 0'i84 mm. (Esophagus 0*04 mm. Ventral sucker 0"i76 mm. in diameter. Pedicle about \ mm? long, may be completely retracted. Genital Pore — opens on the apex of the pedicle in front of the ventral sucker. Its exact position varies with the state of contraction of the parts. In certain cases it actually opens within the cuticular border of the sucker, in other cases it opens externally to the sucker and anterior to it. The opening is covered with scales. The vas deferens and uterus run alongside one another until they merge near the apex of the pedicle into a common sinus. Vitellaria — consist of eight acini on each side, extending from slightly behind the base of the pedicle to the anterior border of the ovary, or as far back as a line separating the posterior border of the ovary from the anterior border of the anterior testis. 256 THE ANIMAL PARASITES OF MAN V. .ex. lat V. seui. P. ex. Fig. 155. — Paropisthorchis caninus : from the bile-ducts of the pariah dog, India. AceL v.y ventral sucker ; £//., uterus ; V. ex. /a/., longitudinal excretory duct ; V. sem., seminal vesicle; Se?n. r««' by 53 //, to 82 /a. Average.— gg'SH'hy 56/^. Habitat. — Gut of man (Filipinos), Philippine Islands. Echlnostoma malayanum, Leiper, 191 1. Twelve millimetres long, 3 mm. broad, 1*3 mm. thick. Ends bluntly rounded. At the anterior end a ventral furrow on either side, one- third the width of the body, marking off the circum-oral collar. Along its edge is a row of forty- three spines extending across the middle line dorsally but not ven- trally. The spines vary in size from Fig. 168. —Echinostoma ilocanum. Vo.^ oral sucker ; Ph.^ pharynx ; Cirre, cirrus ; F.z;., ventral sucker; 6V., uterus; G.c.^ ovary; Ov., shell gland ; T., testes; 7".^., vitellarium ; C.ex,, excretory vesicle. (After Erumpt.) Fig. 169. — Echinostoma ilocanum^ Garrison, 1908 : head end showing collar of spines, ventral view. (After Leiper.) 0*07 mm. in length (ventrally) to 0*05 to o'oi6 mm. (dorsally). Cuticular spines also exist on the ventral side as far back as posterior end of body, but dorsally limited to a triangular area ending in front of the ventral sucker. Oral sucker 0*07 mm. thick, occupying the middle HIMASTHLIN.E 2fy) third of the circum-oral disc; pharynx 0*25 mm. in diameter; oeso- phagus 0-04 mm. long ; gut caeca simple, extending to end of body ; ventral sucker 0*9 mm. long by 075 mm. broad by 07 mm. deep ; wall about 0*25 mm. thick. The sucker is inclined at an angle of 40° to the ventral surface. Testes lobed, one behind the other, behind the ventral sucker. Cirrus pouch well developed, reaching to the posterior edge of the sucker. Genital pore in the angle between neck and anterior lip of ventral sucker. Ovary smooth, 0-3 mm. in diameter, 0*85 mm. behind ventral sucker. Vitellaria very numerous, extending from posterior margin of sucker to posterior end of body, where they intermingle. Eggs few in numb :r, brown and large. Habitat. — Gut of man (Tamils), Malay States. Fig. 170. — Echinostoma malayanum, Leiper, 1912 : anterior end showing collar of spines, (After Leiper.) ventral view. Sub-family. Himasthlinae, Odhner, 1910. Genus. Artyfechinostomum, Clayton-Lane, 1915. Crown of thirty-nine spines, continuous over dorsum. Two corner spines long. Vitellaria extend from posterior margin of sucker to pos- terior end of fluke. Eggs without filament. [Although the possession of strong rose-thorn hooks is given by Odhner as a sub-family charac- teristic, yet in this genus assigned to this sub-family they have not been seen.— J. W. W. S.] Artyfechinostomum sufrartyfex, Clayton-Lane, 1915. Spirit specimens : 9 by 2*5 by o'8 mm. thick. Ventral sucker conspicuous, i mm. in diameter. Cirrus sac 2 mm. long. Testes lobed, about 1*5 mm. in diameter. Posterior border of posterior testes I mm. from posterior end. Vitellaria meet posteriorly behind the posterior testis. Family. Schistosomidae, Looss, 1899. Genus. Schistosoma, Weinl, 1858. Syn. : GyncEcophorus, Dies., 1858; Bilharzia, Cohh., 1859; Thecosoma^ Moq. Tandon, i860. The males have bodies that widen out considerably behind the ventral sucker, the lateral parts of which in-roll ventrally, forming the almost completely closed canalis gynsecophorus, within which the female is enclosed. There is no cirrus pouch. The male has five or six testes, the females are filiform ; the uterus is long. 270 THE ANIMAL PARASITES OF MAN There is no Laurer's canal. The ova almost equally attenuated at either extremity ; they have a small terminal spine, and are not provided with a lid. They contain a miracidium, ciliated on all sides, which is characterized by the possession of two large glandular cells, which discharge anteriorly beside the. gastric sac. They live in the vascular system of mammals. (An allied genus [Bilharziella] lives in the blood-vessels of birds.) Schistosoma haematobium, Bilharz, 1852. Syn. : Distoma hceynatobiujn^ Bilh. ; Distoma capense^ Harley, 1864. The Male is whitish, 12 to 14 mm. in length, but is already mature when 4 mm. long. The anterior end is o*6 mm. or a little over in length. The suckers are near each other, the oral sucker is in- fundibular, and the dorsal lip is longer than the ventral one. The ventral sucker is a little larger, 0*28 mm., and is pedunculated. A little behind the ventral sucker the body broadens to a width of I mm., decreasing, however, in thickness ; the lateral edges in-roll ventrally, so that the posterior part of the body appears almost cylindrical, 0*4 to 0*5 mm. in diameter ; the posterior extremity is somewhat more attenuated. The dorsal surface of the pos- terior part of the body is covered with spinous papillae. There are delicate spines on the suckers, and largerones invest the entire internal surface of the gynaecophoric canal, as well as a longitudinal zone at the edge of that side of the external surface that is covered by the other side rolling over it. The oeso- phagus is beset with numerous glandular cells (fig. 173), and pre- sents two dilatations; the intestinal bifurcation is close in front of the ventral sucker, the two branches uniting sooner or later behind the testes into a median trunk, which may again divide at short intervals. The excretory pore is at the posterior end, but placed somewhat dorsally ; the genital pore is at the beginning of the gynaecophoric canal, thus Fig. \T\.— Schistosoma hcejftatobtum,B\\. : male carrying the female in the canalis gynge- cophorus. 1 2/ 1. (After Looss.) SCHISTOSOMA H^MATOBIUM 271 Fig. 172. — Tiansverse section through a pair of Schistosoma hcematobium in copula. In the male the p6int of reunion of the intestinal forks has been cut across. (After Leuckart.) O.s. behind the venti*al sucker ; into it opens the vas deferens which, posteriorly, broadens into the seminal vesicle and then continues as the vasa efferentia of the four or five testes (fig. 173). The Female — filiform, about 20 mm. in length, pointed at each end, and measuring 0*25 mm. in diameter in the middle. Their colour varies accord- ing to the condition of the contents of the intestine. (Posteriorly they are dark brown or blackish.) The cuticle is smooth except in the sucker, where there are very delicate spines, and at the posterior end, where there are other larger spines. The oral sucker is a little larger than the pedunculated ventral sucker (0*07 and 0*059 mm. respectively). The anterior part of the body, 0*2 to 0*3 mm. in length ; the oesophagus is as in the male. The intestinal bifurcation is in front of the ventral sucker, the two branches uniting behind the ovary and the trunk running in a zigzag manner to the posterior border. There are indications of diverticula at the flexures. The ovary is median. In young females it is of an elongated oval shape ; in older females the posterior end becomes club-shaped, whereas the anterior end becomes attenuated ; the oviduct origin- ates at the posterior end, but immedi- ately turns forwards and joins the parallel vitelline duct in front of the ovary (fig. 174), where the shell gland cells open ; the common canal becomes dilated to form the ootype, and then proceeds as the uterus, with only slight convolutions, along the central field to the genital pore, which lies in the middle line immediately behind the ventral sucker. The single vitellarium starts behind the ovary and extends to the posterior end. The acini are situated at the sides of the excretory duct, which runs a median course. The eggs are compact spindles, much dilated in the middle ; they have no lid, and are provided with a terminal spine (rudimentary filament) at the posterior end, measuring 120 yu, to 150 /I, in length and 40 /^ to 60 /a in breadth, but vary in size and shape (fig. 175;. Oe. _ V.s.~- Fig. 173. — Anterior end of the male Schistosoma hcematobium, Bilh. V,s., ventral sucker ; /., gut cceca ; G.p., genital pore ; T., testes ; O.s.y oral sucker ; Oe., cesophagus with glandular cells; V.s., vesicula seminalis. 40/1. (After Looss.) 272 THE ANIMAL PARASITES OF MAN Distribution. — In order to understand the distribution of the worms and eggs in the body, it may be well to recall the blood supply of the abdominal and pelvic organs. It is generally assumed that the early life (?cercarial stage) of the worms occurs in the liver, and that the young worms travel from here, where they are invariably found, to their various sites along the portal vein and its tributaries and so against the blood stream. The tributaries of the portal vein are : — (i) Superior mesenteric, the tributaries of which are : (a) the veins of the small intestine ; (6) ileo-colic ; (c) right colic ; (d) middle colic ; (e) right gastro-epiploic ; and (/) inferior pancreatic. By these paths infection of the small intestine, ascending and transverse colon and pancreas would occur. (2) Splenic. (Ova have been recorded by Symmers in the spleen.) (3) Inferior mesenteric, the tributaries of which are (a) superior haemorrhoidal veins from the upper part of the haemorrhoidal plexus; (6) sigmoid veins from sigmoid flexure and lower portion of descending colon ; (c) left colic vein draining descending colon. The superior haemorrhoidal veins form a rich plexus in the rectum, and below this level in the upper and middle parts of the anal canal. The plexus forms two networks, an internal plexus in the submucosa and an external on the outer surface. The internal plexus opens at the anal orifice into : (a) branches of the inferior haemorrhoidal vein (from the pudic) ; (b) the external plexus. The external plexus gives off : (a) inferior haemorrhoidal opening into internal pudic (of internal iliac vein) ; (b) mid-haemorrhoidal into internal iliac or its branches ; and (c) superior haemorrhoidal opening into inferior mesenteric. The external plexus further communicates with the vesico-prostatic plexus. The vesico-prostatic (vaginal) plexus opens into the vesical veins, which drain into the interior iliac vein. This plexus also receives afferents from the pudendal plexus, the chief tributary of which is the dorsal vein of the penis. The pudendal plexus also receives branches from the inferior pudic and the anterior surface of the bladder. There is thus a communication between the portal vein and the vena cava by means of these plexuses, viz., through the inferior and middle haemorrhoidals, and by the inferior haemorrhoidals to the bladder and thence by the vesical veins or the pudic to the caval system (interior iliac). It is thus by the inferior mesenteric and its tributaries that the worms reach the descending colon, rectum, anal canal, and eventually the bladder, and in some cases the caval system. Before considering what is actually found post mortem in these veins and the organs drained by them, we may further recall the fact that the calibre of ''medium" veins is 4 to 8 mm., ''small" veins SCHISTOSOMA H^MATOBIUM 273 less than 40//, in diameter and capillaries Syu, to 20 /x. Further, the maximum diameter of the male worm is 1 mm., that of the female 2S0 fjL and eggs /// uiero So fi to 90//, long by 30//, to 40//,. Liver and Portal Vein. — Here worms are most easily found post mortem. Often only males are found and these of the same size, and if females occur only a few worms are found in copula. The worms are frequently not full size and the males may contain no free spermatozoa in their testes, and as regards the females some may be fertilized, others not, as shown by the presence or absence of spermatozoa in the seminal receptacle or uterus. In either case they may contain eggs — lateral-spined — usually one, less often two, but there may be as many as five or six. These eggs may also show some abnormality, which takes the form of : (i) abnormal con- tents, viz., disintegrating yolk cells with or without an ovarian cell ; (2) abnormal shape but with normal contents and probably repre- sented by the collapsed and empty egg-shells which are found in the tissues. As to the interpretation of these facts, Looss believes that these lateral-spined eggs are products of young females whose egg-laying is not at first properly regulated. The shape that the eggs take, viz.y with a lateral spine, is determined by an excess of material — ovarian and yolk cells — being present in the ootype. The shape of eggs depends upon the position they have in the ootype during their formation. In young females an excess of cells — yolk cells especially — accumulates, distending not only the dorsal wall but a portion also of the short duct joining the ootype to the uterus. The result of this is that the axis of the ootype and egg is almost transverse to the body, and the posterior funnel-shaped portion of the ootype, instead of being terminal, has now a lateral or rather a ventral position, so that the spine which occupies this portion, instead of being terminal, is now lateral. It is noteworthy that these lateral-spined eggs are thicker, owing to the excess of material present, and not uncommonly have a curved anterior border, due to a projection of the anterior end into the anterior opening of the ootype. As these eggs are being laid by females in the portal vein they are carried back to the liver by the blood stream. The liver is one of the commonest sites for these eggs ; also terminal-spined eggs may be found here for the same reason. Hcemorrhoidal Veins. — Mature worms, generally in copula, are usually found here, though young not fully grown females may also occur. The tissues of the rectal wall (or colon) show, as a rule, large quantities of lateral-spined eggs, though less often only terminal- spined eggs may be found. Vesico-prostatic Plexus. — Worms in copula are found in the veins 18 274 THE ANIMAL PARASITES OF MAN of the submucosa in the bladder, and the eggs in the mucosa, and those voided are usually terminal-spined, though lateral-spined eggs are not so rare as generally thought. The problem next arises as to how the eggs get to the lumen of the gut or bladder. The female worm is 280 /x in diameter. Veins in the submucosa of the rectum less than 178 /x in diameter are not affected with endo- phlebitis. It is probable that the female even by stretching could not penetrate much beyond this. Eggs are probably then laid in the submucosa as near the muscularis mucosa as possible. Now if the eggs are laid in a vein of larger calibre than the worm fills, the eggs would be carried back to the inferior mesenteric vein, so that pre- sumably the worm must succeed in blocking the vein already narrowed by endophlebitis, so that by the stasis which ensues the eggs may escape from the veins. How this occurs is not exactly known ; it is not necessarily due to the spine, as the same escape into the tissues occurs in spineless eggs, such as those of Schistosoma japonicum. The eggs, then, pass as foreign bodies through the tissues. Another hypo- thesis is that the worms leave the veins in order to lay their eggs, but the evidence is against this. Caval System. — Occasionally worms that have passed through the vesical plexus may be found in the iliac vein, inferior vena cava, and even the lungs. If the worms are young they contain a lateral- spined €^gg ; if adult, numerous (50 to 100) terminal-spined eggs. Lungs. — When the liver is strongly infected with (terminal-spined) eggs it is possible that by passive movements some may pass into the intralobular veins, and thence by the inferior vena cava to the lungs. Gall-bladder. — Similarly terminal-spined eggs pass into the bile- capillaries and gall-bladder (where they may be abundant), and so into the faeces. Detection of Eggs. — Occasionally eggs may be found in various other parts of the body. They are best detected by macerating pieces of the tissue in question in about ^ per cent, hydrochloric acid at 50 to 60° C. (Looss). Pathological changes : — Rectum.— These have been studied thoroughly by LetuUe in the case of an apparently pure infection of the rectum.^ They take the form of a chronic diffuse inflammation, which may result in— (i) ulcera- tion, or (2) hyperplasia of the mucosa, producing adenomata. Ulcerative Form. — The mucosa is transformed into a mass of vascular connective tissue. The connective tissue spaces next be- come invaded by numerous mononuclear cells. The tissue itself ' It is noteworthy that in this almost classical case no worms were found in any of the sections. It is further noteworthy that the eggs in the rectum showe-i great irregularity of form. Eggs with a spine at each end were not uncommon ; exceptionally eggs with two polar spines and one lateral. SCHISTOSOMA H^MATOBIUM 275 undergoes diffuse sclerosis, becoming hard and fibroid. Eventually ulcerative necrosis sets in. During these changes the Lieberkiihn glands are destroyed. The process does not extend to the submucosa, in this respect differing from that in chronic dysentery. Hyperplastic Form. — The Lieberkiihn glands of the mucosa at first hypertrophy ; then there is an actual hyperplasia resulting in adeno- mata. The interstitial tissue of the glands is also greatly hyper- trophied, giving rise to very vascular granulations. These growths are often hollow and contain worms. Many eggs are found in the mucosa on their way to the lumen of the gut. The muscularis mucosa is thickened up to twice or even ten times the normal. Its vessels are dilated (36 fi to 80 /i,), but they do not allow of the passage of worms. The submucosa is profoundly changed ; rigid and hard instead of supple. It is here that the greatest number of eggs occur. A re- markable condition of endophlebitis exists in the veins of the submucosa, not only in the smaller ones but also in the larger ones (370//, by 270 /x). This endophlebitis results in a more or less complete occlusion of the vessels of the lumen. The muscular coats are free from change, also their veins. The Serous Coats. — The veins about 1,900//, also show endophlebitis. Besides the rectum, in extreme cases even the transverse colon, the caecum and small intestine may be affected. Bladder. — In the early stages the mucosa is deep red and swollen like velvet, or there may be localized patches of hyperaemia or extravasation. The subsequent changes take two chief forms : — (i) Sandy Patches. — The mucosa ooks as if it were impregnated with a fine brownish or yellowish powder (myriads of ova). This is accompanied by a gradual hypertrophy and new formation of con- nective tissue, so that dry, hard or plate-like patches with this sandy appearance arise ; the thickening eventually affects all the coats of the bladder. In the older patches many of the eggs are calcified. These patches sooner or later break down, ulcerate and necrose. Phosphatic deposits are abundant and stone is common. These patches are not found in the rectum. (2) Papillomata. — Where the inflammatory change produced by the eggs gives rise to hypertrophy and hyperplasia of the mucosa, papillomata result, the axis of which is formed by connective tissue of the submucosa. These are most variable in shape and form and bleed readily, and sometimes contain cavities of extravasated blood. As in the rectum, it is in the submucosa that eggs are most abundant, and worms in copula occur in the veins of this layer, but endophlebitis is not as general as described in the rectum. Malignant disease of the bladder is not an uncommon sequela of bilharziasis. 276 THE ANIMAL PARASITES OF MAN Besides the bladder, the ureters and kidneys may in advanced cases be involved. The prostate and vesiciilae seminales are commonly diseased. Eggs have been recorded in the semen. The urethra is frequently attacked; the vagina in the female. Eggs also occur in the lymphatic glands of the gut. Geographical Distribution. — East Africa : Nile Valley, Red Sea Coast, Zanzibar, Portu- guese East Africa, Delagoa Bay, Natal, Port Elizabeth. South Africa : Cape Colony, Orange Free State, Transvaal, Mauritius, Bourbon, Mada- gascar. West Africa : Angola, Cameroons, Gold Coast, Gambia, Senegal, Sierra Leone, Lagos, Nigeria. North Africa : Tripoli, Tunis, Algeria, parts of the Sahara. Central Africa : Sudan, various portions. Uganda, Nyasaland. It occurs with varying frequency in these regions. It is probably more widely spread than this list implies, as undoubtedly many cases are seen which are not recorded. Isolated cases have been recorded from Arabia, India,^ Greece, Cyprus. The means by which infection is brought about are still uncertain ; we only know that the miracidia (fig. 175) enclosed in the dis- charged eggs do not hatch if the eggs remain in the urine, but after cooling perish. As soon, however, as the urine is diluted with water the shell swells, generally bursting length- ways, and releases the miracidium from its investing membrane, so that it can swim about with the aid of its cilia. In its structure it differs but little from the miracidium of Fasciola hepatica, as, for instance, in the lack of eyes ; the two large gland cells situated on either side of the intestinal sac are also present in the miracidia of Fasciola hepatica. Sarcode Globules. — This is a term applied to certain globules which at times appear in the miracidium and are later ejected. Some authors V.SC. Fig. 1 74. — Schistosoma hamatobium^ Bilh. : geni- talia of the female. V.s.^ ventral sucker; /., gut caeca ; V.d.^ vitelline duct; V.sc, vitellarium ; (?., ovary ; Oe., oesophagus ; Sh., shell gland ;£/., uterus. Magni- fied. (After Leuckart.) * In a case from Madras, recoided by Stephens and Christophers, the eggs were long and spindle-shaped, quite unlike the eggs of Schistosoma hcematobium. SCHISTOSOMA MANSONI 277 consider them as indicative that the miracidium has developed into a sporocyst, but Looss considers them to be degeneration products. The Bilharzia mission, under K. T. Leiper, sent to Egypt by the War Office early in 19 15, reports that cercariae of bilharzia type were recognized in four of the commonest fresh-water molluscs around Cairo. With material obtained from naturally infected Planorbis boissyi acute bilharziosis was experimentally produced in rats, mice, and monkeys. Infection takes place experimentally through the skin and also through the mucous membrane of the mouth and oesophagus. The miracidium, after entering the mollusc, develops into a sporocyst. This gives rise not to rediae, but to secondary sporocysts, which, in Schistosonna haematobium, Bilharz, 1852. Male, four or five large testes. Gut forks unite late, so that the single gut stem is short. Female, ovary in posterior half of body. Uterus very long, voluminous, with many terminal-spined eggs, some lying in pairs. Vitellaria in posterior fourth of body. Cercariae in Bullinns contorUis and Bullimis dyboivski (syn. : Physa alexandrina) in Egypt. Schistosoma mansoni, Sambon, 1907. Male, eight small testes. Gut forks unite early, so that the single gut stem is very long. Females, ovary in anterior half of body. Uterus very short ; usually only one lateral-spined Qgg at a time /// titero. Vitellaria occupy posterior two-thirds of body. Cercariae in Planorbis. boissyi in Egypt. The above morphological descriptions are founded on worms of each species, derived from experimentally infected mice (Leiper, R. T., Brit. Med. Journ.y March 18, 191 6, p. 411). To Binder : face p. 276 Syn. : TS\ cattoi, BlancEard, 1905. Male. — Eight to 19 mm., but extreme limits are 5 to 22*5 mm. Con- sists of a short fore-body, separated by the ventral sucker from the hind-body. The ventral sucker is stalked and somewhat larger than 276 THE ANIMAL PARASITES OF MAN Besides the bladder, the ureters and kidneys may in advanced cases be involved. The prostate and vesiciilae seminales are commonly diseased. Eggs have been recorded in the semen. The urethra is frequently attacked; the vagina in the female. Eggs also occur in the lymphatic glands of the gut. Geographical Distribution. — East Africa : ^- f®^ Nil^ Valley, Red Sea Coast, Zanzibar, Portu- guese East Africa, Delagoa Bay, Natal, Port Elizabeth. South Africa : Cape Colony, Orange Free State, Transvaal, Mauritius, Bourbon, Mada- p-pj^c^r Sarcode Globules. — 1 nis is a term appiiea to certain giODUies wnicn at times appear in the miracidium and are later ejected. Some authors * In a case from Madras, recoided by Stephens and Christophers, the eggs were long and spindle-shaped, quite unlike the eggs of Schistosoma hcematobium. SCHISTOSOMA MANSON] 277 consider them as indicative that the miracidium has developed into a sporocyst, but Looss considers them to be degeneration products. The Bilharzia mission, under R. T. Leiper, sent to Egypt by the War Office early in 19 15, reports that cercariae of bilharzia type were recognized in four of the commonest fresh-water molluscs around Cairo. With material obtained from naturally infected Planorbis boissyi acute bilharziosis was experimentally produced in rats, mice, and monkeys. Infection takes place experimentally through the skin and also through the mucous membrane of the mouth and oesophagus. The miracidium, after entering the mollusc, develops into a sporocyst. This gives rise not to redise, but to secondary sporocysts, which, in turn, produce cercariae. These, like the adult worm, differ from other distomes in lacking a muscular pharynx. Schistosoma mansoni, Sambon, 1907. According to Manson, Sambon and others, the eggs with lateral spines belong to a species different from Schistosoma hcBmatobium. Infections with this species only are said to occur in the Congo, Southern States of North America, West Indies (Guadeloupe) and Brazil (Bahia). The following characters, according to Flu, differentiate this species : (i) In the male the transition from the anterior portion of the worm to the lateral fields (the infolded portions which form the gynaecophoric canal) is not a gradual one as in Schistosoma hcematobium, but in this case the lateral fields rise suddenly, almost at right angles to the anterior portion. (2) The ovaries have a well-marked convoluted course as in no other schistosome. (3) The ootype is symmetrical in reference to the long axis of the body, its duct being lateral on the ventral side (Looss' explanation of this we have already given). (4) The worms live exclusively in portal vein and tract. (As lateral-spined eggs occur also in the bladder, this is not exactly true.) Fig. 175. — Ovum of Schistosoma hamatobiuniy Bilh., with miracidium, which has turned its an- terior end towards the posterior end of the egg. 275/1. (After Looss. Schistosonna japonicum, Katsurada, 1904. Syn. : S. cattoi, Blanchard, 1905. Male. — Eight to 19 mm., but extreme limits are 5 to 22*5 mm. Con- sists of a short fore-body, separated by the ventral sucker from the hind-body. The ventral sucker is stalked and somewhat larger than 278 THE ANIMAL PARASITES OF MAN the oral sucker. Both suckers are larger than the corresponding ones in S. hcematohium. Body usually smooth, but in the fresh state numerous fairly evident spines along the margin of the canal. (Eso- phagus : two bulbs. The junction of the gut forks more posterior than in S\ hcematohhmij the median united gut stem occupying a quarter to one-fifth to one-sixth of the body length. An excre- tory canal runs along each side of the body, opening into the dorsal excretory pore. Testes irregularly elliptical, six to eight in number, in the anterior part of hind-body. The vasa efferentia unite into a common vas deferens which opens directly behind the ventral sucker. The seminal vesicle lies just behind this. Female. — Up to 26 mm., generally thinner than the male. Surface smooth. Suckers armed with fine spines. Ventral sucker larger than oral. Body thicker behind the region of the ovary. The gut forks unite immediately behind the ovary. The united gut much thicker than in S. liceinatobiujii. Ovary elliptical, almost in the mid-body, its hinder por- tion dilated. The oviduct arises from its posterior end and then runs sinuously forward, where it is joined by the vitel- larian duct ; the vitellarium well deve- loped, extending from behind the ovary almost but not quite to the posterior end as in S. hctmatobitun. Shell gland ducts enter at the junction point of oviduct and vitelline duct. The canal here forms an ootype and then proceeds as the uterus to open directly behind the ventral sucker. The uterus occupies almost half the hind-body. In S\ hcema- tobiiim this is not so. The uterine canal is cleft-like, i.e., its dorso-ventral diameter is much greater than its lateral diameter. The number of eggs varies from about 50 to 300 from observations made in various hosts. Eggs. — In iitero assume various shapes, as they are soft ; the lumen of the uterus is narrow. Outside they are oval, faint yellow, double Fig. 176. — Schistosoma japoni- cum : anterior end with testes ; posterior end with point of union of caeca. Length of worm about 10 mm. (After Katsurada.) SCHISTOSOMA JAPONICUM 279 contoured. In faeces the eggs measure 83*5 /^ by 62*5/1 (man) ; 85 /x by 61-5 ^ (cattle) ; 98*2 fju by 73*8 fi (dog). The eggs have either small lateral spines or thickenings, and Looss at the opposite side has described cap-like thickenings. The eggs in the tissues undergo various de- formities, and may contain a miracidium, as also the eggs in faeces do ; or the contents may consist of granular matter or amorphous masses or they may be calcified. Lympho- cytes and giant cells may also invade the eggs. Mode of Infect ion. ^The miracidia hatch in water in as little as fifteen minutes, but the majority in one to three hours. They will live in water for about twenty-four hours. In water they undergo a transformation into *' larvae/' which then penetrate the skin, as has been shown by Fig. 177. — Schistosoma japo- nicum, male and female in co- pula. X 60. (After Katsurada.) Fig. 178. — Schistosoma japonicum : eggs from human liver, showing "spines" and "hoods" at opposite pole. (After Looss.) Japanese writers to hold good for man, cattle, dog and cat. The penetration of the skin is attended with an eruption on the legs, "■ Kabure." The exact route by which the worms reach the portal vein is uncertain. Infection in Japan takes place from spring to autumn, especially May to July, when the soil is contaminated with manure of cattle infected with S. japonicum. They also appear to develop in molluscs. Leiper and Atkinson found cercariae (in sporocysts) 28o THE ANIMAL PARASITES OF MAN in the liver of a mollusc, Katayama nosophora. They infected mice by immersing them in water containing liver emulsion and so free cercariae, thus confirming the similar results of Miyairi and Suzuki. Habitat. — The worm occurs in Japan, China, and the Philippines. The normal host is man and mammals. Cattle, dog and cat are often found naturally infected. Mice can also be experimentally infected. Their seat of election is the portal vein and its branches, especially the mesenteric veins. They either swim free in the blood or remain fixed by their suckers to the intima of the vessels. They have also been found in the vena cava and right heart of a cat, but not so far in the vesical plexus. Eggs are found in the submucosa and mucosa of the gut, especially the colon, and at times in the serosa and subserosa of the small intestine, where they give rise to new growths. Occasionally eggs are found in the brain. The life of tlie worms is at least two years. Pathogenic Effects. — An?emi3. through loss of blood due to worms ; enlarged spleen, toxic in origin (?) ; phlebitis, thrombosis, due to portal stasis ; the eggs, however, cause the They are carried by the circulation to various Fig. 179. — Schistosoma Ja- ponicum : from dog. Uterine ^gg* "^ c. 800. (After Kat- surada.) greatest mischief. 7/ Fig. v%0.~ Schistosoma japonictim : from dog. X c. 800. (After Katsurada.) Fig. 181. — Schistosoma japonicum : from dog. Egg from faeces, x c. Scx). (After Katsurada.) SCHISTOSOMA JAPONICUM 281 organs where they produce inflammation, granulation tissue, and later connective tissue. Liver. — The eggs reaching this organ give rise to granulomata and hence enlarged liver, and later, when connective tissue is formed, to contraction. The surface is rough and irregularly granular, ''parasitic embolic cirrhosis " of Yamagiwa. ^^ rf% ms€> Q (■ ■ o-}V>xr^^ ■n ^ ^>-\>-; rPS^c;,.., - : ^c^ O' n o L'.^^ '■■ .J Fig. \%2.— Schistosoma japonicum : section through the gut of a Chinaman showing eggs. X 58. (After Catto.) Gut. — The eggs in the mucosa and submucosa cause catarrh and destruction of tissue or new growth. In the small intestine the eggs are mainly in the serosa and subserosa, where they give rise to poly- poid or branched growths. 282 THE ANIMAL PARASITES OF MAN S/)/^^n.— Enlarged, at first due to toxin (?) and later due to portal stasis. Eggs in the spleen are uncommon. Ascites also arises from the portal stasis, and is generally present in advanced cases. Eggs may be found in many other situations : glands (numerous), mesentery, stomach, pancreas, kidney, etc. The bladder remains free. Fig. 183. — Schistosoma japonicuni : liver showing eggs in the intra- and interlobular connective tissue, x c. 80. (After Katsurada.) Class III. CESTODA, Rud., 1808. Tapeworms have been known from ancient times — at all events, the large species inhabiting the intestines of man — and there has never been a doubt as to their animal nature. The large cysticerci of the domestic animals (occasionally of man also) have been known for an equally long period, but they were generally regarded as growths, or "hydatids," until almost simultaneously Redi in Italy, and Hartmann and Wepfer in Germany, concluded from their movements and organi- zation that they were of animal nature. From that time the cysticerci have been included amongst the other intestinal worms, and Zeder (1800) established a special class {Cystici, Rud., 1808) for the bladder worms. Things remained in this condition until the middle of the last century, when Kiichenmeister, by means of successful feeding experiments, demonstrated that the cysticerci were definite stages of development of certain tapeworms. Before Kiichenmeister, E. Blanchard, van Beneden, and v. Siebold had held the same opinion in regard to other asexual Cestodes. CESTODA 283 Since the most remote period another question has again and again occupied the attention of naturahsts, the question of the morphological nature — that of the INDIVIDUALITY OF THE TAPEWORM. The ancients, who were well acquainted with the proglottids {Vermes aecurbitani) that are frequently evacuated, were of the opinion that the tapeworm originated through the union of these separate proglottids, and this view was maintained until the end of the seventeenth century. In 1683 Tyson discovered the head with the double circlet of hooks in a large tapeworm of the dog ; Redi (1684) was also acquainted with the head and the suckers of several Tsenice. Andry (1700) found the head of Tcenia saginata^ and Bonnet (1777) and Gleichen-Rusworm (1779) found the head of Diboihriocephalus latus. Consequently most authors, on the ground of this discovery, considered the tapeworm as a single animal, that maintains its hold in the intestine by means of the head, and likewise feeds itself through it. The fact was recognized that there were longitudinal canals running through the entire length of the worm, and it was thought that these originated in the suckers, and that the entire apparatus was an intestine. As, moreover, the segments form at the neck, and are cast off from the opposite extremity, the tapeworm was also compared with the polyps, which were formerly regarded as independent beings. Steenstrup, in his celebrated work on the alternation of generations (1841), was the first to give another explanation. This has been elaborated still further by van Beneden, v. Siebold and Leuckart, and until a few years ago all authorities adopted his views. According to this view, the tapeworm is composed of numerous individuals, something like a polyp colony, and, in addition to the proglottids — the sexual individuals which are usually present in large numbers — there is ONE individual of different structure, the scolex^ which not only fastens the entire colony to the intestine, but actually produces this colony from itself, and therefore is present earlier than the proglottids. The scolex is a "nurse," which, though itself produced by sexual means, increases asexually like a Scyphistoma polyp ; the tapeworm chain has therefore been termed a strobila. Consequently the develop- ment of the tapeworms was explained by an alternation of generations. In support of this opinion it was demonstrated not only that the adult sexual creatures, the proglottids, can separate from the colony and live independently for a time, but that in certain Taeniae, and especially in many Cestodes of the shark, the proglottids detach themselves long before they have attained their ultimate size, and thus separated continue to develop, grow and finally multiply ; the scolex also exhibits a certain independence in so far as, though not, as a rule, capable of a free life, yet it in some cases lives as a free being, partly on the surface of the body of marine fishes and partly in the sea. With the more intimate knowledge of the develop- ment of the cysticerci, the independent nature of the scolex was recognized. It is formed by a budding of the bladder that has developed from the oncosphere, in some cases (Coenurus) in large numbers, in other cases (Echinococcus) only after the parent cyst has developed several daughter cysts. Released from its mother cyst and placed in suitable conditions, it goes on living, and gives rise at its posterior end by budding to the strobila, the proglottids of which eventually become sexual individuals. In order to make this clearer we will briefly summarize what takes place in the jelly-fishes. V>ymetamorphosis is meant a developmental change in the sa7ne indivi'dual, while alternation of generations, or metagenesis, implies a stage in which reproduction of individuals takes place by a process of budding or fission. This asexual reproduc- tive stage alternates with the sexual mode of reproduction. Thus in the development of the Scyphozoa (jelly-fishes) we have : — (r) The fertilized ^g% cell divides regularly and forms a morula. 284 THE ANIMAL PARASITES OF MAN (2) By accumulation of fluid in the interior this becomes a closed sac with a wall formed of a single layer of cells, forming the blastosphere or blastula. (3) One end of the sac is invaginated, forming a gastrula. (4) The gastrula pore or mouth closes, forming again a sac, the walls of which have two layers, forming 2iplanula. (5) This becomes fixed to a rock, an invagination forms at one end, a depression — the stomodaeum — communicating with the enteric cavity. Tentacles grow out and we have a Scyphozobn polype^ Scyphistoma or Scyphula. It is to this stage that Steenstrup gave the name " nurse " (" wet-nurse ")» because it nourished or produced asexually the succeeding forms. (6) Asexual reproduction by transverse fission occurs in this, forming a pile of saucer- or pine-cone-like animals which before this time had been considered to be a distinct animal, which was called sirobila from its resemblance to a pine- cone. This is the alternate generation. (7) The individuals of the strobila become free and are called Epkyrulcs. (8) These develop finally into adult sexual jelly-fish, Scyphozoa, so that com- paring a tapeworm with this we have {a) t^g, {b) scolex (= Scyphula or " nurse "), {c) asexual reproduction of the tapeworm chain (= strobila), id) development of the individuals of the chain (proglottids) into sexual adults. Van Beneden's terminology for these stages is the following : Ciliated embryo = protoscolex ; scyphistoma — deutoscolex (or scolex) ; free Ephyrula = proglottis. According to this view, as is the case in many endoparasitic Trematodes, asexual reproduction by budding occurs at two stages of the whole cycle of development, viz. (i) in the formation of the scolex by budding from the bladder ("nurse"), (2) in the formation of the strobila by budding from the scolex ("nurse"). But in cysticercal larval forms it appears that the scolex does not arise in this way but is simply a part of the proscolex (hexacanth embryo), becoming invaginated into it for protection, so that there is no asexual gemmation here. It has been questioned also whether the strobila also arises by gemmation. If it does, the tapeworm is a colony of zooids produced by budding from the asexual scolex ; if it is not produced in this way, then the tapeworm is to be regarded as an individual in which growth is accompanied by segmentation. Against the " colony " view are the facts that the muscular, nervous, and excretory systems are continuous through- out the worm, and that some tapeworms, such as Ligula^ are unsegmented. Finally, if the tapeworm is an individual the question arises which is the head end. As new segments are formed at the neck, and as this point in annelids is the antepenultimate segment, the scolex must be the last or posterior segment. The caudal vesicle or bladder of larval forms is consequently anterior. According to this view, in tapeworms as among many endoparasitic flukes, an asexual multiplica- tion occurs at two points of the whole cycle of development, which is as follows : (i) ^%%^ (2) oncosphere or hexacanth embryo, (3) bladder (cysticercus or hydatid), (4) (after digestion of the bladder) by budding, the scolex, (5) by budding from the scolex the sexual proglottids, (6) the %%% ; (4) and (5) being the two asexual stages. Anatomy of the Cestoda. If we except the tapeworms with only one proglottis, the Cestoidea Monozoa, Lang = Cestodariay MonticeUi, we can always distinguish in the Cestodes, in the narrower sense, one scolex or head and a large or small number of segments (proglottids). The scolex serves the entire tapeworm for fastening it to the internal surface of the ANATOMY OF THE CESTODA 285 intestinal wall, and therefore carries at its end. various organs which assist in this function, and which are as follows : (i) Suctorial ORGANS, i.e., the four suckers (acetabula), which are placed crosswise at the circumference of the thickened end of the scolex ; further, the double or quadruple groove-like suckers (bothridia), which are diversely shaped in the various genera and families/ (2) Fixation ORGANS (hooklets)^ that likewise occur in varying numbers and different positions; they may be in the suckers, or outside them on the apex of the scolex ; for instance, in many of the Tceniidce they appear in a circle around a single protractile organ, the rostellum, or the latter may be rudimentary, and is then replaced by a terminal sucker. (3) Proboscis. One family of the Cestodes, the RhynchobothriidcB, carries four proboscides, moved by their own muscular apparatus, on the scolex, and they are beset with the most diverse hooks. (4) Tentacle-like formations are only known in one genus (Polypocephalus). The thickened part of the scolex that carries the suckers is usually called the head ; the following flat (unsegmented) part connecting it with the proglottids is called the neck, and is sometimes quite small. In a few cases the entire scolex (or head) disappears, and its function is then undertaken by the contiguous portion of the chain of proglottids, which is transformed into a variously shaped PSEUDO-SCOLEX. The proglottids are joined to the scolex in a longitudinal row, and are arranged according to age in such a manner that the oldest proglottis is farthest from the scolex, and the youngest nearest to it. The number of segments varies, according to the species, from only a few to several thousands ; they are either quadrangular or rectangular ; in the latter case their longitudinal axis falls either longitudinal or transverse to that of the entire chain, according as the segments are longer than broad or broader than long. When the number of segments is very large, the youngest ones are, as a rule, transversely oblong, the middle ones are squarish, and the mature ones longitudinally oblong. The posterior border of the segments, as a rule, carries a longitudinal groove for the reception of the shorter anterior border of the following proglottis. The two lateral borders of the segment are rectilinear, but converge more or less towards the front, or they are bent outwards. In most of the Cestodes the segments, just as the neck, are very fiat ; in rare cases their 1 They may remain simple, and are then not separated from the remaining muscles of the scolex ; or they project as roundish or elongated structures over the scolex, hollow on their free surface, and often divided into numerous areas by muscular transverse ribs. They may also carry accessory suckers on their surface. 2 The various parts of a booklet are thus named from the point backwards : (i) blade or prong, (2) guard or ventral or posterior root, (3) handle or dorsal or anterior root. 286 THE ANIMAL PARASITES OF MAN transverse diameter is equal to their dorso-ventral diameter. As a rule the segments, singly or several united together, detach them- selves from the posterior end, in many cases only after complete maturity is attained, and in others much earlier ; they then continue to live near their parent colony, to still call it by that name, in the same intestine and continue their development. Even when evacuated from the intestine the proglottids under favourable circumstances can continue to live and creep about, until sooner or later they perish. The first proglottis formed, and which in a complete tapeworm [i.e., sexually complete] is the most posterior, is as a rule smaller and of different shape, it also frequently remains sterile, as likewise happens in the next (younger) segments in a few species ; otherwise, however, sooner or later the generative organs develop in all the segments, mostly singly, sometimes in pairs ; in the latter case they may be quite distinct from each other or possess some parts in common. The term *' mature " is used for a proglottid that has the sexual organs fully developed, while '^gravid" is used for one containing eggs. Most of the species combine male and female genitalia in the same segment, only a few are sexually distinct (Dioecocestus). In the hermaphrodite species one male and one female sexual orifice are always present, and, in addition, there may be a second female orifice, the uterine opening ; as a rule, however, this is lacking, and in one sub-family, the A cole ince, to which also the genus Dioecocestus belongs, the other sexual orifice, the opening of the vagina, is also absent. The position of these orifices varies ; the cirrus and vagina usually open into a common atrium on one lateral border or on a surface of the segments ; the orifice of the uterus may be on the same surface or on the opposite one. The surface on which the uterus opens is termed the ventral SURFACE; if this orifice is absent, one must depend on the ovary, which almost always approaches one of the two surfaces ; this surface is then called the ventral. The length of the Cestodes — independently of their age — depends on the number and size of the segments, as well as on their contraction ; the smallest species {Davainea proglottina) is 0*5 to vo mm. in length; the largest may attain a length of 10 m., and even more. The entire superficial surface of the tapeworms is covered with a fairly resistant and elastic layer, which exhibits several indistinctly limited layers and which is usually called a cuticle, which also covers the suckers, and is reflected inwardly at the sexual orifices. In some species fine hairs appear, either on the entire body or only in the region of the neck, on the external surface. In the cuticle there can be ANATOMY OF THE CESTODA 287 A.m. recognized, besides the pores, which no doubt are concerned with nutrition, spaces in which He the ends of sensory cells. Close under the cuticle lies the external layer of the parenchyma (basal membrane), and below this the circular and longitudinal muscles forming the dermo-muscular coat. The matrix cells of the cuticle occur as in the Trematodes, only on the inner side of the peripheral muscles in the external zone of the parenchyma ; they are fusiform cells, forming one or two layers, but are not arranged in the manner of epithelial cells (fig. 184, Sec). They have fine branching processes which run between the dermal muscles, pass through the basal membrane and penetrate the internal sur- face of the cuticle with small pistil-like enlargements, ex- panding on the internal surface of the cuticle into a thin plasma layer. In addition to the above mentioned, there are other cuticular formations occur- ring on the cuticle of some Cestodes, such as immobile hairs and variously formed hooks, such as are seen prin- cipally on the scolex. Their development is only roughly known in a few species ; they are usually already present in the larval stage, and of the same arrangement and shape as in the fully developed tapeworms ; a matter of importance, because by these Ex.c. F.v.s. Fig. 184. — Schematic representation of a small part of a transverse section o{ Ligula sp. Bs., basal membrane ; Cn., cuticle ; at its base are the end- plates of the subcuticular (epithelial) cells; in the centre a cuticular sense organ, O.s. ; F.v.s., vitelline follicle ; Exc, excretory vessel ; C, calcareous cor- puscle ; L.m., longitudinal muscles ; M.c. myoblast ; P.m., parenchymatous or dorso-ventral muscles ; /*/., plexus of nerve fibres ; A.m., circular muscles; Sc.c.y subcuticular or matrix cell; T.c, terminal flame cell. 500/1. (After Blochmann.) structures larvae can be recog- nized as being those of a certain species of tapeworm. The CUTICULAR GLANDS in Cestodes are scarce. The PARENCHYMA forms the chief tissue of the entire body, and in all essentials its structure is similar to that of the Trematodes. 288 THE ANIMAL PARASITES OF MAN The same doubt exists here also as to the nature of the parenchyma. Recent authors consider that it consists of highly branched cells, the processes of which ramify in all directions. These cells lie in a non- cellular matrix containing fluid vacuoles. This matrix spreads in between and so breaks the continuity of the epidermal cells. In the parenchyma of almost all the Cestodes there are found in adult specimens, as well as in larvae, light-refracting concentrically striated structures, of a spherical or broad elliptical shape, which, on account of their containing carbonate of lime, are termed calcareous CORPUSCLES (fig. 184, C). Their size, between 3 ^ and 30 ^, varies according to the species ; their frequency and distribution in the paren- chyma also varies, but they are chiefly found in the cortical layer. L.m. Sec. M.f. L.m. Sm.f. Fig. 185. — Half of a transverse section through a proglottis of Tcenia crassicollis. Cu., cuticle; Ex.v., external excretory vessel, to the right of which there is the smaller internal one ; T., testicular vesicles ; L.m. longitudinal muscles (outer and inner) ; M./., lateral nerve with the two accessory nerves; Sec, subcuticular matrix cells; Sm.f., submedian nerve; Tr.m., transverse muscles; 67., the uterus, and the middle of the entire transverse section. 44/1. They are the product of certain parenchymatous cells, in the interior of which they lie like a fat globule in a fat cell, but according to others they are intercellular in origin. The MUSCULAR SYSTEM of the proglottids is composed of— (i) the subcuticular muscles (figs. 184 and 185), as a rule consisting of a single layer of annular muscles ; (2) longitudinal muscles; (3) dorso- ventral fibres extending singly from one surface to the other, and at both ends expanding in a brush-like manner, and inserted into the basal membrane, consisting of an outer, more numerous, and an inner, less numerous but more powerful layer (the number of bundles in this layer being in certain cases of specific importance) ; (4) trans- verse fibres, the elements of which penetrate to the borders of the seg- ments, thus passing through the longitudinal muscles and reaching ANATOMY OF THE CESTODA 289 .--C the cuticle. In the region of the septa the transverse and dorso- ventral muscles form a kind of plate. The mass of parenchyma bounded by the transverse muscles is termed the medullary layer, while the mass lying outside them is termed the CORTICAL layer. It was known long ago that the myoblasts adhere to the dorso- ventral fibres as thickenings, but it is only recently that large star- shaped cells (fig. 184), separated from but connected with them by processes, have been recognized as the myoblasts of other fibres (Bloch- mann, Zernecke). Within the scolex the direction and course of the muscular layers change. The suckers are parts of the musculature, locally transformed, with a powerful development of the dorso -ventral muscles, now become radial fibres. The rostellum of the armed Taeniae, like the proboscis of the Rhynchobothfiidce, also belongs to the same category of organs. In the simplest form, the ros- tellum, or top of the head (as in Dipylidiuin caninum), appears as a hollow oval sac, the anterior part of which, projecting beyond the upper surface of the head, carries several rows of hooks (fig. 186). The entire internal space of the sac is occupied by an elastic, slightly fibrous mass, while the anterior half of the surface of the rostellum is covered by longitudinal fibres and the posterior half by circular fibres. On contraction of the latter the entire mass is protruded through the apical aperture, the surface of the rostellum becomes more arched, and the position of the hooks is, in consequence, altered. The rostellum of the large-hooked Ta^niidcv, which inhabit the intestine of man and beasts of prey, is of a far more complicated structure, for, in addition to the somewhat lens-shaped rostellum carrying the hooks on its outer surface, there are secondary muscles grouped in a cup-like manner (fig. 187). Every change in the curvature of the surface of the rostellum induces an alteration in the position of the hooks. In the hookless Tctniidct the muscular system of the rostellum is altered in a very different manner ; in a few forms a typical sucker appears in its place. The NERVOUS system commences in the scolex and runs through 19 Fig. 186. — Dipylidium caninum : from the cat. In the upper figure the rostellum is retracted, in the lower protruded, a, sucker ; <5, hooks of rostellum ; B, enlarged hook ; c, apical aperture on scolex ; d, longitu- dinal muscles; (?, circular muscles. (After Benham. ) 290 THE ANIMAL PARASITES OF MAN the neck and the entire series of proglottids. Within the proglottids it consists of a number of longitudinal nerve fibres of which those at each lateral border are usually the largest. In the Taeniae the lateral nerves are accompanied both dorsally and ventrally by a thinner nerve (accessory nerve) (fig. 185) ; on each surface, moreover, between the lateral nerve and the median plane, there are two somewhat stronger bundles (sub-median), so that there is a total of ten longitudinal nerve bundles. They lie externally to the trans- verse muscle plates, and the lateral and accessory bundles lie externally to the principal excretory vessels, and are every- where connected by numerous anastomoses and secondary anastomoses ; one typical ring commissure is usually found at the posterior border of the segments. ^In the Boihrio- cephalidcv the distribution of the nerve bundles is different (for instance, two lie in the medullary layer), or they are split up into a larger number of branches. In the scolex the nerve bundles are connected in a very remarkable manner by commissures with that which is generally termed the central part of the entire nervous system. There occurs normally a commissure between the two lateral nerves ; at the same level, the dorsal and ventral median nerves are also connected at each surface as well with each other^^as with the lateral nerves, so that a hexagonal or octagonal figure is formed. The so-called apical nerves pass from this commissural system anteriorly, embrace the secondary muscular system of the rostellum semicircularly, and form an annular commissure (rostellar ring) at the inner part of the rostellum. The peripheral nerves arise from the nerve bundles as well as from the commissures situated in the scolex ; some go direct to the muscles, while others form a close plexus of nerves external to the inner longitudinal muscles, which plexus likewise sends out fibres to the muscles, but principally to [{numerous fusiform sense organs Fig. 187. — Longitudinal section of the head and neck of Tienia crassicollis, showing the lens-shaped muscular rostellum, with two hooks lying in the concentric cup-like mass of muscles. L.f/i., longitudinal muscles of the neck ; Lf., left lateral nerve ; C, ganglion ; S.c, subcuticular layer ; W^, external, W^, internal excretory vessel. 30/1. SZZI^^I ANATOMY OF THE CESTODA 291 (fig. 184, PL) ; they lie internal to the subcuticular cells and, piercing the cuticle with their peripheral processes, end as projecting " receptor " hairs. Higher organs of sense are not known. The EXCRETORY APPARATUS of the Cestodes is similar to that of other flat worms. The terminal (flame) cells, which hardly differ in appearance from those of the Trematodes, are distributed throughout the parenchyma, but are more common in the cor- tical than in the medullary layer (fig. 184, T.c). Before opening into a collecting tube, the capil- laries run straight, tortuously, or in convolutions, anastomos- ing frequently with one another or forming a retc niirahile. The collecting tubes, which have their own epithelial and cuti- cular wall, and which also appear to be provided with muscular fibres, occur typically as four canals passing through the entire length of the worm (fig. 1 89) ; they lie side by side, two (a wider thin-walled ventral, and a narrower thick-walled dorsal one) in either lateral field ; in the head the two vessels on each side unite by means of a loop, at the posterior extremity they open into a short pyriform or fusiform terminal bladder w^hich dis- charges in the middle of the posterior edge of the original terminal proglottis. This primitive type (fig. 189) of arrangement of collective tubes is subject to variation in most Cestodes, in the scolex as well as in the segments. Indeed, even the lumen of the four longitudinal tubes does not remain equal, as the dorsal or external tubes are more fully developed and become thicker, whereas the ventral or internal ones remain thin, and in some species quite disappear in the older seg- ments (figs. 185, 187). Moreover, very frequently connections are established between the right and left longitudinal branches, as in the head, where a 'frontal anastomosis" develops, which in the Tceniidce usually takes the form of a ring encircling the rostellum (fig. 190), and in the segments of a transverse anastomosis at each posterior border, Fkj. 188. — Tcenia canurus, head and part of neck showing nervous system. Enlarged. (After Niemiec.) 292 THE ANIMAL PARASITES OF MAN especially between the larger branches, and more rarely between the smaller collecting tubes also (fig. iqO- The so-called '' island " formation is another modification, i.e., at any spot a vessel may divide and after a longer or shorter course the two branches reunite, and this may appear in the collecting tubes themselves as well as in their anastomoses. The above-mentioned ring in the frontal commissure of the Tcemidce is such an island; similar rings also frequently encircle the suckers (fig. 190). In extreme cases [Triceiiophoriis, Ligiila, Dihothriocephaliis, etc.) this island formation extends to all the collecting tubes and their anastomoses. Instead of two or four longitudinal canals only, connected by transverse anasto- moses at the posterior border of the segments, there is an irregular network of vessels, situated in the cortical layer, from which the Fig. 189. — Young Acanthobothrium coronattivi, v. Ben., with the excretory vessels outlined. Slightly enlarged. (After Pintner.) Fig. 190. — Scolex of a cysticercoid from Arion sp., with the excretory vessels outlined. (After Pintner.) longitudinal branches, having again subdivided, can only be dis- tinguished at intervals, and even then not in their usual number. The opening of the longitudinal branches at the posterior end requires more accurate investigation ; it is true that a single terminal bladder is mentioned as being present in many species, but this is also disputed ; when the original end proglottis has been cast off, the longitudinal branches discharge separately. Some species possess the so-called foramina secundaria, which serve as outlets for the collecting tubes ; they are generally at the neck, but may be situated on the segments. The contents of the excretory vessels is a clear fluid, the regurgi- tation of which is prevented by the valves present at the points of origin of the transverse anastomoses. The fluid contains in solution a substance similar to guanine and xanthine. ANATOMY OP THE CESTODA 293 Genital Organs. — With the exception of one genus {DioecocestuSy Fuhrm.), in which the species are sexually differentiated, all the Cestodes are hermaphroditic ; the genitalia develop gradually in the segments (never in the scolex), the male organs, as is usual in hermaphroditic animals, forming earlier than the female. The youngest proglottids generally do not exhibit even traces of genitalia : these, as a rule, develop first in the older segments, and the develop- ment proceeds onwards from segment to segment. In a few ex- ceptional cases (Lignla) the sexual organs are already developed in Vag-. Vsc. Shg. Fig. 191. — Proglottis of Tania sagmata, Goeze, showing genitalia. C, trans- verse excretory canal ; N., lateral longitudinal nerve ; IV., longitudinal excretory canal ; 7"., testicles scattered throughout the proglottis; Ut.^ opposite the central uterine stem (a closed sac) ; Ss.^ genital pore leading into the genital sinus ; above the cirrus and coiled vas deferens {V.d.), below the vagina [Vag.), bearing near its termination a dilatation, the seminal receptacle; Vsc, the triangular vilel- larium, and above it {Shg.) the shell gland ; leading from this to the uterus is seen the short uterine canal, on either side of this the two lobes of the ovary {Ov.). lo/i. the larval stage, but are only functional after the entry of the parasite into the final host. With the exception of the end portions of the vagina, cirrus and uterus, all the parts of the genital apparatus lie in the medullary layer, except only the vitellaria, which in many species are in the cortical layer. The male apparatus consists of the testes, of which, as a rule, there are a large number,^ and which lie dorsal to the median plane (fig. 185, T.) ; a vas efferens arises from each testis, unites with ' There are, however, tapeworms with only one, others with only two or three testes in each segment. 294 THE ANIMAL PARASITES OF MAN contiguous vasa, and finally discharges into the muscular vas deferens that 'is situated in about the middle of the segment. According to the position of the genital pore, the vas deferens opens on the lateral margin or in the middle line in the front of the segment ; it is much Fig. i^2.—Dibothriocephalus laius. Upper figure : female genitalia, ventral view. Lower figure : male genitalia, dorsal view. The central portion only of the proglottis is shown, a, cirrus sac ; b, partly everted cirrus ; c, genital atrium and pore ; d, vaginal pore ; e, uterus ; /, uterine pore ; g, vagina ; h, ovary ; i, shell gland ; j, vitelline duct ; k, lateral nerve ; /, vitellarium ; n, vas deferens (muscular portion) ; /, vas deferens ; q, seminal vesicle ; r and x, vasa efferentia ; s, lateral excretory canal ; /, testicular follicles. (After Benham and Sommer and Landois.) ANATOMY OF THE CESTODA 295 convoluted or twisted, and frequently possesses a dilatation termed the vesicula seminalis. It finally enters the cirrus pouch, which is usually elongated ; within the cirrus pouch lies the protrusible cirrus, which is not uncommonly provided with hooklets. The male sexual orifice almost always opens with that of the vagina into a genital atrium, the raised border of which rises above the edge of the segment and forms the genital papilla (fig. 191). The vagina, like the vas deferens, usually runs inwardly and posteriorly, where it forms a spindle-shaped dilatation (receptaculum seminis) ; its continuation, the spermatic duct, unites with the Fig. 193. — Diagram of genitalia of a Cestode. g.p.^ genital pore; $ ^, male and female ducts opening into genital sinus; c.s.^ cirrus sac; v.d.^ coiled vas deferens {" outer seminal vesicle") ; vag.^ vagina ; sent, rec, seminal receptacle ; sp. d., spermatic duct ; C.c, fertilization canal ; vit. d., vitelline duct ; sk. g. , shell gland; ut. c, uterine canal; ttt., uterus; Ov., ovary; /, pumping organ. Cf. figs. 191 and 233. (Stephens.) oviduct, the common duct of the ovaries (fig. 191). The ovaries, usually two in number, are compound tubular glands in the posterior half of the proglottis, which extend into the medullary layer, but ventral to the median plane. At the origin of the oviduct there is frequently a dilatation pro- vided with circular muscles (suction apparatus), which receives the ovarian cells and propels them forward. After the oviduct has received the spermatic duct the canal proceeds as the fertilization canal, and after a very short course receives the vitelline duct or ducts, and then the numerous ducts of the shell glands (ootype). [Although the nomenclature of these parts varies, we may consider the oviduct as extending from the ovary to the shell gland and as receiving the spermatic duct and then the vitelline duct and the ducts of the shell gland. The short piece into which the shell gland ducts open corresponds to the ootype in the flukes, but in the tape- worms this portion of the canal is seldom dilated. From this point 296 THE ANIMAL PARASITES OF MAN the oviduct is continued as a shorter or longer tube, the uterine canal or true oviduct opening into the uterus proper. — J. W. W. S.] The vitellariuni may be single, but often exhibits its primitive duplication more or less distinctly, in which case it is situated at the posterior border of the segments in the medullary layer (fig. 191). The original position of the double organ is, moreover, the same as in the Trematodes, i.e.j at the sides of the proglottids, and thence eventually extending more or less on both surfaces (figs. 192 and 194) ; the gland is then distinctly grape-like and the follicles lie mostly in the cortical layer. The tgg cell that has been fertilized and supplied with yolk cells receives the shell material at the point of entry of the shell gland ducts, and, as a complete (^gg, then moves onward to the uterus. In those cases in which the uterus in its further course presents a convoluted canal, and may form a rosette (pseudo-phyllidea), there is an external Vvs Fig. 194.— Part of a transverse section through a proglottis of Dibothriocephalus la/us. Ct.^ cuticle; C, cirrus; Vvs., vitelline follicles; L.M., longitudinal muscles; T., testicles j M., medullary nerve ; S.c, subcuticle ; T.m., transverse muscles ; UL, uterus. 20/1. opening which is usually separate from the genital pore, and lies on the same or the opposite surface. In all other cases, however, the uterus terminates blindly and is represented by a longer or shorter sac lying in the longitudinal axis (fig. 191), but in many forms trans- versely. With the accumulation of eggs it becomes modified in various ways : (i) it sends out lateral branches (fig. 241), or (2) forms numerous isolated sacs (parenchymal capsules) containing single eggs or groups of eggs (fig. 217) ; further, (3) in some cases at the blind end one or more special thick-walled cavities are formed (PARUTERINE ORGANS Or UTERINE CAPSULES), in which all or most of the eggs are collected, the uterus then undergoing atrophy. In species in which the uterus lacks an opening, simultaneously with the growth of this organ an atrophy of the male apparatus, at least of the testes and their excretory ducts, takes place ; this atrophy also frequently occurs in the female glands, so that the entire mature segments have besides the uterus only traces of the genitalia left. DEVELOPMENT OF THE TAPEWORMS 297 In the Acolei'iicv the vagina is more or less extensively atrophied, and in any case has no external opening. A number of genera are distinguished by the duplication of the genitalia in every segment ; the genital apparatus in its entirety, or with the exception of the uterus, is double, or the genital glands and the uterus are single, but the cirrus, vas deferens and vagina are double. On comparing the genitalia of the Trematodes and Cestodes the parts will be found to agree, but the vagina of the Cestodes corresponds with the uterus of the Trematodes, and the uterus of the tapeworms to Laurer's canal of the Trematodes, which in most of the Cestodes has lost its external orifice. Development of the Tapeworms. Copulation. — As each proglottis possesses its own genital apparatus, and male as well as female organs are present, the following processes may occur : (i) self- or auto-fecundation (without immissio cirri) ; (2) self- or auto-copulation (with immissio cirri) ; (3) cross-copula- tion between proglottids of the same or different chains (of the same species) ; and (4) cross-copulation in the same proglottis in species with double genital pores. These various modes have actually been observed. In those species w^hich lack the vagina (Acoletnce) it appears that the cirri, which are always furnished with hooks, are driven into the tissues and for the most part reach the receptaculum seminis. The eggs of all Cestodes are provided with shells, but the shells, like their contents, vary. In genera that possess a uterine pore the mature eggs frequently do not differ from those of the Distomata ; they have a brown or yellow shell of oval form provided with an operculum, and contain a number of yolk cells in addition to the fertilized ovarian cell (fig. 128), but in other genera (with a uterine pore) the lid is absent and the egg-shell is very thin, the eggs of these genera resembling those of Cestodes in which the secretion of the vitellarium is a light albumin-like substance that contains only a few granules, and in which the egg-shell is very delicate and without operculum. The eggs of Tmtililce, for example, at first consist of egg-shell (ootype), ovum and yolk cells. The egg-shell is as a rule soft, colour- less and frequently deciduous, and the yolk is scanty in amount and contains few granules. The eggs are, moreover, more complicated than this. They enlarge and change their shape and various envelopes are developed around the embryo. The egg-shell proper often dis- appears, and one or more embryonal envelopes, or protoplasmic 298 THE ANIMAL PARASITES OF MAN layers, arise, so that eventually it is difficult to say whether the whole egg is present, and, if not, what the layers that remain really are. The embryonal development in most species takes place during the stay of the eggs in the uterus; in other species it takes place after the eggs have been deposited and are in water. Separate cells or a layer of cells always separate from the segmentation cells, as well as from the cells of the developing embryo, and form one or more envelopes round the embryo ; usually two such envelopes are formed, the inner one of which stands in intimate relationship with the embryo itself and is often erroneously termed the egg-shell, but more correctly the embryonal shell or emhryophore. In some species it carries long cilia, as in Dibothriocephalus latus, by aid of which the young swim about when released from the egg-shell ; as a general rule, however, there are no cilia and this envelope is homogeneous, or is composed of numerous rods and is calcified, as in Tcenia spp. (fig. 197). The second outer envelope ("yolk envelope ") (fig. 207, 3) lies close within Fig. 195. — ¥,gg o{ Diplogono- porus grandis, showing the morula surrounded by yolk cells and granules. 440/1. (After Kurimoto.) Fig. 196. — Uterine egg of Tania saghiata, G. Uterine shell with filaments ; the oncosphere with embryonal shell (embryophore) in the centre. 500/1. (After Leuckart.) the true (ootype) egg-shell, and remains within it when the embryo hatches out, and in many species, as in Tcenia spp., it perishes at the end of the embryonal development with the delicate egg-shell which was formed in the ootype, so that one observes not the entire egg with egg-shell but only the embryo in its embryonal shell, viz., the embryophore (fig. 197, a.). The embryo (the oncosphere) enclosed within the embryonal shell (embryophore) is of spheroidal or ovoid form (fig. 197, 6.), and is distinguished by the possession of three pairs of spines, a few terminal (flame) cells of the excretory system, and muscles to move the spines. No FURTHER DEVELOPMENT of the oncosphere takes place, either in the parent organism or in the open ; in fact, in all cases in which the oncospheres are already formed within the proglottids they do not become free, but remain in their shell ; it is only when the oncospheres are provided with a ciliated embryophore that they leave the egg-shell, and they even cast this ciliated envelope after having DEVELOPMENT OF THE TAPEWORMS 299 swum about in water by its means for a week or so. Sooner or later, however, all the oncospheres leave the host that harbours the parental tapeworm and reach the open, either still enclosed in the uterus of the evacuated proglottids, after the disintegration of which they then become free, or after being deposited as eggs in the intestine of the host; they then leave it with the faeces. In the former case also, the slightest injury to the mature proglottids while still in the intestine suffices to allow a part of the oncospheres in their embryo- phores to be released and mingled with the faeces. Here they are the generally, but falsely, so-called Taeniae '^ eggs." For, as stated above, the ''yolk" envelope and the true shell deposited in the ootype have before this disintegrated. In other cases, e.g., Hymenolepis spp., the uterine (ootype) shell persists in faeces (fig. 230). In any case the oncospheres must be transmitted into suitable animals to effect their further de- velopment ; in only very rare cases might an active invasion be possible, as, for instance, takes place with the miracidia of many Trematodes. The entry into an animal is, as a rule, entirely passive, that is to say, the oncospheres are swallowed with the food or water. Many animals are coprophagous and ingest the onco- spheres direct with the faeces ; others swallow them with water, mud, or food contaminated by such faeces, artificially by feeding suitable animals with mature proglottids of certain Cestodes or introducing the oncospheres with the food. As the mature tapeworm frequently finds the conditions suitable for its development in only one species of host, or in species nearly related, and perishes when artificially introduced into other hosts, experiment has taught us that to succeed in cultivating the oncospheres certain species of animals are necessary. Thus we are aware that the onco- spheres of Tcvnia solium, which lives in the intestine of man, develop only in the pig, and only quite exceptionally develop into the stage characteristic of all Cestodes — the cysticercus in the wide sense of the word— in a few other mammals. The oncospheres of T. saginaia develop further only in the ox ; those of T. marginata (of the dog) in the pig, goat, and sheep; those of T. serrata (of the dog) in hares and rabbits ; those of Dipylidiiim caniniim (of the dog and cat) in parasitic insects of the dog and cat, etc. It is not unusual that young animals Fig. 197. — a., oncosphere, in its radially striated embryophore (errone- ously termed egg-shell) of Tania africana. Greatly magnified. (After von Linstow.) b., freed oncosphere of Dipylidium caninum. (Alter Grassi and Rovelli.) Both oncospheres show six spines. Infection is easily produced 300 THE ANIMAL PARASITES OF MAN only appear to be capable of infection, while older animals of the same species are not so. Once introduced into a suitable animal, which is only excep- tionally the same individual or belongs to the same species as the one which harbours the adult tapeworm, the oncosphere passes into the larval stage common to all Cestodes, but varying in structure according to the species. In the simplest case — as, e.g., in Dibothrio- cephahis — such a larva resembles the scolex of the corresponding tapeworm, only that the head, provided with suckers, is retracted within the fore-part of the neck. Such a larval form is known as a plerocercoid {ifKrjpT]^, full; KepKo^, tail). They differ from the cysticer- coids in being solid larval forms, elongated, tape-like or oval, with the head invaginated. The conditions appear to be similar in Ligiila, Schistocephalns, Tricenopliorns, but here the larvae are very large, indeed as large iii the first-mentioned genera as the tapeworms originating from them, and the sexual organs are already outlined ; doubtless, however, this stage is preceded by one that corresponds to the scolex of the genus in question, and which represents the actual larval stage. In such cases the development of the body of the tapeworm from the scolex has already begun within the first or intermediate host ; in other cases, except in the single-jointed (monozootic) Cestodes, this only takes place in the definitive host. The direct metamorphosis of the oncosphere into the larval forms termed PLEROCERCOID has hitherto not been investigated, although Ligiila, SchlstocepJialus and Bothriocepliahis are very common parasites, but many circumstances point to the conclusions arrived at by us and by other observers. In the larval stages of other tapeworms we can always distinguish the scolex and a caudal- like appendage, vesicular in the cysticerci (fig. 200), compact in the cysticercoids (fig. 231). The scolex alone forms the future tapeworm, the variously formed appendage perishing. It has now been proved that the appendage, the caudal vesicle, originates direct from the body of the oncosphere, and therefore is primary, and that the scolex only subsequently forms through pro- liferation on the surface of this appendage. On account of this origin the scolex is generally regarded as the daughter, and the part usually designated as the appendage as the mother, originating from the oncosphere. Accordingly, two modes of development of the larval stage may be distinguished ; in the one case, plerocerci and plerocercoids, the oncosphere changes directly into the scolex, thus forming the body of the tapeworm within the primary host ; in the other case, cysticerci and cysticercoids, the scolex only forms secondarily in the trans- formed body of the oncosphere, which later on perishes, the scolex alone remaining as the originator of the tapeworm colony. DEVELOPMENT OF THE TAPEWORMS 301 We may summarize briefly what has been said regarding these larval forms. We have, firstly, sohd larval forms without any bladder. These arise directly from the oncosphere and are of two kinds, plerocercus and plerocercoid. Plerocercus is a solid globular larva with the head invaginated into the posterior portion. Plerocercoid (fig. 208) is a solid elongated larva also with the head invaginated mto the posterior portion, which is sometimes very long. Secondly, we have larval forms with bladders from which the scolices arise thus indirectly from tlie oncosphere. They are of two kinds, cysticercoid and cysticercus. Cysticercoid. — The bladder is but slighUy developed and is usually absorbed again. The anterior portion is, moreover, retracted into the posterior, and in some cases there is a long or a stumpy tail (figs. 220, 231). Cysticercns, or true bladder worms. (These may be divided into [i] cysticercus proper, consisting of a bladder and one scolex ; sec.c c.v. Fig. 198. — Diagram of a cysticercoid. Cf. figs. 220, 227. c.v.y caudal vesicle or bladder (small) ; sec. c, secondary cavity caused by the growth forward of the hind-body; /.,tail bearing six spines. (Stephens.) C.V. Fig. 199. — Diagram of a cysticercus. C.V., caudal vesicle or bladder; i., in- vagination of wall of bladder. (Stephens.) [2] coenurus, a bladder and many scolices ; [3] echinococcus, a bladder in which daughter bladders or cysts are developed, and then in these multiple scolices.) In the case of cysticerci a papilliform invagination forms, pro- jecting into the interior of the bladder (fig. 201). The layer of cells forming the papilla becomes divided into two laminae, the outer^ of which forms a kind of investing membrane (receptaciilum capitis) for the papilla. The head and suckers are now developed on the walls bounding the axial lumen of the papilla. The papilla eventually I.e., regarded from the interior or centre of the invagination. 302 THE ANIMAL PARASITES OP^ MAN evaginates, so that the receptaculum capitis now forms the inner surface of the hollow head, which eventually becomes solid. Our knowledge of the development of cysticerci in the wide sense of the word is limited almost exclusively to that of a few true '^bladder worms" (cysticerci); in other cases we know either only the terminal stage, i.e.j the complete larva, or, exceptionally, one of the intermediate stages, but we are not acquainted with a complete series ; the description must therefore be incomplete. We know from feeding experiments that, after the introduction of mature proglottids or of the fully developed ova of Tcenia crassicollis (of the cat) into the stomach of mice, the oncospheres escape from the shell in the middle portion of the small intestine, and a few hours later penetrate into the intestinal wall by means of a boring movement ; they have been found in this position twenty- seven to thirty hours after the infection. By means of this migration, for which purpose they employ their spines, they attain tke blood- vessels of the intestine; indeed, already nine hours after the infection and later they are found in the blood of the portal vein, and in the course of the second day after infection they are found in the capillaries of the liver, which these larvae do not leave. Leuckart, in experimental feeding of rabbits with oncospheres of Tcenia serrata (of the dog), found free oncospheres in the stomach of the experimental animal, but not in the intestine : however, he came across them again in the blood of the portal vein. The passage through the blood-vessels to the liver is the normal one for those species of Tcvnia the eggs of which become larvae in mammals ; even in those cases in which the oncospheres develop further in the omentum or in the abdominal cavity {Cysticercits temiicollis, C. pisifonnis), there are distinct changes observable in the liver that lead one to the conclusion that there has been a secondary migration out of the liver into the abdominal cavity. Indeed, one must not imagine that the young stages of the Cestodes are absolutely passive ; once they have invaded an organ they travel actively, and leave distinct traces of their passage. In other cases the oncospheres leave the liver with the circulation, and are thus distributed further in the body; they may settle and develop in one or more organs or tissues. Many oncospheres may, by travelling through the intestinal wall, penetrate through it and attain the abdominal cavity direct ; some, perhaps, pass also into the lymph stream. Where there are no blood and lymphatic vessels in the intestinal wall, as in insects, the oncospheres attain the body cavity or its organs direct ; in short, they never remain in the intestinal lumen itself, and only rarely — as in Hyrnenolepis murina of the rat — do they remain in the intestinal wall. DEVELOPMENT OF THE TAPEWORMS 303 When the infection has been intense, and the body is crowded with numerous oncospheres, acute feverish symptoms^ are induced, to which the infected animals usually succumb (" acute cestode tuberculosis ") ; while in other cases the alterations in the organs attacked— as the liver in mice and the brain in sheep — may cause death. Sooner or later the oncospheres of tapeworms come to rest, and are first transformed into a bladder, which may be round or oval according to the species. The embryonal spines disappear sooner or later, or remain close together or spread over some part of the bladder wall (fig. 200). Their discovery by v. Stein in the bladder worm of the ^' meal \vorm " (the larva of a beetle, Tenebrio molitor) first led to the conclusion that bladder worms (cysticerci) actually originate from the oncospheres of Tceniidcv. Fig. 200. — Diagram of dtvelopment of a cysticercus. i, solid oncosphere with six spines ; 2, bladder formed by liquefaction of contents ; 3, invagination of bladder wall ; 4, formation of rostellum (with booklets) and suckers -at the bottom of the invagination ; 5, evagination of head; 6, complete evagination effected by pressure. (Stephens.) The bladder may remain as a bladder, and then by proliferation the .scolex forms on its wall (fig. 202), or it may divide into an anterior so-called "cystic" portion and a solid tail-like appendage of various lengths, on which the embryonal hooks are to be found, and this is particularly the case in those larval forms (cysticercoids), e.^., those of Dipylidhun canimim, that develop in invertebrate animals, such as Arthropoda. As mentioned above one may regard the scolex as an individual that originates through proliferation of the wall of the parent cyst, mostly singly, but in those cysticerci that are termed coenurus (fig. 201) many scolices occur, whereas in those called echinococcus the parent cyst originating from the oncosphere of Tceiiia echinococcus (of the dog) first produces a number of daughter cysts, which in their 3G4 THE ANIMAL PARASITES OF MAN turn form numerous scolices. Echinococcus-like conditions also occur in cysticercoids, as, for instance, in those peculiar to earth- worms ; and similar conditions prevail in a larval form known as Staphylocystis, found in the wood-louse (Gloineris). Thus it happens in these cases that finally one tapeworm egg produces not one, but numerous tapeworms, for, under favourable conditions, each scolex can form a tapeworm. Fig. 201. — Section through a piece of a Ccemiriis cei'ebralis, with four cephalic invaginations in different stages of develop- ment. At the bottom of the invaginations the rostellum, hooks and suckers develop. (From a wax model.) Fig. 202. — Median section through a cysticercus, with developed scolex at the bottom of the invagination. (After Leuckait.) The rudiment of the scolex appears as a hollow bud, the cephalic invagination usually directed towards the interior of the bladder cavity; on its invaginated surface arise the four suckers, and the rostellum with the hook apparatus is formed in its blind end ; we thus get a T.ienia head, but with the position of the parts reversed (fig. 201). In many cysticerci the head rises up from the base of the cephalic invagination and is then surrounded by the latter. A more or less elongated piece of neck also develops, and even proglottids may appear, as in Cysticercus \ fasciolaris (the larva of Tcpnia crassicolUs of the cat) of the Muridas, a process somewhat analogous to that of Ligula, etc. The period that elapses from the time of .^ infection till the cysticercus is fully developed varies according to the species ; the cysticercus of Tcvnia saginata requires twenty-eight weeks, that of T. marginaia seven to eight weeks, that of T. solium three to four months, and that of T. echinococcns longer still. With one single exception {Archigetes) the larvae do not become sexually mature in the organ where they have developed ; they must enter the terminal host, a matter that is usually purely passive, the carriers of the larvae or infected parts of them being usually devoured by other animals. In this manner, for instance, the larvae (Cysticercns fasciolaris) found in mice and rats reach V Fig. ^wj — Cysticercus pisifor?nis in an evagi- nated condition, with neck, fore-body and bladder, with excretory network in its wall. 18/1. DEVELOPMENT OF THE TAPEWORMS 305 the intestine of cats ; those of the hare and rabbit (C. pisiformis) reach the intestine of dogs; those of the pig (C. celluloses) are introduced into man ; those of insects are swallowed by insectivorous birds ; those of crustaceans are ingested by ducks and other water fowl ; perhaps, also, the infection of herbivorous mammals is caused by their accidentally swallowing smaller creatures infected by larvae. Indeed, the researches of Grassi and Rovelli have taught us that such an intermediate host is not always necessary ; Hymenolepis nmrina of rats and mice in its larval stage lives in the intestinal wall of these rodents, and as a larva it passes into the intestinal lumen and develops into a tapeworm in exactly the same way as the larvae of other species that reach the intestine of the terminal host by means of an inter- mediate carrier. Probably this curtailed manner of transmission also occurs in many other species. In some cases the larvae actively quit the body of the intermediate host, as in the case of Ligula and Schisto- cepJiahis, which travel out of the body cavity of infected fish and reach the water, where they may be observed in hundreds in summer, at all events in some localities. The larval stage of Calliobothriuin — wrongly termed Scolex — has been observed swimming free in the sea, and the scolices of Rhynchobolhriiun, without their mother cysts, have been observed free within the tissues of several marine animals. In any case there is almost always a change of hosts, even in the single- jointed Cestodes, for the larva of Caryophyllceus, which lives in fishes of the carp family, is found in limicoline Oligochaetes, that of Gyro- cotyle (Chimaera) in shell-fish (Mactra), and different conditions can hardly be possible for Amphilina. Archigetes alone becomes sexually mature in the larval stage, but the life-history of this creature is not well known, so that it is not impossible that the attainment of sexual maturity as a larva in invertebrates (Oligochaetes) is perhaps abnor- mal, and somewhat analogous to the maturity of some encysted Trematodes. The METAMORPHOSIS OF THE LARVA into the tapeworm is rarely accomplished in a simple manner ; the transformation, however, is not complex in the single-jointed Cestodes, nor in Ligula and Schisto- cephalus ; the latter is swallowed by birds (Mergus, Anas, etc.), produces eggs after only a few days, and very soon quits the intestine of its terminal host. In all other cases it is the scolex only which, by proliferating at its posterior extremity, forms the proglottids, after having invaded as a larva the intestine of a suitable host. The mother cysts, or what corresponds to them, die, are digested, absorbed, or perhaps even eliminated ; on the contrary, segments found on the scolex during the larval stage, also in the case of Cysticercus fasciolaris, are retained. It is not certain whether the larvae of Dibothriocephalus lose any part. 20 306 THE ANIMAL PARASITES OF MAN The time required by the scolex to complete the entire chain of proglottids does not depend only on the number it has to produce, for Taenia echinococaiSy which, as a rule, only possesses three or four segments, takes quite as long a time for their growth (eleven to twelve weeks) as T. solium with its numerous seg- ments ; T. ccenitrus is fully developed in three to four weeks, and the same holds good for DihotJirlocephalus latus, which possesses many more segments than the above-mentioned Taenia of th^ -dog. In a number of species it has been possible to determine fairly accurately the average daily growth ; for instance, in Dibothriocephalus lattis the daily growth is 8 cm., in Tcvnia saginaia 7 cm., etc. The history of the development of the Cestodes demonstrates that persons and beasts harbouring larval tapeworms have become infected by having swallowed the oncospheres of the species of tapeworm to which they belong. In regard to Hymenolepis mnrina alone, it is known that the introduction of the oncospheres into those species of animals which harbour the adult tapeworm leads to the formation of the latter after the development of a larval stage in the intestinal w^all ; nevertheless, only young animals (rats) are capable of infection, for a previous infection, or the presence of mature tape- worms in the intestine, appears to produce a kind of immunity. Biology. In their adult stage, the tapeworms inhabit almost exclusively the alimentary canal of vertebrate animals, with but few exceptions the small intestine, and a few species select definite parts of it. A small number of Rhynchohothriidce of marine fishes live apparently always in the stomach, while in rays and sharks the spiral intestine is their exclusive site. Bothriocephali generally attach themselves with their head on to the appendices of the pylorus of fishes ; other species (Hymenolepis diminuta) occasionally fix their head in the ductus choledochus, and this is more frequent still in the tapeworms of the rock badger (Hyrax), which occasionally penetrate entirely into the biliary ducts. Stilesia hepatica, Wolffh., has so far only been found in the bile-ducts of its host (sheep and goat, East Africa). In the disease of sheep induced by Cestodes, the w^orms have been observed also in the pancreas. Specimens found in the large intestines were probably being evacuated. The Cestodes are looked upon as fairly inert creatures, this opinion having been formed by observing their condition in the cold cadavers of warm-blooded animals. Actually, however, they are exceedingly active, and accomplish local movements within the intestine, for they have been found in the ducts communicating with BIOLOGY OF THE CESTODES 307 the bowel, or in the stomach, and may even make their way forward into the oesophagus. They also invade other abdominal organs through abnormal com- munications, or through any that may be temporarily open between the intestine and such organs ; they thus reach the abdominal cavity or the urinary bladder, or they work their way through the peritoneum. They produce changes in the intestinal mucous membrane at the place of their attachment, the alterations varying in intensity accord- ing to the structure of the fixation organs. The mucous membrane is elevated in knob-like areas by the suckers ; the epithelial cells become atrophied or may be entirely obliterated. Dipylidium caninnni bores into the openings of Lieberkiihn's glands with its rostellum, dilating the lumen to two or three times its normal size, while the suckers remain fixed between the basal parts of the cells. Species with powerful armatures penetrate deeper into the submucosa, and some that are not provided with exceptionally strong armatures, or are even unarmed, may be actually found with the scolex embedded in the muscles of the intestinal walls or even protruding beyond {Tcenia tetragonal MoL, in fowls, etc.). Other species, again, even cause perforation of the walls of the intestine of their hosts. It is generally assumed that tapeworms, which almost without exception live in the gut of vertebrates, get their nutriment from the gut contents, which apparently they absorb through the whole body surface (cuticular trophopores). In favour of this view is the existence of fat drops in the proglottids, the identity in colour in certain forms between that of the fresh worm and the gut contents and the passage of certain substances derived from medicines (iron and mercury preparation) into the worms in the gut, etc. Whether the suckers are concerned in the absorption of nutriment and to what extent is still questionable. The length of life of the adult tapeworm certainly varies ; as a rule it appears to last only about a year ; in other cases (Ligula) it averages only a few days, but we are likewise aware that certain species of Cestodes of man attain an age of several or many years (thirty-five). The natural death of Cestodes often appears to be brought about by alterations in the scolex, such as loss of the hooks, atrophy of the suckers and rostellum, finally the dropping off of the scolex; it is unknown whether a chain of segments deprived of its scolex then perishes or whether it first attains maturity. It has already been mentioned that in a few species the foremost proglottids are transformed into organs of fixation on the normal loss of the scolex. Abnormalities and malformations are encountered relatively frequently in the Cestodes — such as abnormally short or long segments ; the so-called triangular tapeworms, which — if belonging to the Tceniidce — always possess six suckers ; often 308 THE ANIMAL PARASITES OF MAN also club-shaped segments occur between normal ones, or there may be a defect in one segment or in the centre of a number following one another (fenestrated segments) ; bifurcated chains of segments have likewise been observed, as well as incomplete or complete union of the proglottids, abnormal increase of the genital pores, reversion of the genitalia. Besides the above-mentioned increase of the number of suckers on the scolex (in Taeniae), there may be a decrease in the number ; in other cases the crown of hooks may be absent, or abnormally shaped hooks may be formed. Classification of the Cestoda of Man. Order. Pseudophyllidea, Cams, 1863. Scolex without proboscis or rostellum. Head " stalk" absent. Scolex never with four, generally with two (or one terminal) bothria.^ Vitellaria numerous. Uterine opening present. Genitalia do not atrophy when uterus is developed. In large majority of proglottids eggs (or, if formed, their contents) are at the same stage of development. Family. Dibothriocephalidae, Liihe, 1Q02. Syn. : Diphyllobothritdce, Liihe, 1910. Genitalia repeated in each proglottid (polyzootic Cestodes). Ventral and dorsal surfaces flat. Cirrus unarmed. Cirrus and vagina if non-marginal open on the same surface as the uterus. Uterus long, convoluted, often forming a " rosette,"" never dilates into a uterine cavity. Eggs thick shelled, operculated, constantly being formed in mature proglottids. Sub-family. Dibothriocephalinae, Liihe, 1899. Syn. : Diphyllobothriina^ Liihe, 19 10. Segmentation distinct. Scolex unarmed, elongated, sharply separated (generally by a neck) from the first proglottis. Cirrus and vagina open ventrally. Genital pores non-alternating. Vas deferens surrounded by a muscular bulb. Receptacu- lum seminis large, sharply separated from the spermatic duct. Order. Cyclophyllidea, v. Beneden. Four suckers always present. Uterine opening absent. Vitellarium single. Genitalia atrophy when uterus is fully developed. ' Bothridia or '' phyllidia " are outgrowths from the scolex. They are concave and extremely mobile. By some authors the term '' phylliditim " is used for the outgrowth, and the term '' bothridmm" is restricted to the muscular cup. Bothria, on the other hand, are grooves more or less wide, the musculature of which is only slightly developed and is not separated off internally from the parenchyma. Acetabula, or suckers in the usual sense, are hemispherical cups, without lips and with musculature separated internally from the parenchyma. CLASSIFICATION OF THE CESTODA 309 Family. Dipylidiidae, Liihe, 1910. Rostellum if present armed. Suckers unarmed. Uterus breaks up into egg capsules. Paruterine organs absent. Family. Hymenolepididae, Railliet and Henry, 1909. Segment always broader than long. Genitalia single. Longitudinal muscles in two layers. Genital pores unilateral. Testes one to four. Uterus persistent, sac- like. Eggs with three shells. Family. Davaineidae, Fuhrmann, 1907. Rostellum cushion-shaped. Armed with numerous (sixty to several thousand) hammer-shaped hooks in two (rarely one) rows. Sub-family. Davainelnae, Braun, 1900. Suckers armed. Uterus breaks up into egg capsules. Paruterine organs absent. Family. Taeniidae, Ludwig, 1886. Suckers unarmed. Uterus with median longitudinal stem and lateral branches. Female genitalia at the hind end of the proglottis. Genital pore irregularly alter- nating. Testes numerous in front of female genitalia. Ovary with two lobes (wings). Vitellarium behind the ovary. Embryophore radially striated. The Cestodes of Man. Most of the species to be mentioned live in man in their adult stage and occupy the small intestine ; man is the definite host of these parasites, but is not the specific host for all the species ; some of these -species, as well as others (of mammals), may occur in man also in the larval stage. Family. Dibothrlocephalldae. Sub-family. Dibothriocephallnae. Genus. Dibothriocephalus, Liihe, 1899. Syn. : Diphyllobothriiun^ Cobbold, 1858; Bothriocephalus, ^. ^. Rvid., 18 19; Dibothrius^ p. p. Rud., 1819 ; Dibothriimi, p. p. Dies., 1850. Scolex egg-shaped ; dorsal and ventral bothria elongated, moderately strong, -cutting rather deeply into the head ; genitalia single in each proglottis ; papillae in the vicinity of the genital atrium ; the testes and vitellaria are in the lateral fields, the 310 THE ANIMAL PARASITES OF MAN former in the medullary layer, the latter In the cortical layer on both surfaces, and occasionally extending to the median line ; the ovary ventral, the shell gland dorsaL The uterus is in the central field, taking a zigzag course, and frequently forms a rosette. Dlbothriocephalus latus, L., 1748. Syn. : TcBtiia lata, L., 1748; Tcenia vulgaris, L., 1748; Tcenia grisea, Pallas, 1796; Tcenia ineinbrajiacea. Pall., 1781 ; Tania tenella, Pall., 1781 ; Tcenia dentatay Batsch, 1786; Boihriocephalus latus, Bremser, 1819; Dibothrium latinn. Dies., 1850; Boihriocephalus cristatus, Davaine, 1874' ; Boihriocephalus balticusy Kchnmstr., 1855; Boihriocephalus latissimus, Bugn., 1886. Length 2 to 9 m. or more ; colour yellowish-grey ; after lying in water the lateral areas become brownish and the uterine rosette brown. The head is almond-shaped, 2 to 3 mm. in length, the dorso- ventral axis is longer than the transverse diameter ; the head, therefore^ generally lying flat, conceals the suctorial grooves at the borders ; these suckers are deep and have sharp edges (fig. 205). The neck varies in length according to the degree of contraction and is very thin ; there are 3,000 to 4,200 proglottids and there may be more ; their breadth is usually greater than their length, but in the posterior third of the body they are almost square, and the very oldest are not uncommonly longer than they are broad. There are numerous testes situated dorsally in the medullary layer of the lateral fields ; the vas deferens (fig. 192) passes dorsally in transverse loops in the central field anteriorly and forms a seminal vesicle before its entry into the large cirrus pouch. The orifice of the vagina is close behind the orifice of the cirrus ; the former passes almost straight along the median line posteriorly,. and widens into a receptaculum seminis shortly before its junction with the oviduct ; the ovary is bilobed, in shape like the wings of a butterfly, ventrally in the medullary layer ; the shell glands lie in the posterior recess of the ovary ; the uterus, forming numerous transverse convolutions, passes ventral to the vas deferens forwards. Eggs (fig. 207) large, with brownish shells and small lids, 68 /a to 71/1 by 45 A*-; the ovarian cell, which is already, as a rule, in process of segmentation, is surrounded by numerous large yolk cells ; the proglottids nearest the posterior extremity are frequently eggless. The eggs, which are deposited in the intestine and evacuated ^ Until recently this worm, which was understood to belong to a separate species, was proved on examination by R. Blanchard ("Mai. Par.," 1896), to be Dlbothrio- cephalus latus. Compare also Galli-Valerio, in Centralbl. f. Bakt,, Path, und In- fektionskr., 1900 (i), xxvii, p. 308. DIBOTHRIOCEPHALUS LATUS 311 with the faeces, hatch in water after a fortnight or more ; the embryonal integument (embryophore) of the oncosphere is provided with ciUa ; after bursting open the Hd of the egg the oncosphere in its embryo- phore (fig. 207) reaches the water and swims slowly about ; often it slips out of its ciliated embryophore, sinks to the bottom and is capable of a creeping motion; sooner or later it dies in the water. The manner and means of its invasion of an intermediate host are still 1 r^ t a ril rf i?- tt m ffi 33 Fig. 204, — Various chains of segments of Dibothriocephalus la/us, showing the central uterine rosette. (Natural size.) Fig. 205. — Transverse sec- tion of the head of Dibo- thriocephahis latus, 30/1. Fig. 206. — Fairly mature proglottis of Dibothriocephalus lat'iis. The vitellaria are at the sides ; the uterus, filled with eggs, is in the middle, also the vagina (the dark stripe passing almost straight from the front to the back), and the vas deferens (almost hidden by the uterus). Above in the centre is the cirrus sac, and below the shell gland and ovary are seen. 15/1. (From a stained preparation.) unknown ; yet we are aware that the larval stage (plerocercoid, fig. 208), which resembles the scolex and may reach a length of 30 mm., lives in the intestine, in the intestinal wall, in the liver, spleen, genital glands and muscular system (fig. 209) of various fresh-water fish, the pike {Esox lucius), the miller's thumb {Lota vulgaris), the perch (Perca fliwiatilis), Salmo umbla, Triitta vulgaris^ Tr. lacustrls, Thyntallis vulgaris (grayling), Coregonus lavareUiSj C. albula (in Europe) and Onchorhynchiis perryi (in Japan). The transmission of the plero- cercoids from these fish to the dog, cat and man (Braun, Parona, Grassi and Ferrara, Grassi and Rovelli, Ijima, Zschokke, Schroeder) leads to ,12 THE ANIMAL PARASITES OF MAN the development of the broad tapeworm, the growth of which is rapid. In my experiments on human beings the average number of pro- glottids formed per diem averaged thirty-one to thirty-two for five weeks, with a length of 8 to 9 cm. According to Parona the eggs Fig. 207. — Dibothriocephalus lafus : development of egg. i, segmentation complete ; some cells of the blastosphere have migrated through the yolk and have flattened to form f, the yolk envelope ; others form a layer of flattened cells (<;) forming the embryophore ; the remaining cells {d) of the blastosphere form the hexacanth embryo. 2, embryophore [e) is becoming thicker. 3, the ciliated embryo has been pressed out of the shell ; s\ the operculum ; ^, the yolk envelope remaining in the shell [s) ', j, the yolk consisting of separate cells. 4, a free-swimming larva much swollen by the water. (After Benham and Schauinsland.) appear twenty-four days after man has been infected. Zschokke found the average growth in the experimental infection of man between 5*2 and 8'2 cm. per diem, and the person experimented upon by Ijima evacuated a piece of a Dibothriocephalus latus, 22*5 cm. in length, only twenty-one days after the infection. DIBOTHRIOCEPHALUS LATUS 313 The " broad tapeworm " is a frequent parasite of man in some districts, but it also occurs in the domestic dog, and on rare occasions is found in the domestic cat (together with Dibotliriocephahis felis^ Crepl.) and fox. French Switzerland and the Baltic Provinces of Russia are the centres of distri- bution ; from the former districts the distribution radiates to France and Italy (Lombardy, Piedmont) ; from the Baltic Provinces over Ingermanland to Petrograd, over Finland to Sweden (on the shore of the Gulf of Bothnia), in a southerly direc- tion to Poland, and into the Russian Empire and across it to Roumania, and towards the west along the coast of the Baltic Sea to the North Sea, where, however, its frequency considerably diminishes (Holland, Belgium, and the North of France). In Turkestan and Japan the ^' broad tapeworm " is the most frequent parasite of man ; it has been reported in Africa from the vicinity of Lake N'gami as well as from Madagascar ; cases, in Fig. 208.— Plero- cercoid of Dibothrio- cephalus latus. A., with the head evaginated ; B., with the head invaginated. From the muscle of the pike. Fig. 209. — A piece of the body wall of the Burbot, Lo/a vulgaris. The tangential section has exposed the muscles of the trunk, with a plerocercoid of Dibothriocephalus latus. Natural size. part at least imported, have also come under observation in North America. In Germany Dihothriocephaliis latus — apart from the fact that it is undoubtedly imported from Switzerland, Russia or Italy— is 314 THE ANIMAL PARASITES OF MAN particularly frequent in East Prussia amongst the inhabitants of the Courland Lagoon district, on the Baltic ; it is, moreover, also found in the Province and even in the City of Konigsberg. In West Prussia and Pomerania it is very much scarcer. It is also found in Munich and in the vicinity of the Lake of Starnberg (Bollinger). Krabbe found it in 10 per cent, of the sufferers from tape- worms in Denmark; Szydlowski found the ova of this worm in Dorpat in 10 per cent, of the faeces examined; Kruse found the worm in 6 per cent, of post-mortems ; Kessler, in Petrograd, found the eggs in the faeces in 7*8 per cent. ; at post-mortems he found the worms in 1*17 per cent., though Winogradoff only found it in o'8 per cent. In Moscow, according to Baranovsky, 8*9 per cent, of the faeces examined contained the ova of Dihothrio- cephalus. In the interior and southern provinces of Sweden the worm, according to Lonnberg, is only found sporadically, but, on the other hand, in Angermanland about to per cent, of the popula- tion is affected ; while again in Norbotten the majority of persons are affected, and in Haparanda the entire population (with the exception of infants) harbour this parasite. In Switzerland D. latiis is very frequent in close proximity to the lakes of Bieler, Neuchatel, Morat and Geneva (according to Zaeslin 10 to 15 to 20 per cent, of the population are affected) ; the parasite is less frequent in districts one to four hours removed from these lakes. Of the fish from Swiss lakes examined by Schor those from Lake Geneva were most commonly infected, and especially Lota sp. and Perca sp. The frequency and distribution have, nevertheless, decreased perceptibly in places; at the commencement of the eighteenth century the broad tapeworm was very common in Paris, at the present date it only occurs when imported (Blanchard) ; in Geneva, also, according to Zschokke, it has become rarer (formerly 10 per cent., now only i per cent.). The disturbances produced in man by the presence of broad tapeworms are, as a rule, very trifling ; in other cases they produce partly gastric disorders and partly nervous symptoms ; in a number of cases, again, they set up severe anaemia, apparently caused by toxins produced by the worms and absorbed by the host. There is no danger of auto-infection, as the larval stage lives only in fishes, not in warm-blooded animals. The case reported by Meschede (ova like those of Dihothriocephalus latus in the brain of a man who had suffered from epilepsy for six years) must be otherwise explained. Human beings, like other hosts, can only acquire the broad DIBOTHRIOCEPHALUS CORDATUS 3I5 tapeworm by ingesting its plerocercoids with the previously men- tioned fresh-water fishes ; the opportunity for such infection is afforded the more readily by the fact that not only do the lower classes not pay sufficient attention to the cooking of fish, so that all the larvae that are present may be killed, but also in certain localities the custom exists of eating some parts of these fishes in a raw condition ; even the mere handHng of the usually severely infected intermediary hosts may occasionally cause infection. The plerocercoids are as well known as, but differ materially in appear- ance from, the cysticerci {Cysticercns celhdosce) of pig's flesh. In Germany the occurrence of the plerocercoids of Dihothriocephalus latus has been confirmed in the pike, miller's thumb and perch of East Prussia, and more particularly in those taken from the Courland Lagoon. The life of D. latus is a very long one (six to fourteen years), as is deduced from persons who have left D. latus regions after they have been infected. According to the experiments of M. Sciior, plerocercoids of D. latus placed in slowly warmed water completely lose their move- ment at 54*^ to 55' C. ; they survive the death of their host for several days ; they are killed by low temperatures - 3° to -h 1° C. in two days ; strong acids and salt solutions kill them at once, also high temperatures, but all the same at least ten minutes is required in boiling or frying fish in order to kill the plerocercoids with certainty. Dibothriocephalus cordatus, R. Lkt., 1863. Syn. : Bothriocephalus cordatus^ R. Lkt. Length, 80 to 115 cm. ; the head is heart- Fig. 210.— Cephalic end shaped and measures 2 by 2 mm. The °^ Dibothriocephalus corda^ ^ in f ^^^ ' °" ^^^ '^" Viewed suctorial grooves are on the flat surface ; the sideways, on the right from segments commence close behind the head and ^^^ dorsal surface, showing ^ . . a suctorial groove. (After increase rapidly in breadth. At only 3 cm. Leuckart.) behind the head they are already mature ; the greatest breadth attained by them averages 7 to 8 mm., the length 3 to 4 mm. ; the number of proglottids averages 600 ; the most posterior ones are usually square. The uterine rosette is generally formed of six to eight lateral loops. The eggs are operculated and measure 75 /a by 50 />t. Dibothriocephalus cordatus is a common parasite of the seal, the walrus and the dog in Greenland and Iceland, occasionally of man also. No doubt its larva lives in fishes. 3l6 THE ANIMAL PARASITES OF MAN The statement that D. cordatus also occurs in Dorpat in human beings has been proved erroneous {Zool. Anzeiger^ 1882, v, p. 46), as also has the report that this worm lives in hares in the neighbourhood of Berlin, whither it was supposed to have been carried by Esquimaux dogs (Rosenkranz in Deutsch. med. Wochenschr.^ ^^77, iiij P* 620). The parasite stated by the author to i)e D. cordatus is Tcenia pectinata, Goeze, which has been known since 1766. Dibothriocephalus parvus, Stephens, 1908. Largest gravid segments 5 by 3 mm. Uterus forms a central rosette with four to five loops on each side of median Hne. In a pro- glottid measuring 3*5 by 2*25 mm. the genital atrium is situated 0*4 to o'5 mm. behind the anterior margin and the uterine opening the same distance behind the genital atrium. Calcareous corpuscles absent in the preserved specimens. Eggs operculated, 59*2 /x by 407 /x. Distinguished from Dibothriocephalus latus — (i) by the size of gravid segments (the minimum width of gravid segments of D. laUis is 10 to 12 mm., so that D. parvus is a much smaller worm) ; (2) quadrate segments of D. latus measure 6 by 6 mm., those of D. parvus 4 by 4 mm. ; (3) by the eggs. From D. cordatus it is distinguished by — (i) D. cordatus has only fifty immature segments, D. parvus has at least 200, possibly more ; (2) mature segments of D. cordatus measure 7 to 8 mm., maximum width of D. parvus is 5 mm. ; (3) quadrate segments of D. cordatus measure 5 to 6 mm. ; (4) D. cordatus has six to eight uterine loops ; (5) Z). cordatus measures 75/1, to 80//, by 50//.. Habitat. — Intestine of man (Syrian, in Tasmania). Genus. DIplogonoporus, Lonnbrg., 1892. Syn.: Krabbea^ R. Blanch., 1894. The scolex is short and has powerful suctorial grooves ; no neck ; the proglottids are short and broad ; there are two sets of genital organs side by side in each segment, which in all essentials resemble the single one of Dibothriocephalus. Parasitic in whales and seals, occasionally in man. Diplogonoporus grandls, R. Blanch., 1894. Syn. : Bothriocephalus sp., Ijima et Kurimoto, 1894 ; Krabbea gra?tdis, R. Blanch. Scolex unknown ; chain of proglottids over 10 m. in length, 1*5 mm. broad anteriorly, 25 mm. broad posteriorly. The proglottids are very short (0*45 mm.), but 14 to 16 mm. broad. On either side to the right and left of the worm, along the entire ventral surface, there is a longitudinal groove ; these grooves are nearer to each other than SPARGANUM 317 to the lateral margin ; in them, lie the genital pores, and they are in the same sequence as in Dibothriocephalns ; corresponding to the scanty length (0-45 mm.) of the proglottids, the ovary is only developed transversely ; the uterus only makes a few loops. Eggs (fig. 195) thick shelled, brown, 63 yu, by 48 yit to 50 fju. This parasite has hitherto been observed twice in Japanese. Similar species are known in Cetacea and seals. ; Fig. 211. — Diplogonoporus grandis, Llihe, 1899: ventral view of a portion of the strobila, showing two rows of genital pores and partially extruded cirri. (After Ijima and Kurimoto.) ^Z€t-^ zct. ^==^5^ Fig. 212. — Diplogonoporus grandis : ventral view (diagrammatic) of genitalia of left side ; cir, cirrus ; cir.o, cirrus opening ; dtg.^ vitelline duct ; ov.^ ovary ; ovd., oviduct ; j^., receptaculum seminis ; w/., uterus; ut.o., uterine pore ; vag.y vagina; vag.o., vaginal pore; vd, vas deferens. X 150. (After Ijima and Kurimoto.) Sparganum, Diesing, 1854. The term S^ar^^;///m, invented by Diesing, is used as a group name of larval bothriocephalid Cestodes whose development is not suffi- ciently advanced to enable them to be assigned to any particular genus. Sparganum mansoni, Cobb., 1883. Syn. : Ligula mansoni, Cobbold, 1883; Bothriocephalus linguloides,K. Lkt., 1886 ; Bothriocephalus ?nansoni, R. Blanch., 1886. These plerocercoids were discovered in 1882 by P. Manson during the post-iuoriem on a Chinaman who had died in Amoy, 3i8 THE ANIMAL PARASITES OF MAN Fig. 213. — Ce- phalic end of Sparga- num mansoni, Cobb. {After Leuckart.) twelve Specimens being found beneath the peri- toneum and one free in the abdominal cavity. Cobbold described them as Ligula uiaiisoni, and Leuckart, who contemporaneously reported a case in Japan, termed them BothriocephaUis liguloldes. Ijima and Murata reported eight further cases, and Miyake records nine further cases, seven of which are recorded in Japanese literature. The plerocercoid, which hitherto alone is known to us, attains a length of 30 cm. and a breadth of 3 to 6 to 12 mm. The ribbon-shaped body is wrinkled, the lateral borders are often somewhat thickened, so that the transverse section has the form of a biscuit ; the anterior end is usually w^ider and has the head provided w^ith two weak suctorial grooves, either retracted or protracted. The parasite makes migrations within the body, and thus may reach the urinary passages ; then it is either evacuated with the urine or has to be removed from the urethra ; not rarely it causes non-infiam- matory tumours on various parts of the skin, which are at times painful and at times vary in size. Nothing is knov/n of its development and origin. Sparganum proliferum, Ijima, 1905. Syn. : Plerocercoides prolifer, Ijima, 1905 ; Sparganum prolifer^ Verdun, Man son, 1907. These plerocercoids produce an acne-like con- dition of the skin. The condition is really one of capsules in great abundance in the subcutaneous tissue and less so in the corium and in the inter- muscular connective tissue. The encapsuled worms in the corium feel like embedded rice grains and raise the epidermis, giving rise to an acne-like con- dition. Many thousands occur scattered over the body; in Ijima's Japanese case there were over 10,000 in the left thigh. The worms when they first appear in the skin cause itching. The capsules are ovoid, generally about i to 2 mm. in diameter, but they may be smaller and also much larger. The larger ones occur in the subcutaneous tissue. The capsules consist of dense tough connective tissue. Each capsule, as a rule, contains one worm, but as many as seven may occur. The skin of areas that have been long infected is swollen Fig. 21^.— Sparga- num mansoni : on the right in transverse section. Natural size. (After Ijima and Murata.) SPARGANUM PROLIFERUM 3^9 V A A and indurated or adherent, giving a somewhat elephantoid appearance. The subcutaneous tissue is thick and filled with slimy fluid or in other parts sclerosed. The Worm. — The chief peculiarity is its irregular shape and its reproduction in the larval stage by forming supernumerary heads, which are supposed to wander about the body. The simplest forms are thread-like bodies, flat or round, 3 mm. long and 0*3 mm. in diameter, but they may be 12 mm. long by 2'5 mm. broad. The narrow end is the head, which in life invaginates and evaginates, but there is no indication of any suckers, except an inconstant terminal depression. In addi- tion to these simple forms the most compli- cated and irregular forms occur, due to the ^^^- ^is.—Sparganumproii- c \- r 1 J /u J X i. • L f'^ ' •*^" ^'^^ t)uds, right ex- tormation of buds (heads) at various parts, tended, x 4. (After ijima.) "^,1 Fig. 216. — Sparganmn ptolifeiii7n, x lo. (After Stiles.) The detachment and growth of a head account for the presence of more than one worm in a cyst. The irregularity in form is also 320 THE ANIMAL PARASITES OF MAN increased by the presence in the subcuticular tissue of the worm of reserve food bodies. These bodies are supposed to be of this nature and are spherical, lOo /x to 300 fju in diameter, but also much elongated. Calcareous bodies in the Japanese worms were 7*5 /-t to 12 /^ ; in the Florida worms 8-8 fi to 17*6 jx. Mode of Infection. — Probably from eating uncooked fish. Distribution. — Japan, Florida. Family. Dipylidiidae, Liihe, 1910. Genus. DIpylidium, R. Lkt., 1863. Rostellum retractile, with several rings of alternating hooks ; the latter with a disc-like base, having the shape of the thorns of a rose. Genital pores opposite ; genitalia double. Testes very numerous in the central field ; ovary with two lobes ; the vitellaria, which are smaller, behind them ; the uterus forms a reticulum, in the network of which the testicular vesicles lie ; later on it breaks up into sacs enclosing one or several eggs. The eggs have a double shell. \ DIpylidium caninum, L., 1758. Syn. : Tcenia ca?thta, L., 1758, p. p. ; Tcznia moniliformis^ Pallas, 1781; Tcenia ciicu7nerina, Bloch, 1782; Tcenia elliptica, Batsch, 1786; DIpylidium cucimierimim^ Lkt., 1863. This worm measures 15 to 35 cm. in length and 1*5 to 3 mm. in breadth. The scolex is small, rhomboidal, and has a club-shaped rostellum on which hzz:i Fig. 217. — Dipylidium caninum : on the left, the scolex, neck and the first proglot- tids ; on the right, at the top, a packet of ova ; below, hooks of the rostellum, side and front views ; below, an ovum. Various magnifications. (After Diamare.) Fig. 218. — Dipylidium caninum '. egg show- ing a, egg-shell (vitelline membrane of Moniez) j b, albuminous coat ; c, internal shell formed of or secreted by an outer layer of blastomeres (Moniez) ; d^ hexacanth embryo. (After Benham and Moniez.) there are, in three to four rings, forty-eight to sixty hooks resembling rose thorns, the size of those in the foremost being 1 1 />t to 15 /^ and those in the hindmost ring 6 //,. The neck is very short, the most anterior segments broad and short, the middle as long as they are broad ; the DIPYLIDIUM CANINUM 321 Fig. 219. — Dipylidium caninum : central portion of a proglottis. C.p., cirrus sac; V.S., vitellaria ; ^Jtr.z'., excretory vessels ; 7"., testicles lying in the meshes of the uterine reticulum which laterally forms pouches ; O., ovary ; C/., reticulum of uterus ; V., vagina and seminal receptacle (below ovary). Magnified. '_ (After Neumann and Kailliet.) Fig. 220. — Dipylidium caninum : development of embryo, i, solid hexacanth embryo ; 2, primitive lacuna {a) in the embryo ; 3, elongation of hinder part, rudiments of sucker and rostellum appearing ; 4, " body " and " tail " distinct, (d) and (c) excretory system ; 5, fore-body invaginates into hind-body, excretory bladder has a pore; 6, tail has dropped off; scolex growing up into secondary cavity formed by fore-body ; the primitive cavity has been absorbed at stage 4. (After Benham, Grassi and Rovelli.) 21 322 THE ANIMAL PARASITES OF MAN / mature segments are longer than wide (6 to 7 mm. by 2 to 3 mm.), fairly thick, are frequently of a reddish colour, and when cast off resemble cucumber seeds. The genital pores lie symmetrically at the lateral margins ; the roundish egg sacs, arising from the uterine reticulum, contain eight to fifteen eggs embedded in a reddish cement substance (in life). The eggs are globu- /.^'"vr— ^— ->>. lar (43 /t to 50 /t) ; the em- ^^ bryonal shell (embryophore) is ^^^ ^ thin, the oncosphere measures 32 /i to 36/1,. Surrounding the embryophore is an albumin- ous coating, and outside this the thin vitelline envelope (fig. 218). Dipyiidiuin caninuni is a common intestinal parasite of dogs, in which it grows larger {Tccnia ciicnmerlna, Bloch) than in cats {T. elliptical Batsch) ; it has, however, also been found in jackals, as well as in human beings, though in the latter it is of compara- tively rare occurrence (twenty- four cases), and almost always affects children, generally of tender age. One-third of all the cases in children were sucklings, about a quarter of all the cases recorded were adults, and these occurred throughout all Europe with the exception of Spain and Italy. The proglottids, which leave the intestine spontane- ously, are recognizable by the naked eye on account of their form and reddish colour, as As a rule, the presence of this parasite Fig. 221. — Larva (cyslicercoid) of Dipylidium caninum^ consisting of body and tail. The latter is solid and bears on it the embryonal spines. The bladder, which was only slightly developed, has disappeared, and the fore-part of the body bearing the rostellum is now seen invaginated into the hind portion. The booklets are shown in front of the excretory system which has now developed. At a further stage the tail drops off; the head now evaginates, but is still enclosed in a double-walled sac formed by the prolongation upwards on each side of the topmost parts of the body shown in the figure. Cf. fig. 220, 6. Enlarged. (After Grassi and Rovelli.) well as their two genital pores. sets up no marked symptom in the patient. The corresponding larval form (cysticercoid) lives in the louse of the dog {Tricho elect es canis), a fact that was first established by Melnikow and Leuckart ; according to Grassi and Rovelli, as well HYMENOLEPIDID^ 323 as Sonsino, it also lives in the flea of the dog (Ctenocephalus cants) and in the flea of man (Pulex irritans), but not in its larva. The adult segments, which also leave the rectum of dogs and cats spontaneously, creep about around the anus and get into the hair, and are thus partly dried and disintegrated. Part of the segments, or the oncospheres released by disintegration, are then taken up by lice and fleas, within which they develop into larvae (cysticercoids). Dogs and cats are thus infected by then- own skin parasites, which they bite and swallow whilst gnawing at their fur. The infection of human beings must occur in an analogous manner, by transmission of the cysticercoids present on the lips or tongue of dogs when the latter lick them, or it may be that the vermin of cats and dogs harbouring cysti- cercoids are accidentally and directly swallowed by human beings. Family. Hymenolepididae, Railliet and Henry, 1909. Genus. Hymenolepis,^ Weinland, 1858. Accessory sac (opening into genital atrium) usually absent. Vas deferens with an external (outside cirrus sac) and an internal (inside cirrus sac) "seminal vesicle." Three testes in each proglottis. The eggs are round or oval with two to four distinct envelopes. In mammals and birds. Hynnenolepis nana, v. Sieb., 1852. Syn. : Tcenia nana, v. Sieb., 1852, nfc van Beneden, 1867; Tcenia cegyptiaca, Bil., 1852 ; Diplacanthus-nanus, VVeinld., 1858 ; Tcettia {Hy?nenolepts) nana, Lkt., 1863. The worm is 10 to 45 mm. in length and 0*5 to 07 mm. in breadth ; the head is globular, 0*25 to 0*30 mm. in diameter. The rostellum has a single circlet consisting of twenty-four or twenty-eight to thirty hooks, which are only 14 /x to 18 yu, in length. The neck is moderately long; the proglottids are very narrow, up to 200 in number, 0*4 to o*9 mm. in breadth, and 0*014 *o 0*030 mm. in length. The eggs are globular or oval, 30 /^ to 37 /a to 48 /a ; the oncospheres measure 16 yLt to 19 /x in diameter, with two coats, separated by an intervening semi-fluid substance (fig. 224). This species was discovered by Bilharz in Cairo in 185 1 ; it was found by him in great numbers in the intestine of a boy who had ' The genus is by some authors divided into two sub-genera— Hymenolepis, s. str., and Drepanidotaenia, Raill. Drepanidotrsnia.— Body, broad lanceolate, testes three, female genitalia antiporal beside the testes. Scolex small, with eight hooks. Neck very short, longitudinal muscle bundles very numerous. No accessory sac opening into genital atrium. Hymenolepis.— ^d^xxovf , female genitalia ventral to or between testes. 324 THE ANIMAL PARASITES OF MAN died of meningitis. For several years this was the only case, until 1885, since when numerous cases have come to light. Spooner (1873) even reported a case from North America, which may, however, have related to Hynienolepis diminuta. In Europe the w^orm is particu- larly frequent in Sicily, but it has also been repeatedly observed in North Italy; it has, moreover, been reported from Russia, Servia, England, France, Germany, North and South America, the Philippines, Fig. 223. — Hymenolepis nana: head. Enlarged. (After Mettens.) Fig. 222. — Hy- menolepis nana, v. Sieb. About 12 i. (After Leuckart.) Fig. 224. — Hymenolepis nana : an egg. Highly mag- nified. (After Grassi.) Fig. 225. — Longitudinal section through the intes- tinal villus of a rat, with the larva (cysticercoid) of Hy- meno/epis mtirina. Magni-. fied. (After Grassi and Rovelli.) Siam and Japan, in all over 100 cases. Notwithstanding its small size this worm causes considerable disorders in its hosts — mostly children — as it sets up loss of appetite, diarrhoea, various nervous disturbances, and even epilepsy ; all these symptoms, how- ever, disappear after the expulsion of the parasites, w^hich are generally present in large numbers. The development as well as the manner of infection is still unknown ; Grassi is of opinion that Hymenolepis nana is indeed merely a variety of Hymenolepis mnrina, Duj., which lives in rats. HYMENOLEPIS NANA 325 According to Grassi direct development takes place with omission of the intermediate host, but with the retention of the larval stage ; that is to say, rats infect themselves directly with Hymenolepis iniirlna, by ingesting the mature segments or oncospheres of this species, from which subsequently the small larvae originate in the intestinal wall cqrtpar: ry?'/.n., ex.Cdn. :M: ov. Fig. 226. — Hymenolepis nana (murma) : cross section of proglottis from a rat. c.p., cirrus sac; rec. sem., receptaculum seminis ; s.^., shell gland; ov., ovary ;/., testis ; corL par., cortical parenchyma; m.l.n., main lateral nerve; ex. can., excretory canal; y.^., vitellarium. (After v. Linstow.) sec.cay^ ■'pr.cay. pr.ca\/. ':Caud. ¥iG.22'j.— Hymenolepis nana: longitudinal section of an embryo, bl.p., anterior opening of secondary cavity ; caud., caudal appendage ; pr. cav., primary cavity ; sec. cav., secondary cavity. Enlarged. (After Grassi and Rovelli.) (fig. 225); when fully developed they fall into the intestinal lumen and become tapeworms. The identity of the two forms has neverthe- less been disputed (Moniez, R. Blanchard, v. Linstow), though their near relationship cannot be denied. Grassi gave mature segments of Hymenolepis nturina to six persons, but only one person evacuated a tapeworm. This, however, proves nothing in a district where 326 THE ANIMAL PARASITES OF MAN Hymenolepis nana frequently occurs in man ; it was, moreover, not possible to infect rats with segments of Hymenolepis nana (of man). Accordingly this form may represent an independent species, which, however, like Hymenolepis murina, also omits an intermediate host. Hymenolepis diminuta, Rud., 1819. Syn. : TcEuia diminuta^ Rud., 1819 ; Tcenia leptocephala^ Crepl., 1825 ; TcEnia flavopU7ictata, Weinld., 1858 ; Tcenia varesina, E. Parona, 1884 ; Tcenia 7niftima, Grass!, 1886. This species measures 20 to 60 cm. in length, and up to 3*5 mm. in breadth ; there are from 600 to 1,000 segments. The head is very small (0*2 to o*5 mm.), it is club-shaped and has "'""^^^^^^ a rudimentary unarmed rostellum; the neck is short ; ^^ the mature segments are 3*5 mm. in breadth, 066 mm. { > I in length ; the eggs are round or oval. The outer ^ egg-shell is yellowish and thickened, with indistinct '^ radial stripes ; the inner embryonal shell (embryo- \ phore) double, thin ; the outer layer is somewhat pointed at the poles ; oncosphere 28 /x by 36 fju. Between the inner and outer shells is a middle granular layer. Hymenolepis diminuta lives in the intestine of rats — Mus dectimanus (the sewer rat), Mus ratins (the black rat), and Mus alexandrimis, rarely in mice ; it is occasionally also found in human beings. Weinland described it from specimens collected by Dr. E. Palmer in 1842, in Boston, from a child aged 19 months, as T. flavopunctata. A second case relating to a three year old child, from Philadelphia, was only reported in 1889 by Leidy ; a third case was simultaneously reported of a two year old girl in Varese (T. varesina) ; and Grassi described another case relating to a twelve year old girl from Catania (Sicily). Sonsino and Previtera reported Fig. 228. — Hy- menolepis diminuta ; scolex. Magnified. (After Zschokke.) P^IG. 229. — Hymenolepis diminuta : two pro glottids showing testes (3), ovary and vagina Slightly enlarged. (After Grassi.) Fig. 230. — Hymenolepis diminuld. egg from man. (After Bizzozero.) HYMENOLEPIS DIMINUTA 327 the same species in Italy, Zschokke in France, Lutz and Magalhaes in South America, and Packard in North America : a total of twelve cases, five from America, the rest from Europe (Ransom). According to Grassi and Rovelli the larval stage lives in a small moth (Asopia farinaiis), as well as in its larva, in an' orthopteron {Anisolabis anmilipes), and in coleoptera {Acis spinosa and Scaurus striatus). Experimental infections have been successful on rats as /, — -^ Fig. 231. — Hymenolepis diminuta: cyslicercoid from the rat ^Qdi{Ceraiophyllus fasciatus) a, remains of primary vesicle ; d, fibrous layer ; c, radially striated layer resembling cuticle ; d, layer of columnar cells ; e, parenchymatous layer of irregularly disposed cells ; f, parenchy- matous layer. (Stephens, after Nicjll and Minchin.) well as on human beings. In America other species of insects may be the intermediary hosts. Nicoll and Minchin^ found in the body cavity of 4 per cent, of rat fleas {Ceratophyllus fasciatus) the cysticercoid o ^Hymenolepis diminuta. That it belonged to this species was shown by its unarmed rostellum and by feeding; 340 fleas were fed to white rats and fourteen ' Proc. Zool. Soc, 191 1, p. 9. 328 THE ANIMAL PARASITES OF MAN worms obtained, i.e., about 4 per cent., thus corresponding to the infec- tion of the fleas. The development in the flea probably begins in the pupal stage, the eggs being ingested by the older flea larvae. The larva is o'3i by 0*25 mm.; tail o'8 mm., scolex 0*075 by 0*09 mm., suckers, 0*055 mm. in diameter. Microscopically it shows — (i) externally a radially striated layer resembling cuticle, (2) a layer of columnar cells, (3) parenchymatous layer continuous with the tail, (4) fibrous layer around the small caudal vesicle, then the parenchymatous scolex at the bottom of the secondary cavity. Nicoll and Minchin (loc. cit.) found a cysticercoid ^ in the rat flea Ceratophyllus fasciatus which w^as probably that oi Hyiuenolepis miirina. Body o-i6 mm., tail o'lg mm., scolex 0-096 mm. in diameter. Ros- tellum has twenty-three spines in a single row. Length 0*017 mm., handle o-oi mm., guard 0*007 i^tn., prong 0*007 mm. Sucker 0*042 mm. Although this cycle, then, for H. miirina also exists, it is not probable that rats (or man in the case of H. nana if this be considered distinct) infect themselves in this way, as they hardly ingest all the necessary fleas to account for the massive infection which frequently exists in rats (and man), so that Grassi's cycle holds good as the predominant method. Xenopsylla cheopis has also been found by Johnston to harbour both cysticercoids in Australia. Hymenolepis lanceolata, Bloch, 1782. Syn. : To'm'a lanceolata, Bloch, 1782; DrepanidotcEnia lanceolata, Railliet, 1892. The parasite measures 30 to 130 mm. in ^~; length and 5 to 18 mm. in breadth; the head is globular and very small ; the rec. 33/77. Fig. 232. — Hymenolepis lanceolata. Natural size. (After Goeze.) To the right above, two hooks. 120/1. (After Krabbe.) Fig. 233. — Hymenolepis lanceolata : diagram of female genitalia, ov., ovary; ovd., oviduct; rec. sem., receptaculum seminis ; s.g., shell gland; ut., uterus; y.g., vitellarium. (After Wolflhiigel.) 'A third cysticercoid resembling this, but without hooks, has also been found. DAVAINEID^ 329 rostellum is cylindrical, with a circlet composed of eight hooks (31 /x to 35 yit in length). The neck is very short. The short segments increase gradually and equally in breadth, but only a little in length ; the female glands lie on the side opposite to that on which the genital pore is situated; the three elliptical testes are on the same side as the pores ; the cirrus is armed and slender. The eggs have three envelopes and are oval (50 /^ by 35 ytt), the external envelope is thin, the middle intermediate layer or envelope is not so marked as in H. dijjiiniita, and the internal one is very thin and sometimes has polar papillae, as in Hymenolepis diminuta and H. nana. It inhabits the intestine of the following birds : Domesticated ducks and geese, the Muscovy duck (Cairlna moschata), white-headed duck {Erisinatura leticocephala), the pochard (Nyroca nifina), and the flamingo (Phcenicopteriis antiqnonim). It has been recorded from Great Britain, France, Denmark, Austria and Germany. Zschokke reports the receipt of two specimens which a twelve year old boy in Breslau evacuated spontaneously at two different ti mes. The corresponding larva, according to Mrazek, lives in fresh water Cyclops; according to Dadai it is likewise found in another copepod, Diaptomus spinosus, but the hooks of Dadai's larva differed in size. Family. Davaineidae, Fuhrmann, 1907. Sub-family. Davaineinae, Braun, 1900. Genus. Davainea, R. Blanch., 1891. The large scolex is more or less globular, much wider than the rostellum, which is furnished with two rings of very small and numerous hooks. Neck absent, pro- gloltids few, genitalia single. Parasitic chiefly in birds.' Davainea madagascariensis, Davaine, 1869. Syn. : T<^nia madagascariensis^ Dav. ; Tcznia demerariensis^ Daniels, 1895. This worm measures 25 to 30 cm. in length; the head has four large round suckers ; the rostellum has ninety hooks (18 /m in length) ; there are 500 to 700 segments, of which the last 100 are filled with eggs and form half of the entire worm. The segments, when mature, measure 2 mm. in length by 1*4 mm. in breadth; genital pores unilateral; about fifty testes ; the uterus consists of a number of ' [The laival stage of the Davaineas occurs in slugs [Limax] and snails {Helix). ~Y. V. T.] 330 THE ANIMAL PARASITES OF MAN loops, which at each side are rolled up into an almost spherical ball ; when filled with eggs the convolutions unwind, permeate the segment and then lose their wall ; the eggs lying free in the parenchyma become finally surrounded, one, or several together, by proliferating parenchymatous cells ; this is how the 300 to 400 egg masses, taking up the entire mature segment, are formed. The globular oncosphere (8 fi) is surrounded by two perfectly transparent shells, the outer of which terminates in two pointed processes. Davainea madagascaricnsis has hitherto been found in man only (eight times). Davaine described this species from fragments sent to him from Mayotta (Comoro Islands), which were found in two Creole children. Chevreau observed four cases in Port Louis (Mauritius), likewise in children ; Leuckart received the first perfect specimen — it was obtained from a three year old boy, the son of a Danish captain, in Bangkok ; Daniels, at the post-mortem of an adult native of George Town, Guiana, found two specimens {Tcenia dernerariensis) ; and finally Blanchard describes another perfect specimen which was in Davaine's collection of helminthes in Paris, and which was obtained from a little girl 3 years old, of Nossi-Be (Madagascar). The inter- mediate host is unknown. Davainea (?) asiatica, v. Linst., 1901. Syn. : Tcenia asiatica, v. Linstow. There exists only one headless specimen of this species, which is not quite adult, and which is pre- served in the Zoological Museum of the Imperial Academy of Science in Petrograd. It came from a human being and was found by Anger in Aschabad (Asiatic Russia, near the northern frontier of Persia). The specimen measures 298 mm. in length. The breadth anteriorly is only o*i6 mm., the posterior part measures 178 mm. across. The number of segments is about 750. The genita pores are unilateral ; the testes are globular and lie in a dorsal and ventral layer in the medullary layer; the cirrus pouch is pyriform, 0*079 mm. in length and 0*049 mm. in breadth ; the female glands lie in the fore-part of the segments, the ovary reaching to the excretory vessels ; the vitellarium is small and round. The vagina has a large fusiform receptaculum seminis ; the uterus breaks up into sixty to seventy large, irregularly polyhedric eggsacs. Fig. 234. — Scolex of Davainea niada- gascariensis. The hooks have fallen off. 14/1. (After Blanchard.) T^NIID^ 331 Family. Taeniidae, Ludwig, 1886. Genus. Taenia, L., 1758.^ With the characters of the family. In the genus Cladotaenia recognized by some authors, the testes encroach on the mid field and the uterine stem reaches the anterior end of the segment. Taenia solium, L., p. p., 1767. vSyn. : Tcenia cucurbitina^ Pall., 1781 ; Tcenta pelluctda, Goeze, 1782; Tcenia vulgaris^ Werner, 1782; Tcenia dentata, Gmel., 1790; Haly sis solium^ Zeder, 1800; Tcenia humana arrnata^ Brera, 1802; Tcenia {Cystotcenid) solium, Lkt., 1862. The average lengtH of the entire tapeworm is about 2 to 3 m., but may be even more ; the head is globular, o-6 to o'8 to i*o mm. in diameter. The rostellum is short with a double circlet of hooks, twenty-two to thirty-two in number, usually twenty-six to twenty-eight ; large and small hooks alternate regularly; the length of the large hooks is o'i6 to 0'i8 mm., of the small ones O'li to 0*14 mm. The ros- tellum is sometimes pigmented. The suckers are hemispherical, 0*4 to o'5 mm. in diameter. The neck is fairly thin and long (5 to 10 mm.). The proglottids, the number of which averages from 800 to 900, increase in size very gradually ; at about i m. behind the head they are square and have the genitalia fully developed. Segments sufficiently mature for detachment measure 10 to 12 mm. in length by 5 to 6 mm. in breadth. The genital pores alternate fairly evenly at the lateral margin a little behind the middle of each segment. The fully developed uterus con- sists of a median trunk, with seven to ten lateral branches at either side, some of which are again ramified. The eggs are oval, the egg-shell very thin and delicate ; the embryonal shell (embryophore) is thick, with radial stripes ; it is of a pale yellowish colour, globular, and measures 31 //, to 36 /a in diameter ; the oncospheres, with six hooks, are likewise globular, and measure 20 /^ in diameter (fig. 238). ' The Greeks termed the tapeworms eXfiivOes irXarfTai, more rarely x'/P*« ( = fascia) ; the Romans called them tienia, tinea, taniolay later lumbrici, usually with the addition laii, to distinguish them from the Luf?ibrici teretes (Ascarida). The proglottids were called Vermes cucurbitini ; the cysticerci xa^«C«' (hailstones), later hydatids. Plater (1602) was the first to differentiate Tcenia intestinorum { = Bothriocephalus latus) amongst the Lumbrici lati of man from Tcenia longissima (= Tcenia solium). The term solitwi was already used by Arnoldus Villanovanus, who lived, about 1300 ; and, according to him, it signifies "cingulum" (belt, chain), while N. Andry, in 1700, traces this word from "solus," because the worm occurs always singly in man. Leuckart and Krehl derive the word " solium " from the Syrian "schuschl" (the chain), which in Arabian has become *' susl" or *' sosl," and in Latin has become "sol-ium." What Linnaeus described under the term Tcenia solium was really Tcenia saginata ; the latter was first distinguished by Goeze, but was forgotten until Kuchen- meister, in 1852, again called attention to the differences. 332 THE ANIMAL PARASITES OF MAN Malformations are not so common as in T. saginaiaj they consist in two or several proglottids being partly or entirely fused, formation of single club-shaped segments, fenestration of long or short series of segments and so-called double forma- tion, in which the head has six suckers and the segments exhibit a Y-shaped trans- verse section. The oncospheres occasionally also possess more than six booklets. Very slender specimens have led to the description of a particular species or variety {T. tenella). In its fully developed condition T, solium is found exclusively in man ; the head is usually attached in the anterior third of the small intestine and the chain, in numerous convolutions, extends backwards ; a few mature detached proglottids usually lie at the most posterior part, and these are usually evacuated during defaeca- tion. In exceptional cases single proglottids or whole worms may reach contiguous organs if abnormal communications with them Fig. 235. — Two fairly mature proglottids of Tama solium, showing ovaries (one bi-lobed), vitellaria, central uterine stem, cirrus and vas deferens (above), vagina (below), testes (scattered), longitudinal and transverse excretory vessels. Fig. 236. — Head of Tania solium. 4S/I. exist; thus they may reach the abdominal cavity and the urinary bladder, or they may be found in a so-called worm abscess of the peritoneum ; occasionally, in vomiting, single segments or several together may be brought up. Exceptionally it induces severe anaemia. The larval stage (Cysticerciis celhilosce) that gives rise to Tcenia solium lives normally in the intramuscular connective tissue and other organs of the domestic pig, but it is known to exist also in a few other mammals, such as the wild boar, the sheep/ the stag, dog, cat, brown bear and monkey, as well as in man. The cysticercus ' The larvae which on rare occasions are found in the muscular system of sheep are either strayed specimens of Cysticercus tenuicollis^ which normally develop in organs of the abdominal cavity, and appertain to Ti^nia marginata of the dog, or actually Cysticercus yl. canin. Unde- termined Parona (Milan) 1899 150 121 II 4 — 14 Parona (Italy) 1868-99 513 397 71 26 — 19 Krabbe (Denmark) 1869 100 37 53 9 I »» »» 1869—86 200 '53 24 16 8 — >» »> 1887-95 100 89 — 5 6 — j> »» 1896— 1904 50 41 I S 3 — Blanchard (Paris) 1895 ? 1,000 21 — Stiles (United Slates) ... 1895 ( more than 1 300 19 more than) 30c 1 16 — 3 — — Schoch (Switzerland) 1869 I 2 — — Zaeslein (Switzerland) ... 1881 ? 180 19 ? — — Kessler (Petrograd) 1888 ? 22 16 47 Mosler (Greifswaid) 1894 181 112 64 5 — — Bollinger (Munich) 1885 25 16 8 Vieroidt (Tubingen) 1885 121 "3 8 — — — Mangold (Tubingen) 1885-94 128 120 6 8 — — Taenia africana, v. Linst., 1900. This worm measures over i'3 m. in length. The segments are all broader than they are long. The scolex is unarmed and is provided with an apical sucker (o*i6 mm.). The parasite measures 1-38 mm. in breadth, i'03 mm. in width ; the suckers measure 0*63 mm. in diameter. The neck is very short and somewhat broader Fig. 245.— Mature segment of Tcenia africana. The ovary is in the middle, and behind it are the shell gland and vitellarium ; at the sides are the testicles, and externally the excretory canals; the cirrus pouch, the vas deferens and the vagina are on the left. Magnified. (After v. Linstow.) than the scolex; number of segments about 600; the hindmost segments measure 7 mm. in length and 12 to 15 mm. in breadth. The genital pores alternate irregularly in the middle of the lateral margin ; the testes are very numerous and occupy the entire medul- lary layer; the vas deferens is much convoluted; the cirrus pouch T^NIA CONFUSA 343 is pyriform and thick walled ; the cirrus and vagina are beset with bristles directed outwards ; the receptaculum seminis is fusiform ; the ovary is large and double, and consists of radially placed club-shaped tubes that do not anastomose and do not branch : the vitellarium is Fig. 246. — Proglottis of Tcenia africana, wiih uterus. Magnified. (After V. Linstow.) at the posterior border of the proglottids, the round shell gland in front of it ; the uterus consists of a median trunk and fifteen to twenty-four non-ramified lateral branches on each side ; the embryonal shell is thick and has radial stripes — it may be round (31-2//, to SS'B//-) or oval (39/x by 33*8/^); the spines of the oncospheres measure 7 yu, to 8 yn in length (fig. 197). At present only two specimens are known ; they came from a black soldier from the vicinity of Lake Nyasa. The cysticercus is unknown ; perhaps it lives rnlf: ?n5;:;^.nl5 ^^!\t£l' i" thc zcbu, thc fiesh of which thc Askaris t to 36 /x in diameter. When mature this parasite lives in the small intestine of the domestic dog, the jackal, and the wolf, and apparently also in Felis concolor, and is usually present in great numbers ; it can also be transmitted ex- perimentally to the domestic cat, one suc- cessful result out of seven (Deve).^ The larval stage (Echinococcus polymorphus) lives in various organs — chiefly in the liver and lungs — of numerous species of mammals (twenty-seven), especially in sheep, ox and pig, and it is even not uncommon in man, though the Taenia itself has never been found in a human being; accordinglyman can only acquire the echinococcus by ingesting the eggs of the " dog worm." The dogs dis- seminate the eggs of Tcenia echinococcus wherever they go, or carry them to their mouths and coats by biting up the evacuated segments, and are thus able to transmit them directly to human beings (by licking them or making use of the same crockery, etc.). In other cases the onco- spheres, enclosed in the embryophores, must withstand desiccation for a time and then (as when the dogs are '' kissed " or other- wise caressed) are transmitted into or on to 50/1. Fig. 250. — Tcenia echino- coccus : the cirrus sac, the vagina, uterus, ovary, shell gland and vitellarium, and the testicles at the sides are recognizable in the second proglottis ; the pos- terior proglottis shows the uterus partly filled with eggs, as well as the cirrus sac and the vagina. ' In Iceland 28 per cent, of the dogs are infected with this Taenia, in Lyons 71 per cent., in Zurich 3*9 per cent., in Berlin i per cent., and in Copenhagen 0*4 per cent. In Australia even 40 to 50 per cent, of the dogs are affected. It is, however, a question whether, in addi- tion to TcBnia echinococcus, a second analogous form is not involved, as the form from Canis dingo attains a length of 10 to 30 mm. 346 THE ANIMAL PARASITES OF MAN man. As echinococcus disease in man is always very dangerous, it would be a matter of general interest to prevent dogs being infected by destroying the echinococci,^ and all measures would be justifiable which would diminish the superfluous number of house-dogs (for instance, high taxes) ; measures should also be adopted to limit the association of men with dogs, particularly in such frequented places as restaurants, railway carriages and tram-cars. Echinococcus is very common in slaughtered animals ; in Germany, however, the figures in the reports of the abattoirs present an erroneous view in so far as, besides the total number of animals slaughtered, only the numbers of those organs (liver and lungs) are published that were so severely infected with echinococci that, even when the parasites were " shelled " out, the flesh could not be placed upon the market and was therefore "condemned." In Berlin the following animals were slaughtered : — Year 1889—90 1890—91 1891—92 1892-93 1896-97 1902 Oxen ... Sheep... Pigs ... 154,218 430,362 442,115 124,593 37i,Q43 472,859 136,368 367,933 530,551 142,874 355,949 518,073 146,612 395,769 694,170 153,748 434,155 778,538 During the same years the following were condemned in consequence of being infected with echinococci : — Lung Liver Lung Liver Lung Liver Lung Liver Lung Liver Lung Liver Oxen ... Sheep... Pigs ... 7,266 5,479 6,523 2,418 2,742 5,078 5,792 4,595 5,083 1,938 2,059 3,735 4,497 4,435 6,037 1,721 1,669 4,374 2,563 3,33^ 6,785 739 1,161 4,312 3,284 4,561 7,888 1,156 1,939 5,398 2,507 11,138 9,544 791 4,437 9,233 Nevertheless there are statistics that give the total number of animals infected with echinococcus : — Author Place Oxen Sheep Pigs Langrich ... Olt... Steuding ... Prettner Rostock i. M. Stettin Gotha Prague 26- 2 per cent. 7-1 24'6 23-2 37-0 per cent. 258 35*4 5"5 5-4 per cent. 7-3 21-4 ? In Giistrow, in Mecklenburg, half of the animals slaughtered are said to be infected with echinococcus; in Wismar 25 per cent, of the oxen, 15 per cent, of the sheep and 5 per cent, of the pigs are infected ; according to Mayer, in Leipzig, 379 per cent, native pigs, 24*47 per cent. Hungarian pigs, and I3'09 per cent, of sheep were infected with echinococcus ; at the same time it was stated that in regard to the native pigs the liver was more frequently affected than the lungs (s'Si per cent. as compared with 0*26 per cent.) ; in sheep the lungs were more frequently infected ' Mosler, F., '* Ueb. Mittel z. Bekampfg. endem. vork. Echinococcuskrank. , vud. Zeit., 1889, No. 72. Deutsch. STRUCTURE AND DEVELOPMENT OF ECHINOCOCCUS 347 (1271 per cent, to 373 per cent.)> whereas in the Hungarian pigs both organs were almost equally infected (1478 per cent, to I2'03 per cent.). The data of Lichtenheld, in Leipzig, give the frequency with which various organs were affected, as shown in the following table : — Lungs ... Liver ... Spleen Heart ... Kidneys Subperitoneal tissue Cattle 69*3 per cent 27-0 ,, 2-2 „ 075 .. 075 .. Pigs i6*2 per cent. 74*2 ,, 3*2 „ 3'2 ,, 3-2 „ 21*4 percent, 72-0 „ 27 M 1-3 M 1-3 » i'3 >. Sheep 52*2 per cent. 44*9 ,, 2*9 », Horses .S'5 percent. 94*5 .. Structure and Development of Echinococcus (Hydatid). An echinococcus is a spherical or roundish bladder full of a watery liquid, which originates by liquefaction of the oncosphere, and in man may attain the size of a child's head, but remains smaller in Fig. 251. — Echinococcus veterinorwn: the fibrous sac enclosing the echinococcus has been opened and laid back in five parts, so that the surface of the bladder worm may be seen, with the brood capsules, visible to the naked eye, showing through it. Natural size. (After Leuckart.) cattle (the size of an orange or apple). The thin wall of the bladder is composed of an external laminated cuticle (ectocyst) and an internal germinal or parenchymatous layer (endocyst). The latter again 348 THE ANIMAL PARASITES OF MAN exhibits two layers : an outer layer of small cells that are less sharply defined, and an inner layer of larger cells. It contains, in addition, calcareous corpuscles, muscular fibres and excretory vessels. It is rich in glycogen. ^^^^^^Ig ^^Ml^^ Fig. 252. Figs. 252 and 252A. — Diagrams of mode of formation of brood capsule and scolices. (i) Wall of mother cyst, consisting of ectocyst and endocy.>,t ; (2) theoretical stage of invagination of wall; (3) a brood capsule with the layers of the wall in the reverse position to that in the mother cyst ; {4) evagination of wall ; (5) invagination ; (6) fusion to form the solid scolex ; (7) invagination of fore-part of scolex into hind-part. [Note. — The size of the scolex is much out of proportion to the brood capsule.) (Stephens.) STRUCTURE AND DEVELOPMENT OF ECHINOCOCCUS 349 The development in cattle often remains stationary at the bladder stage, and they are then called "acephalocysts," or Echinococciis cysticus sterilis. According to Lichtenheld, sterile cysts occur in 80 per cent, of cases in cattle, in 20 per cent, in pigs, and in 7-5 per cent, in sheep. In other cases large numbers of small, hollow BROOD capsules are formed in the germ layer, but are not arranged in any particular order. The order of the layers is just the reverse in them to what it is in the parent cyst, that is to say, they have inside a thin non-laminated cuticle and the parenchymatous layer on their external surface. These, theoretically at least, may be regarded as invaginations of the bladder wall giving rise to a cavity with the cuticle internal and the parenchymatous layer external. If we should then get an isolated cavity with cuticle internal and parenchymatous layer external, as in the brood capsule (fig. 252). we suppose the orifice to close, in ilMlill iJiiiiiiyiiiiii'iiii^iiiiiilB Fig. 252A. If we next suppose an evagination of the wall of the brood capsule to occur at one point we should get a hollow process lined with cuticle ; at the bottom of this we get the scolex and booklets formed, and a little higher up the tube the suckers (fig. 252, 4). If this hollow scolex is now pictured as being invaginated we get a hollow scolex covered with cuticle and lined by a parenchymatous layer projecting into the cavity of the brood capsule. The two sides of this hollow scolex now fuse and we get a solid scolex projecting into the cavity. Finally, if we imagine once more the rostellum and suckers invaginated into the posterior part of the scolex we get tlie condition as frequently found in the brood capsules,/.^., a scolex covered with cuticle projecting into the cavity, with the rostellum and suckers invaginated into the posterior portion of the scolex (fig. 252A, 7). 350 THE ANIMAL PARASITES OF MAN A large hydatid may contain many thousands of brood capsules. Each brood capsule is about as big as a pin's head, and may contain ten to thirty or more scolices. The delicate wall of the brood capsules may rupture, so that the scolices are now free in the mother cyst. These free scolices and also free brood capsules constitute what is known as " hydatid sand," which settles at the bottom of a glass when hydatid fluid is poured into it. This form occurs chiefly in domesticated animals and is termed E. veierinortuu, Rud., or E. cysticus fertilis. In man, and only rarely in cattle, the mother cyst first forms ^' daughter cysts" {E. honiinis, Rud. [fig. 255]), which, though smaller than the " mother cyst," re- semble it in the structure of their walls ; thus they are covered externally by a lami- nated cuticle and internally by the parenchymatous layer. They originate : Fig. 253. — Section through an invaginated echinococcus scolex. Cf. fig. 252A, 7. X 300. (After Deve.) Fig. 254. — A piece of the wall of an Echinococcus veterinorum stretched out and seen from the internal surface. A few brood capsules (the outline of which is only faintly shown), with scolices directed towards their interior and exterior. 50/1. (i) Between the laminae of the cuticle of the mother cyst from small, detached portions of the parenchymatous layer ; during their growth they bulge inwardly or outwardly and may separate them- selves entirely from their parent cyst. In the latter case they lie between the mother cyst and the capsule of connective tissue formed by the host (E. granulosus or E. hydatidosiis exogeniis) ; when growing inwardly they reach the interior of the mother cyst (£. hydatidosiis endogenus). Their number is very variable and does not depend on the size of the mother cyst. They are as big as, or bigger than, gooseberries. (2) According to some authors, endogenous daughter cysts arise also STRUCTURE AND DEVELOPMENT OF ECHINOCOCCUS 351 from a metamorphosis of scolices that have separated off from the brood capsule. This takes place in the following way : Fluid accumulates in the interior of the scolex, so that eventually nothing remains except a sac consisting of cuticle lined by parenchyma. The cuticle gradually thickens and several layers form (tig. 257). Fig. 255* — Echinococcus hoviinis in the liver. The fibrous capsule and the wall of the echinococcus have been incised, so that the endogenous daughter cysts may be seen. Reduced. (After Oslertag, from Thoma.) (3) Transformation of Brood Capsules into Daughter Cysts. — This is also held to be possible by various observers. New epithelial layers are deposited between the cuticle which lines the brood capsule and the outer parenchymatous layer. This parenchymatous layer gradually disappears and a newparenchymatous layerforms in the interior from the paren- chyma of the scolex or scolices. Although it appears strange that a completely formed scolex with specifically differentiated tissues and organs should retrogress to more primitively organized matter, and again become a proliferating bladder, yet we can hardly doubt that the older observations, regarding such a vesicular metamorphosis, of Bremser (1819), v. Siebold (1837), Naunyn (1862), Rasmusser (1866), Leuckart (1881), Alexinsky (1898), Riemann (1899), Deve (1901), and Perroncito (1902) are correct. Fig. 256. — Section through an echinococcus scolex in process of vesicular metamorphosis, twenty-six days after insertion in the pleural cavity, x 250. (After Deve.) 352 THE ANIMAL PARASITES OF MAN (4) Further, a fourth method of formation of daughter cysts is described by Naunyn as occurring in sterile hydatids, i.e., those con- taining no brood capsules. In this case a portion of the mother wall of the hydatid gets constricted off. Fig. 257. Ficis. 257 and 257A. — Diagram of transformation of a scolex into a daughter cyst (i to 3) : I, scolex in brood capsule ; 2, liquefaction of scolex ; 3, daughter cyst ; and (4 to 6) of a broou capsule into a daughter cyst : 4, brood capsule with scolex ; 5, deposition of new epithelial layers on the inner layer of the parenchyma ; b, disappearance of outer parenchyma and formation of inner parenchyma from the parenchyma of scolex, which has now disappeared. {Note. — The scolices are out of proportion to the brood capsules and to the daughter cysts. (Stephens.) It has also been established that not only daughter cysts transplanted into animals develop further (Lebedeff, Andrejew, Stadnitzky, Alexinsky, Riemann), but that this also holds good if only hydatid scolices from STRUCTURE AND DEVELOPMENT OF ECHINOCOCCUS 353 man or animals are transplanted into animals (rabbits). They develop into echinococci and can then give rise to brood capsules and scolices. As Deve further established, hydatid scolices are not capable of developing in guinea-pigs, while corresponding experiments with rabbits are in the large majority of cases successful where the scolices are introduced subcutaneously or into the pleural or peritoneal cavities. It is only in the case of daughter cysts that further growth is obtained in the case of guinea-pigs. Finally it appears, as has been already stated, that brood capsules can transform themselves into daughter cysts, but according to Deve only within the mother cyst, not aftei transplantation. Daughter cysts that have been formed in the mother cyst of man and animals behave themselves just as the mother cyst does, i.e., they can remain sterile, r™ or give rise to brood capsules l|lL and scolices, or even again ^ to fresh cysts — granddaughter Fig. 257A. cysts. The mother cyst can also die, so that the daughter cysts then lie in the cavity of the connective tissue capsule. The number of the daughter cysts in either case may attain several thousands. The echinococcus fluid, which originally is formed from the blood of the host, is light yellow, with a neutral or slightly acid reaction ; its specific gravity averages 1009 to 1015. It contains about I'S per cent, of inorganic salts, half of which is common salt ; in addition (besides water) it contains sugar, inosite, leucine, tyrosin, succinic acid (associated with lime or soda) and albumens which are not coagulated by heat ; occasionally also the fluid has been found to contain haematoidin and uric acid salts (in echinococcus of the kidneys), which doubtless demonstrates that the echinococcus liciuid originates from the host. It has been generally assumed that echinococcus fluid contains a toxic substance the escape of which into the body cavity (at operation or by bursting of a hydatid cyst) produces more or less severe symptoms (fever, peritonitis, urticaria), so much so that one speaks of hydatid intoxication. The 23 . 354 THE ANIMAL PARASITES OF MAN investigations of Robert, Joest, etc., have, however, shown the harmlessness of fresh undecomposed hydatid and cysticercus fluid for rabbits, mice and guinea-pigs, whether inoculated intraperitoneally, subcutaneously or intravenously. Contrary data or clinical experience must accordingly depend on other factors. According to the researches of Leuckart, the growth of the echinococcus is very slow ; four weeks after infection the average size is only 0*25 to 0-35 mm., at the age of eight weeks it is i to 2-5 mm., and at this period the formation of the central cavity com- mences ; at the age of five months, and with a size of 15 to 20 mm., the first brood capsules with scolices are formed. The conse- quence of this gradual increase of size is that the organ attacked can maintain its functions by vicarious hypertrophy, and that many echinococci induce no special symptoms and cannot even be diagnosed, the latter circumstance being due to their hidden position. The echinococcus cannot be said to be scarce in man, as is shown by the following table for Central Europe : — Place Period No. of post-mortems No. of cases of echino. Percentage Rostock 1861—83 1,026 25 2-43 Greifswald 1862-93 3.429 51 1-48 Jena 1866—87 4,998 42 0-84 Breslau 1866—76 5,128 39 0761 Berlin 1859-68 4,770 33 0-69 Wurzburg — 2,280 II 0-48 Gottingen — 639 3 0-469 Dresden 1852—62 1,939 7 036 Munich 1854-87 I4,>83 35 0-2S Vienna i860 1,229 3 024 Prague — 1,287 3 023 Kiel 1872—87 3,581 7 0-19 Ziirich, Basle, Berne — 7,982 II 013 Erlangen 1862—73 1,755 2 o-ii These, however, are only cases that have become known by post- mortem ; in addition, there are cases that have been treated medically, of which there are a few statements, at all events relating to the principal districts of Germany. According to Madelung, one case of echinococcus occurs in every 1,056 inhabitants in the town of Rostock, in the district of Rostock one to every 1,283, in Schwerin one to every 5,887, and in Ludwigsort one to every 23,685 ; according to Peiper, in Upper Pomerania one case occurs to every 3,336, in the district of Greifswald one to every 1,535 inhabitants. The northern districts of Pomerania are more affected than the southern ones. Accordingly, echinococcus is also considerably more frequent in cattle in Pomerania. On an average in Germany io"39 per cent, oxen, 9-83 per cent, sheep, and 6" 47 per cent, pigs are infected, whereas in Upper Pomerania 3773 per cent. STRUCTURE AND DEVELOl^MENT OF ECHINOCOCCUS 355 oxen, 27-1 per cent, sheep, and I2'8 per cent, pigs are infected ; in Greifswald, indeed, 64*58 per cent, oxen, 51 '02 per cent, sheep, but only 4*93 per cent, pigs are infected. In accordance with these figures Tcc?iia echmococcus must be frequent in dogs in Ponierania, especially in Upper Pomerania ; on the other hand, the conjecture that the frequency of echinococcus in Mecklenburg is explained by the occurrence of TcB7ua echinococcus in foxes has not been confirmed, as the fox does not harbour this worm in Mecklenburg. Beyond the European continent, echinococcus is frequent in the inhabitants of Iceland, Argentine, Paraguay and Austraha. In Iceland, according to Finsen, i in every 43 inhabitants is affected with echinococcus ; according to Jonassen the proportion is i to 63 ; this is due to the habits of the people of Iceland or, in fact, to the frequency of Tcviiia echino- cocciis in dogs, and the prevalence of the hydatid in cattle. In cer- tain districts of Australia it is just as frequent. In Cape Colony, Egypt and Algeria echinococcus is not rare, but it is scarce in America and in Asia, with the exception of the nomadic tribes of Lake Baikal. Echinococcus attacks persons of every age, though it is rare in children up to 10 years of age and in old people. It occurs most frequently between the ages of 21 and 40 years. According to all statistics it preponderates in women (about two-thirds of the cases). The liver is its favourite seat (57-1 per cent, of the cases) ; next in order come the lungs (8 per cent.), kidneys (6 per cent.), cranial cavity, genitalia, organs of circulation, spleen (3*8 per cent.), etc. As a rule one organ only is invaded ; multiple occurrence may originate from one infection, or eventually from a later infection (?), or it may come to pass that from some cause (through the spontaneous rupture of an echinococcus, or the rupture of one caused by an injury or surgical operation) daughter cysts, brood capsules or scolices escape into the abdominal cavity,^ where they settle or become transformed and go Fig. 258. — Hooklets of echinococcus. a, of Echinococcus veterinorum ; b, of Tania echino- coccus^ three weeks after infection ; c, of the adult Tcenia echinococcus ; d, the three forms of hooklets outlined one within the other. 600; I. (After Leuckart.) ' In such cases the toxic effects of the echinococcus fluid usually — if not always — manifest themselves. Such effects are manifested by severe symptoms of poisoning being set up, by urticaria, peritonitis, and ascites, and not infrequently they cause a fatal termination. 356 THE ANIMAL PARASITES OF MAN on growing. In the distribution of this secondary echinococcus the great powers of motiHty of the free scoHces must be taken into account (Sabrazes, Muratet, and Husnot). Human echinococci may also die at various stages of development, become caseous or calcified, or may be absorbed, the cause for this being either disease of the hydatid itself or inflammation of its connective tissue capsule; the discovery of the laminated cuticle, which has great powers of resistance, or the finding of the hooklets of the scolices is sufficient to form a conclusion as to the nature of such formations. Siebold (1853) was the first to rear Tcenia echinococcus in the dog by feeding it with the echinococcus of cattle and especially of sheep. Kuchenmeister, van Beneden, Leuckart, Railliet and others obtained similar results, and Thomas, Naunyn, Krabbe and Finsen succeeded in rearing T. cc/iiiiococciis in dogs from the bladder worms of human beings ; these grow comparatively slowly (one to three months^) and only during the process of growth develop their hooklets in their definite form (fig. 258). It lies in the nature of things that dogs, whether experimentally or naturally infected, almost always harbour T. echinococcus in large quantities. That cats exceptionally harbour these worms has been already mentioned (Deve). Finally, Leuckart infected young pigs by feeding them with mature segments. Echinococcus multilocularis (alveolar colloid). In addition to the form of echinococcus already described, and which is also frequently termed Echinococcus nnilocnlaris, there is a second form which occurs in man as well as in animals, and which is termed E. ninUilocnlaris, s. alveolavis (alveolar colloid). It was originally regarded as a tumour ; its animal nature was first established by Zeller and R. Virchow. The parasite, which varies in size from that of a fist to a child's head, presents a collection of numerous cysts, measuring between o'l and 3 to 4 mm. to 5 mm. in diameter, which are embedded at first in a soft, connective tissue stroma ; the cut surface has therefore a honeycomb appearance. The cysts are surrounded by a pellucid and laminated cuticle, and each according to its size encloses either a small-celled tissue or a cavity lined by a parenchymatous layer ; the fluid contained in such a cavity may be transparent, or is rendered opaque by globules of fat, bile- pigment, haematoidin and fat crystals. According to some authors all or most of these cysts intercommunicate ; others state that this is the case at least as regards the cuticle. The scolices are by no means ' According to Perroncito the scolices had not formed proglottids nine days after feeding, but the latter were present twenty-four days after feeding, although the formation of eggs had not begun. ECHINOCOCCUS MULTILOCULARIS 357 found in all the cysts, and when present only a few, rarely half, of the cysts contain scolices (one or more) ; it is supposed that at least some of these scolices are formed in brood capsules, and that the former are capable of undergoing a cystic metamorphosis. One circumstance is peculiar to the multilocular echinococcus of man, namely, the disintegration that sets in at certain stages ; in the centre of the parasite a cavity forms that frequently becomes very large and is filled with a purulent or brownish or brownish-green viscid fluid ; in this fluid one finds shreds of the wall of the cavity, calcareous bodies, echinococcus cysts, also scolices and booklets, as well as fat globules and crystals of haematoidin, margarine and cholesterin and concretions of lime. Such ulcerative processes, according to Ostertag, are never present in the multilocular echino- coccus of oxen,^ in which the separate cysts are larger and the connective tissue integument less powerfully developed. Fig. 259. — Echinococcus multilocularis in the liver of the ox. Natural size. (After Ostertag.) Hardly anything positive is know^n with regard to the development of the alveolar echinococcus ; its peculiar conformation is attributed by some to enormous infection of oncospheres, by others to the abnormal situation of one oncosphere ; a few authors ascribe it to infection of lymphatic vessels, others to infection of the biliary ducts or to peculiarities of the surrounding hepatic tissue ; Leuckart ascribes it to a grape-like variety of form which continues budding ; a few more recent authors consider multilocular echinococcus to be specific- ally different from unilocular echinococcus, and therefore also different the species of Taenia arising from them. Melnikow- Raswedenkow is also of this opinion. According to this author the oncospheres infect the lumen of a branch of the portal vein in 1 This may perhaps be explained by the fact that the hosts are slaughtered before the parasites have attained the size or other conditions necessary to disintegration. 358 THE ANIMAL PARASITES OF MAN Glisson's capsule of the liver and grow into an irregularly sliaped formation (chitinous coil), which breaks through the vascular walls and thus forms the alveoli. So far the data coincide well with Leuckart's opinion of the original grape-like form of the Echiiiococcns multilocularis ; according to Melnikow-Raswedenkow the *' granular protoplasmic substance" (parenchymatous layer) is not only present in the interior of the loculi but also outside, and, moreover, "ovoid embryos " are supposed to develop in the chitinous coils, which, " thanks to their amoeboid movements, reach the lumen of a vessel, where, under favourable circumstances, they begin to develop further," that is to say, they become '' chitinous cysts with fantastic outlines," or also ^' single-chambered chitinous cysts " ; scolices may develop in both. Deve, however, considers that these embryos are only prolongations of the protoplasmic layer which secondarily cuticularize. The multilocular echinococcus, which in man produces a severe disease and almost always leads to premature death, infects most frequently the liver, but may also be found primarily in the brain, the spleen and the suprarenal capsule ; from the liver by means of metastasis it may reach the most various organs, especially those of the abdomen, but also the lungs, the heart, etc. Up to 1902, 235 cases have been described and up to 1906, 265, being 70 from Russia, 56 from Bavaria, 32 from Switzerland, 30 from the Austrian Alps, 25 from Wiirtemberg ; the remaining cases are distributed over Central Germany, Baden, Alsace, France, Upper Italy, North America. In some the origin is doubtful ; in any case after Russia, the mountainous South of Europe is the principal region of distribution. As to the domesticated animals, the same parasite is found principally in the ox (according to Meyer, in Leipzig, in 7 per cent, of the oxen affected with echinococcus) ; it is rarer in the sheep and very scarce in the pig. It has already been mentioned above that recently the multilocular echinococcus has been stated to be specifically different from hydatid or unilocular echinococcus. To this may be added the fact that Mangold, who fed a young pig with oncospheres of a Taenia reared from the multilocular echinococcus, found two growths in the liver four months later, which he took to be E. multilocularis, and con- sequently one has to assume the existence of two different worms. The chief defender of this view, already put forward by Vogler, Mangold, and Miiller, is Possett. He bases his opinions on (i) the more restricted distribution of the multilocular hydatid, the former occurring in districts where only cattle are raised, the latter where sheep-breeding is established ; (2) that those engaged in looking after sheep are attacked by multilocular, whereas those looking after cattle are attacked by unilocular hydatid ; (3) that among the cases of unilocular hydatid occurring in the distribution areas of multilocular hydatid no transitions between the two forms are observed ; (4) on the difference in the hooks both in the hydatid as-well as in the Taenia SERUM DIAGNOSIS OF ECHINOCOCCUS 359 stage ; the hooks of Tcviiia echinococcus are plump, sharply curved^ and have a short posterior root process the length of which is to that of the total length as i to 47, whereas on the contrary the hooks of the alveolar echinococcus are more slender, slightly bent, and have a long posterior root process (i to 2-5) ; and (5) on the form of the uterus, which in the alveolar Taenia has the form of a spherically distended sac anteriorly. Serum Diagnosis of Echinococcus. (i) Precipitin Reaction. —M\x equal parts of hydatid fluid (of the sheep) and serum of patient. Keep at 37° C. The reaction is not decisive as it may be given by normal sera. (2) Complement Deviation. — Required : (i) Hydatid fluid of sheep (antigen), (2) guinea-pig complement, (3) patient's serum, (4) red cells of sheep, (5) haemolytic serum (of rabbit) against sheep's red cells, (6) o*8 per cent, salt solution. Mix the antigen + patient's serum (heated) + complement + salt solution at 37° C. for one hour. Add red cells of sheep + haemolytic serum. Allow to stand for half an hour at 37° C. It is imperative to make adequate control observations. An example will indicate the method. Salt solution 1*3 c.c. 4- patient's serum (heated) 0*2 c.c. + hydatid fluid 0*4 c.c. + complement ci c.c. of serum diluted to a quarter strength + haemo- lytic serum and red cell emulsion i c.c. Result : no haemolysis, i.e.^ the patient's serum contains specific (echinococcus) antibodies. 360 THE ANIMAL PARASITES OF MAN C. NEMATHELMINTHES. J. W. W. STEPHENS, M.D., B.C., D.P.H. Bil.ATERALLY Symmetrical animals, without limbs and with a body cavity in which the gut or other organs float. They are generally cylindrical. Class. NEMATODA. Nemathelminthes with an alimentary canal. Nematodes are as a rule elongated round worms of a filiform or fusiform shape ; their length varies according to the species from about i mm. to 40 to 80 cm. The outer surface of the body is smooth or annulated, and at certain points provided with papilla?, occasionally also with bristles and alar appendages. The anterior end ■carrying the oral aperture is usually rather slender, occasionally quite thin ; the posterior end is pointed or rounded ; the anus, as a rule, lies somewhat in front of the posterior extremity. The sexes are almost always separate, and the male can ., gut; Dil.^ dilator cells of the gut; F., a process of the dilator cells forming a network over the vas deferens; SL, lateral line; Sp., spicule; Fa^., vas deferens. The anterior end of the worm lies to the right. Magnified. (After Goldschmidt.) formed of epithelium cells derived from the ovarian tubes. These cells subsequently fuse and form a membrane — the CHORION. The shape of the completed eggs is characteristic of the different species, and therefore a single egg often suffices to diagnose the species. According to the species, the eggs may be deposited sooner or later, either before or during segmentation, or with the embryo perfectly developed. Only a few species are viviparous, e.g.^ Dracun- ciilus medinensis, Trichinella spiralis; in the other Nematodes the further development of the extruded eggs takes place after various 372 THE ANIMAL PARASITES OF MAN lengths of time in the open, in moist earth, or in water. Thick-shelled eggs can maintain their developmental capacity for a long time, even after prolonged desiccation. Finally, a nematode-like embryo develops, which usually lies somewhat coiled up within the shell, and varies in its further development according to the species to which it belongs. In the simplest forms, as in the free-living Nematodes, the embryos, apart from their size, resemble their parents, and grow up into these after leaving the egg-shell. In many parasitical Nema- todes, however, the young must be called larvce, as they present characters which are subsequently lost. The manner of conveyance of the eggs or the embryos contained in them after they have left the body into the definite host is very different in the various species. (i) Without Intermediate Host. — (a) In many the conveyance into the definite host is effected directly after the larvae have developed within the eggs ; thus, for instance, the feeding of suitable animals with the embryo-containing eggs of species of Trichocephalus and Ascaris leads to an infection of the gut, for the young Trichocephali or Ascarides only leave the egg-shell when they have attained the intestine of the final host, in which they become adult. In other cases (6) Ancylostoma, Necator, the larvae hatch in the open, and live for a time free, changing their form ; they grow, cast their skin, and finally gain the intestine of the host by means of water or through the skin, when they lose their larval characters and assume the structure of the adult worm. (c) In a number of Nematodes, however, heterogony occurs. This terms signifies a mode of development in which two structurally different sexual generations of the same species alternate with each other. To these appertains, for instance, Angiostomum (syn. : Rhabdonema) nigrovenosumy which lives in the lungs of frogs and toads ; this Nematode measures about i cm. in length and is hermaphrodite (protanrdic). The eggs are deposited in the pulmonary cavity, and through the cilia of the same reach the oral cavity, where they are swallowed and thus conveyed into the intestine. They pass through the entire gut, and are finally evacuated with the faeces ; often, indeed, the young themselves emerge from the egg-shell within the hind-gut of the frogs. These young forms become sexually differ- entiated, remain much smaller than the parent, their oesophagus is differently constructed (rhabditis form), and they are non-parasitic (fig. 266). After having grown in the open they copulate ; the males die soon after copulation, and the females in their own bodies develop a few young, which, given the opportunity to get into frogs, infect them, and are transformed into the hermaphroditic Angiostomum. DEVELOPMENT OF THE NEMATODES 373 The same manner of development occurs in other species of the same genus, and also in the case of Stroitgy hides stercoralis. (2) With Ifiteiinediatc Host. — {a) Frequently, however, the larvae of Nematodes make use of one or even two intermediate hosts ; their condition then resembles that of Cestodes or Trematodes, excepting that there is never a multiplication within the intermediate hosts. The larvae become encapsuled amongst the tissues of the in- termediate host, and wait till they are introduced with the latter into the final host. For instance, OlUdanns tricuspis, the adult form of which is found in cats, previously lives encysted in the muscular system of mice. Cucidlaniis elegans, which at- tains the adult stage in fishes (perch, etc.), is found encysted in species of Cyclops. Other examples of species that require an intermediate host are Filaria barter of ti and Dractmculus medinensis. Peculiar conditions prevail in the case of {h) Trichinella spiralis. This species, which in its adult state lives in the intestine of man and of various mammals, is viviparous ; the young Trichinae, however, do not leave the intestine, but reach the intestinal w^all (Cerfontaine, Askanazy) in the following way : the female intestinal Trichinae bore into the intestinal wall, where they are found in the submucosa, or in the lumen of the dilated lacteal vessels. Here the young are born, in the intestinal wall, and leave this position with the lymph stream. Some of them, no doubt, actively bore through the intestinal wall, reaching the lymph or blood-stream, or even pass into the body cavity. What occurs during their further migrations is difficult to say at present. It has hitherto been maintained that the wandering is entirely active ; for instance, the ligaturing of an artery would be no protection against the part of the body supplied by such artery being invaded by Trichinella. This observation cannot be otherwise explained than by the active progress of the young Trichinella. The question, however, may be mooted as to where and when the worms quit the blood-vessels, which they for the most part reach through Fig. 269. — A piece of the trunk muscle of the pig with encapsuled embryonic Trichinse. Mag- nified. 374 THE ANIMAL PARASITES OF MAN the thoracic duct, the natural connection between tlie vascular system and the lymphatic system, to wander further independently, and ultimately reach the muscular system, in which they become encysted (fig. 269). Thus the progeny does not leave the body of the host inhabited by the parents, as is generally the case amongst helminthes, but uses it as an intermediate carrier to reach another host, which is then the final host. The latter may belong to another species, or may be another individual of the same species. This second migration is, of course, purely passive. Classification of the Nematoda. The Nematodes are usually divided into a number of families, some of which it is at present impossible to define accurately ; moreover, the definition of many genera is also in an unsatisfactory state. Family. Anguillulidae, Gervais and van Beneden, 1859. A "family" name not definable. They comprise a vast number of small forms, most of which live free in fresh water, in soil, or in macerating substances ; amongst them there are some which live parasitically on plants, more rarely on animals. They do not exceed 8 mm. in length. The large majority are only i to 2 mm., or even 0*5 mm. The uterus is straight. Eggs in the uterus at one time, one to four. Genera very numerous, but many of them insufficiently defined (Anguillula, Anguillulina, Rhabditis, Heterodera, etc.). Family. Angiostomidae, Braun, 1895. Small Nematodes undefined morphologically, but characterized by heterogony, i.e., there is a free-living " rhabditic " generation and a parasitic " filariform " genera- tion which succeed one another {e.g., Angiostomum, Strongyloides, Probstmayria). Family. Gnathostomidae. Cuticle covered totally or partly with cuticular laminae fringed posteriorly with multiple points. Head subglobular, covered with simple spines. Two spicules. Vulva behind middle of body, parasitic in vertebrates, especially mammals {e.g., Gnathostoma, Tanqua, Rictularia). Family. Dracunculidae, Leiper, 191 2. Males very small in proportion to females. Anus absent. Vulva absent (.?). Genera : Dracunculus, Icthyonema (in body cavity of eel and other fish). Family. Filariidse, Glaus, 1885. Long thread-like Nematodes. Anus present. CEsophagus without bulb. Vulva usually in anterior half of body. Two ovaries. Generally ovoviviparous. CLASSIFICATION OF THE NEMATODA 375 Development often requires an intermediate host. This family is at present ill-defined, but has been already subdivided into several sub-families, Filariince^ OnckocercincE, Arduennince. Family. Trichinellidae, Stiles and Crane, 1910. CEsophagus consisting of a chain of single cells, the lumen of the fjesophagus passing through their centre. Ovary single. Vulva at junction of anterior and posterior portions. Sub-families: (i) Trichurince^ (2) Trichinellincc. Family. Dioctophymldae. Body anteriorly armed with spines or unarmed ; mouth without lips, with six, twelve, or eighteen papillae in one or two circles ; oesophagus very long without a bulb ; anus terminal in female ; one ovary ; vagina very long ; spicule in male very long ; bursa cup-shaped without rays (Dioctophyme, Hystrichis, Eustrongylides). Family. Strongylidae, Cobbold, 1864. Bursa, supported by rays, always present. Oviparous. Family. Physalopteridae. Mouth with two large lips. Bursa with supporting papillae in form of a lanceolate cuticular expansion, with genus Physaloptera. Family. Ascaridae, Cobbold, 1864. Rather thick Nematodes. Mouth with three lips — one dorsal, two latere- ventral. Sub-families : (i) Ascarince, (2) Heterakince^ etc. Family. Oxyuridae. Smallish forms, 4 to 45 mm., with cuticle thickened on each side for the whole length of body in the form of a lateral flange or wing. CEsophagus long with a well-marked bulb containing a valvular apparatus. Tail end of female drawn out into a long point. Eggs asymmetrical. Males very sniall (about 2 mm.). One spicule. Genera : Oxyuris, Passalurus, Ozolaimus, Atractis, etc. Mermithidcs, greatly elongated " Nematodes," which, in the larval stage, are parasitic in insects, but in their adult condition are free living. Cuticle with diagonal striation. Without an open mouth or anus. Oral papillae present. Characteristic eggs with two processes, ending in a tuft of filaments. Larvas with a movable boring spine at the head end, Gordiidce. — Long, thread-like " Nematodes." Mouth and anterior portion of gut atrophied in adult. Oral papillae absent. 376 THE ANIMAL PARASITES OF MAN The Nematodes Observed in Man. Family Sub-family Genus Species Anguillulida- ... — Rhabditis R.pellio. R. niellyi. Rhabditis sp. Anguillula A. aceti. Anguillulina A. putrefaciens. AngiostotnidcB ... — Strongyloides St. stercoralis. Gnathostomidce .», — Gnathostoma On. siajnense. On. spinigerum. Dracunculidd ... — Dracunculus D. 7nedinensis. FilariidcB ... Filar iince ... Filaria F. bancrofti. F. deinarquayi. F. taniguchi. F. (?) conjunctives. Group. Agamofilaria ... Ag. georgiana. Ag. palpebralis. Ag. oculi huniani. Ag. labialis. F. (?) romanorum- orienialis. F. (?) kilimarce. F{l)sp.l {Mikrqfilaria) Mf. powelli. Mf. philippinensis. Setaria S. equina. Loa L. loa. A canthocheilonema Ac. Persians. Dirofilaria Di. 7nagalhdesi. Onchocercince Onchocerca 0. volvulus. Trichinellida ... Trichurince Trichuris T. trichiura. TrichinellincE Trichinella... T. spiralis. DioctophymidcE ... — Dioctophyme D. gigas. Strongylidce ... Metastrongy lines ... Metastf -ongylus . . . M. apri. Trichostrongylince Trichostrongylus ... T. instabilis. T. probolurus. T. vitrinus. Hcemonchus H. contortus. Mecistocirrus {Ne- M. fordi. matodirus) AncylostomincE Group. CEsophagostomecE Ternidens ... T. deminutus. CEsophagostojnuJTt. . . CE. brujnpti. GE. stephanostomum var. thomasi. CE. apiostomum. Group. Ancylostomece ... Ancylostoma A. duodenale. A. ceylanicum. A. braziliense. Group. Bunostomece Necator N. ainericanus. N. exilidens. Group. SyngamecE Synganius ... S. kingi. ANGUILLULID^ 377 P'amily Physalopterido' . . Ascaridcg Oxyurida Mermithidce Sub-family Genus — ... Physaloptera Ascarincp ... ... As carts Toxascaris Belascaris ... Lagocheilascaris Oxyuris Mermis {Agamomermis) Species P. caucasica. P. mordens. A. lumbricoides. A.sp. A. texana. A, maritima. T. limbata, B. cati. B. marginata. L. minor, O. vermicularis. M. hominis oris. A£: restiformis. Family. Anguillulidae. Genus. Rhabditis, Dujardin, 1845. Buccal cavity elongated, with lips. Its chitinous wall uniformly thick. Lateral lines absent. Males with bursa. Rhabditis peliio, Schneider, 1866. Syn. : Pelodera peliio, Schn., 1866; Rhabditis genitalis, Scheiber, 1880; Rhabditis peliio, Schn., 1866. Males 0*8 to 1*05 mm. in length ; females, 0*9 to 1*3 mm. in length. The posterior extremity of the body of the male has a heart-shaped bursa, and seven to ten ribs on each side ; the bursa may, however, be lacking. The spicules measure o'027 to 0*033 ^^V^- ii^ length, but are never quite alike. The posterior extremity of the female is long and pointed ; the vulva lies somewhat behind the middle of the body, the ovary is single, the eggs are oval, 60 jm by 35 fju. This species was found in Stuhlweissenburg by Scheiber in the acid urine (containing albumin, pus and blood) of a woman suffering from pyelonephritis, pneumonia and acute intestinal catarrh ; the observer was able to convince himself that the Nematodes which were found during the whole period of the illness lived in the vagina, and were evacuated with the urine. Oerley proved that this species had long been known ; during its larval stage {Anguilhila mucronata, Grube, 1849) it lives in earth- worms ; in its adult stage it lives in decomposing matter in the soil. By introducing individuals of this species into the vagina of mice, Oerley succeeded in obtaining infection and multiplication (facultative parasitism). These Nematodes must in some such way have got into the vagina of Scheiber's patient. Two other cases described by Baginsky and Peiper probably belonged to the same or a nearly related species. 378 THE ANIMAL PARASITES OF MAN Rhabditis nieliyi, Blanchard, 1885. Syn. : Leptodera nieliyi, Blanchard, 1885. In 1882 Nielly had a cabin-boy, aged 14, under observation in Brest. The lad had never left the neighbourhood of Brest, and had suffered from itching papules on the skin for five or six weeks ; in the papules the observer found one or several rhab- dites, measuring 0-33 mm. in length by 0*30 mm. in breadth. Their cuticle presented a delicate transverse striation ; the intestine was the only internal organ recognizable, and it opened somewhat in front of the posterior extremity. Therefore, it must have belonged to the rhabditis-like larva of a Nematode, the adult stage of which is unknown. The manner of infection was established almost certainly by a further obser- vation of Nielly's : at the commencement of the illness small Nematodes were found in the blood of the patient ; later on, however, they disappeared, neither were Nematodes found in the faeces, urine or sputum. Therefore it must be concluded that the cabin-boy, who was in the habit of drinking water from brooks, had thus ingested embryo-containing eggs of a Nematode ; the young hatched out in the intestine, perforated it, reached the blood and then settled in the skin ; but, on the other hand, the entry may have been direct through the skin. In connection with the foregoing, reference should be made to a communication by Whittles, insufficient from a zoological point of view. In a case of hypertrophic gingivitis occurring in a female patient, aged 19, who had never left Birmingham, he found Nematode larvae in the periosteum of the upper jaw, which was excised after extraction of the right premolar ; the genital rudiment could be recognized in them. Similar larvae were found in the same patient in abscesses in various regions of the skin, and in the case of her mother in the blood. The author considers that the infection took place through a dog, and refers to the case of O'Neil (1875), ^^^ found Filariae in the skin (in the condition known as "craw-craw"), referred by Manson to Filaria Persians. O'Neil's case was quoted, and attributed to Filaria sanguinis hominis. In conclusion, the author states that he has repeatedly found Nematode larvae in the blood of persons who suffered from pruritus; in his opinion the parasite had been imported through the agency of troops returned from South Africa. Glatzel found true Filaria larvae in a pustule of a cutaneous eruption of the trunk and extremities in a patient at Dar-es-Salam. Skin diseases which are caused by young Nematodes are also observed in dogs (Siedamgrotzky, MoUer, J. G. Schneider, Kiinnemann), foxes (Leuckart), and horses (Semmer). Ziirn found young Nematodes {AnguillulidcB) also in pig's flesh. In Kiinnemann's case it was shown that the adult Rhabdites lived in the straw upon which the dog lay. Rhabditis, sp. In the fluid obtained by lavage from the stomach of a female patient, aged 16, suffering from ozaena, O. Frese found during two consecutive months Rhabdites of various ages, 0*275 to 0-64 mm. in length, the adults all with eggs ; males were not found; transmission into rabbit's stomach failed, but they could be kept alive in much diluted hydrochloric acid (2 : 1,000) for several weeks. Neither eggs nor larvae ANGUILLULA 379 appeared in the faeces of the patient. The nature of the infection, which was perhaps of unique occurrence, remained doubtful. Genus. Anguillula, Ehrenberg, 1826. Buccal cavity very small, without lips. Males without bursa, but with a series of papillae. Lateral lines absent. Anguillula aceti, Miiller, 1783. CuticLile unstriped, body cylindrical, anterior end tapering but little, posterior end long, pointed. Male up to 1*45 mm. long, 0*024 to 0*028 mm. wide; two pre-anal papillae, one post-anal ; spicules equal, curved, 0*038 mm. long ; gubernaculum present ; testis extending in front of mid-line of body. Female up to 2*4 mm. long, 0*040 to 0*072 mm. wide ; anterior uterus reaching to near the oesophagus, posterior to hind gut. Viviparous ; embryos in both or only in one uterine horn, o'22 mm. long, 0*012 mm. broad. The species is a frequent inhabitant of vinegar (prepared by older methods), and was once observed for some time by Stiles and Frankland in the urine of a woman ; the urine had an acid reaction, and once had a distinct odour of vinegar. It was assumed that the patient, who was hysterical and suffered from chronic nephritis, employed vaginal douches with diluted vinegar, perhaps to deceive her physician or to protect herself against conception. According to Ward, Billings and Miller are said to have reported on two other cases. Ill-effects which might be connected with the vinegar eel {Anguillula aceti) were not present. Genus. Anguillulina, Gervais and Beneden, 1859. Syn. : Tylenchus^ Bastian, 1864. Characterized by the possession in the buccal cavity of a spine knobbed pos- teriorly ; bursa present ; uterus asymmetrical. Numerous species parasitic in plants. Anguillulina putrefaciens, Kiihn, 1879. Syn.: Tylenchtis putrefaciens^ YjSSvi\\ Trichina contorta^ Botkin, 1883. In 1883 Botkin {Pet.klin. Wochensckr., 1883) found a small Nematode, which was, however, entirely mistaken, in the material vomited by a Russian ; this was not a species of Trichinella, but an Anguillulina living in onions which had already, in 1879, been described by Kiihn as Tylenchus putrefaciens ; the Nematodes got into the stomach with the onions, causing nausea and vomiting. Family. Angiostomidae, Braun, 1895. Genus. Strongyloides, Grassi, 1879. Syn. : Pseudorhabditis^ Perroncito, 1881 ; Rhabdonema^ Leuckart, 1882, p.p. The genus is insufficiently defined. The parasitic form possesses a simple mouth opening directly into the long cylindrical oesophagus which occupies the anterior third of the body. The free-living forms possess a small buccal cavity ; the oesophagus is short, with a double bulb, in the hinder one there is a Y-shaped chitinous valve ; two spicules of equal size. 38o THE ANIMAL PARASITES OF MAN Syn. Strongyloides stercoralis, Bavay, ^877. Anguillula intestinalis et stercoralis, Bavay, 1877 ; Leptodera intestinalis et stercoralis, Cobb. ; strongyloides, Leuckart Pseudorhabditis stercoralis, Perroncito, 1881 ; Rhabdonema 1883 ; Strongyloides intestinalis, Grassi, 1883 ; Rhabdonema intestinale, Blanchard, 1886. In 1876, a number of French soldiers returned to Toulon from Cochin China suffering from severe diarrhoea. Dr. Normand, under whose treatment they were, discovered a large number of Nematodes in the evacuated faeces, and Bavay described CEsophagus Intestine / Excretory pore Ovary, anterior m Testis Ovary, anterior Ovary, anterior Ovary, posterior ,'»« Testis Anus Intes- tine . Vesicula seminalis Bursal muscles Anal papilla ;'' / . Papills .cU/- Spicule Ovary, posterior ^,-^«Hrw .^-^i^aE^ ^^^^^"^^^ Ovary, posterior Fig. 271. — Strongyloides ster- Fig. 270. — Strongyloides stercoralis^ female: parasitic (:^ra//j, male : free-living generation. generation from gut of man. x 70. (After Looss.) X 170. (After Looss.) them as Anguillula stercoralis. Soon after Normand, at the post-?nortem of five patients who had died of Cochin China diarrhcea. found numerous other Nematodes in the intestine, from the stomach to the rectum, in the bile-ducts and in the pancreas, and these he handed over to Bavay. The latter diagnosed another STRONGYLOIDES STERCORALIS 38 1 species, and described them as A. intestinalis. Both forms were then regarded as the cause of Cochin China diarrhoea until, in 1882, Leuckart was able to demonstrate that the two forms are only two succeeding generations of the same species, of which the one {A. intestinalis) lives parasiiically in the intestine, whereas its young {A. stercoralis) attain the open, where they come to maturity and propagate. The young of these again live parasitically. There thus exists the same heterogony as was discovered by Leuckart in Angiostomutn nigro- venosum of frogs, which heterogony, indeed, according to v. Linstow, appertains to the entile family of the Angiostotnidce. (i) The parasitic generation (strongyloid or filariform ?) is quite colourless and cannot be seen m situ even with a lens. To detect them it is necessary to scrape the mucosa of the jejunum and examine the scrapings microscopically. It measures 2*2 mm. in length, and 34 yu, to 70 yLt in breadth ; the cuticle is finely transversely striated ; the mouth is surrounded by four lips ; the oesophagus is almost cyl- indrical and a third the length of the entire body. The anus opens shortly in front of the pointed posterior extremity ; the vulva is situated at junction of middle and posterior thirds of the body; the uterus has no special ovejector ; the eggs measure 50 //, to 58 yu, in length, and 30 /i- to 34 yu, in breadth, and lie in a chain one behind the other (fig. 270). As in the case of Angiostomum nigrovenosiim^ Leuckart considers this stage to be hermaphroditic, the testes degenerating after having functioned; other authors (Rovelli) regard it as a female reproducing by parthenogenesis. (2) The free-living generation ( ^ and ? ) has a smooth body, cylindrical, somewhat more slender at the anterior extremity and pointed at the tail end. The mouth has four indistinct lips ; the oesophagus is short with a double (rhabditis-like) bulb ; there is a Y-shaped valve in the posterior bulb ; the anus opens in front of the tail end. The males measure 07 mm. in length, 0*035 i^^n^- ^^ breadth. Their posterior end is rolled up ; the two brown spicules are small (38 ft) and much curved. There is also a gubernaculum. The females measure i mm. in length or a little over ; 0*05 mm. in breadth. The tail end is straight and pointed ; the vulva lies somewhat behind the middle of the body. The yellowish, thin-shelled ova measure 70 fi in length and 45 ^ in breadth. As Askanazy has shown, the parasitic form bores deeply into the mucous membrane of the intestine, and frequently into the epithelium of Lieberkiihn's glands, both for nourishment and oviposition. The eggs then develop in the intestinal wall. The eggs which are found in scrapings from the mucosa occur, at least in the case of Strongyloides of the sheep, in chains enclosed in a thin tube or sheath, the origin of which is doubtful ; possibly it is the uterus. The eggs them- selves are only rarely found in stools, e.g.^ after a strong purge. The larvae, which are hatched out, and measure 0*2 to 0*25 mm. long by 382 THE ANIMAL PARASITES OF MAN o*oi6 mm. broad, again reach the lumen of the intestine,^ and grow to double or three times that size, until they are passed out with the faeces. They already differ from the parent ( $ ) in the shape (rhab- ditiform) of the oesophagus. When the external temperature is sufficiently high (26^ to :M Nerve-ring Excretoiy pore Ovary, anterior Ovary, anterior Ovary, posterior Ovary, anterior Anterior oviduct Anterior receptaculum seminis Posterior senainal receptacle M'1 )\aiy posterior Fig. 272. — Strongyloides stercoralis, female ; free-living generation, x 170. (After Looss.) 35° C), they become sexually mature after moulting. In about thirty hours they are com- pletely developed and copulate, now forming the free-living rhabditi- form generation. At lower temperatures the larvae only moult, but do not escape from the old cuticle and do not \ Genital rudiment .\'-V\ \ Fig. 273. — Strongyloides stercoralis : larva from fresh human faeces, x 310. (After Looss.) develop further. At a temperature of about 25° C. only some of the larvae attain maturity. ' As a case published by Teissier shows, they may also abnormally appear in the blood (Arch. mtd. exptr. et d'Aji. path., 1895, vii, p. 675). STRONGYLOIDES STERCORALIS 383 The females of the free-Hving generation (rhabditiform) deposit from thirty to forty eggs, which develop rapidly, sometimes even within the uterus in the case of old females. After the larvae have emerged from the egg-shell, they measure 0*22 mm. in length, and possess the characteristics of the parents (rhabditiform larvae). When they have grown to 0*55 mm. they moult, and while losing their own characteristics they acquire the characteristics of their parasitic grand- parents (strongyloid or filariform). After about eight days the free-living adult generation in the cultures have disappeared, and all the rhabditiform larvae have been transformed into strongyloid or filariform larvae ; they then die off unless they reach the intestine. This cycle of development holds good for Strongyloides stercoralis of tropical origin (Bavay, Leuckart, Leich- tenstern, Zinn). In the European Strongyloides the free-living genera- tion, as a rule, is absent (Grassi, Sonsino, Leichtenstern, Braun) ; the rhabditis-like larvae evacuated with the faeces are transformed into the strongy- loid or filariform type of larva (in cultures which are easily made) which will only become adult if introduced into man. So that we have these two cycles : {A) (i) $ parasitic, (2) eggs, the rhab- ditiform larvae in faeces, (3) free- living strongyloid or filariform larva, (4) $ parasitic. (B) (i) (2) (3) as before, then (4) adult $ and ^ , free living, (5) eggs, (6) rhabditiform larva, (7) strongyloid or filariform larva, (8) ? parasitic. Infection of man results not only Nerve-ring Subventral oeso- phageal glands Excretory pore CEsophagus -- Genital rudiment Anus Fig. 274. — Strongyloides stercoralis: mature filariform larva showing long transparent oesophagus, slender granu- lar intestine and characteristic tip to the tail ending in two small points. X 620. (After Looss). 384 THE ANIMAL PARASITES OF MAN from direct entry into the stomach but also, according to van Durme and Looss, through the skin. Occurrence i?i Man.—k.'s, already mentioned, Strongy hides stercoralis was first observed in persons suffering from so-called Cochin China diarrhoea. From the enormous numbers of parasites evacuated with the faeces, the cause of the disease was apparently evident. It appeared, however, that only some of the soldiers returning from Cochin China and Martinique, and suffering from diarrhoea, harboured Strongyloides (Chauvin). Breton made the same observations in Cochin China and found that only io'4 per cent, of cases of chronic dysentery, and 8*8 per cent, of chronic diarrhoea, show Strongyloides. Normand, moreover, found that only a few of the Europeans residing in Cochin China are exempt from S. intestinalis, yet the people exhibit no intestinal symptoms ; if, however, from any cause a catarrhal condition of the intestine supervenes the condition is changed, the parasites appear in larger numbers, and the disorder is considerably intensified. S. intesiinalis, besides being present in the Indo-China region, also occurs in the Antilles, in Brazil, Africa, and Europe ; in 1878 it was discovered in Italy by Grassi and C. and E. Parona ; in 1880 it was also found in the labourers working at the St. Gothard tunnel. It was imported into Germany, Belgium, and the Netherlands by Italian labourers. One sporadic case has been observed in East Prussia, and the worm has also been reported from Siberia. In mammals the following species are found : Probstmayria {Strongyloides) vivipara, Ransom, 1907, in Equus caballus ; Strongyloides fulleborni, \. Linst., in Anthropopithecus troglodytes and Cynocephalus babuin. Their development is, so far as is known, the same as that of Strongyloides stercoralis (v. Linstow, Centralbl. f. Bakt.^ Path. u. Infektionsk.., 19055 Orig. xxxviii, P- 532). Family. Gnathostonriidae. Genus. Gnathostonna, Owen, 1836. Syn, : Ckeiracanthus, Diesing, 1839. Easily recognizable by the numerous spines which cover the entire body or only the anterior extremity, and terminate in several points ; head globular and beset with bristles ; mouth with two lips ; two spicules ; vulva situated behind the middle of the body. Gnathostoma siamense, Levinsen, 1889. Syn. : Ckeiracanthus siamense^ Lev., 1889. Female measures 9 mm. in length, i mm. in breadth. There are eight rows of simple spines on the head ; the armature of spines extends over the anterior third of the body only ; each spine on the anterior region of the body spreads into three points, of which the middle one is the longest ; the posterior spines are simple ; they gradually become smaller and then disappear entirely. The vulva is situated behind the middle of the body. Male. — 10*5 mm. long by 0*6 mm. broad. Head terminates in a GNATHOSTOMA SPINIGERUM 385 globular swelling with two large lips. Neck 3 mm. broad. In front of neck eight rows of simple spines directed backwards. Anterior half of body with cuticular laminae, posterior unarmed. Two pre-anal and two post-anal papillae. Bursa wanting. Spicules I* I and 0*4 mm. respectively. Leiper considers Gnathostonia siainense to be identical with Guathostoma spinigerum. The single specimen described by Levinsen was found by Deiintzer in Bangkok (Siam), and was obtained from a young Siamese woman who suffered from a small tumour of the breast which had developed in the course of a few days. After the disappearance of the tumour, nodules the size of beans were found in the skin ; out of one of these the worm was obtained. The same observer saw this affection in two other persons. A closely related species, Gnathostoma spinigerum^ Ow., lives in the stomach of wild cat {Felis catus), puma {Felis con- color), tiger {Felis ti^ris), and domestic cat (India) ; another species, Gnathostoma hispidutn, Fedsch., 1839, in the stomach of pigs in Turkestan, Annam, Hungary, Congo, and by Collin in the stomach of an ox (Berlin). Gnathostoma sp. in pariah dogs, Calcutta. G?tathostoma sp. in monkeys, French Guiana. They produce large fibrous thickenings in the stomach wall. Gnathostoma spinigerum, Owen, 1836. Cuticle of bulb with eight rows of chitinous laminae with their posterior edges notched into spines. The laminae on the anterior portion of the body are similar trident laminae. In the middle 'of the body, the laminae are; simple and conical, cuticle posteriorly is un- armed. Mouth with two fleshy lips. Male 5 mm. long byo'S mm. broad; tail spiral,four pairs of papilla. Female about twice as long ; tail straight, trilobed. Fig. 275.— Gnathostoma siainense : to the left, the entire worm (8/i) ; to the right the head seen from above, with two fleshy lips (about 40/1). (After Levinsen.) Family. Dracunculidae, Leiper, 191 2. Genus. Dracunculus, Kniphoff, 1759. Anterior end rounded with a cuticular thickening or shield. Mouth triangular with two lips. Alimentary canal atrophied. 25 THE ANIMAL PARASITES OF MAN Dracunculus medinensis, Velsch, 1674. Syn. : Vena ?nedmensis,V ehch, 1674; Dracunculus persarum, Kampfer, 1694; Gordius medinefists, Linne, 1758 ; Filaria dracu?tculus, Bremser, 1819 ; Filaria cBthiopica, Valenciennes, 1856; Dracunculus medinensis^ Cobbold, 1864; Guinea worm^ Medina worm. The females attain a length of 50 to 80 cm., or even more, and average 1-5 to 17 mm. in diameter. They are whitish or yellowish in colour. The anterior extremity is roundish and bears a cuticular thickening or shield. The triangular mouth opening is surrounded by two projections or lips, behind which on the shield there are two lateral and four sub-median papillae ; the posterior end terminates in a spine, ventrally directed, and about i mm. in length ; the alimentary canal below the oesophagus is atrophied, but not entirely obliterated ; anus absent ; the lateral lines are very flat. The greater part of the body is occupied by the long uterus, in which a great number of young larvae are always found. The ovaries probably lie at the ends of the uterus ; the vulva lies just behind the cephalic shield. During parturition the uterus is prolapsed through this opening. The male is almost unknown. Leiper in an experimentally in- fected monkey found two males 22 mm. long, one from the psoas muscle, the other from the connective tissue behind the oesophagus. Occurrence. — Filaria medinensis has been known since the most remote period. The " fiery serpents " that molested the Israelites by the Red Sea, and which Moses mentioned, were probably filariae. The term Apa/c '^vnov occurs in Agatharchides (140 B.C.). Galen called the disorder dracontiasis ; the Arabian authors were well acquainted with the worm. It is found not only in Medina or Arabia, but also in Persia, Turkestan, Hindustan. The Guinea worm is also widely distributed in Africa, on the coasts as well as in the interior. It occurs in the Fiji Islands. It was carried to South America by negro slaves, but is said at the present lime to exist in only quite a few places (British Guiana, Brazil [Bahia]) ; it is also observed in mammals (ox, horse, dog, leopard, jackal \Canis lapuster\ etc.). Dracunculus medinensis in its adult stage lives in superficial ulcers on the body surface ; it is seen most frequently on the lower extremi- ties, more especially in the region of the ankle, but it also occurs in other parts of the body — on the trunk, scrotum, perineum, on the upper extremities, and in the eyelids and tongue. Sometimes there is only one ulcer and one worm, but more commonly several. It attacks man without distinction of race, age or sex. It is observed most frequently during the months of June to August. Life history.^ — When about a year old the worm seeks the surface ' The larvae resemble those of Cticullanus elegaus parasitic in the perch {Perca Jluviatilis). The larv£)e of this species develop in Cyclops sp. Fedschenko in 1870, at Leuckart's suggestion, succeeded in observing the invasion of Cyclops by Guinea worm larvae. They penetrate not per OS but through the exoskeleton. Newly hatched larvae (in bananas) will cause infection of monkeys. DRACUNCULUS IIEDINENSIS 387 Fig. 277. — Anterior ex- tremity of Guinea worm, showing dorsal and ventral lips, one lateral and two sub- median papillae and the lateral line. (After Leuckart.) Fig. 276. — Guinea worm {Dracunculus tnedi- nensis. (After Leuckart.) Fig. 278. — Dracunculus medinensis. a, anterior extremity seen end on ; O, mouth ; P, papillae ; b, female reduced more than half; er, larvae enlarged. fAfler Claus.) 388 THE ANIMAL PARASI'l ES OF MAN of the body and produces there a thickening as big as a florin. Over this a vesicle forms which eventually ruptures, and at the bottom of the ulcer can be seen a hole from which a part of the worm may project. On bathing the sides of the ulcer with water, a drop of fluid, at first clear then milky, exudes. This contains numerous larvae. In other cases a thin tube an inch long is prolapsed (through the vulva). This is probably the uterus, but the mechanism of parturition is not clearly known. It lasts for about a fortnight. An abundant supply of larvae can be got by placing wet compresses on a fresh ulcer. In a few hours a mass of larvae is obtained. The larvae are 500 yu< to 750 yit by 15/1, to 25 yit, with a long slender tail about one-third of the total length. The cuticle is transversely striated. The body is flattened. They possess an oesophagus and gut. At the anus there are apparently glandular structures. The larvae live and move actively in water for about two days, the majority dying on the third (Leiper). If a num- ber of Cyclops sp. have been collected and isolated in clean water, and the larvae are now added, the further development can be traced. The larvae enter the Cyclops, accord- ing to most authori- ties, by penetrating the exoskeleton, but according to Leiper this is impossible ; they must enter by the mouth and penetrate the gut in order to reach the body cavity. In eight days moult I takes place, the striated cuticle bemg cast off. In ten days moult 2 takes place. In five weeks the larva is mature. If now the infected Cyclops is placed in 0*2 per cent. HCl solution the Cyclops is killed immediately, but the larvae are stirred into activity, escape from the body, and swim about in the acid. This suggests that infection in nature probably takes place by the swallowing of infected Cyclops ; Leiper, by feeding Cyclops containing mature larvae to a monkey, found in it, post mortem six months later, two immature females 30 cm. long and two males 22 mm. long. In certain areas the new cases occur principally in June. Five Fig. 279. — Transverse section of temale Guinea worm ; u., uterus containing embryos ; i., intestinal canal ; ^., ovary. (After Leuckart.) DRACUNCULUS MEDINENSIS 389 weeks later the larvae will become mature in Cyclops, so that infection of Cyclops is taking place in July or August, and from then to June about ten months elapse, giving the period of development in man. Pathology. — The initial induration is accompanied by itching. Urticarial eruptions are described in Dahomey and Mauretanis^ accompanied by fever, rigors, blood-shot conjunctiva, and prostration resembling fungus poisonmg. Symptoms last for one to two days, later the worms appear on the surface. If the worm is ruptured in an attempt to extract it, disastrous results may occur through the escape of the larvae into the tissues : Fig. 280. — Cyclops virescetis^ $. 8, Female, ventral view, x 120; 9, anterior antennas X 240; 10, urosome and last thoracic segment, x 240; ii, foot of first pair, x 320; 12, 15, 16, foot of second, third and fourth pairs, x 240; 14, foot of fifth pair, x 440; 13, last thoracic segment and first segment of urosome of male, x 240. fever, inflammation, abscess, sloughing, ankylosis, even death from sepsis. Eosinophilia is often marked, 11 to 13 or even 50 per cent. Extraction. — (i) The native method consists in rolling the worm round a stick ; i in. to 2 in. are extracted each day, the process taking about a fortnight ; (2) Emily used injections of i in i,coo sublimate into the swelling or into the worm itself fixed by a ligature. 390 THE ANIMAL PARASITES OF MAN (3) Beclere chloroforms the worm ; (4) the worm can be more easily removed when all the embryos have been deposited (two to three weeks). Cyclopidce. — Cephalothorax ovate, clearly separated from abdomen. Anterior antennae of female when bent back scarcely ever stretch beyond the cephalothorax. The second antenn^e are unbranched. First four pairs of feet two-branched, outer branches three-jointed. The fifth pair of limbs are rudimentary alike in both sexes, usually one-jointed. There is no heart. The female has two egg sacs containing about fifty eggs. Genus. Cyclops, Miiller, 1776. Mandible palp rudimentary, reduced to a tubercle bearing two branchial filaments. Maxillary palp rudimentary (obsolete). Lower foot-jaw non-prehensile. Head ankylosed to first thoracic segment. Family. Filariidae. Sub-family. Filarllnae. The residue after exclusion of the ArduenniiKX and Onchocercincv. Genus. Fllaria, O. Fr. Miiller, 1787. Very long, slender Nematodes, without excretory vessels or excretory pore, the males of which are usually considerably smaller than the females. Mouth round, without lips, unarmed. The lateral lines occupy one-sixth of the circumference of body. The tails of the males are bent or spirally rolled, and bear little wing-like appendages. The two spicules are unequal ; almost always there are four pre-anal papillae, but the number of post-anal papillae varies. The vulva is always situated at the anterior extremity. Parasitic chiefly in the serous cavities and in the subcutaneous connective tissue. Insufficiently defined. Filaria bancrofti, Cobbold, 1877. Syn. : Trichina cystica, Salisbury,^ 1868 {iiec Filaria cystica, Rud., 1819) ; Filaria sanguinis hofninis, Lewis, 1872 ; Filaria sanguinis hominis cFgyptiaca^ Sonsino, 1875 ; Filaria wiichereri, da Silva Lima ; Filaria sa7tgui?tis hominum^ Hall, 1885; Filaria sanguinis hominis nocturna, Manson, 1891 ; Filaria nocturna, Manson, 1891. These parasites of man were for a long time only known in their larval stage. They were discovered in 1863 in Paris by Demarquay, in the hydrocele fluid of a Havanese emptied by puncture ; they were next observed by Wiicherer, in Bahia, in the urine of twenty-eight cases of tropical chykiria ; they were likewise observed in North America by Salisbury, who gave them the name of Trichina cystica. The next discoveries (in Calcutta, Guadeloupe, and Port Natal) related to rhyluria patients, until Lewis discovered the larvae in the blood of man (India), and * C. W. S iles ("American Medicine," 1905, ix, p. 682) is of the opinion ihat Salisbury's Trichina cystica is idenlical with Oxyuris vermicularis. FILARIA BANCROFT! 391 found they were almost always present in persons suffering from chyluria, elephan- tiasis, and lymphatic enlargements ; he also, in exceptional cases, found them in apparently healthy persons {Filaria sanguinis ho minis). Lewis and Manson studied the disease and the filariae of the blood very minutely, and became aware that the filaricC were sucked up bv mosquitoes with the blood. Manson described the Fig. 2%i.—FiIaria bancrofti. 1, Anterior portion of male; 2, two rows of papillse on head ; 3, papilla of tail of male ; 4, cloaca of male showing tips of spicules and gubernaculum ; 5, the spicules and gubernaculum of male. (After Leiper.) metamorphoses that take place within the body of the mosquito. The adult female was discovered in Queensland by Bancroft, and soon after Lewis found it in Calcutta ; it was described by Cobbold as F. bancrofti. The male was first seen by Bourne in 1888. 392 THE ANIMAL PARASITES OF MAN Head bougie-like, i.e.^ separated by a narrowing from the neck, having two rows of minute papillae. Cuticle has extremely fine striations. Female. — 50 to 65 mm. long by 1*5 to 2 mm. broad. Vulva 0-4 to 07 mm. behind the head. Anus about \ mm. from the tip of the tail (vulva i to 1*3 mm. from head, and anusciyto 28 mm. from tail according to other authors). The vagina' is a muscular tube form- ing three bold loops, and has terminally a pyriform enlargement. Uterus double (or single). Ovoviviparous. Male. — 25 to 30 mm. long by o'l mm. thick (40 by o*i mm. ac- cording to various authors). Probably two pairs of pre-anal papillae, eight pairs of peri-anal, two pairs of post-anal papillae, and one pair terminal. Tail curved. Two spicules, 0*2 and o'6 mm. respectively, and a cup-like gubernaculum. The long spicule is cylindrical, expanded proximally and tapering distally to a filament with wings. At the tip it is spoon-like. The short spicule is of the same diameter throughout. It is gutter-like, coarsely marked. Testis uncoiled, terminating in a snowdrop-like process (Leiper). ^^^' — 40 /^ by 25 jjb. They do not appear to possess a true shell, but only an embryonal or vitelline membrane secreted by the ovum. Embryos. — In the posterior part of the uterus eggs occur, in the anterior part embryos; the larvae at birth measure 127 fxio 200/1 by 8/A to 10 fji. In the blood they measure in the fresh 260 fju by 7*5 fx to 8/^. In stained films, owing to shrinkage, there is great variation in size, from 154 /A to 311 /A. Probably 260 yu, to 285 /a is the average in stained films. Geographical Distribution. — Europe : Two cases recorded, one from near Barcelona. The patient suffered from h^mato-chyluria and enlarged scrotum wqth mikrofilariae in the blood. A second case from Siena. Africa : The filarial index has not been estimated for various parts. In Nigeria it is about 10 per cent. //<76/7(7/.— Lymphatic glands : e.g., inguinal, femoral, iliac, lumbar, mesenteric, bronchial, superficial cervical, epitrochlear. Lymphatic vessels : e.g., those draining into the receptaculum chyli of the spermatic cord, in the thoracic duct and in various different parts. Organs, etc. : Testis, epididymis, spermatic cord, tunica vaginalis, mammary cyst, and in abscesses. They may occur in masses, but usually only a few (one to eight). Females are commoner than males. Dead and calcified worms are common in the various sites. Distribution of Larva: in Body,— ThtSQ are by no means uniformly distributed, but occur in greater number in tiie capillaries of the lungs. Besides the lungs they occur in the capillaries of other organs, as the following data of Rodenwaldt show:— Mikrofilariae Lungs ... I34,82i' Spleen Liver 4,884 Brain Kidneys ••• 15,253 Glands Glomeruli 8,008 Marrow Parenchyma 7,245 Blood FILARIA BANCROFTI 393 Mikrofilariae 1,666 3,833 o o 3>ooo The following data of Rodenwaldt refer to the larvae of Filaria itnmitis m the dog. They are commoner in organs than in vessels, and especially in the capillaries of the organs, but in the lungs they appear to be equally distributed in capillaries, arteries and veins. The length of life of larvae is unknown, but they appear to be destroyed in the kidneys, as dead calcified specimens are fairly numerous in the capillaries of the vasa recta of the medullary substance. Kidneys : mainly in the glomerular capillaries and those of the vasa recta. Liver : in the capillaries of the portal system, especially in those between the interlobular and the central intralobular veins. Periodicity of Larva^.^ — Roughly speaking, the larvae of Filaria bancrofti are found in the peripheral blood only during the night, disappearing (but not entirely) during the daytime. Their periodicity and that of Loa loa larvae is shown by the table on p. 394, based on that of Smith and Rivas (Amer. Jouru. Trop. Dis. and Prev. Med., 1914, vol. iii, p. 361). It was discovered by Mackenzie that this periodicity could be reversed by making the patient sleep during the daytime, showing that the phenomenon was in some way dependent on sleep or its attendant phenomena. Rodenwaldt gives the following explanation of the phenomenon of periodicity : — Mikrofilariae come to rest in capillaries. After passing up the thoracic duct they would reach the capillaries of the lungs by the superior vena cava. Here they occur in immense numbers. In the case of Loa loa larvae (which have a diurnal periodicity) some of these are forced out by the increased force and rapidity of the pulmonary circulation during the day, but are able to rest (owing to their sticky sheath ?) in the peripheral capillaries on their way to the capillaries of the organs. During the night the force of the current through the lungs is relaxed and consequently they are able ' These figures refer to i c.c. of each organ, and were estimated by cutting sections of definite thickness (30 fi to 40 fi) and counting the filariae in a definite area of section, e.^., I cm.^ The organs before removal from the body have their vessels tied, and are then fixed in hot alcohol. ^ For determining periodicity measured quantities of blood, e.^, 20 mm.*, should be used. A thick film is made of the whole quantity. The numbers present in this quantity may vary from three or four to 300 or 400. 394 THE ANIMAL PARASITES OF MAN to remain in the pulmonary capillaries and do not appear in the capillaries of the systemic circulation. If it is true that the periodicity of Loa loa cannot be reversed by changing the hours of sleep, then the explanation is incomplete. In the case of the larv?e of Filar ia bancrofti (which have a nocturnal periodicity), in order to apply the same explanation we must further assume that the mikrofilariae have less power of resisting the force of the capillary current {i.e., are less sticky). They are washed out of the pulmonary capillaries by day and by night, but it is only at night, when the blood-stream in systemic capillaries is less rapid, that they are able to rest there. In the day- time they are washed on until they reach the capillaries of the organs (possibly again the lungs). The reversal of tlie periodicity by sleeping Larvae of L. Average 132. Deviations fiom average Case i. Average 1,000 Case 2. Average 1,570 loa in equal F. bancro/ti (about). F. bancro/ti (about). quantities of larvae in i c.c. Deviations larvae in i c.c. Deviations blood of blood from average of blood from average 2 a.m. ... 9 - 123 3,500 + 2,500 6,500 + 3.930 4 a.m. ... 6 a.m. ... 1 1 - 121 3,200 + 2,2O0 5,200 + 3.630 41 - 91 2,800 + 1,800 2,000 + 430 8 a.m. ... 1 68 + 36 900 - 100 1,100 - 470 lo a.m. ... 298 + 1^6 210 - 790 350 — 1,220 12 noon ... 531 + 389 30 - 970 50 - 1,520 2 p.m. ... 252 + 120 20 - 980 4C - 1,530 4 pm. ... 6 p.m. ... 8 p.m. ... 146 + 14 10 - 990 30 - 1,540 91 - 41 40 - 960 40 - 1,530 2.1 - 99 60 - 940 ICO - 1,470 lo p.m. ... 5 - 127 600 - 400 800 - 770 1 2 midnight 5 - 127 750 - 250 2,600 + 1,030 Total ... 1,580 — 12,120 — 18,810 — during the daytime admits of a similar explanation. If this explana- tion be true, then a prolongation of the day conditions, e.g., by continued exercise, should result in still keeping the larvae out of the circulation, but this does not appear to be the case. In certain countries, e.g., Fiji, Samoa, Philippines, West Africa, larvae, apparently those of Filavia bancrofti, show no periodicity. In Fiji the usual intermediate host is Siegoinyia psendoscutellaris, a day- bitmg mosquito, so that possibly, as Bahr suggests, the mikrofilari^ have partly adapted themselves to the habits of their intermediate host, as the nocturnal mikrofilariae are adapted for transmission by a nocturnal feeding mosquito, e.g., Culex fatigans, but how this could come about is a mystery. It is not certain in all cases whether the non-periodic mikrofilariae really belong to Filaria bancrofti; some may be L. loa larvae, or possibly unknown larvae. An exact morphological description of these larvae is therefore always necessary. FILARIA BANCROFTI 395 Preservation of Living Larvce. — Blood from the vein (or finger puncture) is shaken up with twenty times its volume of sterile 0*9 per cent, salt solution, and kept in an ice cupboard (F'iilleborn). Concentration of Larvce. — {a) The above mixture is haemolysed with water and then sufficient salt solution added to make up to 0*9 per cent. The solution is allowed to stand or can be centrifugalized. (6) The blood is mixed with sodium citrate and centrifugalized ; the larvae are found in the leucocytic layer (Bahr). (c) Allow blood to clot in a small tube ; the larvae appear on the surface of the clot and are so got in pure serum. A drop of blood may also be allowed to clot on the slide ; the larvae are found in the clear areas of serum. {d) Haemolyse blood with water or acetic acid. Centrifugalize, make smears from, or examine the sediment. Removal of Red Corpuscles. — The blood film is allowed to stand for some minutes in a moist atmosphere. The staining solution is sucked through with blotting paper : the larvae stick to the slide, while the corpuscles are washed out. Morphology of Larvce. — Wet staining: Azur II one part, 0*9 per cent., salt solution 3,000, or very dilute Giemsa or ripened methylene blue or neutral red solutions. Place a drop on the slide and add a drop of blood to this. The larvae remain alive for one or more days ; it sometimes takes twenty-four hours to stain some particular structure. Differentiation by drawing through weak eosin solution is often useful. This method is the best for finest details. The excretory pore, anal pore, excretory cell, and chief " genital " cell stain hrst, then the matrix cells and finally the column of nuclei. Wet fixation and staining : The blood is spread on a large cover- glass — floated on the surface of 70 per cent, alcohol heated to about 70° C. Wash in water, (i) overstain with i in 1,000 azur II solution, warming slightly ; (2) differentiate with {a) absolute alcohol (con- taining, if necessary, a trace of HCl), or (6) with absolute alcohol 96 per cent, ninety parts, anilin oil ten parts ; (3) clear in origanum, bergamot or cajeput oil ; (4) mount in balsam. Or stain with haema- toxylin, e.g., Mayer's glycerine alumhaematein, heating till slightly steaming. Differentiate with acid (2 per cent. HCl) alcohol if ov^er- stained. Clear and mount as above. Dry fixation and staining: (i) With azur II as above, or (2) with haematein (warm). Exarnine the dried films in the usual way without a cover-glass. The azur stains the excretory and genital cells clearly. Thick films : (i) The blood is smeared out fairly thickly over an area as big as a sixpence. (2) Dry quickly to prevent shrinking, using carefully a spirit lamp in a moist climate. (3) Place films downwards in water for a few minutes. 396 THE ANIMAL PARASITES OF MAN (4) Fix in alcohol. (5) Stain withazur II, i in 1,000. Differentiate as above. Examine as a dry film. This method suffices for showing the excretory cell and the Gi cell ; or (6) Stain with haematein (slightly steaming), especially for the column of nuclei and the sheath. The fixation in alcohol in this case may be omitted. (7) The removal of the haemoglobin and the fixation may be combined by using Ruge's mixture (formalin 2 per cent., containing I per cent, acetic acid) or acetic alcohol (glacial acetic i, alcohol 3).^ Striidnre of Larvcv. — (i) Subcuticular cells : By vital staining, at intervals underneath the cuticle are seen a series of spindle-shaped cells — the sitbctiticular matrix cells of Rodenwaldt, the muscle cells of Fulleborn. There are thirty or forty or more of these. (2) Nerve ring : Appears as a break in the nuclear column about 20 per cent, of total length from the head. (3) Excretory system : Consists of a lateral spherical hollow excretory pore which shows a radial striation. Connected with the pore is an excretory cell which appears to be canalized. Excretory pore, 29-6 per cent, of length from head. Excretory cell, 30-6 per cent, of length from head. (4) " Genital " cells and anal pore : Consists of a pore opening ventrally on a very fine papilla with which are connected four other cells in series, the chief ^^ genital " cell (Gi) being some distance from the three others, which lie close to the pore. Gi, 70*6 per cent., anal pore, 82-4 per cent, of length from head. (5) Internal body, viscus, or reserve material : Best shown by vital staining with neutral red. This is agranular strand-like body extend- ing from 527 per cent, to 65 per cent, of length from head. (6) Tail end : (i) Rod-like structures resembling those in the head, 90 per cent, of length, (ii) The column of nuclei extends to 95 per cent, of length, so that the terminal portion is free from nuclei. (7) Mouth : Terminal according to some authors, lateral according to others. Some describe a fang on the head, others not. By vital staining and eosin differentiation two rod-like structures with mush- room-like caps can be seen behind the head. (8) Cuticle : Transversely striated. There is a longitudinal break in the striation on each side corresponding to the lateral lines. The stria- tion is best shown by vital staining with azur II and eosin differentiation. (9) Column of nuclei : These nuclei of the gut cells form the main feature in ordinary dry films stained with hematoxylin. They are separated by a space from the subcuticular cells. ' [Acetic alcohol does well for detecting crescents in thick films of malaria blood.— J. W. W. S.] FILARIA BANCROFTI 397 J3 a> 10 ^— ^ i ^ 398 THE ANIMAL PARASITES OF MAN Distinction between Mikrojilaria bancrofti and Mikroloa ha. Dry Films^ HcEmatoxylin Staining : — Mf. bancrofti. Ml. loa. (i) In graceful curves (but only if (i) Kinked, quickly dried). (2) Tip of tail free from nuclei. (2) Nuclei extend to tip. (3) Column of nuclei separated by (3) Not so distinctly. a space from the cuticle. (4) Gi cell small, easily overlooked. (4) Gi cell large, stains deep blue, cell protoplasm = twice width of larva, easily seen. (5) Excretory cell close to excretory (5) Excretory cell farther from pore, pore, 2 per cent, of length. 4 per cent, of length. Vital Staining with Neutral Red :— (6) Internal body or reserve mate- (6) Not shown. rial clearly shown. Life History. — In the stomach of the mosquito the larvae cast their sheath in the thickened blood in one to two hours. In twenty- four hours the majority have reached the thoracic muscles, where development proceeds. They are at first immobile and of a ^' sausage " form (iiOyLt by 13ft), with a short spiky tail. In three to five days the oesophagus is formed, the larva now being 0*5 mm. long. The larva appears to moult at this time. After the gut is formed papillae, three or four in number, appear at the tail end. In two to three weeks the larvae are i'5 mm. long. They now leave the thorax and reach the labium, but they may be found in various parts of the body, e.g.j the legs. They bore through Button's membrane and so arrive on the surface of the skin, which they rapidly enter. Their development in man is unknown, but it may be very long, as children are not infected till 4 to 5, or even lOj years old, but this may be due to unknown causes. , Development takes place in numerous mosquitoes. Anophelines : Myzomyia rossii, Pyretophorus costalis, Myzorhynchiis sinensis^ Myzo- rhynchus barbirostris, Myzorhynchiis peditceniatus. Culicines : Culex pipiens, Culex fatigans, Ciilex skusei, Ciilex gelidus, Culex sitiens, Culex albopictus, Stegomyia fasciata, Stegomyia pseudoscutellaris, Stegomyia gracilis, Stegomyia perplexa, Mansonioides imiformiSy Mansonioidesiannulipes, Scntomyia alboliiieata, Tceniorhynchtis domesticiis. Partial development takes place in other species. Pathology. — Among the conditions which Filaria bancrofti is FILARIA BANCROFTI 399 400 THE ANIMAL PARASITES OF MAN believed to produce are lymphangitis, varicose glands, especially inguinal and epitrochlear, chyluria, chylocele, lymph scrotum, orchitis, abscess, and elephantiasis. The evidence that these so-called ''filarial diseases" are produced by F. bancrofti is (i) geo- graphical and statistical ; (2) pathological. Bahr has contributed evidence of the former kind from his researches in Fiji, on which we may base the following statements: — (i) The prevalence of filarial diseases is proportional to the prevalence of Mikrofilaria bancrofti in the blood. Thus in four villages examined by him he got the following figures: — Village A Village B Village C Village D Mf. bancrofti 12*5 per cent. . . . 25 per cent. . 31 per cent. . 33 per cent Filarial diseases .. 29 „ •• 39 ... 58 „ .. 34 M Tolal population .. 168 .. 114 ... .. 425 ... .. 222 (2) Out of 257 people with Mf. bancrofti in the blood, 153 were suffering from filarial diseases, i.e., 59 per cent. (3) Whereas of 672 people without Mf. bancrofti in the blood, only 263 were suffering from filarial diseases, i.e.y 37*6 per cent. (4) Again out of 416 people suffering from filarial disease, 153 showed Mf bancrofti in their blood, i.e., 367 per cent. It is generally assumed that all people suffering from filarial disease show at some (presumably early) stage larvae in the blood ; but we do not consider that this must necessarily be so. It appears to us quite possible that living adult filariae may be present in the body, producing disease, without their larvae appearing in the blood. The absence of larvae from the blood in 63*3 per cent, of persons suffering from filarial disease is, however, generally explained otherwise. The adults which occur in enlarged glands, etc., get eventually destroyed by inflammatory reaction, so that larvae are no longer being produced, while the enlarged gland, etc., which the adults have produced remains. This explanation assumes that the larvae of the original worm die in the circulation or elsewhere, e.g., kidney, but we have no evidence as to the duration of life of larvae in the human body; but also it assumes that a person cannot be reinfected with filaria, for otherwise there is no reason why the diseased should not be infected in the same proportion as the non-diseased. But assuming the explanation to be true, it would explain why a diseased population show larvae in only about one-third of the cases. It must be borne in mind also that the figures are rather small. Pathology. — In order to explain the effects which do or may be expected to occur from obstruction of lymphatics, it is necessary to have an accurate knowledge of the distribution and connections of FILARIA BANCROFTl 4OI lymphatic vessels (and glands) and the anastomoses of these vessels. We can onlv briefly summarize our knowledge here. We should recall also that considerable destruction or obstruction of lymphatics or glands may occui- without necessarily producing any lymphatic obstruction, at least, of a permanent nature, e.g.^ when a mass of lymphatic glands is destroyed by a bubo in the groin or, again, when a carcinomatous mass of glands is removed from the axilla. Again, to take the case of chyluria — where it is generally assumed that obstruction must occur higher up than the point at which the intestinal lacteals enter the juxta-aortic glands — this disease may occur, e.g., in temperate regions, quite apart from such obstruc- tion. It is true that some of these cases of chyluria are not cases of chyle in the urine, but, as little or no fat is present, lymphuria. These do not require the above assumption, but seeing that true chyluria may apparently occur without such obstruction, we should be cautious about explaining this and other symptoms on the basis of obstructions which theory may demand, for only too often there are no post-inorteni facts at our disposal. Lymphangitis: What this is due to is unknown. There is no actual evidence of the occurrence of adults in the inflamed vessel. Complete disappearance, not to reappear, of (non-periodic) mikro- filariai from the blood has been shown by Bahr and others to occur within twenty-four hours after an attack of lymphangitis, orchitis adenitis or simply a high temperature. This mysterious phenomenon requires explanation. If the mikrofilariae were being killed by the attack, their dead bodies should still be found in the blood ; or if the adults were being killed, for all we know to the contrary, the larvae might well survive. We consider there is no evidence that either are affected, but that for some reason, as little understood as in periodicity, the larvae now remain in the organs. Abscess : In Fiji, by Bahr, they have been found in the substance of various muscles, e.g., quadriceps extensor, latissimus dorsi, serratus magnus, in the popliteal space, groin, axilla, and over the internal con- dyle of the humerus, and in the upper extremity they are frequently infected with cocci. They not infrequently contain fragments of dead adult filarial. Their mode of origin is not clear. They form nearly 30 per cent, of cases of filariasis in Fiji. Of 95 cases, 41 showed mikrofilariae in blood, 54 did not. Hydrocele and enlarged testis : In Fiji they form about 10 per cent. (36 out of 343) of cases of filariasis. The fluid is usually sterile ; mikrofilariae were present in the fluid in i out of it cases. In the wall numerous calcified adult filariae may be found. The walls consist chiefly of hypertrophied muscle with fibrous tissue, dilated blood- vessels and lymphatics, the lining epithelium of which appears to be 26 402 THE ANIMAL PARASITES OF MAN absent ; of 38 cases 14 had mikrofilariae in the blood, 24 had not. Most of the cases are associated with elephantiasis of the scrotum (11 out of 12 cases). Enlarged glands form over 40 per cent. (153 out of 343) of cases of filariasis, so that they are the commonest expression of filariasis met with in Fiji. The glands are enlarged, fibrotic, and the trabeculae are thickened. The lymphatics are thickened or represented merely by fibrous tissue. The gland also shows dilated blood-vessels and numerous spaces filled with lymph. Giant-cells are common in those glands which contain remnants of filariae. Masses of lympho- cytes enclosed by inflammatory or fibrous tissue are common. Eosinophile cells are also extremely common, not only in the fibrous tissue of the glands, but in other inflammatory or fibrotic conditions : in other organs living or calcified filariae are " usually " present. Only about 33 per cent, show mikrofilariae in the blood. The epitrochlear gland is frequently enlarged in Fiji. Breinl has examined enlarged glands and finds loose vascular fibrous tissue with lymphocytic invasion. In parts, the lymphocytes collect into areas 200 //, to 800 /i in diameter. The lymph tissue surrounding the spermatic cord showed abundance of vessels — large, (2) small. The large had thick walls and wide lumina. In other cases the lumina were nearly filled by a thrombus of newiy formed, fine, loose connective tissue. Varicose glands: In about 7 per cent. (24 out of 343 cases) of filariasis, mikrofilariae are found in the blood in 50 per cent. (12 out of 24). Elephantiasis.— E\eph3.ni'i2isih scroti is associated with hydrocele in 50 per cent, of cases (12 out of 23) ; in 65 per cent, of cases (15 out of 23) there are associated enlarged glands in one or both groins, though also hydrocele and enlarged glands occur without elephantiasis scroti. In 13 out of 27, i.e., about 50 per cent., cases of elephantiasis in various regions, no associated enlargement of glands is found. Elephantiasis forms in Fiji less than 10 per cent, of cases of filariasis. Mikrofilariae are present in the blood in 36 per cent. (12 out of 33) of cases. C/^y/z/r/a.— Exceedingly rare in Fiji. Theory would demand an obstruction above the point t)f entry of the lacteals, viz., the pre-aortic lymphatic glands, but in cases in temperate regions it may occur without any such lesion. In some of these cases the fluid is not chyle (fat absent), but presumably lymph. A discussion of the mode of production of chyluria, lymph scrotum, elephantiasis, etc., is at present premature ; theory has far outrun fact. Too much stress had been laid on the mechanical action of the worms to the almost total exclusion of their (or possibly their larval) toxic action. The above FILARIA DEMARQUAYI 4O3 analysis has been made in the hope of acquiring more extended observations similar to those made by Bahr. Geographical Distribution. — Filaria hancrofti is known in nearly all tropical countries. It occurs in India, China, Indo-China, Japan, Australia, Queensland, the Islands of Polynesia (with the exception of the Sandwich Islands), Egypt, Algeria, Tunis, Madagascar, Zanzi- bar, Sudan, etc., the south of the United States of America, Brazil, the Antilles, etc. Whether it is the same species in all cases is questionable. Filaria demarquayi, Manson, 1895. Syn. : F. ozzardi, Manson, 1897. The adult female F. demarquayi measures from 65 to 80 mm. in length by 0*21 to 0*25 mm. in breadth. The head has a diameter of from 0*09 to 0*1 mm. The mouth is terminal. The genital pore opens at 076 mm. from the head. The alimentary canal is nearly straight and terminates in an anus, which is subterminal. The opening of the anus is marked by a slight papilla. The tail is curved. It rapidly diminishes in size just below the anal papilla. A characteristic pair of fleshy papillae project from the tip of the tail. The diameter near the tip of the tail before its termination is 0*03 mm. F. deniarqtiayi is a thicker worm than Ac. perstans. It differs from F. hancrofti in the greater size of the head, in the smaller tail, and particularly in the F1G.284.— /^.^t to 80 /a in breadth. The diameter of the head is 0*04 mm. The tail is much curved. There are four pairs of pre-anal papillae and two pairs of post-anal papillae. Spicules very unequal in size. Cloaca 121 /a from the tail end. At the tail end two triangular cuticular appendages. 4i6 THE ANIMAL PARASITES OF MAN The adalt worms inhabit the connective tissue at the base of the mesentery, especially in the region of the pancreas, abdominal aorta and suprarenals. To find them the mesentery should be removed, placed in a 2 per cent, solution of formalin, and then carefully examined at leisure. Mikrofilaria perstans. — 160 fiio 210 fi by ^ fi to 6 /jl broad. Has no sheath. Cuticle transversely striated. Tail rounded off, not pointed. Nerve ring at 34 fi. Excretory pore 49 fi, genital pore 125 fju from head. Smaller larvae 90 fju to no /a by 4 yit broad. A " fang" is also described on the head. M/. Persians. Mf. demarquayi. (i) Tail stumpy. (i) Tail pointed. (2) Column of nuclei extends to tip of (2) Does not extend to tip. tail. Periodicity. — N one. Life-history. — Unknown. Geographical Distribution. — Very common in many parts of Africa : Sierra Leone, Dahomey, Northern Nigeria, Southern Nigeria, Came- roons, Ivory Coast, Gold Coast, Old Calabar, Congo, Uganda. Absent from Zululand, Basutoland. On the East Coast of Africa it is not found in the towns of Zanzibar and Mombasa, neither is it found in the country of the Masi, nor amongst the Kavirondo, who dw^ell along the north-east shores of Lake Victoria. In South America, Ac. perstans is very common amongst the aboriginal Indians in the interior of British Guiana. However, it is not found in Geoi-getown and in New Amsterdam, neither is it found in the cultivated strip of coast lying between these two towns, but it is common on the coast farther north near the Vene- zuelan boundary, where the forests stretch to the sea. The Waran Indians, who live at the mouth of the Waini river, harbour this parasite. It is absent in the West Indies. Topographically, Ac. perstans is found only in areas covered by dense forest growth and abounding in swamps. In Kavirondo, where the forest disappears and the land is covered with scrub and short grass, it is not found ; likewise it is not found on tlie grassy plains of the highlands of British East Africa. Towns and cultivated areas are free from it. Genus. Dirofilaria. Railliet and Henry, 191 1. Body very long, thread-like, cuticle transversely striated. Mouth with six papillae. Male tail spiral with voluminous pre-anal and some large post-anal papillae ; spicules unequal. Vulva near the anterior hundredth of body; viviparous. Parasitic in heart or blood-vessels and subcutaneous tissue. ONCHOCERCIN^ 417 DIrofilaria magalhaesi, R. Blanchard, 1895. Fig. 300. — Dirojilaria magalhaesi : posterior ex- tremity. (After V. Lin- stow.) Syn. : Filaria bancrofti^ v. Linstovv, 1892 ; Filaria bancrofti^ P. S. de Magalhaes, 1892 inec Cobbold, 1877). The male measures 83 mm. in length by 0*28 to 0-40 mm. in breadth. The anterior extremity is rounded, and has no papillae (? 6) ; the posterior extremity exhibits a double curve, with four pre-anal and four post-anal papillae on each side. These are large and have a villous appearance. The mouth is round and unarmed, the pharynx measures i mm. in length, is cylin- dfical, very muscular, and its hinder part is dilated. The anus is situated o'li mm. in fron of the hind end. There are probably two unequal spicules; one only, however, is known — apparently the shorter one — the length of which is given as o'ly to 0*23 mm. The female measures 155 mm. in length and 0*6 to o*8 mm. in breadth ; the rings of the cuticle are 0*005 nam. apart (in the male 0-003 mm. apart) ; the anterior extremity is slightly thickened and club-like, the posterior extremity is slender, and terminates obtusely ; the lateral line is o'i27 mm. in breadth (that of the male 0*007 to o*oo8 mm.); the anus opens o'i3 mm. in front of the hind end, the vulva is 2*5 mm. distant from the mouth, the ovaries are two much convoluted tubes. The eggs measure 38 /><, by 1 1 /i,. This species was first discovered at a post-mertem^ in the left ventricle, by J. P. Figueira de Saboia in Rio de Janeiro, and has been described by P. S. de Magalhaes. D. immitis occurs in the right ventricle of the heart of the dog in Europe and the Tropics. D. repens is also a common subcutaneous Nematode in dogs in Annam. Sub-family. Onchocercinae, Leiper, 191 1. Cuticle with spiral thickenings. Genus. Onchocerca, Diesing, 1841. Male with four pre-anal papillae. Female with vulva situated anteriorly. Onchocerca volvulus, R. Leuckart, 1893. Syn. : Filaria volvulus^ R. Leuckart, 1893. The adult male measures 30 to 35 mm. in length by 0*14 mm. in breadth. The body is white, filiform, attenuated at both ends. The head is rounded and has a diameter of 0*048 mm. The cuticle 4l8 THE ANIMAL PARASITES OF MAN is distinctly transversely striated. The mouth is unarmed. The alimentary canal is straight, the anus opening q-o; mm. from the tip of the tail. The tail is strongly curved and somewhat flattened on the concave surface. There are three papillae, one large and two small, on each side of the cloaca and one large and two post-anal small papillae. Two curved spicules, 0*166 and 0*08 mm. respectively. The adult female is of uncertain length, but much longer than the male, probably about 10 to 12 cm. The head is rounded and truncated ; it measures 0*065 mm. in diameter. The tail is curved. The vulva opens 0*55 mm. from the head. The hand-like cuticular thickenings are well marked. Eggs ovoid with a prolongation at each pole 'Mike an orange wrapped in tissue paper." The larva measures about 300 yu, by 7 /a to S fi; it has no "sheath." The body tapers from about the last fifth of its length, and terminates in a sharply pointed tail. At about the anterior fifth of the body there is a V spot. 0. volvulus is found in peculiar subcutaneous tumours, the size of a pea to that of a pigeon's Qgg. The same patient may present one or several of these tumours. The regions of the body most fre- quently affected are those in which the peripheral lymphatics con- verge. Thus they are usually found in the axilla, in the popliteal space, about the elbow, in the sub-occipital region and in the inter- costal spaces. The tumours are never adherent to the surrounding structures, and can be easily enucleated. They are formed of a dense connective tissue wall and internally a looser fibrous meshwork. This is traversed by a series of canals in which the worms lie, but they are also partly embedded in the denser wall. The canals apparently dilate into Cavities filled with slimy pus-like fluid con- sisting largely of larvae. According to Brumpt the posterior extremity of the male, and the anterior extremity of the female with its vaginal opening, are free in one of the spaces for the purpose of copulation and parturition. If a tumour be cut into and placed in salt solution, Rodenwaldt states that the undamaged males wander out into the solution. The formation of the tumours is elucidated by Labadie-Lagrave and Deguy's case. The authors found an immature female Onchocerca volvulus in a lymphatic vessel partly obstructed by an infiltration of fibrin and leucocytes. It appears, therefore, that the presence of the parasites within the lymphatics gives rise to an inflammatory process, and that the consequent fibrinous deposit envelops the parasites, obliterates the lumen of the vessel, and ultimately isolates the affected tract. At any rate, in young tumours the worms appear to lie in a structureless substance permeated by leucocytes in which connective tissue is gradually organized from the periphery, thus isolating the worms. TRICHINELLID^ 419 In cases of infection with 0. volvulus larvae have been found by Onizilleau, Fiilleborn, and Simon in lymph glands, and in the finger blood if considerable pressure is used so as to squeeze lymph out of the tissues. They are sheathlesSj and the following are the dimensions in ordinary dried films: Length, 274 /^ ; nerve ring, 23-7 per cent. ; Gi cell, 6()'6 per cent. ; end of last tail cell, 96*3 per cent. The dimensions of larvae of 0. volvidiis taken from the uterus and prepared in the same way are: Length, 224*5 yu,; nerve ring, 24-3 per cent. ; Gi cell, 68*9 per cent. ; end of the last tail cell, 95*5 pci' cent. In all probability the larvae in the glands and blood are those of 0. volvulus. According to the natives, the tumours may last indefinitely and never ulcerate. Some old patients told Brumpt that their tumours had been present since childhood. Probably Onchocerca volvulus, like some other Filariidce, may live for many years. 0. volvulus occurs in various parts of West Africa : Gold Coast, Sierra Leone, Dahomey, Lagos, Cameroons. Brumpt, on the banks of the Welle between Dongon and M'Binia (Belgian Congo), found about 5 per cent, of the riverine population affected. Family. Trichinellidae, Stiles and Crane, 1910. Sub-family. Trichurinae, Ransom, 191 1. Male with a single long spicule, with sleeve-like sheath. One ovary. Eggs with an opening at each pole closed by a plug-like operculum. Eggs hatch on being swallowed by a new host. Genera : Trichuris, Capillaria. Genus. Trichuris, Roderer and Wagler, 1761. Syn. : Trichocephalus, Goeze, 1782 {7iec Trichiurus, L., 1758); Mastigodes^ Zeder, 1803. The anterior part of the body is very long and thread-like : the posterior, much shorter part, is thicker, rounded posteriorly, and the anus is terminal. The males have the posterior extremity spirally rolled ; the vulva is situated at the commence- ment of the posterior part of the body. The Trichocephali live in the large intestine of mammals, the caecum by predilection ; their development is direct, infection occurs through the ingestion of embryo-containing eggs. Trichuris trichiura, Linnaeus, 1761. Syn.: Trichocephaliis trichiurus., L., 1771 ; Ascaris trichiura, L., 1771 ; Trichocephalus hominis, Schrank, 1788; Trichocephalus dispar., Rud., 1801. The male measures 40 to 45 mm. in length, the spicule is 2*5 mm. long, its retractile sheath is beset with spines. The female measures 45 to 50 mm. in length, of which two-fifths appertain 27 4^0 THE ANIMAL PARASITES OF MAN to the posterior part of the body. The ova are barrel-shaped and have a thick brownish shell which is perforated at the poles. Each opening is closed by a light-coloured plug. The eggs measure 50 fi to 54 yLt in length and 23 fi in breadth ; they are deposited before segmentation. Trichuris trichinra usually lives in the caecum of man, and is also occasionally found in the vermiform appendix and in the colon, exceptionally also in the small intestine ; usually only a few specimens are present, and these do not cause any parti- cular disturbance, although, as Askanazy found, they feed on blood ; in other cases cerebral symptoms of more or less severity are observed when Trichocephali are present in large numbers. At posi-nwrtems performed soon after death the filiform anterior extremity of the worm is frequently found embedded in the mucous membrane (Askanazy). The whip worm is one of the most common parasites of man and appears to be distributed over the entire surface of the globe ; it is, how- ever, more frequent in the warmer regions. It is found in persons of both sexes and all ages with the exception of infants. In autopsies it is found in the following numbers : In Dresden in 2*5 per cent., in Erlangen in ii'i per cent., in Kiel in 31 '8 per cent., in Munich in 9-3 per cent., in Petrograd in o'i8 per cent., in Gottingen in 46"i per cent., in Basle in 237 per cent., in Greenwich in 68 per cent., in Dublin in 89 per cent., in Paris in about 50 per cent., and in Southern Italy in almost 100 per cent. On examining the fasces the eggs of the whip worm were found as follows : In Munich in 8*26 per cent., in Kiel in 45*2 per cent., in Greifswald in 45 per cent., in North Holland in 7 per cent., in Novgorod in 26*4 per cent., in Petrograd in 5 per cent., in Moscow in 5*3 per cent. Fig. 301. — Trichufis trichiui-a : on the left, male ; on the right, female with the anterior extremity embedded in the mucous membrane of the intestine ; below, egg. The development of the eggs is completed in water or in moist soil, and occupies a longer or shorter time according to the season ; the eggs possess great powers of resistance, as do the larvae, which, according to Davaine, may remain as long as five years in the egg- shell without losing their vitality. Leuckart proved by experiment that direct infection with Trichuris ovis {Ovis aries) and T. crenata (Siis scrofa dom.) was produced by embryo-containing eggs ; Railliet obtained the same results with T. depressiusciila of dogs, and Grassi subsequently, by rrieans of two experiments, demonstrated the direct development of Trichuris trichinra. In one case embryo-containing eggs were swallowed on June 27, 1884, and on July 24 the ova of Trichocephali were found in the faeces for the fii-st time. TRICHINELLIN^ 42I Trichuris trichiiira is found not only in man, but also in various monkeys {T. palceformis, Rud.), as well as in lemurs (T. lemur is, Rud.)- Other species are T. crenata in pig ; T. ovis in cattle, sheep, goat, and pig (?) ; T. depressiusciila in dog ; T. campanula in cat ; T. unguiculata in rabbit and hare ; T. cameli in camel; T. discolor in humped cattle ; T. nodosus in mouse ; T. alcocki in the thamin (India) ; T. globidosa in camel ; T. giraffce in giraffe. Sub-family. Trichinellinae, Ransom, 191 1. Male without spicule ; females ovoviviparous. Larvae penetrate muscles of host and become encysted. Genus : Trichinella. Genus. Trichinella, Railliet, 1895. Syn. : Trichina^ Owen, 1835 (^^^ Meigen, 1830). Very small Trichinellincs, the males of which have two conical appendages at the •caudal extremity ; the vulva is situated at the border of the anterior fifth of the body. There is only one species. Trichinella spiralis, Owen, 1835. Syn. : Trichina spiralis^ Owen, 1835. The male measures i"4 to i'6 mm. in length and 0*04 mm. in diameter. The anterior part of the body is narrowed, the orifice of the cloaca is terminal and lies between the two caudal appendages ; internal to these are two pairs of papillae, dorsal one behind the other. .The cloaca is evertible for copulation. The females measure 3 to 4 mm. in length and o'o6 mm. in diameter ; anus terminal. Trichinella spiralis in its adult stage inhabits the small intestine of man, pig, wild boar, rat. The young do not leave the body of the host but become encysted in the muscles. Experimentally it develops in the black rat (Mus rattus), the sewer rat {M. decumamis), the ■domestic pig (Sus scrofa dom.), the wild boar {Siis scrofa ferox), the domestic dog {Canis familiaris) , the fox (C. vulpes) the badger (Meles iaxiis), the polecat (Putorius fcetidits), the marten (Mustela /oina), the raccoon (Procyon lotor), the hippopotamus and the cat, and many other mammals (rodents and carnivora) ; Trichinellae have been arti- ficially introduced, by administering the encysted stage, into the dog, the mole {Talpa europcea), the mouse (Mus musculiis), the hare {Lepus timidus), the rabbit (L. cuniculiis), the hedgehog {Eriiiaceus europcBus), the marmot {Cricetus vulgaris), the vole, the dormouse, the sheep, the calf, the horse, etc. Human beings and the pig, rat, mouse, guinea- pig and rabbit are most easily infected ; less easily the sheep, calf and horse ; with difficulty the cat, dog and badger. Trichinella can also be reared in birds (fowl, pigeon and duck), but the young do not 422 THE ANIMAL PARASITES OF MAN encyst in the muscular system, but are expelled with the faeces. By cold-blooded animals as well as by insects (Calliphora vomitaria) Fig. 302. — Trichinella spiralis. ?, mature female : E^ embryos; V^ vulva; Ov, ovary. i , mature male : 7", testes, c, newly born larva, d., larva in the muscles. 4 P^r cent.) were invaded by Trichinella. The cases, according to the nationality, are divided as follows : — Trichinella Examined Percentage of positive results Absent Present Americans : — {a) Whites 207 201 6 2-89 (6) Negroes 70 65 5 7-14 British and Irish 62 57 5 8 -06 Canadians 12 10 2 16-66 Germans 49 43 6 12-24 Italians 12 10 2 16-66 Other nationalities 27 27 Nationality unknown 66 65 I I-5I Total 505 478 27 5'34 It is worthy of remark that half of all the positive cases were mental patients^ who were found to be affected with Trichinella to well-nigh 12 per cent. Trichinosis was not, however,"the cause of death in any case. Very frequently the Trichinella^ were found calcified and dead. Conditions are similar in most countries of Europe, where, of course, the number of infected pigs is considerably smaller, but the disease depends less on this than on the way in which the pork is prepared. Cases of trichinosis have been known to occur in nearly all the countries of Europe; further, in Egypt, Algeria, East Africa. Syria, India, Australia, and America. North Germany, more TRICHINELLA SPIRALIS 429 especially the Saxe-Thiiringian states, is the classical land for epidemics of trichinosis ; the mortality varies, but it may be very high.^ Prophylaxis. — The grave nature of the disease and the comparatively high mortality relating to trichinosis led the authorities to adopt certain preventive measures, which are the more necessary as national customs cannot be altered in a short time. As the usual process of pickling and smoking, even when long continued, does not certainly ensure the death of the Trichinellse contained in the meat, and also because in roasting and boiling large pieces of pork a considerable time is necessary to permit the temperature required to kill off the parasites (62° to 70° C.) to penetrate to the middle of the joint, it appeared to be most practical to have all pigs microscopically examined for Trichinellae before they, or parts of them, were placed on the market, and all infected meat condemned, no matter whether the Trichinellae were present in large or small numbers, still undeveloped or calcified. Since 1877 obligatory examination of pork has been introduced in Prussia, though as yet it is not thoroughly carried out; other states of North Germany as well as the larger towns of South Germany soon followed ; a complete army of trichina inspectors, officially examined and periodically con- trolled by experts, and whose number in Prussia amounted to 27,602 in 1896, this being even increased to 28,224 in 1899, have the charge of examining pork on certain lines laid down. These are at the present time uniformly administered. The proceeding is usually that the trichina inspector himself goes to the slaughterhouses, or special samplers take pieces of the muscles that are known to be the favourite seats of the parasite (pillars of the diaphragm, the costal part of the diaphragm, muscles of the tongue and larynx, intercostal and abdominal muscles) ; six small portions are separated from each piece, pressed between slides or special compressors, and carefully gone through by examining them with a low power of the microscope. The pigs free from Trichinellae are passed for commerce; trichinous pigs, on the other hand, in Prussia, are only allowed to be used for industrial purposes, i.e.^ the hide and bristles are used, the fat is allowed to be melted down, or certain parts are used for the manufacture of soap or glue. In Saxony, however, it is still permitted to place trichinous flesh on the market, fully declaring its nature, and after having been heated to its deepest strata at a temperature of 100° C. in a suitable apparatus, and under the supervision of a veterinary surgeon. As TO THE PROPORTION OF TRICHINOUS PIGS to healthy ones, the following tables give the figures for Prussia : — ' For instance, extensive epidemics occurred in Hettstadt in 1863 (160 patients, 28 deaths) ; Hanover, 1864 — 1865 (more than 300 patients) ; Hadersleben, 1865 (337 patients, loi deaths); Potsdam, 1866 (164 patients); Greifswald, 1866 (140 cases, i death); Magdeburg, 1866 (240 cases, 16 deaths) ; Halbersiadt, 1867 (100 cases, 20 deaths) ; Stassfurt, 1869 (over 100 cases) ; Wernigerode, 1873 (100 cases, i death) ; Chemnitz (194 cases, 3 deaths); Linden, 1874 {400 cases, 140 deaths); Niederzwohren, near Cassel, 1877 (half the population); Diedenhofen, 1877 (99 cases, 10 deaths); Leipzig, 1877 (134 cases, 2 deaths) ; Ernsleben, 1883 (403 cases, 66 deaths) ; Strenz-Neuendorf, 1884 (86 cases, 12 deaths), etc. According to Johne, 109 epidemics, with 3,402 cases and 79 deaths, occurred in Saxony between i860 and 1889. Stiles, in a work recently published, states that there were 8,491 cases of trichinosis with 513 deaths (6*04 per cent.) in Germany from i860 to 1880 ; and 6,329 cases and 318 deaths (5'02 per cent.) between 1881 — 1898. Of these latter, 1881 — 1898, 3,822 (225 deaths) occurred in Prussia, 1,634 (76 deaths) in Saxony, and 873 (17 deaths) in the remaining states. There is, however, no doubt that many deaths from trichinosis were not recognized, as proved by experience at post-vwrtems. 430 THE ANIMAL PARASITES OF MAN Number of Number of Year pigs examined trichinous pigs Proportion 1878 2,524,105 1,222 I : 2,065 1879 3,164,656 1,938 I : 1,632 1881 3,118,780 1,695 I : 1,839 1882 3,808,142 1,852 I : 2,056 1883 4,248,767 2,199 I : 1,932 1884 4,611,689 2,624 I : 1,741 1885 4,421,208 2,387 I : 1,852 1886 4,834,898 2,114 I : 2,287 1887 5,486,416 2,776 i: 1,988 1888 6,051,249 3,111 I • 1,945 1889 5,500,678 3,026 I : 1,818 1890 5.590,510 1,756 I : 3,183 189I • 6,550,182 2,187 I : 2,996 1892 6,234,559 2,085 I : 2,992 1896 8,759,490 1,877 I : 4,666 1899 9,230,353 1,021 I : 9,040 1902 9,093,210 725 I : 12,397 The proportion, however, is not only subject to variation in separate years, but differs according to the district ; thus, in 1884, in the state district of ?vlinden there was one trichinous pig to 30,146 healthy animals, in Erfurt i to 14,563, in the district of Gnesen i to loi, in Schrimm i to 86, and in Schroda i to 68. In Germany Trichinella is becoming LESS COMMON in pigs (Ostertag) :— {a) Piiissia. Year 1878— 1885 1886—1892 1896 ... 1899 ... 1902 Year 189I 1892 1893 1894 1895 1896 1899 1902 Year 1883-1893 1893— 1897 1902 {b) Saxony. (c) City of Berlin. Pigs found to be trichinous o-o6i- -0-048 per cent 0-033- -0-043 0-02I 0014 O'OII ,, Num found to ber of pigs be trichinous 0*014 per cent. o-oii ,, o*oo8 ,, 0-007 ,, 0-012 ,, 0-0I02 ,, 0*004 0-0056 " Number of pigs found to be trichinous 0*035- -0*064 per cent. 0*022- -0-015 „ 0*006 There is no doubt that the excellent preventive measure of official inspection for Trichinella has led to the avoidance of grave disasters; its introduction has not yet caused an entire cessation of trichinosis in man, because inspection of pork is not obligatory DIOCTOPHYMID^ 43 1 everywhere, so that human beings may become infected by unexamined trichinous pigs from their own country or from abroad^ and also because an infection may occasionally escape notice. For these reasons the meat imported into Berlin from abroad as free from Trichinae is examined again and not always in vain ; finally, also^ gross negligence may at times occur, or fatal errors may be made. In addition private prophylaxis must not be neglected, and its chief aim should be directed to the suitable preparation of pork. Family. Dioctophymldae. Genus. Dioctophyme, Collet-Megret, 1802. Syn. : Eustrongyhts, Dies., 1851. Large worms. Anterior extremity unarmed ; the mouth is surrounded by six papillae. One ovary. The vulva is in the anterior region of the body. Dioctophyme gigas, Rudolphi, 1802. Syn. : Dioctophyjjie renale, Goeze, 1782 ; Ascaris caiiis et mariis, Schrank, 1788 j Ascaris visceralis et renalis^ Gmelin, 1789 ; Sirongylus gigas ^ Rud., 1802 ; Eustrongy- lus <^igas. Dies., 1851 ; Strongylus renalis^ Moq. Tand., i860 ; Eustrongylus visceralis y RailL, 1885. Colour blood-red ; the anterior extremity somewhat slender ; there is a series of about 150 papillae along the lateral lines ; the sub-median lines are strongly developed, and from them spring the radial muscles for the intestine. The males attain a length of 40 cm. and a diameter of 4 to 6 mm. ; the posterior extremity is transversely truncated ; the anal orifice is within the base of the collar-like bursa, the thickened edges of which are beset with papillae ; the spicule measures 5 to 6 mm. in length. The females attain a length of 100 cm. and a breadth of 12 mm. The anus is crescent-shaped and terminal. The vulva is 50 to 70 mm. distant from the anterior extremity. The eggs are oval and have a thick shell presenting numerous depressions ; the shell itself is brownish, but it is colourless at the somewhat thickened poles ; it measures 60 //, in length by 40 /a in breadth. The larva measures 240 /x by i4yL6. Dioctophyme gigas lives in the pelvis of the kidney, more rarely in the abdominal cavity of the seal, otter, dog, wolf, fox, horse, marten and polecat, exceptionally also in human beings. It also occurs in tumours of the mamma and perinaeum. Most of the cases in which this parasite has been reported as occurring in man may be traced back to unrecognized Ascaris hunbricoides or to clots of fibrin ; seven certain cases, eight more or less doubtful, however, remain. 432 THE ANIMAL PARASITES OF MAN The source of infection is unknown, but according to Balbiani the eggs develop an embryo in water or moist soil, and this embryo may remain alive several years without hatching ; the infection of dogs with embryo-containing eggs did not succeed ; an intermediate stage in fishes is conjectured, but still the infection of cattle and horses is unintelligible. Famity. Strongylidse. Sub-family. Metastrongylinae, Leiper, 1908. Buccal capsule absent or slightly developed, vagina elon- gate, uteri convergent^ and have a simple musculature. Parasitic in the respiratory or circulatory system. Genera : Metastrongy- lus, Synthetocaulus. Genus. Metastrongylus, Molin, 1861. Mouth with six lips, of which the two lateral are the largest. Postero- and postero- external rays- of bursa thin, the rest thick. Only the median ray double. Spicules very long and slender, striated. Vulva immediately in front of anus. Eggs contain an embryo when laid. Fig. 306. — Diocto- phyme gjgas, male. Natural size. (After Railliet.) Fig. 307-— Eggs of Dioctophyme g'gas ; above seen from the flat, below in optical section. 400/1. (After Railliet.) Metastrongylus apri, Gmelin, 1789. Syn. : Gordius puhnonalis apri, Ebel, I777 ; Ascaris apri, Gmelin, 1789; Strongylus suis, Rud., 1809; Strongylus paradoxus, Mehlis, 1831 ; Strongylus elongatus, Duj., 1845; Strongylus longevaginatus. Dies., 185 1. The male measures 12 to 25 mm. in length ; the bursa is bilobed ; there are five rays in each lobe ; the spicules are thin and up to 4 mm. in length. The females measure 20 to 50 mm. in length, the anus is close in front of the posterior extremity, which * Convergent : i.e., the uteri are parallel, converging from the anterior part of body to the vagina, which is near the anus, this position being associated with convergence of the uteri. Divergent : Uteri run anterior and posterior, diverging from the vagina, which in this case is near middle of body, ''■ For nomenclature of rays vide p. 449. TRICHOSTRONGYLIN^ 433 CL.yrt has a recurved, hook-like process ; the vulva is close in front of the anus. The eggs are elliptical, 57//, to 100 yit in length, 39 /a to 72 yu, in breadth ; when the eggs are deposited the embryo is already formed, 220//, to 350//, by lOyLt to 12 fJb. Metastrongylus apri frequently lives in the bronchial tubes — usually the smaller ones — of the pig^ and wild boar ; it is also found occa- sionally in sheep and in man ; in young pigs it is apt to set up a bronchitis, which frequently causes death. The first communication as to the occurrence of this species in man was that of Diesing, who, in 1845, in Klausenburg, had the opportunity of examining Strongylidce found by Jortsits in the lung of a little boy, aged 6, in Transylvania; probably also the Nema- todes found in the trachea and larynx of man, and described by Rainey and Bristowe as specimens of Filaria tracliealis^ belong to this group ; according to Chatin, Metastrongylus apri may also occur in the intestine of man ; this oc- currence, however, may in all probability have been due to an accidental introduction of adult worms into the intestine, and should not be attributed to an infection by the larval stage. No experiments to induce in- fection have been made ; it is prob- able, however, that infection is direct and without the aid of an intermediate host. Fig. 308. — Metastrongylus apri: one side of bursa, a., anterior; a.e.^ antero-external ; a.m., antero-median ; p.m., postero-median ; p.e., postero- external ; /. , one division of posterior ray. (Stephens.) Sub-family. Trichostrongylinae, Leiper, 1908. Strongylidae with buccal capsule absent or slightly developed, vagina short, uteri divergent (/.^., anterior and posterior), ovejectors differentiated. Parasitic in the alimentary canal. Contains the genera Trichostrongylus, Haemonchus, Ostertagia, Nematodirus, Cooperia, Dictyocaulus.'^ 1 The reports of the city inspection of meat in Berlin state that Strongylidce in the lungs of pigs are by no means rare; therefore the lungs of 1,941 pigs were condemned between 1885— 1886, of 1,641 between 1886— 1887, of 3,237 between 1887—1888, of 4,855 between 1888 — 1889, of 7,197 between 1889— 1890, and of 5,574 pigs between 1890— 1891, etc. Ostertag found Stro7igyliis apri in 60 per cent, of the pigs examined in the Berlin abattoir ; Meyer, in Leipzig, found the parasite in 15 per cent, of the native pigs and in 52 per cent, of the Hungarian pigs. ^ Dictyo:aidns is parasitic in the bronchi. ' 434 THE ANIMAL PARASITES OF MAN Genus. Trichostrongylus, Looss, 1905. Very small Strongylidce, Mouth with three small lips and nodular or punctiform papillae. Cervical papillae absent. Bursa entirely closed; with large lateral lobes, and median lobe not distinctly defined. Anterior^ rays double, the branches widely divergent, one thin, the other thick, and close to the antero-median. The postero- median ray is thin and close to the postero-external. Posterior ray bifurcate, each branch bifid at the tip (fig. 311). Spicules short, spoon or spatula-like, with on the broad anterior end a lateral knob or disc and in h'ont of the point an angular projection. Gubernaculum of a peculiar canoe or shoe shape in profile. Vulva in the hinder half of the body. Tail with two minute papillae just in front of tip. Eggs thin shelled ; when laid they show eight to thirty-two segments. Parasitic in duodenum, seldom in the stomach of herbivora. Syn. Trichostrongylus instabllis,^ Railliet, 1893. Stro7igylus instabilis^ Railliet, 1893; Strongylus subtilis, Looss, 1895. Male 4 to 5*5 mm. long, o'o8 mm. thick in front of bursa. Spicule 0*135 to o'i45 mm. long, accessory piece (gubernaculum) 0*07 mm. thick. Antero-external ray usually thickest of all, occa- sionally only as thick as the antero-median ; postero-median far more slender than the antero- external and antero-median and ; Fig. 309. — Trichostrongylus instabilis : left, posterior end of male ; right, spicule and gubernaculum, side view. C/. fig. 311. Mag- nified. (After Looss.) Fig. 310. — Trichostrongylus instabilis : posterior end of female. Magnified. (After Looss.) nearer to the postero-external than to the antero-median. Female 5 to 6 mm. long, vulva 1-05 to 1*2 mm. distant from the tip of the • When the anterior ray is double, the branches of it are called an tero -anterior and latero- anterior. ^ Identical with T. colubriformis of the sheep according to Leiper. If so, this latter name has priority. TRICHOSTRONGYLUS PROBOLURUS 435 tail, placed longitudinally, 50 /x to 55 /a long, always shorter than the unpaired portion of the canal formed by the union of the two ovejectors ; anus 0*055 to 0*07 mm. distant from tip of the tail ; ova 73 yLt to 80 /A by 40 /a to 43 yu,. This species lives in the duodenum, exceptionally also in the stomach of Oris aries, 0. laficaudn, Ant Hope dorcas, Camelus dronie- darius (Egypt), CynocepJialns haniadryas (North Africa), sheep and goats (France), and has been found by Looss in bodies of fellaheen at Alexandria and in the stomach of a Japanese female by Ijima. Trichostrongylus probolurus, Railliet, 1896. Syn. : Strongylus probolurus^ Railliet, 1896. Male 45 to 5'5 mm. long, in front of bursa o'o8 mm. thick ; spicule o'i26 to o'i34 mm. long, gubernaculum 0*075 to o'o8 mm, long. Bursa : latero-anterior rib thickest ; antero-external thicker than antero-median, postero-median and postero-external very short and close together. Female 4*5 to 6 mm. long, vulval opening I'oS to 1*25 mm. from tip of tail, placed longitudinally , and slightly curved, 76 /a long. Fig. 312. — Trichostrongylus probolurus ." spicules and gubernaculum of male ; on left, ventral view ; on right, lateral view. X c. 300. (After Looss.) Fig. 311. — Trichostrongylus probolurits : tail of male from left side, d., posterior; e.d., postero- external ; /./., postero-median ; m.l., antero- median ; e.i., antero-external; /.z/., latero-anterior; V.V., antero-anterior ; g-zib., portion of guberna- culum ; sp., portion of spicules. x c. 300. (After Looss.) always longer than the unpaired portion of the ovejector ; anus 0*040 to 0*05 mm. distant from tip of tail. Posterior end thick, point of tail short. Ova 76 //, to 80 yu, by 43 yu, to 46 /ju. Habitat. — In the duodenum of Ovis aries, 0. laticatida, Antilope dorcas, Camelus droniedarius (Egypt) and occasionally also in man (Egypt). Trichostrongylus vitrinus, Looss, 1905. Male 4 to 5*5 mm. long, in front of bursa 0*085 mm. thick. Bursa larger than in the other two species, antero-external rib thickest, 28 436 THE ANIMAL PARASITES OF MAN antero-anterior and postero-median equally thick, straight. Spicule o*i6 to 0*17 mm. long, gubernaculum 0*085 to 0*095 ^^' long- Female 5 to 6-5 mm. long, vulval opening 1*15 to 1*25 mm. distant from tip of tail, crescent shaped, oblique to body axis, and around it irregular thickenings. Ova 84 yit to 90 /x by 46 fi to 50 //,. In duodenum of Ovis aries, 0. laticatida, occa- (7u^if. sionally in Camelus dronie- dariiis and in man (Egypt). v.v- ^-&.d. YiG. ZiX.— Trickostrofjgylus vitrinus : tail of male Y\G.Z\i^.-Trichostrongylus from left side. «'., posterior ; e.d., postero-external ; z'^/rz^z.^ : spicules and guber- /./., postero-median ; ^^/./., antero-median; ^./.,antero. naculum ; on left, ventral external; l.v., latero-anterior ; v. v., antero-anterior; ^^^w ; on right, lateral view. gub., portion of gubernaculum ; sp.^ portion of spicule. ^ ^' 3°°' (Alter Looss.) X c. 300. (After Looss.) Genus. Haennonchus, Cobb., 1898. Small mouth cavity contains a '' tooth " or ^* lancet " arising from the dorsal side. Cuticle of head and neck not inflated. Cervical papillae well marked. Bursa bilateral, with large lateral lobes and a small dorsal lobe not median, but lateral, attached to the base of one of the lateral lobes (fig. 316). Posterior ray bifurcate, each branch tifid apically. Each lateral lobe six rays. Anterior rays separated distally, curving forward. Antero-median and postero-median rays distally curve away from the antero-external. Postero-external ray long and slender. Spicules less than i mm. Gubernaculum present. Vulva in posterior part of body covered by a prominent tongue-like flap. Eggs ellipsoidal. Haemonchus contortus, Rudolphi, 1803 ; Cobb., 1898. Dorsal ''tooth" or "lancet" 10 /x to i5yLt long. Cervical papilla C3 mm. from head. Male 20 mm. long by 400 /x thick (maximum). Asymmetrical lobe H^MONCHUS CONTORTUS 437 of bursa 150 yu, by 125 //, attached to left lateral lobe. Posterior ray bifurcate ; each branch bifid. Stem of ray less than twice as long as its branches. Spicules 300 yu, to 500 /^ with knobbed tips, and the left spicule with a barb 20 fju from the tip, right spicule with a barb 40 /a from tip. Gubernaculum 200 />«, by 25 yL6 to 35 /x, fusiform with thick- ened edges. Female 18 to 30 mm. by 500 fj. (maximum). Vulva 3 to 4*5 mm. from tip. Linguiform flap 0*5 mm. (a second one exists, according to Brumpt). Anus 400 fi to 630 //, from tip. Tail acutely pointed. Eggs 75 /a to 95 yU< by 40 yLt to 50 fJL. :^;..'OV^ — int. A — e^eL Fig. 315. — Hcemonchus contortus: vulval region of female viewed from left side, int.^ intestine; lab.^ lingui- form process covering vulva ; ov.^ ovary ; ovij.y ovejector ; ut.^ uterus; vag., vagina ; z/«/., vulva. X 75. (After Ransom.) m.l. Fig. 316. — Hcemonchus contortus: tail of male, dorsal view, d., posterior ray of the asym- metrically placed posterior lobe; ^.o'., postero- external ; /./., postero-median ; mj., antero- median ; o^u:d^ Fig. 319. — (Esophagostornum stephatiostomum var. thomasi. i, male, natural size ; 2, female, natural size; 3, head of female, ventral view, showing cephalic vesicle and ventral cleft limiting it behind, x 55 ; 4, head of female, dorsal view, x 225 ; 5, head of male, end view, showing external and internal leaf crowns, x 225 ; 6, tail of male, lateral view [cf. fig. 318, D), X 20; 7, tail of female, lateral view, x 20 ; 8, CEs. thomasi, posterior ray of bursa, x 150; 9, (Es.dentigerum, from chimpanzee, posterior ray of bursa, X 150; 10, CEs. stephanostomum, from gorilla, posterior ray of bursa, x 150. GESOPHAGOSTOMUM STEPHANOSTOMUM 443 muscularis mucosae of the gut wall. The cysts project internally and externally, and contain inmature adults, which eventually escape into the lumen of the gut. Fig. 320. — CEsophagosiomum stephanostomum var. thomasi : csecum and ascending colon. Subperitoneal cysts are seen on the top right hand, and in the lumen of the gut numerous cysts arranged transversely. The small roundish patches are areas of necrosis in the cyst walls. ^ After Thomas.) CEsophagostomum stephanostomum var. thomasi, Raill. and Henry, 1909. Body thick, pointed only at the ends. Buccal capsule much reduced. External crown of thirty-eight leaflets (the *^ crown " nearest the centre of lig. 319, 5). Male 17 to 22 mm. long by 750 //, thick. Spicule 1*380 to 1*475 mm., slightly curved at the tip. Female, 444 THE ANIMAL PARASITES OF MAN iinmature, i6 to 20 mm. long by 900 yu, thick, tail ending in a little conical appendage. Anus 230 jju, vulva 500 jx to 525 fi from tip. Ovejectors close together. Uteri very short in form of oblong pouch, CEs. stephanostomuni, Stossich, 1904, m the large intestine of gorilla. (Es. stepliauostomuui var. dentigera, Raill. and Henry, 1909, in the chimpanzee. Habitat. — In large and small intestine of man, Brazil. Pathology. — Nodules occur in the gut wall ; 187 were found by Thomas in his, the sole case. The tumours contain each a single worm. Fig. 321. — (Esophagoslomuvi Stephanos totmim var. thomasi : portion of the ileum, showing a cyst with protruding worm, x 8. (After Thomas.) Fig. 322. — CEsophagostomum ste- phanosto77iuvi var. thomasi : colon with oesophagostome withdrawn from its cyst cavity, x 20. (After Thomas.) CEsophagostomum apiostomum, Willach, 1891. According to Leiper, CEs. hnimpti is identical with, and hence a synonym of, this species. Parasitic in large intestines of monkeys, producing dysentery, and in man (Northern Nigeria). According to Walker this species is common in Philippine monkeys. Ova are scanty in the faeces. They measure 73 /a to 84 //, by 44 /x to 57 fi and are in the morula stage. They are easily cultivated. The rhabditiform larva is 340 yLt by 16 /x and has a long filiform tail. It moults twice, and at the second moult becomes a filariform larva retaining the skin of this moult, this stage being that of the mature larva. It.now measures 9 mm. long by 30 /x thick. Walker suggests that the mode of infection is similar to that of ancylostomes. ANCYLOSTOME^ 445 Group. Ancylostomeae, Railliet and Henry, 1909. Bursa with anterior ray cleft, median double,^ postero- and postero- external arising from a common trunk, posterior bifurcate, each limb being tridigitate. Vulva in posterior third of body. Uteri divergent. Contains the following genera : (i) Strongylus,' (2) Ancylostoma, (3) Uncinaria, (4) Characostomum, etc. Genus. Ancylostoma, Dubini, 1843, emend. Looss, 1905. Ventral margin of mouth capsule armed with teeth, the "roots" of which are continued backwards and appear on the external surface of capsule as rib-like thickenings. Terminal third of dorsal ray cleft. Genital tubes very long, with short, closely packed diagonal convolutions. Ancylostoma duodenale, Dubini, 1843. Male 9 mm. long by 0*45 mm. thick, female 12 mm. long by 0*6 mm. thick. Pale flesh colour, or an intense red in posterior third. Anteriorly may be more or less black due to (blood) pigment in the cells of the chyle intestine (= stomach + small intestine). The worm is about the same thickness all through and is plump and rigid. Cuticle striated. The body has a marked torsion, so that if the ventral side of the head is upwards the anus appears to open laterally and vice versa. The dorsal curve of the head end is only slight and the oesophagus is roughly cylindrical. Buccal Capsule. — The buccal capsule is bent dorsally, 0*2 1 mm. long, 0'i9 mm. broad. If a worm is rolled under the cover-glass so that the dorsal side is upwards, we observe the following features (fig. 325) : In the dorsal edge of the chitinous capsule there is a gap as if a U-piece had been punched out. This is the " dorsal gap or incision." The so-called ^' dorsal teeth " are simply the rounded edges of the tips of this gap. They project beyond the skin which covers the capsule externally. Below this gap is seen a curved line which, if followed along the sides of the capsule on each side, merges into the base of the most posterior ventral tooth. This line is the optical expression of a very shallow groove on the inside of the capsule. The skin on the outside of the capsule, which is reflected over the edge of and into the capsule, dips into this groove, which gives it a firm attachment. Below the middle (dorsally) of this curved line there is a thickening in the capsule wall, which is perforated by the opening of the dorsal oesophageal gland. This is the ''dorsal ridge" ; in optical section it has a conical appearance with a lumen (of the duct). ' I.e.^ with a distinct space between the limbs. * Strongylus (Syn. : Sclerostomum) differs slightly in its posterior ray from the other genera of the group. Each bifurcation is trifurcate rather than tridigitate. 446 THE ANIMAL PARASITES OF MAN On the ventral wall one sees the two pairs of strong teeth, their points being directed somewhat backwards. They are covered by cuticle above and below, but their points are free, piercing the cuticle. Gh Cdr Sp Gcph Ut I. ^ Bni- Fig. 323. — Ancylostonia duodtjiaie, male. B, bursa ; Bm, bursal muscles ; Cdr^ cement gland surrounding the ejacu- latory duct ; Glc, cervical glands ; N, nucleus of cephalic gland ; AV, nerve ring ; Ty testes ; Sp, spicule ; Vs, vesicula semina- lis. X 15. (After Looss.) Fig. 324. — Ancylostoma duodenale, female. Ay anus ; Gcph, cephalic gland ; Ny nucleus of cephalic gland ; Glc, cervical gland ; Ov, ovary ; Pex, excretory pore ; Rs, receptaculura seminis ; Ui, uterus ; F, vagina, x 15. (After Looss.) The "roots" of these teeth followed backwards appear as two thickenings or ribs on the outside of the capsule w^all, so that the outside wall is not smooth — a characteristic of the genus Ancylo- stoma. In the space between these ribs lies the ventral nerve papilla, ANCYLOSTOMA DUODENALE 447 and lying against the outside of the outer root the lateral nerve papilla. The nerve papillae are thus, as it were, concealed by these roots, and not conspicuous as they are in Necator. Following the ventral curve of the capsule on the inside, posteriorly we next find two triangular ventral lancets.^ These stand straight up into the capsule on either side of the longitudinal axis, converging at their summits. So that to sum up, the cutting apparatus is entirely ventral, consisting of two pairs of cutting teeth and a pair of lancets. Cervical Papillcv. — Two, one on each side behind the head at the level of the excretory pore. They consist of ''pulp," i.e., extensions of the substance of the lateral bands covered by cuticle and supplied with a nerve (fig. 326). CEsopJiageal Glands (3). — The chitin of the triradiate oesophagus is continuous with that of the buccal capsule. In its muscular walls are three glands — one dorsal, two sub- ventral. The dorsal gland opens into the buccal cavity through the dorsal ridge ; the two others into the lumen of the oeso- phagus at the nerve ring. They branch freely amidst the muscle? They are probably digestive in function. Cephalic Glands (2). — Lie in the lateral lines or bands on either side. They begin about the middle line of the body, and their ducts open at the base of the outer ventral tooth on the surface of the skin on each side. Each is 0*15 mm. thick in the middle, and has a single nucleus about as big as an ancy- lostome egg (N, fig. 323). They probably function as poison glands. Excretory System and Cervical Glands (2). — The excretory pore lies in the mid line ventrally behind the oesophageal nerve ring (figs. 324 and 326). It opens into the excretory vesicle, a cavity in a large cell with lateral appendages which fuse with the lateral lines, this cell thus forming the '' bridge " of the excretory system. Adhering to this (bridge) cell are the spindle-shaped cervical glands (G/c, fig. 324), and branches from the excretory vesicle enter the glands, which are excretory in function ; the vesicle also receives branches Fig. 325. — Ancylosto7na duodenal e : showing ventral teeih, dorsal cleft, and behind it the dor- sal ridge with duct of dorsal oesophageal gland. X c. 200. (After Looss.) The ventrallancet (of one side) of Necator is seen in fig. 335. 448 THE ANIMAL PARASITES OF MAN from the lateral excretory canals (fig. 326) running in the lateral lines or bands. The cervical glands are swollen anteriorly, forming the so-called ampullae just in front of the bridge. They extend backwards a little beyond the anterior loop of the testis. Lateral Lines. — (i) Are broad elevations of the subcuticle, in which, here and there, a nucleus occurs. (2) Near the bursa in the male they increase in volume, and finally divide into branches which form the "pulp" of the different rays. (3) In addition to the lateral lines or bands, there is also a dorsal and ventral band. (4) The 7*» Fig. 326. — Ancylostoma duodenale : diagrammatic representation of excretory system ex.p., excretory pore; e.c.g., excretory cervical gland ; Ex. ves., excretory vesicle in B.c.y bridge cell, which is connected with c.g.^ cervical gland, and /./., lateral lines; ceph.g.^ cephalic gland ; l.ex.c, lateral excretory canal passing into the bridge cell ; /./., lateral line containing excretory canal and cephalic gland; c.p., cervical papilla; w., nuclei of bridge cell. (After a drawing of Looss.) ventral band is well developed caudally, forming a large pad dorsal to the cloaca, "pulvillus post-analis." The bursal rays are outgrowths of the lateral lines. Beside this ^' pulp " they contain a nerve, and at their bases complex muscles. The Bursa is closed on all sides with a short median (ventral) lobe, which may be tucked inwards. It is an outgrowth of the inner layer of the skin pushing the outer layer before it, so that it consists of three layers, not four, as it would be if it were a fold. The bursa is twice as broad as long. It is supported by a variety of rays, the arrange- ment of which is best followed from the figure (fig. 327). The ANCYLOSTOMA DUODENALE 449 different terminology for these rays as used by various authors should be noted : Ventral = anterior ; externo-lateral =: antero- external ; medio-lateral + postero-lateral or antero-median + postero- median = median (doubled); externo - dorsal = postero - external ; dorsal = posterior. All the rays end in tactile papillae, seven, on each side ; the postero-external and antero-external on the outer surface of the bursa, the five others on the inner surface/ Of the six terminal digitations of the dorsal ray, only the external two contain tactile papillae. In the male there are prebursal papillae and minute caudal papillae in the female. In the female the inner layer of the cuticle projects at the posterior end as a sharp spike, 20 /x long, which may sometimes be broken off. Ovaries. — The anterior tube runs from the cephalic to the posterior end and back again. The posterior tube begins anteriorly, runs to the Ci/e Fig. 327. — Ancylostoina duodenale : bursa enlarged. Ca, anterior rpy cleft ; cle, antero-external ; els, antero-median ; dp, postero-median ; Cde, postero external ; Cd, posterior bifurcated, each bifurcation tridigitate. After Railliet.) posterior end of the body, and then back to the cephalic end, form- ing a vulval loop before ending. The ovaries on the whole run in oblique coils. The uterus is the thicker portion of the tube, 5 mm. long. A short tube connecting the ovary and uterus is the oviduct. The two uteri unite to form a single duct, the vagina, opening i mm. behind the middle line. The portion of the uterus next to the oviduct functions as a seminal receptacle, whereas the part next the vagina functions as an ovejector. Testis. — The blind end begins a little behind the beginning of the cement gland. The transverse coils occupy the middle third of the body. About the middle of the body it passes into the spindle- shaped seminal vesicle, which, with the spicular canal and rectum, opens into the cloaca. An anterior longitudinal coil pushing in between the cervical glands is characteristic of Ancylostoma. The cement gland surrounds the ejaculatory duct for practically its whole course, and it occupies nearly the posterior half of the body and 1 This also occurs in other Strongylida, e.g., in the genus Strongylus (Syn. : Scleroatomum). 450 THE ANIMAL PARASITES OF MAN secretes a brown or black cement. The spermatozoa are curved rods about 2 fi long. Spicules are 2 mm. long, ending in a fine point. They are moved by exsertor and retractor muscles. At first they lie free in the body cavity; next in a groove in the dorsal wall of the cloaca; then in an isolated canal, and finally in two canals. Anteriorly each has two longitudinal crests on its inner surface. These meet the correspond- ing crests of the other spicule, and so form a canal along which the sperm passes into the female. The gubernaculum is a thickening of the dorsal wall of the cloaca. It is not a free piece, but is moved by various muscles. Fk;. 328. — Ancylostoma duodeiiale : bursa of male. The rays from left to right are: (i) anterior clef t ; (2) antero-external ; (3) and (4) median doubled, i.e., antero-median and postero-median ; (5) postero-external arising from a common trunk with the posterior. X c. 120. (After Looss.) Genital Cone is a prominence on the floor of the bursa on the ventral side of the body, on which the genito-anal orifice opens. The cone is only slightly marked in Ancylostoma duodenale, but is much more prominent in Necator aniericanus. ^ Distribution. — Africa, Egypt, Europe,, Japan, China (mainly), but in association with Necator aniericanus in Southern States of America, British India, Assam, Burma, Hongkong, Liberia, Jamaica, Martinique, Costa Rica, Colombia, Antigua, Guadeloupe. Habitat. — The worms live in the jejunum, less frequently in the duodenum, of man only. Food. — The worms feed on the mucous membrane of the gut," attaching themselves to the base of the villi, sucking these in; and when these are destroyed they attack further the submucosa. As a rule the worms have no blood in the gut, but in their attack on the submucosa a blood-vessel may be eroded, and so the gut of the worm filled with blood. ANCYLOSTOMA DUODENALE 451 Development. — The eggs are oval with broadly rounded poles, 56yLt to 61 fx by 34//, to 38 yu-. In fresh faeces they contain four granular nucleated segmentation masses of the ovum (fig. 329) separated by a clear space from the shell. Egg of Ancylostonie appears to have a single contour. Under high powers this appears double, but they are the outer and inner surface of the true (chitinous) egg-shell. Internal to this is the extremely delicate yolk-envelope, a kind of skin secreted by the Qgg cell around itself for protection. The function of this is probably to absorb water to swell and burst the outer chitinous shell. The embryos when hatched are termed larvae. Embryos which are ready to hatch have their bodies almost free from granules ; others, though apparently mature, that have granules will not hatch. ^ a h c d Fig. 329. — Ancylostonia auodenale : eggs in different stages of develop- ment, a to c, in fresh faeces ; d, containing a larva, only in old faeces. X 336. (After Looss.) Larva. — Stage I : Average length, 25 mm. Maximum thickness in oesophageal region, 17 fi. Head end fairly blunt, from behind the anus (the tail) tapering in an uniform manner. Buccal cavity is characteristic, 10 /jl to 12 fi by 1 fi to S /jl, longer and narrower than the corresponding larvae of Strongyloides stercoralis. CEsophagus " rhab- ditic " in character, i.e.y it has three sections, but they are not so clearly marked off as in larvae of the genus Rhabditis. The posterior bulb has a Y-shaped valve, the function of which, according to Looss, is to prevent regurgitation of food. The granules of the gut serve as a reserve of food, and are used up if the larvae are starved. The genital rudiment consists of two cells half-way between the end of the ' Table of Differences between Larv^ of A. duodenale and S. stercoralis. (i) Vestibulum oris (2) Genital rudiment (3) Thickness (4) CEsophagus ... (5) Tail (6) Motion (7) Gut A. duodenale i'8 fi broad 3 fi to 5 /i long Thicker One-fourth body length Pointed Less active than Soon fills with dark granules S. stercoralis 25 /* to 33 /t Half body length Two fine points Rhabditiform. Filariform. 29 452 THE ANIMAL PARASITES OF MAN oesophagus and the anus in the mid-ventral Hne. The larva lives on f^cal matter and grows to about 0-4 mm., then moult^ I takes place in two days or more, the skin being ruptured by the activity of the larva. Stage II : The larva is now in this stage, which does not differ much from the previous one. It grows to 0-5 mm. The mouth Fig. 330. — Ancylostoma duodenale larva on fourth day of culture on right Strongyloides stercoralis larva on left. (After Leichtenstern.) Opening closes. The oesophagus elongates, becoming cylindrical or " filariform " ; a new skin is formed underneath the old one, and in about a week moult II takes place. Stage III : The mature larva remains enclosed in the old skin. Its movements are now much more active and of a boring character. Length is now o'6 mm. This mature stage has been erroneously called the encysted larva, because there is no cyst secreted from its ' Moults take place by the formation of a new skin below the old one, the two being in close apposition at first. ANCYLOSTOMA DUODENALE 453 surface by the larva, but it is simply the old skin, which is not cast off, but is retained for purposes of protection, as the larva is free living, but casts it as soon as it assumes parasitic life again. From the egg to this mature stage is thus six to ten days. Fig. 331. — Ancylostoma duodenale: left, four days after transmission into dog, 190/1 ; in the centre, at the commencement of the second stage of develop- ment (five to six days), 105/ 1 ; on the right, fourteen to fifteen days after trans- mission. 42/1. (After Looss.) Bionomics of Development. — Air : Eggs can develop when shut off from the air for a '' comparatively long " time. Temperature : Hatching takes from eight hours upwards. Eggs develop best at 25° to 30° C, but will not develop below 8° to 10° C. The larvse, however, will stand freezing. 454 THE ANIMAL PARASITES OF MAN Moisture : Eggs and larvae do not live long under water, because they suffocate or starve, but mature larvae will live for months (six to twelve) in water ; they require no food — in fact, can take none in — but live on their reserve granules, and in course of time become as clear as glass. Thiguiotropisni : The mature larvae, after casting their skin, will penetrate pith, climb up stems, stalks, etc., and creep into any pore. It is important to recognize that this third stage of the mature larva is the only infective one. Mode of Entry into the Body. — Infection is effected through the mouth (Leichtenstern and others), and also through the skin, as was first discovered by Looss and afterwards confirmed from the most diverse quarters, partly in the case of Ancylostoma duodenale, partly in that of A. caninuni in dog, man, and monkey. The larvae that gain access to the intestine partly through contaminated food, or through unwashed hands, or under some circumstances through water, first throw off their " sheath " — that is, they complete moult II. Moult III takes place four to five days after they have reached the gut, and they now have a mouth capsule supplied with four small teeth arranged crosswise, enabling them to fasten on to the intestinal epithelium, upon which they feed. On this food the worms grow in four to six days to 3 to 5 mm. in length, and now moult IV. takes place, thus attaining their definite shape and distinctive character. About eight days later the sexual organs commence to function ; at this time the first copulation should be taking place — it will later be frequently repeated — and a few days later the first ova are laid, first in less and later in larger numbers, so that they appear in the faeces about four to five weeks after the infection.^ Infection by the Skin. — Mature larvae, which are placed on the skin of man or suitable animals, cast their ^' sheath " and bore their way through delicate fissures either horizontal in the superficial scales of the epidermis, or through vertical fissures into hair follicles where these exist, and then they invade the cutis. Now according as they migrate further into the lymphatic vessels or the small vesicles, the final path to the gut differs to some extent. The blood path leads to the right heart, and from there into the lungs ; here the larvae leave the blood stream and enter the air passages, over the mucosa of which they travel further headwards, through the bronchi into the trachea and larynx, and from hence through the oesophagus to the stomach ; in some cases also they are swallowed. The ' From the number of eggs present in a given quantity of faeces, the number of female Ancylostomes present in the gut can be reckoned by a formula of Leichtenstern's (x = ^'L, in which a signifies the number of eggs counted in a single gramme of fseces). ANCYLOSTOMA DUODEXALE 455 lymphatic path leads finally also into the blood stream, but the lymphatic glands must first be passed, and in these many larvae are retained and perish. In the cutaneous infection seven to ten weeks elapse till the time of appearance of the first ova in the faeces. The penetration of the skin by the larvas also in man causes reddening and burning at the affected points, and this is followed in a few days by transitory swell- ing in the subcutaneous connective tissue. Skin affections can also be set up by such Ancylostoma (and Strongyloides) larvae as do not gain access to the blood or lymphatic vessels or gut ; such larvae apparently wander further in the connective tissue, and, as Looss has in his own person observed, gain access to the cutis at different points, thus causing progressive swellings (accompanied by intense itching), which cease when the worm again penetrates into the deep tissues. Skin affections such as " ground-itch " or " pani-ghao " occurring in the tropics and only attacking the feet, or other affections (^.^., sump bunches) are now well recognized as being due to the invasion of Ancylostoma larvae. Other names for these skin affections are water-sore, sore feet of coolies, maza- morra, bunches, botches, quaddeln, kratze, ampoules, gourmes, taons, pitirr. Whether oral or dermal infection is the more important one further observation must decide. The duration of life of Ancylostotna duodenale, which is a specific parasite of man and has not been observed in other mammals, amounts to about five years, as strayed larvae according to Looss wander for this extent of time in the body. Cultivation of Larvce. — (i) Mix the faeces (free from drugs such as salines or thymol) with animal charcoal, adding water if necessary till a consistence of porridge is obtained. If the stools are very fluid, allow to sediment first and pour off the fluid. The best charcoal is that made from bones, and should not have an acid reaction. Char- coal is necessary in order to prevent fermentation, which kills the larvae. Spread in layers 2 to 3 mm. thick in Petri dishes. Incubate at room temperature. To extract the larvae from the culture allow the surface thoroughly to dry, then pour on water ; the larvae wander out and are poured off and subsequently further purified by sedimentation or filtering through blotting paper, the larvae passing through. (2) A funnel is plugged with cotton wool, then filled with washed sand to within a centimetre or two of the rim. Stand this in a jar of water so that the level of the water is slightly below that of the sand. On the surface of the wet sand now place layers of blotting paper, and spread the faeces, diluted if necessary, on this in layers of a few millimetres thick {vide p. 474). Detection of Eggs. — Vide p. 473. Dermal Infection of Dogs. — Infection with larvae of A. caninum. In two hours most of the larvae are free in the cutis and in four hours in the subcutaneous tissue. By scraping a few days later the mucosa of the trachea large numbers of larvae are found there. 456 THE ANIMAL PARASITES OF MAN Ancylostoma ceylanicum, Looss, 191 1. At the anterior edge of mouth capsule one large tooth ; below or behind this towards the middle line a very small tooth, the tip only of which is seen. Male 5 mm. average. Lobes of bursa almost as long as broad, strongly projecting towards the ventral side. Rays short and relatively thick. Female 7 mm. Habitat. — Intestine civet cat {Viver- riciila nmlacensis), Ceylon, and man in Bengal according to Clayton-Lane. Other species are : A. caninum (Ercolani), in cat and dog, Europe and Africa; A. inalayamun (Ales- sandrini), 1905, in the Malay bear (Helaretos lualayanus) ; A. pluriden- tatmn (Alessandrini), 1905, in Felis Fig. 332. — Ancylostof/ia ceylanicum : head end, two teeth on each side, the inner almost concealed by the outer. X c. 200. (After Looss.) niitis, Brazil. Ancylostoma braziliense, Gomez de Faria, 1910. In cats (and dog), Brazil. Female 8*5 mm., male 7*5 mm. long. Eggs 65 fi by 32 /jL. Leiper considers it to be identical with A . ceylanicum. Fig. 333.^ — Ancylostoma braziliense: bursa of male. (After Gomez de Faria.) Group. Bunostomeae, Railliet and Henry, 1909. Bursa with median double, postero- and postero-external arising from a common trunk, posterior bifurcated, each limb bidigitate (fig- 336)- Vulva in middle of body or a little in front. Uteri divergent. NEGATOR 457 Contains the following genera : (i) Bunostomum (= Monodontus) ; (2) Necator ; (3) Bathmostomum ; (4) Gaigeria. Genus. Necator, Stiles, 1903. Mouth capsule small, narrowed anteriorly (ventrally) by chitinous plates, as in Uncinaria. On each side of the base of the dorsal cone a lateral chitinous plate or lancet with smooth edge (not serrated), ventral lancets as in Ancylostoma. No ridges on outside of ventral wall. Aperture of dorsal oesophageal gland on tip of a cone pro- jecting freely into the buccal capsule. Bursa closed. Posterior ray cleft to its root. Necator americanus, Stiles, 1902. Syn. : N, africanus^ Harrison, 1910. Male 8 mm. long, female 10 mm. The head is strongly bent dorsal- wards so that almost by this character alone it can be distinguished horn Ancylostoma duodenale. The buccal capsule is markedly small — Fig. 334. —Necator americanus. Showing cutting plates and the projecting dorsal ridge, and deep in the cavity the edges of the ventral lancets, x c. 475. (After Looss.) in the male, 0*093 by 0*084 mm., in the female o'li by 0*097 mm. There are no teeth anteriorly on the ventral side of the capsule, but instead there are two cutting chitinous plates, the anterior portions of which are prominent and angular, and meet in the middle line in front. Posteriorly on each side the plate projects less, while between the anterior and posterior parts there is a deep angle. The inner (posterior) ventral lancets which also occur in A. duodenale are large, and project far into the lumen, the tips of these, of the lateral lancets, and of the dorsal cone almost meeting in the centre of the lumen. As already stated in the definition of the genus Necator, there are also lateral lancets which start from the base of the dorsal cone. This dorsal ridge, or rather in this case cone, is a striking object in 458 THE ANIMAL PARASITES OF MAN the mouth, and projects right out into the cavity, and on its summit opens the dorsal oesophageal gland. Fig. 335. — Necator americamis : lateral view, showing the dorsal ridge per- forated by the duct of the dorsal oesophageal gland, the lateral lancet and ventral lancet and the nerve papillae, x c. 475. (After Looss.) The bursa is about as long as broad, but has the lateral lobes strikingly lengthened, giving a trilobed appearance (fig. 336), but as in Ancylostoma diiodenale it is closed on the ventral side. The distribution of the rays is best understood from the figure. The genital aperture lies on a marked conical pro- tuberance ; the cement gland is bilobed in transverse section. In the female the opening of the vulva is in front of the middle line, in A. duodenale it is behind. The spicules, 0*92 mm. long are hooked at the extremity. Eggs more pointed at the poles than those of A. duoden- ale, 6^ Ijuio ^2 fi by 36 yLfc, so that it may not be possible to dis- tinguish single eggs owing to individual variations, yet on comparing a number they can be distinguished. Fig. 336. — Necator americanus : bursa of male. The rays from right (top) to left are: (i) pos- terior, (2) postero-external, (3) and (4) median doubled, i.e., postero-median and antero-median, (5) antero-external, (6) anterior (cleft), and above it on left a pre-bursal ray. x c. 120. (After Looss. ) SYNGAME^ 459 Geographical Distribution. — Brazil, Porto Rico, Cuba, Central Africa, East Africa, Victoria Nyanza, Gold Coast, Uganda, North- Western Rhodesia, Ceylon, Mysore. For other localities where A. duodenale is also found see p. 450. Habitat. — In small intestine of man and gorilla {Troglodytes gorilla). Necator exilidens, Cummins, 1912. Syn. : A^. africanus, Looss, 1911. Male 7 mm., female 9 mm. long. The edges of the cutting plates are rounded, not angular, and do not meet in the middle line. Inner (posterior) ventral lancets very small. Lateral lobes of bursa broader than long. Rays thick and plump. Habitat. — In the chimpanzee {Anthropopithcciis troglodytes), Ancylostomiasis. Morbid Anatomy. — Organs pale and bloodless. Abdominal organs sodden, and there is fluid in the serous cavities. Lungs: oedema. Kidneys : fatty changes, especially large pale kidney. Liver and heart also show fatty changes— there is much haemo- siderin in the liver cells. Blood : early stages, a leucocytosis 20,000 upwards, and eosinophilia 50 per cent. Later, anaemia (hydraemia). The number of worms found varies from ten to 1,000. They are rare in the duodenum, but occur as far as 6 ft. from the pylorus. Group. Syngameae, Railliet and Henry, 1909. Bursa with anterior and median ray cleft ; antero-external, close to median; postero-external, arising separately from posterior; pos- terior bifurcate to base, each branch bifurcate or trifurcate. Vulva in the anterior fourth of body. Uteri divergent. Genus. Syngamus, von Siebold, 1836. Head thickened, not tapering ; broad mouth with gaping buccal capsule. Male and female often in permanent copula. Parasitic in respiratory passages of birds and mammals. Habitat. — S. trachealis in poultry ; S. bronchialis in goose ; S. laryn- getis in cattle ; S. vasicola in goats, etc. Syngamus kingi, Leiper, 1913. Buccal capsules of male and female on same level. In S. trachealis and S. laryiigeiis, that of male in front of that of female. In S. dispar, that of male behind that of female. QCsophagus of male one-sixth, that of female one-ninth of total length. Mouth capsule in male and female terminal ; it is dorsal in S. trachealis and in mammalian 4t)0 THE ANIMAL PARASITES OF MAN Species. Tail of female bluntly pointed. Ovary reaches to anus. Excretory pore opposite the middle of the bulb of oesophagus. In S. trachealis it is opposite the oesophageal valves. Habitat — Found in sputum of patient by King in St. Lucia. Normal host probably a carnivore. Fig. 337. — Syngamus kingi : anterior end of male. (After Leiper.) Fig. 338. — Syngamus kingi : anterior end of female. (After Leiper.) Family. Physalopteridae. Genus. Physaloptera, Rudolphi, 1819. Mouth surrounded by two large lateral lips bounded posteriorly by a cuticular band projecting anteriorly, forming a collar. Each lip bears anteriorly and inwardly a cuticular appendage, the external tooth. Immediately below and internal to the external teeth the internal teeth, one on each lip. Each lip bears two large submedian papillae. Tail of male with four pairs of pedunculated papillae in a row on each side external to the six pairs of unpedunculated papillae. Spicules unequal. Vulva in the anterior region of the body. Eggs with a characteristic thick smooth shell. Parasitic in the intestine, more especially the stomach, of mammals (twenty species), birds (twelve species), reptiles (fourteen species). ASCARID^ 461 Physaloptera caucasica, v. Linstow, 1902. The male measures 14*2 mm. in length and 071 mm. in breadth ; the bursa is broad, rounded off in front and narrower at the back ; the right spicule measures 0*62 mm. in length, the left spicule 1*76 mm. ; there are two papillae in front of the orifice of the cloaca, four behind it and six unpedunculated on the tail. The female measures 27 mm. in length, 1*14 mm. in breadth ; the caudal extremity is rounded off ; the vulva is on the border of the first and second sixth of the length of the body; the eggs have thick shells, and measure 57 //, by 39 //,. It has hitherto only been observed once, by Menetries in the intestine of man (Caucasus). Fig. 339. — Bursa oi Synga??ius trachealis. a., anterior ray cleft ; a.e. , antero-external ; w.«.,antero-me(lian; w./.,postero-median ; p.e., postero-external ; /., one branch of posterior (trifurcate). (Stephens.) Physaloptera mordens, Leiper, 1907. Large worms resembling an im- mature Ascaris lumhricoides. The inner lancet-shaped teeth have a sharp cutting edge towards the lumen. Below each is a cuticular boss projecting into the mouth (fig- 340)- Male 30 to 50 mm., bursa with four pairs of pedunculated papillae, the second and third lying external to the first and fourth on each side. Spicules unequal, one slender (4-6 mm.), the other stouter (6 mm.). Female 40 to 55 mm. Tail sharp. Vulva opens between the anterior fourth and fifth of the body. Eggs 43*6//, by 35-3 //, with a thick smooth shell. Habitat. — (Esophagus, stomach, small intestine of man (several cases). Nyasaland and Portuguese East Africa. Family. Ascaridae, Cobbold, 1864. Sub-family. Ascarinae. Without oesophageal or intestinal diverticula ; spicules without flanges. Genus. Ascaris, L., 1758. Intermediate lips and auricles absent. Lips edged with fine teeth. Lips triangular in cross section. Not grooved on internal surface. 46: THE ANIMAL PARASITES OF MAN .; Slii.(». ves.Sim. Fig. 340. — Physaloptera mordens, Leiper, 1907. (1) adult male : o.e., oesophagus; ch.i.^ chyle intestine; t.c, testicular coils; ves. sem., vesicula seminalis ; sp.i, long spicule; sp.2, short spicule; B., bursa. (2) Mouth parts: r., cuticular collar embracing the two lips posteriorly ; c.b., cuticular bosses guarding the mouth laterally ; e.d., external tooth ; i.d., internal tooth ; stn.p., submedian papillae. (3) egg of P. caucasica. (4) egg of P. inordens. (5) bursa enlarged : fed.p., pedunculated papillae ; ses.p., sessile papillae. (After Leiper.) ASCARIS LUMBKICOIDES Ascaris lumbricoides, L., 1758. 463 The colouring, in the fresh condition, is reddish-yellow or greyish- yellow ; the body is of an elongated spindle shape. The oral papillae are finely toothed. The dorsal papilla carries two sensory papillae, the two ventral papillae each one sensory papilla. The male measures from 15 to 17 to 25 cm. in length, and about 3 mm. in diameter ; the posterior extremity is conical and bent hook-like ventrally ; the spicules measure 2 mm. in length, are curved, and somewhat broadened at their free end ; on each side around the orifice of the cloaca there are seventy to seventy-five papillae, of which seven pairs are post-anal. The testicular tube is much folded, showing through the body integu- ment, and is about eight times the length of the body. The female measures 20 to 25 to 40 cm. in length and about 5 mm. in diameteir ; the posterior extremity is conical and straight. The vulva is at the junction of the anterior and middle thirds of the body, which, at Fig. 341. — Ascaris lumbricoides. extremity of the male with the spicules pro- truding from the orifice of the cloaca {Sp.) ; b, anterior extremity from the dorsal surface, the two lobes of the pulp of the lip separated by the " saddle " ; ^, anterior extremity from the ventral surface; /*., excretory pore. (From Claus.) Fig. 342. — Ovum of Ascaris lumbricoides^ with shell and albuminous envelope. 400/1. this point, has a slight ring-like constriction ; the convoluted ovaries measure ten times the length of the body. The ova are elliptical with a thick (4 />t) transparent shell (fig. 342) and an external albuminous coating which forms protuberances ; the ova measure 50 /i, to 70 yu, in length, 40 ytt to 50 //, in breadth ; they are deposited before segmentation ; the albuminous coating is stained yellow by the colouring matter of the faeces, but is sometimes absent. The egg cell is unsegmented, it almost completely fills the shell, and its nucleus is concealed by the large amount of coarse yolk granules. Abnormal or unfertilized eggs also occur in faeces. They are distinguished by their elongated form (80 \ju by 45 />t), irregularly cylindrical, its contents consisting of refractive granules. Ascaris lumhricoides is one of the most frequent parasites of man ; it is distributed all over the inhabited parts of the world, and though it is particularly frequent in the warmer regions, yet it also occurs 464 THE ANIMAL PARASITES OF MAN in Finland, Greenland, etc. In temperate climates A. lumhricoides occurs most frequently in young children ; it is, moreover, more common amongst country dwellers than amongst the inhabitants of towns, but is not lacking in infants, adults and aged persons. As a rule only a few specimens are present in the intestine, but many cases are known in temperate zones in which several hundreds of worms have been found in the same patient. This species is particularly numerous in the negroes of Africa and America. It occurs also in the monkey, dog and pig {1 A. suilla). The parasite was known in ancient times ; the Greeks called it eXjjLLv^ crrpoyyvXrj, Plinius termed it Tinea rotunda, ater on it was named Luntbricus teres. The aaKapi^ of the Greeks is our Oxyuris. The small intestine is the normal habitat of Ascaris lumhricoides ; the worms, however, often leave this part of the intestine and wander into the stomach, whence they are frequently evacuated by vomiting, or they may creep through the oeso- phagus into the pharynx and crawl out through the nose or mouth ; very rarely they may find their way into the Eustachian tube or into the naso-lachrymal duct, or into the excretory ducts of the liver and pancreas ; exceptionally they may gain the trachea, and they have also been found in the abdominal cavity. They may bore through adhesions between the intestinal wall and the omentum (worm abscess) ; they occasionally penetrate the urinary apparatus and are passed with the urine ; in febrile diseases ^. /«;«MV<9/^«, apart. Cephalic wings long, narrow, semi- lanceolate. Male, 4 to 6 cm. Spicules, 1,002 y, and 1,005 M- Female, o'5 to 10 cm. Eggs, 75 /x to 85 /a, shell thick and smooth. Host : dog, occasionally man. Fig. 343. — Ovum Fig. 344. — Transverse section through the head of Toxascaris livi- part of Belascaris cati from the cat, with the lateral bata, with thin albu- wings. In addition, one n)ay note the four fields of minous envelope. muscles, the longitudinal lines with the oesophagus Magnified. in the centre. Magnified. (After Leuckart.) Genus. Belascaris (/Se\o9, an arrow), Leiper, 1907. Body anteriorly bent ventrally, cuticle coarsely striated. (Esophagus with a distinct bulb. Tail of male conical. A papillce- bearing protuberance behind the anus. Testis in anterior half of body. Vulva in anterior part of body. Eggs corrugated. Belascaris cati, Schrank, 1788. Syn. : Belascaris mystax, Leiper, 1907 ; Ascaris mystax. Striations 12 /x to 16 /i, apart. Cephalic wings lanceolate. Male 3 to 6 cm. Spicules 17 to 1*9 mm. Female 4 to 10 cm. Eggs, 65 /t to 75 //< in diameter, surface finely honeycombed. Host : domestic cat, and man, eight or nine cases. Belascaris marginata, Rudolphi, 1802. Striations 16 /a to 22 yu- apart. Cephalic wings long, narrow, semi- lanceolate. Male, 5 to 10 cm. Spicules, 750 /x and 950 /x. Female, 9 to 18 cm. Eggs, 75 /A to 80 II. Shell finely honeycombed. Host : dog. Genus. Lagocheilascaris, Leiper, 1909. Thick lips separated by a furrow from the body ; between the lips small intermediate lips without "pulp." The cutting angle of each lip bifurcated. Along each lateral line a cuticular wing extending the whole length of the body. Eggs, thick shell with a mosaic pattern. OXYURID^ 467 Lagocheilascaris minor, Leiper, 1909. Male, 9 mm., tail sharply curved. Spicules colourless, 3*5 and 4 mm. long. More than twenty-four pairs of pre-anal papillae, at least five pairs of post-anal. Female, 15 mm. Straight posteriorly. Vulva 6 mm. from head with two lips. Eggs, 65 />t in diameter. Host : man, cutaneous abscesses. Trinidad. Family. Oxyuridae. Genus. Oxyuris, Rudolphi, 1803. Mouth unarmed. The three labial papillae are only slightly protuberant, the oesophagus is long and presents two well-marked bulbs. The vulva is in the anterior part of the body. Oxyuris vermicularis, Linnaeus, 1767. Syn. : Ascaris vermicularis^ L. ; Fusaria, Zeder, 1803. Colour white, the striated cuticle forms projections at the anterior end w^hich extend some distance back along the middle of the ventral and dorsal surfaces ; the longitudinal lateral flanges of the skin corre- sponding to the lateral lines are well seen in transverse sections ; there are three small retractile labial papillae around the mouth. The male measures 3 to 5 mm. in length, and shortens on death; the posterior extremity of the body is curved ventrally and presents six papillae. Spicule 70/x, long, hook-like. The female is 10 mm. m length and 0*6 mm. in diameter ; the anus is about 2 mm. in front of the tip of the tail ; the vulva is in the anterior third of the body ; the eggs are oval, asymmetrical, with double-contoured shells, and measure 50 ^ to 55//, by 16 fjb to 25//,; they are deposited with clear, non-granular tadpole-like embryos already developed. Habitat. — Adults in large intestine of man. Young forms in small intestine and often in the appendix. The worm lives in the lower part of the small intestine, caecum and vermiform appendix, and before becoming adult undergoes two or three moults (Heller). According to Wagener the worms at times live in the gut wall, giving rise to calcareous nodules. When the uterus of the fertilized females begins to fill with eggs they leave the caecum and travel through the colon to the rectum. The uterus is now packed with eggs which contain a tadpole-shaped embryo. Egg-laying now takes place, partly in the rectum, partly outside, the mode of exit being not only passive through defsecation but also an active one on the part of the worms when the patient is in bed. In this case the worms crawl out of the anus, producing a most intolerable itching as they scatter their eggs between the nates and the perinasum. From here in the case of girls they may get occasionally into the vulva and vagina, and even into the oviducts and so into the body cavity. The worms also may wander through the alimentary canal in the opposite direction, getting out occasionally through the mouth. Recently a role has been assigned to them, as to other gut parasites, in appendicitis and typhlitis. 30 468 THE ANIMAL PARASITES OF MAN It is stated that the males die after fertilizing the females, thus explaining why they are so rarely met with in faeces [but it is probable that they often escape notice from their small size.— J. W. W. S.]. Development. — The eggs, which often adhere together, contain a tadpole-hke embryo, the thin tail of which is bent upwards ventrally ; Fig. 345--^, male, and B, fe- male, of Oxyuris vermicularis. 5/ 1 . Fig. 346. — On the left, female ; on the right, male. A, anus ; M, mouth ; F, vulva. Greatly enlarged. (After Glaus.) Fig. 347-— Oxy- uris vermicularis : egg freshly deposited, vk^ith tadpole-like embryo, x 640. Fig. 348. — Oxy- uris vermicularis : egg twelve hours after deposition, with ne- matode-like embryo. X 640. the embryo in a short time, given a sufficiently high temperature, passes into a second folded nematode-like embryonal stage, lying in the egg- shell, either in the faeces, with which also numerous females pass out, or in the moisture of the groove between the buttocks, and they there await the opportunity of being reintroduced into man per os. It is MERMITHID^ 469 very improbable that infection takes place directly in the large intestine, as is occasionally stated, because although the harbourers of Oxyuris are frequently liable to auto-infection, this takes place exclusively through the mouth, and is conveyed by the fingers, on which the ova of Oxyuris, and occasionally the female worms, have clung. The opportunity for this is afforded every evening, as naturally the troublesome itching caused by the wandering of the worms is met by scratching and rubbing with the fingers. It is therefore possible that the eggs may even thus be introduced into the nose, where the young Oxyuris are perhaps hatched out, if they get high enough up on the moist pituitary mucous membrane. As a matter of fact, the larvae of Oxyuris have been found in the nose. Moreover, one can understand that the eggs of Oxyuris are transferred from person to person by the hand, directly or indirectly. This again explains the wholesale infections which occur in collective dwellings, after a person harbouring Oxyuris has been admitted into boarding-houses, etc. The primary infection may be also caused in other ways — by foods, fruits, vegetables and other articles that are eaten raw, and are polluted with the ova. Perhaps also flies or their excrement play a part in the distribution of the parasite, similar to that demonstrated by Grassi as taking place in the spread of the ova of Trichocephalus and Taenia. The assumption of a direct development without an intermediary host was first substantiated by Leuckart by experiments on himself and three of his students ; about fourteen days after swallowing the eggs the Oxyuris has attained 6 to 7 mm. in length ; Grassi, and later on Calandruccio, infected themselves by swallowing adult female Oxyuris, with the same results. Heller found worms in the gut {appendix vermiformis) of a male child five wrecks old. Other species are : O. co7npar in the cat ; O. curvula and O. mastigodes in horse, ass, mule ; O. ambigua in the rabbit ; O. poculunt in the horse ; O. tenuicauda in the horse. Many species occur in insects, especially in Blattidce and Hydrophilidce (aquatic beetles). Family. Mermlthidae. Genus. Mermis, Dujardin, 1845. With characters of the family. Mermis hominis oris, Leidy, 1850. Fourteen centimetres in length, o-i6 mm. in breadth; mouth terminal ; posterior extremity obtuse and provided with a recurved hook 50 /^ long. The parasite was " obtained from the mouth of a child." Stiles considers it to be probably a Mermis, possibly swallowed in an apple. 470 THE ANIMAL PARASITES OP^ MAN Agamomermis, Stiles, 1903. Group name for immature Mermithidce. Agamomermis restiformis, Leidy, 1880. This worm measures 65 cm. in length, pointed anteriorly, the posterior extremity broadened and rounded off (1-5 mm. in breadth) ; the mouth is terminal, without lips. Behind the mouth six papillae ; the oesophagus measures i'i25 mm. in length ; the intestine appears to terminate blindly. This parasite was obtained in West Virginia from the urethra of a young man, aged 20, who for a few days previous to expelling the worm passed turbid and bloody urine. TECHNIQUE 471 TECHNIQUE. Preservation and Examination of Flukes. Fixation. — (Method a.) (i) Place the flukes in a test tube or small bottle a quarter full of normal saHne. Shake the contents as hard as possible (the object of this is to extend the flukes) for half a minute. (2) Add immediately an equal bulk of saturated aqueous solution of corro- sive sublimate and shake again as vigorously as possible for a few minutes. (3) Transfer when convenient to 70 per cent, alcohol. (Before staining and mounting remove the sublimate with tincture of iodine.) (Method B.) In case of large flukes, e.g.^ Fasciola hepatica, Fasciolopsis buskij compress the flukes between two glass slides with rubber bands or thread. Fix in sublimate or in absolute alcohol, or in 10 per cent, formalin. (Method c.) Place the flukes in 10 per cent, formalin solution. Staining is successfully effected by using quite dilute solutions of carmine or haematein overnight. This is far preferable to using strong solutions, as it may be almost impossible to remove a too intense stain. Almost any dilute carmine solution suffices. One of the best is acetic-alum carmine (boil excess of carmine in a saturated aqueous solution of potash-alum for about fifteen minutes ; add glacial acetic acid to the extent of 10 per cent. ; let it stand for a week ; filter). For use, dilute about thirty times with water. Place the flukes directly in the stain. Stain overnight or longer. Differentiation, — In order to get the sharpest picture, it is best now to differentiate (but this may often be omitted) with acid alcohol (70 per cent, alcohol 100 parts, HCl 5 drops). Allow to act from one to twenty-four hours, according to the appearance of the flukes. Similarly, in staining with hematoxylin solution, dilute twenty to thirty times so that the water is merely tinged with the stain. Differentiate as before. After staining, dehydrate, clear, and mount in balsam if required. Clearing and Mounting. — (i) Carbolic acid (carbolic acid 94, water 6) is a very convenient clearing agent. It may be used for stained or unstained specimens. It will clear rapidly without previous dehydration. If it is required to mount a specimen permanently, transfer from carbolic to alcohol, then cedar-wood oil (or xylol, etc.), then balsam. (2) Creasnte. — Dehydrate the specimen, stained or unstained, transfer to creasote. If it is desired to mount permanently, transfer back to alcohol, then cedar-wood oil, then balsam'. (3) Cedar-wood Oil. — Preferable to xylol or oil of cloves. Dehydrate the specimen in alcohol. To mount permanently, transfer to balsam. 472 THE ANIMAL PARASITES OF MAN (4) Glycerine. — Vide under methods of preservation of ova; to mount permanently, transfer to glycerine jelly ; subsequently to harden the jelly, expose to formalin vapour. Of these media, carbolic acid has the greatest refractive index excepting that of balsam. The latter may, in some cases, render structures too transparent, and it may be advisable to use only glycerine or glycerine jelly. Preservation of Ova in F^ces, Urine, Bile, etc. Heat some 70 per cent, alcohol in a basin to about 60 to 70° C. (until bubbles begin to appear). Add the faeces, etc., in the proportion of one part to about nine of fixative ; keep stirring. Allow the sediment to settle. Transfer to a bottle with some fresh 70 per cent, alcohol. Transference to Glycerine. — Prepare 5 per cent., 10 per cent., 20 per cent, solutions of glycerine in 70 per cent, alcohol. Pour off the alcohol in the bottle of faeces, etc., and replace by 5 per cent, glycerine solution. Allow to stand an hour or so. Then in the same way replace the 5 per cent, by a 10 per cent, glycerine, and finally by a 20 per cent, glycerine solution. When in this latter expose freely to the air (protecting from dust), so as to allow the alcohol and water to evaporate. Add a few drops of glycerine from time to time till eventually the ova are in pure glycerine. (In a very moist climate it may be necessary to use lime or calcium chloride to dry the air.) To mount permanently transfer some of the sediment to glycerine jelly. Preservation and Examination of Cestodes. Fixation. — (i) Saturated aqueous corrosive sublimate. — Add to this glacial acetic acid to the extent of i per cent. (Note this fixative will dissolve the "calcareous corpuscles"; 10 grammes of sublimate to 160 c.c. of water will give a saturated solution.) Warm the fixative to 70° to 80° C. (Avoid the use of needles.) Use plenty of fixative. Allow to act for a quarter of an hour or so. (a) Transfer to 70 per cent, alcohol. (It is advisable to remove the sublimate by the use of Lugol's solution, or a solution containing tincture of iodine, adding this until the iodine colour is permanent.) Or (b) transfer for preservation to 10 per cent, formalin. Or (2) 10 per cent, formalin. — In order to prevent contraction it is advisable to extend the tapeworm and keep it fixed by glass plates, or wind the worm around a wide glass tube or bottle, and then fix it. Or (3) fix in hoi alcohol. Staining. — As under flukes. It is necessary to sacrifice portions of the tapeworm for this purpose, cutting out, e.g., mature segments, so as to study the topography of the genitaHa. Clearing. — As under flukes. To examine the hooks satisfactorily it is best to cut off" the head with a sharp knife and mount. A certain amount of pressure is then advisable in order to view the hooks completely so as to measure them. TECHNIQUE 473 Preservation of Ova in F^ces, etc. As under flukes. Preservation and Examination of Nematodes. Fixation. — (i) Thoroughly wash the worms to get rid of mucus, etc., by shaking up in warm saline (or water) till the washings are clean. Then transfer to 70 per cent, alcohol heated to about 70° C. It is absolutely necessary to use hot fixatives in order to extend the worms. If no alcohol or spirit is immediately available, drop the worms into hot water, or saline, and transfer later to 70 per cent, alcohol. (2) Drop into hot 10 per cent, formalin. Cleaving. — (i) Carbolic acid, vide p. 471. (2) Creasote, vide p. 471. (3) Glycerine, vide p. 472. Staining. — In case of quite small Nematodes, e.g., Anguillulida, carmine may be used, but as a rule staining is not advantageous. Rolling. — In order to study the mouth parts, or bursa, etc., it is neces- sary to place the worm in any desired position. This is done as one would roll a penholder along the table by one's finger placed on top of it. In the case of a worm, one edge of the cover-glass is placed over the worm, the other is supported by a strip of cardboard. By tapping the cover-glass the worm will now revolve as much as required provided it is round and straight. In certain cases it may be necessary for this purpose to cut off the head or tail. Roll these separately. When a suitable position is got, the worm may be fixed in this position by pressure on the cover-glass, so as slightly to flatten it. Mounting the Head. — If it is required to get an end view, it is necessary to cut off the head transversely as near the end as possible, and then mount. Detection of Eggs (Bass and Hall). — Mix the faeces thoroughly with ten times the volume of water. Filter through gauze. Centrifugalize the filtrate. Wash the sediment and centrifugalize. Repeat twice. To sedi- ment add CaCl2 solution, sp. gr. 1250. The eggs float to surface. Pour off surface fluid. Dilute to sp. gr. 1050. Centrifugalize. Examine sedi- ment, which contains practically all the eggs in the stool. Detection of Small Nematodes. — Mix the faeces thoroughly with water. Allow to settle for five minutes. Carefully decant off, or better, syphon off the fluid. Mix the sediment again with water. Allow to settle. Remove the fluid. Repeat several times. Examine the sediment in a Petri dish. As the fluid is poured off, the worms will be seen collected in the backwater. Remove them with a brush. Fix in hot 70 per cent, alcohol. 474 THE ANIMAL PARASITES OF MAN Cultivation of Larval Forms of Ancylostoma and Strongyloides. A modification of the second method of Looss (p. 455) is that of Fiilleborn. A glass filter funnel is lined with linen or with cotton wool dyed black with iron-tannin. On this is placed a layer of sterile sand, and on top of this the faeces. The whole is moistened. The larvae hatch out and wander through the meshes of the wool, appearing on the edges of the same as white threads visible to the naked eye. With a platinum needle these can be easily removed. The glass filter can be placed on a glass cylinder, and this in another large stoppered cylinder containing caustic potash solution at the bottom, so that any larvae escaping from the funnel are killed. ACANTHOCEPHALA 475 D. ACANTHOCEPHALA, Rud. Gutless, neraatode-like worms that carry at their anterior end a retractile rostrum beset with hooks. In their adult condition they only live in vertebrate animals. During their larval stage they are often parasitic in invertebrate animals. The Acanthocephala are elongated cylindrical worms, with a rounded posterior end. In some species an annulation is distinctly recognizable ; they are, however, not segmented. The size varies according to the species, between about $ to lomm. and 40 to 50 cm. ; in general, however, there is a preponderance of the small species. The sexes are separate, and the males can easily be distinguished from the females without examination of the genitalia, as the females are both larger and thicker. The body wall of Echinorhynchus is limited by a thin cuticle, which is attached inwardly to the hypodermis. In only exceptional cases a syncytium with large nuclei, even in the adult condition, is represented by the hypodermis ; and in it fibre systems, the elements of which run in layers in various directions, appear, and it is only towards the interior from these strata of fibres that the nuclei of the hypodermis are found. As a rule, these fibres, at all events the radiary fibres, are regarded as muscles. Hamann desciibes them as elastic fibres, which lie in a viscid gelatinous connective substance (transformed protoplasm ?) ; a lacune system filled with a granular fluid, the central part of which are two longitudinal lacunes lying at the sides, also belongs to the cutaneous strata, as do the sc-called lemnisci, two short, flat organs suspended in the body cavity, and the pedicles of which are attached anteriorly at the border between the rostrum and body ; their structure as well as their origin permit them to be traced to the skin (fig. 348A). Finally, inwardly below the skin there follows a layer of annular, and after these a layer of longitudinal muscles, the structure cells of which remain present in the residues, carrying nuclei. The motor apparatus of the rostrum, the sheath of the rostrum, and the lemnisci also belong to the muscular system. The rostrum represents a finger-shaped hollow process of the cutaneous layer ; but, according to Hamann, it originates from the entoderm and passes through the skin secondarily. It is covered by a thin cuticle, and as a rule contains a large number of regularly placed chitinous hooks that adjoin a granular formation tissue. From the base of the rostrum springs a tubular hollow muscle extending into the body cavity ; this is the RECEPTACULUM PROBOSCIDIS, from the base of which again bundles of longitudinal muscles originate, which pass along its axis and that of the rostrum itself, and are inserted at the inner surface of its anterior end (RETRACTOR PROBOSCIDIS). These muscles when they contract invaginate the proboscis and draw it into the receptaculum ; when reversed they act again as PROTRUSOR PROBOSCIDIS. The whole of the anterior body, however, can be invaginaled, and for this purpose there is a muscle that originates from the body wall at a variable distance back, and which is joined to the receptaculum (retractor receptaculi) ; there is also a bell-shaped muscle which springs from the body wall behind the lemnisci in rings, and passes forward to the spot of attachment of the lemnisci. The nervous system consists of a cluster of ganglia situated at the base of the rostrum, from which three nerves pass towards the front and two towards the back. No sensory organs are known. The excretory organs, according to Kaiser, lie at the upper border of the ductus ejaculatorius in the male and at the so-called bell in the female. Here they 30- 476 THE ANIMAL PARASITES OF MAN represent the long-known villous tufts, placed on disc-like cushions. In each of the cylindrical villi — which terminate blindly towards the body cavity— there lies a cilium, which springs from the membrane lining the villus, and which lies in a space cavity of the villus, which ultimately proceeds as a little canal. There are three canals discharging into the uterus that serve to conduct the excretory materials from the body cavity ; special glandular cells corre- sponding to the terminal cells of the Platyhelminlhs, at the commencement of the system, are not present in the Acanthocephala, \T. Fig. 348A. — The male of Echinorhynchus augustatus. L.^ lemnisci ; T.y testicles; A, prostatic glands ; /*.?-., sheath of proboscis, with ganglion ; R.r.^ retractor of sheath of proboscis. 25/1. Sexual Organs. {a) Male Organs. — The greatest part of the male genital apparatus is contained in a muscular sheath^— the ligament — which originates at the posterior end of the receptaculum proboscidis, passes along longi- tudinally through the body cavity, and is inserted at the posterior end of the worm. The two oval testicles usually lie one behind the other ; their vasa efferentia unite sooner or later into a vas deferens which passes backwards, and finally terminates in the penis ; the terminal portion of the conducting apparatus is sur- rounded by six large glandular cells (prostatic glands) Fig. 348B. — ^Anterior portion of the female apparatus of Echinorhyvchus acus. On the left seen from behind, on the right seen from the front, /^.inferior orifice of the bell; B, bell; IJg, ligament; M, mouth of bell; Ut, uterus. Magnified. (After Wagener.) the excretory ducts of which open into the vas deferens. The penis itself is placed atthe base of a bell-shaped invagination of the posterior end, the bursa, which is everted during copulation. {b) Female Organs. — There are only two ovaries present in the ligament during the larval staged During the course of growth they divide into accumulations of cells (placentula^, loose or floating ovaries), which finally cause the ligament to burst ECHINORHYNCHUS GIGAS ^ 477 and they thus attain the body cavity. Thence a peculiarly constructed apparatus finally conveys the eggs out. This apparatus consists of the uterine bell and vagina, the latter discharging at the posterior extremity of the body. The bell is a muscular canal provided with apertures at both the anterior and posterior extremities. Its interior space is in direct communication with the body cavity, and the anterior orifice takes up all materials floating in the cavity— egg-balls, mature and immature eggs— and pushes these further backwards. The continuation of the bell lumen is now narrowed by a number of large cells in such a manner that only bodies of a certain form can pass through this tract and attain the uteius; everything else is conveyed back into the body cavity through the posterior opening of the bell. The eggs are already fertilized in the body cavity, and in this position go through their development to the formation of the embryo. Completely developed eggs are surrounded by three shells, and are generally fusiform. The eggs agglomerate in masses in the uterus until they are finally deposited through the vagina and vulva. For the further development, the transmission of the eggs into an intermediary host — usually a crustacean or an insect — is necessary ; the metamorphosis is very compli- cated ; but this transmission may be very easily effected artificially by feeding suitable crustaceans {Asellus, Gammarus, etc.) with the eggs of Aca7ithocephala ; this being the only method of inducmg the larva to hatch out so that its structure may be studied. The larva appears in the form of an elongated, somewhat bent body, at the stumpy anterior end of which there is a crown of hooks or spines, whereas the posterior end is pointed. Especial retractors draw in the hook-beset anterior surface, and an elastic cushion beneath them jerks them forward again when required. In the middle of the body a roundish heap of small cells is seen, from which the entire body of the Echinorhynchus originates, even to the cutaneous layer ; the latter is also the larval skin in which the small Echinorhynchus gradually grows. The development of all the organs takes place within the intermediary host, and the parasite only needs to be imported into the terminal host to attain the adult stage after a certain growth. In some cases, however, a second intermediary host is utilized. Species of Acanthocephala only occur exceptionally in human beings. Echinorhynchus gigas, Goeze, 1782. Syn. : Tcenia hirudinacea^ Pallas, 1781. The body is elongated, gradually decreasing in thickness towards the back. The rostrum is almost spherical, and is beset with five or six rows of recurved hooks. The males measure 10 to 15 cm. in length, the females 30 to 50 cm.; the eggs are provided with three shells, of which the middle one is the thickest. The eggs measure o'o8 to 01 mm. in length. The giant Echinorhynchus occurs especially in the intestinal cana^ of the domestic pig ; it is less common in other mammals. It bores deep into the mucous membrane with its rostrum, and causes an annular proliferation around the perforated spot ; occasionally also it causes perforation of the intestine. It is doubtful whether the giant Echinorhynchus occurs in man. Leuckart admitted that there were a few positive cases. According to Lindemann, Ech. gigas occurs in human beings in South Russia, and its presence is not rare. This statement, however, has not been confirmed. Its, ^^^' }^'^^'7~ ^^^ presence in man is by no means impossible, as its inter- 01 Echinorhynchus ^ .. , , / \ y, c 1 njr 1 1 ^i \ ^igas. 300/1. (After mediary host, the cankerworm, or cock-chater {Meiolontha)^ Leuckart.) is, according to Schneider, occasionally eaten raw by human 47^ THE ANIMAL PARASITES OF MAN beings. According to Kaiser, the golden beetle {Cetonia aurata) and, according to Stiles, another beetle in America {Lachnosterna arcuata) are also intermediary hosts. Echinorhynchus hominis, Lambl, 1859. This term is applied to an Echinorhynchus found by Lambl in the intestine of a boy who had died of leuccEmia; the worm was 5*6 mm. in length, and the almost spherical head was beset with twelve transverse rows of hooks. Echinorhynchus moniliformis, Bremser, 1819. The male is 4 cm. in length, the female 8 cm. long. This species lives in the intestine of field-mice, rats, marmots and Myoxus quercitius. A beetle {Blaps mucronata) is the intermediary host. This species has also once been artifically cultivated in man (Grassi and Calandruccio). a:f i-^n : :■ ": - e^f-.t.. .n\i . .: -'«tiv. ,^'^Aim': '^ '..,-..,.: .J .;..v.; .r:. *,.. !,,!.<.. •./ /■:•'.. '^ ! ..^•..u .-.. : " ■.-.■^4 . \ : .J, . ' .:ru.i?i: on"i .. ?: c^'iU L..-.. , •?^0!.-i.i,\' / .U'-:.'[\.':i 'ej ;■• ^^;^ _.)■. -rtj^.v y, ' ■ :-:;'nfjf' C'l ' h-.r; :!!j GORDIID^ 479 E. GORDIIDAE. Very long thin worms similar to Filariae, which, in their adult condition, hve free in brooks, pools and springs ; the mouth and the commencement of the intestine are obhierated ; there are no lateral ridges, and the muscular system presents a structure different to that of the Nematoda . The posterior end of the male is split, and spicules are lacking ; there are two testicles. In both sexes the genitalia discharge through the terminal gut. The larvae, which carry a rostrum beset with hooks, force themselves into the larvcC of water-hisects : more rarely they invade molluscs, and they then become encysted within the body of the host. According to Villot, at least a part of them attain the intestine of fishes, where they again become encysted, and after a period of rest they travel into the tissues of their hosts, and finally again reach the exterior by way of the intestine, where they then become adult. In most cases, however, the gordius larvas are taken up by predacious water insects ; they live for a while in the body cavity of these insects, undergo a metamorphosis, and finally wander into the water. A few species invade man accidentally with water, in which case they are usually vomited up : — Gordius aquaticus^ Dujardin, 30 to 90 cm. in length (Aldrovandi, Degland, Siebold, Patruban). Gordius tolosanus^ Duj., 11 to 13 cm. in length (Fiori). Gordius varius, Leidy, 10 to 16 cm., female, up to 30 cm. in length (Diesing). . Gordius chilensis^ Blanch. (Guy). Gordius villoti^ Rosa (Bercutti, Camerano) ; Gordius tricuspidatus^ L. Def. (R. Blanchard), Gordius violaceus^ Baird (Topsent), and Gordius pustulosus^ Baird (Parona). j ■J ■ ^ ■ • -i ... i 1 • it.-.- 4%> THE ANIMAL PARASITES OF MAN F. HIRUDINEA s. DISCOPHORA (Leech). The Hirudinea^ which have been appropriately included amongst the Annelida, differ in many respects from the typical members of the group; their body is long and flat, it lacks the parapodia that are characteristic to all forms of Annelida ; but, on the other hand, possesses a terminal posterior sucker, and in many species there is also an anterior sucker. The mouth is terminal at the anterior end, the anus lies dorsally above the posterior sucker (fig. 3480). The body is segmented, but this is less manifest in the body covering than it is in the arrangement of the internal organs ; the segmenta- tion, nevertheless, is also indicated exteriorly by the appear- ance of the cutaneous sensory organs which correspond to the segments. This shows what the condition of the ganglia in the abdominal ganglion chain has taught us, that the anterior and the most posterior segments are considerably abbreviated— a part of the latter taking part in the formation of the suctorial organs. In a great many species the skin is distinctly annu- lated, four or five of such rings, at least in the central region of the body, appearing on one segment of the body. The condition of their body cavity is another peculiarity of the Hirudineaj it is narrowed by the powerful development of the connective tissue and the muscular system into four tubular sinuses, which have the appearance of blood-vessels. There are usually one dorsal and one ventral median trunks, as well as two lateral trunks ; in addition, a particular blood-vessel system exists. » The skin consists of a very thin cuticle that is cast off from time to time ; it is secreted by the underlying cylindrical epi- thelium, which contains numerous goblet cells. The muscular system is strongly developed ; it consists of long tubular fibres, which run circularly, longitudinally and in the dorso-ventral direction ; the muscular system , is subject to a particular expansion in the clinging organs and at the commencement of the intestine. On the whole, the alimentary canal represents a tube running straight from the mouth to the anus, which possesses a number of blind sac-like protuberances at the sides varying according to the species. The most anterior section, the pharynx, in the leeches with maxillae carries three chitinous, semicircular plates furnished with teeth —the jaws — which serve to tear up the epidermis in order to open the blood-vessels ; in the leeches with rostra a long protractile proboscis rises from the base of the elongated pharynx. Numerous salivary glands, the secretion from which possesses toxic properties, discharge into the pharynx. The oesophagus, which follows the pharynx, and to the exterior of which numerous radiary muscles are fixed, is a suctorial organ in its entire structure. The nutriment in the larger species con- sists of the blood of vertebrate animals, in smaller species and in the young stages the food consists of small invertebrate animals. Fig. 348D.— The internal organs of the leech. The creature has been opened from the dorsal surface, and part of the intestine has been removed. The testicles, with vas deferens, may be seen between the blind ducts of the intestine ; beyond these on either side the segmental organs. The female genital organs are in front of the most anterior pair of testicles. (After Kennel.) GNATHOBDELLID^ 481 The NERVOUS SYSTEM exhibits the typical structure of other segmented worms ; the sensory organs consist of the previously mentioned goblet-shaped cutaneous sensory organs, of the organs of taste, and of eyes, the latter frequently being present in large numbers. The EXCRETORY or segmental organs exhibit many peculiarities, which cannot^ however, be detailed here. They commence with funnels in the lacunes of the body cavity, and usually discharge on the ventral surface. ' Almost all the Hirudmea are hermaphrodite and copulate reciprocally. The two ovaries are very small, and the oviducts that proceed from them soon unite into a common duct, which then passes into the uterus and discharges through the short vagina in the median line of the ventral surface behind the male, organs into the so-called clitellar region. The male sexual apparatus consists of symmetrically arranged testicles, varying in number according to the species, the short vasa efferentia of which, one by one, run into the vas deferens, passing towards the front on each side. In frqnt, at about the level, or a little in front, of the female. gejiitalia, the two vessels pass into a convoluted mass of tubes to the so-called epididymis, and then discharge into the single protractile penis (fig. 348D). All leeches deposit so-called COCOONS. These are small barrel-shaped or pouch- like bodies, which are surrounded by a thicker shell and contain a number of eggs in a large mass of albumen ; the albumen originates from glands of the generative organs, the shell substance from cutaneous glands of the clitellar region. Family. Gnathobdellidae (Leeches with Jaws). ^ ,.; These are distinguished by the possession of usually three jaws in the pharynx"; the body consists of twenty-six segments. The posterior sucker is large and flat; the anterior sucker is smaller. The Hirudinea have five pairs of eyes, the NephelititE have four pairs. Genus. Hirudo, L., 1758. The entire body consists of 102 annulations, five appearing on one segment in the central region of the body. The pharynx has three semicircular jaws, the arched border of which is beset with numerous teeth (50 to 100). The male sexual orifice lies between the thirtieth and thirty-first rings, the female orifice between the thirty - fifth and thirty-sixth. There are numerous species, some of which are utilized for medicinal purposes. ^ • ^ ' J ^ Hirudo medicinalis, L., 1758. It occurs in numerous colour varieties, one of which has been designated Hirudo officinalis, Moq.-Tandon. Usually the dorsal surface is greyish-green and is marked with six rusty-red longitudinal stripes. The ventral surface is olive-green, more or less spotted with black, and marked at the sides with a black longitudinal line. The length averages 8 to 12 to 20 cm. This leech lives in swamps, ponds and brooks, overgrown with plants and having a muddy bed. The cocoons are deposited in the soil at the sides. Europe, as well as North Africa, is its home. At the present day it has been exterminated from most parts of Central Europe, but it is still very common in a 1) Hungary. Its use for medicinal purposes is well ¥iG, 24^E.—JItni(fo medicin- known. A large leech can suck about 15 grs. of ''''^- , '^' anterior end with :,,«>,,, • , f ■ open buccal cauity, with the jaws, blood, and about the same amount is lost through j ^^ ^j^g. ^ ^^g j^^ isolated. secondary haeriiorrhage. (After Claus ) 482 THK ANIMAL PARASITES OP^ MAN Hirudo troctlna, Johnston, 1816. Syn. : Hirudo ittterrupta^ Moq.-Tandon, 1826. This species measures 8 to 10 cm. in length. The back is greenish, with six rows of black spots surrounded by red ; the lateral borders are orange-coloured ; the abdomen spotted or unspotted. Its habitat is in North Africa and Sardinia. It is applied medicinally in England, Spain, France, Algeria, etc. Genus. Limnatis, Moq.-Tandon, 1826. Nearly related to Hirudo, but is differentiated by a longitudinal groove on the inner surface of the upper lip of the anterior sucker. The jaws are furnished with over 100 very sharp toothlets. Limnatis nilotica, Savigny, 1820. Syn.: Bdella nilotica, Sav. ; L. nilotica, Moq.-Tandon; Hcemopis {vorax), Moq.-Tandon, 1826,/. p.\ Hcefnopis sanguisuga^ Moq.-Tandon, 1846 {nee Hir. sanguis^ Bergm., 1757). This species measures 8 to 10 cm. in length, and becomes gradually more pointed towards the front ; the body is always soft. The back is brown or greenish, and has usually six longitudinal rows (rarely only two or four) of black dots. The abdomen is dark ; but numerous colour variations occur. The native place is North Africa, especially the coastal regions ; it is also found in the Canaries, the Azores, Syria, Armenia, Turkestan, perhaps also Southern Europe. It is taken into the mouth with drinking water, and may settle in the pharynx, larynx, oesophagus, and nasal cavities of human beings. This species has also been observed in the vagina and on the conjunctiva. It is equally fond of attacking domestic animals. Hirudo mysomelas (Senegambia) and Hirudo granulosa (India) are placed with this genus, and, like our leech, are also used for medicinal purposes. J Genus. Haennadipsa, Tennent, 1861. These leeches live on land, and measure 2 to 3 cm. in length. About a dozen species are known. They are a veritable scourge to persons in the tropics (Asia, South America), as they attack them to suck their blood. They are able to force their way even under close-fitting garments, so that it is difficult to protect oneself from their assaults {Hcemadipsa ceylonica, Bl., and other species). Family. Rhynchobdeliidae (Leeches with Rostrum). These are furnished with a proboscis in lieu of the jaws ; the segment consists of three annulations. Genus. Haementaria, de FiHppi, 1849. Haennentaria officinalis, de Fil. Inhabit Mexico, where they are used for medicinal purposes. Genus. Placobdella, R. Blanch. Placobdella catenigera, Moq.-Tandon. Indigenous to South Russia, Hungary, Italy and South France. It is a parasite of the swamp turtle, but frequently attacks human beings. ARTHROPODA 483 G. ARTHROPODA (Jointed-llmbed Animals). BY FRED. V. THEOBALD, M.A. Bilaterally symmetrical segmented animals which are covered with a thick •cuticle that is frequently calcareous {Crustacea)^ but always thinner between the segments ; they carry (primitively) a pair of jointed appendages on every segment.^ The segments of the body are uniform in certain regions, but differ from those of contiguous regions, so that it is easy to distinguish three parts (head, thorax and abdomen), each composed of segments. The cephalic segments are always formed into a uniform head, the segmentation being scarcely recognizable at either end ; the thoracic segments may also fuse, or part or all of them may coalesce with the head ; the abdomen, as a rule, retains its segmentation, but this may possibly also be lost, in which case it is [sometimes] united to the cephalothorax. The structure of the three regions depends mostly on the varying form and function of the appendages : those on the head are primitively locomotive organs (and frequently are still so in the early stages), but they become transformed into feelers and mouth- parts (mandibles, maxillae) ; the limbs of the thorax, however, usually retain their ambulatory functions, as frequently do those of the abdomen ; sometimes, however, the abdominal limbs disappear, entirely or partly; in the latter case they are then utilized for other purposes. In their organization the Arthropoda approach the segmented worms. The Arthropoda are generally divided into five groups {Crustacea^ Protracheaia^ Arachnoidea, Myriapoda^ and Insecta or Hexapoda\ of which only the Arachnoidea and the Hexapoda interest us here. A. ARACHNOIDEA (Spiders, Mites, etc.). The head and thorax are always united together ; the abdomen is either segmented or without exterior segmentation, in which case it is united with the cephalothorax.^ The number of pairs of appendages ' [In most Arthropoda the skin is hardened by a deposit of chitin {Hexapoda, etc). — F. V. T.] 2 Parasitic or free-living Crustaceans may now and then invade man abnormally. Thus, according to Betten, C aligns curtus invade the cornea (Betten, R. A., ** Par. Crust, as a Foreign Body on the Cornea," Lancet, 1900, i, p. 1002; and Centralbl. f. Bakt. u. Far., xxix, p. 506). According to Laboulbene, also Gammanis pulex (Laboulbene, A., "Obs. d'accid, caus. par le G. ful. apport. avec I'eau de boison dans I'estomac d'un homme," Bull. Acad, med., 1898, p. 21). ^ R. Blanchard has compiled thirty-five cases in which Myriapoda have been observed in the intestine as well as in the nose of human beings ("Sur le pseudopar. d. myriap. chez I'homme," Arch, de Par., 1898, i, p. 452). E. Munoz Ramos reports an additional case (ibid., p. 491). A few years ago a doctor in East Prussia sent me a rain worm out of a lady's nose (of. Hanan, A., " Wahrsch. Pseudoparas. v. Schweiss fliegenlarv. u. angebl. Para?. V. Regenwurmern b. ein Hysterisch," Arch, de Par., 1899, ii, p. 23). * [This is only so in the Acarina or mites, not in the Aratieida or spiders. — F. V. T.] 484 THE ANIMAL PARASITES OF MAN amount to six, of which the two front pairs, the chelicerae and the pedipalpi, are attached to the head region and the four remaining pairs to the thoracic region.^ The abdomen in the adult condition has no appendages. The Arachnoids are air-breathers, and for this purpose are either provided with tracheae or with so-called lung-sacs, or they breathe through the surface of the body. Some aquatic forms breathe by gills. There are eight or ten orders of Arachnoids,^ of which, however, only two, the Acarina and the Linguatiilida, have to be considered here.^ Order. Acarina (Mites). Small Arachnoids, the three parts of the body of which are, as a rule, coalesced ; it is only rarely that a faint line indicates the division between a cephalothorax and abdomen. The two appendages on the head are designed for biting or puncturing and sucking, and vary according to their use. The chelicerae * are fang-like jaws or puncturing bristles forming a kind of rostrum, the pedipalpi are claw-like or shear- shaped, or form a suctorial proboscis.^ The four pairs of legs are usually well developed, more rarely they are rudimentary or have partly vanished ; many parasitic forms are provided with pedunculated suckers [ambulacra — F. V. T.]. Respiratory organs (tracheal tufts) may be present or absent. The nervous system is reduced to a minimum, eyes are usually lacking. The intestine, situated in the central part, generally has three blind appendages ; the anus is situated on the venter above the posterior end. Sexes separated ; nearly all the species deposit eggs, from which six-legged larvae hatch. The Acarina live either free in the water or in moist soil, or they are parasitic on plants and animals.^ ^ [The true character of the Arachnoidea is the presence of four pairs of ambulatory appendages. This number is reduced to two pairs in the gall-making Phytoptid^^ and they differ from all other Arthropoda in having no antennae. — F. V. T.] ^ Twelve orders are now recognized, as follows : Pentastomida or Linguatulids ; Tardi- grada or bear-animalcules ; Phalangidce or harvest-men ; Acarina or ticks and mites ; Palpigradi ; Solifugce ; Pseudoscorpiotiidea or book mites; Pedipalpi or false scorpions; Scorpionidea or true scorpions ; Aratieida or spiders ; Xiphosura or king crabs ; and Pycnogonida^ marine Arachnoids. ' Chelifer cancroides has also been observed as a pseudoparasite in man (Arnault de Very, S., "Pseudopar. du. Chel, cancr. chez I'homme," Conipt. rend. Soc. de Biol., 1901, liii, p. 105). * [The chelicerae are sometimes regarded as modified antennae, but it is more natural to regard them as the morphological equivalent of the mandibles of Hexapoda. — F. V. T.] * [The pedipalpi, or second pair of jaws, consist of a stout basal segment and a palp, which may have the appearance of a leg in Arachnida ; this may end with or without a claw, or with a chela (scorpions) ; they may also form a tube enclosing the styliform chelicerae (mites). — F. V. T.] ^ [Acarina are also found living upon trees, feeding upon other Arthropods and also upon spores of lichen and fungi {Oribatid(B or beetle mites) ; they also swarm indoors amongst stores and provisions {Tyroglyphidce and Glyciphagi, household, sugar and cheese mites). This order is very important, as many are parasites upon man, his domestic animals and his cultivated plants, and attack his provisions and stores. Some live on blood, and in some of the ticks distribute various protozoal and other blood parasites and germs. — F. V. T.] TROMBIDIID^ 48: Family. Trombidiidae (Running Mites). Soft-skinned Acarina with tracheae and with two eyes, usually pedunculated ; they are often brightly coloured ; chelicer^e lancet- or claw-shaped ; pedipalpi claw-like ; legs composed of six segments, with suctorial discs between the terminal ungues.^ Larvae six-legged. To the latter belong the larvae of several species of Trombidium such as : — Genus. Tronnbidium, Latreille (and Leptus). Leptus autumnalis, Shaw, 1790. Leptus occur as parasites in the human skin and cause a cutaneous disease known as autumn erythema, and produce a very unpleasant sensation on account of the trouble- some itching ; in children it is very often accompanied by fever.^ Formerly these mites were considered adult forms, but when they were recog- nized as mite larvae they were taken for those of the spider-mite {Tetranychus telarius) ; the investigations of Hanstein, however, showed this to be a mistake. Fig. 349. — Leptus autumnalis, with so-called mcking proboscis. Enlarged. (After Gudden.) Fig. 350. — Leptus autumnalis : the so-called proboscis is formed around the hypopharynx sunk into the skin, loo/i. (After Trouessart.) When Henking first investigated the development of Trombidium fuliginosum, parasitic in the larval stage on vine-fretters, he demonstrated the occurrence of a form very similar to Leptus autumnalis^ and the "autumn, grass, or gooseberry" louse was commonly designated the Trombidium larva. Even before Henking's work it had been described by Megnin as the larva of Trombidium holosericeum^ a red-coloured species, frequently occurring in spring and summer on the ground, trees, etc. This assumption, however, as Moniez was the first to explain, is not correct ; ^ [Some have seven segments to the legs. — F. V. T.] ' [This minute parasite is especially obnoxious in barley fields. In walking over barley stubble one is sure to be attacked by this Acarus in many districts. Trombidium is often very prevalent in gardens, especially along rows of peas, and in spring they may be found on fruit trees and bushes. Nut-pickers are frequently attacked by Leptus, and also pickers in other fruit plantations. It is often called the harvest mite. — F. V. T.] 486 THE ANIMAL PARASITES OF MAN indeed, as many as three species come under consideration : T. gymnopterosum^ L., T. fultginosufu, Herm. (according to Brucker), and two species known hitherto only in the early stage, T. siriaticeps. Helm, et Oudem., and T. poriceps, Helm, et Oudem., which are not only parasitic on mammals, but on birds, on Arthropods and especially on insects. Arthropods appear to be the normal hosts for the larvae. The above-mentioned forms invade the skin of man by means of their oral apparatus, by preference invading the orifices of the sebaceous glands so as to suck the blood ; around the point attacked there arises a wheal about the size of a lentil, and around the inserted hypopharynx a fibrinous secretion, the ^'proboscis," which, however, is a product of the host, just as chitinous secretions are provoked by Trombidia parasitic on Arthropods. Further species, analogous in habit to Leptus auiumnalis, are described by Riley from Central and South America as L. americanus and L. irritans. [L. autumnalis attacks small mammals by preference, such as moles and hares, which are often literally covered with them. Dogs are also subject to their attack, and cats suffer similarly. This mite also frequently appears in colonies on cows ; cavalry horses after autumn manoeuvres often suffer from an erythematous affection about the hocks and knees due to this pest. [A number of Leptus, so far undescribed, occur abroad which attack man in the same way as L. mitiimnalis in Europe. Dr. Durham has brought me specimens from British Guiana called hetc rouge ; this species works under the skin much as does our European species, but it is very distinct, being considerably larger. — .p. V. T.] Trombldium tialsahuate, Lemaire, 1867. T. tialsahuate occurs in Mexico under conditions similar to those of Leptus here. It also frequently attacks men, and especially fastens itself on to the eyelids, in the axillae, navel, or on the prepuce ; it induces itching and swelling of the parts affected, and sometimes even causes suppuration ; the symptoms, however, generally disappear after a week and remain localized. ^ Other species of mites which attack man are reported, mostly by travellers, from various other places ; zoologically, however, there is little known about them. The pou d'agouti in Guiana, niaibi in New Granada, colorada in Cuba, mouqui in Para, and the buschmucker in New Guinea represent a few of these. ' Lemaire, •'Import, en France du tialsahuate," Compt. tend. Acad. Sci., Paris, 1867, Ixv, p. 215. AKAMUSHI OR KEDANI 487 Akamushi or Kedani. In a few districts of Japan there occurs a serious illness, with a mortality of 40 to 70 per cent. It is called river or flood fever, and the Japanese doctors have connected it with a small mite (akamushi, kedani). Baelz has opposed this opinion on the grounds that he has repeatedly observed the same species of mite in his dwelling without any subsequent illness occurring. Accord- ing to Keisuke Tanaka, however, a connection certainly does exist, inasmuch as the akamushi, like Leptus, attacks persons to suck blood. If the mite is not removed, or if the spot attacked is injured by scratching, etc., a papule surrounded by a red area Fig. 351. — The kedani mite. Enlarged. (After Tanaka.) forms, and a pustule ensues ; and finally a black scab covers the seat of injury. The lesion becomes the point of entrance of bacteria, especially a species of proteus which produces river fever. If the mites are carefully removed no general illness takes place. The orange-red mites, which we only know in their larval condition, measure o*i6 to 0*58 mm. in length by o*io to 0*24 mm. in breadth. They have leg-like palpi with three joints, hirsute bodies, and very hairy legs composed of five segments, terminating with three ungues. 488 THE ANIMAL PARASITES OF MAN Family. Tetranychidae (Spinning Mites). These have tracheae and eyes ; the palpi are composed of four segments, of which the last but one has a powerful claw. The legs have six segments with sucker discs between the claws. [Tiie red spiders or spinning mites (Tetranychi) are usually placed in the family Trombidiidce. — F. V. T.] Genus. Tetranychus, Dufour. Tetranychus molestlssimus, Weyenbergh, 1886.^ Found in Argentine and Uruguay on the under surface of the leaves of Xanthium macrocarpiun ; it attacks mammals and men, producing severe itching, accompanied by fever in the latter. It has been asserted by Haller that the Cape ailment (Port Natal ;, \v M sickness) is caused by mites, but this statement has been contested. Tetranychus telarius, L., 1758,^ var. russeolus, Koch. This common spinning mite like- wise attacks human beings, but the papules produced by it very soon disappear. Family. Tarsonemidae. A family distinguished by complete sexual dimorphism, the species of which are provided with tracheae ; the legs have five segments ; the terminal segments of the front pair of legs of both sexes possess a claw ; the terminal segment of the posterior pair of legs of the male likewise has a claw. In the female this pair of legs, like the second and third pairs of both sexes, is provided with two booklets and a sucking disc. The cuticle of the body on the back is " annulated." [This family of small transparent mites live normally as plant parasites. The last two pairs of legs are widely separated from the two front pairs. — F. V. T.] * [This species is also known as Bicho Colorado. It spins a web under the lower surface of the leaves, and it is only from December to February that it attacks warm-blooded animals and man. — F. V. T.] 2 [There is something wrong here, probably in the identification. T. telarius is purely a plant-feeder, and it is extremely unlikely a variety would attack man. Anyhow, it will not do so in Great Britain.— F. V. T.] Fig. 352. — 7 eiranychus telarius var. rus seolust Koch. Enlarged. (After Artault. PEDICULOIDES 489 Genus. Pediculoides. Pediculoides ventrlcosus, Newport, 1850. Syn.: Heteropus ventrlcosus^ Newport, 1850; Acarus iritici^ Lagreze-Fossot, 185 1 ; Physogaster larvarmn, Lichtenstein, 1868; Sphcerogyna ventricosa, Laboul- bene and Megnin, 1885. Males are oval in shape, 0*12 mm. in length and q-qS mm. in breadth, flattened. There are six pairs of chitinous hairs on the dorsal surface and a lyre-shaped lamella on the posterior part. The female in the non-gravid state is cylindrical in form, 0*2 mm. in length and 0-07 mm. in breadth ; when gravid the posterior part of the body becomes enlarged into a ball, which may attain 1-5 mm. in size, as in the case of Pnlex penetrans and of the female Termites. On emerging the young are already provided with four pairs of legs and copulate soon after birth. Fig. 353. — Pediculoides ventricosus, a, male ; b, young female ; c, gravid female. Enlarged. (After Laboulbene and Megnin.) These animals live on the stalks of cereals, and feed on vegetable and animal juices ; they are also found on corn-infesting insects. They invade the barns and seek out the insects living in the dry grains of corn, or wait for an opportunity of obtaining food. They have been repeatedly observed on human beings, particularly labourers occupied in handling grain ; their bite causes severe irritation, local elevation and reddening of the epidermis, as well as fever. It cannot be positively asserted that all cases of the occurrence of cereal mites on man relate to P. ventrlcosus^ as the descriptions are often insufficient. Geber states that one form is Chrlthoptes 7nonungulculosus^ or Acarus hordel; Flemming mentions Tarsone?nus unclnatus ; Koller Orlbates sp. ; and Karpelles Tarsonemus Intectus. [The pregnant female Pediculoides has a large round inflated abdomen, in which the ova hatch and the young mature. Later they escape from the parent as adults— F. V. T.] 490 THE ANIMAL PARASITES OF MAN Genus. Nephrophages. Nephrophages sanguinarius, Miyake and Scriba, 1893. Males measure 01 17 mm. in length and 0-079 mm. in breadth; females up to 0*360 mm. in length by O' 1 20 mm. in breadth. The head is provided with two very large scissors-like jaws and two large round eyes. The legs are composed of five segments and are all of equal length ; the three anterior pairs of legs have pedunculated ambulacra, the posterior ones terminate in a claw. The cuticle on the back is thickened in three places, shield-like; the abdominal surface without scutellum is longitudinally striped and is beset with chitinous hairs. Colour greenish to brownish- ^^ s^ \ jr '1 $*. . V \ yellow. Eggs 0*040 mm. 0*046 to Fig. 354. — Nephrophages sanguinarius : male, ventral surface. Enlarged. (After Miyake and Scriba.) Fig. 355. — Nephrophages sanguin- arius : female, dorsal aspect. En- larged. (After Miyake and Scriba.) The authors discovered these mites, but always dead, in the urine of a Japanese suffering from fibrinuria complicated with chyluria and haematuria. They surmised that they were endoparasites, probably situated in the kidney ; but this view is not convincing, though they also report that for a week, day after day, the mites were found in the patient's urine, as well as in urine drawn off by means of a catheter, and in the water used to wash out the bladder (one or two specimens and an egg). The statement that these mites have large eyes makes the discovery suspicious, to say the least. The significance of the discovery is not supported by the further statement that Disse is supposed to have found an encapsuled mite closely related to the Tyroglyphides on the wall of the vena cava. In the case of Marpmann, who found a dead Acarid in the urine of a man suffering from chronic nephritis, and in whom later examinations proved negative. EUPODID^, GAMASID^ 491 the author himself was of opinion that the mite had reached the urine from outside. We are certainly acquainted with mites living endoparasitically, namely, the Cysticolce, AnalgesincB^ of which Laminosioptes galli?iarum live in the intramuscular and subcutaneous connective tissue of fowls, and Cytoleichus sarcopto'ides in their air sacs. Another kind of mite {Halarachne halichoeri) is occasionally found in the nasal mucous membrane of the seal {Haltchcerus grypus)^ and, quite recently, Pneumonyssus simicola^ which is more nearly related to Halarachne, has been found in the lung of Cynocephalus sp. It is therefore not improbable that endoparasitic mites are found in man ; but no definite discovery has yet been made. Family. Eupodidae. Small tracheate mites, with moderately long or short pedipalpi, composed of four segments, of which the last segments bend ; chelicer^e forceps-shaped, with serrated edges ; legs with two claws, more rarely with one, and terminating in a tuft ornamented with fine hairs; genital orifices on the abdomen, surrounded by a circle of little hairs. Most species live free — one lives parasitically on the bodies of slugs. Genus. Tydeus, Koch. Tydeus molestus, Moniez, 1889. Male, o'2 mm. in length, o'i25 mm. in breadth. Females, 0*225 mm. in length, o'i35 mm. in breadth; gravid female 0*315 to 0*360 mm. in length and 0*180 mm. in breadth. They were observed by Moniez on an estate in Belgium, whither the creature had apparently been im- ported twenty-five years previously with Peru- vian guano ; they ap- peared regularly in the summer and remained until the first frost set in; they were found on grass-plots, on trees and bushes in masses; they regularly attacked human beings, mammals and birds, tormenting their hosts in a terrible manner. Fig. 356.- Tydeus molesHis : seen in profile. (After Moniez.) Enlarged. Family. Gamasidae (Coleopterous or Insect Mites). Chelicer^E chelate or styliform ; pedipalpi filiform ; the legs are composed of six segments with two terminal ungues and a bladder-like sucking disc [caruncle — F. V. T.]. Stigmata situated between the third and fourth pairs of legs ; the cuticle thickened, leather-like ; no eyes ; the larvas have six legs. The GamasidcB are predaceous on small insects and other mites ; some are parasitic on insects, and one is noticeable as a pest on birds, etc. 31 492 THE ANIMAL PARASITES OF MAN Genus. Dermanyssus, Dnges. Dermanyssus gallinae, de Geer, 1778. Syn. : Pnlex gallincu, Redi, 1674; Atarus gallincB^ de Geer, 1778 ; Dermanyssus avium^ Duges, 1834. The male measures o*6 mm. in length by 0-32 mm. in breadth; the female 07 to 075 mm. in length by 0*4 mm. in breadth. The body is somewhat pear-shaped ; the colour whitish, reddish, or reddish- black, according to the contents of the intestine. The legs are. fairly short and strong. During the day they live concealed in the nests, cracks, etc., of the hen-house, and at night attack the inmates in order to suck their blood ; they rarely remain long on the birds. They have been repeatedly found on persons, on whose skin they produce an itching eruption. Dermanyssus hirundinis, Hermann, 1804. Syn. : Acarus hirimdi?tts, Herm., 1804. Of a brownish colour, i-2 or 1*4 mm. in length ; lives in the nests of swallows and is occasionally found on man. [The red hen mite {Dermanyssus galHnce) not only attacks poultry and man, as stated above, but is found on all birds and Fig. 357. — Dermanyssus gallince. En- larged. (After Berlese.) Fig. 358. — Dermanyssus hirundinis. 40/1. (After Delafond.) many mammals. The D. gallmce is the same as D. avium. The species found in swallows' nests is also said to be the same. This mite can remain for weeks without any food from its normal host. They only attack man when entering or cleaning dirty and neglected fowl-houses ; they do not produce a true dermatosis. They, chiefly attack the backs of the hands and forearms of those who con- stantly attend poultry and give rise to symptoms similar to the papular eczema of scabies. That they may remain some time upon IXODID^: 493 the human body we know from the following cases out of many recorded : Geber observed that the Dermanyssus had caused a diffused eczema on a woman, which lasted four weeks and then disappeared. The tlque of F. V. Raspail is the bird Dermanyssus ; he records children and adults being attacked not only when handling pigeons, but even when walking in a garden manured with pigeons' dung. The affection soon disappeared wiien the pigeons were destroyed and the excreta buried. I have frequently heard of poultrymen being seriously attacked by this pest. — F. V. T.] Genus. Holothyrus. Holothyrus coccinella, Gervais, 1842. Measures 5 mm. in size ; lives on birds in the Island of Mauritius ; ducks and geese frequently fall victims to its bite ; it also attacks human beings, on whose skin it causes severe burning and swelling, but no reddening ; it may be dangerous to children, especially by settling in the oral cavity. Other Gamasides occasionally occur in man, for instance, according to Moniez, Leignathiis sylviariivi^ Canestr. et Fanzago ; according to Neumann Lcelaps siabularis. The former live normally in the nests of various species of Sylvia^ Lselaps on dried vegetable substances, also in houses. [Marchoux and Conoy {Bull. Soc. Path, exot., 1912, v, No. 10, pp. 796-798) found Leishman granules in Lcelaps echidnintis. It is assumed that Leishman granules may be found in most Arachnoids, and have no connection wath Spirochaeta. — F. V. T.] Family. Ixodidae (Ticks). Comparatively large Acarines with a leathery skin ; they are flattened in form, but after sucking blood the abdomen becomes spherical ; the cheliceras are rod-like and possess a serrated terminal joint, bent hook-like ; the median parts of the pedipalpi (maxillae) form a rostrum furnished with barbed hooks (fig. 359) ; the maxillary palpi themselves are club-like or rounded ; the legs are composed of six segments with two terminal ungues, often also with "sucking discs" ; the stigmata are at the sides of the body, posterior to the fourth or third pair of legs. The larvae are six-legged. [The true ticks (Ixodidcv) are all blood-suckers, and as far as is known they do not take vegetable food at all. Not only are the Ixodidcv important as actual parasites, but they are most so on account of the fact that they are the active agents in carrying various diseases in animals and apparently in man. It has been conclu- sively proved that the bont tick {Amblyouiina hebrceiun) is the carrier of the fatal ''heart-water fever" so rife amongst sheep in South Africa, that the dog lick {Hcvmaphy salts leachi) is the agent by which the protozoa that cause malignant jaundice in dogs are distributed, that 494 THE ANIMAL PARASITES OF MAN Texas fever in cattle is spread by Rhipicephahis anniilattis, and Coast or Rhodesian fever by R. appendiciilaUis and R. siinns. Their im- portance as disease carriers amongst mammals is therefore consider- able, and it may prove to be so for man.^ They frequently attack man, but chiefly, according to my observations, in their early stages in Europe ; this is not so, however, abroad. The life-history of a number of ticks has been clearly demonstrated. Mr. Wheler has shown that in Ixodes reduvins it is as follows : the female deposits her eggs in masses upon the ground, gradually reducing in size as the eggs pass out, until she finally remains a mere shrivelled empty bag and then dies. The eggs are oval, golden brown in colour and smooth; in length they are 0*59 mm.; as in all Ixodidcv they are covered with a glutinous secretion, by means of w^hich they adhere together in masses. These egg masses may be deposited anywhere on the ground, but amongst rough, coarse herbage seems to be the favourite place. The egg stage may last as long as twenty-two weeks, or it may only take eight weeks. In the case of the bont tick a single female may deposit 15,000 or more eggs. The process of egg-laying is described as follows by Mr. Wheler : "When egg-laying is about to take place, the head is further depressed till it rests close against the under side of the body. In this attitude the end of the rostrum actually touches the genital orifice, the palpi being at the same time widely opened out. Behind the head and from beneath the shield, at what for the purposes of explanation may be described as the back of the neck, a white, perfectly transparent, delicate gelatinous membrane is brought dow^i through inflation, either with air or with a transparent fluid, above the head, which it temporarily conceals. The end of this mem- brane terminates in two conical points which appear to be covered with a glutinous secretion, and at the same time an ovipositor of a somewhat similar character, but only semi-transparent, is pushed forward from the genital orifice. This latter is a tube, within which is the Qgg. As the ovipositor projects it turns itself inside out, like the finger of a glove, leaving the egg protruded at the end and lying between the two finger-like points of the membrane. The membrane and the ovipositor are then withdrawn each from the other. The tgg adheres to the former, which collapses through the withdrawal of its contents, dragging the head forward and depositing it on the top of the head. Neither legs, palpi, nor the organs of the mouth take any part in oviposition, but after the collapse of the membrane the palpi are closed and the head is raised, by which action the egg is pushed forward to the front edge of the shield, forming ' This has been proved in Uganda— so-called tick fever in man. IXODID.E 495 in time an adherent mass of eggs, which are deposited in front of the tick." [The egg gives rise to the larval form, the so-called "seed-tick" stage. At first these minute specks are pallid and soft, but they soon harden and darken in colour. These larvae are six-legged and crawl up grasses and various plants, and there await a passing host, weaving their two front legs in the air and becoming attached by this means. The larval ticks feed upon the blood of the host, and when replete fall to the ground, the body becoming inflated in the meanwhile. These larvae may remain on the host only two days, or they may remain much longer. Eventually they moult on the ground and change to the nymph or pupal stage, wiiich has four pairs of legs. This pupa acts just as the larva, crawls up plants and waits to regain the host. After a time the nymphs, having gorged themselves with blood, fall off and remain on the ground for nearly three months ; they then moult and become adult males and females. In about ten days they assume their normal colour and regain the host afresh ; the female gradually swells until she attains that large inflated form so characteristic of ticks. The male does not swell, but nevertheless feeds upon the host and fertilizes the female. [The act of coitus is strange : the male tick inserts its rostrum and other mouth organs into the sexual orifice of the female, between the base of the posterior pair of legs. The males then die and the females fall to the ground and deposit the ova. There are variations in the different species, of course, from those given above, which apply solely to Ixodes reduvius. The larvae and nymphs seem to attack most animals, but the adults mainly keep to the same host. The periods in the life-cycle of ticks not only vary in the different species, but in each species according to climatic conditions ; for instance, in the bont tick {Amblyounna hehrceum, Koch), Lounsbury has shown that the development is rapid in summer, slow in winter. The period from the time that the female drops to the time she commences to lay eggs varied in specimens observed by him from twelve days in summer to twelve weeks in winter, and the complete period from the dropping of the female to the hatching of the eggs, from eleven weeks in summer to thirty-six weeks through the winter. Other stages vary in a similar manner. [Ticks may live a long time away from the host provided they are supplied with a certain amount of moisture. Mr. Wheler kept dog ticks {Ixodes pUunhetis) in the larval stage for ten months ; the pupae, male and female, of /. reduvius for six months. [I have kept Oriiithodoms mouhata alive for eighteen months without food. [In many species moulting takes place off the host, but in /. hovis^ 496 THE ANIMAL PARASITES OF MAN now known as Rhipicephahis annulatus^ Say (the carrier of Texas fever), moulting takes place on the host, and in many other species also.^ Some species of ticks leave their host on its death (as the dog tick, Hcemaphysalis leaclii), but others die with the host (bont tick, Aniblyonifna hebrceum). [Two species are of special importance, namely Oniithodorus moubata, Murray, which may infect human beings with the spirillum of African tick fever, and Dermatocentor reticulatus var. occidentalism which is said to be the carrier of Rocky Mountain spotted fever. Classification of Ixodida\ [The TICKS, or Ixodidce, are divided into two groups, known as (1) Argantiiicv, (2) Ixodince. The Argantince are told from the Ixodiiice by the absence of dorsal or ventral shields in both sexes, and also by the rostrum being placed beneath the cephalothorax, which covers it over : except in the larval stage, in which it is sub- terminal, and in the pupal, when it partly projects. Legs nearly equal in length. The sexual orifice is situated between the two first pairs of legs. The males usually smaller than the females. [The Ixodince have the legs unequal, of six segments with two false segments, making them look as if composed of eight segments. The rostrum is terminal and never hidden beneath the body. The sexual orifice is situated between the bases of the first three pairs of legs. In the males the orifice is obsolete or very rudi- mentary, sexual intercourse being effected by the rostrum. The males are smaller than the females. The shield in the females never covers so much as one-half of the body even when fasting, also in the larvae and nymphs ; but in the males, which do not distend, the shield covers the body entirely, or all but a narrow margin. The Ixodince are divided into two groups : (i) the Ixodce, and (ii) the Rhipicephalce. The former have a long proboscis reaching nearly to the end of the palpi or even a little longer than the palpi. The palpi are longer than broad. The Rhipicephalcv have short palpi, nearly or quite as broad as long, more or less conical or subtriangular. They were called Conipalpi by Canestrini. Synopsis of Genera. [A. ArgantincE : Rostrum concealed in adult, partly exposed in larvae and nymphs. No dorsal and ventral shields. Body flat with thin edges, finely shagreened and punctate Argas. Body with numerous small round granules and with thick sides ... ... ... ... ... ... ... ... Ornithodorus. ^ Some ticks require only one [R. decoloratits), others two (A', evertsi), and some three hosts {R. appendiciilatus) in order to reach maturity. IXODES 497 Body with dorsal shield over [B. TxodmcE : Rostrum terminal, some part of it. I. Rostrum and palpi longer than broad (/;ir{S'^^^onyssus, Kolenati, 1857). Amblyomina, Koch, 1844 {Ixodes, Latreille, 1795). Hcemaphy sails, Koch, 1844 {Rhipistoina, Koch, 1844 ; Go?itxodes, Duges, 1888 ; Opitodon, Canestrini, 1897). Rhipicephalus, Koch, 1S44 {A carus, Linnaeus, 1758; Ixodes, Latreille, 1795; Phanloixodes, Berlese, 1889 ; Boophilus, Curtice, 1890). Dermacentor, Koch, 1844 {Ixodes, Latreille, 1795 5 Pseudixodes, Haller, 1882). — F. V. T.] Genus. Ixodes, Latreille. Ixodes reduvius, L., 1758.^ Syn. : Acariis reduvius and ricinus, L. ; Ixodes ricinus, Latreille, 1806. The males are oval ; their length i*2 to 2 mm. ; they are brownish- red or black in colour ; the females are yellowish-red, 4 mm. long; ' Ixodes reduvius and /. ricinus are synonymous. [The above should read Ixodes ricinus, Latreille, 1806.— F. V. T.] 498 THE ANIMAL PARASITES OF MAN when gorged they are lead-coloured, and may attain 12 mm. in length by 6 to 7 mm. in breadth. The dog tick (fig. 360) lives in thickets on leaves, etc., and attacks sheep and oxen, and more rarely dogs, horses, and human beings, into the skin of which the female bores with the rostrum in order to suck blood ; the bite is not dangerous, and sometimes is not even felt. Inflammation, however, is set up if the creatures are forcibly removed from the wound, as the rostrum as a rule is torn off in the process. If left alone or smeared over with some grease— vaseline, oil, butter, etc. — the creatures drop off spontaneously. Sometimes the entire tick bores itself into the skin ; they also appear to be permanent inmates of kennels. [The species /. rednvius is the same as /. riciniis, Latreille. The male is 2*35 to 2*80 mm. long; the body is dark brown, almost black; with a pale, almost white, margin ; there are also traces of reddish mottling. Coxae of the first pair of legs with a short spine. Rostrum much shorter than that of the female ; shield oval ; anal shield small, about one-third the length of ventral shield. The adult female varies from 2*80 to 3*5 mm. when not distended, but w^hen gorged may reach A.^ ) ]( \ 10 mm. long. The shield and Fig. 359. — A., the rostrum of Ixodes ricinus (male) ; B., the terminal joint of the maxillary palpi of the female. Enlarged. (After Pagenstecher.) Fig. 360. — Female of Ixodes ricinus , gorged full, dorsal and ventral surfaces. 2/1. (After Pagenstecher.) legs are dark blackish-brown, body deep orange-red with four dark longitudinal lines, paler beneath and light grey in front. When dis- tending it is pale red to grey or white ; when fully gorged olive-green, or dark red to black, with irregular yellow streaks on the back and sides just before egg-laying. Sexual orifice opposite fourth pair of legs. The nymph varies from 1*60 to 170 mm. long when fasting; the body is olive-white, opaque, with four distinct brown posterior markings and similar anterior ones, leaving a pale centre to the shield. When fully gorged it is 3 mm. long. As the nymph distends, it changes from opaque white to blue-black, and finally black. The little larva is o'8o to 1*50 mm. long, transparent with olive-green intestinal markings ; as it becomes inflated it changes to blue-black, and then black. There are no eyes. It is widely distributed, and chiefly attacks sheep ; sometimes it occurs on dogs and also attacks IXODES HOLOCYCLUS 499 man. Megnin records it from horses in the nymph stage. Amongst its other num.erous hosts are goats, cattle, deer, hedgehogs, moles, bats, birds, and lizards. It is usually known as the grass tick and bottle- nosed tick. This species occurs in Europe, Asia, North Africa, and North America. [Synonyms. — Considerable confusion exists over the name of this and other common ticks, owing to the same species having been described under a great many names. Observers have taken the same species on different animals and in various stages to be distinct, and have described them accordingly. [The name Ixodes rednvitis, Leach, does not stand, as Leach was describing quite a different parasite. The name /. ricinus, Latreille, 1806, is now substituted by Neumann and Wheler. [The synonyms given by Wheler are as follow : Rednvnis, Charleton, 1668 ; Ricinns caninus, Ray, 1710 ; Acarns ricinoides, de Geer, 1778 ; Acarus ricinns, Linnaeus, 1788 ; Cynorhcestes rediiviiiSy Hermann, 1804; Cynorhcestes ricinns, Hermann, 1804; Ixodes inegatJiyrens, Leach, 1815 ; Ixodes hipnnctatns, Risso, 1826 ; Cynorhcestes hermanni, Risso, 1826; Crotonns ricinns, Dumeril, 1829; Ixodes traheatns, Audouin, 1832 ; Ixodes plnmbens, Duges, 1834 ; Ixodes rednvitis, Hahn, 1834; Ixodes fnscns, Koch, 1835 (^) J Ixodes lacertce, Koch, 1835 (?) J Ixodes pnstidarnni, Lucas, 1866 ; Ixodes fodiens, Murray, 1877; Ixodes rnfus, Ixodes snlcatns, and Ixodes scinri, Koch. — F. V. T.] Ixodes holocyclus, Neumann, 1899. [Under the name /. holocyclns, Cleland {Jonrn. Trop. Med. and Hyg., 1913, xvi, No. 3, pp. 43-45) says that: ** This tick is common in man where there is dense scrub and tropical jungle along the east coast of Australia at certain times of the year. It may cause severe symptoms in children resulting in death." He records a child being attacked in 1884 which died, and another case from which 200 ticks were removed, the symptoms being weak heart, collapse, syncope, but the patient recovered under treatment ; again, in the same journal (pp. 188, 189), the case of a 4i-year-old girl who was bitten showed widespread muscular paralysis, and other cases resembling conium poison. [Taylor {Rep. Ent. Anst. Inst. Trop. Med., 191 1, p. 21, 1913) refers to this species as the scrub tick of New South Wales. The partially fed female has a dark reddish-yellow scutum and is almost as broad as long, punctations very numerous, equal and confluent in places, long white hairs on the lower half of each coxa. He records it as attacking man commonly, mentioning Kamerunga, Cairns district, Queensland, and Sydney, N.S.W., as localities. — F. V. T.] 500 THE ANIMAL PARASITES OF MAN Ixodes hexagonus, Leach, 1815. Syn. : Ixodes sexpunctatus, Koch, 1897; /. viilpis, Pagenstecher, 1861. Lives in the same manner as the foregoing; especially attacks hounds, but also other mammals and even birds. The difference consists in the shape of the legs, the shorter rostrum, and the larger size of the male. It also occasionally attacks man, but is usually confused with the previously mentioned species. [The synonyms of this species are as follow : Ixodes antiimnaliSj Leach, 1815; /. m//(^c^/, Audouin, 1832; /. rediivhis, Audouin, 1832; /. crennlatns, Koch; /. erinaceus, Murray, 1877; /. ricinus, Megnin, 1880. Two other synonyms are given above by Braun. [The female when fully replete is 11 mm. long, when fasting 3-86 mm. ; the shield is heart-shaped and punctate, body finely hairy ; palpi short and broad ; labium shorter, and tarsi of all the legs more truncate than in /. ricinus. The colour of the distended body is drab and somewhat waxy ; rostrum, shield and legs light testaceous. The male varies from 3*5 to 4*0 mm. long, and is reddish-brown in colour with lighter legs; the shield is punctate and leaves a narrow margin around the body ; the body is elliptical, almost as large in front as behind. There is a spine on the coxae of the first pair of legs, which is shorter than in the m.ale /. ricinus and longer than in the female. The genital orifice is opposite the interval between the second and third pair of legs. The fasting nymph is 176 mm. long, light bluish-grey, margined and transparent, with four large posterior intestinal marks joined together behind the shield and smaller ones extending to the front and sides. When fully distended it is uniformly brownish-white ; shield, legs and rostrum pale testaceous. The larva varies from o"88 mm. when fasting to 176 mm. when gorged. Its body is light, but gradually becomes darker, with similar intestinal marks to ricinus. [This tick is very common, especially on ferrets, stoats and hedge- hogs. It is also found on sheep, cattle, etc. The males do not generally occur in company with the females on the host. Pairing probably takes place on the ground. — F. V. T.] Genus. Amblyomma, Koch. Amblyomma cayennense, Koch, 1844. Syn. : Amblyomma mixtum^ Koch, 1844 ; Ixodes herrerce^ Duges, 1887 ; Amblyomma scuiptum^ Berlese, 1888. Characterized by the possession of eyes. The male measures 3-8 mm. in length by 3 mm. in breadth ; the female 4 mm. in length by 3 mm. in breadth, but when full of blood may become 13 mm. ^ Neumann, G. L., "Rev. de la fam. des Ixodides," /// Mem. Soc. Zool. France, 1899, xii, p. 129. HYALOMMA 501 in length and 11 mm. in breadth. They are common in the whole of Central America (Carrapatas); and attack mammals, amphibious animals and man.^ [This species was described by Fabricius. It occurs in Cayenne, Guiana, in Southern Texas, Florida, California, Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Bermuda, Cuba, Jamaica, Trinidad, Colombia, Venezuela, French Guiana, Brazil, Paraguay and the Argentine. It is called the silver tick. It frequently attacks man. Schwarz and Bishopp {Bull. 105, U.S. Dept. Agric, p. 158) heard of one man w^hose legs were well covered with suppurating sores and w^ho was ill from the attack of these ticks and the wounds produced by scratching, and records other cases of their swarming on man. Newstead {Ann. T.rop. Med. and Par., 1909, iii, No. 4, p. 442) records it as the worst pest to man in Jamaica. — F. V. T.] Amblyomma americana, Linnaeus. The so-called long star tick, from the silvery spot on the apex of the scutum of the female. It will attack any mammal and even birds and also man. It occurs in North America, and also in Brazil, Guiana and Guatemala. Its punctures frequently end in suppuration. In the Eastern and Southern States man is more frequently attacked by this species than any other. Moss-gatherers in Louisiana are badly attacked by it.^ It also attacks the milkers in dairies. Attempts to transmit Texas fever failed with this species. Amblyomma maculatum, Koch. The so-called Gulf Coast tick, of the Gulf Coast, occurs on birds,^ mammals and man, especially cattle, and attacks the ears. Genus. Hyalomma, Koch. Hyalomma aegyptium, L., 1758. Syn. : Acariis cegyptius^ L., 1758 ; Ixodes camelinus^ Fischer, 1823. A species frequently found in Africa, particularly in Egypt and Algeria, and which also occurs in France and Italy, as well as in Asia. Male 8 mm. in length, 4*5 mm. in breadth. Female up ta 24 mm. in length and 15 mm. in breadth. It infests large and small animals as well as human beings.^ [This is one of the largest ticks, nearly reaching the size of the bont tick. It is known in Africa as the bont leg-tick; all farm stock ^ Neumann, G. Y^.^Joc. cit.^ p. 205. 2 Morgan, " Ticks and Texas Fever," Louisiana State Bull. 55, pp. 134, 135, P^- 59- 3 Neumann, G. L., loc. cit., p. 285 ; Ronsisvalle, " Sui fenomeni morb. prodotti nel uomo da un Ixodide denominate Hyal. ceg.,'' Boll. Ace. Gioenia sci. vat., 1891, xvii. 502 THE ANIMAL PARASITES OF MAN is attacked, but sheep and goats suffer most. Only one generation appears to occur each year. The male is almost black with a pale marginal stripe; the replete female brown with irregular light blue stripes. It is abundant in parts of South Africa.] Genus. Haemaphysalls, Koch. Haemaphysalis punctata, Canestrini and Fanzago, 1877 — 1878. Syn. : Hcemaphysalis sulcata^ Canestrini and Fanzago, 1877 — 1878 ; Rhico- cephaltis exposiiicius, Koch, 1877 ; HcEinaphysalis peregriniis^ Cambridge, 1889 ; Herpetobia sulcata, Canestrini, 1890. [This species does not appear to be common. It occurs on sheep, goats, horses and cattle. I have seen a female taken from man in Britain. The female when fasting is 3*44 mm. long, when gorged 12 mm. long. Colour, reddish-brown, leaden -grey when gorged; dorsal shield deeply indented in front ; rostrum, shield and legs brownish ; body finely punctate, both above and below ; sexual opening opposite the coxae of the second pair of legs in both sexes. Palpi a little longer than the labium ; first segment short and narrow, second and third widened on the dorsal face. Coxae with a short, broad blunt spine ; tarsi short, terminated with a spur on the first pair. The male is 3*10 mm. long. Body rather narrow, yellowish to reddish-brown ; dorsal shield nearly covers the whole body; numerous punctures over the whole surface. Eleven indentations on the posterior margin of the body ; peritremes lighter in colour, large and comma-shaped. The three anterior pairs of legs with a short spine on the haunches, the fourth with a very long one directed backwards. The nymph varies from 2*5 to 3*0 mm., is oval, and light yellow to dark red in colour. Dorsal shield rounded with few punctations. No spur on tarsi, and sexual orifice nearly obsolete. Larva short and oval. Length 1*20 mm. — F. V. T.] Genus. Dermacentor, Koch. Dermacentor reticulatus, Fabricius, 1794. Syn. : Acariis reticulatus, Fabr., 1794; Ixodes reticulatus, Latreille, 1806; /. uiarmoratus, Risso, 1826. This tick is provided with eyes, but it is distinguished from Ixodes and analogous genera by the lack of the abdominal plastron in the male, which measures 5 to 6 mm. in length by 3-5 mm. in breadth. The female may attain 16 mm. in length and 10 mm. in breadth. It is found in the South of Europe, in Asia, and in America; it attacks chiefly oxen, sheep and goats, and occasionally man.^ ' Neumann, G. L., " Rev. de la fam. des Ixodides," J//w. Soc. Zool. France, 1897, x, p. 360. DERMACENTOR VENUSTUS 503 [This tick sometimes causes much annoyance to human beings. It was once most troublesome at Revelstoke. Specimens have recently been found on fowls, turkeys and pheasants in Kent. [Other synonyms are as follow : Cynorhcestes pichiSf Hermanur 1804; Crotonus variegatiis, Dumeril, 1829; I. pictus, Gervais, 1844; Deiinacentor albicolUs, Koch, 1844 — 1847; D. pardalinus, Koch, 1844 — 1847; ^' ferruglneus, Koch, 1844 — 1847; Ixodes holsattis, Kolenati, 1857 > Psetidixodes-holsatuSf Haller, 1882 ; Hcemaphysalis inarmorata, Berlese, 1887. [The female when fasting is 3'86 mm. long by 2 mm. wide. The body is depressed, larger behind and reddish -brown in colour. The shield is very large and extends to the level of the third pair of legs, with a few large and many small punctations, milky white, variegated with reddish-brown. Sexual orifice opposite the coxae of the second pair of legs. Coxae of the front legs are deeply bifid, the others with a moderate spine. When gorged light brown, and may reach 16 mm. When depositing eggs the female is mottled with dark brow^n above and below. The male is like the female. The shield is reddish-brown, variegated with a milky white pattern. Coxae of the fourth pair of legs three times the size of the third. There is a sharp backwardly pointing spine on the second palpal segment, also seen (but smaller) in the female. Length 4*20 mm. [According to Mr. Wheier this is a very variable species both in size and colour. It occurs in England on sheep, but not com- monly. It has probably been introduced into Britain. Besides the animals mentioned above it is also found on deer. — F. V. T.] Dermacentor venustus, Banks. [The Rocky Mountain tick fever tick. This species has been wrongly called Dermatocentor reticidaUis var. occidentalis. The correct name of the carrier of Rocky Mountain tick fever is Dermacentor venustus, Banks (Hooker, Bishopp and Wood, Bidl. 106, U.S. Dept. Agric, Div. Ent., p. 165). [The female is from i3'8 by 10 by 6*4 mm. to 16*5 by 11*4 by 6*9 mm. when gorged ; the male from 2*1 by 1*5 mm. to 6 by 3*7 by 1*4 mm. The male reddish-brown ; scutum with an exten- sive pattern of white lines, usually but little white on the mid- posterior region, legs slightly lighter than scutum, joints tipped with white. Female with scutum mostly covered with white, abdomen reddish-brown, legs as in male. The nymph when unengorged reddish-brown, when gorged dark bluish-grey ; the larva is yellowish- brown when unengorged, slate blue when engorged. The ova light brown, shinv and smooth. 504 THE ANIMAL PARASITES OF MAN [The chief wild hosts are the brown bear, coyote, woodchuck, rabbit, wild cat, badger and mountain goat for the larvae ; practically all small mammals act as hosts for larvae and nymphae, whilst the adults are seldom found on other than large domestic animals ; horses and cattle are preferred. It occurs in British Columbia, southward to Northern New Mexico, and from the foothills of the Rocky Mountains in Colorado to the base of the Cascade Range in Oregon and California; abundant in Western Montana, Idaho, Eastern Washington, Oregon, North Utah, West Wyoming and North-west Colorado. [Of great importance in the Bitter Root Valley of Montana, where a number of cases of fever occur each year, with a mortality of about 70 per cent. In British Columbia this tick causes tick paralysis in man and sheep. Only the adults seem to attack man and animals there (Hadwen and Nuttall, Parasitology, 1913, vi, No. 3, pp. 288- 297 and 298-301) according to the Canadian Medical Associa- tion Jonrnal, December, 1912. The symptoms are unlike spotted fever. For full details of this tick vide Bulls. 105 and 106, U.S. Dept. Agric] Dermacentor occidentalis, Neumann. This tick only occurs in the Pacific Coast region of the United States. Owing to the fact that it frequently attacks man as well as occurring in great abundance in Oregon and California, it is of con- siderable economic importance. It is spoken of as the wood tick, and in the regions where found is the most common tick to attack man. Hooker, Bishopp and Wood {Bnll. 106, U.S. Dept. Agric, Div. Ent., 1912, p. 189) state that a number of cases have been brought to their notice where the bite of this tick has caused considerable local inflam- mation, which in some cases required physicians' attention. It has been supposed to be connected with Rocky Mountain spotted fever, but it is doubtful if it is concerned in its transmission. The engorged female is steel grey, the dorsum with an olive-green surface colour, which covers the grey except in small spots, giving a mottled appearance. The unengorged males and females are reddish-brown, scutum covered with a whitish bloom, interrupted by many red punctures. The female is 9 by 6-i by 3-3 mm. to ii'S by y6 by 5-6 mm. ; the male 2*8 by i-6 mm. to 4*2 by 2-3 mm. The larvre are bluish-grey when engorged, reddish-brown when unengorged. The nymph is light brown, sides of scutum darker, and the intestines dark brown. It is confined to the Coast Range and Sierra Nevada Mountains in California and Oregon and the small mountain range to the south-west. MARGAROPUS, RHIPICEPHALUS, SPECIES OF ARGAS 505 Dermacentor variabilis, Say. The American dog tick has also been found on man, but it is of Httle economic importance as it is easily removed from its host. Genus. Margaropus, Karsch. Margaropus annulatus australis, Fuller. The so-called Australian cattle tick. Newstead ^ reports this as a great pest to man in Jamaica in its larval stage. Its chief hosts are cattle, horses, goats, sheep, dogs and rabbits. Margaropus microplus, Canestrini. Recorded by Aragao (Mem. Inst. Osivaldo Cruz, 1911, iii, fasc. 2, p. 163) as occurring in larval stage on man in Brazil. Genus. Rhipicephalus, Koch. Rhipicephalus sanguineus, Latreille, 1804. Syn. : Ixodes sanguineus, Latr., 1804; /. rufiis, Koch, 1844 ; Rhipicephalus limbatus^ Koch, 1844 ; Rh. siculus, Koch, 1844 ; Rh. stigmaticus, Gerstacker, 1873. Spread over almost the entire tropical and sub-tropical regions, occurring in Europe in the South of France and in Italy ; it infests dogs and more rarely sheep; oxen, cats, foxes and human beings are also attacked.^ Neumann's Table of Species of Argas. J Body elliptical (sides curved) 2. i Body oblong (sides straight), ending anteriorly in a point ... 7. J Body transversely oval vespertilionis. ( Body elongate oval 3. f Margin of body striated 4. t Margin of body formed by quadrangular areolae PERSICUS. {Body flat, integument plainly wrinkled ... ... ... 5. Body tumid, elongate ; integument finely wrinkled ; coxae of fourth pair of legs near anterior third of body ... ... hennanii. J Body oval, narrowed anteriorly rEflexus. 5 ( Body elliptical, blunt, hardly narrowed anteriorly 6. f Body twice as long as broad ... ... ... cucu77ierinus. \ Body hardly longer than broad transgariepinus. Dorsal integument with large polygonal depressions ; tarsi appearing bifid brumptii. Dorsal integument almost smooth ; tarsi not appearing bifid ... ... ... ... ... ... ... ... cBqualis. The Argantince are distinguished from the Ixodince by the head, which in the former is situated on the inferior aspect of the ' cephalothorax, while in the Ixodince it projects freely ; also by the very short proboscis, the small club-like palpi, the lack of suckers on the legs, as well as by the scutellum, which covers the entire back and is bent up round the borders. Two genera are dis- tinguished : Argas, Latreille, 1796 (Rhynchoprion, Hermann, 1804), and Ornithodorus, Koch, 1844, The species live on mammals, but more especially on birds. ' Ann. Trop. Med. and Par., 1909, iii, No. 4. ^ Neumann, G. L., he. cit., 1897, p. 385. 5o6 THE ANIMAL PARASITES OF MAN Genus. Argas, Lalreille. Argas reflexus, Fabricius, 1794. Syn. : Acarus 7'cjlexus, Fabricius, 1794 ; A. ina7'gi7iatus, Fabricius, 1794; Rhy7ichoprio7i coluDibcr^ Hermann, 1804. The European marginated tick, Argas reflexus (length of male 4 mm., breadth 3 mm., length of female 6 to 8 mm., breadth 4 mm.), is of a yellowish colour and has yellowish-white legs. The ingested blood shows red or brown through the intestine, which is provided with blind sacs. It lives in dovecots. It remains hidden during the day and at night crawls on to the sleeping pigeons to suck their blood. It has been observed in France, England, Italy, Germany, and Russia. Persons sleeping near infected dovecots, or in apart- ments formed from pigeon-lofts, are also attacked, even when the room in question has not been used for sheltering pigeons for years, as '^marginated ticks" can live in a fasting condition for a very long time. The bite sometimes gives rise to serious symptoms, such as general erythema and sudden oedema. [This pest more often feeds on the blood of man than is imagined. Blanchard states that he has received them from men's clothes in Strasburg. Boschulte, of Westphalia, re- cords these parasites in a bedroom inhabited by children and connected with a pigeon- house. The children were bitten during sleep on the hands and feet. The result of the bite was intense itching along the nerves, the bite ^ ^ , ^ only beini^ marked by a red spot. In a girl from the dorsal surface, the of 14 or 15, vesiclcs wcrc lormed Similar to intestine showing through thosc produced by burns, and in an old man the integuments. (After ^ n ,^., 1 • r 1 Pagenstecher.) an ulcer formed. Others record painful punc- tures and persistent oedema produced by this pigeon pest. It was once abundant in Canterbury Cathedral, and often caused much annoyance, I am told, to the worshippers ; the ticks falling down from the roof, where they were living, deriv^ed from the numerous pigeons that breed in the towers. This Acarus has enormous powers of vitality, living without food for months at a time.— F. V. T.] Argas persicus, Fischer de Waldheim, 1824. Of oval form and brownish-red colour. The male measures 4 to 5 mm. in length by 3 mm. in breadth ; the female 7 to 10 mm. in length by 5 to 6 mm. in breadth. It frequents the entire north-west ARGAS BRUMPTI 507 and north-east of Persia (the gerib-gez or malleh of the Persians, the miana bug of travellers). It lives concealed in houses and attacks man at night to suck his blood. Its bite is much dreaded, but the serious results may probably be attributed to unsuitable treatment of the wound or its invasion by bacteria. [This tick, sometimes called the tampan and wandlius in South Africa, is mainly a fowl parasite. Fow4s and ducks frequently die under its attack, particularly young ones, death being due to loss of blood. This tick remains attached to its host during its larval stage for about five days ; it then leaves and moults in concealment. In its subsequent stages it visits its host by night and remains for about an hour only, during which time it distends itself fully with blood. As a nymph it moults twice, not once as do the cattle ticks. This tick and other Argas become larger with each moult, but retain their same general appearance. The female visits the host every now and then, and, between, deposits eggs in sheltered crevices. About fifty to 120 are deposited at once. Four weeks seems a necessary period to intervene between visits to the host, and the interval may be extended to upwards of a year according to Lounsbury.'* [It is found in the Sudan, where Balfour has found granules derived from the seg- mentation of spirilla in their digestive tract. Fantham and Hindi have confirmed this. It has been assumed that these granules carry infection. [This so-called Persian tick, the miana, which is such a scourge to travellers in Persia, appears to infest the huts of natives in that country. It has been sent me from Quetta, where it has invaded houses to such an extent Fig. 362.— ^;x«.f perskus: . . i 1- • ii rrM • 1 ventral aspect. 7/1. (After the natives cannot live in them. 1 he virulence Megnin.) of its bite is probably due to the tick trans- mitting fever germs from natives, probably inured, to strangers, who would be susceptible. — F. V. T.] Argas brumpti, Neumann. [Found in Somaliland, by Brumpt, and in the Sudan. This tick attacks man as well as wild animals and produces a painful swelling, according to King,^ but as pointed out by that naturalist it probably relies on other than human food. — F. V. T.] ' «* Report of Government Entomologist, Cape of Good Hope, for 1899," 1900, p. ZZ- 2 "Fourth Report Wellcome Res. Labs.," 1911, p. 128. 32 5o8 THE ANIMAL PARASITES OF MAN Argas chinche, Gervais, 1844. This Acarus, a native of the temperate parts of Colombia, is very troublesome to man. It is probably identical with A. americaiuiSf Packard, which infests domestic fowls and turkeys, and occasionally also cattle, and is differentiated from A. reflexus by the sculpturing of the cuticle. Genus. Ornithodorus, Koch. Neumann's Synopsis of the Genus Ornithodorus is as follows : — Hypostome unarmed ; integument in nymph stage and partly in adult spinulose ... megnini. Hypostome armed with recurved teeth ; integument not spinulose ... ... ... ... ... ... 2. Camerostome with movable lateral flaps talaje. Camerostome without movable lateral flaps ... ... ... 3. ■ Anterior border of distal segments of legs with tubercles or festoons 4. Anterior border of segments of legs without tubercles or festoons ... ... ... ... ... ... ... ... 8. Body not much contracted anteriorly 5. Body pointed anteriorly ... ... ... ... ... ... 7. Tubercles of distal segments of legs higher than broad, distant 6. Festoons of distal segments of legs as broad as high, con- tiguous ... ... ... ... ... ... ... ... pavimefiiosus. Eyes present SAVIGNYI. 1 No eyes moubata, J Eyes present coriaceus. 1 No eyes turicata. f Integument with fine radiating wrinkles lahorensis, \ Integument granular 9. j Tarsi appearing bifid at apex furcostts. 1 Tarsi not appearing bifid at apex 10. ( Tarsi of first pair of legs with three dorsal tubercles, of other \ legs with one caTiestrinii. (^ Tarsi without dorsal tubercles or with only one 11. Tarsi of last three pairs of legs with pronounced dorsal protuberance tholozanii. . Tarsi of legs with indistinct dorsal protuberance errattcus. Ornithodorus moubata, Murray, 1877. An abundant African tick which is one of the caixiers of the spirillum of African relapsing fever and can also carry Filaria perstans (Christy). Its body is oval, yellowish-brown when young, greenish- brown when mature The integument is covered with mamillose tubercles. No eyes jid the stout legs granular above, the tibiae and tarsi fringed with tubercles on the upper side. Pocock ^ records it from 1 *• A System of Medicine," AUbutt and RoUeston, i, pt. 2, p. 195. ORNITHODORUS SAVIGNYI, CORIACEUS, TALAJE, AND TURICATA 509 Uganda and German East Africa, Congo and Angola, to Namaqualand and the Transv'aal in the south. It is called bibo in Uganda, moiibata in Angola, and tampan on the Lower Zambesi. It feeds on animals and birds as well as man. Its bite is very painful. This tick is found in native huts, living in cracks and crevices and in the thatch roofs. The female tick infected with the spirillum transmits the infection to the eggs and the next generation. They appear to be able to live without food a long time, and probably live for years. They lay their eggs in masses on the ground or in crevices, and when they hatch they are in the nymph stage with four pairs of legs. 0. moubata also occurs in Madagascar with recurrent fever (Lamoureux, Bull. Soc. Path, exot., 1913, vi, No. 3, pp. 146-149). Ornlthodorus savignyi, Audouin, 1827. At one time considered the same as the preceding species, but can be easily separated by the presence of two pairs of eyes. It is widely spread over Africa and has been found in South India and at Aden. In the Sudan it occurs in large numbers. King ^ records that a few miles N.N.E. of Khartoum 370 specimens were collected in two hours under a single tree by a well. It is found in Somaliland, where relapsing fever occurs and no 0. moiibata^ which it probably replaces as a transmitter (Drake-Brockman, " Rep. Col. Office," April 6 and April 16, 1913). It also occurs in Tunis, where the natives call it "tobbiah" (Weiss, Arch, de I' Inst. Pasteur de Tnnis, 1912, pt. 4, p. 226). Ornlthodorus coriaceus, Koch. Found in Mexico, Paraguay and California. Attacks man. Ornithodorus talaje, Guerin, 1849. An eyeless species with somewhat elongate pentagonal body found in Mexico and South America, called the '^chinche." A variety of it {coniceps) is found at Venice, etc., and another variety on various islands in the Indian Ocean and South Atlantic. Its bite is very painful to man. Ornithodorus turicata, Duges, 1876. Without eyes. Indigenous in Central America ; attacks human beings and pigs. The bite is painful and is often followed by serious consequences. [So virulent is this species that pigs put in an infested sty often die in a night. This " turicatas " of Mexico often reaches 6 mm. in length.— F. V. T.] ' "Fourth Report Wellcome Res. Labs.," 191 1, B, p. 129. 510 THE ANIMAL PARASITES OF MAN Ornithodorus tholozani, Laboulbene and IVregnin, 1882. Syn. : Argas tholozani^ Lab. and Meg., 1882. Without eyes. Males 4 to 6 mm. in length and 2 to 4 mm. in breadth ; females 8 to 10 mm. in length and 4 to 5 mm. in breadth. It especially attacks sheep. Native of Persia and Asia Minor. [This species is reputed as being very dangerous to man. It is locally know as the kene, or sheep-bug. In its fully gorged state it is deep violet.— F. V. T.] Ornithodorus megnini, Duges, 1883. Syn. : Argas viegnini^ Duges, 1883. Length 8*5 mm., breadth 5*5 mm. Native of Mexico. [Another synonym for this species is Rhynclioprion spinosiun, Marx. The adult males and females are grey to dark brown, the male somewhat the smaller ; female 5 by 3*5 by 2-5 mm. to 10 by 6 by 3*5 mm. The larvae at the seed tick stage are dark grey, turning to pink, then to a whitish grey when engorged. The nymph when young is blood-red in front, rest pearly white ; later they turn reddish-brown. [Intense pain may be caused by its presence in and around the ears. [Two specimens in the nymphal state were taken from the ears of a visitor to Cambridge by Dr. ]. Christian Simpson. They were supposed to have entered the ears when the gentleman was camping out in Arizona (Lancet, 1901, i, No. 4,052, p. 1198, fig. 3). [This species attacks the horse, ass, dog, cats and oxen, generally around the ears, and also attacks man. It is well known in the United States as infesting the ears of children (New York Ent. Soc. Journ., 1893, pp. 49-52). [It occurs in Texas, Arizona, New Mexico and California as well as Mexico, Brazil, and possibly many parts of South America ; and recently Bedford (*' Sec. Report Div. Vet. Res., S. African Union," 1912, pp. 343, 344) has shown it to occur at Vryburg and Fauresmith, in the Transvaal, on stock. It also occurs in the Sudan. — F. V. T.] Other Literature on Ixodida. (i) " Penetration de V Ixodes ricinns sous la peau de rhomme," Conipt. rend. Soc. de Biol.^ 1891, xliii, ser. 9, iii, pp. 689-691, R. Blanchard. (2) "Notas sobre Ixodidas brazileiros, " Mem. Inst. Oszvaldo Cruz, 1911, iii* fasc. 2, pp. 145-195, pis. II and 12, Dr. H. de Beaurepaire Aragao. Table of Brazilian Species. (3) ** Contribuicao para a sistematica e biolojia dos Ixodidas," Alein. Inst. Oswaldo Cruz., 1912, iv, fasc. I, pp. 96-120, pis. 2 and 3, Dr. H. de Beaurepaire Aragao. TYROGLYPHID^ 511 Family. Tyroglyphidae. Very small mites without eyes and without tracheae, with smooth skin. The males usually have a suctorial pore on either side of the anus, which is used during copulation, or suckers may be found in both sexes near the sexual orifice. The mouth parts form a cone with chelate cheliceras, and three-jointed pedipalpi ; the legs are usually short, have five segments with a terminal claw and suckers, or either one or other of these organs. The numerous species and genera live free and from choice in slowly decomposing vegetable and animal matter (cheese, cereals, flour, sugar, presel'ves, dried anatomical preparations, bacon, dried fruits and fungi), also in the corners of dwellings, elc. ; they incidentally get into or on to man, or are found in chamber utensils and in spittoons ; actual parasites are rarely found amongst them. [The chief genera are Tyroglyphus, Rhizoglyphus, Glyciphagus, Aleurobius and Histiogaster. The first three have typical characters referred to, but are distinguished from each other by the two former having the hairs on the dorsum smooth, whilst in Glyciphagus they are hairy, plumose, or feathered. Rhizoglyphus can be told from Tyroglyphus by having claws on the tarsi without any suckers ; Tyroglyphus has both claws and suckers. The larvae are hexapod and may become adult in the usual way by repeated moults, or they enter the so-called hypopial stage. In this the eight-legged nymph becomes quiescent, and during this stage it fixes itself to some insect or other animal by a patch of suckers on the lower surface of the hind end of the body, and is so carried from place to place. The hypopus does not feed and has a hard shell and short legs. When it has reached a new home it moults and development proceeds in the normal way. Canestrini and Kramer treat the TyroglyphidcB as a sub-family of the Sarcoptidce^ calling them sub-family Tyroglyphi7Ke,i\\Q other sub-families being Sarcoptince, CanestriniiticE and Analsmce. — F. V. T.] Sub-family. Tyroglyphinae. Genus. Aleurobius, Canestrini. Aleurobius (Tyroglyphus) farinae, de Geer (part), Koch. The male measures o'33 mm. in length by o*i6 mm. in breadth; the female o'6 mm. in length by 0*3 mm. in breadth. These mites possess five pairs of suctorial organs of a light colour ; the legs are reddish. Moniez observed them in Lille on the skin of labourers who had been unloading Russian corn. A few of the species generally mentioned under the designation of Tyroglyphus slro are probably the common flour-mite, which also occurs on dry cheese. [The farince of de Geer is an Aleurobius described by him in 1778 (" Mem. Hist. Ins.," vii, t. 5, f. 15, p. 97) as Acariis farincv. — F. V. T.] Genus. Tyroglyphus, Latreille. Tyroglyphus siro, L., 1756. (Defined by Gervais, 1844.) Male 0-5 mm. in length by 0*25 mm. in breadth ; female 0-53 mm. in length by 0*28 mm. in breadth; the males have two suckers on the tarsi of the fourth pair of legs. Penis straight, colour whitish or reddish. 512 THE ANIMAL PARASITES OF MAN Tyroglyphus longior, Gervais, 1844. White or yellowish, with two black spots on the abdomen. Male o*55 mm. in length, 0*28 mm. in breadth ; penis bent. Female o*6i mm. in length and 0*28 mm. in breadth. r. siro and T. longior live on dry cheese, in flour, on dried fruits, etc., and have been occasionally observed in the stools, urine, or pus of human beings, and also on their skin. The so-called vanillismus is to be attributed to these species. [T. siro and T.farincc of Schrank (non Geer) are the same. They are described under other names, such as Acarus lactis, Linn. ; A. favormrij Herm., etc. ; A. lactis in milk, farince in flour, and siro in cheese; and as A. dysenterice, Linnaeus (^' Syst. Nat.," ed. 12, pp. 1024-1767).] Fig. 363. — TyroglypJnis fariuLC : male. Enlarged. (After Berlese.) Fig. 364. — Tyroglyphus' longior^ Gerv. (After Fum. and Robin.) It is to these species that a case of dysentery was referred. Rolander, who studied under Linnaeus, was attacked by what was called dysentery. The complaint soon gave way to treatment, but eight days after it returned, soon disappeared, but again came a third time. All the time Rolander had been living like the other inmates of the house, who all escaped. Linnaeus, aware that Bartholemy had attributed dysentery to insects which he said he had seen, advised his student to examine his stool. The result was that innumerable mites were found to be present. Their presence was easily accounted for by the fact that they were found in numbers in a cup made of juniper wood from which the student alone drank of a night, and they were found to be of the same species. What this species is we do not know. Linnaeus called it Acarus dyseniericv, but it was the same as his Acarus siro. No records have GLYCIPHAGUS 513 occurred since. It cannot be, as Latreille supposed, the cheese mite, for they have been eaten by milhons since, and it is strange no such case has occurred again. [Tyroglyphus minor var. Castellani, Hirst, causes the copra itch in persons employed in the copra mills in Ceylon. The skin of the hands, arms, legs and even body becomes covered with pruriginous papules, papulo-pustules and pustules near the head. The eruption begins as a rule on the liands. The mites live in the copra dust. They produce dermatitis. Castellani produced the disease experimentally by rubbing copra dust containing mites on the skin of healthy people. Beta-naphthol ointment (5 to 10 percent.) proved useful in treatment (Jonrii. Trop. Med. and Hyg., December 16, 191 2, Castellani and Hirst). — F. V. T.] Genus. Glyciphagus, Hering, 1838. Glyciphagus prunorum, Her., and G. domesticus, de Geer. The Glyciphagi are differentiated from the Tyroglyphi in that the chitinous hairs on the body are fringed or feathered, and that they lack a furrow dividing the cephalothorax from the abdomen. They live under similar conditions to the Tyroglyphi and are occasionally found on man or in faeces. [Sugar merchants and grocers are frequently troubled by swarms of G. dojuestiais, which leave the stores when being handled, and especially shopmen, who handle sugar kept in small stores for some time. These are the Acari that cause that irritating temporary affec- tion known as " grocer's itch." — F. V. T.] Glyciphagus cursor, Gervais. Under this name Signor Moriggia figures a horny excrescence of great length growing from a woman's hand, and containing in its cavities quantities of Acarus. This species is really G. domesticus, de Geer. G. domesticus has also been described by Gervais (Ann. Sci. Nat.f 1841, ser. 2, xv, p. 8) as G. hippopodes. Glyciphagus buski, Murray.^ [This is a mite found by Busk and named after him by Murray. It was taken from beneath the cuticle of the sole of the foot of a negro in the Seamen's Hospital Ship on the Thames in 1841, in large sores of a peculiar character confined to the soles of the ' Cooper and Busk's Micros. Jonrn., 1842, and "Economic Entomology," Murray, p. 280. 5H THE ANIMAL PARASITES OF MAN feet. It appeared that the disease was caused by its burrowing beneath the thick cuticle. The disease was attributed to the wearing of a pair of shoes which had been lent to another negro whose feet had been similarly affected for nearly a year. The negro to whom the shoes were lent came from Sierra Leone. Mr. Busk stated that some water brought by Dr. Stranger from the River Sinoe, on the coast of Africa, contained one nearly perfect specimen, and fragments of others very similar to if not identical with this Acarus. Mr. Busk adds that he had been informed by Staff- Assistant Surgeon P. D. Murray that at Sierra Leone there is a native pustular disease called craw-craw — a species of itch breaking into open sores. [From Busk's original figure I see no reason to doubt that this is a Glyciphagus.— F. V. T.] Genus. Rhizoglyphus, Claparede, iJ Rhizoglyphus parasiticus, Dalgetty, 1901. The Rhizoglyphii are to be recognized by their short legs, which are beset with spines, and by the tarsi, which terminate in a claw. They live on plants, roots and bulbs, especially the bulbs of lilies. YiG. T,es.— Rhizoglyphus parasitiats. a,, male ; ^., female. Enlarged. (After Dalgetly.) This species has been observed on the feet of Indian coolies working in the tea plantations; they produce a skin disease which always commences with blebs between the toes, and which almost always extends to the malleoli, but not beyond. The HISTIOGASTER 5^5 Acari have an elliptical body, which is grey, but varies from greenish-yellow to greenish-brown when the stomach is full. Eyes are absent. The legs are composed of five segments and terminate with a claw. The males measure o'i8 mm. in length by o'o8 mm. in breadth, and possess genital and anal pores ; the females measure 0'2 mm. in length by 0*09 mm. in breadth.^ [This is also known as coolie itch and is common in Indian tea plantations. — F. V. T.] Genus. Histiogaster, Berlese, 1883. Histiogaster (entomophagus?) spermaticus, Trouessart, 1900. The genus Histiogaster, which also approaches the Tyroglyphince^ is characterized by the circumstance that the males possess suctorial pores used in copulation, as well as leaf-shaped appendages at the posterior end of the body. They feed on vegetables, especially on small fungi. Fig. 366. — Histiogaster (entomophagus ?) spermaticus : on left, male; on light, female — both from the abdominal aspect. 200/ 1. (After E. Trouessart.) This species has been described by Trouessart,^ who found numerous specimens, some adult, others in the developmental stage (larvae, nymphs), and ova, in the fluid removed by puncture from ^ Dalgetty, A. B., "Water-itch; or Sore Feet of Coolies," Journ. Trop. Med., 1901, iv, p. 73. ^ Trouessart, E. R., Comft. renJ. Soc. Biol., Paris, 1900, lii, pp. 742-744, 893, 894; Arch, de Par., 1902, v, pp. 449-459. 5l6 THE ANIMAL PARASITES OP^ MAN a cyst of the right testis. The males measure 0-25 mm., the females 0*32 mm., and the larvae o'l mm. in length. The author is of opinion that the animal — perhaps a fertilized female — was introduced by a catheter, and, as a matter of fact, it was afterwards found that the patient had once had the catheter passed in India while suffering from pernicious fever. It would here rather appear to be the case of a facultative para- sitism of an otherwise free-living species. Histiogastcr ciitoiiiopliagtts, Laboulbene, is found occasionally in collections of insects feeding on larger species containing much fat ; the species also occurs on dry cantharides; it appears to belong to the region of South Europe, where, however, it is widely spread. [Entomophagus occurs all over Europe and in America. It has been described under the following names : Acanis tnaliis, Shimer, 1868 {Trans. Illinois Hort. Soc.) ; Dernialeichus niali, Riley, 1873 {Rep. Ins. Missonri, v, p. 87) ; Tyroglyphns mali, Murray, 1877 (" Eco. Ent. Apt.," p. 275) ; T. coriicalis, Michael, 1885 {Trans. Roy. Micros. Soc, ser. 2, v, 3, p. 27, figs. I to 14) ; Histiogaster coriicalis, Canestrini, 1888 {Prosp. Acarof., iii, p. 397) ; H. alenrophagns, Sicherin, 1894, Canestrini, Prosp. Acarof., vi, p. 815. Trouessart's species is evidently distinct. — F. V. T.] Genus. Cheyletus. Cheyletus nnericourti, Lab. Acaropsis mericourti^ Moq. Tand. [This mite has been described from three specimens found in pus which flowed from an abscess in the ear of a naval officer, produced by inflammation of the auditory passage. Where the mites came from we do not know, as they were found near the Bank of New- foundland. This genus of Acari has enormous mandibles and a peculiar tracheal system ; two ungues and appendages to the tarsi. — F. V. T.] Family. Sarcoptidae (Itch Mites). Small mites without eyes and tracheae, and with delicate, transversely striated cuticle. The mouth parts form a cone, over which the shield-shaped upper lip protrudes ; the chelicerae are chelate ; the pedipalpi (or maxillary palpi) have three joints ; the legs are short and compact, and composed of five segments ; the terminal joints have pedunculated suckers (ambulacra) or a long bristle. The larviis are six- legged. They live on or under the skin of birds and mammals, on which they produce the skin disease known as scabies, or itch. [The Sarcoptid(E attack the hairs, feathers or epidermis of birds, animals and man, living as permanent parasites. The punctures they produce are followed by the formation of more or less thick crusts or scabs, beneath which the mites live and breed (so called scab, mange and itch). Most are oviparous, some ovoviviparous. The eggs are minute, ovoid, with a thin semi-transparent shell. They incubate in a few days, varying from two to ten or eleven, as a rule. Generally sarcoptic Second ,, „ 30 Third » 45 Fourth „ ,, 60 Fifth M 75 Sixth M 90 SARCOPTID^ 517 diseases lie dormant in winter and revive in spring and summer in man ; but in animals with long wool, such as sheep, they are most active during winter, although revival of active reproduction takes place in spring. [Speakmg generally, for the Sarcoptidce there are three distinct stages in the development of the male, four in the female, as follows : — [(i) The larva. In this stage only three pairs of legs occur. [(2) The nymph, in which a fourth pair of legs appear, and which thus approaches the adult ; but so far no sexual organs occur. Nymphs are of two sizes — the smaller being future miles, the larger females. [(3) The next stage in the female is the age of puberty^ the female now being provided with a vulvo-anal slit ; this so-cdW^di pubescent female is fertilized by the male. The male then dies. But the female again casts her skin and enters another stage — [(4) The ovigerous female — the egg-laying female— which has differently modified legs. [The rate at which these Acari breed is very great. Gerlach has found that roughly, in each Sarcopt gallery, a female produces fifteen individuals — ten femailes and five males — and that the progeny reproduce again in fifteen days. The table given below thus shows that one pair may produce the enormous number of 1,500,000 descendants in three months : — First generation after 15 days ... 10 females ... 5 males 100 ,, ... 50 ,, 1,000 ,, ... 500 ,, 10,000 ,, ... 5,000 ,, 100,000 ,, ... 50,000 ,, ... 1,000,000 ,, ... 500,000 ,, := ly^oOyOOO individuals. [These Acaririce are divided into three distinct sub-families, namely the Cyto- lichince^ Sarcoptince, Caiiestriniino'. [The Sarcoptince 2\o\\^ interest us here, and of the nine genera the three following are the most important : — [(i) Sarcoptes, Latreille ; Eusarcoptes. [(2) Psoroptes, Gerv. ; Dermatodectes, Gerlach ; Dermatocoptes, Fiirstenberg. [(3) Chorioptes, Gerv. ; Symbiotes, Gerlach ; Dermatophagus, Fiirst. ; Sarco- dermatocedes, Del. [The following are the main characters of these three genera: — \Sarcoptes — round or slightly oval; the two posterior pairs of legs being nearly or quite concealed beneath the body; the tarsi end in simple long pedicles, with ambulatory suckers. {Psoroptes — oval; the legs are all visible outside the niargin of the body; the ambulatory suckers are carried on long triangulated stalks ; the male has copulatory suckers and abdominal prolongations. [Chorioptes — oval ; legs long, thick, all visible ; ambulatory suckers very wide, carried at the end of simple, short pedicles. [Sarcoptes make channels or furrows beneath the epidermis, and in these the female lays her eggs. This form of acariasis is thus difficult to cure. It is the cause of human itch {vide Sarcoptes scabiei). [Psoroptes do not make sub-epidermic galleries ; they live and breed in colonies beneath crusts or scabs formed by the changes they produce in their host's skin. Sheep scab is a common type of disease produced by Psoroptes. This genus is of little importance as a parasite to man. [Chorioptes live as Psoroptes ; they also do not affect man. Otodectes, Can., affecting cats and dogs, and others occur, but do not affect man as far as we know at 5i8 THE ANIMAL PARASITES OF MAN present ("Demodicidae und Sarcoptidae," von Professor G. P. Kramer, Das Tierreich, 1899).— F. V. T.] Canestrini und' Sub-family. Sarcoptinae. Genus. Sarcoptes, Latreille. Sarcoptes scabiei, de Geer, 1778. Syn. : Acarus scabiei^ de Geer, 1778 ; A.psoricus^ Pallas, 1760 ; A. siro^ L., 1778 ;; Sarcoptes exulcerans, ? Linn,, 1758, Nitsch, 1818 ; S. hommis. Raspail, 1834, and Hering, 1838 ; S.galei, Owen, 1853 ; 5. communis^ Delaf. et Bourg., 1862; S. scabiei \2iX. hominis^ Megnin, 1880. The body is oval or nearly circular and whitish in colour, with trans- verse rovN^s of striae partly interrupted on the back. There are trans- verse rows of small bristles on the dorsal surface, and groups of trichomae on the front, sides and back. There are chitinous hairs at the base of the legs ; the two first pairs are provided with pedunculated ambulacra in both sexes, the two posterior pairs terminate each with a long bristle in the female ; in the male the third pair of legs terminate in a bristle, the fourth pair with a pedun- culated ambulacrum. The anus is situated at the pos- terior border of the dorsal surface. At one time numerous species were dififerentiated, according ta the form of the Acarus, tlie num- ber, position and size of the hairs and spines, even according to the hosts, etc. All these characteris- tics, however, fluctuate so con- siderably that absolute differentiation is impossible; the supposed species may be regarded in the same light as Megnin did, as varieties. It is also hardly possible to distinguish the mite of human scabies {S. hominis) from that of a number of domestic animals {S. squamiferus). It is best, therefore, to accept one single species {S. scabiei)^ which may give rise to different races or castes by living in the skin of man and mammals, but can pass from one host to the other. [Canestrini and Kramer, in their monograph of the SarcoptidcCy enumerate eighteen distinct species of this genus, from the dog, goat. Fig. 367. — Sarcoptes scabiei : female, dorsal aspect. 200/1. (After Furstenberg.) SARCOPTES SCABIEI 519 •camel, horse, ferret, lion, wolf, sheep, pig, etc., and two species parasites of man (scabiel and scabiei-criistoscv). There is no doubt that they are distinct species. — F. V. T.] The S. scabiel of man (S. scabiei var. Iwminis) (length of male 0-2 to o'3 mm., and breadth 0*145 ^^ 0*190 mm. ; length of female c*33 to o'45 mm., and breadth 0*25 to 0*35 mm.) lives in the tunnels that it excavates in the epidermis, and attacks by preference places with thin skin, such as between the fingers, in the bend of the elbows and knees, in the inguinal region, on the penis, on the mammae, but may also affect other parts. The tunnels, which vary from a few millimetres to a centimetre and more long, do not run straight, but are somewhat tortuous ; the female is found at the terminal end. The tun- nels contain the excrement and oval eggs (0-14 mm. in length) of the parasite ; the males are rarely met with, as they die off after copulation ; the females die after depositing their eggs. The six-legged larvae hatch out after four to eight days, and after about a fortnight, duringwhich time they change their skins three times and undergo metamorphosis, they begin themselves to burrow. Transmission from person to person rarely is effected through linen, but by direct contact (as in coitus) ; trans- mission can be artificially effected on horses, dogs and monkeys, but not on cats. The smaller S. scablei-cnis- toscv, Fiirstenberg, is the cause of the itch that occurs chiefly in Norway ; it is not certain whether this is a distinct species of itch mite. [This is quite a distinct species, which is recorded from Germany and France. Megnin {Parasitology ^ 1880, p. 165) described this as S. scablel var. lupl. The female is 140 fx long, 340 ^i broad ; the male is 170 yit long by 150 fji broad. In Science (March 3, 1893, p. 125) is recorded that at the Indiana Academv of Science Dr. Robert Hessler referred to ** a case of that extremely rare and almost extinct form of itch Fig -Sarcoples scabiei : male, ventral aspect. 203/I. (After Fiirstenberg.) 520 THE ANIMAL PARASITES OF MAN known as 'Norway itch,' the scabies norvcgica of Hebra, 1852. The afflicted man was covered with thick, creamy white, leathery scales ; some of these scales measured over an inch in diameter and JL in. thick. A constant shedding of scales went on, a handful being gathered daily. They were found full of mites and eggs and riddled with passages. Under treatment the mites were killed and the skin became normal. Dr. Hessler made a calculation of the number of eggs and mites, amounting to ova and shells 7,004,000, mites in all stages 2,009,000. — F. V. T.] The following forms may be transmitted from domestic animals to man : — (i) S, scabiei var. eqiii. Male, o'2 to 0-23 mm. long, 0*16 to 0*17 mm. broad. Female, 0*40 to 0*42 mm. long, 0*28 to o'32 mm. broad. The horse is the normal host. (2) S. scabiei var. ovis. Male, 0*22 mm. long, o"i6 mm. broad. Females, 0*32 to o*44mm. long, 0*24 to 0*36 mm. broad. This mite lives on sheep, and passes over to goats and human beings; it may also be artificially transferred to horses, oxen and dogs.^ (3) S. scabiei var. caprcc. Male, 0-24 mm. long, o'lSS mm. broad. Female, o'345 mm. long, 0*342 mm. broad. On goats, passing from them to horse, ox, sheep, pig and man. On the latter, in contradistinction to the varieties (i) and (2), it pro- duces a severe affection. (4) S. scabiei var. caineli. Frequently observed in man, chiefly in Africa. A few cases have been observed in Europe ; the affection induced by it is severe. (5) S. scabiei vviY. auchefticr. Male, 0*245 ^'^^' ^o^R' o"i82 mm. broad. Female, 0*34 mm. long, 0264 mm. broad. It lives on the llama, and may be transmitted to man. (6) S. scabiei var. suis, Male, 0*25 to 0*35 mm. long, o"i9 to 0*3 mm. broad. Female, 0*4 to 0*5 mm. long, 0*3 to 0*39 mm. broad. In the domestic pig and wild boar; occasionally also in man. The settlement, however, is usually of short duration. (7) S. scabiei var. canis. Male, 0*19 to 0*23 mm. long, 0*14 to 0*17 mm. broad. Female, 0*29 to o'38 mm. long, 0*23 to 0*28 mm. broad. In the house-dog, and also, not unusually, in human beings. (8) and (9) .S*. scabiei var. vuipis and S. scabiei var. leofiis of the fox and lion have likewise been observed on man. These are all distinct species and should read as follows : S. canis, Gerl. ; S. ovis, Megn. ; S. egtii, Gerl. ; .S". dromedarii, Gerv. {cameli, Megn.) ; S. auchenice, Raill. ; S. suis, Gerl. : S. vuipis, Fiirst. ; S. leonis. Can. Sarcoptes minor, Furstenberg, 1861. Anus situated on the back, legs short, pedunculated ambulacra broad ; living on cats (S. minor var. cati) and rabbits (S. minor var. cunicnli). In cats this mite usually lives in the cervical region, and thence spreads to the ears and head ; it usually causes the death of the infected animals; it is easily transferable from cat to cat, is difficult to transmit to rabbits, but once settled on them can easily ' [This mite produces the so-called '* black muzzle " of sheep.— F. V. T.] SARCOPTES MINOR 521 infect other rabbits. On the other hand, the transmission of the itch mite of the rabbit to the cat does not succeed. In man S. minor induces an eruption that disappears after about a fortnight. [S. minor, Fiirstenberg, i86t ('' Kratzm.," viii, p. 218), comes in Railhet's sub-genus Notoedres, 1893 {'' ZooL," ed. 2, p. 660). Canes- trini raised this to generic rank in 1894 (Pros/). Acarof.,\\, p. 724). [There are three species : (i) iV. notoedres, Mcgnin = Sarcoptes alepis, RaiUiet and Lucet {Compt. rend. Soc. de BioL, 1893, xlv, p. 404), and Sarcoptes notoedres var. mnris, Megnin {Parasitology, 1880, pp. 172-174). This occurs on the black and brown rats and the water-vole. [(2) N. cati, Hering, 1838 {N. acta. ac. Leop., ii, 18, xliv, p. 605, figs. 9 and 10), = Sarcoptes minor, Fiirstenberg (" Kratzm.," 1861, viii, p. 215). Found on the cat in Germany, France, Italy, and Britain. [(3) iV. cnniculi, Gerlach, 1857, ^'Kratzm.," iii, figs. 20, 21. It lives on the rabbit and is found in Germany and France. — F. V. T.] Fig. 369. — Sarcoptes minor var. call : on the left, female (lying on its abdomen) ; on the right, male (lying on its back). (After RaiUiet.) The itch mites of domestic animals, which belong to the genera Psoroptes (= Dermatodectes = Dermatocoptes) and Chorioptes (Symbiotes = Dermatophagus), as a rule do not infest and live on man, even when artificially transmitted. It is, however, possible for this to occur. Moniez ("Traite de par.," 1896, p. 559) mentions that a species of Chorioptes — probably C^. bovis — had been found on man, as had also Demodex folliculorum. This author also includes Derniatophagoides scherevieteiuskyi, Bogdanofif {Bull. soc. imp. d. natural., Moscou, 1864, xxxvii, p. 341), which has repeatedly been found on man in Moscow and Leipzig (Zurn, Ber. d. ined. Ges., Leipzig, 1S77, p. 38), as Chorioptes bovis. Other References to Scabies crustoscz and norvegica, etc. (1) "Ein P'all von Scabies crustosa norvegica,'" Wiirzb. med. Zeitschr., 1, pp. 134-139, pi. 3, H. Bamberger. (2) " Ueber die Kratzmilbe {Acarus scabiei),'" Notiz. a. d. Geb. d. Nat. u. Heilk., Weimar (1913), xlii (11), Oct., pp. 161-166 (1834), de Blainville. (3) " Rapport sur le ciron de la gale {Acariis scabiei),''^ Ann. de Mus. d' Hist, nat.y 1831 ; Parasitology, iv, pp. 213-232, dc Blainville. 522 THE ANIMAL PARASITES OF MAN Family. Demodlcidae (Mites of the Hair-follicles). Small Acarina, elongated in worm-like fashion, with annulated abdomen, and without eyes or tracheae. The mouth parts consist of a suctorial proboscis and three-jointed palpi ; the legs are short, and have three segments with small terminal ungues. The anus is situated on the anterior border of the abdomen ; oviparous ; the larvae have six stumpy legs. These mites live in the hair-follicles of mammals. Genus. Demodex, Owen. Demodex folliculorum, Simon, 1842. Syn. : Acarus folliculoriini^ Sim., ] Macrogaster platypus^ Miescher, 1843; Steatozoon folliculorum^ Wilson, 1847. 843; Demodex folliculorum^ Owen, 1843; Simonea folliculoi^um^ P. Gervais, 1844 ; As in Sarcoptes sccibiei, numerous varieties of this species are known ; the form parasitic on man lives in the hair-follicles, the meibomian and sebaceous glands, and hardly ever causes incon- venience; the male measures o'3 mm. in length and the female about 0*4 mm. in length. The eggs o'o6 to o'o8 mm. in length, 0*04 to 0*05 mm. in breadth, and are thin-shelled. The creatures are always attached with the head end downwards in the parts mentioned ; they are most frequent in the sebaceous glands of the face, by the nose, lips and forehead, but they may be present on the abdomen and on other parts of the body. They may occasionally obstruct the excretory gland ducts, thus causing infiammation of the gland (comedones) ; their agglomeration in the meibomian glands sets up inflammation of the margins of the eyelids. There are generally only a few specimens in a gland. According to some statements Demodex occurs in 50 per cent, of mankind and even in children ; they survive the death of their hosts by several days. Fig. 2>1o. — Demo- dex follictilorum of the dog. (After Megnin.) The variety living in the dog {D. folliculorum var. canis) is smaller than the variety living in man, and produces a skin disease resembling scabies in these animals. According to Ziirn they may also live on man ; nevertheless, no other investigator has recorded a similar observation, and attempts at artificial infection have proved negative.^ [Ten distinct species of Demodex are given by Canestrini and Kramer (" Demodicidae und Sarcoptidae," Das Ticrreich, 1899, vii). The species are certainly distinct. [The species living on the dog (D. canis, Leydig, 1844) is cosmo- politan. According to the British Medical Journal (February 22, 1913, ' [This mite causes what we know in England as red mange in dogs. — F. V. T.] PENTASTOMIDA 523 p. 407), dog mange may be caught by humans. Whittield and Hobday describe in the Veterinary Journal seventeen cases which have come under their observation. — F. V. T.] Order. Pentastomida. Family. Linguatulidae. Aradmida grfeatly altered in consequence of their parasitic manner of life ; for a long time they were regarded as helminthes. The body is elongated, vermiform, flattened or cylindrical, and more or less distinctly annulated. The head, thorax, and abdomen are not defined from each other (fig. 371). The elliptical mouth, surrounded by a chitinous ring, is situated at the anterior end, on the ventral surface, and the intestine leading straight through the body opens at the posterior end. Two retractile hooks are at the sides of the mouth (fig. 372) ; these are usually considered to be the terminal joints of two pairs of legs, but it appears to be more correct to regard them as the remains of the antenna? and palpi (Stiles). According to this opinion, the legs in the adult state are completely degenerated. The nervous system is reduced to an oesophageal ring. No organs of sense are recognizable except the papillae at the anterior end. There are neither organs of circulation nor of respiration.^ The sexes are distinct. In the small male the sexual orifice is situated ventrally in the anterior part of the body ; in the female it is placed near the anus. The Linguatulidcc lay eggs, and from each ^%%^ after being conveyed into an inter- mediate host, a four-legged larva, with rudimentary mouth parts, hatches out. It goes through a series of metamorphoses, and passes through a second larval condition, which, however, possesses the essential characteristics of the fully developed form. Sooner or later it migrates during this stage, and reaches its final host, mammal or reptile, in the nostrils or lungs of which the adult Linguatulidce live. I [As adults they live as internal blood feeders in various birds, reptiles and mammals, especially in the nasal and respiratory- passages. The larval stage occurs in another host in an encysted condition ; this host is usually an animal preyed upon by the species in which the sexual forms are found. The larvae bore through the walls of the host's stomach and enter liver and spleen or brain, where they encyst ; here they grow until they assume almost the appearance of the adult. These encysted larvae on being eaten later make their way into the nasal passages and lungs, where they mature. Both adults and larvae occur in man, as mentioned later. [Three genera are recognized in this family : — [(i) Lingiiafula. — Body flat, annulated. Adults live in the nasal sinus. [(2) Porocephalus. — Body cylindrical, elongate, with often deeply cut rings. Adult in respiratory organs of snakes, larvce in animals and man. [(3) Reighardia.—Q,y\\ndx\Q.d\^ but not ringed. Not found in humans. — F. V. T.] ' What are designated as stigmata in the Linguatulides are the orifices of sebaceous glands. 33 524 THE ANIMAL PARASITES OF MAN Genus. Linguatula, Frohlich. Linguatula rhinaria, Pilger, 1802. Syn. : Tcenia rhinaria^ Pilger, 1802 ; Polysto7na tcrmoides^ Rud., 18 10; Linguatula tcrnioides, Lam., 1816 ; Pentastoma ia-nioides^ Rud., 18 19. The male is white in colour, 18 to 20 mm. in length, anterior portion 3 to 4 mm. in breadth, posterior part 0-5 mm. in breadth. The female is of a yellowish colour, 8, 10, or 13 cm. long, anterior part 8 to 10 mm. and posterior part 2 mm. wide. The brownish eggs can be seen in the median line. The body is elongated, rather flat, and exhibits about ninety rings or segments with crenellated borders. The hooks round the mouth are strongly curved and are articu- lated to a basilar support. Eggs oval, 0*09 /a in length, 0*07 yLt in breadth. L. rhinariay in the adult con- dition, lives in the nasal cavity and frontal sinus of the dog, wolf, fox, horse, goat, and occasionally of man ; it causes severe catarrh, epistaxis and suppuration. Development. — The ova, which are found in masses in the nasal mucus, already possess an embryo ; they are expelled with the nasal secretion, and are swallowed by herbivorous mammals with their food, mostly by hares and rabbits, but also by sheep, goats, oxen, horses, antelopes, fallow^ deer, pigs, cats, and occasionally also by human beings. The young larvae hatch out in the stomach; they possess a thickened an- terior body with rudimentary mouth parts and two pairs of limbs ; the body gradually tapers to a short tail. The larv^ of the Linguatulidcv bore through the intestinal wall and reach the liver, more rarely the mesenteric glands, etc. ; they Here become encysted and enter a sort of pupal stage in which they lose their limbs ; after several moultings and gradual growth the second larval stage, having the appearance of the adult Linguatula, Fig. ^71.— Lingua- tula rhinaria : fe- male. Natural size. Fig. 372.— Larva of Linguatula rhinaria {Pentastoma denticula- tum). Enlarged. (After Leuckart.) LINGUATULA RHINARIA 525 sets in. About five to six months after infection the creatures have become 4 to 6 mm. long, possess eighty to ninety rings, which have a series of fine points on their posterior border; the mouth and intestine are formed, the sexual organs mature and the two pairs of hooks are near the mouth. This larval stage (fig. 372) has been known for a long time, but it was regarded as an independent species of animal, and therefore had a separate name {Liiigitafiila scrrata, Fr. ; Pentastonia denticnlatiun, Rud., etc.). Later these Linguatula larvae make an attempt to escape from their hosts, and this, of course, can only be effected by means of an active migration ; they leave the cysts, and according to their respective positions in the abdominal or pleural cavities they reach the bronchi Fig. 373. — Linguatula rhinaria: on left, eggs in gelatinous covering, iio/i. On right, first larval stage. 300/1. (After M. Koch.) or the intestine, and finally pass out ; they may be again sniffed up by dogs and settle in their nasal cavities. Still this outward migra- tion does not appear to be necessary for further development. A portion of the larvae gain access to the nasal cavities directly through the trachea, and thus herbivorous mammals certainly become directly infected. In most cases the infection of dogs, wolves and foxes, that is, of carnivorous mammals, takes place through consuming the bodies of mammals, or parts of them, such as the liver and lungs, which are alTected with the second larval form ; in any case most larvae obtain access first to the stomach of their host, from here they make an. active migration through the oesophagus to the oral and nasal cavities, in which they settle. It is possible also that ^26 THE ANIMAL PARASITKS OF MAN the same larvae which are free in the oral cavity when the food is being eaten migrate into the nasal cavities. After being stationary a fresh skin is formed and the spine-bearing cuticula are thrown off. The male attains its full size in the fourth, and the female in the sixth month. The duration of life is stated to be from fifteen months to several years. L. rhinaria has been observed in man in the adult as well as in the larval condition {Pentastoma denticulatum). Zenker first called attention to the occurrence of the larva in man, having found it nine times in the liver in i68 autopsies. Heschl found it twice in Vienna in twenty autopsies, Virchow found it in Wiirzburg and Berlin, Wagner in Leipzig (lo per cent.), and Frerichs in Breslau five times in forty-seven autopsies. The parasite is nmch less frequent in Switzerland. According to Klebs, one case occurs in 900 autopsies, and according to Zaeslin two cases occurred in Basle to 1,914 autopsies. In the Seamen's Hospital in Kronstsidt P. den^i'cu/a/i^m has been found six times in 659 autopsies. It was almost always the liver that contained one or a few specimens. The parasite was very rarely found in the kidney or spleen, or encysted in the intestinal wall. The adult L. rhinaria is far more rarely observed in man. A case reported by Landon that related to a blacksmith of Elbing is particularly interesting. This man accompanied the campaign of 1870; he soon, however, fell ill with pains in the liver, accompanied by icterus and intestinal disorders. Soon after the war, and after the symptoms were reduced to icterus and weakness, bleed- ing of the nose set in and continued with slight intermissions for seven years ; an unpleasant sensation of pressure in the left nasal cavity set in, with inflammatory swelling of the mucous membrane. At last, in the summer of 1878, when the pressure in the nose had considerably increased, a Linguatula was expelled from the nose with a violent attack of sneezing, and lived for three days longer in water. The bleeding of the nose then ceased and the patient soon recovered. There can be no doubt that the first illness was connected with the invasion in the liver of numerous larvae of Pentastoma, and disappeared after their encystment ; one or a few of these must subsequently have found its way to the nose and settled there. Genus. Porocephalus. Porocephalus constrictus, v. Siebold, 1852. • Syn. : Nematoideutn honmiis^ Diesing, 185 1 ; Pentastomuni constrictuin^ V. Sieb., 1852 ; Porocephalus constrictus, Stiles, 1893. Porocephalus is distinguished from t inguatula by its cylindrical body and by certain internal structures. Porocephalus constrictus is at present only known in its larval stage. It is milk white in colour with golden-yellow booklets. Number of rings, twenty-three. Length 13 mm., breadth 2*2 mm. There are no prickles on the posterior border of the annulations of the body. This species was first discovered by Pruner encysted in the livers of two negroes in Cairo. Bilharz reported two further cases in which the parasites were encysted in the liver and in the mucosa of the intestine ; a few other observa- tions have been made by Fenger, Aitken, Giard and Chalmas. Aitken's report deals with soldiers of the British Colonies in Africa. The parasites were discovered in POROCEPHALUS CONSTRICTUS 527 the liver as well as in the lung, and appear to have been the cause of death in one case (pneumonia, peritonitis), Pruner has found the same parasite also in the Hver of the giraffe. It has recently been assumed that Porocephaliis constrictus is the larva of Pentasto7na fnoniliforme, Diesing, 1835, that attains a length of 70 mm. and lives in the lungs of African Pythonides. The larva is known to have been ejected from monkeys {Cercopithecus albo ocularis ^ Cynocephaliis maifnon)^ from the giraffe {Cajnelopardalis ^iraffd)^ from a species of hyaena {Proteles cristatus)^ and should be expected to occur frequently in smaller mammals which have been swallowed by African serpents of enormous size. [The three species of Pcntastoniidce, or tongue worms, found in man are LinguaUtla serrata, FroHch ; Porocephaliis aruiillatuSy Wyman ; and Pentastoma iiionilifon/iis, Diesing. [(i) Lingiiatiila serrata has been referred to under a great number of names. ^ It is a frequent parasite in dogs, oxen and sheep ; as an aduh in the dog and also in the fox and wolf. The nymphal stage is found in rats, hares, rabbits, the horse, oxen, sheep, goats, pigs, camels, deer, the African and long-eared hedgehogs, porcupine, guinea-pig and peccary. In man it is found in both adult and nymphal stages. Sambon says the nymphal stage is of frequent occurrence, but is usually overlooked. Zenker, who first found it in man, obtained it in nine out of 160 post-morteuis, usually encysted in the liver. It is then said to be harmless. Landon, in 1878, found the adult in man, the patient suffering from epistaxis for about seven years ; in the end during a fit of sneezing the living parasite was ejected through the nostril. This case is of particular interest as it appears to suggest that this Acarid may now and then pass its entire development in the same host, or at any rate may actively migrate from the liver to the nasal cavities after a period of encystment in the liver or elsewhere, which has also been observed in herbivorous animals {vide also p. 526). [It is recorded from man in Central America (Darling, Bull. Soc. Path, exot., 191 2, v, p. 118; and again Arch. Int. Med., 1912, v, p. 401), also from Rio de Janeiro {Mem. Inst. Oswaldo Cruz, 1913, fasc. ii, p. 125) by Faria and Travassos. [(2) Porocephaliis aruiillatus, Wyman, is also known under a variety ' [Synonymy given by Sambon : Adult form, Tenia lanceole, Chabert, 1787; Ver rhinaire, Chabert, 1787; Tcenia rhinaris, Pilger, 1805; Tania lanceolata, Rudolphi, 1805; Cochlus r/iinariiis, Rudolphi, 1805; Prionoderma rhinaria^ Rudolphi, 1808; Polyslovia t(tnioides, Rudolphi, 1809; Linguatula tcEuioides, Lamark, 1816 ; Prionodervia lanceolata, Cuvier, 1817 ; Pentastoma ta;moides, Rudolphi, 1819; Linguatnla lanceolata, de Blainville, 1828; Lingiiatiila rhinaris, Railliet, 1885 ; Linguatnla caprina, R. Blanchard, 1900. Nymphal form : Linguatnla serrata, Frolich, 1789; Tcvnia capraa, Abildgaard, 1789; Tcvtiia caprina^ Gmelin, i8co ; Poly stoma serrata, Gueze, 1800 ; Halysis caprina, Zeder, 1803 ; Linguatula denticulata, Rudolphi, 1805 ; Echinorhynchus caprece, Braun, 1809; Tetragulus caprice, Bosc, 1810; Pentastoma denticulatum, Rudolphi, 1819 ; Pentastoma e/narginatum, Rudolphi, 1819; Pentastofnafera, Creplin, 1829; LAnguatulaferox, Gros, 1849. 528 THE ANIMAL PARASITES OF MAN of names.^ This species is widely spread over tropical Africa. The adult sta<:^e is found in pythons and puff-adders, the nymphal in the chimpanzee, Sykes monkey, mandrill and other monkeys, the lion, leopard, banded ichneumon, Aard wolf, dog, black rat. South African reedbuck and the giraffe. The adult has never been found in man or any mammal. No fewer than sixteen cases of the nymphal form,. Sambon tells us, have been found in man, and it is probably much more widespread than at present know^n. So far it has only been found in the African natives. This species has sixteen to seventeen body rings in the male, eighteen to twenty-two in the female, and the body does not taper as much as in the next species. [(3) Peiiiasionia uioiiiliforjiiis, Diesing,'Ms an Oriental species, found in India, Indo-Chma and South China, and the Malay Archipelago. The adult occurs in both the Indian and reticulated pythons. The nymphal stage has been found in monkeys, the tiger, the civet and the Indian otter. [The nymph has twice been found in man ; in one case in the liver of a Filipino, the other in the serous coat of the small intestine of a native of Sumatra. [This species can be told by the female having twenty-nine to thirty-three body rings, the male twenty-six, and the annulations are more bead-like and less prominent than in the African species. [In addition to these three, Sambon thinks it probable that others occur m man. — F. V. T.] Other References to Pent astomi dee. (i) "Die Wanderung des Pentastomum deniiculahim beim Rinde," Centralbl. f. Bakt . u. Parasitenk., Jan. 2, 1889, v (i), pp. 1-5, V. Bates. (2) "II Pentastoma vicnilifor7ne. Dies., nella pantera," Med. -vet. Torino, 1877, 4 s., vi (12), pp. 529-532, R. Bassi. (3) " On the Organization and Development of Linguatula (Pentastoma), accompanied with the description of a new species from the abdominal cavity of the mandrill," Ann. and Mag. Nat. Hist., 1848, 2 s. ii (7), 2, pp. 69-70, v. Beneden. (4) "De la Linguatula ferox [Pentastoma denticulatum aut serratnm),'' Bnll. Acad. roy. d. Sci. d. Belg., 1855, xxii, pt. I (I), pp. 4-10, V. Beneden. * Adult form as Linguatula armillata, Wyman, 1847 ; Pentastomum poly zonum , ITarley, 1856; Pentastoftmmarmillatum, Leuckart, i860; Pentastomum armillatum, Diesing, 1864; Porocephaliis armillatus. Stiles, 1893 ; Porocephalus polyzonus, Stiles, 1893 : Porocephalus moniliformis, Neumann (in part), 1899. Nymphal form : Linguatula diesingii, van Beneden, 1849; Pentastomum euryzomim, Diesing, 1850; Nematoideum hominis, Diesing, 185 1 ; Pentastomum constrictum, von Siebold, 1852 ; Linguatula constricta, Kuchenmeister, 1855 ; Pentastoma leonis,V^e6\., 1863; Pentastoma fornatum, CohhoXei, 1879; Pentastomum protelis, Hoyle, 1883 ; Porocephalus constrictus. Stiles, 1893 5 Linguatula constrictor, Galli-Valerio, 1896; Pentastomum diesingii, Shipley, 1898. 2 The synonomy is as follows :— Adult form : Pentastoma moniliforme, Diesing, 1835 ^ Linguatule mofiiliforme, Megnin, 1880 ; Porocephalus moniliformis, Stiles. Nymphal form : Pentastoma fornatum, Creplin (in part), 1849 ; Pentastoma wedlii, Cobbold, 1866 ; Penta- stoma aonycis, Macalister, 1874 ; Porocephalus armillatus. Stiles (in part), 1908. INSECTA 529 (5) "Note sur quelques pentastomes," Btill. Acad. roy. d. Set. de Belg., 1857, 26, 2 s., ii (5), pp. 29-30, V. Beneden. (6) " Ueber das Petitastoma in de gekrosdrusen den Schafe," Repert. d. Thierh. Stuttg.^ 1861, xxii, pp. 37-38, Collin. (7) " Eine Linguatula aus der Mesenterialdriise des Schafes und Dromedars als zweites ungesche. Stadium von Pent, taenioides ,'' Notiz. ti. Tagsber. 11. d. Geb. d. Nat, u. Heilk. Jena, 1862, v, pp. 127, 128, Colin. B. INSECTA (Hexapoda). Three separate regions can always be distinguished in the body of insects, namely, the head, thorax and abdomen. The head is a roundish unsegmented capsule and possesses four pairs of appendages. The first pair are the various shaped feelers (antennae), which are placed on the superior surface of the head next to the eyes ; then more ventrally placed a pair of upper jaws (mandibles) without palpi and without articulations ; they are powerful masticatory organs.^ The first pair of lower jaws (maxilhe) are jointed and bear a palpus (palpus maxillaris) ; the second pair of maxillae are soldered together and form the lower lip (labium), and likewise carry a palpus labialis on each side. The upper lip (labrum), as well as the other parts (which, however, are only appendages), belong to the mouth, which IS really formed of a number of closely united pieces. The mouth parts are modified according to the functions required of them. Coleoptera, Neuroptera^ and Ortho- ptera have biting or masticatory mouth parts which conform with the scheme described above. In the licking mouth parts of the Hymenoptera the maxillae and under lip are considerably elongated, while the mandibles retain their form and are used for triturating the food ; in the Lepidoptera nearly all the mouth parts are shortened except the maxillae, which form a long and sometimes spirally rolled suctorial proboscis ; the Dipiera and Rhynchota have piercing and sucking mouth parts. The mandibles and maxillae are metamorphosed into needle-like structures, while the suctorial apparatus is formed by the labrum. The thorax consists of three segments, which are frequently united ; ventrally it carries three pairs of legs, which consist of a definite number of articulated pieces joined together. Their form also changes according to their function, so that legs for running, walking, digging, swimming, jumping, and preying are seen. A pair of wings are respectively attached to the last and last but one thoracic rings, and these may be traced back, not to metamorphosed appendages, but to tracheal branchia. They are composed of chitinous membranes supported by branched structures (veins or ribs). Their size and formation vary ; they are seldom of equal size and form {S europterd) ; often the posterior wings are larger than the anterior wings, the former then only serving as protective coverings for the latter {Coleopterd), or the anterior wings are larger {Lepidoptera), or the posterior wings are shortened or are entirely absent {Dipiera); and finally there are insects in which both pairs of wings are lacking.^ The abdomen retains its segmentation, but, with the exception of a few groups related to the primitive forms of insects, has no appendages in the imago condi- tion ; the abdomen usually consists of ten segments, on the last of which the anus is situated. ' [The mandibles are only powerful masticatory organs in biting-mouthed insects {Mandibulata) ; in the sucking or piercing-mouthed insects they may be absent, or in the form of needle-like stylets {Haustellata).—Y . V. T.] ^ [As in the order Aptera, which includes the Thysanura and Collembola, and also exceptions in other orders, as the fleas amongst Diptera, the Mutillus and ants amongst Hymenoptera. — F. V. T.] 530 THE ANIMAL PARASITES OF MAX We need only observe the following characters in considering the anatomy of insects : — The EPIDERMIS consists of the chitinous cuticle, which is separate from the cellular layer beneath (hypodermis) ; the various appendages are supported by the chitinous layer. The INTESTINAL CANAL usually consists of the anterior, median and terminal intestine, and as a rule passes straight through the body ; salivary glands discharge into the anterior part, and, in some cases, yield a stiffening secretion which serves for spinning webs ; numerous or scanty hepatic tubes are appended to the median intestine, while on the border between the median and terminal intestine open four to six long tubes (vasa malpighiana), which act as urinary organs. Finally the end portion of the intestine carries various glands (anal and rectal glands, etc.). The CENTRAL NERVOUS SYSTEM agrees in structure with that of the Annelids, but is more highly developed. The pharyngeal ring surrounds the front part of the intestine ; the sensory nerves originate from its superior pharyngeal ganglia and are the seat of the higher psychical functions; the inferior pharyngeal ganglia govern the mouth parts, and in addition appear to regulate the movements (cerebellum) of the vertebrates. The chain of GANGLIA lying on the ventral side of the abdomen consists primi- tively of pairs of ganglia corresponding with the twelve segments, which are connected by longitudinal and transverse commissures. But many changes in the ganglia may be seen in insects caused by partial or entire amalgamation of single ganglia, so that in a few cases only one abdominal ganglion is present. In conclusion, a definite INTESTINAL NERVOUS SYSTEM is always present. Of the organs of sense the FACETTED EYES, situated at the sides of the head, deserve special mention, as do also the ORGANS OF TOUCH and SMELL, situated on the antennae, and the organs of hearing and taste, or finer sensations, situated at the mouth and in the buccal cavity. The sounds emitted by insects are, as a rule, produced by the friction or beating of certain chitinous parts, but sounds are also produced in breathing (flies). The organs of respiration, the so-called trachea?, are highly developed ; there are openings (stigmata) at the sides of the body which draw in air by means of the active participation of the muscles of the body. The number of stigmata varies between two and ten pairs ; the tracheae themselves branch off from the trunks in the most varied manner, and carry air to the internal organs. The colourless BLOOD circulates between the tissues and organs, and is kept circulating by the contraction of a chambered dorsal vessel provided with ostia, and which terminates in a short aorta opening at the anterior end. Insects are SEXUALLY distinct ; their sexual glands are in pairs and have a tubular structure, but the testicular tubules are united together by a capsule into an oval testicle ; exceptionally, also, the excretory canals are double, as also the sexual orifices ; usually the paired canals unite into a single oviduct or spermatic duct which terminates at the posterior end of the body after receiving the products of various glands. As to the history of the development of insects, all that is necessary to mention here is that the young hatched from eggs only exceptionally (as in Apterygota) resemble the adult parent (insecta ametabola) ; as a rule they differ from them not only in the shape of the body, but also more or less by their manner of life, and only attain the form of the parent through metamorphosis. This is a gradual process (insecta hemimetabola) in the Rhy?ichota and Orthopiera, or a sudden one with a stage of inanition (insecta metabola) in the other orders. This stage of rest or inanition, the PUPA, concludes the larval life (caterpillar, maggot, INSECTA 531 etc.) ; during the pupal stage no nourishment at all is taken, but the internal organs undergo changes ; in some forms the rest is not absolute, as voluntary local movements may take place (pup^ of gnats). The insects are divided into numerous orders according to the form of the mouth parts, the structure of the wings, as well as the manner of the development ; with the exception of the lowest group {Apterygotd)^ which is most nearly related to the ancestors of the insects, and which has no wings and undergoes no metamorphosis, all the remaining orders, which are termed Pterygoia^ have wings on the thorax, though there are, of course, a few species and families of this group which have lost their wings. The Pterygota include — (i) Orthoptera. — Biting mouth parts, anterior wings leathery, posterior wings thin, folded longitudinally ; metamorphosis incomplete (grasshoppers, crickets, cockroaches). (2) Pseudo7ieuroptera. — Biting mouth parts, wings of equal size, thin, not folded up (dragon-flies, hair and feather lice, termites). (3) Rhynchota or Hemiptera. — Mouth parts formed for puncturing and sucking ; wings alike, or the anterior wings may be thickened, parchment-like at their base (plant lice, cicadae, bugs and true lice). (4) Neuroptera. — Biting mouth parts ; wings alike, thin ; metamorphosis complete (ant-lions, lace-wing flies, etc.). (5) Trichoptera. — Licking mouth parts ; anterior wings narrow, posterior wings longitudinally folded, both ornamented with little hairs ; the larvae are worm-like in form, live in water, and breathe through tracheal gills (may flies, etc.). (6) Lepidoptera. — Suctorial mouth parts ; wings covered with scales (butterflies). (7) Coleoptera. — Biting mouth parts ; anterior wings thickened and differ in colour, appearance and function from the thin, folded posterior wings (beetles). (8) Hyinenoptera. — Mouth parts for licking and biting ; the wings alike, membranous (ichneumon flies, ants, wasps, bees, humble bees). (9) Dipiera. — Mouth parts formed for puncturing, sucking or licking ; posterior wings degenerated (gnats, flies, gadflies, fleas). (10) Strepsiptera. — Anterior wmgs shortened ; the female without wings and living parasitically (fan-wings). The parasites of man occur amongst the Rhynchota^ Coleoptera, and amongst the Diptera. [The most usual and recent classification of the Hexapoda is the following : — (i) Apte7'a. — Wingless insects ; scarcely any metamorphosis. (2) Neuroptera.— Y QMX membranous wings, frequently with much network ; the front pair not much, if at all, harder than the under pair ; the latter with but little or no fan-like action in closing ; mandibulate ; meta- morphosis variable, but rarely complete. (3) Orthoptera. — Four wings ; front pair coriaceous or leather-like, usually smaller than the other pair, which are of more delicate texture and contract in repose like a fan ; mandibulate ; metamorphosis complete. (4) Thysanoptera. — Four very narrow fringed wings ; mouth imperfectly suctorial ; metamorphosis slight. (5) Hemiptera. — Four wings ; the front pair either all transparent or with the basal half leathery; mouth suctorial; metamorphosis slight. 532 THE ANIMAL PARASITES OF MAN (6) Diptera. — Two membranous wings only ; mouth suctorial, very varied ;; metamorphosis complete. (7) Lepidoptera. — Four large wings covered with scales ; mouth suctoriaU metamorphosis great. (8) Hymenoptera.—Your membranous wings ; front pair larger than hind, which do not fold up in repose ; mandibulate, sometimes with a tubular proboscis ; metamorphosis complete. (9) Coleoptera. — Four wings, the front pair hard and horny (elytra), meeting in a line over the back and covering the delicate hind pair ; mandibulate ; metamorphosis complete. [There are two other well-known arrangements, namely, Packard's and Brauer's, of recent date, but the one given here, which is based on Linnaeus' grouping by Dr. Sharp, is by far the simplest. — F. V. T.j Order. Rhyncota.^ The lower lip forms a long thin tube that can be turned back (rostrum), and within which lie the setaceous mandibles and maxillae ; the first thoracic segment is not united with the two posterior ones ; the anterior wings are usually leathery as far as the centre. {a) RhYNCOTA APTEKA PARASITICA. Family. Pediculidae (Lice). The lower lip is transformed into a projecting rostrum provided with barbed booklets in which the hollow extensile sucker (maxillae and mandibles) is situated ; no wings ; no metamorphosis ; only simple eyes ; the antennae are five-jointed, the feet possess hook-like terminal structures ; the barrel-shaped eggs (nits) are deposited on the hair of the host. [The lice or PediculidcE are also known as Anoplura and Siphunculata. [They have been split up into a number of families and sub-families and a number of genera, but as far as this work is concerned it is best to retain the single family Pediculidcr. [Only the three species mentioned here are common parasites of man, but now and then horse and cattle and sheep lice, Hcematopinus^ may cause transitory annoyance.— F. V. T.] Genus. Pediculus, Linnaeus. Pediculus capitis, de Geer, 1778. Male I to 1-5 mm. in length, female 1*8 to 2*0 mm. in length. The colour varies from light grey to black according to the colour of the hair of the human race upon which they are parasitic. The abdomen has eight segments, of which the six central ones are ' [Usually known as Hemiptera, There are two sub-orders, Heteroptera and Homoptera. The former have the base of the front wings coriaceous ; the latter have all four wings membranous. The Homoptera are Aphides or plant lice and scale insects {^Coccida), none of which attack man. Recently an interesdng case has been reported to me where certain Aphides had been passed in human urine. One species was Rhopalosiphiim dianihi, the other found in the urine was the hop aphis {Phorodon humuli). I cannot believe, however, that they had been actually passed, in spite of the case being reported by a medical man.— PEDICULUS VESTIMENTI 533 each provided with a pair of stigmata. The thorax is as broad as the abdomen. Eggs o"6 mm. in length; about fifty are deposited by a female head louse. The young can propagate when eighteen days old. The head louse lives especially in the hairy parts of the head of human beings ; more rarely it is found on other hairy parts of the body. It is spread over the entire surface of the globe, and was present in America before the arrival of Europeans. Quite exceptionally it is said that it bores itself deep into the epidermis and can live in ulcers, [The eggs are pear-shaped and are attached to the hairs near the roots by means of a clasping collar. They hatch in about seven days. The young are like the adults and mature in a month. Its general colour varies with that of its host. In West Africans nearly black, in Hindoos dark and smoky, on Chinese and Japanese yellow, on Hottentots orange, on South American Indians dark brown (Murray).— F. V. T.] Fig. 374.— Mouth parts of Pediculus vestimenii. Enlarged. (After Denny.) Fig. 375.— Ovum of the head louse. 70/1. Fig. 376.— Head louse, male. 15/1, Fig. 377. — Pediculus vesti- mentis Burm. : adult female. iS/i. Pediculus vestimenti, Nitzsch, 1818. The head in front is somewhat rounded. Antennae longer than in the head louse ; 2 to 3-5 to 4 mm. in length ; whitish-grey ; the abdomen is broader than the thorax; stigmata as in P. capitis. Eggs 07 to o'9 mm. in length ; about seventy are deposited. P. vestimenti lives on the neck, throat and trunk of persons, and the clothing next the body, in which also the eggs are deposited. The louse of so-called pedicular disease {P. tabescentiuin) is, according to Landois' researches, only the usual P. vestijnenti ; moreover, many cases of phthiriasis are attributable to mites or fly maggots. [This parasite has often been a great pest amongst soldiers during long campaigns, especially amongst the Russians during the Crimean War. Vide also notes in Addenda (p. 615) under " Body, Head and Clothes Lice." — F. V. T.] 534 THE ANIMAL PARASITES OF MAN Genus. Phthirius, Leach. Phthlrius inguinalis, Redi, 1668. Syn. : Pedicidiis pubis^ L. Male o*8 to i*o mm. in length ; female 1*12 mm. in length ; colour greyish-white ; form subquadrate ; the two posterior pairs of legs are strong ; the abdomen has nine segments and six pairs of stigmata ; and still another pair of stigmata is situated between the two anterior limbs. Eggs pear-shaped, o'S to 0*9 mm. in length, 0*4 to o'5 mm. in breadth, and are deposited in rows of about ten on the hairs. Fig. 378. — Phthirius inguinalis. Leach : they are distinguished by the larger tracheal trunks originating from the stigmata. Enlarged. Pediculus pubis ^ which is found almost exclusively in the Caucasian race, lives on hairy parts of the body, but hardly ever on the skin of the head ; the pubic region is its favourite place of abode. [This species reproduces more rapidly than other lice, and is communicated much more freely. The eggs are often laid singly attached to the hairs near their apex. It is known as the " crab louse."— F. V. T.] (6) Rhyncota HEMIPTERA. Family. Acanthiadae. Body flattened, antennae four-jointed, rostrum three-jointed, wings atrophied. [This family, the CimicidcB, includes the bed bugs ; the proboscis, which lies in a groove, is of three segments ; the front wings are shown by two small elytra, there is no trace of hind wings. Two species are known commonly to attack man.— F. V. T.] Genus. Cimex, Linnaeus. Cimex lectularius, Linnaeus. Syn. : Aca?tthia lectidaria^ Fabricius, 1794. It measures 4 to 5 mm. in length, 3 mm. in breadth ; brownish- red ; eight abdominal segments. The female deposits fifty whitish eggs at a time (1*12 mm. in length) three or four times a year ; the CIMEX LECTULARIUS 535 entire development up to complete maturity takes about eleven months. [They will breed all the year round, but less so in cold weather.— F. V. T.] The bed bugs live in the cracks and fissures of human habitations, under carpels, behind pictures, in furniture, bedsteads, etc. ; hidden during the day, they attack persons at night to suck their blood. The alkaline secretion of the salivary glands dropped into the wound around the separate bites causes the so-called " wheals." The bed bugs were known in bygone days by the Greeks (/capjo) and the Romans (cimex). They were first mentioned from Strasburg in the eleventh century, and in England about 1500. [This is the common bed bug of northern latitudes and must not be confused ;vith the tropical bed bug {C. rotu7tdatus). The bed bug can migrate from one house to another; this especially takes place when a house is uninhabited. They escape from windows and pass along walls, water-pipes and gutters, and so reach adjoining houses. This noxious pest accompanies man wherever he goes ; ships and trains become infested, especially the former. [A characteristic feature in this animal is the peculiar odour it produces, like many others in the same group of insects. This odour comes from a clear, oily volatile liquid secreted by glands in various parts of the body. Although he normal food is man's blood, the bed bug can subsist upon moist wood, dust and dirt that collects in crevices in floors, walls, furniture, etc. The puncturing mouth consists of Fig. 379. ■Head of the bed bug from the ventral surface, a, the rostrum ; b^ the antenna ; and r, the eye. ^o\\. a fleshy under lip, within which lie four thread-like hard filaments which pierce the flesh, the blood being drawn up through the beak. [The eggs are oval, white, with a projecting rim around one end, with a lid which is pushed off when the young hatch ; they are laid in cracks and crevices in batches of from twelve to fifty. The t-gg stage lasts from seven to ten days. The larval stage so gradually passes into the adult that one scarcely notices the change ; during its growth the skin is cast five times, and at the change the little wing-pads are seen, showing that the adult stage is reached. The young larva is at first pale yellowish- white. It resembles the parent, but has no trace of elytra. Although eleven weeks is said to be necessary for their development, the stages may be gone through much more rapidly ; Howard and Marlatt ^ give seven weeks in some instances. It seems pretty certain that these Cimex only take one meal of blood between each moult and another preceding egg laying. — F. V. T.] ' " Household Insects,' 1896, p. 37. Howard and Marlatt, Bull. 4 (N.S.), U.S. Dept. Agric, 536 THE ANIMAL PARASITES OF MAN Cimex rotundatus, Signoret, 1852. [This bug is common in warm climates; it is an abundant insect in India, and King has found it in the Sudan, where C. leciularius is, however, the common species. It is usually known as the tropical bed bug. Signoret's bug can be told from the other common species by the shape of the pronotum. In C. rotundatus it is uniformly convex, whilst in C. lectularins the lateral edges are flat and sometimes even concave. The abdomen of rotundatus is also rather more elongate. [This species is of considerable importance, as according to Patton it may act partly as the intermediary host of the piroplasma of kala-azar. [Wenyon found at Bagdad that Cimex sp. would take up Leishmania from Oriental sore, and that the parasite developed into flagellate form. Patton came to the conclusion that the bed bug transmitted Oriental sore in Cambay, India, but Wenyon contests this view {vide Journ. Loud. School Trop. Med., 191 2, ii, pt. i, pp. 13-26). Franchini {Bidl. Soc. Path, exot., 1912, v. No. 10, pp. 817-819) was unable to connect Cimex with this disease. At present nothing seems proved. Besides their possible connection with kala-azar, it has been shown by Howard and Clark {Journ, Exp. Med., 19 12, xvi, No. 6, pp. 850-859) that they can carry the virus of poliomyelitis. [This bed bug was originally described from the Island of Reunion in 1852 by Signoret. A similar insect was described from Burma by Fieber, in t86i, as C. macrocephalus. This is the same as Signoret's species. [The distribution given by Patton^ is as follows : India, Burma, Assam, Malay, Aden, Islands of Mauritius and Reunion. Patton in this paper refers to an erroneous statement made in a recent edition of this book (the last English edition). As I have personally kept ledidarius in moist dirt, wood and refuse for over two years, the statement as far as I am concerned is not erroneous. Moreover, since his doubting this fact the same experiment has been twice repeated with the same results. What they did and do persist on I cannot say.— F. V. T.] Cimex columbarius, Jenyns. [This is common in parts of Europe in pigeon nests, and also amongst poultry {vide Report Econ. Zool. for year ending September 30, 1913, pp. 142-144, Theobald). It occurs in Britain on the latter and will attack man. I have personally been badly bitten whilst collecting them. It is rounder and has shorter antennas than ' Indian Med. Gaz., February, 1907, xlii, No. 2. REDUVIID^ 537 C. lectularlus. Jenyns also described a more pubescent species from swallows as C. hinindinis. 1 have recently received an account of the swallow bug invading a house in Kent and causing much annoyance. — F. V. T.] Cimex ciliatus, Eversmann, 1841. 3*3 mm. in length, yellowish-red, thickly covered with hair ; indigenous in Russia (Kasan). [From a single specimen seen it is evidently distinct. — F. V. T.] Family. Reduviidae. Head long, narrowed behind into a neck ; eyes large, prominent ; rostrum thick and curved ; antennae moderately long, slender at the tip ; legs long and stiff; carnivorous. Amongst the Rediiviidce one genus is of particular importance, namely the genus Conorhinus, which has a long head and the first segment of the beak very much shorter than the second, and the posterior tibiae longer than the femora. These large bugs have a wide distribution, the Oriental region, North and South America, and the West Indies, Madagascar and West and Central Africa. These large bugs may cause very nasty wounds by their bites, but beyond that it has recently been shown that one interposes in the life- cycle of a trypanosome, namely — Genus. Conorhinus, Lap. Conorhinus meglstus, Burm. This large bug has recently been shown by Chagas to be the agent in the development of the trypanosome (7. cruzii) which is the cause of the well-known disease in many parts of Brazil called Barheiro (Barbier). This insect is about i in. long, black, with four red spots on the pronotum, and six red lateral lines on the abdomen, black legs, head and beak. The insect is figured in a coloured plate (No. 9) in Mem. hist. Oswaldo Cruz, IQ09, i, fasc. 2, pp. 158-218. A further account is given by Neiva.^ Conorhinus sanguisuga, Lee. (Blood-sucking Cone-nose). This bug is also known as the Texas or Mexican bed bug, also as the big bed bug. It is particularly troublesome in the Mississippi Valley in bedrooms. The bite is very severe and results in more pronounced swelling and inflammation than that of the Cimex. ^ Mem. Inst. Oswaldo Cruzy 1910, 2, fasc. 2, pp. 206-212. 538 THE ANIMAL PARASITES OF MAN Normally this genus feeds upon the blood of mammals and insects. Its fondness for human blood appears to be quite a new habit, and appears limited to the mature insect only. It is nearly an inch long, flat, head very narrow and long, the rostrum short and thick. In colour it is dark brown with pink markings. They are fully winged when adult, and they fly with ease, entering houses on the wing, especially being attracted by lights in windows ; they also run swiftly. Like the bed bug they conceal themselves during the day and come out at night and bite the sleeper. The effect of the bite is very varied, but as- a rule a sore, itching wound, accompanied by burning pain and swellings, which may extend over a good deal of the body, occur. A specific poison is undoubtedly injected into the puncture ; but no doubt serious results are also due to the beak being contaminated through the insects feeding upon foul carrion. Mr. Lembert, when bitten by a Coiiorhimis sp. (?) on the Pacific slope, exhibited the following symptoms : an itching sensation extending up the leg, large blotches manifesting themselves on the upper part of the limb and extending up to the hands and arms ; his lips swelled and the itching and swelling extended over the head ; there was also much nausea. Similar results are recorded from other regions.^ The eggs of the C. sanguisuga are at first white, then become yellow, then pink ; the young hatch in twenty days. There are two larval and two pupal stages, the latter showing wing-pads. The eggs are laid and the young feed out of doors, chiefly upon insects. It is particularly abundant in April and May indoors. Conorhinus, sp. novum (Monster Bug). Another species; acts in a very similar way in California, the bite being very poisonous. Conorhinus rubrofasciatus, de Geer^ (Malay Bug). This large bug attacks man in Malaysia and elsewhere. It is recorded as inflicting "a very nasty sting, which is done by the huge proboscis." Acute pain and inflammation follow^ in a few minutes. In one case the whole leg became swollen. This species occurs over the whole Oriental region, in Madagascar and Sierra Leone. It is dark brown in colour with dusky yellow or brick-red markings on the pronotum and elytra. Donovan suggests that it may be connected with the kala-azar piroplasma. ' " Household Insects," p. 42. 2 \_First Report Econ. Zool.y 1903, p. 130.— F. V. T.] REDUVIUS 539 Conorhinus renggeri, Herr-Schaff (Great Black Bug of Pampas). This large black bug is mentioned by Darwin/ who states as follows : ^' At night I experienced an attack (for it deserves no. less a name) of the benchuca, a species of Reduvius, the great black bug of the Pampas. It is most disgusting to feel soft wingless insects, about an inch long, crawling over one's body. Before suck- ing they are quite thin, but afterwards they become round and bloated with blood, and in this state are easily crushed. One which I caught at Iquique (for they are found in Chili and Peru) was very empty. When placed on a table, and though surrounded by people, if a finger was presented the bold insect would immediately protrude its sucker, make a charge and, if allowed, draw blood. No pain was caused by the wound. It was curious to watch its body during the action of sucking, as in less than ten minutes it changed from being flat as a wafer to a globular form. This one feast, for which the benchuca was indebted to one of the officers, kept it fat during four whole months, but after the first fortnight it was quite ready to have another suck." Mr. Kirby'^ also refers to this species. Conorhinus variegatus (Variegated Cone-nose). Occurs in Florida in houses, and chases bugs (Cimex) and flies ; not definitely known to bite man. Conorhinus nigrovarius. This species occurs in South America. It is one of the forms known as bichuque. Its bite makes a troublesome swelling. Conorhinus protractus also attacks man in Utah.^ It has been called the '^big bed bug." Genus. Reduvius, etc. Reduvius personatus, Linne. Syn. : Reduviics personatus, Leconte, 1855' European, but also found in the United States. The bite causes mtense pain. It bites when caught or handled, but does not seem to do so voluntarily. Swelling and irritation result which may last a week, and may even cause death.* In 1899 it was very abundant ' Charles Darwin, "A Naturalist's Voyage" (Voyage of the Beagle), 1888, p. 330. '-2 " Text-book of Entomology," 1885, p. 205. ■^ " The Big Bed Bug of the Far West," Bull. 18 (N.S.), U.S. Dept. Agric, 1898, p. loi. ^ " Insects to which the name ' Kissing-bug ' became applied during the summer of 1899/' Bull. 22 (N.S.), U.S. Dept. Agric, 1900, p. 24. 34 540 THE ANIMAL PARASITES OF MAN at Washington and elsewhere; other species occurred, and so no definite opinion existed as to the actual biter, but some people took R. personatus actually biting. It was first described as a parasite of man in America by Walsh and Riley.^ A popular name for this bug is the wheel or masked bug— a black insect, three-fourths of an inch long. The larva of this hug is carnivorous and covers its body with dust so as to conceal itself from its prey. The adult is active on the wing. Coriscus subcoleoptratus, Kirby, 1837. Syn. : Nabicula subcoleoptrata, Kirby, 1837; Nabis subcoleoptratus, Reuter, 1872; Coriscus subcoleopij-aius^ Stal, 1873. Northern United States. Howard was bitten by one between the fingers— the pain was intense, like a needle prick, but the swelling ■was small. No other case known. Rasahus biguttatus, Say, 1831. Syn. : Pirates biguttatus, Stal, 1862 ; Callisphodrus biguttatus, Stal, 1866 ; Rasahus biguttatus, Stal, 1872. Common in southern United States, and found in Cuba, Panama and Para, etc. Known as the two-spotted corsair on account of the great spot on the hemielytra. Frequently found in houses, where it chases the bed bug. It also bites man frequently. From 1869 Walsh and Riley placed it amongst the parasites of man. In the United States Davidson^ is of opinion that all cases attributed to spider bites are due to this insect. Melanolestes morio, Erichson, 1848 (Non-walker). Syn.: Pirates morio, Erichson, 1848 ; Melanolestes morio, Stal, 1866; Pirates picipes, Herrich-Schaffer, 1848 ; Melanolestes picipes, Howard, 1900. Guiana and Mexico and eastern and southern United States. Length 20 mm., hides under stones and logs during daylight, and flies at night. Attracted by lights into houses. Very abundant in 1899 at Washington. Howard cites cases where it was proved to bite man. Melanolestes abdominalis, Herrich-Schaffer, 1848. Syn. : Pi7'ates abdominalis, Herrich-Schafter ; Mela7tolestes abdominalis, Uhler, 1875. Allied to the former ; some say similar, but can be told by the shorter wings on the female. It occurs in the same localities as M. morio. * American Entomolbgist, 1 869, i, pp. 84-88. 2 R. Blanchard, " Sur la Piqiire de quelques Hemipteres," Arch, de Par., 1902, p. 145. •' "So-called Spider-bites and their Treatment," Therap. Gaz., February 19, 1875. ARADID.^., LYG^ID^ 54I Phonergates bicoloripes. This reduvid attacks man in Africa. Family. Aradidae. Broad and very flat bugs, with antennas of four segments and the beak of three ; scutellum short, no cuneus to elytra and the tarsi of two segments. They normally live under the bark of trees, etc., and are found in most parts of the world. Dysodius lunatus, Fabr. (Pito Bug). A large species which is found in South America, frequenting houses, and bites very severely. The Ochindundu. The bug is described by Wellman {Joiini. Trap, Med., April 2, 1906, p. 97) as not only feeding on ticks, such as Ornitliodorits moubata, but as also attacking man. It is called by the Angola Bantus Ihe ochindundu. It is black in colour ; the first two pairs of legs are of a bright red hue. It has curious paddle-like structures on the front four legs, which seem to be designed for securely holding the ticks. It infects native kraals for the sake of preying on ticks. The natives also state that it inflicts a bite which far exceeds in painfulness that of the tick. They compare the bite with that of a poisonous snake. Family. Lygaeidae. Scutellum short ; antennjE four-jointed ; ocelli present ; membranous part of hemielytra with never more than five nervures. Nearly all vegetable feeders . A few are recorded here as bitinsf man. Lyctocorls campestris, Fabricius. Syn. : Acafithia campestris, Fabr. i^Lyctocoris doinesticus). Rare in habitations, lives on human blood. Found by Blanchard in a bed at an hotel at Liverpool. The bite is undoubtedly worse than that of Cimex ; cosmopolitan. In colour it is ferruginous, shining, legs testaceous ; hemielytra slightly shorter and narrower than the abdomen ; membranous portion transparent, the apex broadly fuscous. Length 3*8 to 4*8 mm. Rhodinus prolixus, Stal, 1859. Sometimes attacks man, and the bite is very painful. It is 25 mm. long and 8 mm. broad, and occurs in Colombia. It^ is 542 THE ANIMAL PARASITES OF MAN found also in Cayenne and Venezuela. This like other species is known in South America as bichuque or benchuca. [A few other unimportant species are also recorded as biting man, such as Harpactor cnientas, in the South of France ; Eulyes amcena, from Borneo and Java ; Ariliis carinatiis, Forster, from Brazil. The latter appears to be the same as the Acanthia serrains, Fabricius.— F. V. T.] Order. Orthoptera. [The only Orthoptera recorded as doing actual harm to man are certain wing- less locusts found in Africa. The cysticercus stage of a small tapeworm found in rats and man has been found in an earwig (Alcock). [The strange Hemi77ierid(P found in West Africa, resembling wingless cock- roaches, are parasitic on rats {Criceto/nys). PhasinidcE^ or stick insects, are said to be able to eject a fluid which may cause blindness if it comes in contact with the eyes. Locusts Injurious to Man. [A wingless locust — Enyaliopsis durandi^ Luc— is recorded by Wiggins^ as injurious to man in Uganda. "The bite of this insect," it is said, "gives rise to a very nasty eruption, which may extend over the whole body, with high temperature and general malaise. The skin at the site of the bite sloughs away, and generally leaves a large deep cavity, which heals very slowly." [An allied species — E.petersi^ Schaum — emits a clear yellow fluid, but accordmg to Marshall this does no harm.^ Stannus writes that " for some years I have been cognizant of the fact that among the natives of Nyasaland an allied if not the same species is held to cause skin lesions by the emission of a fluid on the bare skin surface of the body. I have seen cases of ulcers on various parts of the body, for which the ' nantundua ' was assigned as the cause." He then describes the destruc- tion of the superficial layers of the skin which he observed after the yellow fluid had been on the skin twelve hours. — F. V. T.] Order. Coleoptera. The larvae of beetles, similarly to those of some other Arthropoda (myriapods and the larvce of gnats), have sometimes been observed in man as purely accidental guests. In one case or another, such accounts may have originated through a mistake of the observer. Thus English doctors report the presence of the larvae of Blaps niortisaga in the stools of human beings, Sandberg of the larva of Agrypnus murinus in his ten year old son, and Blanchard mentions the larva of a beetle that was vomited by a child. All these cases, however, do not represent actual parasitism, although there are beetles living parasitically.^ Silvanus surinamensis, Linnaeus (Saw-toothed Grain Beetle). [Taschenberg records this beetle as having invaded some sleep- ing apartments adjoining a brewery where stores were kept, and annoying the sleepers at night by nipping them when in their beds. 1 Bull. Ent. Res., 1910, i, pt. 3, p. 227. '^ Ibid., 191 1, ii, pt. 2, p. 180. 3 [Dr. Daniels has sent me a small coleopterous larva found in an abscess on a man in British Guiana.— F. V. T.] DIPTERA 543 [This beetle is common in many parts of the world amongst groceries, corn, meal, seeds, dried fruits, etc. It is about ^^ in. long, much flattened and chocolate-brown in colour. The thorax has two shallow grooves and bears six minute teeth on each side. The jaws are strong, but the bite cannot be very serious. — F. V, T.] Order. Diptera or Siphonaptera. Aphaniptera (Fleas). Wingless, the thoracic rings distinct and free ; antennae of three segments ; legs very powerful ; abdomen with nine segments. [Ten segments are present, but only nine are visible. — F. V. T.] The mandibles transformed into serrated puncturing organs, which are situated in the split sheath of the rostrum ; the maxillae are laminated and have palpi, and more or less conceal the other parts. The importance of fleas lies mainly in the fact that they act as plague carriers. About 150 species have already been described. The only ones of importance for this work are those found on man and those on rats and mice. The two families in which these are found are known as Pulicidce and Sarcopsyllidce. The eggs of fleas are laid on the ground, on rugs, etc., and in birds* and rodents' nests. They hatch rapidly in warm weather and in warm climates, varying from two to five days ; in cold countries they may take two or three weeks to incubate. The larva is a footless creature, pearly white in colour, the head sometimes being darkened, composed of fourteen segments including the head, and although apodal can move with considerable agility. It lives amongst dust and dirt, and feeds upon any organic matter it can find. In about two weeks it is said to become mature, and then spins a cocoon in which pupation takes place. The cocoons of the common human flea and the fowl flea become covered with dust and dirt. The period of pupal life seems varied, for I have had the fowl flea hatch out in ten days, and others in three weeks at the same time of year. The adults are a blood suckers and cause considerable irritation as well as acting as disease carriers, and in the Sarcopsyllidce the females attach themselve:> permanently to their hosts, embedding themselves under the skin, where they become pregnant. Some kinds harbour the cystic stage of tapeworms, and the rat trypanosome passes certam stages in the rat flea. Most fleas have definite hosts, but some, like the rat and fowl fleas, attack man. The fleas which can carry the bacillus of plague are Xenopsylla cheopis, Pulex irritans, Ceratophyllus fasciatus and Hoplopsyllus anomalus. The two families, PulicidcB and Sarcopsyllidcr, can be distinguished as follows : Thoracic segments much foreshortened, coxae and femora of hind legs very slightly enlarged Sarcopsyllidce. Thoracic segments normal, coxae and femora of all the legs much enlarged ... ... ... Pulicidce. Family. Sarcopsyllidae (Jiggers). The members of this family are not confined to one host. Three genera are known and tabulate as follows : — a. Hind coxa without a patch of spines on the inside. aS. Hind femur simple \. Dermatophilus. or. Hind femur with a large tooth-like projection near the base 2. Hectopsylla. j3. Hind coxa with a patch of short spines on the inside 3. Echidnophaga. 544 THE ANIMAL PARASITES OF MAN Genus. Dermatophilus, Guerin. Dermatophilus caecata, Enderl. The eyes of the female vestigial. Taken on and behind the ears of Mus rattus in Brazil. Dermatophilus penetrans, L., 1758 (Jigger, Chigoe). S y n . : Sarcopsylla petieirans. About I to i"2 mm. in length; brown in colour. Eyes distinct. The males only occasionally visit man to bite ; the fertilized female, on the other hand, bores into the skin with her head, particularly about the toes of the host, and then attains considerable dimensions. The eggs develop on the I soil with a metamorphosis ! similar to common flea. Fig. 380. — Dermatophilus fetietraus : young female. Highly magnified. (After Moniez.) FiCt. 381. — Dermatophilus penetrans: older female. Enlarged. (After Moniez.) The sand flea (nigua) particularly infests Central and South America, and, in 1873, was carried by ships from Brazil to the West Coast of Africa. In a com- paratively short time it has become disseminated throughout Africa and has also appeared in Madagascar ; recently also it has been reported from China. Besides attacking man, it also settles on mammals, for instance, on dogs, pigs, etc. According to Jullien the wound or little swelling caused by the female has no particular significance, as children infested with ten or eleven sand fleas quietly proceeded with their ganjes. It will be understood, however, that the wound easily affords the opportunity for the setting up of inflammation or even septic processes, as is the case in any kind of wound. [The jigger is also well known in the West Indies.— F. V. T.] Genus. Echidnophaga, Ollifif. Four species found on rats, etc. Echidnophaga gallinacea, Westwood (Chigoe of Fowls). [This flea is a native of tropical Asia and Africa. It lives on the fowl chiefly, attacking the neck and around the eyes. Specimens PULICID^ ^545 Pulex, Xeiiopsylla. Hoplopsylhcs. Ctenocephalus. were sent me from Texas, where they not only attack poultry but also children, the latter somewhat severely. It also occurs on cats, iand is found on rats in Africa. It has been introduced into North America. [Three other species are found on rats, viz. : E. jiiyrmecobii, Rothsch., from Australia ; E. imirina, T'lrdh, irom. Southern Europe ; and E. liopns, Rothsch,, India and Western Australia. — F. V. T.] Family. Pulicldae (True Fleas). Rothschild's classification is as follows : — Section I. — Club of antennas distinctly segmented only on the hind side. A>K /-. Hind edge of tibias with about twelve short and three long bristles, the short ones forming a kind of comb b^. Abdomen with at least one comb Ceratophyllus. Pygiopsylla. Chiastopsylla. Neopsylla. Ctefiophthalmus . Ctenopsylla. Hystricopsylla. Genus. Pulex, Linn. Pulex irritans, L., 1758. Male 2 to 2*5 mm. in length, females about 4 mm. ; reddish or dark brown ; head without bristles; thoracic and abdominal rings of bristles on the dorsal aspect, and small hairs directed backwards at he posterior margin. The barrel-shaped white eggs are deposited 54<> THE ANIMAL PARASITES OF MAN in cracks in the boards, sweepings, spittoons, etc. ; they produce legless larvae consisting of fourteen segments, which, after about eleven days, are transformed into pupae; after another eleven days the flea emerges. ,^--v"''' ! ' m^ C;; ^■" \. "I ■ I Fig. 382. — Piilex irritans. 14/1. Fig. 38^. — Larva of flea. Enlarged.' (After Railliet.) Fig. 384. — Pulex serraticeps. 22/1. BonL Fleas live in human dwellings all over the world, and periodically pass on to persons to suck their blood. They may deposit their eggs on very uncleanly individuals, and even undergo development, therefore it is possible to find larvae and pupae on such persons. The dog flea, Pulex serraticeps, is easily distinguished from the flea of man by the large thick bristles on the posterior margin of the first thoracic ring (fig- 384). Genus. Xenopsylla, Glink. Xenopsylla cheopis, Rothschild. This is the common rat flea of tropical countries. Rothschild ^ says : "Although practically cosmopolitan, it cannot apparently flourish in temperate and cold cHmates." ' Bzill. Ent. Res,, 191 1, i, pt. 2, p. 92. CTENOCEPHALUS, HOPLOPSYLLUS, CERATOPHYLLUS 547 In the male the bristles of the flap-like process of the clasper all slender ; in the female the narrow portion of the receptaculum seminis long. Originally discovered in Egypt. This is apparently the chief plague flea. The Indian Plague Committee have proved that this flea is easily infected when fed on plague rats, and that the bacillus multiplies rapidly in the flea's stomach and that the fleas may remain infective for fifteen days. How the flea infects man does not apparently seem to have been proved, as it does not do so through its bite, but the excrement is highly infec- tive. It is probable that this poisoned faecal matter gets to the wound caused by the piercing mouth. Xenopsylla brasillensis, Baker, occurs on rats in West Africa and has been introduced into Brazil. Genus. Ctenocephalus, Kolen. Includes the cat and dog fleas. The dog flea, C. canis, Duges, is found on the dog all over the world, but especially in temperate climates. It also occurs on rats. Man is often badly bitten by this insect and it overruns houses. The eggs are laid on rugs, carpets and dust and dirt and amongst dogs' hair, but are not fastened to it and fall anywhere. The ova may hatch in about fifty hours and the larvae live for seven days and then spin their cocoons amongst dust and dirt. The pupal stage lasts about eight days. The cat flea (C. fells) is widely distributed over the world, and occurs on many mammals beyond the cat, and is also found on rats. Genus. Hoplopsyllus, Baker. A genus found in North America related to Pulex, but at once recognized by the prothorax bearing a comb. Hoplopsyllus anomalus, Baker, which is found on the ground squirrel (Citellus heecheyi) in California, an d according to Rothschild once found on the rat, has been proved to carry the plague bacillus and to play an important part in plague infection in California. Genus. Ceratophyllus, Centis. Ceratophyllus fasciatus, Bosc. This flea is also found on the rat in Europe and will attack man. It is a plague carrier. It has eighteen to twenty teeth on the ' •' Report United States Public Health, 1909," xxiv, No. 29. 548 THE ANIMAL PARASITES OF MAN prothoracic comb and no black spines on the head. The genus Ceratophylkis of Centis has a pronotal comb and three ante-pygidial ch^ta3 on each side. Two other specimens recorded : C. londiniensis, Kothsch., and C. anisiis, Rothsch. The former on rats and mice in London, the hitter on Felis sp. from Japan and Mus iiorvegicus in Cahfornia. Genus. Ctenopsylla, Kolen. This genus contains a very abundant rat and mouse species, C. rniisciili, Duges, which is widely distributed over the globe. Genus. Hystrichopsylla, Tasch. Large hairy fie^s, with no eyes and one or more combs on the abdomen. In the Mediterranean area one species, H. tripediiiata, Tirah, is common on rats and mice, and also in the Azores. Several others occur on rats and mice. For information concerning these the reader is referred to Rothschild's papers. Pulex pallipes is another species found on the rat and man. Systematic, Anatomical, and Biological Remarks on Mosquitoes. Mosquitoes come in the Nejuatocera^ one of the sub-orders of the Diptera^ and are divided into numerous families, of which, however, the Culicidct are of most interest to us here. Other families as the Psychodidcc and Chironomidce are also of considerable importance, vide following pages. The head is small, the facetted eyes are placed laterally, but there are no accessory eyes (ocelli). In front of the eyes are situated the comparatively long antennas, the differences of which strongly mark the distinction of sex.^ The antennas are composed of fifteen or sixteen segments. In the male they are covered with long whorl-like hairs, while in the female the antennal hairs are short— differences that are perceptible even with the naked eye.- The proboscis, which is longer than the antennit, protrudes from the inferior aspect of the head and is composed of the following parts (figs. 387 and 388) : Two grooved half tubes, facing one another, of which the upper one is the upper lip (labrum) and the lower one the lower lip (labium), which represents a pair of coalesced maxillas. Within the tube formed by the labrum and labium are the mandibles and maxillae, trans- formed into instruments for piercing, and a single puncturing organ, the hypo- pharynx. On the right and left, next to the proboscis, are placed the straight five- jointed palpi, the final joint of which is thickened in the male.' In biting, the labrum. which is swollen at its free end, is not introduced into the wound like the * [This is by no means always the case ; in the genera Deinocerites, Wyeomyia, Limatus, Theobald, and in Sabethes, Robineau Desvoidy, they are nearly the same in both sexes. — F. V. T.l '^ [This is not always the case, vide previous note. — F. V. T.l •'' [This is only so in Anophelina and in the genus Theobaldinella, Neveu-Lemaire, Grabhamia, Theobald, Acartomyia, Theobald, elc. In true Culex and many other genera the male palpi are pointed. — F. V. T.] REMARKS ON MOSQUITOES 549 other mouth parts, but is bent backwards. The labium and hypopharynx push direct into the skin ; the maxillae and mandibles, however, which are needle-like and serrated; at the tips, penetrate with a saw-like movement. [The swollen free end of the labrum really means the labellce^ two articulated pieces, supposed by some to be the labial palpi. In most species the mandibles are not serrated at their ends. — F. V. T.] The saliva is introduced into the wound through the lumen of the («) (^) Fig. 385. — Head of a male {a) and of a female [b) Anopheles. Slightly enlarged. (After Giles.) {a) (^) Fig. 386.— Head of a male {a) and of a female {b) Culex. (After Giles.) hypopharynx, while the blood is sucked up by the mosquito in the groove of the labium. The three thoracic segments are soldered together. The central one carries the membranous wings on the sides of the dorsal surface ; the posterior somite carries the small halteres (rudimentary posterior wings). There are three pairs of long slender legs on the lower side. 550 THE ANIMAL PARASITES OF MAN The abdomen has no limbs, is composed of eight (rarely nine) distinct segments ; the sexual and anal orifices are at the posterior end, the stigmata on the sides. The intestinal canal (fig. 389) is composed of three principal divisions ; the anterior part reaches as far as the front pair of legs, and consists of the oesophagus, which md. ■■ y 7 H -— , -^^ipf J_^ 1 tr PI \ i\ li J - 4\ \\ i Iwli J Wi ^i w^y^ \VWl// L ,-- u.i. ind. Fig. 387. — Mouth parts of Anopheles claviger.^ h., hypo- pharynx; md., mandible; mx., maxilla; u.L, upper lip; /./., lower lip ; f.., palpi. (After Grassi.) 7nd. Fig. 388. — Anopheles maailipennis : transverse section through the proboscis of a female {a) and a male {b). hy., hypopharynx, with duct of the salivary gland; ?«., muscles; md., mandibles; 7?tx., maxillae; /., labium ; /./., labrum. (After Nuttall and Shipley.) is provided with two small lateral diverticula. [At the commencement of the (eso phagus are one or mo re diverticula, which vary in size ; they contain air, food ' [This should read Anopheles maailipennis, Meig. ; there was no type of A. claviger. — F. V. T.l REMARKS ON MOSQUITOES 551 and bacteria.— F. V. T.] The mid gut reaches as far as the fifth and sixth abdominal ring ; in front it is thin, and has numerous small supra-oesophageal ganglia ; the posterior part is, however, more dilated. Four or five Malpighian tubes, the excretory organs, discharge at the place where the mid gut passes into the terminal gut. The pair of salivary glands have one common excretory duct leading into the hypopharynx. These glandular bodies are situated in the thorax ; each consists of three slightly serpentine tubules, the dorsal and ventral tubes being long, the central one shorter. The above-named characteristics apply to both genera Culex and Anopheles, but in the genus Culex is smaller, Anopheles larger. [In Anopheles the ends of the ducts in the lobules are dilated, whilst in most of the genera the ducts are the same size all along. The lobules may bifurcate, and in Psorophora there are five lobules. — F. V. T.] The legs of the genus Culex are about the same length as the whole body ; in Anopheles they are double that length.^ In Anopheles the palpi and proboscis are of equal length; in Culex the condition is different, according to sex. In the male the palpi are longer than the proboscis ; in the female considerably shorter and the number of segments diminished. The venation of the wings exhibits furthur points of differentiation, as also their adornment, though this last sign is not by any means always conclusive ; most species of the genus Culex have unspotted wings, whilst those of Anopheles are usually spotted. More important is the fact that in Culex the Fig. 389. — Longitudinal section of an Anopheles, showing alimentary canal. In the forepart of the thorax is the salivary gland consisting of three tubules ; ventrally, the suctorial stomach extending into the abdominal cavity ; the stomach, and at the posterior end of the abdomen the Malpighian vessels. (After Grassi.) abdomen is decorated with small scales, similar to those on butterflies, whereas there are small bristles on the abdomen of Anopheles. [This cannot be said to be a character by which an Anopheline may be told from a Culicine, for in such common Anopheline genera as Cellia and Neocellia we get plenty of scales on the abdomen. — F. V. T.] An experienced observer can, however, separate the two genera by the difference in size and their manner of resting. When settled they either touch the resting place with all the legs or only with the four anterior legs. In consequence of the different length of the legs, the body of Culexj approaches the resting place more closely ; moreover, Culex holds the abdomen parallel or at an acute angle to the resting surface, whereas Anopheles carries the abdomen directed upwards (at an angle of about 145°) and holds the head down. Both genera, however, usually only rest on the four anterior legs, and then, as has long been known, Culex carries the third pair directed towards the dorsum, while those of Anopheles hang down. In regard to the differentiation of the species, I must refer you to the special literature, and content myself by observing that about 150 species of Culex and about fifty species of Anopheles have been described, of which fifty about four are ^ [This is certainly not always the case. — F. V. T.] 552 THE ANIMAL PARASITES OF MAN found in Europe. [The number of known Anophelines now is more— loo species— of other CuhcidiE over 700.— F. V. T.] According to our present knowledge it appears that the entire genus Anopheles can transmit malaria to man ; this observa- tion has been confirmed in Anopheles clavi'oer, Fabr. ; A. inacidipennis, Meig. ; A. bifurcaius, L. ; A. superpictus, Grassi ; A. pseudopictiis, Gr., all of which are found in Italy,^ Germany, etc., as well as in the tropics. Moreover, in A. costalis^ Loew ; A.funestus, Giles (Africa) ; A. quadrimaciilaius. Say (North America), and A. rossii^ Giles ; the latter is perhaps identical with A. superpictus, Gr., as well as with A. culidfacies (India). {Anopheles maciilipennis and A. claviger are the same. Certainly neither viaculipenftis nor bifurcaius has been found in the tropics. Anopheles quadrimacidatus, Say, is the same as A. macnlipe7i7tis. There is no Fkx. 390. -Anopheles maculipennis, Meigen. Enlarged. (After Grassi.) evidence that all Anophelines carry malaria, but there is much to show that certain species only are capable of so doing. A list of known carriers is given later. — F. V. T.] Everyone is aware that mosquitoes swarm at sunset in fine weather, and then seek out human beings and other warm-blooded animals to take food. In this regard, however, the sexes differ, for it is almost without exception that the females only suck blood, while the males subsist on the juices of plants (blossoms or fruits).^ ' Compare Ficalbi, E., " Venti spec, di zanzare {Culicidce) ital. . .," Btdl. Sor, ent. iial., 1899, xxxi ; abstracted in Centralbl. f. Bakt., Par. u. Infektionsk., 1900, xxviii, p. 397. ■^ Both males and females may be kept alive in captivity for a long time if given fruits, or even only sugar and water. . REMARKS OX MOSQUITOES 553 After sucking, and when night has fallen, the mosquitoes find a place of refuge, for which purpose they utilize the grasses or foliage of trees and bushes, or inhabited or uninhabited rooms of houses, also cellars, stables, verandahs, etc., where they also pass the day. [Some mosquitoes bite in the daytime— Stegoymia and some Anophelines ; some bite right into the night, as Cule.x fatzgans and C. pipiens.—Y . V. T.] The period required for digestion varies according to the temperature. It takes two days in summer, and may take up to ten days or more in cool weather. After digestion is complete more food is taken up, this being necessary [in some species only— F. V. T.] for the maturing of the sexual products in the female. It is still unknown under what circumstances copulation takes place ;^ in any t^ Fig. 391. — Larva of Anopheles niaculi- pennis, Fabr. Enlarged. (After Grassi.) Fig. 392. ■LTrvaof Culex. Enlarged. (After Grassi.) case, sooner or later the females are fecundated, and when the ova have become mature, and the season is not too far advanced, they seek a suitable place in which to deposit them.^ These are larger or smaller, permanent or temporary, collections of standing water, pools, puddles, lakes, pits, water in rain-water barrels, basins, etc. Nevertheless, certain kinds prefer certain waters ; thus Anopheles {claviger) maculi- pennis and several of the Culices seek stagnant water overgrown with swamp vegeta- tion and decomposing vegetable matter; A. bifurcatus and certain Culices, clear water with some vegetation (such as fountains and the lakes in gardens and parks) ; Ciilex pipiejis has a preference for rain-water barrels, even though the water be dirty and evil-smelling. [I have found the larva? of Anopheles bifurcakis living in great numbers in ponds and lakes completely overgrown with floating water-weeds, and those of Culex pipieiis in liquid manure. ' The act of oopulaiion in many species is now known. The female Culex has three receptaculae seminalis, while ihe female Anopheles has one receptaculum seminis. - It is certain that the females perish immediately after deposidng the ova ; but this does not always hold good, as a part of them survive for a few days. The males die soon after copulation. cc^ THE ANIMAL PARASITES OF MAN Sexua/ Organs of the Mosquito.— 1\\^ female has a pair of ovaries, opening into a single tube by the ovarian tubes ; into the single tube opens a duct coming from the spermatheca;, and also a mucous gland. The spermathecae store up the male cells. The male organs consist of two testes joined by ducts (vasa deferentia) to the ejaculatory duct formed by their union. Each vas deferens is joined by a bhoit tube with the sac-like vesicula seminalis.— F. V. T.] There is also a difference in the manner in which Culex and Anopheles deposit their ova. Culex deposits two to three hundred eggs in compact heaps that float on the water, and in which the eggs stand perpendicularly one next the other ; whereas Anopheles jjiaculipenms deposits only three or four up to twenty eggs, united in groups that float horizontally on the water ; the eggs of A. bifurcatus, again, are arranged in star-like groups. The eggs are about 075 mm. in length, and assume a dark hue soon after being laid. The development only occupies a few days. The young larvae grow rapidly, changing their integument several times ; Fig. 393. — Pupa oi Anopheles inacuiipennis, Meig. Enlarged. (After Grassi.) the larvae also differ in the various genera, though they have a general resemblance (figs. 391 and 392). The long legless larva has a flattened head, a fairly broad, rectangular, or trapeziform thorax, on which there are bristles, and an abdomen distinctly seg- mented, and on the segments of which there are also lateral bristles. The situation of the stigmata marks the difference between the two genera. Though in both genera the stigmata are at the posterior end and on the dorsal surface, they are in Anopheles close to the surface of the body ; in Culex, however, they are on the free end of a long tube (siphon). The position of the larva in the water also differs. The larva of Anopheles lies almost horizontally beneath the surface of the water, the posterior border of the penultimate abdominal segment, upon which the stigmata are situated, being on the surface ; whereas the larva of Culex hangs head downwards perpendicularly in the water, the point of the siphon only touching the surface. In about a fortnight the larva is fully grown and becomes a pupa. The pupa CULICID^, OR MOSQUITOES 555 (fig"' 393)5 which moves in jerky movements, remains in the water, but partakes of no food. In shape it somewhat resembles a tadpole, that is to say, it consists of a bulky anterior portion, on the surface of which the head, with its appendages, is recognizable, and a more slender segmented abdomen. Above, on the thorax, there are two small trumpet-shaped breathing tubes for the conveyance of air to the tracheal system. After three or four days the perfect mosquito hatches out, remains a short time on the surface of the water until its chitinous integument is hardened, and then flies away. The females that are fertilized in the autumn hibernate in sheltered spots in the open air, or in houses, cellars, under stairs, in stables, barns, etc., and are the progenitors of the first generation of the following year. In accordance with the climate of a country, or the kind of weather of a year, the conditions in regard to the manner of life and the duration of the development of the mosquito vary. At all events, the life-history of the mosquito elucidates many points relating to malaria which were hitherto not understood. [The length of the egg, larval and pupal life varies so much that it is not possible to give an account of any value here. Frequently the eggs may incubate in two days, whilst I have had StegoiJiyia fasciata eggs from Cuba that have hatched out under abnormal circumstances more than two months after they were laid (" Mono. Culicid.," iii, p. 6). Some larvae, as Aftopheles bifurcatus, live for months during the winter. Some mosquitoes therefore hibernate as larvae. The larvae and pup^e of the different genera present very marked characters, mainly in regard to the structure of the siphons. Specific differences may be found in the frontal hairs of Anopheline larvae and in the number and arrangement of a group of spines at the base of the siphon in Culicines. — F. V. T.] Gulicidse or Mosquitoes. The importance of these insects to man is very great. They not only produce painful bites, which may become inflamed and give rise to a considerable amount of cedema, but they are more important on account of the part they play in the distribution of various diseases. CidicidcE may not only carry disease germs, but act as intermediate hosts for certain parasites, such as some of the Anophelina for malarial parasites, Culex for Filariae, and Stegomyia for yellow fever, etc. ; the last- named is in any case the distributor of that fatal disease. It is therefore very necessary to know the life-history, habits and characters of these pests. Mosquitoes exist in almost all parts of the world from the Arctic circle to the tropics ; temperate regions suffer from them less than the two extremes, but even there they form not only a source of great annoyance but of danger as malaria and possibly now and again yellow fever carriers. A few years ago comparatively few species were known, now some 800 odd have been described. Their number will probably not stop far short of 1,000, in spite of the fact that many have been described under different names, yet really the same species. Some are purely domestic, others entirely sylvan ; the former, as we might expect, often have a very wide distribution, having been taken from place to place in boats and trains. The more rapid transport becomes, the greater becomes the possibility of this wide distribution of many species increasing, and the spread of other species from their natural home to foreign parts by sea and then by trains further inland. All CulicidcE are aquatic in their larval and pupal stages. Almost all small collections of water, both natural and artificial, may form breeding grounds for these pests. Some even breed in pitcher plants and many in bromelias. The 35 556 'HE ANIMAL PARASITES OF MAN YiG. 3^^— Heads of Culex and Anopheles : (i) Culex trale ; (2) Culex fema]( (3) Anopheles male ; (4) Anopheles female. (After Daniels.) CULICID^ OR MOSQUITOES 557 favourite resorts for the larvae of Anophelina are small natural collections of water, such as puddles, ditches and small pools around swamps; certain species (A. viaculi- pennzs, etc.) live in rain barrels as well. They may also occur in the sluggish water at the edges of rivers or even in mid river, where the flow is checked by masses of water weeds {Myzomyia funesta, etc.). The Stegomyias prefer artificial collections of water, but also occur in natural pools. The yellow fever species {S. fasciaid) prefers small collections, such as in barrels, pots, jars, etc. Culex occur in all manner of places — rain barrels, tanks, cisterns, ponds and ditches. Some of the South American species of Culex, Wyeomyia, Joblotia, etc., breed in the collections of water at the base of bromelia leaves.^ Very few Culicid larvae live in salt water except in Australia, where Dr. Bancroft has found them in salt water of specific gravity 1*040 {Mucidus alternants and Culex ati7iulirostris). Other salt water mosquitoes are known in America. The food of the larvae is very varied ; the majority appear to feed upon confervae, small Crustacea and insects; some are Fig. 395. — a, eggs of Culex ; <5' b^, eggs of Anopheles ; r, eg^ ^, ^goi Tseniorhynchus ; e, egg of Psorophora , egg of Stegomyia ; cannibals, readily devouring others of their own kind. The larger larvae of Mega- rhinus, Psorophora, Toxorhynchites and Mucidus are extremely ravenous and devour one another. There are two main types of larvae, the Anopheline and Culicine ; in the former there is no respiratory siphon, in the latter the siphon is long or moderately long. The head offers certain marked peculiarities which are of specific value ; this especially applies to the Anophelina^ in which the frontal hairs are of great service in distinguishing the larvae,^ whdst in Culex ihe number and position of the spines at the base of and on the siphon are characteristic. The position assumed by the larvae in the water also varies in the different groups ; most of the Anophelines lie horizontally, most of the Culicina and ALdeomyina hang head downwards. The 1 " Wald Mosquitoes and Wald Malaria," Dr. Lutz, Centralbl. f. Bakt., Par., 21. Infektionsk., i Abt. Grig., xxxiii, No. 4. 2 Informaii n sent me by Dr. Grabharn shows this statement to be not quite correct, as the frontal hairs may vary in different stages of the same larva. This he has shown in Cellia albipes, Theob., and I have noticed it in a Nyssorhynchus from Africa. 558 THE ANIMAL PARASITES OF MAN pupee also vary, but not to the same extent; the chief differences to be noticed are in the form of the two respiratory trumpets. The eggs, which may be laid separately {Ajtopheles macuhpennis^ Stegoviyia Proboscis Antennae 1 Wing scaies"! Proboscis Palp Antennae. Basal lobes ofantennqe Yrorts Vertex Eyes Oaipul— Nope Fig. 396. — Diagram showing the structure of a typical mosquito. (Theobald.) 5^'?t:arsci fasciaia, Joblotia nivipes, etc.), or in rafts {Ctdex pipiens^ C.fatigans) or in chains {Pseudotceniorhynchus fasciolatus), present a great variety of forms. The most peculiar are shown in fig. 395 (Ttisniorhynchus, Culex, Stegomyia, Anopheles, Psorophora). CULICID^ OR MOSQUITOES 559 As in all insects, they differ very materially in each species of one genus. Those best known are the Anopheline eggs. The eggs always float on the surface of the water; immersion soon destroys them, but many may occur in mud and can resist desiccation. Fig. 397. — Types of scales, a to k ; head and scutellar ornamentation, I to 5 ; forms of clypeus, 6. (Theobald.) i, head and scutellum of Stegom)ia, etc.; 2, of Culex and Mansonia ; 3, of Howardina, yEdes, etc. ; 4, of Megarhinus and Toxorhynchites, etc. ; 5, of Cellia and some other Anophelines ; 6, a', clypeus of Culex; d', of Stegomyia ; <:', of Joblotia. C/tarac/ers of Adul^ Cuh'cidcF.— The ch\e( ch2iY2iC\.eYS by which true mosquitoes, or Ciilicidce^ are known are the following : — (i) Wings always with the veins covered with scales; the longitudinal veins, 56o THE ANIMAL PARASITES OF MAN usually six in number (in one genus seven) ; the costal vein carried round the border of the wing. (2) Head, thorax and abdomen usually, but not always (Anopheles, etc.), covered with scales. (3) Mouth parts formed into a long piercing proboscis. As a rule the males may be told from the females "by their antennae being plumose, whilst in the females they are pilose {vide fig. 394), but this does not invariably hold good, for in Deinocerites, Theobald, and Sabethes, Desvoidy, and others, they are pilose in both sexes. The labial palpi are very variable in regard to their form and the number of segments ; in the Aiiophelina they are long in both sexes, as long or nearly so as the proboscis, more or less clubbed in the males ; in Culicma^ Joblotiiia and Heptaphlebomyia^ they are long in the males, short in the females ; in A£deo?nyina, short in both sexes. Scales. — The most important structural peculiarities in Culicidce are the scales, which form the chief and most readily observed characters for separating genera and species. The importance of scale structure has been recently ignored by some Fig. '^98. — Neuration of Wing. Explanation of JVing, Veins and Cells. — A, costal cell ; B, sub-costal cell ; C, marginal cell; D, first sub-marginal cell (= first fork cell) ; E, second sub-marginal cell ; F, first posterior cell ; G, second posterior cell (= second fork cell) ; H, first basal cell ; I, second basal cell; J, third posterior cell; K, anal cell; L, anxiliary cell; M, spurious cell ; c, costal vein ; 1st — 6tk, first to sixth longitudinal veins ; a, a' and a', incrassa- tions (a' called by Austen the sixth vein, a" the eighth vein); y, supernumerary cross vein; z, mid cross vein ; /, posterior cross vein ; s.c.^ sub-costal. (Theobald.) workers, who are probably right academically, but as a means of separating groups^ and so more easily running down a species, the practical man is strongly advised to follow this method. As to what a genus is, is purely a matter of personal opinion. If one examines any recent standard work on entomology one will find a species being placed in varied genera by the varied authorities. The head, thorax, abdomen and wings are in nearly all cases clothed with squamce of varied form, of which the following are the main types (fig. 397) : — (i) Flat, spade-shaped scales (a). (2) Narrow curved scales (e). (3) Hair-like curved scales (d). (4) Spindle-shaped scales (/). (5) Small spindle-shaped scales (£■). (6) Upright forked scales {/i) and (i). (7) Twisted upright scales (J). (8) Inflated or pyriform scales (/'). THE CLASSIFICATION OF CULICID^ 561 (9) Mansonia scales (d). (10) Small broad asymmetrical scales (c). Various other varieties are found on the wings, such as : — (i) Narrow linear lateral scales. (2) Narrow lanceolate scales. (3) Broad lanceolate scales. (4) Elongated, broad, truncated scales (= Pseudotaeniorhynchus-like scales). (5) Pyriform scales. (6) Asymmetrical broad or Taeniorhynchus scales. (7) Flat spade-like scales.^ The wings have a series of scales along the middle line of the veins, and also lateral scales to all or nearly all the veins. The wing is also fringed by a series of scales (fig. 396), which, however, are of little systematic importance ; the so-called " border scales " (b.s.) vary, however, to some extent, and are useful characters in separating some of the Taeniorhynchus. The Classification of Culicidcv. Section A. — Proboscis formed for piercing ; metanotum nude. Scutellum simple. I. Wings with six-scaled longitudinal veins. A. Palpi long in the male. a. Palpi long in both sexes, clavate in S Aftophelina, \. First submarginal cell as long or longer than the second posterior cell. Anlennal segments without dense lateral scale tufts. /Wing scales lanceolate ... Wing scales mostly long and narrow ... Wing scales as above, but f o u r t h long vein near base of third and outstanding scales on pro- thoracic lobe Wing scales partly large and inflated Thorax and abdomen with hair- like scales Prothoracic lobes sim- ple; no' flat head scales Anopheles^ Meigen. Myzomyia^ Blanchard. Neomyzomyia, Theobald. I ^ \ I Prothoracic lobes mammil- ' lated ; some flat head scales. . Basal lobe of $ genitalia of \ two segments Prothoracic lobes with dense outstanding scales ... Thorax with some narrow curved scales ; abdomen hairy Wing scales small and lanceolate. Wing scales broad and lanceolate Cycloleppteron^ Theobald. Stethoinyia^ Theobald. Feltinella, Theobald. PyretopJwrus^ Blanchard. Myzorhynchella^ Theobald. Heart-shaped scales occur on the wings of Etiorlepiiomyia. 562 THE ANIMAL PARASITES OF MAX II, Thorax with hair-Hke curved scales, some narrow curved ones in front ; abdomen with apical lateral scale tufts, scaly venter ; no ventral tuft Thorax with hair-like curved scales ; abdo- minal scales on venter only, with a distinct ventral apical tuft Much as above, but abdomen with long spine-like dense lateral tufts Thorax with very long hair-like curved scales ; abdomen pilose, except last two segments which are scaly ; dense scale tufts on third femora ; wings with broadish, blunt, lanceolate scales /Abdominal scales as lateral dorsal patches of small flat scales ; thoracic scales nar- Thorax and row and curved, or spindle- abdomen { shaped with scales Abdomen nearly completely covered with irregular scales and with lateral tufts ^ No lateral scale tufts ... Thoracic scales ha[r-like except a few narrow curved ones in front ; abdominal scales long," broad and irregular Thorax with hair-like curved scales and some broad straight scales, others spatu- late on sides. Abdomen covered with fine hairs except last three segments, which are scaly. Tufts of scales on hind femora. Wing scales lanceolate Antennal segments with many dense scaly tufts Antennas with outstanding scales on second segnient, more appressed ones on the first. At least one segment of abdomen with long flat more or less spatulate scales ... First submarginal cell very small With a distinct cylindrical tubercle project- ing obliquely from the prothoracic region Scutellum trilobed. A rribalzagia^ Theobald. My2'.orhynchus^ Blanchard. Chrystia, Theobald. Lophosceloniyia, Theobald. Nysso?'hynchus, Blanchard. Cellia^ Theobald. Neocellia, Theobald. Kerieszia. Theobald. Mano^uinhosia^ Cruz. Chagasia, Cruz. Calver/hia, Ludlow. Bironella, Theobald. Dactyloinyia, Newstead and Carter.^ ' T"^e following genera of Anophelites have been founded by James* : — (i) Abdomen with hairs but no scales. Thorax with dorsum with long narrow curved scales, which form on the anterior promontory a thick bunch projecting over the neck. Protho- racic lobes with a tuft of rather broad true scales, upright forked scales of head of usual broad expanding type: Patagiamyia, James. Includes Gigas, Giles, and Lindesayi, Giles. Both seem to me typical Anoplieles. (2) Abdomen as above. Thorax very similar. Prothoracic lobes with hairs, no scales. Upright * Records of Indian Museum^ 1910, iv, No. 5, p. q8. THE CLASSIFICATION OF CULICIDJi: 563 First submarginal cell much smaller than the second posterior cell ; proboscis long and bent Palpi long in both sexes Last segment of ^ palpi blunt. Last seg- ment of (^ palpi long and pointed Palpi short in the female First submarginal cell longer than the second posterior cell Legs more or less densely scaly ; head not entirely clothed with flat scales ; all the legs densely scaly. Wings with large pyriform scales Wings with narrow scales Hmd legs only densely scaled Head entirely clothed with flat scales. . Legs uniformly scaled with flat scales. Head and scutellar scales all flat and broad. Palpi of 2 short, of ^ thickened apically and tufted Palpi of 2 longer than in Stegomyia and in (^ long and thin, acuminate, simple Head scales mostly flat, but a median line of narrow curved ones ; scutellar scales flat on mid lobe, narrow curved on lateral lobes and palpi longer than proboscis Head scales mostly flat, irregular, narrow curved ones behind ; mid lobe scutellum with flat scales, lateral with narrow curved ; ^ palpi shorter than proboscis Head scales mostly flat, but a few narrow curved ones in middle in front ; scutellar scales all flat Head scales all flat ; scutellar scales all narrow curved Megarhiniiice. Megarhinus^ Rob. Desvoidy. Ankylorhynchiis^ Lutz. Toxorhyjichites^ Theobald. Culiciftce. Mucidtis, Theobald. Psorophora^ Rob. Desvoidy. Ja?tthi?iosoma, Arribalzaga. Stegomyia^ Theobald. Desvoidea^ Blanchard. Macleayia^ Theobald. Catageiojnyia, Theobald. Scuto?nyia, Theobald. Skusea. Theobald. forked scales of head rod-shaped: Neostelhopheles, James. Includes Atkenii, James; Immaculatus, Theobald ; Culiciformis, James and Liston. These seem to me to be true Anopheles. ■(3) Abdomen with hairs and scales on dorsum of each segment ; ventrally there are six scaly tufts on the apices of six segments. Thorax with scales and a tuft of outstanding ones on prothoracic lobes : Christophersia, James. Type Halli, James. Very close to if rot identical wiih CelJia. {4) Head with narrow curved scales lying rather flat upon head and flat lateral scales, upright forked ones behind. Central lobe of scutellum with tuft of narrow curved scales, lateral lobes with large flat oval scales ; male palpi longer than proboscis, two large apical seg- ments with long projecting hairs : Leslieomyia, Christophers. Type Leslieomyia tcEuio- rhynchoides, Christophers, from Amritsar, India. -(5) Abdomen with first six or seven segments with hairs only, eighth and seventh (?) with scales, also genital processes. Thorax with hairs and narrow curved scales sharp pointed, blunt-ended broad scales on each side of anterior third. No tufts of scales on prothoracic lobes. Head usual type of upright forked scales : Nystomyzomyia, James. Type Rossii, Giles. 5^4 THE ANIMAL PARASITES OF MAN Head with flat scales, except a small median area of narrow curved ones ; scutellar scales all narrow curved ... Head with all flat scales except a thin line of narrow curved ones behind ; scutellar scales all narrow curved ... Head with small flat scales over most of surface, with median line and line around eyes of narrow curved ones ; scutellar scales bluntly spindle or club-shaped ... Head and scutellar scales narrow curved. Wing scales long, narrowly lanceolate, col- lected in spots ; palpi clubbed in S ; five-jointed and rather long in $ Wing scales (lateral) long and narrow, and $ palpi three-jointed, ^ not clubbed and hairy Wing scales at apex of veins dense and rather broad, femora swollen ; small dark species ... Wings with short, thick, median scales and short, broadish lateral ones on some of the veins ; scales mottled ; fork-cells rather short Wings with dense, broadish, elongated, trun- cated scales ... Wings with broad, short, asymmetrical scales Head covered with rather broad, flat, spindle- shaped scales ; scutellum with small flat scales to mid lobe Head clothed with flat, irregularly disposed scales all over, with patches of narrow curved ones ; ^ palpi clubbed Abdomen with projecting flat lateral scales with deeply dentate apices ; wings not ornamented ... ... ... Wings ornamented ; scutellum with flat and narrow curved scales 7. Palpi short in ^ and $ Wings unornamented. Antennae pilose in ^ and ? ; second joint very long Antennas plumose in the <^. Head clothed with narrow curved and flat scales. Mid-lobe of scutellum with six border- bristles. Scutellum with narrow curved scales. Palpi in ? four-jointed, in ^ two-jointed ... Mid-lobe of scutellum with four border bristles. Ho7uardt?ia, Theobald. Danielsia^ Theobald. Hulecoctoinyia^ Theobald. Theobaldia, Neveu-Lemaire. Ciilex^ Linnaeus. Melanoconio?!^ Theobald. Grabhamia^ Theobald. Pseiidotceniorhy7ichus^ Theobald. Tceniorhynch us^ A r r i balzaga. Giles ia, Theobald. Acarlowyia^ Theobald. Lasioco7iops^ Theobald. Finlaya, Theobald. A^deoiiiyi7ia. Deinocerifes, T'heobald. JLdes^ Meigen. HE CLASSIFICATION OF CULICID^ 565 Scutelliim with flat scales. Head clothed with flat scales only. Fork-cells normal length. Mid-lobe of scutellum with four border- bristles. Palpi of $ two-jointed Palpi of ? five-jointed, metallic Fork cells very small or small. Scutellar scales flat. First SLibmarginal cell longer than the second posterior cell; no flat scales on mesothorax ... First submarginal cell smaller than the second posterior cell ; flat scales on meso- thorax... Scutellar scales narrow curved. First submarginal cell as in Uranotaenia ... Wings ornamented with Mansonia-like scales VeraUi7ia^ Theobald. Hcemagogus^ Willis ton. Ficalbia. Theobald. UraiioicEnia^ Arribalzaga. Mi^nofnyia, Theobald. ^■Edeoinyia, Theobald. Section B. — Metanotum ornamented with chastse, squamce or both. o. With chaetae only. Proboscis longer than whole body ; lateral wing scales Tasniorhynchus-like ... Proboscis as long as whole body in 2 ; frons drawn out into a prominence ; wing scales rather broad and long Proboscis not as long as the whole body ; lateral vein scales narrow ... Proboscis not as long as whole body, swollen apically ; wing scales long and broad )8. Metanotum with squam.e and chsetce. Palpi short in S^ and ? . Proboscis straight in $ and S^ ; legs with scaly paddles Venation like Sabethes. Legs simple Venation like Culex Proboscis in ^ elbowed, with two scaly tufts Palpi long in c^, short in $ II. Wings with seven-scaled longitudinal veins: Culex type Phoniomyia^ Theobald. I Binolia, Blanch ard = ( Runchiomyia, Theobald. IVyeomyi'a, Theobald. Dendriomyia^ Theobald. ■Sabethes^ Rob. Desvoidy. Sabethoides, Theobald. Goeldia, Theobald. Limattcs^ Theobald. Joblotina, Blanchard. Heptaphlebo7nyia, Theobald. Section C. — Proboscis short, not formed for piercing Metatarsus longer than first tarsal joint Metatarsus shorter than first tarsal joint ... Coreth^-ina. Corethra, Linnaeus. Mochlonyx^ Ruthe.^ ' Many other genera have been created ; these will be fT)und in my catalogue of Cidicidce ly "Monograph of the Mosquitoes of the World," 1901-10, 5 vols., in my " Novse in my Culicidae," family CulicidiV Genera Insectorum, etc. 66 THE ANIMAL PARASITES OF MAN Notes on the Different Genera. Sub-family. Anophelina. The following Anophelines have been recorded as malaria carriers : — ^^- Anopheles maculipennis^ Meigen. Anopheles bifurcatus,lAViv\dS:MS. * Myzomyia funesta^ Giles. Myzomyia lutzii, Theobald. *Myzoinyia rossii, Giles. Myzomyia listoftii^ Liston. Myzomyia culicifacies^ Giles. Pyretophorus superpictus, Grassi. * Pyretophorus costalis, Loew. Pyretophorus chaitdoyei, Theobald. *Cellia argyrotarsis^ Robineau Desvoidy. Myzorhynchtts pseudopicitis^ Grassi. Myzorhynchus barbirostris^ Van der Wulp. Myzorhynchus sinensis^ Wiedemann. Myzorhynchus paludis, Theobald. Myzorhynchus maiiritianus^ Grand pre. Neocellia stephensii, Liston. Neocellia willmori^ James. Nyssorhynchus theobaldii^ Giles. Nyssorhynchus fuliginosus^ Giles. Nyssorhynchus annulipes, Walker. Those marked with an asterisk (*) also carry the larvae of Filaria bancrofti^ as also do Myzorhynchus minutus, Theobald, and Myzorhynchus Jiigerrinius, Giles_ Genus. Anopheles, Meigen. " Syst. Beschr. Europ. zwei. Ins. I," 1818, ii, p. 2, Meigen ; "Mono. Culicid.," 1903, i, p. 191 ; iii, p. 17 ; and 1910, v, p. 3, Theobald. This genus contains a few large species found either in temperate climates or in hills and mountains of warm climates. The type is the European and North American A. maculipennis. Fig. 399. — ^'xugoi Atiophele'j maciilipejinis, MQ\^t\\. A. 7naculipennis, Meigen. This species and A. bifurcatus are malaria carriers. True Anopheles only occur in Europe, North America, the North of Africa and in the mountains of India, and one has been found by Bancroft similar to A. bifur- catus in Queensland. They are easily told by the absence of scales on thorax and abdomen, and by the ratlier densely scaled wings with lanceolate scales. NOTES ON THE DIFFERENT GENERA 567 Genus. Myzomyia, Blanchard ; Grassia, Theobald. Coinp. re?id. heb. Soc. Biol., No. 23, p. 795, Blanchard ; "Mono. Culicid.," 1910, iii, p. 24 ; V, p. 16, Theobald. This genus occurs in Asia, Africa and South America, Europe and East Indies. The type is M.funesta^ Giles, found in Central and West Africa. Although structur- ally there is not much difference between this genus and Anopheles, they differ greatly in appearance, and there are usually a few narrow curved thoracic scales projecting over the head, whilst the wing scales are much smaller in proportion, and the wings more uniformly spotted, always so along the costa. Ftmesta and lutzii are undoubtedly malaria bearers and also rossii. Genus. Neomyzomyia, Theobald. "Mono. Culicid., "1910, v, p. 29. A single species only occurs in this genus, N. elegans^ James, from India. In this genus, which is near to Myzomyia, the fourth long vein is very near the base of the third, and there are outstanding scales on the prothoracic lobes, and there is a marked tuft of dense scales at the posterior angles of the head. Genus. Cycloleppteron, Theobald. " Mono. Culicid.," 1903, ii, p. 312 ; 1903, iii, p. 58 ; 1910, v, p. 33. Two common species only occur in this genus, C. grabhamii, Theob., from Jamaica, and C. mediopiinciatus^ Theob. (Lutz., ms.), from South America. The chief character is the presence of large black inflated pyriform scales on the wings. The palpi are densely scaled. Neither have been shown to be malaria bearers. Genus. Feltinella, Theobald. " Mono. Culicid.," 1907, iv, p. 56. A single species, so far only found in this genus. The basal lobes of the male genitalia of two segments, the prothoracic lobes with dense outstanding scales. The species, F. pallidopalpi^ Theob., occurs in Sierra Leone. Genus. Stethomyia, Theobald. " Mono. Culicid.," 1903, iii, p. 13 ; 1907, iv, p. 59 ; iQio, v, p. 35. Four species occur in this marked genus — one S. Jiimba, Theob., from British Guiana and Para, another S. fiagalis, Theob., from the Malay States, S. culici- fonnis, James and Liston, from India, and S. pallida, Ludlow, from India. The former may be a malaria carrier, for Dr. Low says : " Malarial fever is got amongst the Indians and often of a severe type. In that connection it is interesting that in the interior, at a place called Corato, I got an entirely new Anopheles in large numbers." The genus is easily told by its unornam-ented wings, flat head scales, mammillated prothoracic lobes and long thin legs. Genus. Pyretophorus, Blanchard; Howardia, Theobald. Compt. rend. heb. Soc. Biol., No. 23, p. 705, Blanchard ; Journ. Trop. Med.,v, p. 181 ; and " Mono. Culicid.," 1903, iii, P- 13 ; 1910, v, p. 36, Theobald. Forty-four species come in this genus, of which Anopheles costalis, Loew, is the type. ^68 IHE ANIMAL PARASITES OF MAN This genus is found in Africa, India, Europe and in Australia. Three species are proved malaria bearers, namely, P. costalis, Loew, P. chaudoyei, Theob., and P. superpictus, Grassi. Members of this genus can be told by having narrow curved thoracic scales, hairy abdomen, and much-spotted wings. Genus. Myzorhynchella, Theobald. "Mono. Culicid.," 1907, iv, p. 78. In this genus the thorax has distinct, narrow curved scales, and the abdomen is hairy, the wing scales broad and lanceolate, and the head with broad scales not closely appressed, but not forked or fimbriated. Five species are known : lutziy Cruz ; parva, Chagas ; nigritarsis, Chagas ; tibiomaculata, Neiva ; gilesi, Neiva ; and nigra, Theobald. They are all recorded from Brazil, and 7iigra also from Mexico. Genus. Manguinhosia, Cruz, in Peryassu. •*Os Culicideos do Brazil," 1908, p. 112. A single marked species from the Brazils. The thorax has piliform curved scales, and some narrow curved and flattened ones on the sides. Abdomen pilose, except the last three segments which are scaled. No tufts of scales on posterior femora. Allied to Lophoscelomyia, but at once told by the absence of scale tufts on the hind femora. M, lutzi, Cruz, Brazil. Genus. Chrystya, Theobald. " Rep. Sleeping Sickness, Roy. Soc. Eng.," 1903, vii, p. 34. A very marked genus in which the hairy abdomen has very long, dense, hair-like, apical, scaly tufls to the segments. A single species only so far known, C. implexa^ Theobald, from Africa (Uganda, Sudan, etc.). Genus. Lophoscelomyia, Theobald. Entomologist^ 1904, xxxvi, p. 12. A single species only, from the Federated Malay States. The hind femora have dense, apical scale tufts ; the thorax long, hair-like curved scales ; abdomen pilose, except the last two segments which are scaly ; wings with broad, blunt, lanceolate scales. Genus. Arribalzagia, Theobald. " Mono. Culicid.," 1903, iii, pp. 13 and 81 ; and 1910, v, p. 48. Two species only occur, found in South America. The thorax and abdomen have scales and hairs respectively, as in Fyretophorus, but the abdomen has in addition prominent lateral apical scale tufts to the segments and a scaly venter. Wings with membrane tinged in patches and wing scales bluntly lanceolate and very dense. The type is A. maculipes, Theob. found in Trinidad and Brazil; A. pseudoinaculipes^ Cruz, also in Brazil, Genus. Myzorhynchus, Blanchard ; Rossia, Theobald. Compt. rend. heb. Soc. Biol., 1902, No. 23, p. 795, Blanchard ; Journ. Trop. Med., 1902, p. 181, Theobald ; "Mono. Culicid.," 1903, iii, p. 84; 1907, iv, p. 80; 1 9 10, V, p. 49. A very marked genus of large, dark, densely scaled species, found in Europe, Asia, Africa and Australia. The thorax with hair-like curved scales; the abdomen NOTES ON THE DIFFERENT GENERA 5O9 with ventral and apical scales, and a median ventral apical tuft, and with very densely scaled palpi in the female, and densely scaled proboscis. It seems to be mainly an Asiatic and East Indian genus, but three species occur in Africa and one in Australia. They are mostly sylvan species and bite severely. Fourteen species are known. Five are malaria carriers {vide list, p. 566). Genus. Nyssorhynchus, Blanchard ; Laverania, Theobald. " Mono. Culicid.," 1910, iii, p. 14 ; v, p. 55, Theobald ; Compt. rend. heb. Soc. Biol.^ No. 23, p. 795, Blanchard. A group of small, closely allied species found in Asia, Africa and Australia, twelve out of the twenty species coming from India. The thorax is covered with narrow curved and spindle-shaped scales, abdomen with small, flat or narrow curved dorsal scales, especially on the apical segments or in patches ; the legs are always banded or spotted with white, and the tarsi have as a rule one or more pure white segments. • (This banding and spotting is of no generic value, however.) The species show considerable seasonal variation. The type of the genus is N. maculaius^ Theobald. Three are malaria carriers {vide list, p. 566). Genus. Cellla, Theobald. " Mono. Culicid.," 1903, iii, p. 107 j 1910, v, p. 67. Very marked Anophelines, with densely scaly abdomens, the scales irregularly disposed on the dorsum and forming dense lateral tufts ; thorax with flat spindle- shaped scales ; palpi densely scaled and also the wings. The type of the genus is the African C. pharoensis^ Theob. It is represented in Asia by C. kochii, Donitz ; in West Indies and South America by C. argyrotarsis , Desvoidy, and C. bigotii, Theob. ; in Africa by C. squamosa^ Theob., etc. C. argyrotarsis , Desvoidy, and C. albimana, Wiedemann, are undoubtedly malaria bearers. Genus. Neocellia, Theobald. "Mono. Culicid.," 1907, iv, p. in. Allied to Cellia, but has no lateral scale tufts. Three species recorded froui India. Genus. Kert^szia, Theobald. "Ann. Mus. Nat., Hung.," 1905, iii, p. 66. This genus has the thoracic scales hair-like, except a few narrow curved ones in front ; abdominal scales long, broad and irregular. A single species, K. boliviensis, Theob. from Bolivia. Genus. Manguinhosia, Cruz. The thorax has narrow hair-like curved scales and some broad straight scales ; others spatulate on the sides. Abdomen with fine hairs, except the last three segments which are scaly. Tufts of scales on the hind femora. Wing scales lanceolate. The type is M. lutzi, Cruz, from Brazil. cyo THE ANIMAL PARASITES OF MAN Genus. Chagasia, Cruz. "Brazil-Medico," 1906, xx, pp. 20, 199. This genus can at once be told by the antennal segments having many dense scaly tufts. Type, C.fajardoi, Lulz, from Brazil. Genus. Calvertina, Ludlow. Canadian Entomologist, 1909, xli, pp. 22, 234. The antennae in this genus have outstanding scales on the second segment, more appressed ones on the first. At least one abdominal segment with long, flat, more or less spatulate scales. Type, C. lineata, Ludlow, from Philippine Islands. Genus. Birdnella, Theobald. "Ann. Mus. Nat. Hung.," 1905, iii, p. 69. At once told by the first submarginal cell being very small. Type, B. gracilisy Theob. from New Guinea. Sub-family. Megarhininae. Three genera occur in this marked sub-family ; they are the largest of all mos- quitoes, and are very brilliantly coloured, and many have tail fans. They occur in North and South America, Asia, Africa, and Australia. The long curved proboscis is very marked. They are usually spoken of as elephant mosquitoes ; some are vicious blood-suckers at times. The three genera tabulate as follows : — a. Palpi long in both sexes. &. Last segment of $ palp round or blunt as if broken Genus A/if^^^r/zZ/mi-, R. Desvoidy. )8;8. Last segment of $ palp long and pointed Genus Aiikylorhyjichus, Lutz. 00. Palpi of female short of male long. Palpi of female not more than one-third length of proboscis ... Q^xvws roxorhyfichites,T\i^o\i2\d.. Genus. Megarhinus, Robineau Desvoidy. " Mem. Soc. d'Hist. nat. de Paris," 1827, iii, p. 412 : " Mono. Culicid.," 1901, i, p. 215 ; 1903, iv, p. 163; 1907, iv, p. 128 ; 1910, v, p. 89. All large brilliant mosquitoes with long palpi in both sexes and, as a rule, with a caudal fan of scales ; the proboscis is long and bent. They are all sylvan species, and are not so far recorded as biting man. Genus. Toxorhynchites, Theobald. " Mono. Culicid.," 1901, i, p. 244 ; 1903, iii, p. 1 19 ; 1907, iv, p. 140 ; 1910, v, p. 95. Differs from the former genus in that the female palpi are short. The palpi may have one, two or three minute terminal segments. Banks's genus Worcesteria has three. The elephant mosquito of India {T. iimnisericors), Walker, bites very severely. They are sylvan species. NOTES ON THE DIFFERENT GENERA 571 Sub-family. Culicinae. Genus. Mucidus, Theobald. "Mono. Ciilicid.,'" 1901, i, p. 268 ; 1910, v, p. 125. This genus is so far confined to Australia, West and Central Africa, India, East Indies and Malay Peninsula. They are all large mosquitoes, easily told by the whole body being more or less covered with long twisted scales, giving them a mouldy appearance, and the legs densely scaled with outstanding scales ; the wings .with large parti-coloured scales. The Australian M. a//ernans, Walker, occurs in larval form both in fresh and salt water. The adults bite man. Genus. Psorophora, Robineau Desvoidy. " Mem. de la Soc. d'Hist. nat. de Paris," 1827, iii, p. 412, R. Desvoidy ; " Mono. Culicid.," 1901, i, p. 259; 1903, iii, p. 130; 1907, iv, p. 158 ; i9io,v, p. 123, Theobald. This genus is confined to the Americas and the West Indies. Several species exist which can easily be told from Mucidus by the absence of long twisted scales and the narrower wing scales. The legs are densely scaled and the thorax orna- mented with flat spindle-shaped scales. F. ciliata^ Robineau Desvoidy, occurs in both North and South Americfj, and bites man. Genus. Janthinosoma, Arribalzaga. "Dipt. Arg.,' 1891, p. 52, Arribalzaga; "Mono. Culicid.," 1901, i, p. 253 ; 1903, iii, p. 124 ; 1907, iv, p. 152 ; and 1910, v, p. 118, Theobald. Hind legs only densely scaled ; some of the hind tarsi are always white. The venation is as in Culex. The abdomen is metallic and iridescent. They all bite man and occur only in the Americas and West Indies. Genus. Stegomyia, Theobald. " Mono. Culicid.," 1901, i, p. 283 ; 1903, iii, p. 130 ; 1907, iv, p. 170; 1910, v, p. 151. This, the most important genus in the Culicince^ can be told by the head and scutellum being clothed with flat scales and the thorax with narrow curved ones. About forty species are known in this genus, occurring in Southern Europe, Asia, Africa, Australia, the Americas, East and W^est Indies, and on most oceanic islands. Many of them seem to be vicious blood-suckers. They are mostly black and white mosquitoes, and several seem to go by the name of tiger mosquitoes. The genus contains the yellow fever mosquito (6*. fasciata^ Fabricius), the only one that need be dealt with in detail here. The chief known species tabulate as follows : — A. Proboscis banded. a. Legs basally banded. Thorax brown, with scattered creamy-white scales ... ... ... ... ... annulirostris^ Theobald. Thorax black, with narrow, curved golden scales ... ... ... ... ... periskelta^ Giles. «o. Legs with basal and apical banding. Fore legs with no bands ; mid with apical and basal bands on first and second tarsals, hind with basal bands. Thorax white in front, with a brown eye- like spot on each side ///i?;;/^^?///, Theobald. 36 572 THE ANIMAL PARASITES OF MAN AA. Proboscis unhanded. /3. Legs basally banded. % Abdomen basally banded. Thorax with one median silvery-white line Thorax as above, but pleurae with white lines Thorax similar, but two white spots near where line ends Thorax with two median yellow lines and lateral curved silvery lines Thorax with two short median lines and a white patch on each side Thorax with large lateral white spots in front, smaller ones by wings, two narrow median lines and two posterior sub- median white lines... Thorax with a white W-shaped area in front, a prolongation curved on each side enclosing a brown eye-like spot ... Thorax with white frontal median spot, two large lateral spots, a small one in front of the wings, a narrow median white line and narrow sub-median ones on posterior half. Last two hz?id tarsi white ... Thorax brown, with broad white line in front extending laterally towards wings, where they swell into a large patch, a white line on each side just past wing roots. Last two hind tarsi white Thorax with silvery white spot on each side in front, small one over roots of wings and white over their base. Last two hind tarsi white Thorax with two lateral white spots, front ones the largest, a small median one near head, two yellow median lines, a short silvery one on each side before the scutellum Thorax with silvery white scaled area in front and another on each side in front of wings Thorax with median yellowish-white line, a silvery patch on each side in front of wings extending as a fine yellow line to the scutellum, and another silvery spot before base of each wing Thorax with small grey-scaled area in front of wing roots and three short creamy lines behind scutellaris^ Walker. Pseudosctitellaris^ Theobal d. geleln7tensis, Theobald. fasciata^ Fabr. nigeria^ Theobald. Ulii^ Theobald. W-alba, Theobald. wellinannii^ Theobald. albipes^ Theobald. pseudonigeria, Theobald. simpsoni, Theobald. argenteojnaculata^ Theobald. poweri^ Theobald. ininutissima^ Theobald. NOTES OX THE DIFFERENT GENERA 573 Thorax (?) denuded ; abdomen black ; fifth segment with yellow basal band ; sixth unhanded ; seventh, two median lateral white spots ; eighth, two basal lateral white spots ; second hind tarsal nearly all white ... 77. Abdomen unhanded. First hind tarsal all white, second basally white, last two dark. Thorax chestnut brown, with a broad patch of white scales on each side in front and a median pale line 80. Legs with white lines as well as basal bands. Thorax brown, with white lines ; abdomen with basal bands 000. Fore and'mid legs with apical bands, hind basal. Fourth tarsal of hind legs nearly all white Mid metatarsi with basal pale banding, base and apex of hind, also base of first tarsal pale 0000. Legs unhanded. 5. Abdomen basally banded. Thorax with front half white, rest bronzy- brown ... ... Thorax deep brown, with scattered golden scales, showing two dark eye- like spots ; head white, dark on each side and behind ... Thorax brown with golden abdomen with narrow basal fifth and sixth segments only ... 55. Abdominal banding indistinct. Thorax with broad silvery white on each side in front ?55. Abdomen unhanded. Thorax with six silvery spots Abdomen with apical white lateral spots. Thorax unadorned, except for pale scaled lines laterally Abdomen with basal white lateral spots. Thorax with two pale median parallel lines and two silvery lateral spots Thorax unadorned. A white spot middle of head ... No white spot ... AAA. Proboscis yellow basally, dark apically. Abdomen with apical pale bands ... A \AA. Proboscis with median interrupted white line on basal half. Head black, anterior margin grey 2555. stripes ; bands on patch 55355. {/uh'a, Theobald. terreus., Walker. grantii^ Theobald. mediopunctata., Theobald. assamemis, Theobald. pseudonivea, Ludlow. albocep/iala^ Theobald. nuriostrjaia., Banks. albolateralis., Theobald. argenteopmtcfafa, Theobald. punctolateralis., Theobald. iiiinuta, Theobald. tripunctata., Theobald. amesii, Ludlow. crassipes, Van der Wulp. albomarginata^ Newstead. 574 THE ANIMAL PARASITES OF MAX Stegomyia fasciata, Fabriciiis (Yellow Fever Mosquito). This insect, which is the proven carrier of yellow fever, is commonly called the tiger, brindled, spotted day or striped mosquito. It is also referred to by some writers as S. ca/opus, Meigen. It is subject to considerable variation in colour, but the thoracic markings are generally very constant. The general colour is almost black to deep brown, the head with a median white area, white at the sides and in front around the eyes ; the thorax has two median parallel yellow lines, a broad curved silvery one on each side and white spots at the sides ; tlie scales on the intervening spaces of the thorax are brown. The dark abdomen has basal white bands and basal white lateral spots. The dark legs have basal white bands, the last segment of the hind legs being all white except in a variety from South America and the West Indies {luciensis), which has the tip of the last hind tarsal dark. The abdomen may also vary in colour, some having pale scales over most of the surface {queenslandetisis). The food of the adult female consists mainly of man's blood, but she will also feed on dogs and other animals. The male has been said to bite, but such is very unusual. This mosquito bites mainly in the daytime up till about 5 p.m. The adults breed the first day after emergence. They may live a consider- able time, Bancroft having kept females for two months in confinement. The ova are laid separately, often in chains ; they are black, oval, with a reticulated membrane outside, some of the reticulated cells containing air. They may hatch in from six to twenty hours, the larval stage nine days, the pupal stage three ; thus the whole cycle may be completed in from twelve to thirteen days. The ova when dry can remain undeveloped for a considerable time. The larvae are greyish-white, with short, thick siphon, and feed at the bottom of the water, only coming to the surface now and again to breathe. This is almost entirely a domesticated gnat, seldom being found far from man's habitations. Its larvne occur in such small collections of water as old sardine tins, jam-pots, calabashes, puddles, barrels, wells — in fact, wherever water is held up, even to the gutters of houses. Not only are they found breeding on land, but also on board ship, although they prefer artificial collections of water. They may also breed in larger natural collections. This insect is easily transported by steam and sailing ships and by train, and this doubtless explains its very wide distribution. The adults may live for fifty days, and it is on this account and their frequent occurrence on ships that danger lies in regard to the Panama Canal. An infected insect may leave that endemic centre of yellow fever and live until the vessel arrived at the Philippine Islands and fly ashore, and so introduce the disease for the wdi^w^ fasciata possibly to spread. Roughly the distribution of this pest is as follows : Africa from South to North, but especially along the coast and up the Nile. In Asia, in India, Ceylon, Burma, Siam, along the ports of the Malay Peninsula, in French Cochin China, Philippine Islands, the Andaman and Nicobar Islands, Japan, Malay Archipelago, and East Indies, Turkey in Asia, Arabia and Palestine. In Australia it occurs in Queensland, New South Wales, Victoria and South Australia. In Europe in Italy, Spain, Portugal, Greece, in the Mediterranean Islands. In South America, Central America, Mexico, North America, and the W^est Indies it is very abundant, and it also is found in the Bahama Islands, Fiji, Sandwich Islands, Samoa, the Azores, Teneriffe and Santa Cruz, Pitcairn Islands and Bermuda. For a full account of its distribution the reader is referred to the following : "The Distribution of the Yellow Fever Mosquito {Stegomyia fasciata^ Fabricius) and General Notes on its Bionomics;" "Mem. le"" Congres international THEOBALDIA, CULEX 575 d'Entomologie, 191 1, ii, pp. 145-170, F. V. Theobald. In addition to being the yellow fever carrier, it is supposed by Wenyon to be the intermediate host of the parasite of Bagdad sore. Stegomyia scutellaris, Walker. A vicious biter, found in India, China, Malay, East Indies, and Ceylon. The thorax has one median silvery stripe, and so can easily be told from S.fasciata. A very similar species occurs in Fiji, but can be told by the pleurae having white lines, not spots {S. pseudoscutellaris^ Theobald). It is the intermediate host of filaria in Fiji (Bahr). A number of nearly allied genera occur here (c^'ide synoptic table). Genus. Theobaldia, Neveu-Lemaire. Theobaldinella, Blanchard. Includes several large Culicines, of which T. a?tmilata^ Meigen, is the type. The wings are usually spotted {aniiulata^ i?tcidens, etc.), but may be nearly plain {spathz- palpis). The males have the palpi swollen apically, and the females have long five-jointed palps. Several of these are vicious biters. Theobaldia annulata, Meigen. This large gnat (6 mm. long) can be told by its wings having five large spots of dark scales and by its legs having broad basal white bands to the tarsi. The larvas occur in rain barrels and small pools. It is essentially a domestic form, occurring in houses and privies. Its distribution is Europe generally and North America. The bite is very severe, and in some districts gives rise to painful oedema.' Theobaldia spathipalpis, Rondani, occurs in Italy, Mediterranean Islands, Palestine, the Himalayas, Khartoum, and in South Africa. It is about the same size as T. annulata, but is yellowish-brown in colour, with striped thorax and mottled and banded legs. It occurs in privies and bites very severely. Genus. Culex, Linnaeus. " Syst. Nat. Ed.," 1758, x, Lmn.neus ; "Mono. Culicid.," 1901, i, p. 326 ; 1910, V, p. 322, Theobald. This large genus still contains many forms which should be excluded. The species normally have narrow curved median head- scales, and similar ones on the Fig. 400.— Wing of a Culex. scutellum ; the female palpi are shorter than in the former genus and the male palpi are pointed ; the lateral vein-scales are narrow and linear. The type is Culex pipie?ts, Linn., the common gnat of Europe. The thorax Theobald, "Second Report on Economic Zoology," 1903, p. 9. 576 THE ANIMAL PARASITES OF MAN is covered with narrow curved golden-brown scales, the abdonien has basal pale bands to the segments and the legs and proboscis are unhanded. The stem of the first submarginal cell is always less than one-fifth the length of the ceil. It lays its eggs in rafts in water-butts, etc., and even in the foulest water. They are first deposited in England in June and July, and again soon after hatching in August. In some districts this gnat bites man viciously, in others not at all. The common tropical gnat {Culex fatigans, Wied). This resembles the European Culex pipiens, but can always be told by the stem of the first submarginal cell always being much longer than it is in C. pipie?ts. This is one of the species that has been proved to transmit filariae to man, etc. Varieties of it occur in almost every country between 40° N. and S., having a very similar range to S. fasciata. In all countries it appears to be connected with the transmission of Filaria bancrofti^ and it is also said to carry the micrococcus of dengue fever. Genus. Melanoconion, Theobald. " Mono. Culicid.," 1903, iii, p. 238 ; 1907, iv, p. 507 ; 1910, v, p. 455. This genus is composed of eight species, most of which are small black gnats which bite viciously and which occur in swamps and jungles. They can at once be told from Culex by the veins of the wings having dense broadened scales on their apical areas and along the upper costal border. The femora and apices of the tibiae are swollen. The black mosquito, Melanoco7iion atraius, Theob. This small gnat is a very troublesome pest in swamps in the West Indies. The female bites both by day and by night, and the bite causes severe irritation. The larvie live in permanent ponds. It is almost black in colour, but sometimes presents a dull coppery sheen ; each segment has small lateral basal white spots. Length 2*5 to 3 mm. It occurs in Para and British Guiana as well as in the West Indies. Ordinary mosquito netting is no use for keeping off this pest. Genus. Grabhamia, Theobald. " Mono. Culicid.," 1903, iii, p. 243.; 1907, iv, p. 284 ; and 1910, v, p. 277. Allied to Culex, but separated by the wings having short fork-cells, mottled scales, the median ones thick and also some of the lateral ones short and broad ; the last two joints of the male palps are very slightly swollen. The eggs are laid singly, not in rafts, and the larvae have short, thick siphons. Ten species occur and are found in Europe, North America, West Indies and Natal. G. dorsalts, Meigen, bites severely in Europe. G. sollicitans. Walker, is a great scourge along the New Jersey Coast and at Virginia summer resorts and in Florida. It breeds in brackish water and is the most common mosquito of the Atlantic seaboard. Genus. Pseudotaeniorhynchus, Theobald ; Taeniorhynchus, Theobald, non-Arribalzaga. Differs from the former in having the whole wing veins clothed with dense, broadish elongated scales. 1 hey occur in South America {T. fasciolatus, Arri.), in Africa (T. tenax, Theob.), in Europe {T. richardii, Ficalbi). The latter bites very severely. OTHER NEMATOCERA 577 Genus. Taeniorhynchus, Arribalzaga ; Mansonia, Blanchard ; Panoplites, Theobald. Compt. rend. heb. Soc. Biol., 1901, iii, -^j^ p. 1046; "Mono. Culicid.," 1901, ii, p. 173; and 1910, v, p. 446, Theobald. A very marked genus, easily told by the broad asymmetrical wing scales. It occurs in Africa {T. africana and T. major, Theob.) ; in Asia {T. uniformis, Theob. ; T. annuHpes, Walker, etc.) and in Australia (7". aiistraliensis) ; in the Americas and West Indies (7". titillans, Walker). The eggs (fig. 395, d) are peculiar in form and are laid separately ; the larva has not been described ; the pupa has long curved siphons. They mostly occur along rivers, in swamps and forests, and bite very severely. They also enter houses {T. titillans). T. imiforinis is most troublesome during the rains. The saliva is strongly acid. Both these species carry the larvae of Filaria bancrofti. Genus. Chrysoconops, Goeldi. " Os Mosq. no Para," 1905, p. 114, Goeldi; "Mono. Culicid.," 1910, V, P- 433, Theobald. Bright yellow or yellow and purple mosquitoes, with rather dense wing scales. Numerous species occur in Africa {aurites, amtettii, fuscopentiatus, etc.), others in India, Australia and South America. Low found filarias in the thoracic muscles o^ fuscopennatus in Uganda. Several of the Aideojuyina bite, especially the small Uranotccnias. They are all sylvan species, seldom entering houses. They need not, therefore, be referred to here. For full details of the Culicid genera and species the reader is referred to my monograph' and other works mentioned below. Other Nematocera. Other nematocerous flies are midges, daddy-long-legs and sand-flies. The ones which cause annoyance to man besides Cidicidce are the followmg : — Sand-flies {Simididcr), certain midges {Chironoinidce), and a few owl midges {Psychodidcp). The Neinatocera have long thread-like jointed antennas and their pupae are, as a rule, naked ; the larvae have a distinct head and can thus be told from the next section {Brachycerd). Family. Simulldae. This family consists of a single genus, Simulium, Latreille, which Roubaud has recently divided into two sub-genera called Pro-Simulium and Eu-Simulium. These insects, which are frequently spoken of as sand-flies, are found in all parts of the world; they are all small insects varying from 1-5 to 3 mm. The females are very bloodthirsty, but the males appear to be incapable of sucking blood. The head sunk under the humped thorax ; antennae short, straight ; palpi short » ** A Monograph of the Cnlicidtt of the World," 5 vols, and atlas, 1901 to 1910, British Museum (Nat. Hist.) ; and the following : Howaid, Dyar and Knab, "The Mosquitoes of North and Central America and the West Indies," 1912 ; James and Liston, "The AnophelinK of India," Leicester, 1908; "The Culicidcc of Malay," Inst. Med. Res., Fed. Malay States, iii ; Ann. Trop. Med. and Par., papers by Ne\\ stead and Carter; Mem, Inst. Oswaldo Cruz, papers by Luiz, Neva, Chagas ; and the Bulletin of Entomological Research, etc 578 THE ANIMAL PARASITES OF MAN and broad, of four segments, bent ; wings broad and in some iridescent, legs stout. The male has holoptic eyes, whilst in the female they are small and widely separate. The sucking proboscis is short. The thorax and abdomen are clothed with short hairs which may form spots and markings ; these are golden, silvery, grey, or brown- ish. In the sub-genus Pro-Simulium the second segment of the hind tarsi in both sexes is elongate, linear, and without a basal notch ; in Eu-Simulium it is short, curved, and dorsally notched at the base. Simulidce often occur in swarms, and attack not only man but cattle, horses, and poultry. In some districts they are more annoying than mosquitoes. Their life-cycle has been most completely worked out by King, in Africa. The larvae and pupae occur in swiftly flowing water, by waterfalls, in rapids, etc. The ova are laid in gelatinous masses on plants or rocks close to or overhanging the water. The larva is cylindrical, enlarged posteriorly, where it is provided with a sucker, by means of which it attaches itself to a rock, water weeds, debris, etc. ; anteriorly it has a proleg close behind the head on the lower surface. The head is dark and chitinous. The respiration takes place by means of branched tracheal gills which protrude from the dorsal surface of the last body segment ; they are retractile. The colour varies from deep green to yellow or almost black. Their food consists of algae and other organisms in the water brought to their mouth by two fan-like organs .placed on the head. The larva? can crawl from place to place by means of the thoracic proleg ; they occur in masses, usually in a more or less erect attitude. A net- work of threads is spun on their support, by means of which King tell us " they are enabled to maintain their position against the strongest current ; frequently they will leave their support and let themselves out into the stream anchored by threads of silk and enabled by them to return." When full fed the larva spins a pocket-shaped cocoon on the support, within which it pupates. The pupa is motionless and has a pair of branched spiracles pro- jecting from behind the head. When the adult emerges, a bubble of air collects around it, and in this it floats to the surface and at once takes wing. The European species take a month to complete larval life, a week being spent in the pupal stage. The flies are most restless, and even when stationary continually move their legs about like feelers. Sometimes the swarms consist entirely of females, sometimes early in the season mostly of males. The females pierce the skin of humans on tender spots, such as ears, the fore- head, around the eyes and nose, and crawl into the cavities. They are quite harmless at night, mainly attacking about sunrise and sunset. Some crawl up the arms and legs and down the neck, and leave behind little red weals which itch intensely {S. dammsum, Theob.), and blood may flow freely from the wounds. The following are some of the worst species : — Simulium columbaschensis, the " Kolumbatz fly," which abounds in the damp marshy lands along the Danube, and is a great plague toman and beasts in Hungary, and is also abundant in Austria and Moravia, and is most numerous after inunda- tions from the Danube. They sometimes appear in such swarms that it is impossible to breathe without getting them into one's mouth. There are instances of children being killed by these flies when left on the ground by their mothers when working in the fields. S. damnosum, Theob. This occurs throughout Equatorial Africa and is known as the «'jinja fly" in Uganda, the "fouron" in the French Congo, the "kilteb" m the Sudan. It is a most vicious biter, and in some parts occurs in "belts'"; Dr. Christy found one such extending from the shores of the Victoria Nyanza northwards along the right bank of the Nile for twelve or fifteen miles or more, and perhaps three or four miles wide. In this area the flies swarm in millions at certain CHIRONOMID^ 579 seasons, so much so that the natives have to leave their plantations. The bite causes a weal, marked by a drop of blood. S. gnseicolHs, Becker. The so-called " nimitti " occurs in Upper Egypt and the Anglo-Egyptian Sudan. It lives near the river and is not found more than half a mile from it. Human beings are bitten on the face and hands, animals in the region of the pudenda. S. lattpes, Meigen. This is a European species, also found in Natal. S.zvellmanni, Roubaud. The " ohomono " of Angola, where it bites viciously and is dreaded by the naked porters. S. buissoni, Roubaud. Occurs in abundance in the Marquesas Islands. It has been suggested that this species may help to propagate leprosy.' A large number of these insects have been described by Lutz in Brazil.-* A Simulium sp. (?) is very harmful to poultry in Cape Colony.=^ In America, Simiilidce are most annoying. One, S. meridio)iale, Riley, also known as the turkey gnat in the Mississippi Valley, has been supposed to be the carrier of chicken cholera ; anyhow, it has caused the death of thousands of chickens and turkeys in Virginia annually.'' In Mexico Townsend found a Simulium which was named S. occideittalis^ which caused great annoyance to man, many people being so susceptible to them as to Fig. 401.— Wing of Simulium. Fig. 402.— Wing of Chironomus. preserve through the gnat season a chronic inflammation of the exposed parts of face and neck, resulting from the repeated bites giving rise to sores.'* Men and horses have been partially incapacitated by the bites of sand-flies or Simulium in a Hampshire wood (Cantlie, Brit. Med. Jour7i., April 28, 1900, v. No. 2,052, p. 1023). Family. Chlronomidae (Midges). The Chirotioinidce or midges are not only frequently mistaken for mosquitoes, but some are very annoying to man by biting him as mosquitoes do. They are easily distinguished from true mosquitoes {Culicidcs) by the following characters : (i) head small, often retracted under the cowl-like thorax ; (2) no scales to the wings or body; and (3) the different arrangement of veins on the wings (fig. 402). Two genera are important as annoying man, namely, Culicoides, Latreille, and Johannseniella, Williston. The larvae of Chironomidce are either aquatic, both fresh water and marine, and help to make the former foul,*^ according to Slater, or may, as in Ceratopogofiifice, live beneath the bark of trees, etc. The pup£e are very varied and also the life-histories of the different genera.^ The blood-sucking habit is confined to the sub-family Ceratopogofiince . 1 Btdl. du Mus. d'Hist. nat,, 1906, xii, p. 522. ^ Mem. Inst. Oswaldo Cruz, 1910, ii, fasc. 2, pp. 211-267. '^ C. Fuller, "A New Poultry Pest," 1899, Leaflet No. i, Dept. Agric. 4 Insect Life, 1888, i, p. 14. ^ Ibid., 1893, v, p. 61. " Entomologist, 1879, p. 89. ' Theobald, "An Account of British Flies," i, p. 172. 58o THE ANIMAL PARASITES OF MAN Sub-family. Ceratopogoninae. This sub-family of midges consists of very small species varying from i to 2 mm. in length ; the wings have darkened areas, and the second longitudinal vein is wanting, and the first and third veins are stouter than the others and placed close to the anterior margin, the fourth and fifth are forked ; the antennas in both male and female are composed of fourteen segments, six or eight in the males bearing long hairs. The chief blood-suckmg species belong' to the genera Culicoides, Latreille, and Johannseniella, Williston. The latter genus differs from the former in the absence of an empodium or median appendage on the last segment of the tarsi. The genus Ceratopogon, as restricted by Kieffer, is not supposed to take vertebrate blood, but Austen has recently noticed that the type specimen of C. castafieus, Walker, and a new species described by him, apparently have their bodies distended with blood. The wings in the Ceraiopo^onincu are carried flat when at rest. Fig. 403. — A Ceratopogon, or midge. Greatly enlarged. In spite of their small size the females are the most bloodthirsty and annoying of all insects. The Culicoides, which are often called " sand-flies," bite during the day and rarely at night. Usually they are most troublesome between 3 and 6 p.m. They frequently attack in swarms, especially in the open, and owing to their minute size can get through fine mosquito netting. Some of them produce a distinct " buzz " when on the wing. These insects are found in all parts of the world. No species has been definitely connected with any disease, but Culicoides has been suspected of carrying the germs of Delhi boil. The larvas of Culicoides are elongate in form and have smooth bodies composed of thirteen segments including the head, which is horny ; there is no proleg on the first segment as seen in Chironomus, and on the anal segment are retractile gills. They are very active and live in the sap of various trees which saturates diseased bark. The pupas are smooth, but the abdominal segments bear a transverse row of small spines. Austen describes a number of Culicoides and one Johannseniella and three Ceratopogons from Africa,^ and Lutz^ a number of this sub-family from Brazil, ' Bull. Ent. Res., 1912, iii, pp. 99-108. - Mem. Inst. Ostvaldo Cruz, 1913, v, fasc. pp. 81.99. I, pp. 45-72, pis. 6-8 ; and 1914, vi, fasc. 2, PSYCHODID.^i: 581 including a new genus, Centrorhyncus. Another genus, Terseslhes, Townsend (Centrotypus, Giassi ; Mycleiotypus, Noe), also occurs in Brazil. Culicoides omatus^ Taylor, is described from Townsville, Australia, found in mangrove swamps. It is a very vicious biter and causes considerable irritation, settling on hands and wrists (Taylor, Rep. Ent. Aust. Insi. Trop. Med. [191 1], 191 3, p. 24). Family. Psychodidae (Owl Midges). This family of diptera is of considerable importance, not only on account of the blood-sucking habits of some species, but especially on account of one at least having been proved to be the carrying agent of " papataci " fever, a three-day fever very prevalent in Malta and several parts of Southern Europe in the autumn. It is also possible that these small flies are connected with the formation of ^' Delhi boil," caused by a protozoan parasite. PsycJiodidce are all very small flies, many of which have a moth-like appearance, and owing to their fluffy nature are spoken of in Britain as "owl flies," sometimes also as " window flies." Their bodies and wings are covered with hairs, densely in some (sub-family Psychodino'), and in a few with patches of flat scjuamas. In the non- blood-sucking Psychodiitce the wings are carried in a peculiar manner downwards over the body, to a slight extent resembling the Hepialidce, or swift moths. The Fig. 404.— An owl midge, Phlehotomtis sp. Greatly enlarged. (From Giles's '* Gnats or Mosquitoes.") wings may be ovoid or lanceolate, and have a marked venation as seen in the figure. The proboscis is short and non-suctorial in the majority of genera, but in the sub- family Phlebotojuiiice it is elongated and hard. The antennas are long and of sixteen segments, and bear whorls of fine hair. There are two sub-families, Psychodincc 2ir\d Phlebotonii?icc ; in the former the mouth is not suctorial ; the female has a horny ovipositor and the second longitu- dinal vein is branched at the root of the wing ; in the second sub-family the proboscis may be formed for sucking, the female has no horny ovipositor, and the second long vein has its first fork near the middle of the wing. The sub-family Phlebotomince contains the genus Phlebotomus, which occurs in South Europe, South Asia, Africa, North and South tropical America. They are all small grey, brown, or dull yellow-coloured flies, and carry their wings when at rest upwards like a butterfly. The proboscis is moderately long and the legs long and thin. The females are most vicious blood-suckers, but in some species anyhow the 582 THE ANIMAL PARASITES OF MAN males also bite {P. duboscii). They are mainly nocturnal feeders and hide away during the day in any dark corners or crevices. The life-cycle has been worked out by Newstead^ and Grassi- in Europe, and by Howlett^ in India. The larv£E have been found in crevices in rocks and caves, in dirty cellars, and dark damp places containing rubbish, and are also said to live in crevices in the walls of privies and cesspits. The minute larva is very marked ; as figured by Newstead it has two long chaetcC projecting upwards, in some stages branched, in others simple, and on the seg- ments a few blunt spine-like processes. The pupae are found in similar situations. The ova are very minute, elongate, translucent white, and covered with a thin coating of viscous matter when first laid ; soon after they become dark brown, shiny, with long black wavy lines. Newstead found the incubation period in Malta to last for about nine days in P.papatacii. Five species are known in Europe, five in Africa,^ two in North America, and eight are described by Annandale"* in the Oriental region. Lutz and Neiva have described three species from Brazil^ {P. longipalpis^ inter- inediiis and sqitamiveniris). Brachycera (Flies). The antenna? as a rule have three segments, and are usually shorter than the head. The first segment of the antennae is frequently very small, and the third one is gener- ally the largest, and sometimes possesses a terminal annulated bristle. The palpi have from one to three segments ; the mandibles are covered by the labium. The three thoracic rings are coalesced ; wings are almost always present, the posterior ones being rudimentary and covered with a little scale. From the ova legless maggots are hatched, which as a rule have not a distinct head, but occasionally possess two claw-like booklets. These maggots live in decomposing organic matter ; they rarely live in water and some of them are parasitic. They either become barrel- shaped pupas within the last larval integument or, after casting it, are trans- formed into naked pupae. The larvae of numerous Brachycera have been observed in man, some in ulcers or on mucous membranes, others in the skin or in the intestine, etc. In many cases the report only mentions the presence of the larvae of flies ; in other cases the species has been determined ; whilst in still other cases the corresponding adult creature is unknown. We must therefore confine ourselves to describing the most common varieties. Family. Phoridae. These flies belong to the same division of Diptera, the Aschiza, as the family Syrphidcp or " hover flies." They are all small insects with marked antennae and wings ; the former have the third segment globular and enlarged, and thus hiding the first two ; the wings are short and broad, the venation shows two short, thick, long veins with four thin ones running out from them. The larvae normally live in decaying animal and vegetable matter, but one species, Aphiochata ferruginea^ Brun., has been found as an intestinal parasite of man. ■ BtiU. Ent. Res., 191 1, ii, pt. 2, pp. 47-78. ^ "Ricerche sui Flebotomi/' Mem. delta Soc. ital. delta Scienze, 1907, ser. 3, xiv. PP- 353-394- » " Indian Sand-flies," Ind. Med. Cong., 1909, sec. Ill, pp. 239-242. * Newstead : Btttt. Ent. Res., 191 2, iii, pp. 361-367. ^ J^ec. Ind. Mus., v, pt. 3, Nos. 13 and 14. " Mem. Inst. Oswatdo Cruz, 191 2, iv, fasc. I, pp. 84-95. V SEPSID^, SYRPHID.E. 583 Aphiochaeta ferruginea, Brun. This small fly belonging to this family is of an orange-ochreous colour, the upper part of the thorax tawny, and with dark bands on the abdomen, legs pale yellow, the hind femora tipped with dark brown. It measures only 2 to 3 mm. in length. This insect is shown by Austen to be widely distributed in the tropics, being found in India, Burma, West Africa, and Central America. The larvae breed in decaying animal matter, such as putrid meat, decomposing shell-fish, etc. Heusner bred out sixty-three flies from larvae taken from an Indian's foot. Baker {Proc. Binnia Brafich Brit. Med. Assoc, 1891, p. 11- 16) found that the maggots of this fly were passed per aiium by a European at intervals during a period of ten months. Baker found that the larvre fed on human faeces; from the egg stage to the deposition of eggs from the resultant brood of flies occupied twenty-two days. He concludes that they are capable of propagating, and do so while living within the human intestines. He also records the larvae in two girls. The larva does not seem to have been described, but Austen describes the pupa {Trans. Soc. Trap. Med. and Hyg.^ iii. No. 5, p. 229). Phora rufipes, Meig. The larvie of the "hump-backed fly" live in rotting potatoes, muslirooms, radishes, etc., and when accidentally introduced into the intestine of man can, like other larvae, live there twenty-four hours and even more, and may set up serious gastric disturbances. P. rufipes is the same as P. pallipes., Latr. Family. Sepsidae. Small blackish flies, elongate, with abdomen narrowed at the base, thickened and curved downwards towards the extremity. Larvae often found in decaying vegetables, ham, cheese, etc. The larvae have the power of skipping ; conical in form, pointed in front, truncated behind, about 5 mm. long, shiny and smooth, the anal segment with fleshy protuberances. The genus Piophila has a short proboscis and the cross-veins of the wings approximate. Piophila casei, L. Cheese flies. The larvae live in ripe cheese, with which they are sometimes introduced into human beings (Meschede). The larvae of the cheese flies {Piophila casei) may pass through the alimentary canal of human beings alive, and have been occasionally referred to in cases of internal myiasis. It also breeds in dead bodies in adipose tissue. Howard records it on human excrement. It is thus possible that some of the recorded cases of thi^ pest being passed alive may be due to eggs deposited on human faeces. Family. Syrphidae (Hover and Drone Flies). Amongst the large family of Syrphidce is found a section known as the Eristalince or drone flies, whose curious long-tailed larvae are popularly called " rat-tail larvae," on account of the end of the body being drawn out into a long telescopic tail of two segments, at the end of which are placed the breathing pores. These larvae 384 THE ANIMAL PARASITES OF MAN live in water, no matter how foul, and in liquid manure. They have occasionally been obtained in foul drinking water by human beings and from eating watercress improperly washed or from badly kept beds. Austen {Trans. Soc. Trop. Med. and Hyg., iii, No. 6, p. 221) records that in the autumn of 1907 a number of the larvae of the common drone fly {Eristalis tenax) were passed per reciiun by a woman in Hampshire who had recently arrived from France. The patient had eaten a con- siderable quantity of watercress before leaving France. I have twice found small Eristalis larva? clinging by their long tails on watercress served at table. Family. Drosophilidae. Small, rather plump flies, with short, broad abdomen, with bristles on the head and legs. Often abundant in decomposing fruit, and may occur in dense masses. Drosophila melanogaster, Br. The larva? of this fly occur in over-ripe fruit and in fungi, often also in human habitations, and live in substances undergoing acid fermentation (vinegar, decaying fungi, rotting fruit, in damaged spots in diseased trees), much more rarely in animal substances, and they occasionally gain access to the human intestine (for example, by the medium of sour milk). When introduced in any quantity, they cause vomiting or attacks resembling colic ; when taken in the pupal stage no unpleasant results are produced. Family. Muscidae. Teichomyza fusca, Macq. Syn. : Scatella in'inaria., Rob. Desv. ; EpJiydra loncripetinis., Meigen. The larvae live in the urine in privies. Several authors state they have found them in fresh fasces or in vomited matter. Pruvot states that they continue for three days in the stomach of rats into which they have been intentionally introduced. (Pruvot, G., " Contrib. k I'etude des larves de dipt, trouv. dans le corps humain," These de Par.., 1882; Chatin, J., in Coinp. rend. Soc. de BioL., Paris, 1888 [8], v, p. 396 ; Roger, H., ibid.., 1851 [i], iii, pp. 88, etc.) Homalomyia canicularis, L., etc. Hoiiialoniyia nianicata, Meig., live as larvc-e in decomposing vegetable matter or in cultivated vegetables (cabbage) ; they are easily recognizable by their plumed bristles, which are situated laterally on the body segments. They obtain access fairly often to the human intestine and give rise to very uncomfortable symptoms. Cases have been recorded from Germany, Austria, France, England, North America (Wacker, in Artzl. Intelligenzbl., 1883, xxx, p. 109; Florentin, in Compt. rend. Soc. de Biol, Paris, 1904, Ivi, p. 525; and other authors). The larvae of an allied genus (Anthomyia), which, however, does not possess plumed bristles, has been found in the external auditory meatus of a man {A.plicvialis, according to Danthon). [//. cafiicularis is common to Europe and North America, and is an;abundant house-fly. It is the small house-fly so often seen on windows. Besides living on vegetable matter, they have also been found in the nests of the humble bee. Larvae of this species (fig. 405) were sent to the British Museum, taken from the f^ces o MUSCID^ 585 a woman suffering from cancer.' They were found at Shrewsbury. Hagen^ reports the larvc^ of this fly as occurring alive in the urethra of a patient. — F. V. T.] Homalomyia scalaris, Fabr. [This is not a synonym of the above, but a distinct species. [//. )iianicata^ Meigen, is also distinct.— P". V. T.] Anthomyia desjardensii, Macq. This fly, allied to Homaloinyia, is the cause of both intestinal and cutaneous myiasis at Bihe, Angola (Wellman, Joiirn. Trap. Med. and Hyg., June, 1907, X, p. 186). Hydrotaea meteorica, L. The larvae live in decaying vegetable substances, also in dung, and have been evacuated in some cases by man (Zetterstedt, Joseph). Fig. 405. — Larva of Homalomyia canictilaris. Enlarged. Fig. 406. — Larvse of Calliphora vojuitoria. Enlarged. Fig. 407. — Larva of Chrysomyia nmcellaria. 4/1. (After Conil.) Cyrtoneura stabulans. Larvae in fungi, but occasionally also on larvae of butterflies and Hynienoptera_ occasionally introduced into the human intestine (Joseph). Musca domestica, L., and M. {Calliphora) vomitoria^ L., and allied species ; larvae of these have been repeatedly found in the intestine and nose of man (Mankiewicz, etc.).* ' Theobald, " First Report Economic Zoology," Brit. 3fus. {Nat. Hist.), p. 55. '^ Hagen, Froc. Bost. Soc, N.H.., xx, p. 107. ^ " Larvae of a Musca, probably M. corvina, were passed in numbers/^;- rectiim by a child in Liverpool with Homalomyia larvre," — "Second Report Economic Zoology," Theobald, 1903, p. 16. 586 THE ANIMAL PARASITES OF MAN Musca domestica, Linn. (Common House-fly) It is not only on account of a few larvae of the common house-fly {Musca domestica) being found in the intestines of man that it is of importance medically. It is far more important on account of the part it plays in the spread of diseases of the intestines, such as typhoid fever and cholera, infantile diarrhoea and dysentery. Howard and Clark {Journ. Exp. Med., 1912, xvi. No. 6, pp. 850-859) have shown that the house-fly is capable of carrying the virus of poliomyelitis for several days on the surface of the body and for several hours in the gastro-intestinal tract. The house-fly may also distribute the ova of Tccfiia solium and the white worms (Oxyuris and Ascaris). It has also been proved that they may carry the germs of tuberculosis, and it is said that they play an important part in the spread of infectious ophthalmia in Egypt. This insect is found in all parts of the world. In warm countries it breeds all the year round, and it may do so even in temperate climates in warm places, such as stove houses. Most, however, die off in the autumn ; but some survive the winter as adults, in such places as kitchens, restaurants, and warm houses. I have never failed to find a few Musca domestica in houses during the winter. The majority, however, hibernate as puparia. The females deposit from 120 to 150 eggs in a batch in stable manure, rotting vegetation, house refuse, spent hops, old soiled bedding, etc. A single female may lay as many as six batches of ova during her life. The eggs are shiny white, and hatch in from eight to twenty-four hours in warm weather to three or four days in cool weather. The while footless maggots are cylindrical, tapering to a point at the head end, truncated posteriorly. The head consists of two dark mandibular hooks and two short antennse. ,On the tail end are two plates, the stigmata, in which the main tracheal trunks open ; in the second segment are a small pair of projecting stigmata. The larval stage lasts from seven to five days in hot weather ; but in cold weather in temperate climes it may last six or eight weeks. The larva on reaching maturity becomes a barrel-shaped puparium of a dark brown to black colour, and in this case changes to the pupa. This stage lasts from three days in the tropics to four or five weeks in cold weather, the life- cycle thus varying from ten days in the tropics io fourteen in warm weather in Europe up to three or four months under unfavourable conditions. All breeding grounds should be burnt or otherwise done away with, such as stable manure, house and kitchen refuse, human excrement and soiled substances, also decaying vegetation as soon as possible, certainly by every sixth day. Stable manure should be kept in closed receptacles and should be removed by every sixth day to at least one mile from habitations and sprinkled with chloride of lime. All kitchen and household refuse should be burnt at once or buried in pits and covered with soil. Latrines should be as far as possible from hospitals, mess rooms and tents. Food — especially milk, sugar and fruit — should be kept screened with muslin when house-flies are about. Mess rooms and tents and hospitals should have doors and windows screened with fine wire gauze during the fly season. All possible steps should be taken to prevent them contaminating man's food and from breeding in human excrement and from entering hospitals. When present in dwelling-houses in numbers they may be killed by fumigation with pyrethrum or sulphur. CHRYSOMYIA 587 Genus. Chrysomyia, Rob. Desv. Chrysomyla (Compsomyla) macellaria, Fabr. ; Lucilia macellaria, Fabr. Syn. : Lucilia /lofninivorax, Coq. ; Calliphora itifesta, Phil. ; Calliphora anthropophaga, Conil. A species distributed from the Argentine to the south of the United States which deposits its ova on ulcers, in the aural meatus or in the nasal cavities of persons who sleep in the open air. The larvae are yellowish white, 16 mm. long, are armed with two strong mouth hooks, and provided with spinous rhigs (screw-worm) ; they lie hid in the nasal and frontal sinuses, in the pharynx, larynx, etc. ; they per- forate the mucOus membranes, even cartilage, migrate into the eyes, the cranial cavity, middle ear, and cause severe disturbances ; after the mature stage, in which the larvae leave the host to enter the pupal state, these symptoms often spontaneously Fig. 408. — The screw-worm fly {Chrysomyia niacellaria), abate after a lapse of eight days, leaving behind greater or less cicatrices, and con-^ sequently also defects in function of the organs attacked. Very often, however,, sepsis sets in, usually with a fatal termination. (Coquerel in: Arch. gen. de med., 1858 (5), p. 513; 1859, xiii, p. 685; Ann. Soc. enl. France^ 1858 (3), vi, p. 171 ; 1859, vii, p. 234. Weber in : Rec. de mem. de mtd. milit., 1867 (3), xviii, p. 159. Franclus, A., in: Arch. f. path. Anat., 1868, xliii, p. 98. Conil in : Bol. Acad. nac. cienc. Cordoba^ i88r, iii, p. 296. Humbert, Fr., in : Proc. U.S. Nat.- Mns.,. 1883, vi, p. 103; Amer. Nat., 1884, xviii, p. 540. Lindsay in : Journ. Trap. Afed., 1902; v, p. 220, and other authors.) [This species is known as the screw-worm fly. It attacks animals as well as. man, especially laying its eggs on wounds formed by barbed wire. It may also be found on dead flesh. Dr. St. George Gray sent me speciniens from St. Lucia, from 37 588 THE ANIMAL PARASITES OF MAN the nose and mouth of a patient in Victoria Hospital. Others were found in the vagina of another patient. Out of the four patients attacked, two occupied the same bed, one after the other, and a third the next bed to it. The other case was in a more remote part of the hospital. There are numerous records of this fly attacking man. It occurs from the Argentine to Texas. — F. V. T.] Chrysomyia viridula, Rob. Desv. [This species is somewhat larger than the former ; the body is metallic bluish- green, the dorsum of the thorax with three blackish, longitudinal stripes, and the face ochraceous ; about lo mm. long. Austen records this species from man, Dr. Daniels having bred it from larvae from a sore on a human being in New Amsterdam, British Guiana. Dr. Laurence also bred it in Trinidad. In the latter case between loo and 150 maggots were discharged from the nose of a woman suftering from facial myiasis (^Brit, Med. Joicrn., January 9, 1909, p. 88 + fig.). — F. V. T.] Genus. Lucilia, Rob. Desv. Lucilla nobilis, Meig. The larvae were observed by Meinert in Copenhagen in the auditory meatus of a person who, after taking a bath, fell asleep in the open air, and on waking felt singing in the ears, and had a sensation as if there were water in the auditory canal. During the next days severe pains set in, and there was a discharge of blood and pus from both ears, as well as from the nose. On washing out the meatus the maggots made their appearance. Lucilia ccesar and Z. sericata have also been observed in the larval state in man (Thompson, Hope, Henneberg and Calendoli, Napoli, 1907). [This golden-green fly usually lays its eggs on decomposing organic matter ; now and again it lays its eggs in wounds on man. — F. V. T.] Genus. Pycnosoma, Brauer and v. Bergenstamm. The species of this genus have a general resemblance to the Lucilias and Chrysomyias, but the body is stouter and the abdomen banded. The genus can be distinguished from Chrysomyia by the absence of the three thoracic stripes and by the eyes of the male, in which the facets forming the upper portion are much enlarged, whereas in Chrysomyia they are not noticeably larger. Austen also points out that the sterno-pleural bristles in Pycnosoma are i : i, in Chrysomyia 2:1. The genus is found in tropical Asia and Africa only. All records of Chrysomyia (Compsomyia) in India must be referred to this genus. Bezzi and Stein (" Katalog der Palaarktischen Dipteren," 1907, iii, p. 543), however, regard the two as synonymous. The larvae are frequently found in the nostrils of man and burrow into the sinus, but normally they live on decaying animal matter. Pycnosoma forms the so-called Indian screw-worm. Patterson {Ind. Med. Gaz.^ October, 1909, xliv, No. 10) records the case of a woman at Tezpin, Assam, from whom as many as 100 larvae were removed at one time, and later the left orbital cavity was found packed with hundreds of maggots; eventually the patient died. It is possible that this, however, was due to a species of Sarcophaga. Austen undoubtedly records this genus causing nasal myiasis in India {Trans. Soc. Trop. Med. and Hyg., iii, p. 235). At Dehra Doon, U.P., a woman discharged 100 larvae from her nose, with great pain in the nasal region and frontal sinuses. The so-called " peenash," a common malady in Rajputana, is a true nasal myiasis. SARCOPHAGA 589 Genus. Sarcophaga, Mg. Sarcophaga carnosa, L., 1758. Larvae of flesh-flies provided with two claws at the anterior end, which settle on raw or cooked meat, and in the open on carcases of animals ; they are often observed in man, both in the intestine (introduced with food) and in the nasal cavities, frontal sinus, conjunctiva, aural meatus, anus, vulva, vagina, prepuce, and open ulcers, often migrating further from the regions first attacked. (Gayot in Compt. rend. Acad. Set., Paris, 1838, vii, p. 125. Grube in Arch. f. Naiurg., 1853, xix, I, p. 282. Legrand du Saulle in Compt. rend. Acad. Set., Paris, 1857, xlv, p. 600, and other authors.) [This fly is viviparous. The fly varies from 10 to 30 mm. in length, and is of a general ash-grey colour ; the thorax with three dark stripes, the abdomen light grey with three black spots on each segment ; legs black ; base of wing.s yellow. It also attacks animals and birds, especially geese. The genus Sarcophaga is universally distributed. The maggots are whitish or yellowish footless larvas of twelve segments, tapering to a point in front, broadened posteriorly. There are two mouth liooks, by means of which they rasp their food. The breathing pores are at the end and consist of two groups of three slits, each surrounded by a hardened area. They pupate in their old skin, which turns brown. — F. V. T.] Sarcophaga magnifica, Schiner, 1862.^ Syn. : Sarcophaga wohlfahrti., Portschinsky, 1875. A species widely distributed over the whole of Europe, occurring especially in Russia (Mohilew) ; the presence of the larvse in man was first observed by Wohlfahrt (1768). The larvae settle in the pharynx, in the nose, the aural meatus, the conjunctiva, and in other regions of the human body ; they also attack domestic animals and birds. As Portschinsky has shown, they cause severe inflammations, haemorrhages and suppurations in the organs in which they occur ; children are especially attacked. A number of cases have been observed also in Central and Western Europe. [The fly has a light grey abdomen with shiny black spots which do not change their shape and appearance according to the angle in which the fly is viewed. — F. V. T.] (Wohlfahrt : " Observ. de vermibus per nares excretis," Halae, 1768 ; Nov. Act. Acad. Caes. Nat. curios.., 1770, iv, p. 277. Gerstacker in: Sitzicngsb. Ges. nat. Frde. Berl., 1875, p. 108. Portschinsky in: Horce soc. efttom. ross., 1875, 1884, p. 123. Laboulbene in : Ann. Soc. ent. France, 1883 (6), iii ; Bull., p. xcii. Leon in : Bull. Soc. des MM. et Nat. de /assy, 1905, xix, p. i. Freund, L., in : Verh. Ges. deutsch. Naturf. u. Arzte, Homburg [1901], 1902, ii, 2, p. 450, and other authors.) [Probably most cases of attack in Europe are due to this species. — F. V. T.] The above cited do not exhaust the number of observations of diptera larv;e parasitic in man ; there are yet to be mentioned the larvae of .S". hcemorrhoidalis , S. hcEinatodes (of G. Joseph), those of S. ruficornis (excitants of a cutaneous myiasis in the East Indies), those of species of Eristalis (of Hanby and others), and those of Fhora rufipes (of Kahl, of Warsaw, and others). In many cases the determination of the diptera laivaj has been omitted (or must be omitted) ; such is the case with diptera larvae in the eye (Schultz-Zehden in : Berl. klin. Wochenschr., 1906, p. 286. OUendorf in : Med. Korrespo7idenzbl. d. wiirt. drtzl. Landesver., 1904, p. 1017. Kayser in : Kli7t. Monatsbl. f. Augenheilkunde, 1905, xliii, i, p. 205. Ewetzky and V. Kennel in : Zeitschr.f. Augenheilktuide, 1904, xii, p. 337, and other cases). Austen ' [The correct name for this fly is Wohlfahrtia magnifica, Schiner. — F. V. T.] 590 THE ANIMAL PARASITES OF MAN records several cases of myiasis due to Sarcophaga {vide Trajis. Soc. Trap. Med. and Hyg., 1910, iii, No. 6). The larvae of African Muse Ida' have now become of greatei- interest; Hke several Oestrid larvae they live normally in the skin of mammals, but also attack man. The knowledge of these species is certainly very insufficient, but this is not likely to be the case much longer, as medical men practising in the Colonies are giving their attention to these parasites. At the present time four distinct forms are recognized according to Gedoelst.^ Sarcophaga chrysdtoma, Wied. [This species is recorded as attacking human beings at New Amsterdam, Britisli Guiana. The fly is 15 mm. long, has a golden-coloured face, three broad black thoracic stripes and ochraceous buflf anal segments. It was bred from larvife obtained by Dr. Roland from a sore on a girl's foot. It is known to occur in the Brazils and the West Indies. Another species was also bred which Austen was unable to identify. — F. V. T,] Sarcophaga plinthopyga, Wied. [This and other species of Sarcophaga are called "yaw flies" in Dominica, as they are believed to be concerned in the dissemination of frambcesia or yaws (Nicholls) {vide Austen, Trans. Soc. Trop. Med. and Hyg.^ 1910, iii, p. 239).— F. V. T.] Ochronnyla anthropophaga, E. Blanch. ; Cordylobia arthrophaga, Griinberg. Indigenous to the Senegal and neighbouring districts ; in the district of Cayor (between the mouth of the Senegal and Cape Verde) the larva is known as the " ver de Cayor."' It lives under the skin, especially at the lower extremities and the lower region of the trunk, producing small boils, which cause pain, but after about eight days, when the larva leaves the body to enter the pupal stage, the pain discontinues. Besides man the larva occurs in dogs, goats, cats, and in the jackal. It is still questionable whether the fly deposits its eggs direct or on the ground, from whence the larvae as they emerge gain access to animals and man. Larvae yellowish-white, 14 mm. long, 4 mm. wide, eleven segments^ ; head with two globular antennas-like appendages, two black curved mouth hooks, and two wart- shaped, finely spinous structures at their base. Body evenly covered to the seventh segment with small black prickles, which are stronger at the sides and the anterior borders of the segments ; from the seventh they increase in size, on the two hindermost they are wanting ; on the last segment two deep yellow spiracles, each with three markedly curved fissures ; in ' [The following are known to cause myiasis in man in Africa : Cordylobia anthropophaga, Griinb. ; Aiichmeromyia liiteola, Fabr. ;" A. rodhani, Gedoelst ; Oestrus ovis, Linn. ; and Anthomyia desjardensii, Macq. The anthropophaga, Blanchard, and the depressa, Walker, referred to here are Griinberg's anthropophaga. — F. V. T.] 2 [Austen gives the length as 12 to 12-5 mm. and the breadth as 5 mm. ; he describes the larva as follows : Bluntly pointed at the anterior extremity, and truncate behind ; from third Fig. 409. — Ochromyia larva on the skin of man, South Africa. 3/1. (After Blanchard.) CORDYLOBIA 59 1 addition two stigmata on the posterior border of the first segment. Duration of the larval stage about eight days. Upon the construction of roads in Guinea the larva is spread by dogs far into the interior. Auchmeromyla (Bengalia) depressa (Walker).^ Distributed in the region of Natal and apparently over the whole of South Africa. The " larva of Natal," as one may still term the species provisionally, as its identity is not certain, possesses on its head (besides the mouth hooks) lateral protuberances beset with a row of chitinous spines. The cuticle of the body is spinose. The spines are difficult to recognize on account of their transparency and want of colour; they are longest over the anterior segments, from the fifth they become smaller, and over the hindermost they are very small. Apart from the foremost segment, the position they take is that of rows running transversely or obliquely, two to four generally in juxtaposition ; the number of spines in the groups gradually increases posteriorly, attaining the number of eight to twelve on the sixth segment, and this number is maintained to the end of the body. Isolated spines are found over the head ; over the second, third and fourth segments single ones are still found adjoining the groups of -n^ tj~~^ j c ui r -nt . . r \ r ° ^ Fig. 410. —Head end of " larva of Natal, spmes, from the filth onward they Magnified. (After Gedoelst.) are wantmg. From here the spines cover the whole free surface of the segments ; over the fourth the anterior three- quarters, over the third two-thirds and over the first and second only the anterior half. The stigmata found at the anterior end also serve as distinguishing characters. The parasitic stage appears to last about fourteen days. [Fuller {Agric. Joiirn.^ Dept. Agric. and Mines, Natal, 1901, iv, p. 606) refers to this as Bengalia depressa also. — F. V. T.] Genus. Cordylobia, Griinberg, 1903. Cordylobia grlinbergi, Donitz. Syn. : Ochroniyia anthropophaga^ Griinberg, 7iec Blanch. ; Cordylobia ajithropophaga^ Griinberg. Endemic in German East Africa and neighbouring regions. Larva up to 14 mm. long, 4 to 5-5 mm. wide, of cylindrical shape, slightly narrowed behind, truncated, gradually tapering in front ; antenna-like processes, cone-shaped, blunt. to eleventh segments thickly covered with minute recurved spines of brownish chitin, usually arranged in transverse series of groups of two or more, which can be seen to form more or less distinct undulating and irregular transverse rows. In each of the two posterior sligmatic plates, the respiratory slit on either side of the median one is characteristically curved, resembling an inverted note of interrogation. The barrel-shaped puparium is on an average iO'3 by 4*6 mm. ; its colour varies from ferruginous to nearly black. — F. V. T.] ' [According to Austen this is Cordylobia anthropophaga^ Griinb. Bengalia depressa^ Walker, is a very different insect, whose life-history is unknown. — F. V. T.] ^g2 THE ANIMAL PARASITES OF MAN Smaller cylindrical formations at the base of the mouth hooks surrounded by a circle of chitinous hooks. Body from the first segment covered with small brown squamous spines which are disposed in numerous irregular transverse rows. The spines are small over the two first segments, the two posterior thirds of all the segments, as well as from the eighth ; over the third to the seventh they are larger, but between these there are very small spines. The breathing pores of the stigmata at the anterior end are kidney-shaped ; the orifices are elongated and very tortuous, each divided into three. The larval period appears to last several weeks. Cordylobia anthropophaga, Grunberg. This well-known cutaneous African parasite seems to have been the cause of much confusion in regard to names. It belongs to the genus Cordylobia of Grunberg, and is one of the family Muscidcp, and differs from Auchmeromyia in that the second abdominal segment of the female is of normal size, whilst in Auchmeromyia it is more than half the length of the whole abdomen, and in the male the eyes are holoptic or close together, whilst in Auchmeromyia they are wide apart. The flies of this genus (three so far described) attack man in their larval stage (anyway two of the three), and also dogs and other animals, by burrowing into the skm and producing painful boils. \C. anthropophaga, Griinberg, is widely distributed in Africa, extending from Senegal, where its maggot is known as the "ver de Cayor," and is referred to on p. 590 as Ochromyia anthropophaga, E. Blanchard, to Natal, where it is known as the " Natal worm," and referred to erroneously on p. 591 as Bengalia depressa, Walker. [It is a thick-set Muscid of a general straw-yellow colour, with blackish markings on the dorsum of both thorax and abdomen, about 9*5 mm. long. The larva is fat and when mature about 12 mm. long, bluntly pointed in front, truncate behind ; from the third to eleventh segments it is thickly covered with minute recurved spines of a brownish colour, arranged in transverse series of groups of two or more, which form more or less distinct irregular transverse rows. On each of the two posterior stigmatic plates, the respiratory slit on either side of the median one is characteristically curved, resembling an inverted note of interrogation. The puparium is brown to ferruginous or black and about 10 mm. long. The maggots are found in both natives and white men, and occur as a severe pest in dogs, also in monkeys, rats, and other mammals. In Sierra Leone it is called the " tumba fly." The larvae have been frequently found as true subcutaneous parasites, each larva living singly and forming a boil or warble in the skin, with an opening just as in an ox-warble, through which the maggot breathes and eventually escapes. Although they more usually occur as isolated specimens, Marshall found in Salisbury, South Rhodesia, that sixty were extracted from one lady, and Berenger-Feraud, in Senegal, that more than 300 occurred in a single spaniel puppy. [Neave {Bull. Ent. Res., 1912, iii, p. 217) records it from ulcers in a native at Lourengo Marques in 1908, and at the same time from ulcers in a dog, and that it is a severe pest to man in Mozambique and parts of the Transvaal. It seems to be more abundant in North Rhodesia and Nyasaland than to the north (Neave, Bull. Ent. Res., 191 2, iii, p. 310). It is also recorded in Zanzibar, German East Africa, Uganda, East Tropical Africa (Neave). [Simpson {Bull. Ent. Res., iii, p. 170) records a Muscid larva taken from the breast cf a European in South Nigeria that was probably Cordylobia. [It is not known how infection takes place. Neave {Bull. Ent. Res., iii, p. 310) says : " Many instances in human beings would preclude the possibility of eggs LUND S LARVA, AUCHMEROMYIA LUTEOLA 593 having been laid direct on the skin : in these cases they have probably been laid on the clothing put out to dry. [Gedoelst has described another species, C. rodhani^ and Austen a third species, C. prcr^ra7idis^ from Nyasaland, Cape Colony, Transvaal, Natal, North-west Rhodesia, and German East Africa. [The following are some papers dealing with this subject : Proc. Ent. Soc, London, for year 1907, p. xlvii ; Jomit. R.A.M.C., 1908, pp. 5-1 1, figs, i and 2, by Austen; Journ. R.A.M.C.^ 1908, pp. i and 2, by Major F. Smith ; Trans. Soc. Trop. Med. and Hyg.., 1910, iii, pp. 223-225, by Austen. — F. V. T.] Lund's Larva. Endemic in the region of the Congo State ; called after Commander Lund, from the skin of whose arm it was extracted ; 12*5 mm. long, 4*5 mm. broad ; colour yellowish, with brown rings, on account of the division of the brown spines ; head cone-shaped, with two hemispherical smooth antennae, two thick black mouth hooks and wart-shaped bodies, between which are situate two to three longitudinal rows of dark brown chitinous laminas. The body segments are covered over their whole Fig. 411. — Lund's larva : on the left, the whole larva, magnified six times. On the right, the head end, much enlarged. (After Gedoelst.) surface with irregularly distributed triangular yellow spines, the points of which are coloured dark brown. Its size increases from, the second to the sixth segment, diminishes from the seventh to the ninth, at the tenth it is reduced, and at the eleventh quite small. The posterior stigmata are bean-shaped, each with three markedly tortuous openings. Duration of the larval stage unknown ; the same applies to the pupal and imago stages. Auchmeromyia luteola, Fabricius. [This fly, the parent of the so-called Congo floor maggot,^ belongs to a nearly allied Muscid genus to Cordylobia, but which can at once be told by the great length of the second abdominal segment. The maggot occurs in numbers in the ' Dutton, Todd and Christy, "The Congo Floor Maggot," Metn. xiii Liv. Sch. Trop. Med., p. 40. 5Q4 THE ANIMAL PARASITES OF MAN native huts in the Congo region and is fairly common in central and northern parts of Mozambique ; it is also recorded from the Zambesi River and the vicinity of Barberton in the Eastern Transvaal {Bull. Ent. Res., 191 2, iii, p. 216), in German East Africa, in Nyasaland, and British East Africa. It is also recorded from Bara, Kordofan,^ where they occurred on the floor of the men's prison and bit the prisoners. They were destroyed by sprinkling Jeyes' fluid on the floor. Neave states {ibid., p. 310) that it occurs in the more neglected huts in native villages throughout tropical Africa, and frequently enters a tent when pitched near a village. It is also found in West Africa. The fly is thick-set and about the size and build of a bluebottle fly; length 10 to 12 mm.; tawny in colour to dirty yellowish-brown, with dusky hairs, giving it a smoky appearance ; the flattened thorax has long dark stripes and the abdomen a dusky line in the centre of the second segment, which meets a dark line on its posterior border ; the dusky third segment has a narrow yellowish anterior line; the fourth segment is also dusky; legs bufl"wiih black hairs ; the fifth tarsal segment black. The larviie are whitish, becoming reddish after a feast of blood, with much wrinkled skin and rather flat and broad. They live in crevices of the mud floor, under sleeping mats during the daytime, and come out at night and suck the blood of sleepers and then retire to shelter again. Dutton, Todd, and Christy noticed that where people slept on beds or platforms raised above the floor the maggots were not so numerous as under the sleeping mats laid on the ground. They turned up many of the maggots from a depth of three inches or more.' — F. V. T.] Family. Oestridae. [The family of Oestridcp or warble flies are all parasitic in their larval stage, usually termed the "bot" stage. They are found as parasites in warm-blooded animals, and man is frequently attacked by them. The members of this family have the mouth rudimentary, many of them are hairy and bee-like, with large eyes and the head large, the lower part more or less swollen. The thorax is large with a distinct transverse suture, and the abdomen short and stumpy or very slightly elongated. The male genitalia are hidden, whilst the female ovipositor is often elongated. The wings may be transparent (Hypoderma) or mottled (Gastrophilus), and have muscid- like venation ; the tegulas usually large, the legs moderately long. [As a rule each species is confined to a particular host, but as we see recorded here those that attack animals may also attack man. The flies occur in warm weather and usually during the warmest part of the day, and have a strong dislike to shade and water. The genus Hypoderma attack oxen, sheep, goats, antelope and musk deer; Oestrus, sheep, antelope and horses; Gastrophilus, the horse and ass; Cephenomyia, the deer ; Cepholomyia, the camel and bufifalo ; Dermatobia, dogs, cats, oxen, deer, apes and man ; Cuterebra and Rogenhofera, rodents and opossums- [Some live as parasites in the stomach and intestines (Gastrophilus) ; others infest the skin (Hypoderma, Dermatobia and Oestromyia, the latter on Lagomys and Hypodaus) ; (Edemagena tarandi also infests the skin of the reindeer in Siberia and boreal America. Oestrus lives in the nasal sinus, and Cephalomyia in the throat as well, Cuterebra and Rogenhofera, the skin or scrotum, so that we have really three groups of parasitic oestride larvae : (i) cutaneous, (ii) intestinal, and (iii) facial. [No species seems confined to man, but the so-called "creeping disease," caused by Hypodermae, and the attack of sheep nasal fly are comparatively common, as also is the Dermatobia attack.— F. V. T.] . ' Balfour, Journ. Trop. Med., 1909, xii, No. 4, p. 47. 2 Journ. Trop. Med., 1905, viii, No. 6, p. 90. cutaneous oestrid^: 595 Cutaneous Oestrid^. The eggs are deposited on the surface of the body ; the larvns burrow in the skin, which they reach after somewhat long peregrination. Genus. Hypoderma, Latreille, Hypoderma bovis, de Geer. The cattle fly or warble fly, which swarms during the hot season, settles on the head or on the hair of grazing cattle ; through the young being licked off they gain access to the mouth and are swallowed.^ The larvae appear first in the commencing portion of the stomach, to escape, as some state, into the preceding sections of the alimentary canal ; at any rate, they are found from July onward regularly in the submucous tissue of the pharynx, in which they travel about for several months (up to November, and in isolated cases up to February) ; they then penetrate the muscularis and migrate by way of the subserosa along the mediastinum, the crura of the diaphragm, the renal capsules, and the intermuscular connective tissue of the psoas muscle in the direction of the spinal canal, into which they penetrate by way of the muscles and nerves, through the intervertebral foramina. Here they stay for about two to three months, then they leave the spinal canal again through the vertebral foramina and make their way (from January to March) through the inter- muscular connective tissue of the muscles of the back to the skin of the back, where sooner or later (from January to June) they arrive and enter a resting stage, which commences with penetration of the skin and terminates with outward migration from the boils due to the wound set up by the maggot. At the commencement of this period the larvae cast their skin, and their form, hitherto cylindrical, becomes oval. After about a month, a second moulting of the skin takes place — the third larval stage, which lasts about two and a half months (up to June). The approaching end of the same is indicated by a change of colour on the part of the larva from the hitherto yellowish-white to brown and finally to blackish-brown. When they have become mature the larvre leave the warbles, drop on to the ground and pass into the pupal stage in the superficial layers of the soil within twelve to thirty-six hours. After about a month the flies emerge. Irregularities with regard to the time and direction of the migrations of the larvss take place (Jost, H., in Zeitschr. f. wiss. ZooL, 1907, xxxvi, p. 644). In a number of cases the larva of the cattle fly has been observed in the human integument, usually in the winter months, that is, during the migration period ; consequently, it is not surprising that the larvae before they enter on the resting stage and produce a warble undergo migrations. But that this takes place subcutaneously — w^hich does not appear to be so in the case of cattle — is perhaps explained by the fact that in man, on account of the short space that has to be traversed, the larvae are not sufficiently developed to enter on the resting stage simultaneously upon having obtained access to the integument. Whether the Oestrid larvae in Bulgaria that similarly migrate beneath the skin in man belong to the cattle fly or to another species, or even another genus, has not yet been 1 [This is not the case, for Carpenter has shown that muzzled calves become infected ("Mem. First Int. Cong. Ent.," pp. 289-293). ]o%\.{Zeiischr. f. wiss. ZooL, 1907, xxxvi, pp. 644-715) thinks that the ova, not young larva;, are ingested (vide note in Supplement. — F. V. T.J- 596 THE ANIMAL PARASITES OF MAN ascertained. (Doctorow, in Arch, de Par., 1906, x, p. 309 ; Spring, A., in Bull. Acad. set. Belg., 1861 (2), iv, p. 172; Walker, R., in Brit. Med. yoiirn., 1870, i, p. 151 ; Kjelgaard, in IJgeskr. f. Laeger, 1904, p. 535 ; Condorelli, M., in Bull. Soc. Zool. Italy 1904, xiii, p. 171.) Hypoderma lineata, de Villers. The larvie of this species, that occurs not only in Europe but in North America, live under similar conditions in the skin, very rarely in man; also migrating subcutaneously (Topsent in Arch, de Par., 1901, iv, p. 609). [In Sweden, the ox warble fly {H. bovis) is well known to attack man. Schoyen states " that over 100 years ago up to the present time cases of travelling grubs under the human skin in some districts of Sweden were well known." The species appeared to be H. bovis, many of which he had examined. They accomplished long ramblings under the skin, always in an upward direction, previous to their appearance through an opening in a tumour on the upper part of the body, on the head, neck, or shoulders. An interesting case is recorded in Insect Life., ii, pp. 238-239. A bot similar to H. diafia was taken from the eye and cheek of a child at Kane, McKean County, Pa., U.S.A. It was said to have travelled in five months from the elbow to the eye. Riley later {Insect Life, iv, p. 31c) was inclined to think the maggot was that of H. lineata, the common American ox warble, which is also found in Europe in great numbers. I have recorded another case in England {Rept. Econ. Zool. for year ending September 30, 1910, p. 128), where Dr. Menzies removed the larva of H. bovis from the upper eyelid of a patient. It caused considerable swelling of the face, much pain and distress ; but the case did well, and the wound healed at once. The larva was nearly mature. Numerous other references to this so-called creeping disease will be found in the Supplement. [It is quite probable that bovis and lineata are confused in the latter accounts. The larvae are, however, easily distinguished if carefully examined. — F. V. T.] Hypoderma diana, Brauer. In its larval stage it lives like other species of Hypoderma, attacking the red deer {Cervas elaphas) and roe deer {Cervas capreolus) ; it is occasionally also found in man (Joseph, in "Myiasis externa dermatosa," Hamburg, 1800; Volkel, in Berl. klin. IVochenschr., 1883, ^x, p. 209). Genus. Dermatobia, Brauer. Dernnatobia cyan iventris, Macq. Syn. : Dermatcbia noxialis, J. Goudot. The genus Dermatobia represents the subcutaneous Oestridce of Europe in warmer parts of America. Both domesticated and wild mammals are attacked, according to one statement birds also {Ramphastus), and man with fair frequency.^ It is assumed that in all cases one and the same species is concerned, for which recently a name originating from C. Linne, jun. {Oestrus ho?ninis), has been employed. Three larval stages are recognized in the skin ; the two first appear to Duprey advances the opinion that Dermatobia deposits its eggs not only on the skin of man and animals, but also on the leaves and twigs in the bush, where, too, young larvie have been met with which gain access from hence to men and animals {Jonrn. Trop. Med. and Hyg., 1906). DERMATOBIA CYAXIVENTRIS 597 resemble one another in the club-shaped or tadpole-like appearance (called macaque in Cayenne, mayacuil [mayoquil] in Mexico), the third is swollen spindle-shaped (Berne, called torcel). Segments 2 to 4 in the club-shaped larvae are closely beset with small black spines, segments 5 to 7 bear at the anterior border a complete ring of strong black hooks, segments 4 to 6 a similar ring, which, however, is interrupted at the ventral surface. The four last segments forming the tail are smooth, only at the posterior end are there small spines. The arrangement of spines of the third stage Fig. 412. — Dervialobia noxialis, Goudot. Fig. 413. — Larva of Der- viatobia cyaniventris in its natural size and magnified. (After Blanchard.) Fig. 414. — Larva of Der- matohia cyaniventris. Enlarged' (After Blanchard.) differs from this. Italian workmen that have been employed in Brazil show the presence of Dermatobia larvae on their return (Blanchard, in Bull. Soc. Ent. France, 1893, P- 24; Bull. Soc. centr. de Med. vet., 1896; Ann. Soc. Ent. Fra?tce, 1894, Ixiii, p. 142 ; Ward, H. B., in Mark Annivers. Vol., Article 25, p. 483, New York, 1903). l^Dennatobia cya7iiventris, Macquart, 1843, is said not to be the same as noxialis {vide Brauer, " Mono. Oestriden," 1863, p. 266). It is known by various other names, 598 THE ANIMAL PARASITES OF JMAN as niiche or giisano in New Granada, the ura in Brazil, and the macaw fly in Cayenne. It occurs in Central and South America and the West Indies. According to Goudot the fly is found in great numbers on the borders of large woods and lands covered with underwood. [It is seldom that more than one larva is found in each individual. It is generally found in the arm and leg, but now and then the face. The perfect insect has never been bred from a larva removed from a human being, so that there is still uncertainty as to the actual species. D. cyanive?itris is 11 to 12 mm. long, has an ochraceous buff-coloured face, dark grey thorax, metallic dark blue to purple abdomen, and brownish wings. D. noxialis is somewhat larger. [In the Journal of Tropical Medicine and Hygietie, January 15, 1905, viii, p. 23, reference is made to this Oestrid in Trinidad, where it is called the "mosquito worm." One case here recorded showed no fewer than four worms on the chin and one on the hand. It is here stated that the fly never attacks man or animals directly, as it is said to do by Scheube, but that the eggs are deposited on leaves and branches in wooded lands and forests, and thus man, hunting dogs and wild animals in passing through get the larvae deposited on them accidentally. The affection is common in Trinidad. Mention is made that a little i in 40 carbolic lotion syringed into the aperture in the skin over the worm quickly killed it. [The cattle worm, or founzaia ngombe, is the name given to a larva which develops beneath the skin of oxen and men in Central Africa, especially amongst the natives and stock of Unyamonezi. According to P. Dutrieux, the egg is laid by a large fly that accompanies cattle. It is unknown between the central plateau or the Ugogo and the East Coast.— F. V. T.] Cavicolous Oestrid^. The forms belonging to this group inhabit as larvae the nasal and frontal sinuses of ruminants, Eqnidce and Prohoscidcc, which they leave for the pupal stage. The larva of — Genus. Oestrus, Linnaeus, Oestrus (Cephalomyia) ovis, L., occurring in sheep, has also been observed in man in six cases in the nose and larynx (Saitta in Gaz. d. Osp. d. Clinic, 1903, No. 128). So far as is known, the eggs are deposited in the nasal cavity. {^Oestrus ovis frequently occurs in man. MM. Sergent (^«;/. de VInst. Pasteur, 1907, pp. 392-399) mention that they lay their ova on the noses, eyes and mouth of humans in Algeria whilst flying, but that they disappear after three to ten days or the inflammation produced by them. Portschinsky {Mem. Bur.Ent. Sci. Com. Cent. Bd. Land Adm. and Agric, 1913, x. No. 3, p. 63) also gives cases. He doubts that ova are laid on the nose ; evidently the Russian habit is anomalous, for the Sergents, Collings and myself find ova laid as a common occurrence. I have often seen them on the nose of sheep. This fly also occurs in the Argentine (Serres, in Gaceta Rural, April, 1913, vi, pp. 759-761). [The tamne or thimni of the Kabyles, a human myiasis of the Tuareg mountains in the Sahara, is caused by Oestrus ovis. Here the larvcE are said to be ejected on to the conjunctival and nasal mucous membrane of humans. [Ed. and Lt. Sergent [Bull. Soc. Path, e.xot., 191 3, vi, No. 7, pp. 487-488) report their attack from the Ahaggar mountains, in Central Sahara. The Tuareg name for .the fly, tamne, is the Targui form of the word thimni used by the Kabyles.— F. V. T.] GASTRICOLOUS OESTRID^ 599 GASTRICOLOUS OESTRID^. The eggs are deposited on the hairs of Eqiiidcv, and the larvae escaping from them are Hcked up and swallowed. They pass their larval stage, according to the species, in various parts of the intestine and stomach, and when mature, pass out per aiiiim in order to undergo the pupal stage. Genus. Gastrophilus, Leach. One of the most frequent species is Gastrophilus eqiii^ Fabr. ; the eggs are laid on the hairs ; the larvae live some ten months in the stomach, living attached to the inner surface. The eggs of G. hceniorrhoidalis, L., are deposited on the lips or the long hairs on them. The larvae adhere to the cardiac end of the stomach, to the stomach itself, and finally to the terminal portion of the intestine. Here, however, and elsewhere in the intestine, the larva: of G. pecoruni', Fabr., are also niet with, whilst the larva: of G. nasalis (so called because the eggs are deposited in the nasal orifices) almost exclusively inhabit the anterior section of the duodenum. Cholodkowsky attributes the " wormlet " observed by Samson and Sokolew {Wratsch, 1895, Nos. 48 and 57) and others {ibid., 1896-98) to Gastrophilus larvae. It burrows into the epidermis of man by minute passages. This observation should, however, be verified. The phenomenon is designated as skin- mole, larva migrans, and creeping eruption. Other Papers on Dipterous Larv/e, etc., in Man. (i) " Ein Fall von lebenden Fliegenlarven im menschlichen Magen," Deutsch nied. Wochenschr., Leipz. and Berl., xxiv (12), pp. 193-194. Bachmann, and review of same, " Living Fly Larvre in the Human Stomach," fhiladelphia Med. fota-n., 1898, i, 18, p. 773. (2) "Siidi una larva di dittero parassita della congiuntiva umana," Ami. di ottal., Paira, 1895, xxiv (4), pp. 329-336, I fig., E. Baquis. (3) " Sur quelques dipteres suceurs de sang, observe a Terre-Neuve," Arch, de Par., Patis, 1900, iii (i), pp. 202-204, E. Barret. (4) "An Account of the Larvae of two Species of Insects discharged from the Human Body," Edin. Med. and Surg, /ourn., January I, i8n, vii {25), pp. 41-48, I pi., figs. I to 8, T. Bateman. (5) " Un cas de myiase par la Sarcophaga magnifica en Roumanie," Bull. Soc. ZooU de France, Par., 189 1, xvi (2), pp. 25-26, R. Blanchard. (6) " Sur les oestrides americains dont la larve vit dans la peau de I'homme," Ann. Soc. ent. de France, 1892, v, pp. 109-154, figs. It 12, R. Blanchard. (7) " Note additionnelle sur les oestrides americans dont la larve vit dans la peau de I'homme," Bull. Soc. ent. de France, Pari--, 1894, xiv, pp. 209-211, R. Blanchard. (8) " Note sur des larves de Dermatobia provenant de Bresil," Bull. Soc. ent. de France, Paris, 1893 {2), pp. 24-27, R. Blanchard. (9) " Larven der Wohlfahrtfliege {Sarcophila wolfahrtii) im Zahnfleische eines Menchen," Wratsch., St. Petersburg, 1888, 5-6, E. K. Brandt. (10) '• Ueber den sogenannten Oestrus honiinis und die oftmals besichteten Verirrungen von Oestriden der Saugetheieie zum Menchen," Verhandl. d. k. zool.-bot. Gesellsch., i860, x Abhandl., pp. 57-72, Brauer. (11) "Ueber die Larven der Gattung Cuterebra, Clk.," Verhandl. d. k. zool.-bot. Gesellsch., i860, x Abliand., pp. 777-786, Brauer. (12) " Des desordres produits chez I'homme par les larves de la Lucilia hominivorax,''^ These, Paris, 1864, 43 pp., V. Audouit. (13) " Note on the 'Flesh Worm,'" Med. Press and Circ, London, April 12, 1882, Ixxxii (N.S. xxxiii), p. 314, P. S. Abraham. OOO THE ANIMAL PARASITES OF MAX (14) *' Larvas de la Calliphora liniensis en fosas nasalis," 1855, 18 pp., F. Aguirre. (15) " Raro caso di parasilismo nell 'uomo dovuto alia larva di una mosca {Sarcophaga affinis, Meigen)," Boll. d. Soc. Rom. fer gli Stud. Zool., Roma, 1893, i^ (5-6), pp. 278-289, I pi., 3 figs., Giulo Alessandrini. (16) " Observations sur I'espece de ver nomme Macaque (Oestrus)," Mem. Acad. Sci- par Hist., 1753, p. 72, F. Artur, (17) " Contribuicao ao estudo da biologia da Dermatobia cyanivent?-is,'^ Trav. do Inst, de Maiigufjihos, 1908. BlTlNG-MOUTHED AND OTHER NOXIOUS DlPTERA WHICH MAY BE Disease Carriers. [Amongst the division Brachycera (as meant in this work) we get several groups of flies which, like the fleas and mosquitoes, are partially parasitic on man, the adults, mainly in the female sex, being provided with a piercing mouth with which they extract the blood of man and animals. The importance of these parasites is net the mere fact that they feed upon our blood, but that they often carry germs from man to man (tsetse-flies and trypanosomiasis, Tabaiiidce and anthrax). Amongst the most important biting-mouthed Diptera in this section are the following : Tabanidcp., or gad-flies ; Glossitice, or tsetse-flies ; and certain other Muscidcr. Some of the exotic Asilidic and a few Leptidcr also bite man. Family. Tabanidae (Gad-flies). [The Tabanidce have a broad, rather flattened body and a large head ; eyes united in the male (except in some Chrysops). The antennae are composed of three seg- ments, have the third joint composed of five to eight annuli — in Chrysops they are fairly long. The proboscis is projecting, and sometimes much elongated. The legs are moderately stout. The venation of the wings is shown in fig. 415. [This family of gad or horse flies contains a great number of genera, all of which may bite animals and man more or less severely. The female alone is blood-sucking, the males feed upon the juices of flowers. The females deposit their spindle-shaped white, black, or brown eggs on leaves, stems of plants that either overhang or stand in water, and amongst rushes ; they are at first white, but become brown or black. The eggs are laid in rounded, flattened or conical masses com- posed of layers one upon the other. The larvne are carnivorous, feeding upon snails, worms, other larvce, etc., and have a distinct head ; they are cylindrical, com- posed of eleven segments, the last with a vertical breathing pore, or the last two segments may form a breathing tube. The majority taper to a point at each end, in colour shining white or dull grey to yellowish, many of the larger specimens mottled or banded with dark brown or black. The first seven abdominal segments are encircled near the anterior margin with a ring of fleshy protuberances consisting of a transverse dorsal ridge which may be divided by a depression into two. The young larvae burrow into any soft vegetable substance ; they live both in the water and under damp soil surrounding water, also in damp earth generally. The larvte are not only carnivorous, but they are cannibals, frequently devouring their own species. They may take more than a year to mature. [The pupcx are found close to the surface of mud and earth, and are mostly dull yellowish to brown in colour, with rows of spines on the distal third of each abdominal segment ; the thorax bears a pair of ear-shaped spiracular structures, and there are also six denticles at the apex of the abdomen. [A habit common to the adults of most of the Tabaiiidce of considerable economic TABANIDi^i: 6oi importance is that of the adults coming to water to drink. Portschinsky' has found that by applying kerosene to the pool they frequent the adults are killed, and Hine- that the same oil kills the larvas that fall into the water from eggs laid on plants above, \Tabanida' are not only of importance as purely biting insects, for they may often convey pathogenic organisms from one animal to another, such as the bacillus of anthrax, which they are known to carry, and possibly also trypano- somes in regard to man. Chrysops also acts as a host of Filaria loa in South Nigeria (Lei per, Brit. Med. Joiirn.^ January, 191 2, pp. 39-40). Two species are incriminated, viz.^ C. silacea and C. dimidiata. With animals these flies play a more important part, for MM. Sergent, in Algeria, have proved that species of Tabanus are able to transmit three forms of animal trypanosomes by biting a healthy animal as long as twenty-two hours after having bitten an unhealthy Fig. 415. — The ox gad fly {Tabanus boviims, Linn.). one. In India they have also been shown to transmit the parasite of "surra" in dogs and rabbits by Rogers. Other observers have since corroborated these results, and Mitzmain, who has recently performed valuable work in this connection, states that T. striaius is undoubtedly the carrier of this disease in the Philippine Islands. Certain members of the genus Haematopota have also been shown to be capable of the direct transmission of Trypanosoma evansi. Martoglio {Ann. dig. sper.^ 191 3» xxiii, N.S., No. 3, pp. 363-366) states that the trypanosome disease of dromedaries known as salaf is transmitted by Tabanidce., especially Pangonia {P. magretti and P. beckeri) in Italian Somaliland. It is quite likely that these flies play a much greater part in the spread of such diseases than is imagined at the present time. [The TabanidcE are divided into two groups or subfamilies : (i) The Pangonince.^ ' Vide Btdl. 20, N. Sc, U.S. Div. Ent. - '* Tabamda of Ohio," Ohio State University Bidt. 19, 1903, sec. 7, p. 14. 6o2 THE ANIMAL PARASITES OF MAN and (2) the Tabaniiice ; the former have spurs on the hind tibia" and usually ocelli ; the latter have neither tibial spurs nor ocelli. [The Paiigoni7ice contain two main genera, Pangonia and Chrysops. In the former the proboscis is much elongated, and the third antennal segment is composed of eight rings, and is never angulated or ungulated at the base. The proboscis is often very long. [In Chrysops, the so-called blinding storm flies, all the three segments of the antennae are long, the third having only five annulations, and the proboscis short but very strong. [There are many genera in the Taba7iince, which are found in all parts of the world, of which two only are shown here — viz.^ Tabanus and Ha^matopota. The former has the first two segments of the antennas short, the third angulated at the base, sometimes spurred and composed of five annulations ; the second has the second segment short, and the third composed of four annulations — never angulated nor spurred at the base — and the wings are adorned with grey or brown markings. These latter are usually called "brimps" and "clegs" in Britain, the former gad or horse flies, the seruts and mangrove flies of tropical countries. Fig. 416.— The brimp {HcBinaiopotaphivialis, Linn.). Family. Asilidae (Wolf Flies). [These flies are of little importance in regard to the subject dealt with in this book ; but I have notes sent concerning the biting habits of one or more species belonging to this family from the Malay States and Africa. {^Asilidce, or wolf tiies, are easily told by the following characters : Large or moderate-sized flies, thickly hairy ; head separated from thorax by a narrow neck ; eyes separated in both sexes; proboscis firm and horny, adapted for piercing; abdomen long, pointed, and composed of eight segments. Legs strong and bristly, of moderate length. Wings sometimes mottled, lying parallel over the abdomen when at rest. There are nearly 3,000 species. They live mostly upon insects, but some are said to bite animals and man. They are, however, of little importance in this respect. BLOOD-SUCKING MUSCID^ 603 Family. Leptidae. [This widely distributed family of flies has a few species which suck the blood of man, and the writer has been personally badly bitten in Norway by a Leptis which was apparently Leptis scolopacea. [The Leptidce have usually blotched wings and similar venation to Tabanus ; they are elongated flies of moderate or large size, and of dull colours. The antennas are varied and consist of three segments, either with or without a terminal bristle or with the third segment compound, and in a few they may be almost nematocerous. The wing veins are distmct, very crowded anteriorly, the third long vein is furcate, basal cells large, and there are usually five posterior cells, the anal cell being open in some ; the squanicC are always small, sometimes only rudimentary, [Four are known to be blood-suckers, namely the American Symphoromyia, Trichopalpus obscurics in Chili, and Leptis strigosa and L. scolopacea in Europe. The genus Symphoromyia has a single spur on the hind tibicE, none on the fore or, raid tibic\2, the third segment of the three-ringed antennae kidney-shaped, and a short proboscis. In the genus Leptis the hind tibiae have two spurs, and the third antennal segment is not reniform. [The other biting genus Trichopalpus can be told at once by the elongated proboscis. Most of this family live upon other insects. The larvas live in earth, decaying wood, sand, stagnant waters, and the nests of wood-boring beetles ; they are usually cylindrical and may have fleshy abdominal legs ; the anal segment has a transverse cleft, and often two posteriorly directed processes and two stigmata between them. They are all predaceous, and in one genus (Vermileo) make pitfalls in sand like the ant lions {Mynneleon). Bloods sucking Muscidse. [The blood-sucking MuscidcE are mainly contained in the following genera : Glossina, Stomoxys, Haematobia, Lyperosia, Stygeromyia, Philsematomyia and Bdellolarynx. [The first is the most important genus on account of the part it plays in the spread of trypanosome diseases. Stomoxys may also serve as a disease carrier. The remainder and a few more genera cause considerable annoyance by their bites, and may also act as occasional carriers of pathogenic organisms. All these flies have their mouth parts elongated to some extent, forming a distinct proboscis, which becomes more or less strongly chitinized ; the labella are usually serrated or spiny, and thus form a structure easily capable of piercing the skin. Unlike the Culicidcc^ the blood-sucking Muscidce have the sanguinary habit common to both sexes. Genus. Glossina, Westwood. [This genus contains sixteen species,^ all of which are confined to the Ethiopian region. Glossina may be distinguished from other allied genera by the proboscis, the antennae, wings, and male genitalia. The proboscis projects forwards and has 1 swollen bulb-like base to the slender labium which holds the two structures, the needle-like epipharynx and the thread-like hypopharynx ; the whole proboscis is ' This does not include G. mactdata, Newstead, which is regarded by Austen as a synonym of G. palpalis, Rob. Des. ; according to this authority the curiously spotted appearance of the type and only example of G. maculata is due to foreign matter. 38 604. THE ANIMAL PARASITES OF MAN ensheathed in the maxillary palpi. The antennas have the first two segments small, the third large with a marked pore, the orifice of the sense organ near the base ; from the base of the third segment also arises the three-jointed arista, the first two segments being, however, minute ; the third bears a series of from seventeen to twenty-one fine branched hairs on one side. The male genitalia or hypopygium is more or less oval and tumid, its long axis lying in the antero-posterior direction, with a vulviform median groove (the anus) running from the anterior margin to beyond the middle. [Newstead has shown the importance of the study of the genitalia in separating species {vide Bull. Ent. Res., ii, pp. 9-36 and 107-110, and iii, pp. 355-360; and Anji. Trop. Med. and Par., vii, No. 2, PP- 331-334)- [The tsetse-flies reproduce differently from all other Miiscidce. The female pro- duces at each birth a single full-grown larva, which is retained within the oviduct and there nourished by the secretion of special glands, and on being born crawls to some hiding place and at once becomes a puparium. Fig. 417. — Head of Glossina longipalpis, Wied. (After Griinberg.) Fig. 418. — Antenna of Glossina pallidipes, male. (After Austen.) [The larva is a yellowish footless maggot nearly as large as the mother's body, the skin shagreened and the anal extremity having a pair of large, black, granular prominences separated by a depression containing the breathing pores. [The puparium is brown of various shades, the tumid lips of the larva being con- spicuous,^the size and shape of the lips enabling the puparia to be identified. SPECIES OF GLOSSTNA 605 [These puparia are often found in masses at the base of trees, in hollows in trees and rocks just buried under vegetal debris. These insects are generally confined to definite tracts known as " fly-belts." They usually occur in damp, hot places on the borders of rivers and lakes, and never far from water in the case of the palpalis group, although others of the morsitans group may be found a considerable distance from water. They are usually absent on grass plains, but may now and then occur there (Kinghorn, vide Hindles' " Flies and Disease, Blood-sucking Flies," 1914, p. 274) ; cover of trees, shrubs, or thick reeds is essential to them. [Their range in Africa extends roughly from 18° N. to 31° S. Sfilossina palpalis is the chief carrier of the more prevalent type of sleeping sickness. Two distinct types of parasites can produce this disease, viz.^ Trypano- soma gambiense^ which produces the ordinary sleeping sickness, transmitted by G. palpalis, and Trypanosoma rhodesiettse the Rhodesian or Nyasaland sleeping sickness, transmitted by G. 7norsitans, and possibly identical with T brucei^ the parasite of N'agana. Koch has also shown that G. pallidipes^ Austen, and G.fiisca^ Walker, can be artificially infected with the human trypanosome. It appears probable that Koch used G. brevipaipis^ not G. fiisca^ in his transmission experi- ments, as at that \\m^ fusca included nearly all the large tsetses, but brevipalpis is its Eastern representative. [A Table of Species (modified after Austen) is appended here : — tachinoides, West wood. I. Glossifta palpalis GROUP. Dorsum of abdomen ochraceous buff or buff ; third and following segments exhibiting sharply defined, dark brown or clove brown, interrupted transverse bands ... Dorsum of abdomen not so marked ... ... ... 2. Third joint of antenntc pale (cream buff to ochraceous buff), clothed with long and fine hair, forming a conspicuous fringe on front and hind margins pallicera^ Bigot. Third joint of antennas entirely dark (mouse-grey) except at extreme base on outer side, and without a conspicuous fringe of long and fine hair Dorsal surface of abdomen dark sepia brown , median paler area on second segment broad, and more or less quadrate or irregular in outline hypopygium of $ buff or ochraceous buff Dorsal surface of abdomen blackish-brown ; median paler area cuneate {i.e.^ triangular in outline) hypopygium of $ grey 3- caliginea^ Austen. palpalis^ Rob. Desv. II. Glossi?ia morsitans Group. Hind tarsi entirely dark ; small slender species ; abdomen bright ochreous or reddish ochreous with dark lateral markings Hind tarsi not entirely dark ; abdomen drab-grey, buff or ochreous buff with conspicuous dark interrupted transverse bands ausfenii, Newstead. 2. 6o6 THE ANIMAL PARASITES OF MAN Last two joints of front and middle tarsi with sharply defined clove brown or black tips Last two joints of front and middle tarsi without sharply defined clove brown or black tips (front and middle tarsi either entirely pale or, at most, last two joints of front tarsi faintly brownish at the tips), and last joint and distal half of penul- timate joint of middle tarsi light brown, never so dark as to form a sharp contrast with the remaining" joints Third joint of antennas with a distinct fringe of fine hair on front margin ; dark brown or clove-brown bands on abdommal segments extendmg close to hind margins (z>., pale ground colour, apart from the median interspace, confined to a very narrow hind border) Third joint of antennae without a distinct fringe of fine hair on front margin ; dark brown or clove- brown bands on abdominal segments not extend- ing close to hind margins pallidipes, Austen. lo?jgipalpis, Wiedeman. morsitaiis^ Westwood in. Glossina fusca GROUP. Third joint of antennae fringed with fine hair on anterior and posterior margins ; fringe on anterior margin conspicuous under a hand lens magnify- ing 1 5 diameters (nominal) when head is viewed in profile ... Third joint of antennae with fringe of fine hair on anterior margin so short as to be scarcely notice- able under a hand lens magnifying 15 diameters (nominal) when head is viewed in profile (longest hairs m fringe in length not exceeding one-sixth of width of third joint) ; palpi long and slender ... Longest hairs in fringe on front margin of third joint of antennas, in length equal to from one- fourth to one-third (not exceeding one-third) of width of third joint ; palpi of moderate length ... Longest hairs infringe on front margin of third joint of antennas in length equal to from one-half to three-fourths of width of third joint ; palpi noticeably long and slender Pleurae drab-grey or isabella-coloured, hind cox^ bufif or greyish-buff .... Pleurae dark grey ; hind coxae mouse-grey 3- tabaniformis^ Westwood. nigrofusca^ Newstead. fusca. Walker. fuscipleuris, Austen. IV. Glossina brevipalpis GROUP. I. Dorsum of thorax with four sharply defined brown, more or less oval or elongate spots, arranged in a parallelogram, two in front and two behind the GLOSSINA PALPALIS 607 transverse suture ; proboscis bulb with a sharply defined brown or dark brown tip ... Dorsum of thorax without such spots ; proboscis bulb not brown or dark brown at tip Wings with upper thickened portion of anterior transverse vein much darker in colour than adjacent veins and thus standing out con- spicuously against the rest of the wing ... Wings with upper, thickened portion of anterior transverse vein not much darker in colour than adjacent veins, and thus not standing out con- spicuously against the rest of the wings (wings practically unicolorous) longipennis, Corti. brevipalpis^ Newstead. jnedicorum^ Austen. Glossina palpalis, Rob. Desv. [This is the chief carrier of sleeping sickness in Nature. It is found in places over the whole of West x^frica from the mouth of the Senegal River to Angola, and extends eastwards into the Bahr-el-Ghazal. The eastern boundary follows the valley of the Nile and includes the eastern shores of Lakes Victoria and Tanganyika ; from the southern end of the lake the boundary tends south-west, approximately following the frontier between North-eastern Rhodesia and the Congo Free State, and passing through the Katanga district of the latter country into Angola (Austen). It may occur up to 3,000 ft.; but, accord- ing to Bagshawe, it has not been recorded above 4,000 ft. It feeds on the blood of many animals, including reptiles, amphibia, birds, and even amphibious fishes, as well as all the wild mammals. It seems, how- ever, to possess a decided predilection for man, and undoubtedly thrives better upon mammals and birds than upon cold-blooded animals. [It is not usually found far from water, requiring a humid atmosphere and tem- perature of about 85° F. (shade). But a marked seasonal distribution is shown, the flies considerably extending their range during the rainy season, and thus visiting districts which are dry for the greater part of the year; as the rains diminish the fly gradually leaves the temporary haunts and returns to the more permanent ones. It bites only by day, and then only in sunny weather, and usually lives in shade. [Roubaud has shown that the first larva produced is about three weeks after copulation, and that others are produced at an interval of nine or ten days. The puparium stage is rapidly produced after the expulsion of the larva, often in Fig. 419. — Glossina palpalis and pupa rium. (After Brumpt.) ' Newstead has recently described another species as G. severini {Ann. Trap. Med. and Par., 19 1 3, vii, No. 2, pp. 331-334). It is allied to G. fuscipleuris, Aust n. 6o8 THE ANIMAL PARASITES OF MAN three-quarters of an hour. The puparium stage lasts from thirty-two to thirty-five days. The puparia occur in well-drained humus close to water, sheltered by trees or bushes, in crevices in rocks, and between the exposed roots of trees, sometimes in sand. [Bruce has shown that only a very small percentage of flies fed experimentally on infected animals ultimately become infective, and that the infectivity of this small percentage depends upon a delayed infection of the salivary glands. [A variety, wellmani of Austen, is found in Angola, Gambia, the Katanga district of the Congo Free State, the Matondwi Islands of Tanganyika, etc. Glossina morsitans, Westwood. [This species has been shown by Kinghorn and Yorke, and also by Bruce, to be responsible for the transmission of Trypanosoma rhodesiense, the micro-organism producing sleeping sickness in man in Rhodesia and Nyasaland and also in parts of German and Portuguese East Africa. Fisher and Taute have demonstrated Fig. 420. — The tseise-fly {Glossina viorsilans, Westwood). experimentally that Trypatiosoma gainbiense — the sleeping sickness parasite of other parts of Africa— may also be transmitted by this fly, and in addition it is known to be capable of disseminating several species of trypanosomes pathogenic to animals. Of these, T. brucei {= ? T. rhodesie7tse), the parasite of tsetse disease, first incriminated by Bruce, is perhaps the most important. [It is the most widely spread of all tsetse-flies ; its range extends from Sene- gambia in the north-west to Southern Kordofan and Southern Abyssinia in the north-east, and then southwards to the Bechuanaland Protectorate, North-eastern Transvaal and Zululand. The actual localities given by Austen are Gambia, French Guinea, Gold Coast, Togoland, Dahomey, Northern Nigeria, Congo Free State, the Bahr-el-Ghazal, the Uganda Protectorate, German East Africa, and Portuguese East Africa. [This species is confined to " belts," often of very limited extent, and appears to prefer regions where there is sufficient vegetation for moderate but not excessive cover and a hot, moderately dry climate. It is not. nearly so dependent upon STOMOXYS 6C9 water as is G.'palpalis^ and generally is most active in a dry atmosphere ; some observers, however, state that in certain districts it is more common along the banks and edges of rivers. This tsetse-fly has been taken as high as 5,500 ft. altitude. It infests native villages as well as the bush. Like other tsetse-flies it bites not only during the hottest part of the day, but also on bright warm moon- light nights, and it feeds on the blood of all mammals. [The structure of the male genitalia of those representatives of G. inorsitans occurring on the West Coast of Africa and in parts of the Soudan presents certain constant differences from that of the typical form of this species ; this form is known as G, morsitajis^ race sub?norsitans, Newst. Genus. Stomoxys, Geoffroy. [The members of this genus which occur in temperate and tropical countries are provided with a hard, slender, shiny black proboscis which projects horizontally from beneath the head ; by means of this structure they can bite severely. In general appearance they resemble house flies, but the proboscis at once Fig. 421. — The stinging fly {Stomoxys calcitrans, Linn.). distinguishes them. In many parts of Britain they are known as storm flies on account of their frequent appearance indoors previous to a storm of rain or wind, which I have invariably found to be correct ; they are also called stinging flies. In colour they are greyish, dusky or brownish-grey or black, varying from 5 to 7 mm. in length; the thorax has dark longitudinal stripes and the abdomen dark spots or bands. In the male the eyes are closer together than in the female. These flies usually occur in stables and farmyards, along woods and in lanes, and mainly attack mammals. [One s^G:c\Qs{Slo?noxys calcitrans, Linnaeus) occurs practically all over the world. The female lays her eggs in moist, warm, decaying vegetation ; as many as eighty may be laid by a single female. The ova are white, banana-shaped, with a broad groove on the shorter curvature ; they may hatch in two or three days. The creamy-white larva tapers to a point at the head end, and is truncated at the tail end. Two black mouth hooks are plainly visible at the cephalic extremity. There are two plates on the posterior surface of the last segment which bear the respiratory pores, nearly circular in outline. It reaches maturity in fourteen to twenty-one days; when mature it is 11 mm. long. The pupal stage is passed in the old larva 6io THE ANIMAL PARASITES OF MAN skin and lasts from nine to thirteen days ; it is barrel-shaped, 5 to 8 mm. long, and of a bright reddish-brown to dark chestnut-brown colour. [This insect may act as a carrier of anthrax, and has been proved to be the agent of an extensive epidemic of malignant pustule in the Isle of Pines, New Caledonia.^ [Noe's- experiments tend to show that it is an intermediate host and transmitter of Filaria labiato-papillosa of the ox. [Surra is generally stated to be transmitted by Stomoxys as well as Tabanus, and yet Nitzman in the Philippines obtained uniformly negative results in exhaustive experiments. Others have also been unsuccessful. Certainly Stomoxys can transmit the disease in French West Africa (Bonet and Roubaud), and mechanically has been proved to be capable of disseminating other trypanosomes (experimentally) : sleeping sickness (J", gcunbiense) ; nagana (7". brucei) ; sou ma (7". cazalboui) ; and el debat (Z'. soudane7ise). [5. calcitrajis may also be a carrier of poliomyelitis (Rosenau and Brues, Harvard Alumni Bulletin, 191 2, xv, No. 9, pp. 140-142). Several species are ow known (5. brunnipes^ Griinb. ; S. inornata, Griinb. ; S. fiigra^ Macq. ; S. omega^ Newst. ; S. ochrosoma^ Speiser, etc.). Genus. Lyperosia, Rondani. [A genus of small flies which bite man and animals, but are not so far connected with the transmission of any disease in man, but in Java it appears to carry surra (P. Schat, Meeledeel Praefstatioti Oost-Java, 1903, 3e ser.. No. 44), the species being Lyperosia exigua, Meijere. These flies can be told from Stomoxys by the palpi being broader, flattened laterally, and as long, or nearly so, as the proboscis. When not feeding the palpi enclose the proboscis, as in Glossina. They are usually about half the size of Stomoxys, and are the smallest blood-sucking Muscidcc. They frequently swarm around and upon domesticated animals. [The life-history of the horn fly in America (Z. irritans, Linn.) is well known. It lays its ova singly in freshly dropped cow-dung, and there the maggots feed, pupating in the soil beneath. [Patton and Cragg also give some details as to the life-history of Liperosia exigua ("Medical Entomology," p. 375) as follows: " Z. exigiia, whose habits have been observed in Madras, usually lays twelve eggs at a time. The flies immediately return to the cow and the process is repeated when the dung is again dropped. The larvae migrate from the dung when about to pupate, and the puparia are always found in the earth at some distance away or under the sides of the patch of dung. The fly usually hatches out in five days, though sometimes as late as the eighth. Weiss has studied the life-history of irrila?ts var. weisii from Algeria ; its larval stage lasts five days, and the flies hatch out of the puparia in another five days." [The other biting genera oi Muscidce, Hsematobia, Hasmatobosca, Bdellolarynx, Stygeromyia, and Philaematomyia, although sometimes annoying to man, have not in any way been connected with any disease. [The horse fly {Hcnnatobia irrita?is, L."^) attacks cattle chiefly, but now and then man is bitten. The different species can be told from Stomoxys by the palpi being nearly as long as the proboscis. ' Bull, des Seances de la Soc. ent. de France, 1878, pp. cxliv, cxlv. == Atti della Reale Accad. del Lincei, Anno CCC. Se Quinta, 1903, xii, 2 sem. fasc, PP- 387-393- 2 This is apparently the stimulans of Meigen. PUPIPARA 6ll [The genus Philaematomyla, Austen, is intermediate between Stomoxys and Musca in structure, and between the non-blood-sucking Musca, as M. domestica^ and the blood-sucking Musca pattoni^ Austen, which feeds on the blood exuding from the bites of true blood-suckers. They occur in Central Africa and India, Ceylon and Cyprus {vide " The Life-history of PhilcEmatomyia insignis^ Austen," Ann. Trap. Med. and Par. ^ 191 2, v, p. 515). [Two flies belonging to the family Anthomyidae also attack man, namely : — \HydrotcEa jneteorica, L. (the meteoric fly). This fly attacks man as well as animals. They especially bite around the eyes and nostrils of animals, but are not so particular with man ; the head, however, is usually chosen. Linnaeus called it the meteoric fly because it often forms clouds around horses' heads at the approach of rain. The Hydrotasas are usually black or blue-black in colour with bare eyes and simple abdomen, the front femora peculiarly constructed. H. meteorica, L., occurs in Britain. [The members of the genus Hydrophoria, Desvoidy, also bite man. Pupipara or Eproboscidae. [The Pupipara are all blood-suckers, the majority occurring as parasites on mammals and birds, where they are more or less permanent parasites. Occasionally some may attack man. They all produce their young fully formed, and they assume the pupal stage immediately after extrusion. The puparia are large. They are mostly flat, louse-like flies which may or may not be winged. In the case of Melo- phagus I have found the puparia are often passed by the female. The winged forms have a short quick flight, and when disturbed will seek shelter in man's hair or beard. Two main families occur : (i) the HippoboscidcF, and (2) the NycteribiidcE. The former occur on animals and birds, the latter on bats only, but may invade man. Two other families are known — the Braulidce (bee parasites) and the Streblidce (bat parasites). [The mouth of the Hippoboscidce is long and sharp, forming a proboscis. The thorax and abdomen are flat and leathery. The legs are stout and strong, and terminate in large dentate claws and other structures of use in holding on to the hair or feathers of their host when blood-sucking. [Austen says it is probable that the Hippoboscidce are descended from ancestors belonging to the MuscidcB., which underwent modification in bodily structure as the consequence of the adoption of a parasitic mode of life. [Two wings are present in the true Hippoboscse, Hippoboscaequitia (of the horse), H. camelina (of the camel), H. maculata (of oxen), and H. capejtsis (of dogs), but are absent in Melophagus, the sheep tick or ked fly {M. oviniis). [In two genera, Lipoptena and Echestypus, wings are at first present, but are lost as soon as the fly finds its permanent host. [With regard to their biting man, such is only occasional. I have known sheep shearers to be badly bitten by Melophagus ovinus, and have more than once been attacked myself when standing where shearing is taking place. Sharp records the grouse parasite, O^^nithomyia lagopodis^ as once biting severely a gamekeeper in Scotland. There are also records of H. inaculata biting man in Africa and India. [Although so far not connected with any human disease, it is interesting to note Theiler has shown that Hippobosca rufipes^ v. Olfers, and H. maculata^ Leach, are capable of transmitting Trypanosoma theileri^ Laveran, the cause of gall sickness amongst cattle in the Transvaal. It is now considered, however, that Trypano- soma theileri is non-pathogenic, and that the cause of gall sickness is a piro- plasma-like organism known as A7iaplasma margi7iale. Theiler, Laveran and 6l2 THE ANIMAL PARASITES OF MAN Mesnil all hold this view {vide Laveran and Mesnil, " Trypanosomes and Trypano- somiases," second edition, 191 2, p. 330). [Zjw^-^m.— Three members of this genus have been shown to transmit the non-pathogenic (?) organism, HceDioproteus columbcE amongst pigeons in Algeria and S. America. Insects and Epidemic Poliomyelitis. [In a recent number of the Joicrftal of Economic Entomology} Brues and Sheppard point out the possibility of acute epidemic poliomyelitis (infantile paralysis) being an insect-borne disease. They summarize as follows : — [Many facts connected with the distribution of cases and the spread of epidemics of this disease with histories of insects bites, suggest at least that the disease may be insect-borne. Field work during the past summer, together with a consideration of the epidemiology of the disease so far known, points strongly towards biting flies as possible carriers of the virus. It seems probable that the common stable fly {Stomoxys calcitrmts^ L.) may be responsible to a certain extent for the spread of acute epidemic poliomyelitis, possibly aided by other biting flies such as Tabanus lineola. No facts which disprove such a hypothesis have as yet been adduced, and experiments based upon it are now in progress. [If the disease should prove to be common to any species of domestic animals, as is now strongly suspected, a secondary connection of ticks in spreading the disease among such animals seems probable, as has been mentioned. [The following is some of the more important literature on Diptera in general : Meigen, J. W., " Syst. Besch. d. bek. europ. zweifliigligen insecten," 1818-1838, 7 vols. ; Brauer, F.» "Monographic der Oestriden," Wien, 1863; Idem, "Nachtr. hiersu, " Wien. ent. Zeit.i 1887, vi, pp. 4, 71 ; Schiner, J. R., " Fauna austriaca : die Fliegen," Wien, 1860-64; Low, Fr., " Ueber Myiasis und ihre Erzeuger," Wien. vied. Wochenschr., 1882, xxii, p. 247; 1883, xxxiii, p. 972; Joseph, G., " Ueb. Fliegen als Schadlinge und Parasiten des Menschen," Deiitsch. vied. Zeit., 1885, i, p. 37 ; 1887, iii, pp. 713 and 725 ; Peiper, E., " Fliegenlarven als gelegentl. Paras, d. Mensch.," Berlin, 1900; Theobald, F. V., "Monograph of the Culicidai of the World," 1901-1911, 5 vols, and i atlas, plates ; Austen, E., "A Monograph of Glossina Tsetse-flies," 1903, i vol. ; Van der Wulp, " Diptera neerlandica," 1877 ; Walker, " Insecta Britannica : Diptera," 1851-53 and 1856; Lundbeck, "Diptera danica," 1907-12; Zetterstedt, "Diptera scandinavire," 1850 ; Theobald, "British Flies," 1892; Aldrich, " N. American Diptera," 1905 ; Loew and Osten Sacken, "Monographs of the N. American Diptera,'' 1862-63 and 1869; Macquart, "Diptera exotique," 1830-47; Rondani, "Diptera exotica et Italica," 1863-68 ; Williston, " Manual of Famiiies and Genera of N. American Diptera," second edition ; Verrall, " British Flies." A fuller literature will be found in Peiper, as well as in Ruber's " Bibliographie d. klin. Ent.," 1899, iii, Jena, in the Bibliography at the end of this work and in the Rev. of App. Ent. (Dulau and Co., London), where all references to modern research can be found. — F. V. T.] ' Charles T. Brues and Philip A. E. Sheppard, "The Possible Etiological Relation of certain Biting Insects to the Spread of Infantile Paralysis," /^«r;/. Econ. E71L, 1912, cciv, pp. 305-324- ADDENDA 613 ADDENDA. Akamushi or Kedani Sickness (vide also p. 487). — Schutfner (Far East. Assoc. Trop. Med., Conipi. rend. Trois. Cong. Biennial^ 19^3, Saigon, 1914, pp. 309-315) states he observed a peculiar fever in Deli, Sumatra, somewhat resembling typhoid. This he traced either to a mite or tick. He figures the possible carriers, namely, a Trombidium and Cheyletida'. He calls this disease pseudo-typhus — a variant of Japanese kedani sickness, which, he says, also occurs in the Philippines. Ticks. — African Tick Fever : Marzinovsky (Proc. of Conference of Bacteriologisfs and Representatives of Medical Sanitary Authorities on the Campaign against Infections Diseases in connection with the War, Soc. Rnss. Physicians in mem. Pirosov, Moscow, 191 5, pp. 56-68), states that African tick fever has been imported into Persia, and that it is there carried by Ornithodorus tholosani. Tick Pakalysis : Todd (" Paralysis and Tick-bite," Can. Med. Assoc. Jonrn., 1914, iv. No. 9, pp. 825-826) refers to paralysis ascribed to the bites of ticks in children, and possibly adults, in America, British Columbia and Australia. He states that a young child, perfectly well one day, has more or less complete paresis or paralysis on the next, fever, a rapid pulse, and other constitutional symptoms. The child may be dull and stupid, and may have convulsions. If the tick is not found and removed the child may die, but if it is removed, the symptoms disappear and recovery is complete in a few hours. The tick must be entirely removed. Diptera. — Psychodid^ : Bolt {China Med. Jonrn., Shanghai, xxix. No. 2, pp. 78-86) states that sand-flies (Plebotomus) and the fever due to them are common m North China, May and June being the worst months. The natives of the region appear to be immune, but all others suffer. Old ruined buildings are the favourite haunts of the Phlebotomus. The species of Phlebotomus has not been determined. Pulicidae. — Dermatophilus (Sarcopsylla) penetrans, or the *' Jigger." — This flea (r/t/d p. 544) is believed by Lama {Giorn. Ital. Mai. Yen., Milan, 1914, xlix, pp. 465-472) frequently to carry leprosy and he points out that the early lesions of leprosy usually appear on the uncovered parts of the body. This flea also attacks rats. Brachycera. — Leptid^ {vide p. 603) : White, A. {'' The Diptera- 6l4 THE ANIMAL PARASITES OF MAN Brachycera of Tasmania," part I, Papers and Proc. Roy. Soc. of Tasuiaiiia for 1914, 191 5, pp. 35-74), describes a new blood-sucking Leptid; Spaniopsis tahaniformis, which resembles a small gad fly (Tabanus) in appearance. Pycnosoma putorium : This is believed by Roubaud (" Les Pro- ducteiirs de Myiases et Agents similaires chez I'homme et les ani- maux," Paris, 191^, part I) to be largely concerned in the spread of amoebic dysentery in French West Africa. LtLciUa argyrocephala, Macquart : This green-bottle fly is described by Roubaud as producing myiasis in Africa (" Les Producteurs de Myiases et Agents similaires chez I'homme et les animaux," 19 14, Paris, part I). It attacks ulcers and sores in man and animals. Aiichmeromyia liiteola, Fabr. : Schwetz (Ann. Trop. Med. and Par., 19 14, viii. No. 3, pp. 497-507), collected a large quantity of this insect at Kabinda. He placed them in flasks with sand and a few days later they pupated, and in fifteen days several flies hatched out. The larval period varies from an unknown minimum up to several months. The larva may live for at least two months without food. A female oviposited on the 17th, and on the i8th one larva hatched. The pupal stagei seems to last eight to fifteen days. The larvae appear to bite by day as well as night according to native m format ion. Cordylobia anthropophaga, Griinb. : Roubaud (" Etudes sur la Faune parasitaire de I'Afrique occidentale frangaise," part I, ''Les Producteurs des Myiases et Agents similaires chez I'homme et les animaux," Paris, 1914) gives the life-history of this species. One fly laid 150 ova in a glass vessel, on the sides, and on some rotten fruit, and died the following day. He found that fifteen larva? just hatched placed on sand in a glass vessel with a guinea-pig gave rise to characteristic tumours on the ventral surface of the body and the anus. Other experiments failed. It thus seems that infection takes place from larva? which have hatched apart from the host. Infection of man is regarded as accidental; no positive infection of horses, oxen, sheep or pigs is known— it is rare in goats, and poultry never seem to be attacked. The result of experiments tends to show that the apparent choice of a host is mainly a question of body temperature. The larva, whether freshly emerged or eight to ten days old, penetrates the skin immediately, boring obliquely between the epidermis and dermis. Once removed from the tumour the maggot cannot bore again. The first moult takes place about three days after penetration, and the total period of residence in the host is seven to eight days. Upon emerging the larva falls to the ground and buries itself. In two or three days it pupates and this stage lasts no longer than twenty days. High temperatures, such as 95° F., appear to be fatal. ADDENDA 615 Myiasis.— Coates, G. M., " A Case of Myiasis Aurium accom- panying the Radical Mastoid Operation," Jotirn. Amer. Med. Assoc, Chicago, 111., 19 14, Ixiii, pp. 479-480 : Apparently C. macellaria, forty to fifty coming away with the gauze after the operation. Huber, G. U., and Flack, F. L., "An Unusual Case of Screw- worms in the Nose and Nasal Accessory Sinuses/' Journ. Amer. Med, Assoc, Chicago, 1914, Ixiii, pp. 2288-2289. Auricular Myiasis. — Francaviglia, M. C, "An cora sulla myiasi auricolare," Boll. Sedute Accad. Gioenia, Catania, 1914, No. 31, pp. 15-23. This writer mentions the following parasites in the human ear : Sarcophaga carnaria, L. ; Wohlfartia uiagiiifica, Schiner ; Chrysomyia macellaria, F. ; CalUphora vomitoria, L. ; and Anthomyia pliivialis, L. He refers to a severe myiasis in Russia, due to a fly variously recorded as Sarcophaga zvohlfarti, Rond. ; S. ruralis, Meig. ; or Sarco- pliila meigeni, Portsch. These are all probably synonyms of W. magnifica. Chrysomyia macellaria, in Central America and South America, is quite as harmful as S. carnaria, causing perfora- tion of the tympanum and meningitis. Lucilia nobilis and L. ccesar have also been incriminated. Of the sub-family Anthomyince, the larvae of Fannia scalaris, Meig., F. canicidaris, Meig., F. incisiirata, Zett, and Hydrotcea meteorica, L., are chiefly associated with myiasis. He recommends, if the larvae are outside the tympanum, an injection of chloroform vapour by a few drops of water saturated with chloroform, by an emulsion of 5 per cent, carbon bisulphide or with benzine. When detached they may be removed with forceps or a solution of boric acid. If the tympanum has been perforated, the larvae must be removed at once. Francaviglia also records the larva of Oestrus ovis in the human ear {Boll. Sedute Accad. Gioenia, Catania, 1914, No. 31, pp. 23-27). Body, Head, and Clothes Lice. — Lobaczewski (IVien. klin. Wochenschr., Vienna, 191 5, xxviii, pp. 373-374) recommends the impregnation of body linen with a 30 per cent, solution of oleum betas in 96 per cent, alcohol as an efficient method of keeping the body free of lice. But the process must be renewed each time the linen is washed and it takes fifteen minutes to carry out. On adding the oil to the alcohol, a portion of the former is precipitated, the super- natant fluid is decanted and poured over the linen, which is wrung out in it and dried. The garments retain their lice-proof properties until washed. Three days after wearing the clothes thus treated no lice remain on the body. Portnikov, Proc. of Conference of Bacteriologists and Representatives of Medical Sanitary Author ities on the Campaign against Infectious Diseases in connection with the War, Soc Russ. Physicians in mem. Pirosov, Moscow, 191 5, p. 131). 6 [6 THE ANIMAL PARASITES OF MAN Pediculus capitis and Plithinis pubis are shown to be successfully controlled by applying spirit extract of sabadilla and both white and grey mercury ointment, solution of corrosive sublimate of a strength of I in 250 to I in 100, amyl and ethyl alcohol, benzine, chloroform, carbon tetrachloride, methane, birch tar, liquid of malinin, etc. The control of Pediculus vestiuieuti by the mixture of tartaric acid and sodium sulphite slightly moistened with water is advised. It is placed HI small linen bags underneath the shirt ; the heat of the body produces a reaction which continues for two days, giving off a large amount of SO2, which spreads beneath the shirt and kills all the parasites but does not affect the skin. Marzinovsky, in the same Pioceedings (pp. 56-68), gives a number of remedies for Pediculus i^estiuienti (called liuuiauus), and mentions quinine or mercury, which latter the natives in Tur- kestan carry on their hands and legs in bracelets soaked in mercury compounds. He also mentions ethereal oils, the most effective being clove oil, eucalyptus, oil of anise and camphor. He recommends for disinfecting clothing for army purposes the chamber used by the Japanese on a large scale. Kummerfelds' wash is advised, and is pre- pared as follows : 20 parts of precipitated sulphur are incorporated in a mortar with 50 parts of glycerine ; 2 parts of camphor are separately ground with 50 of eau-de-Cologne and 20 of borax, and 870 parts of distilled water are added ; the whole is mixed together and 3 drops of an extract of musk are added ; shake in order to prevent the sulphur settling down; 50 parts of ether are added to the mixture. This sounds an expensive and troublesome preparation to make. Shipley A. E., '* F'lowers of Sulphur and Lice," Brit. Med. fount., 1915, p. 295. It is here stated by Dr. Lounsbury that the^ South African troops were supplied by the Government with bags of flowers of sulphur sewn in small calico bags and secured to the underclothing next the skin as a preventive of lice. The bags were 2 in. square, one on the trunk and one against each leg. This is a generally accepted preventive, but is best mixed with equal parts of creosote and naphthalene. Shipley, A. E., '' Insects and War," Brit. Med. Jouru., September 19 to November 14, 1914. General advice given re lice. SUPPLEMENT 617. SUPPLEMENT: CLINICAL AND THERAPEUTICAL NOTES. PROTOZOA. INTRODUCTION. The aim of the present volume is to give an account of the animal parasites of man, the number of which is very large. The Protozoa that infest man are very important, and the literature relating to them and to the treatment of the diseases that they produce is very exten- sive. All that can be done in this Appendix is to give a very brief outline of some of the more recent and approved methods of treat- ment, for further details of which the reader should refer to standard medical works, among which the following are noteworthy : — AUbutt and Rolleston (1907): "System of Medicine," vol. ii^ part 2, "Tropical Diseases and Animal Parasites," London. Castellani and Chalmers (191 3): "Manual of Tropical Medicine" (second edition), London. Laveran and Mesnil (1912) : " Trypanosomes et Trypanosomiases '" (second edition), Paris. Manson (1914) : "Tropical Diseases" (fifth edition), London. Mense (1905) : " Handbuch der Tropenkrankheiten," Leipzig. Ross (191 1 ) : " The Prevention of Malaria," London. Scheube (19 10) : " Die Krankheiten der Warmen Lander," Jena. References to the treatments tried in many parasitic diseases can be found in the Sleeping Sickness Bulletin and Kala-azar Bulletin, both now superseded and greatly extended in scope in the Tropical Diseases Bw/Zd/Z/f, published by the Tropical Diseases Bureau, Imperial Institute,. London, S.W. The following diseases, due to protozoa and allied forms, are. discussed : — I. Amoebic Dysentery. II. 'Trypanosomiases. III. Flagellate Diarrhoea and Dysentery. IV. Leishmaniases — Kala-azar and Oriental Sore. V. Spirochaetoses — Relapsing Fevers, Yaws, Syphilis and Bronchial. VI. Malaria. VII. Balantidian or Ciliate Dysentery. 6l8 THE ANIMAL PARASITES OF MAN I— AMCEBIC DYSENTERY. Amoebic dysentery, due to Eniamceba hisiolytica (see pp. 34-41), is present throughout the tropical world and also occurs in temperate zones. Walker and Sellards^ (1913) conducted important experiments with amoebae on prisoners in the Philippine Islands. They showed experimentally that cultural amoebae are non-pathogenic. As regards experiments w^ith Entamoeba coli, after feeding to twenty individuals they concluded that E. coli is a parasite of the human intestine but non-pathogenic and non-culturable. In a third series of experiments, after feeding with motile Entamoeba histolytica, tetragena cysts were found in the stools later; when tetragena cysts were administered, motile E. histolytica were present in the subsequent stools. Some of the histolytica cases developed dysentery after a time. They lay stress on the necessity for the frequent examination of stools in order to detect carriers. The incubation period of entamcebic dysentery is usually long. With regard to the symptomatology of amoebic dysentery, Castellan i and Chalmers distinguish four types — the acute, chronic, latent, and mixed types. The acute type has an abrupt onset ; pain is felt in the lower part of the abdomen, and the motions, rarely exceeding thirty daily, are accompanied by much griping and straining. Blood and mucus are present in the motions, and occasionally greyish material, consisting of leucocytes, mucus, Charcot-Leyden crystals, amoebae, and bacteria, sometimes with particles of tissue. Nausea and vomiting may occur, Digestion is usually deranged. The abdomen is sunken, the liver and spleen are normal, but tenderness is felt along the course of the large intestine. The urine may be diminished in quantity. The chronic type may succeed the acute, or appear like diarrhoea, the motions being faeculent and containing mucus. Between exacerba- tions, constipation may occur. The number of motions may only be twelve to fourteen per diem. Gangrenous complications may occur at any time, and chronic dysentery may persist for many years. The latent type is important, as the patients, though free from dysenteric symptoms, harbour amoebae and act as parasite carriers. The latent condition may lead to acute attacks or to liver abscess. The mixed type occurs where amoebic and baciilary dysentery are combined. There is much fever, nausea, and vomiting. The motions are numerous and often very offensive. Treatment. — The most modern method of treatment, due to Leonard Rogers, is by emetine. According to Castellani and Chalmers, it is w^ell to relieve griping and straining by either a hypo- dermic injection of morphia or by small enemata of 40 minims of ^Philippine Journ. Sc, B, viii, p. 253. SUPPLEMENT 619 laudanum in 1 oz. of mucilage of starch or by using ^ gr. morphia or i gr. codeine suppository. A dose of castor oil (5iv to 5vi) with or without a few minims of liquor opii sedativus or a few doses of saline may be given during the first twenty-four hours. After the castor oil has acted or simultaneously, emetine treatment should be commenced ; ^ to i gr. of emetine hydrochloride, dissolved in sterile normal salt solution, is injected hypodermically three times a day for two or three days. If emetine cannot be obtained, 5 gr. doses of ipecacuanha every three to six hours in the form of membroids, or as pills coated with salol or keratin, can be substituted. After acute symptoms have disappeared, intestinal irrigations once or twice daily, on alternate days, are useful. A solution of tannic acid (3 ^o 5 P^^ 1,000) or of quinine bihydrochloride varying in strength from I in 5,000 to i in 750 is very slowly injected in quantities of ^ to 3 pints by means of a long, soft, rectal tube. For gangrenous dysentery Castellani and Chalmers state that appendicostomy, with irrigation of the whole lower bowel with quinine lotion (i in 1,000) or collargol (i in 500), is the only chance. The use of emetine should be continued in smaller doses after the dysenteric symptoms have ceased, in order to prevent relapses and as a possible safeguard against the development of a liver abscess. Recently (July, 1914), Dr. W. E. Deeks^ has given an account of his successful procedure in dealing with the dysenteries in the Ancon Hospital, Panama Canal Zone, of which medical clinic he is the chief. With regard to amoebic dysentery he advocates : (i) Rest, to increase the patient's resistance ; (2) a generous milk diet, which is practically all absorbed before it reaches the large bowel ; (3) saline or plain water irrigations, one to three daily ; (4) the administration of bismuth sub-nitrate in heroic doses ; 180 gr. is given mechanically suspended in about a tumbler of plain or effervescent water every three hours, day and night in severe cases, only lessening the amount when improvement takes place. Mechanical suspension in a large quantity of water is essential. When the stools begin to decrease in number and the tongue becomes clean, the number of doses is reduced to three or four daily. In very chronic cases one or two doses daily for a month after convalescence are recommended. In exceptional cases of extreme emaciation and exhaustion, showing marked toxic symptoms, surgical treatment is necessary, and at Ancon a wide, open caecostomy is performed. The treatment of dysentery with bismuth sub-nitrate has been in use for some years at Ancon. Latterly, a combined treatment by hypodermic injections of emetine and bismuth sub-nitrate by the ' Annals Trop. Med. and ParasitoL, viii, pp. 321, 353. 39 620 THE ANIMAL PARASITES OF MAN mouth has been used, and the authorities there consider that it is better to combine the two drugs rather than use each singly. Emetine probably acts as a direct poison to the amoebae, while the bismuth probably acts by destroying the symbiotic organisms necessary for their growth. With regard to preventive measures, all drinking water should be filtered and boiled, and uncooked vegetables and salads avoided. Scrupulous care w^ith regard to personal cleanliness, and avoidance of touching the mouth or lips after contact with dysenteric patients, are essential. Isolation of parasite carriers is of great use in combat- ing and controlling outbreaks of amoebic dysentery. The pollution of soil and water must be rigorously prevented. Liver abscess due to amoebae must be localized by exploratory punctures, and then opened and drained. Intramuscular injections fo emetine hydrochloride, ^ gr. to i gr. every day, will reduce the temperature and afford relief. Oral endamoebiasis has been recently investigated by Bass and Johns, Smith and Barrett and colleagues (see pp. 43, 733). It responds to treatment with emetine, and ^ gr. of emetine hydrochloride administered hypodermically each day is of service. Rinsing the mouth with a solution of fluid extract of ipecacuanha is also useful. Rogers^ (iQiS) recommends a combined treatment of emetine and streptococcal vaccines for pyorrhoea alveolaris. II.— TRYPANOSOMIASES. The human trypanosomiases are those occurring in Africa, due to Trypanosoma ganibiense and T. rhodesiense and spread by Glossinae, and that due to T. cnizi, occurring in South America and spread by the Reduviid bugs, Triatoma spp. These trypanosomiases present different clinical features and are best dealt with separately. African Sleeping Sickness. Sleeping sickness, due to Trypanosoma gamhiense or varieties thereof, was first reported from West Africa and is now present, not only along the West Coast and in Nigeria, but throughout the Congo basin into Uganda, north of which it exists in the Bahr-el-Ghazal province of the Sudan. In Nyasaland and Rhodesia a more virulent but less widely distributed disease is produced by Trypanosoma rhodesiense. There is a general similarity between the two diseases, and the symptoms as described by the leading authorities agree in the main. The malady due to T. rhodesiense has been known only since 1910 and the differences between the malady due to it and to T. gamhiense will be indicated. ^ Ind. Med. Gazette^ April, 1915, I, p. 121. SUPPLEMENT 621 The course of the disease may be roughly divided into three stages, the incubation, the febrile or glandular, and the cerebral stage. The exact incubation period is not known with certainty in man. Probably, in most cases, it does not exceed two to three weeks, but disease signs may not appear for months. The bite of the Glossina gives rise to local irritation, which may be overlooked. The irrita- tion usually subsides in the course of a few days. The febrile, or glandular stage, is marked by attacks of fever of an intermittent type. An erythematous eruption is often found on Euro- peans. This rash begins as irregularly shaped pinkish patches which clear in the centre until a ring is produced. It may occur on any part of the body but is more frequent on the trunk. A typical symptom is the enlargement of one or more of the lymphatic glands, especially those of the neck. A general, deep hyperaesthesia, known as Kerandel's sign, may be present, and if the patient strikes a limb against any hard object, a feeling of acute pain is felt, the sensation being slightly delayed. As repeated attacks of fever increase, the patient may become anaemic. The febrile stage may last for years, and cure may be brought about at tliis phase, but frequently, after the febrile stage has lasted some time, the cerebral stage is reached. Tachycardia is also a symptom. Auto-agglutination of the red blood corpuscles is another useful characteristic, as it is said to occur rarely in other tropical diseases, but some workers doubt its value. The cerebral, or true sleeping sickness stage is marked by a great change in the habits of the victim, who becomes apathetic and dull, careless and dirty in habits, and begins to experience difficulty in walking. Tremors of varying degrees of severity are common and the gait is peculiar. There is usually fever with rise of temperature from 100° F. to 104° F. in the evening, becoming subnormal in the morning. For some days before death, it often becomes permanently subnormal. Congestion and oedema of the lungs, with patches of pneumonia, are not infrequently observed before death. The torpor gradually deepens, and the patient loses flesh. Frequently the lips swell and saliva dribbles. The patient usually becomes comatose and death ensues. Mania and delusions, and psychical and physical symptoms resembling those found in general paralysis of the insane, sometimes occur, and death may arise from secondary complications such as pneumonia or dysentery. Pathologically, the disease seems* to consist of a chronic inflam- mation of the lymphatic system. The trypanosomes reach the lymphatic glands which become inflamed, and gradually invade the blood and the cerebrospinal fluid. Sooner or later, as a result of the lymphatic disease, changes occur in the membranes and substances of the brain and spinal cord. There is round-celled perivascular infiltration of the pia-arachnoid of the brain and spinal 622 THE ANIMAL PARASITES OF MAN cord. These changes cause compression of the blood-vessels, and so lessen the supply of blood to the brain and spinal cord. Further changes in the latter organs result in the production of the symptoms that have given the disease the name of ** sleeping sickness." The disease due to Trypanosoma rhodesieiise generally runs a more rapid course than that due to T. gainbiense. The torpor and sleepi- ness may not be obvious or be very slight, and the enlargement of the lymphatic glands of the neck also may not be marked or may appear to be absent. The duration of . the disease often appears to be from three to six months. Treatment is only of use if commenced in the earlier stages of the disease. The substances of most value so far are arsenic in the form of atoxyl (introduced by Wolferstan Thomas in 1905) and antimony in the form of tartar emetic. Castellani and Chalmers and Manson recommend treatment by combining the use of both substances. The combined treatment is recommended not only because both sub- stances have been proved of service independently, but also because certain strains of trypanosomes resistant to arsenic are known, and trypanosomes can develop a resistance to arsenic. Such forms, that would not be affected by the atoxyl, are left open to attack by the antimony salt. Daniels also recommends combined arsenic and antimony treatment, and (19 15) uses atoxyl and antiluetin. Atoxyl is best given intramuscularly in 10 per cent, solution in sterile normal saline solution. Galyl is also said to have given good results. Castellani and Chalmers recommend : (i) Hanson's method of administration of atoxyl, viz., 2 to 3 gr. of atoxyl are given by intramuscular injection every third day for at least two years; or (2) Broden and Rodhain's method, 7^ gr. of atoxyl by intramuscular injection every fifth day. For the combined therapy by atoxyl and antimony they recommend the following : — '' An atoxyl injection (3 gi'-) is given every third day or 7^ gr. every fifth day, and sodio- tartrate of antimony (Plimmer's salt) is administered daily, 2 gr. dissolved in a large quantity, of water (2 pints) by the mouth or by the rectum. Tartar emetic, however, is best given by intravenous injections, using solutions of i in 100 or i in 1,000. The dose of the drug to be given is 5 to 10 eg. per injection. It is important that none of the fluid of the injection should escape into the surrounding tissues, as a violent inflammation may result. These injections should be administered monthly on ten consecutive days for a long period." Macfie and Gallagher (1914) injected 6 gr. of atoxyl intra- muscularly every week in cases infected with 7 . nlgeriense in the Eket district of Southern Nigeria. Large doses of atoxyl were often said to cause distressing results SUPPLEMENT ' 623 such as optic atrophy, and when the onset of such occurred the drug was usually discontinued. However, Daniels^ (July» iQ^S) points out that eye troubles, such as iridocyclitis, are symptoms of trypanosomiasis. - Other arsenical preparations such as soamin and arsenophenylglycin have been used, but less successfully than atoxyl. Fowler's solution, well diluted, has been given by the mouth when treatment by injection was not possible, the doses commencing with 5 minims and increasing to 15 minims. Salvarsan and neo-salvarsan have also been tried for sleeping sickness. Plimmer recommended powdered antimony suspended in sterile olive oil. Ranken used precipitated metallic antimony in normal saline solution injected intravenously. Laveran and Thiroux have recommended a combined treatment of atoxyl and an inorganic salt of arsenic such as orpiment. The orpiment is given as pills, in doses of 2 gr. of orpiment two or three times daily. Opium is added to the orpiment to prevent diarrhoea. This treatment is said to have been used in man with good results. Trypanosoma rhodesieiise seems less amenable to treatment than T . gambiense. The main preventive measures seem to he in segregation of the sick in areas not infested with Glossinae, and in measures against these flies, such as bush clearing and destruction, to some extent, of proved reservoirs in big game. South American Trypanosomiasis. The chief clinical features of the trypanosomiasis occurring ni Brazil have already been indicated (see p. 87). With regard to treatment, according to Castellani and Chalmers the indications are the same as those for African trypanosomiasis, together with treat- ment for hypothyroidism. Preventive measures are directed against the Reduviid bug, Triatoma rnegista, that transmits the disease. The bugs occur in numbers in the cracks of the houses of the poor of Minas Geraes, and may be destroyed by sulphur fumigation, lime- washing or whitewashing. III.-FLAGELLATE DIARRH(EA AND DYSENTERY. The chief causal agents are Trichomonas hominis {T. intestinal is), CJi ilomastix (Tetramitns) mesnili and allied organisms (see pp. 54 to 57), and Lamblia intestinalis (see pp. 57 to 60 and Appendix pp. 734 to 736). These parasites and the associated diarrhoeas occur in temperate as well as in warm climates. Probably some of the diarrh(jeas in India are thus caused. The same, or similar parasites occur in various Muridae, especially rats and mice, which may act as reservoirs. ' lourn. Trop. Med. and Hyg.y xviii, p. 157. 624 THE ANIMAL PARASITKS OF MAN (i) Mello-Leitao^ (1913); writing from Rio de Janeiro, states that there is a primary flagellate dysentery, due to Trichoinonas iniestinalis (Leiickart) and to Lamblia iniestinalis (Lambl), either separately or in combination. He considers it a benign disease, and the most frequent form of dysentery in young children. Trichomonas and Lamblia were found to be pathogenic to children under 3 years of age. EscomeF (191 3) collected 152 cases of dysentery in Peru due solely to Trichomonas. Examination of the reservoirs containing the water used for drinking purposes showed the presence of Tricho- monas. After the reservoirs were cleaned no more Trichomonas was found and the cases of dysentery ceased. Brumpt^ (191 2) described a colitis due to Trichomonas intestiiialis in a patient returned from Tonkin. Cases of infection by Cliiloniastix (Tetramitus) niesnili, with colitis or dysenteric symptoms, are recorded by Brumpt (1912) from France, and by Nattan-Larrier^ (191 2) from the Ivory Coast respectively. Marques da Cunha and Torres'^ (1914) describe five cases of chronic diarrhoea in Brazilian children due to the Chilomastix (Tetramitus). GabeP (1914) described a case of seasonal diarrhcjea contracted in Tunis and caused by a Tetramitid parasite which he named Difdmus tnnensis, as the discoverer considered that it lacked an undulating membrane in its large cytostome. Derrieu and Raynaud^ (i9H) I'ecord a case of chronic dysentery in Algeria due to a Trichomonad possessing an undulating membrane and five free flagella. The parasite was named Hexamastix ardin- delteilif but the generic name Hexamastix is pre-occupied. Chatterjee's Pentatrichomonas bengalensis (191 5) is possibly the same organism. Treatment. — Escomel (1913), finding ipecacuanha and calomel useless, recommends turpentine for Trichomonad dysentery. Two to 4 grm. of essence of turpentine in an emulsion are given by the mouth, and enemata containing 15 to 20 drops of turpentine emulsified in the yolk of an egg to which is added a little water and tincture of opium. Derrieu and Raynaud found this treatment effective in Algeria. Smithies^ (19 12) reports two cures of cases of severe dyspepsia, in which Trichomonads were found in the stomach contents, after administration of a single dose of 50 to 60 gr. of thymol, given at bed-time, together with 2 gr. of calomel, and followed by an ounce of Carlsbad salts in the morning. The patients came from the ' BriL /ourn. Child} en's Diseases, x, p. 60. - Bttll. Soc. Path. Exol., vi, p. 120. ' Ibid., v, p. 725. * Ibid., V, p. 495. 5 Btazil Medico, xxviii, p. 269. ^ Arch.f. Protistenkimde, xxxiv, p. i. ^ Bull. Soc. Path. Exot., vii, p. 571. ^ Amer. fottrn. Med. Sci., cxliv, p. 82. SUPPLEMENT 625 Southern United States, and had been in the habit of drinking unfiltered surface water in the locaHties in which they Hved. Mello- Leitao^ used magnesium sulphate and water or milk diet. Sometimes enemata of coUargol (i per cent.) or electrargol were required. Rosenfeld recommended calomel. Methylene blue has also been tried. Recently, EscomeP (1914) recommends enemata of an aqueous solution of iodine (i per 1,000) and farinaceous diet. Lynch^ (19 15), working in South Carolina, recommends a mouth wash of saturated solution of bicarbonate of soda three times daily in oral infections. A similar solution was used as a douche in vaginal trichomoniasis. Stiles (19 1 3) points out that when amoebae or flagellates are found in a large percentage (10 to 40, or even 60) of the members of a community, means should be taken to improve the methods for the disposal of the dejecta, so that the food-supply may be carefully protected against faecal contamination. Cysts of the parasites may be air-borne or conveyed to food on the bodies of house-flies. (ii) Lainhlia intestinalis in man may cause diarrhoea with dysenteri- form stools. The diarrhoea may be of a chronic recurrent character. The flagellate, or a variety of it, is fairly common in the digestive tract of rats and mice. Mathis^ (1914) gives an interesting account of cases in Tonkin. In a child, aged 3, the stools were at first glairy and blood-stained, containing many encysted Lamblia. The child's home was infested with mice. In another case, the house of the patient harboured numerous rats. According to Mathis, prognosis is favourable, but emetine hydro- chloride is without action on Lamblia. Prowazek and Werner^ (1914), however, state that emetine will act upon the flagellates, but not upon the cysts. They recommend uzara (two tablets, three times daily) and extract of male fern as useful in certain cases. Martin Mayer (1914) found emetine hydrochloride successful in a case in the Hamburg Seamen's Hospital, but Assmy (19 14) points out that a suitable diet and daily doses of magnesium sulphate are sufficient, in his experience, to effect an improvement, and he doubts the specific action of emetine. Escomel (1914) recommends milk diet, then calomel succeeded by castor oil. According to Noc, Lamblia may also be water-borne. Healthy carriers of Lamblia cysts are known. Food should be protected from being soiled by rats and mice. ' Brit. Journ. Children's Diseases, x, p. 60. ^ ^^^//^ Soc. Path. Exot., vii, p. 657. 3 Atner. Journ. Trop. Dis. and Prevent. Med., ii, p. 627. * Bull. Soc. Med. Chirurg. Indo-Chine, v, p. 55. 5 Beihefte z. Arch. f. Schiffs- ti. Tropen-Hyg., xviii, 5, p. 155. 626 THE ANIMAL PARASITES OF MAN lY.— LEISHMANIASES. A. Kala-azar. (i) ^^ Indian " Kala-azar cine to Leishmania donovani. Indian kala-azar due to Leishmania donovani is a very fatal disease with a rate of mortality varying from 70 to 98 per cent, of the cases. The incubation period is very variable and the early symptoms not well defined. The incubation period seems to range from three weeks to several months after exposure to infection. The onset seems to commence with a rigor and attack of irregular, remittent fever, which may show two remissions per day in a four-hourly temperature chart. Rogers considers the daily double remission almost diagnostic. The duration of this first attack is from two to six weeks. The spleen and liver enlarge, especially the former, and are painful and tender. Towards the end of the time the temperature declines and the first period of the disease ends. After this period an apyrexial interval occurs, which, after some weeks, ends in an attack of fever resembling the first. Periods of pyrexia and apyrexia alternate. Anaemia com- mences and asthenia appears and deepens steadily. The patient is now thin and wasted, the abdomen much swollen and protuberant, the ribs show clearly, the limbs are wasted and skin and tongue darker than normal. In Europeans the skin is of a remarkable earthy hue, and in natives of India darker than normal, approaching black. Intestinal disturbances, often in the form of very obstinate and intractable diarrhoea or dysenteric attacks, are common. Papular eruptions often appear, particularly on the thighs ; haemorrhages also may occur. The disease lasts for periods varying from seven months to two years, and usually ends fatally. Treatment, unfortunately, has not been very successful up to 1915. Manson has reported two cases of cure by intramuscular injections of atoxyl daily or every other day in doses of 3 gr. Rogers has advocated large doses of quinine, 60 to 90 gr. daily until the temperature falls and then 20 gr. daily. Castellani and Chalmers consider the best results are obtained by large doses of quinine given intramuscularly, supple- mented by a course of quinine cacodylate injections or atoxyl injec- tions. Tartar emetic should be tried (see pp. 627, 629), especially as L. Rogers (July, 191 5) has had promising results in ten cases. Castellani (1914) and Mackie (1915), have also had successful results. Leishman states that the administration of red bone-marrow, either raw or in the form of tablets, may be beneficial. Good nursing and careful diet are essential, and diarrhoea or dysentery must receive the appropriate treatment. With regard to preventive measures, the extermination of bugs SUPPLEMENT 627 and other biting insects seems to be of most service. Domestic and personal cleanliness is of great importance. Patients should be segregated. It would probably be as well if houses in which many cases of kala-azar occurred were destroyed. Dodds Price, in Assam tea gardens, moves the coolie lines 300 to 800 yards from old infected ones, with satisfactory results. (ii) Infantile Kala-azar due to Leishmania infantum. This malady is found among children, rarely in adults, along the Mediterranean littoral. The disease commences insidiously and is often unrecognized until some intestinal disturbance occurs. The spleen is then found to be somewhat enlarged, and the case has often been regarded as one of malaria. The child becomes anaemic, suffers from diarrhoea, alternat- ing with constipation, and has attacks of irregular fever. The spleen continues to enlarge and protrudes from under the cover of the ribs. Haemorrhages from the nose and gums and into the skin occur. Anaemia and wasting set in. The abdomen then becomes very enlarged. The child becomes much less active both physically and mentally, and looks prematurely old. Death often occurs from exhaustion, though some cases of spontaneous recovery are known. Treatment up till recently has been unsatisfactory. Some of the remedies tried, as quoted by Castellani and Chalmers, are 15 eg. doses of atoxyl, benzoate of mercury (2 to 4 mg. as a daily injection), thiarsol (5 to 15 mg. by subcutaneous injection), salvarsan, etc. Recently Cristina and Caronia (1915)^ have given repeated intravenous injections of i per cent, aqueous solution of tartar emetic, the dose varying from 2 to 10 eg. The treatment in various cases has lasted from 15 to 40 days. Prophylactic measures seem to lie in the destruction of infected dogs and diminishing the breeding of fleas (see p. in). B. Oriental Sore, due to Leishmania tropica. Oriental sore, known under many other names (see p. 107), is a local infection of the skin due io Leishmania tropica. The incubation period varies from a few days to some weeks, or even months, and then one or several small itching papules appear. Each spot becomes red and shotty, the papules increase slowly in size and the surface becomes covered with papery scales. After a variable time, usually not exceeding three to four months, ulceration occurs and a yellowish secretion is exuded that soon dries into a scab. Under the scab ulceration continues by erosion of the edges, and subsidiary sores arise around the parent ulcer and usually fuse with it. Healing com- mences after six to twelve months. Granulation begins at the centre lull. Soc. Path. Exot.y viii, p. 63. 628 THE ANIMAL PARASITES OF MAN and spreads outwards, and when healing is complete, a depressed, whitish or pinkish scar remains. Many treatments for Oriental sore have been devised but do not seem particularly satisfactory. Castellani and Chalmers state that the scabs should be removed by boracic acid fomentations, and the ulcers thoroughly disinfected once or twice daily with a i per i,ooo solution of perchloride of mercury, after which an ordinary antiseptic ointment is applied. The use of permanganate of potash has been advocated both by French and English doctors. Both large and small sores can be treated. The patient's skin around the sore is protected by a thick layer of vaseline, and the surface of the ulcer powdered with potassium permanganate, which is kept in position by a pad of gauze and a bandage. The treatment is said to cause great pain for six to eight hours, but at the most, three treatments are necessary before the sore becomes a simple ulcer, well on the way to healing. The perman- ganate may also be used in ointment. Excision of the ulcer when small is advisable when the site of the ulcer permits of this. According to Manson, reports on treatment by radium, salvarsan and carbon dioxide snow are decidedly promising. Mitchell (1914)^ reports favourably on the use of carbon dioxide snow in the form of a pencil, in India. In Brazil several workers (1914) record successful results from the intravenous injection of a i per cent, solution of tartar emetic in distilled water. Low (191 5) has successfully treated a case by •direct local application of tartar emetic. Row (191 2) has treated cases of Oriental sore by inoculation of killed cultures of the causal organism. As the disease is very contagious, the slightest wound, and any insect bite, should be thoroughly disinfected with 5 per cent, carbolic acid or iodine. Destruction of bugs, lice, and other biting insects should be enforced. As dogs may contract the disease (see p. 108), it is well not to allow them in the house and not to encourage undue contact with them. Naso-oral Leishmaniasis {Espundia) due to Leishmania tropica. This form of Leishmaniasis has been reported from South America and recently by Christopherson^ {1914) fi'oi^i the Sudan. In South America it is often called Espundia, also Buba and Forestal Leish- maniasis. The primary lesion is found usually on the forearms, legs, chest or trunk. This ulcer is of the Oriental sore type, and after some months, or even as long as two years, heals up, leaving a thick scar. While the ulcer is open, or more often after it has healed, lesions appear on the mucosa of the mouth and nose. The hard and ' Journ. Roy. Army Med. Corps, xxiii, pp. 440-446 (see Trap. Dis. BiilL^v, No. 5, p. 276). "^ Annals Tiop. Med. and Paras itoL, viii, p. 485. SUPPLEMENT 629 soft palate, gums and lips all may be attacked. The mucosa of the nose is usually attacked and the cartilages become destroyed, produc- ing great deformity. In bad cases the pharynx and larynx may become infected. Till recently it was believed that treatment was of little use unless the case could be investigated early. Escomel considered that if the primary cutaneous lesion w^as excised or destroyed, further progress of the disease was prevented. When lesions have appeared on the mucosa of the mouth or nose, little could be done. The ulcers might be cauterized and mild antiseptic mouth washes used. In 1913 Vianna, working in Brazil, introduced treatment by tartar emetic, which is now becoming more wadely known and proving efficacious. Carini^ (1914) applies it thus. Tartar emetic (that is, potassium antimonyl tartrate) in i per cent, aqueous solution is intro- duced slowly into a vein, such as the vein at the bend of the elbow, in doses of 5 to 10 c.c. daily or on alternate days according to the tolerance of the patient to the drug. Eighteen to forty injections have been used. In some of the memoirs on the subject, the drug is referred to as antimony tartrate. The course of the disease is chronic and may last for twenty to thirty years, death usually resulting from some intercurrent disease. At present the actual transmitter of Espundia is not known with certainty. Various sand-flies (Simulidae) have been suspected of transmitting the disease, though so far proof is wanting. It has also been suggested that the natural food sources of some Simulidae known to bite man, namely, certain snakes^ and lizards,^ are possible reservoirs of the disease. Prophylactic measures would seem to consist in the immediate disinfection of insect bites by tincture of iodine, and by avoidance of areas known to be infested with snakes and lizards, and insects that prey on them and man indifferently. The destruction of the primary lesion as soon as detected is essential, and the isolation of advanced cases of the disease seems advisable. Y.— SPIROCHiETOSES. A. Relapsing Fevers. The relapsing fevers of Europe and of America, due to Spirochceta recur reiitis and S. novyi (probably a race of S. recurrent is), present much the same symptoms, w^hich differ in some respects from those • Bu//. Soc. Path. Exot., vii, p. 277. 2 Lindsay (1914), Trans. Soc. Trop. Med. and Hyg., vii, p. 259. ' Sergent (Ed. and Et.), Lemaire and Senevet (1914), Btdl. Soc. Path. Exot.y vii, P- 577. 630 THE ANIMAL PARASITES OF MAN due to S. duitoni, the excitant of '' tick " or '' relapsing " fever in Africa (see pp. 1 16-122). The incubation period of S. recnrrentis varies from two to twelve days, during which time a very slight indisposition may be noticed. The onset is usually sudden, with severe headache, pains in the back, limbs and stomach and a feeling of weakness. There is a rise of temperature to 103° F. or 104° F., and the temperature continues high till about the sixth or seventh day. The skin is yellowish, hot and damp ; a rash, disappearing on pressure, may occur on the trunk and legs, nausea is always present and thirst is usual. The liver and spleen both enlarge. The number of respirations and pulse-rate become increased. On the sixth or seventh day a crisis occurs. There is violent perspiration, with a rapid fall of tempera- ture, pulse and respiration become normal and the patient sleeps and awakes better. Improvement continues for some days, and recovery may ensue, but usually about the fourteenth day relapse occurs, lasting usually three or four days. A second relapse is unusual. Numerous complications are known, e.g., bronchitis, pneumonia, diarrhoea and dysentery. With regard to treatment, the specific appears to be salvarsan. Castellani and Chalmers recommend salvarsan administered intra- venously. Intramuscular inoculations (for example, into the buttock) of a suspension of " 606 " in oil can also be given. The drug is very efficacious, but large doses should not be given. An intravenous injection of 4 or 5 gr. does not give rise to unpleasant symptoms but is sufficient to effect a cure. The incubation period for the American form of the disease is at least five to seven days, and the first attack lasts about five to six days. The treatment is by salvarsan as detailed previously. As relapsing fever is spread by body lice and possibly by bugs, preventive measures are directed against these insects. Strict cleanliness of person, clothing, bedding and dwellings is essential. Furniture, e.g., wooden bedsteads, liable to harbour such insects should not be used. The principal and best-known relapsing fever of Africa is that excited by Spirochcvta dutioni, and transmitted to man by ticks, chiefly Ornithodorus monbata. The incubation period is usually about seven days but may be longer. The patient is dull and lethargic, perspires freely and is often constipated. The temperature rises to 103° F. or 105° F., there is headache, pains in the back and limbs, general chilliness and great pain in the region of the spleen, which often enlarges. The symptoms become w^orse, there is a fall of temperature with improvement in the morning, and a rise, with increase of pain, in the evening. Spirochaetes are now found in SUPPLEMENT 63 1 the blood in greater numbers. The symptoms last three to four days and end in a crisis with profuse sweating and fall of temperature below normal. The day before the crisis there is a pseudo-crisis, when the temperature falls but there is no improvement. The patient is left weak and tired. Recovery may follow, but more usually a relapse occurs. The intermission period varies ; five to eight days is common. The symptoms of the relapses are like those of the first attack. The number of relapses varies, five to eleven may occur. The treatment recommended is by salvarsan, as for the European relapsing fever. With regard to prophylaxis, localities w^here ticks abound must be avoided and the parasites themselves destroyed. Native huts should be avoided. Mosquito nets, a bed well off the ground and the use of night lights are advised by Manson to avoid attacks by ticks, which are often nocturnal in their habits. In North Africa (Algeria, Tunis, Tripoli, Egypt), and sometimes in the Anglo-Egyptian Sudan, a spirochaetosis due to 6>. berhera occurs. According to Castellani and Chalmers, the incubation period v^aries somewhat. The fever reaches its height during the first twenty- four hours, and afterwards shows a morning remission. Jaundice is often absent, but there may be hepatic tenderness and splenic enlargement. One or two relapses usually occur. The treatment is on the same lines as for the other spirochaetal fevers. Sergent and Gillot^ (1911)5 working at the Institut Pasteur of Algeria, have had good results by using injections of salvarsan in doses of 075 to T"o eg. per kilogramme weight of the patient. The prophylactic measures are directed against lice and other biting insects. Personal cleanliness is most necessary. In Asia, a relapsing fever, due to the spirochaete named S. carteri by Manson in 1907, producing a mortality of about 18 per cent., occurs. The symptoms have a general resemblance to those produced by S. recurreiitls, but on the fall of temperature to subnormal on the sixth or seventh day, when profuse perspiration and polyuria occur, instead of improvement following, the patient often becomes collapsed, with a clammy skin and feeble pulse. Improvement is slow. The first relapse occurs about the fourteenth day of the attack, when the temperature may be higher than for the first attack. There are seldom more than four relapses. The treatment is by salvarsan, of which doses of not more than 5 gr. intravenously should be given. Sudden heart failure being common, Castellani and Chalmers state that cardiac stimulants should be given. Prophylaxis is the same as for European relapsing fever. * Bu//. Soc. Path. Exoi., iv, p. 440. 632 THE ANIMAL PARASITES OF MAN B. Yaws or Framboesia tropica. Yaws is essentially a tropical disease, though it is found in the tropical and subtropical zones in all parts of the world, except in the mountains and cold districts. In 1905, Castellani found the causal organism, Treponema pertenue (sometimes called Spirochccta pertennis) (see p. 127). The disease shows three periods : (i) The primary stage, consisting of the development of the primary lesion or papule, which is usually extragenital. The papule dries into a crust beneath which an ulcer lies. (2) The secondar}^ or granulomatous stage, which com- mences from one to three months after the primary lesion is first seen. It consists of a general eruption of small papules, some of which enlarge and become granulomatous nodules covered with a yellowish crust. They are common on the limbs and face. (3) The tertiary stage, in which deep ulcerations and gummatous nodules appear. Any of the tissues may be involved. Osseous lesions may occur. The disease does not appear to be hereditary ; it is usually spread by contact. The best treatment appears to be by salvarsan or neo-salvarsan. Castellani and Chalmers recommend intramuscular and intravenous injections. For intramuscular injection an alkaline or neutral solu- tion of the drug is preferable, or a suspension of the drug in oil may be used. The dose varies from o'3 to 0*5 grm., according to the age and sex of the patient. For use intravenously, a slightly smaller dose is required. Galyl is also being used. In countries where frambcesia is endemic, slight skin abrasions should be carefully treated with antiseptics. Yaws patients should be isolated till cured, and their dwellings and personal possessions disinfected. C. Syphilis. Syphilis, due to Treponema pallidum (sometimes called Spiroduvta pallida), is prevalent throughout the tropics as well as in temperate zones. The disease is amenable to treatment by salvarsan and neo- salvarsan, for administration of which see relapsing fever and yaw^s. Galyl is also being used with favourable results. Lambkin's mercury cream has been found useful in treating numerous cases in Uganda. The life-history of the parasite is given on p. 124, and further medical details hardly come within the purview of this book. D. Bronchial Spirochaetosis. Bronchial spirochaetosis, due to Spirochceta hronchialis (see pp. 122, 739) is probably of wide distribution in the tropics. The spirochaetes have been found in cases of chest complaints, especially those with bronchitic symptoms. The disease may be suspected in atypical cases of pneumonia and bronchitis, and may be mistaken for incipient phthisis. SUPPLEMENT 633 Chalmers and O'Farrell^ (1913); writing from Khartoum, re- commended rest in bed, good food and ventilation, coupled with treatment by arsenic in some form, preferably associated with glycero- phosphates. These may be given by the mouth, or intramuscularly as an injection of : — Sodium cinnamate ... ... ... ... 0*05 grm. Sodium cacodylate ... ... ... o'lo ,,' Sodiutn glycerophosphate ... ... ... o'lo ,, Taylor^ (1913-14), writing from Entebbe, Uganda, prescribes arsenious acid by the mouth in increasing doses. Creosote has been used in West Africa. YL— MALARIA. Malaria, known also under the names of ague, paludism, marsh fever, remittent fever, intermittent fever and climatic fever, among others, is a very widely spread disease. It is most prevalent in the equatorial regions and gradually diminishes north and south of the equator. The various malarial parasites (see pp. 155 to 172) are spread by species of Anophelines, and hence malaria is present in districts favourable to these intermediate hosts, that is, in places where there is a considerable amount of atmospheric moisture and rain, as well as heat. The principal malarial parasites are : Plasmodium vivax, the agent of simple tertian fever ; Plasmodium uialarice, the parasite of quartan malaria, and Laverania malarice or Plasmodium falciparum, producing malignant tertian or sub-tertian malaria (and quotidian, see p. 167). These various malarial fevers present certain clinical features in common, which will be stated here (see also pp. 155 to 157). For further particulars regarding malaria in all its aspects the reader is referred to the book by Sir Ronald Ross on "The Prevention of Malaria," to the ** Manual of Tropical Medicine," by Drs. Castellani and Chalmers, and to the ** Tropical Diseases " of Sir Patrick Manson. Typical malarial fevers consist of a series of pyrexial attacks which recur at definite intervals of twenty-four (quotidian), forty-eight or seventy-two hours, according to the parasite present in the patient's blood. Each attack shows three stages, a stage of rigor, a heat stage and a stage of profuse perspiration. Following on these three stages, there is an interval relatively or actually without pyrexia. Then the fever returns again. A rise of temperature, often accompanied by a general feeling of malaise, may precede the initial stage of rigor. When the latter sets in, the patient feels intensely cold, shivers violently, the skin becomes cold and the features pinched. There may be violent ' Journ. Trop. Med. and Hyg.^ xvi, p. 329. ^ Annual Med, and Sanii. Kept., Uganda^ for 1913, p. 80. 634 THE ANIMAL PARASITES OF MAN vomiting and convulsive attacks in young children. The temperature, however, is really above the normal, and continues to rise. After about an hour, the shivering abates and the heat stage succeeds it. The temperature rises rapidly, even to 106° F. The patient becomes very flushed, the pulse is rapid, headache may be intense and the skin dry and burning. This stage, that causes acute distress to the patient, may last for one or often three to four hours, and then the patient commences to perspire profusely, the clothing and bedding often being saturated with sweat. After this, the fever rapidly declines, and when the sweating ceases, the patient may feel almost well although somewhat languid. The sweating stage persists from two to four hours, so that the attack lasts as a rule from six to ten hours. After an interval of one, two or three days, a recurrence takes place. During the early part of the attack, especially at the stage of rigor, there is great splenic enlargement. At first the enlargement disappears in the interval, but in the case of repeated attacks the spleen tends to become permanently enlarged. During malarial attacks and during the intermission period, there is a great increase in the amount of nitrogen excreted by the kidneys, while the excretion of iron and bile in the faeces is increased. Stitt^ (1914) points out that it is characteristic of malignant tertian paroxysms that they set in with chilly sensations rather than a frank, definite chill, and that the fever is of the remittent type. Plasmodium malarice and P. %'ivax rarely produce marked lesions in the bodies of their hosts, as they sporulate in the circulating blood and so do not accumulate in any one organ. On the other hand, Laverania malarial (Plasmodium falciparum) multiplies within the internal organs of its host, and consequently aggregates or clusters of the pai-asites occur therein. The organ in which most sporulation occurs suffers most. The liver is generally enlarged, soft and con- gested. The capsule of the spleen is tense, but the splenic consistency is less than normal. The bone-marrow is often dark and congested in the spongy bones and brownish-red in long bones. The blood- capillaries of the brain and spinal cord are often filled or blocked with sporulating parasites and large quantities of pigment are found in these organs. Even if the parasites are absent, the pigment is present in the endothelial cells. Pigment is found in most organs of the body. Atypical forms of malaria may occur in v/hich some or all of the symptoms are much modified. Irregular fevers also may be produced by successive infections by the same parasite, or by the presence of two different malarial parasites. ' '* The Diagnostics and Treatment of Tropical Diseases." London : H. K. Lewis. SUPPLEMENT 635 As regards the diagnosis of malaria, according to Manson the three pathognomonic signs are — periodicity, the effect of quinine, and the presence of the malarial parasite. Trcatineiit. — The great specific for malaria is quinine. It attacks the merozoites or asexual generation. The drug can be adminis- teied by the mouth, by the rectum, by intramuscular injections or by intravenous injections, the two latter methods being adopted in serious infections or where gastric complications are present. When quinine is taken by the mouth, the more soluble acid salts, e.^., quinine bihydrochloride and bisulphate, are better than the sulphate, the form in which quinine is usually sold. Tablets, pills and cap- sules are convenient means of taking quinine but must not be old or hard, or they may pass unchanged through the body. In the case of mild tertian or quartan malaria, Castellani and Chalmers recommend the administration of a dose of quinine four hours before the sporulation of the parasite is due. Another modification is to give 10 gr. of quinine by the mouth in the morning and a second dose of 10 gr, as above. In many cases they give 5 to 10 gr. of the drug three times a day. Administration of qumine per reciiim may be useful but they recommend intramuscular inocula- tion. The solutions used must be sterile, and the '^ sterilettes,'' small, hermetically sealed vials, containing i grm. (15 gr.) or \ grm. (72 &') o^ quinine in solution, are recommended. A deep injection into the deltoid or gluteus muscle is usual. For pernicious infections, intravenous inoculation with not less than I grm. at a time is recommended. After the fever has subsided, the administration of quinine in smaller doses must be continued for some time, in order to avoid relapses. Stitt (19 14) writes that '' there now seems to be a tendency to use the alkaloid itself instead of its salts, it having been found that the alkaloid and its very insoluble tannate are absorbed from the digestive tract equally as well as the soluble salts." Euquinine or ethylcar- bonate of quinine contains 81 per cent, of quinine, but is expensive. During malarial attacks, constipation must not be allowed. Headache can be relieved by cold applications, and perspiration must be encouraged in the early stage by hot tea, warm lime drinks, etc. After bad attacks, a change to a cooler climate is desirable, but the quinine treatment must not be discontinued. Preventive measures take two main forms, directed respectively against the malarial parasites in man, and against the mosquitoes that convey the parasite from man to man. With regard to man, houses should be built away from low- lying marshy ground, and kept free from vegetation such as grass or 40 636 THE ANIMAL PARASITES OF MAN brush which furnishes shelter to the mosquitoes. In the tropics, the chief reservoirs of the malarial parasites are the native children, hence European quarters should be away from native dwellings as far as possible. Mosquito nets, having tw^enty to twenty-four meshes per square inch, should be used invariabty, and houses should be screened. Malaria-conveying mosquitoes bite chiefly towards evening. Quinine treatment for preventive purposes is important. A dose of 5 gr. of quinine daily, with a dose of 10 gr. on the seventh day (Castellani), is efiicacious. Some workers, however, recommend a large dose (15 gr.) on two consecutive days every eight or ten days for three months, while others recommend 10 gr. twice a week. Celli administered 3 gr. of quinine morning and evening. The second line of attack is directed against mosquitoes, especially Anophelines, on the lines so well set forth by Sir Ronald Ross.^ The accumulation of small quantities of water in various vessels, many of them unnecessary, should be prevented, as Stegomyia (Culicines) breed in such receptacles. Anophelines breed in small pools. All drinking water and household vessels, water-butts and cisterns must be effectively screened with wire gauze. Cesspools, etc., must also be screened, and they, and all collections of water, should be oiled with crude petroleum sprays every week or ten days, or fortnight according to some workers. The petroleum is a good larvicide and suffocates the Anopheline larvae, while its presence renders the site obnoxious to the adult mosquitoes. The amount of crude petroleum or kerosene will vary according to the locality concerned, due regard being paid to its powers of spreading on the surface treated. Different authorities have used different quantities, such as i oz. of oil to i square yard or to 15 square feet. Others have used i pint of the petroleum to a circle of 20 feet in diameter, while ^ pint for every 100 square feet of surface has also been recommended. The larvicide used so successfully in Panama consisted of : — Average mixture Crude carbolic acid {containing 15 per cent, phenol) ... 300 j;allons Caustic soda ... ... ... ... ... 30 lb. Resin ... ... .,, ... ... ... 200 ib. One part of this mixture in 5,000 parts of water containing mosquito larvae destroys them within five minutes ; i part in 8,000 of water kills larvae in thirty minutes. Small fish, such as the ''millions" fish, that feed on the larvae, can be introduced into collections of water and are of local service. Ducks may also act as destroyers of larvae. The growth of water-weeds and rank vegetation, tliat affords shelter to the larvae, must be prevented as far as possible. Wherever possible hollows should be filled up, swamps and roads ' " The Preven'irn of Malara." Second Er iticn (191 1). London : John Murray. SUPPLEMENT 637 should be well drained. Much good has followed the use of such measures in Panama, Egypt, British Guiana and other places. The ideal conditions for malaria reduction appear to consist in a combina- tion of general quinine prophylaxis with anti-mosquito measures. YII.— BALANTIDIAN DYSENTERY. This disease is also known as ciliate or ciliary dysentery. The chief causal agent is Balantidinin coll. Others are Balantidiuin niinutnin, Nyctof herns fciba, etc. (see pp. 200-206). Balantidiasis is insidious and is marked by alternate attacks of diarrhoea and constipation with vomiting, while mucus is passed in the motions, which are foul smelling. There may be chronic ulcera- tion of the colon. (Edema of the face and limbs and ana3mia may occur. Treatment is at present rather unsatisfactory. Castellani and Chalmers state that ''the symptomatic treatment for entamoebic dysentery may be tried." Various treatments, more or less empiri- cal, by calomel, quinine, carbolic acid in pill form, salicylic acid, extract of male fern, methylene blue, iodine solution, rice water and tannin enemata are mentioned by Prowazeld (191 3) and by Seifert. E. L. Walker^ (1913) found, from experimental work, that organic compounds of silver, e.g., protargol, were most effective. Local treat- ment by large enemata of collargol or protargol seems to be indicated. Behrenroth^ (19^3) successfully treated a Prussian case with thymol, given in 4 grm. doses every two days, followed at the end of a fortnight by de-emetinized ipecacuanha, given in pills containing 6 eg. each, to the number of thirty a day. In about another fort- night the symptoms had subsided. The thymol checked the diarrhoea, but it was necessary to give the de-emetinized ipecacuanha to kill off the balantidia still present, Phillips (1915) also recom- mends thymol. Ardin-Delteil, Raynaud, Coudray and Derrieu (1914) found neither emetine hydrochloride nor protargol of use. As regards prophylaxis Walker states that pigs '* should be confined and not allowed to run in yards and dwellings." Behren- roth considers that dirty hands, for example, those of farm workers brought into contact with pigs, are probably the medium of infec- tion. The personal cleanliness of such persons is, then, of the greatest importance. i ' Beihefte z. Arch. f. Schiffs- u. Tropeu-Hyg.^ xvii, 6, p. 371. 2 Pkilippiiie Jl. Sc, Sect. B, viii, pp. 1-15, 333-349- ^ A'ch. f. Verdattuu^s Krankhdten^ xix, p. 42. 638 THE ANIMAL PARASITES OF MAN PLATHELMINTHES (Flat Worms) BY J. W. W. STEPHENS, M.D., B.C., D.P.H. FASCIOLIASIS. Fasclola hepatica. The symptoms of disease evoked by Fasclola hepatica are rarely observed in our part of the world, whereas Kermogant^ states them, to be of frequent occurrence in Tonkin^ ; the parasites are there called " Douv-es." In our experience they are only accidentally found post inortein in a certain number of cases, as no changes are manifested during life which would permit of any conclusion being drawn as to the presence of these parasites. In three cases (Bierner,^ Bostroem^ and Sagarra^) icterus was present ; in a fourth case, recorded by Duffek/ the parasites had led to a severe and acute distomiasis of the liver, combined with chronic purulent and ulcerative cholecystitis^ with purulent cholangitis and dilation of the bile-ducts and numerous small abscesses of the liver. The total number of flukes found in these cases amounted to about fifty. The parasites passed from the duodenum into the bile-ducts, and first obstructed the flow of bile and then set up icterus, followed by cholecystitis and cholangitis. As regards localization of the liver fluke in the pharynx, see p. 242. The treatment must be directed to the principal symptoms; prophylaxis is especially important in districts where distomiasis is of frequent occurrence. As the embryos live in water, only boiled or filtered water should be drunk. The attempts of Tappeiner^ to discover an effective remedy against liver-fluke disease (liver rot), so prevalent among sheep, were unsuccessful. Fasciolopsis buski. This parasite lives in the intestine, not in the liver of man ; it produces bloody stools and typical symptoms — high fever and a condition of apathy (Odhner).^ ' Kermogant, Soc. vied, des HSp., February 7, 1905. ^ [The distomiasis of Tonkin is due to Clonorchis sinensis find not to /'. hepatica. — J. W. W. S.] 3 Bierner, Schwtiz, Z.eitschr. f. Heilk.^ 1863. --* Bostroem, Deutsch. Arch.f. Jdiu. Med., 1883. ^ Sagarra, quoted by Dufiek. ^ Dutifek, Wiev, /Jin. IVcchenschr., 1902, xxx. ■^ Tappeirer, JMiinch. vied. Wccheinchr.^ 1900, I. « Oeilu er, CeuhaliLf. Bald., 1502, xxxi. SUPPLEMENT 639 PARAGONIMIASIS. Paragonimus ringeri. The disease produced by the lung fluke is specially endemic in Japan, also in isolated parts of China, Formosa and Korea. The fact that the lung-fluke disease is most frequently found in mountain- ous districts (Katsurada') is worthy of special attention. The onset of pulmonary paragonimiasis is generally insidious (Looss^) ; generally the only symptom is a slight cough, occurring at first at longer, and later at shorter intervals ; it is accompanied by the expectoration of discoloured sputum, frequently blood-stained. Though now and then severe haemorrhages result, up to the present no case has been established in which they have been the direct cause of death. Examination of the thorax frequently fails to reveal anvthinr^ This section, except for minor corrections, is practically a translation of the original. To Binder : face p. 638. ^yinpioms in Tnese tracts. ' The most dangerous locality is in the brain. Otani,''^ Inouye,^ Yamagiva,^ and recently also Taniguchi,^ have found post mortem the worms and their ova in tumours of the brain, or^ in areas of softening in cases of Jacksonian epilepsy ; in Taniguchi's case the eggs were found in masses in the inflammatory areas of softening. In the nineteen cases of paragonimiasis of the brain collected by Inouye, the following symptoms were observed : general convulsions on eight occasions, unilateral convulsions on six occasions, convul- sions with paralysis on the same side and hemiplegia, five times each ; ' Katsurada, Ziegler^s Beitr. z. path. Anat., igoo, xxviii. ^ Looss, " Handb. d. Tropenkrankh.," von Mense, iQc;, i. ^ Inouye, quoted by Looss. ^ Scheube, " Die Krankh. d. warm. Lander," 1896. ^ Otani, quoted by Looss. ^ Inouye, quoted by Looss. "^ Yamagiva, quoted by Looss ** Taniguchi, Arch. f. Psych, ti. Nervenkrankh.., xxxviii. 638 THE ANIMAL PARASITES OF MAN PLATHELMINTHES (Fiat Worms) BY J. W. \V. STEPHENS, M.D., B.C., D.P.H. FASCIOLIASIS. Fasciola hepatica. The symptoms of disease evoked by Fasciola Jiepatica are rarely observed in our part of the world, whereas Kermogant^ states them to be of frequent occurrence in Tonkin- ; the parasites are there called "Douves." In our experience they are only accidentally r. .1 L .^j. v._. ;,, ,, ^^,-fain nnmhprof cascs. as uo chauges are prevalent among sheep, were unsuccessful. Fasciolopsis buski. This parasite lives in the intestine, not in the Hver of man ; it produces bloody stools and typical symptoms — high fever and a condition of apathy (Odhner).^ ' Kermogant, Soc. vied, des HSp., February 7, 1905. ■2 [The distomiasis of Tonkin is due to ClonorcJiis sinensis nnd not to /". hepatica. — J. W. W. S.] ^ Bierner, Schweiz. Zeitschr. f. Heilk.^ 1863. '1 Bostroem, Deutsch. Arch. f. kliu. Med., 1883. ■'• Sagarra, quoted by Dufilek. ^ Dutifek, Wien. klin, IVcchenschr., 1902, xxx. "^ Tappeii er, A flinch, mcd. IVcchenschr. , 1900, 1. * Oiihi er, Ceiih all I. f. Bait., 1502, xxxi. SUPPLEMENT 639 PARAGONIMIASIS. Paragonimus ringerl. The disease produced by the lung fluke is specially endemic in Japan, also in isolated parts of China, Formosa and Korea. The fact that the lung-fluke disease is most frequently found in mountain- ous districts (Katsurada^) is worthy of special attention. The onset of pulmonary paragonimiasis is generally insidious (Looss^) ; generally the only symptom is a slight cough, occurring at first at longer, and later at shorter intervals ; it is accompanied by the expectoration of discoloured sputum, frequently blood-stained. Though now and then severe haemorrhages result, up to the present no case has been established in which they have been the direct cause of death. Examination of the thorax frequently fails to reveal anything abnormal. Inouye^ states that the most frequently observed changes consist in retraction of the thorax and in a contraction of its infra- scapular portion. Scheube* repeatedly observed that the one side, presumably that which harboured the worm, moved less freely than the other. The physical changes are not uniformly spread over the whole lung, but are localized. The disease may come to a standstill for long intervals and then set in again, lasting on the whole from ten to twenty years. In addition to paragonimiasis of the lungs, cysts are frequently found on the eyelids, which occasionally extend deeply into the orbit and hinder the movements of the eyes. Post mortemj cysts the size of hazel nuts containing one, two, or three adult worms are found in the lungs, and in addition, not uncommonly there exist pulmonary emphysema and bronchiectasis. Besides being present in the lungs and in the eyelids, the parasites have also been found in the pleura, the liver, the intestinal wall, the peritoneum, the cervical glands, and in the scrotum, without actually occasioning any actual symptoms in these tracts. The most dangerous locality is in the brain. Otani,-'* Inouye,^ Yamagiva,^ and recently also Taniguchi,^ have found post mortem the w^orms and their ova in tumours of the brain, or, in areas of softening in cases of Jacksonian epilepsy ; in Taniguchi's case the eggs were found in masses in the inflammatory areas of softening. In the nineteen cases of paragonimiasis of the brain collected by Inouye, the following symptoms were observed : general convulsions on eight occasions, unilateral convulsions on six occasions, convul- sions with paralysis on the same side and hemiplegia, five times each ; ' Katsurada, Ziegler's Beitr. z. path. Anat., 1900, xxviii. ■2 Looss, " Handb. d. Tropenkrankh.," von Mense, 190S, i. ^ Inouye, quoted by Looss. "* Scheube, " Die Krankh. d. warm. Lander," 1896. ' Otani, quoted by Looss. ^ Inouye, quoted by Looss. ^ Yamagiva, quoted by Looss ^ Taniguchi, Arch. f. Psych, u. Nervenkrmikh.^ xxxviii. 640 THE ANIMAL PARASITES OF MAN in Taniguchi's case, attacks of cortical epilepsy, choreiform twitchings in the right extremities, which gradually become athetotic. The following were symptoms of rarer occurrence : paresis of the right upper extremity, vertigo, dementia, and amnesic aphasia, disturb- ances of vision. Paragonimiasis of the brain appears to arise by embolism from a primary pulmonary lesion. The diagnosis depends upon the finding of ova in the sputa ; if together with ova in the sputa, cerebral disturbances make their appearance, in all probabihty the cause is the presence of woruis or ova in the brain. The prognosis of pulmonary paragonimiasis is favourable ; on the other hand, that of cerebral paragonimiasis is very doubtful. The treatment of the pulmonary lesion consists only in paying attention to the general condition (good food, rest, cough remedies), as all attempts to destroy the w^orms in the lungs by means of vermi- cidal drugs administered internally or by way of inhalation have so far been without result. The treatment of the cerebral lesion is entirely hopeless. Trephining has been proposed for cases the con- dition of which is more favourable, but it has not reached the stage of performance. Prophylaxis consists in general management : cleansing and if need be boiling of everything that is eaten or drunk. Clonorchls sinensis. According to our present knowledge Cloiiorcliis sinensis is only found in China and Japan ; even the post-mortem case reported by Laspeyres^ was that of an Asiatic sailor who was admitted into the General Hospital St. George, Hamburg, in a moribund condition with the clinical diagnosis of beri-beri. The bile-ducts are the usual site of the parasite, though Katsurada- has found them also in the pancreatic ducts. In addition, it is found not uncommonly in the upper portion of the small intestine, especially in the duodenum, also, though decidedly rarely, in the stomach. As these sites, however, do not afford the conditions necessary to life, they are only found here on their way out of the body of the host. The initial stage of infection with this fluke generally runs a symptomless course ; in proportion as the worms multiply the following symptoms are manifested : First there is a morbid sense of hunger and irregularity in defaecation ; at the same time the patient experiences a feeling of pressure and pain in the epigastrium and right hypochondrium, or just a dull pain. Pressure increases the pain considerably. The liver appears to be enlarged, sometimes the ^ Laspeyres, " Dissert. Kiel," 1904. '^ Katsurada, Ziegler's Beitr. z. path. Anat.^ 1900, xxviii. SUPPLEMENT 64 1 enlargement is specially perceptible over the left lobe of the liver. The patients maintain a proportionately good general state of liealth in this state for a long time and may hope to recover. In severe cases there occurs copious and generally bloody diarrhoea, also icterus. The next stages are anaemia, emaciation, epistaxis, ascites, enlarged spleen, and cachexia, to which the patient finally succumbs. In general the course of the disease is very chronic and irregular; m winter and spring there is generally improvement, in the summer and autumn the patient gets worse. At post-niorteni the bile-ducts are enlarged and thickened, there is interstitial hepatitis with enlargement of the liver, but not to such an extent as in hypertrophic cirrhosis. After the initial enlargement contraction of the liver sets in, the peritoneal coat and capsule proper of the liver become more or less thickened in places. In the pancreas also dilatation and thickening of the ducts occur, as well as interstitial inflammatory processes. Obstructions in the portal circulation may lead to catarrhal changes in the stomach. The diagnosis is based on the demonstration of ova in the faeces. As a radical treatment is still unknown, consequently it can only be purely symptomatic. Prophylaxis consists in the prohibition of drinking unboiled water or eating uncooked molluscs, fish, etc., of canal water. Leaving the epidemic region may bring about gradual recovery. BILHARZIASIS. Schistosoma haematobium. The symptoms of bilharziasis are manifested chiefly in the urinary apparatus, and above all as haematuria, at the outset without any special troubles. Later, however, it is accompanied by subjective symptoms in the shape of feelings of pain, and of vague pains in the perinaeum and lumbar region, and of burning in the urethra during the passing of urine. All the symptoms are usually aggravated after excesses in eating and drinking, and after considerable bodily exertion. Another condition found, but not often mentioned, is lipuria (Stock^) ; the highest amount has been 2 per cent, fat in the urine. Stock found 6 to 20 per cent, of eosinophile cells in ten cases examined by him. They appear to be increased, especially in the early cases ; Kautsky^ also called attention to the excessive degree of eosinophilia, whilst Goebel'^ expresses the opinion that a specific toxic action on the organism generally is not developed in bilharziasis. Kautsky* assumes a toxic anaemia as in the case of ancylostomiasis. English authors also have called attention to the eosinophilia and to a considerable ' Stock, Lancet, September 29, igo6. '^ Kautsky, Wien. klin. Rundschau^ iQOjj xxxvi. ^ Goebel, Arch. f. Schiffs- u. Tropen-Hyg.^ I903> vii. ^ Kautsky, Wien. klin. Rundschau, 1903, xxxv 642 THIi ANIMAL PARASITES OF MAN amount of leucocytosis (Balfour/ Douglas and Hardy-). The severe fonus occur almost exclusively in men ; symptoms of catarrh of the bladder make their appearance, vesical calculi are frequently found, whilst the formation of stone in the kidneys and ureters is rare. Urethral fistula occurs in bilharziasis, often without stricture, and if granulations occur the fistula is distal to them. GoebeP regards the bilharzia fistula as a chronic burrow^ing of pus, caused by the irritation set up by the ova as foreign bodies and consecutive restricted suppuration ; and secondly as due to the passage of urine through the defect in the epithelium or the wall of the urethra. The fistulae, which are generally situated at the neck of the bladder and al the membranous portion, are very tortuous and frequently very numerous; they often lie embedded in well-marked tumours — in fact, in granulation tumours with marked inclination to excessive formation of cicatricial tissue. The opening generally is in the perineal and scrotal regions. In the case of a patient, aged 21, from the Transvaal, Kutner* found by cystoscopic examinations the whole summit and walls of the bladder covered with l^uge and small tumours. In addition lo smooth glistening tumours, others were more or less dis- integrated, and scattered large and small cauliflower-like growths occurred. Like malignant growths, the tumours were inclined to break down, the process extending from within outwards towards the surface. Whether the hydrocele so frequent in Egypt has any connection with bilharzia is not known. A frequent sequela of bilharziasis is complete sexual impotence (Petrie^). Bilharziasis of the rectum is manifested by symptoms of dysentery ; the repeated violent attempts at defaecation lead in time to prolapse of the rectum, which sooner or later induces septic infection and so death. In the mucosa of the rectum, polypoid growths similar to those in the bladder are met with, due to the ova of the parasites in the mucosa and submucosa. In the case of a man, aged 36, who had lived for a long time in South Africa, Burfield*^ found in the excised vermiform appendix ova of Scliistosonia luvinatohimn ; he assumed this to be a gradual secondary infection of the appendix, whilst Kelly'' mentions a case of primary bilharziasis of the appendix; the eggs lay in the submucosa directly above the muscularis. Tumours containing numerous ova are frequently found in the region of the genitalia, thighs and scrotum. In one case Symmers^ found numerous male schistosomes in the portal blood and a copulating * Balfour, Lancet, December, 1903. 2 Douglas and Hardy, ibid., October, 1903. ^ Goebel, CentralbL f. d. Kiankh. d. Haim ii. Sexualorgane, xvii. * Kutner, ibid., xvi. '" Petrie, Brit. Med. Joum., July, 1903. ^ Burfield, Lancet, February 10, 1906. ^ Kelly, quoted by Burfield. ^ Symmers, Lancet, January 7, 1905. SUPPLEMENT 643 pair in the left lung. Though schistosome eggs have been found by some observers in the lung tissue, this is nevertheless the first case in which living parasites have been found in the lesser circulation. Perliaps they got there by way of the external iliac vein from the veins of the bladder and rectum. In the female sex bilharziasis is incomparably rarer than in the male and is generally limited to haematuria. Biliiarziasis of the vagina, which takes the form of an acute vaginitis, is frequent according to Milton.^ Horwood^ found in one case a polypoid tumour of the cervix uteri, and in the connective tissue of the tumour Schistosoma ova, both in masses and singly, It could not be established whether the ova reached the vagina and thence the cervix directly, or through the urine from the bladder. Tlie course of the disease is chronic, and in slight cases, provided fresh infections do not occur, is not unfavourable; in severe cases the cachexia caused by loss of blood, or intercurrent diseases to which the patients easily succumb — e.g., pyelitis, pyelonephritis, pyaemia, or ur^emia — lead to a fatal issue. In regions in which Schistosoma luvmatohiiun is endemic, or in patients from such regions, the diagnosis is easy by microscopically linding the eggs in the urine. As regards the treatment of the affection this much must be said, that so far there is in existence no certain remedy. In countries where bilharziasis is endemic copaiva balsam is considered a specific. Kutner {loc. cit.), however, in the case of his patient who for a long time had taken no inconsiderable amounts of copaiva, had no success wortli speaking of to record. Urotropin (three times daily, i grm.) has similarly failed, salol (075 grm. several times daily) perhaps affords relief in affection of the bladder (Milton). Methylene blue, oil of turpentine with extract of male fern (Brock^), or the latter alone and santonin given in small doses for a week at a time, in the morning, are said by Petrie'' to be of value. Sandwith^ and Harley^ were not very successful. By way of experiment Kutner for some time used collargol per rectum, proceeding on the assumption that this prepara- tion, which has proved of such remarkable service in bacterial infec- tion, would perhaps render a continuance of life difficult for the bilharzia worms. But this hope proved illusory. In order so far as possible to limit the loss of blood, Kutner regularly employed stypticin for long periods (three times daily, two tabloids of o'oi grm.) Milton, quoted by Loo?s, " Handb, d. Tropenkrankh.," v. Mense, 1905, i, p. 95. 2 Horwood, Brit. Med. Journ., March 10, 1906. 3 BxocV, Jotmi. of Path, and Bad., 1893. ^ Petrie, loc. cit. ^ Sandwiih, Annal. of Surgery, 1904, xxxix. " Harley, Lancet, 1870. 644 THE ANIMAL PARASITES OF MAN with undoubted success, in so far that the haemorrhages became considerably less in amount. As two patients in the course of enteric fever lost their haematuria, Stock accordingly recommends subcu- taneous injections of Wright's typhoid vaccine. In the early stages of the rectal lesion suppositories of iodoform, ichthyol, or narcotics might possibly be of use. In the case of urethral hstulae, division, excision and scraping out of the granulation tissue are recommended ; in cystitis with formation of tumours high resection with curetting of the tumours or their destruction with the cautery ; in the case of vesical calculi, high resection, curetting the bladder, and then drainage. Tumours of the rectum must also be removed by operation. Prophylaxis is important ; it should be extended to all modes of using water, only filtered water being drunk, and only boiled w^ater being used for washing. This advice should be given to tourists who travel through the infected districts, and is also recommended to soldiers and officials who are despatched to the Colonies. The favour- able influence of change of climate can only show itself where fresh infections are avoided. CESTODES. GENERAL. It seems advisable to preface the section on the Cestodes with some general observations on the symptoms of disease provoked by tapeworms, especially so far as they relate to the question of toxic effects, and to include the Nematodes in this discussion. After this will follow a brief exposition of the most important intestinal lesions causally connected with intestinal parasites. It is known to every experienced practitioner that the different intestinal parasites can give rise to a series of nervous symptoms, slight or severe, and produce, above all, blood changes — anaemia of the most varied nature, to the extent of severe progressive anaemia. These symptoms are regarded by many authors as reflex, or, as in the case of ancylostomiasis, the main feature from the loss of blood caused by the habit of life of the intestinal parasites. More fre- quently, however, they are regarded as toxic conditions produced by the parasites. In view of this divergence of opinion there appears to be some advantage in defining clearly the present position as to the toxic action of parasites. Most interesting in this respect are D ibothriocephalns latiis and Ancylostoiim diiodenale. We are indebted to the clinic at Helsingfors for our most detailed knowledge of bothriocephalus anaemia. Reyher^ was the first to Reyher, Deutsch. Arch. f. klin. Med., 1886, xxxix. SUPPLEMENT 645 demonstrate that this parasite under certain circiunstances can pro- duce a severe, progressive and sometimes fatal anaemia, which can be cured, generally in a surprisingly short time, by expulsion of the worm. Among the various hypotheses which have been advanced as to the mode of origin of bothriocephalus anaemia, the greatest importance has been attached to the assumption already mentioned by Reyher, but definitely expressed by von Shapiro/ to the effect that Bothriocephalus latns produces a poison which is absorbed by the intestine and exercises a deleterious influence on the composition of the blood, especially on the erythrocytes, perhaps also on the blood- forming organs. This assumption is supported by no slight number of clinical and experimental investigations. Podwissotsky^ observed severe blood changes in a child, aged 4^, affected with B. latus. In the case reported by Pariser^ the severe anaemia in a girl dis- appeared fairly soon after expulsion of the worm. In that reported by Schaumann'* high fever accompanied the bothriocephalus anaemia ; he also proved the haemolytic properties of the broad tapeworm. The case reported by F. Miillef^was one of sev^ere anaemia. Also, in the first of the cases described by Kurimoto^ of DiplogONOporiis grandis there were present the same symptoms of anaemia as in the case of B. latus. Meyer'^ observed severe anaemia in two youths caused by 5. latus. Rosenquist^ has discussed the proteid metabolism in anaemia. The presence of B. latus produces in the majority of cases an increased proteid consumption, to which the blood change generally corresponds — toxic anaemia ; in a further communication he reports on twenty cases of bothriocephalus anaemia, nineteen of which were cured by expulsion of the worms, while one case proved fatal, and he again emphasizes the toxic properties of the intestinal parasites. In the case reported by Bendix,^ that of a girl, aged 4^^, the anaemia was moderate, whilst in the case of Zinn^^ (a woman, aged 30) the anaemia was so excessive that the patient succumbed five days after expulsion of six bothriocephalus heads. Isaac and van den Velden^^ have established that in the serum of patients who suffer from anaemia due to B. latus, parasitic products are dissolved,. ' von Shapiro, Zeilschr.f. klin. Med., 1888. ^ Podwissotsky, /a/^r(5. y. Kinder krankh.., 1S89. '^ Pariser, Deulsch. vied. IVochenschr., 1892. ^ Schaumann, Berli.n, 1894, and Deiitsch. med. PVochenschr., 1898. '' Miiller, Chariie- Annul., xiv. 6 Kurimoto, Zeitschr. f. klin. Med., xl, and Kongr. f. inn. Med., Karlsbad, 1899. '^ Meyer, Motint Sinai Hosp. Reports, 1903 and 1904, iv. ^ Rosenquist, Vereinf. innere Med. in Berlin, May 6, 1901 ; and Zeitschr. f. klin. Med. xlix. ^ Bendix, Deutsch. Aerzte Zeitg., 1904, i. '° Zinn, Deutsch. med. Wochenschr., 1903. " Isaac and van den Velden, Deutsch. med. Wochenschr., 190 , xxvii. 646 THE ANIMAL PARASITES OF MAN as shown by a distinct precipitin reaction. Galli-Valerio^ considers it likely that toxic substances are secreted by the living helminthes which produce a lowering or raising of the body temperature, nervous disturbances and haemolysis. Tallqvist^ succeeded in extracting from B. latus a lipoid-like body which had a strong haemolytic action. The experimental anaemia thereby produced differed in no respect from the severe chronic bothriocephalus anaemia of man. The question as to under what special conditions severe, and sometimes fatal bothriocephalus anaemia is developed is answ^ered by Leichten- stern^ and by Lenhartz,* by the assumption that among the Bothrio- cephali some are toxic, that is, manufacture a poison which, when absorbed by the host, produces a severe anaemia. Certain factors lead him to conclude that an accumulation of poison, dependent on time and place, occurs in the Bolhriocephali. In the case of ancylostome anaemia, experience so far, according to Leichtenstern,^' by no means supports the hypothesis of a difference in virulence of the worms according to time and locality, ancylostome anaemia being rather, so far as is known at present, in all races of man, everywhere and at all times, simply and solely dependent on the number of ancylostomes, the duration of the disease and — within certain narrow limits — on the individual capability of resisting the loss of blood and the toxic effect of the parasites. As is shown by a short historical resume of the toxic action that has to be con- sidered in ancylostome anaemia, we must admit that doubtless here, as in the case of bothriocephalus anaemia, the toxins secreted by the parasites exercise a haemolytic action, even while admitting Leichten- stern's contention that the significance of the loss of. blood due to ancylostomes must not be underrated. The toxic hypothesis acquired a definite standing through a series of experiments of Lussana^ on rabbits, where he succeded in producing anaemia by injecting urinary extracts of ancylostome patients. Arslan^ extracted toxins from the urine of two ancylostome patients and injected them into rabbits, which thereupon sickened and show^ed the same blood changes as the ancylostome patients. Retinal haemorrhages, so frequent in ancylostome anaemia, which, according to Fischer^ and Samelsohn,^ are not due to direct loss of blood, must also be ascribed to a para- sitic toxin. A further argument in favour of the toxic hypothesis ' Galli-Valerio, Therap. Monatsh., 1905. - Tallqvist, Zeitschr. f. klin. Med., 1907, Ixi. ^ Leichtenstern, " Handb. d. Therap. v. Pentzoldt.-Stintzlng," 1898, 2nd edition, iv. ^ Lenhartz, ibid., 1903, 3rd edition, iv, p. 607. •' Leichtenstern, Deutsch. med. IVochenschr., 1899. ^ Lussana, Rivista Clin. Arch. ital. di din. Med., 1890. ^ Aislan, Rev. mens, des Mai. de VEnfance, 1892. *" Fischer, Versamml. d. ophthal. GeseUsck., 1892. ^ Samelsohn, ibid. SUPPLEMENT 647 is furnished by the blood changes recorded by Zappert,^ Miiller and Rieder,2 Bucklers,^ and Neusser/ which must be regarded as the expression of toxic action, especially with reference to eosinophilia. The striking increase in proteid destruction in ancylostomiasis observed by Bohland,^ and which ceased after the parasites had been expelled, also gives additional support to the assumption of toxic action. The observation of Daniels^ also deserves consideration in this connection, according to which the presence of yellow pigment in the liver and kidney cells is to be attributed to blood destruction by a verminous toxin absorbed from the gut. Looss^ considers it not at all improbable — in fact, almost certain — that Ancylostoma, in addition to withdrawing blood, exert a kind of toxic action on their host. Scheube^ attributes almost equal importance to the loss of blood, the digestive disturbances, and the intoxication induced by certain metabolic products of the parasites. According to v. Jaksch^ ancy- lostome anaemia is not- induced solely by loss of blood, but by the fact that the parasites produce a ferment which has a toxic action and produces stimulation in those organs in which the eosinophile cells arise. The haemolytic action of ancylostomes has frequently been observed by Galvagno'^ in men employed in sulphur mines. According to Loeb and Smith^^ the anterior half of the body of ancylostomes contains a substance which probably causes anaemia. Bauer^^ found in the urine of ancylostome patients glycuronic acid, which he considers to be a sign of metabolic disturbance due to parasitic toxins. As has been demonstrated by Allessandrini,'^ the secretio n of glands in the anterior part of the body has a distinct haemolytic effect on the erythrocytes. While the worm attaches itself to the mucosa by means of its teeth, these glands discharge their secretion, producing hyperaemia. The extravasated blood is acted on by this secretion, so that it can serve as food for the parasites, Hynek^^ attributes eosinophilia (up to 20 per cent.) to a toxic action. Goldmann^^ expresses a similar opinion, though he ' Zappeit, IVien. klin. IVochenschr., 1892. - Muller and Rieder, Deiitsch. Arch. f. klin. Med., xcviii. 3 Bucklers, Munch, ined. Wochenschr., 1894. * Neusser, Wien. klin. Wochenschr., 1892. ^ Bohland, Milnch. vied. Wochenschr., 1894. ^ Daniels, Lancet, No. 3,725. ' Looss, Cenlralvl.f. Bakt.^ 1897. " Scheube, " Die Krankli. der warm. Lander," 1896. " V. Jaksch, Milnch. med. Wochenschr., 1902. '0 Galvagno, Arch, di Patol. e Clin, inf., 1902-1904. " Loeb and Smith, Cenlralbl. f Bakt., xxxvii. '- Bauer, Wien. klin. Wochenschr., 1904. 1^ Allessandrini, Policlinica, 1904. ^* Hynek, Klin. Chron., 1904. '^ Goldmann, Wien. klin. Rundschau, 1905. 648 THE ANIMAL PARASITES OF MAN assumes that the anaemia is secondary, as the toxin of the cephaHc glands, as the parasites bite, penetrates the mucosa and thence into the blood, where it dissolves the red blood corpuscles. Homani' discusses the agglutinating haemolytic action of the serum of ancy- lostome patients. Whether Ancylostoma produce toxins and what is their nature, or whether the loss of blood causes the anaemia, Liefmann^ was unable definitely to determine ; haemolytic substances do not appear to take any part in it. Berti^ also is inclined to attribute the anaemia to metabolic products of the ancylostomes; he found, in fact, that a serum obtained from a sheep (after subcutaneous injections of the culture fluid of ancylostome larvae) was efficacious in the treatment of ancylostome anaemia. Peiper'^ likewise assumes that the parasite secretes a cell toxin. Lobker^ at the present day still maintains that the cause of the disease must be looked for really, if not perhaps entirely, in the continued withdrawal of blood by the parasites ; the secretion of toxins by ancylostomes has not yet, in his opinion, been conclusively proved. Except in the case of Botliriocepkalus laUts, referred to previously, toxic action appears to be of quite subordinate importance for the other Cestodes occurring in man — especially Tcenia solium and T. saginata, which are most frequently found ; thus Cao^ flatly denies the presence of toxins in the body of Taeniae, while others, such as Messineo and Calmida,^ Jammes and Mandoul,^ consider they are justified from their investigations in concluding that Taeniae contain a specific toxin. Messineo'-' injected, with all bacteriological pre- cautions, extracts of Taenia, dissolved in physiological salt solution. He invariably obtained severe motor disturbances and frequently death. The observation by Pereira^^ of a case of chorea in which rheumatic and cardiac symptoms were absent and which after expul- sion of a Taenia was quickly cured, also favours the view of a toxic action. Barnabo," however, was unable to obtain a toxin from Taenia saginata. Gagnoni,^^ on account of a marked eosinophilia which, after expulsion of a Tcenia saginata, fell within fourteen days to I per cent., assumes the formation of a Taenia toxin. Dirksen's^^ observation has reference to a sailor affected with serious anaemia, who, after expulsion of twelve pieces of Taenia soliniii, was rapidly 1 Romani, Gaz. d. Osp., 1904. ^ Liefmann, Zeitschr.f. Hyg., 1905, 1. ^ Berli, Gaz. d. Osp., 1906. •* Peiper, Deutsch. med. lVo:henschr., 1897. ^ Lobker and liruns, Arb. atis dem kaiserl. Keichsgesundheitsamt, 1906, xxiii. " Cao, Ri/orjua Med., 1 901. ■^ Messineo and Calmida, Centralbl. f. Baki., xxx. ** Jammes and Mandoul, Acad, des Sciences, 1904. ^ Messineo, Gioru. med. del regio cserc, 1505. '" Pereira, Lancet, September, 1903. " Barnabo, Speiimenfale, 1906, v. " Gagnoni, Pediafric, 1903. '' Dirksen, De^iisch. med. IVochenschr., 1903. SUITLEMENT 649 cured. A portion of the worm was already breaking down, the absorption introducing into the body highly toxic ha^molytic pro- ducts, to which the anaimia must be ascribed. How far the serious disturbances of the nervous system, frequently to be observed in cases of Hymenolepis nana, are to be considered as of purely reflex nature or toxic must remain an open question ; the same applies to Dipylidinni caninnni, in which case Brandt^ observed serious central nervous symptoms. Caution is necessary in judging as to any con- nection between worm stimulus and nervous symptoms in cases of Ascaris infection. Peiper^ is inclined to regard such nervous symptoms not as reflex, but rather as due to a toxin contained in the helminthes, or metabolic in origin. In cases of pernicious anaemia when the symptoms disappear after expulsion of Ascaruhv a toxic action must be assumed (Demme^). Additional clinical observations do not, indeed, lead to any definite conclusion as to the question whether Ascaridce produce a toxin which is capable of causing more or less injury either to the nervous system or to the blood, yet it may be worth while to give a brief review of this question. In a case of Kutner's,^ that of a girl, aged 12, there was a haemolysis which was cured after expulsion of twenty- four Ascaridcv. Attacks of opisthotonos in a girl, aged 16, ceased after seventy-eight Ascaridcv had been expelled (Lutz'^). Unusually serious disturbances were observed in a man, aged 26, who was rapidly cured by Drouillard*^ by the removal of a great number of Ascarida^. The observations on pseudomeningitis are of especial interest ; they are evidently toxic in origin as in the case of Annaratone,^ of a man who was taken ill with gastro-intestinal symptoms and who died with meningitic symptoms. Post niortcin the brain was normal, but the stomach contained a great coil of Ascaridce. The cases of Delille,^ Meriel,^ Papi^^ (the occurrence of Cheyne-Stokes respiration has been ascribed to the action upon the centre in the medulla oblongata of the products of the Ascaridcv), and Taillens^^ related to children in which the meningitic symptoms (meningismus), partly serious, dis- appeared with the removal of the Ascarida\ Mareo^^ designates this disease helminthiasis meningitiformis, which exhibits all the symptoms * Brandt, quoted by Pollak in Cent^albLf. Bakt., 1889, v. ^ Pciper, vide Seift-rt, " Lehrb. d. Kinderkrankh.,'' 1897, p. 243. ^ DeiTime, vide Seifeit, ibid. * Kutner, Be'l. klin. Wochenschr., 1865. •5 Lutz, Centralhl. f. Bakt. ^ Drouillard, Journ. de Med., 1900. xi. "^ Annnratcne, Giom. vied, del regio esetc, 1900. * Dflille, Journ. de Med., May 10, 1907. " Meriel, Annul, de Med. et Chir. inf., 1900. '0 Pani, Gaz. d. Osp., 1901. ^^ Taillen?, Arch, de tited. d'Enf., 1906. '^ Mareo, Allg. VVien. med. Zeitg,, 1902. 650 THE ANIMAL PARASITES OF MAN of meningitis, but which is caused by the metaboUc products of A scar i da'. Schupfer/ Duprey- (observations in the West Indies, where such symptoms are said to be of very frequent occurrence), Naab^ (the flow of w^ater from the mouth at night is mentioned as a remarkable fact), and Hammiss* assume the action of an Ascaris toxin in the clinical observations made by them, mostly children with fever and intestinal symptoms. Schupfer assumes in such cases, as he observed it once in a man, aged 23, that the disease termed Lonibricolse a forme iyphoide by Chauffard was due to B. coli of marked virulence due to the action of the Ascaridcv. The Widal reaction was negative. Koneff^ reports a case in which acute attacks of cramp, trismus, and rigidity of the pupil disappeared after expulsion of seven Ascaridce. Tetanus, as observed by Buchholz*^ in a girl, aged 17, and rapidly cured after expulsion of sixteen Ascaridce, is manifestly rare, since only Rose^ mentions this as a cause in his article on Tetanus. Only a few experimental data exist. Cattaneo^ could detect only a very weak toxin in Ascaris, while Messineo,'' by injecting into animals extracts in physiological salt solution, invariably succeeded in producing serious motor disturbances and frequently death. Interesting also are the observations of Huber,^^ who, after working with Ascarida% suffered from itching of the head and neck, blisters, swelling of the ear, con- junctivitis, ecchymosis and troublesome palpitation in the head. He consequently assumes that Ascaridcv can induce irritation by chemical (toxic) means. In the case of Trichocepliahis dispar no more than in the case of Ascaris Inmhricoides can we speak with certainty of a toxic effect, even though a number of observations are available which might justify such an assumption as regards these intestin?.! parasites. Barth^^ found the brain normal in a man who had d ed with meningitic symptoms, but the intestines were full of Trichocephalus dispar; Gibson^^ records the rapid cure of serious cerebral symptoms after expulsion of Tricho- cephalus, so also Pascal,'^ Burchhardt^^ and Rippe.^^^ Moosbrugi^er^^ Schupfer, Gaz. d. Osp. 1901. 2 Duprey, Lamet, 1903. ^ Naab, Munch. meL Wochenschr., 1902. ^ Hammiss, IVien. med. Wochenschr.^ 1904, iii. ^ Koneff, quoted by Liesen, " Dissert. Eoiin," 190^. 6 Buchholz, Norsk. Mag. for Lcige, 1903. ^ Rose, Billroth and Piiha, '' Chirurgie." " Caltaneo, Arch.f. Kinder heilk., xliv. " Messineo, Giorn. vied, del regio eserc. , 1905. '" Huber, Deutsch. Arch.f. klin. Med., 1870, vii. " Earth, reported by Valleix, Paris, 1845. '" Gibson, Lancet, 1862. '^ Pascal, quoted by Kahane, KorrespondeJizhl. f. Schiveizer Aerz/e, 1907, viii. " Burchhardt, Deutsch. med. IVochemchr., 1880. '•' Rippe, St. Petersb. med. Wochenschr., 1907, i. "* Moosbrugger, Med. Corresfondeuzbl. f Wurttemberg, 1890. SUPPLEMENT 651 was the first to draw attention to grave anaemic conditions induced by Trichocephalus, Morsasca^ and Becker^ to progressive grave anaemia (trichocephalus anaemia is accompanied by marked reduction of the number of red blood corpuscles, of the specific gravity and of the haemoglobin, well-marked morphological changes of the red cell, micro-, macro-, and poikilocytosis and nucleated red cells). Sandler,^ in his case of a boy, aged 11, who died of anaemia, assumes a tricho- cephalus toxin to be the cause of the disease, and Kahane also reports on anaemic conditions induced by Trichocephalus. Girard,'* in addition to symptoms in the gastro-intestinal tract, calls attention to those arising in tlie blood — anaemia and its sequelae — and also to nervous symptoms : cerebral phenomena, headache, giddiness, aphonia, sym- ptoms of meningitis. In a case of Schiller's'^ high fever was present, which probably set in when the Trichocephali present in the gut in great numbers commenced their parasitic activity. Hausmann,^ in order to explain the adaptability of Trichocephalus, assumes that according to the locus niinoris resistenticc, at one time the reflex at another the toxic action is effective, now on one organ, then on another ; anaemia being present in most cases, frequently general and local neuroses and cerebral symptoms of various kinds. With regard to the toxic action of Oxyuris there is only the single record of Hartmann,^ who noticed the disappearance of epileptic fits and psychic disturbances in a gu'l, aged 13, after the removal of Oxyuris. Nervous disturbances and blood changes can but rarely be attributed to Strongyloides. Silvester^ and Valdes^ report on giddi- ness, headache and anuria in cases observed by them ; whether the eosinophilia recorded by Biicklers^^ and Bruns^^ is due to the toxin of Strongyloides must remain an open question. Reference has already been made to the possibility that intestinal ciliates {Balantldluin coll) can also produce toxins. The contents of echinococcus cysts appear to contain a substance only moderately toxic, giving rise to urticaria, in a series of cases where the fiuid has escaped into the abdominal cavity (during puncture). D. Muller^^ has collected nine such cases out of the literature, to which may be added six cases of Finsen^^ in which the I ' Morsasca, abstract in Centralbl. f. innere Med., 1897. '^ Becker, Deutsch. j?ied. Wochenschr., 1902. ^ Sandler, ibtd.y 1905. ^ Girard, Anual. deTlnst. Pasteur, 1901. ^ Schiller, Beiir. z. klin. Chir., 1902, xxxiv. ^ Hausmann, St. Peter sb. med. Wochenschr., igoo. ■^ Hartmann, Naturforschervers., Koln, 1889. " Silvester, quoted by Schliiter, "Dissert. Kiel," 1905. " Valdes, quoted by Schluter, op. cit. '" Bucklers, Miinch. med. Wochenschr., 1894. " Bruns, MiUich. med. Wochenichr.^ 1907. '- Milller, D., " Dissert. Wurzburg," 1885. " Finsen, quoted by D. Muller. 41 652 THE ANIMAL PARASITES OF MAN escape of fluid into the peritoneal cavity led to severely itching urticaria, which usually disappeared again after one or two days. On one occasion, nideed, urticaria occurred after rupture into the pleural cavity. In the case recorded by Caffarena^ of echinococcus of the right lobe of the liver, widespread urticaria developed as the result of the exploratory puncture. In the case of an echinococcus of the liver rupturing into the abdominal cavity La Spada^ ascribed the symptoms leading to death to toxic influence while the peritoneal symptoms w^ere less marked. Eosinophilia in hydatid disease is slight according to the investigations of Bindi^ and Santucci,* and is, according to Welsh and Barling,^ no certain sign of echinococcus ; it is independent of the age, sex and temperature of the patient, but upon rupture of the cyst eosinophilia invariably sets in. The question as to the importance of helminthes in relation to certain diseases of the gut requires special discussion, but it concerns only Ascaris liimhricoideSj Oxyuris vermicular is y and Trichocephaliis dispar, and the question of appendicitis first of all. The entrance of intestinal parasites into the vermiform appendix was already known to medical men in the fifties of last century, as is showai by the works of Merling« (1836), Zebert^ (1859), Platonor^ (1853), and Schachtinger^ (1861). Most of these authors have considered intestinal worms, together with other foreign bodies, to be the cause of appendicitis. As regards the part played by these intestinal parasites in the etiology of appendicitis, so much discussion has taken place during the last few years that it is worth while to give a resume of the later views on this question, even though at the outset it must be admitted that the matter is not cleared up. Bergmann^^ records a case in which an Ascaris perforated the appendix and got into the peritoneal cavity. StriimpelP^ reckons among the symptoms of Trichocephalus the possibility of a *' typhlitis." On account of the marked sensitiveness of the ileo-caecal region, Boas^^ mentions the possibility of confusing it with appendicitis. StilP^ regards Oxyuris as a principal cause of catarrhal affections of the appendix. Arbore-Rally^* regarded severe symptoms of appendicitis in a boy, aged 10, as due to Ascarides. In all cases of appendicitis Metschnikoff^^ requires a microscopical examination to be made for eggs, and considers treatment for worms ' Caffarena, Convers. din. Genova, 1902. ^ l^ Spada, Gaz. d. Osp., 1904. ' Bindi, ibid.y 1907. ^ Santucci, ** Clinica moderna," 1905. ^ Welsh and Barling, Scot. Med. and Surg, yotirn., 1907. » > > ^0 Bergmann, Prag. med. Wochenschr.^ 1890. " Striimpell, '* Lehrb. d. spez. Path. u. Therap.," 1894. '2 Boas, Deutsch. vied. IVochenschr., 1895. '' Still, Brii. Med. Journ., 1899. ^'» Arbore-Rally, Arch, de Med. des En/., 1900. '^ Metschnikoff, Bull, med., 1901. SUPPLEMENT 653 carried out otherwise as a cause of the frequency of perityphHtis. Matignon^ does not agree with this opinion, as in spite of the extra- ordinary frequency of intestinal worms in China, he has only seen one case of appendicitis in four and a half years, and Des Barres^ expresses himself in similar fashion. Out of twenty-one cases of appendicitis Kirmisson^ discovered the ova of Trichocephalus eighteen imes and the ova of Ascarides in three of these cases ; in twelve cases of enteric fever the examination for eggs was negative nine times. Moty^ considers Oxyuris to be the sole cause in his three cases of appendicitis. Girard^ ascribes to Trichocephali the role of more or less septic foreign bodies which may bring about the entry of intestinal bacteria into the appendix, and Triboulet^ describes a case of appendicitis which he considers was due to Ascaris. In Morkowitin's'' case numerous Oxyuris had clearly caused the appendicitis, von Genser^ records the case of a boy, aged 5, who was operated on for appendicitis, and who passed through the opera- tion wound a living Ascaris on the eighteenth day after the operation. In the first case communicated by Schiller^ the disappearance of the typhlitic swelHng after the discharge of the Ascarides pointed to the etiological significance of the parasites, and the same obtained in a further case published at an earlier date by Czerny and Heddaus.^*^ In a case abstracted by Kaposi^^ Trichocephali appear to have been a contributory cause in the production of the appendicitis. In a further case reported by Schiller, where the appendix was removed, was shown that Oxyuris had given rise to a pronounced appen- dicular colic. In a girl, aged 13, who died from diffuse peritonitis, Schwankhaus^^ found that an Ascaris had perforated the appendix. Ramstedt^^ found in an extirpated appendix a whole '^tangle" of Oxyuris, and believes in the possibility of their having provoked the inflammation ; he recommends an examination for entozoa before the operation, without, however, after Metschnikoff's example, sub- stituting worm treatment for the operation. Rostowzeff^^ ascribes only a minimal direct etiological significance to intestinal worms ' Matignon (abstract), Munch, rued. Wochenschr.^ 1901. ■2 Des Barres, Gaz. des Hop., 1903. ^ Kirmisson, Annul, de Med. el Chir, des Enf.y 1901. 4 Moty abstract), Mx'inch. med. Wochenschr., 1901, p. 910. G irard, Amtal. de VInst. Pasteur, 1901. 6 Triboulet, .S'iJc-. med. des H6p. de Paris, 1901. ■^ Morkowitin (abstract), Centralbl. f. d, Grenzgebiele, 1902. " V. Genser, Wien. med. Wochenschr. y 1901. Schiller, Beitr. z. klin. Chir., 1902, xxxiv. ^° Czerny and Heddaus, Und., xxi. " Kaposi, ibid., xxviii. '^ Schwankhaus, Amer. Pracl., 1901. '3 Ramstedt, Deulsch. med. Wochenschr., 1902. 1* Rostowzeff, Russ. vied. Rundschau, 1903. 654 THE ANIMAL PARASITES OF MAN .in the origin of appendicitis ; in 163 cases he found worms in three -instances. Wirsaladze^ expresses himself in a similar fashion. Oppe^ observed Oxyuris six times in excised appendices, and emphasizes the opmion that in appendicitis the question of a worm cure ought to be taken into consideration. Ascaris and Oxyuris, if no contra-indication exists, may be expelled, but in the case of Trichocephakis, which frequently defies all expulsive treatment, no attempt should be made, but operation proceeded to forthwith. In a case briefly reported by Hanau^ Oxyuris was undoubtedly the etiological starting-point; in a case of Galli-Vallerio* Oxyuris and Trichocephakis. In the opinion of Ssaweljews^ in some cases of appendicitis, in addition to other causes, intestinal parasites play a prominent part. The case recorded by Nason^ is an interesting one ; m this an Ascaris in the appendix became twisted with it round a coil of gut, causing obstruction. Spieler' argues against the under- estimation by many authors as to the part played by intestinal worms in producing appendicitis, although he also does not regard them as a frequent, to say nothing of an exclusive, cause of the disease. In a case recorded by Begonin^ fifteen Oxyuris w^ere found in the excised appendix (the mucosa showed some ulceration), and in another recorded by Putnam^ twenty Oxyuris were present in the appendix, in which there was no evidence of any change. The standpoint Schilling^*^ takes is to the effect that entozoa irritate the mucosa and can increase an already existing inflammation, but he considers it very questionable whether they can produce appendicitis. Blanchard'^ assumes the possibiHty of a secondary infection arising from lesions of the mucosa produced by helminthes (Ascaris and Oxyuris). Moore^^ considers Trichocephakis the excitant of the appendicitis in his case. In a second case of appendicitis recorded by Auley'^ operation became unnecessary owing to the passage of the Ascaridcv. Page's^* case is an interesting one ; it was that of a man who came up for operation with a diagnosis of appendicitis. On incising the abdominal wall numerous Ascarides were found at the base of the ' Wirsaladze, Bcbnlsch. Gaz. Botkina^ 1902. ^ Oppe, Munch, vied. Wochenschr.^ 1903. ^ Hanau, ibid., 1903. ^ Galli-Vallerio, CentrMl. f. Bakt.y 1903, p. 1094. '" Ssawtljews, Deutsch. vied. Zeifg., 1903. ^ Nason, Jotirn. Amer. Med. Assoc. ^ 1904. "' Spieler, Wien. klin. Wochenschr., 1904. ^ Begonin, Journ. de Mid. de Bordeaux, July, 1902. ^ Putnam, quoted by Spieler. "o Schilling, " Wurzb. Abhandl.," 1905. v. " Blanchard, Acad, de Med., July 3, 1904. ■'2 Moore, Brit. Med. Journ., Augu t 18, 1906. '^ Auley, ibid., i(^o6. '^ Page, New York Med. Journ., Januaiy 20, 1906. SUPPLEMENT O55 wound, lying in cavities ; even after eight days Ascarides escaped from the wound. The author assumes there was a perforation of the gut wall ; it is strange that the worms were able to exist a proportion- ately long time in the muscular tissue. Schoeppler^ states that there is the danger of an appendicitis even after the death of an Oxyuris that has found its way into the appendix. Oui^ met with two specimens of Trichocephalus which had become embedded by their thin ends deep in the mucosa. Frangenheim^ is not in a position to pronounce any opinion as to what part intestinal parasites play in the etiology of appendicitis. In a case recorded by Kahane'^ many Trichocephali were found partly free in the appendix and partly embedded in the mucosa ; microscopically appendicitis was dia- gnosed. At a laparotomy for salpingitis Heekes^ found the appendix elongated, thickened, and containing about eleven Oxyuris without the mucosa being in any way changed. In one case Andrews^ claims Ascarides to have been the direct cause of the appendicitis. The literature dealing with this question, so important in our time, has been collected almost without any omissions, but, unfortunately, no decisive opinion as to the significance of parasites in appendicitis can be inferred from it. The vexed question whether intestinal parasites, especially Ascaris, are able to penetrate the intestinal wall is just as little finally decided. Leuckart,^ Heller,^ Mosler and Peiper,^ Henoch/^ Davaine," Klichenmeister,'"^ and Bremser^^ are opposed to the idea that the healthy intestinal w^all can be penetrated by intestinal worms, especially Ascarides, whilst a whole series of other authors are of the opinion that even the healthy intestinal mucosa can be perforated. Among these is numbered Mondiere,^* who is of the opinion that Ascaris, by violent pressure against the mucosa, forces it so much apart that it is enabled to escape through the gap thus formed into the peritoneal cavity; this opinion is shared by V. Siebold.^'^ Rokitansky^^ considers perforation of the gut by Ascaris ^ Schoeppler, Centralbl. f. Bakt.^ 1906. 2 Oui, Rev. prat. d'Obstet. ei de Paed., 1906. •^ Frangenheim, Samml. klin. Vortr.^ 1906, No. 424. ^ Kahane, Schweiz. Korrespondenzbl.^ 1907, viii. ^ Heekes, Brit. Med. Journ., March 16, 1907. ^ Andrews, ibid.^ 1906. '^ Leuckart, " Die Parasiten des Menschen." ^ Heller, *• Handb. d. spez. Path./' v. Ziemssen, vii. '•^ Mosler and Peiper, ** Spez. Palh. u. Ther.," v. Nothnagel, vi. '0 Henoch, " Vorlesungen iiber Kinderkrankheiten." " Davaine, " Traite des Entozoaires." '^ Kiichenmeister and Ziirn, " Die Parasiten des Menschen." '=^ Bremser, *'Lebende Warmer im lebenden Menschen." '^ yionAihxe, Schmidt'' s Jahrb., 1840. '» V. Siebold, *' Parasiten" in Wagner's " Handworterbuch," 1845. '« Rokitansky, " Path. Anat." 656 THE ANIMAL PARASITES OK MAN as at least a rare occurrence. Gerhardt^ does not doubt that the worms can actively perforate the intestine. Cases like those of Abrault,- Apostolides,^ Marcus'' (recorded by Perls as a valid example of '* ascaridophagous" gut perforation), Wischnevvsky,^ Galvagno/ Salieri' certainly show that perforation of the healthy gut wall cannot be denied, but at the same time that this occurrence, compared with the frequency of Ascaridce, should be regarded as exceedingly rare. It is another matter as to whether it is possible for the worms to penetrate an intestinal wall already diseased, especially when ulcerated ; a whole series of observations are in favour of this. In Lini's^ case (fifty-six Ascarides escaped from the umbilicus of a girl, aged 7), in Gralfe's^ (eighty Ascarides escaped from an inguinal tumour), in Nicolino's^^ (perforation of the intestinal wall with strangulated hernia), in Liesen's" (a living Ascaris in the peritoneal cavity in a woman suffering from a peritoneal abscess) — in these it is clear that disease processes in the intestine preceded the exit of the worms. In a case described by Boloff^^ the Ascarides appear to have produced, by forming a tight coil, necrosis of the gut with perforative peritonitis. In a case recorded by Lutz^' the perforative peritonitis w-as without doubt provoked by Ascaris, and in one by Schiller^* the Ascaris had clearly gained access to the peritoneal cavity through a gunshot wound opening. In a case observed by Rehn^^ the w^orm probably entered through a gangrenous portion of the intestine in a hernial sac. Broca^^ is unable to deter- mine whether in his case the intestinal perforation was primary (a worm escaped from the abdominal wound about two months after a laparotomy for suppurative peritonitis). The case reported by Lutz^^ is of special interest : it was that of a young man who had shot himself in the region of the abdomen, and w^ho died after fifteen days. At the post-mortem two Ascarides were found in the pulmonary artery ; they had probably escaped from the intestine, and had gained access to the inferior vena cava. Froelich^^ assumes that in his case ' Gerhardt, quoted by Liesen, *' Dissert. Bonn." 2 Abrault, quoted by Seifert, •' Lehrb. d. Kinderkrankh." ' Apostolides, Lancet, 1898. ^ Marcus, quoted by Seifert, •* Lehrb. d. Kinderkrankh." •'■' Wischnewsky, quoted by Seifert, ibid. ^ Galvagno, Arch, de Patol. et Clin, inf., 1902. ■^ Salieri, Rif. vied., 1902. ^ Lini, Schmidt's Jahrb., 1838. " Graffe, F/otokoll d. Ges. f. Natur u. Heilkiinde, Dresden, 1853. '° Nicolino, Clin, mod., 1902. " Liesen, "Dissert. Bonn," 1904. '2 Boloff, quoted by Seifert, " Lehrb. d. Kinderkrankh." '8 Lutz, Centralbl.f. Bakt. '^ Schiller, Beitr. z. klin. Chir., xxxiv, p. 200. ''^ Rehn, 5^1? Schiller, loc. cit., p. 201. "^ Broca, Rev. mens, des Mai. de VEnf., 1904. '■^ Lulz, IVien. klin. Wochenschr., 190$, xv. '^ Froelich, Rev. mens, des Mai. de I'Enf., 1897. SUPPLEMENT 657 (a boy, aged 11) the Oxyuris were able to penetrate the whole intestinal wall, but Vuillemin^ considers this improbable, and is more inclined to think that the Oxyurides penetrated the rectum at small ulcerated points, and thus gained access to the perirectal connective tissue. In females Oxyuris not only have the power of penetrating far into the sexual organs (Marro^), and perhaps causing a parasitic endometritis (Simons^), but also clearly of gaining access to the peritoneal cavity by way of the tubes, as is to be assumed in the case recorded by Kolb^ (that of a woman, aged 42, in whom post inortein nodules were found over the peritoneum of Douglas's pouch, in which the pressure of encapsuled Oxyuris could be demonstrated), in that reported by Chiari^ (adult Oxyuris in Douglas's pouch) and by Schneider^ (an Oxyuris encapsuled in the pelvic peritoneum). Sehrt's^ case is worthy of attention ; in this an abscess was found in the omentum with numerous Ascaris ova in the pus and a nodular lesion of the peritoneum, with Ascaris ova encapsuled in the nodules. Massive accumulation of Ascarides may give rise to a complete occlusion of the gut. Such an occurrence is not so surprising as might be thought when one reflects that the number of Ascarides in one individual may amount to several hundreds. For instance, one boy evacuated within a single day 600 Ascarides (Fauconneau- Dufresne^) and within three years 5,126 worms. In the case recorded by Tschernomikow^ a boy, aged 2^, evacuated during a day 208 worms, partly through the stomach, partly through the intestine. Coil-formation of such masses of Ascarides renders possible not only constipation, but also complete obstruction with symptoms of ileus, as shown by the five cases quoted by Mosler and Peiper,^^ as w^ell as from observations made by Raie,^^ Schulhof,^^ Rehberg,^3 Rocheblave,^* Heller,^^ Leichtenstern,^^ Huber,^^ and VVilms.^^ In two cases of Black^^ and Parkinson^*^ the intestinal obstruction was caused by a coil of tapeworms. In the earlier history of medicine the helminthes played a great part as the excitants of many intestinal diseases and of enteric as well. Even if to-day they no longer be regarded as such, the conception that they represent the predisposing factor in typhoid ^ Vuillemin, Centralbl. f. Bakt., 1902, ^ Marro, Arch, per le Set. med., 1901. » Simons, Centralbl.f. Gyndk., 1899. ^ Kolb, Ceniralbl.f. BakL, 1902. '"' Chiari, Prag. me J. Wochenschr., 1902. ^ Schneider, Centralbl.f. Bald., 1904. "^ Sehrt, Beitr. z. klin. Chir., li. ^ Fauconneau-Dufresne, quoted by Seifert. 9 Tschernomikow, quoted by Seifert. " Raie, Lancet, 1899. '3 Rehberg, ** Dissert. Konigsberg," 1907. '^ Heller, loc. cit. '^ Huber, quoted by Rehberg. ° Mosler and Peiper, loc. cit. 2 Schulhof, Milne h. med. Wockemchr., 1903. ' Rocheblave, Gaz. des Hdp., 1898. 6 Leichtenstern, " Ziemssen's Handb.," vii. Wilms, Deutsch. Zeitschr. f. Chir., xlvi. '9 Black, Brit. Med. Joiirn., 1872. ^o Parkinson, quoted by Rehberg. 658 THE ANIMAL PARASITES OF MAN infection through the injury they inflict on the mucosa (Guiart/ Blancliard,^ Vivaldi and Tonello^) must not be summarily rejected. Vivaldi and Tonello found helminthes in 80 per cent, of their typhoid patients, numbering among these Trichocephahis dispar, Oxyiiris vennicularls, Ancylostoma diiodenale, and Ascaridct. The report of Leuckart"^ is here worth citing, to the effect that Thiebault never failed to find Trichocephalus in his cholera patients at Naples. Blanchard^ goes so far as to express the desire that in every febrile affection of the intestine an anthelmintic treatment with thymol should be undertaken as early as possible, even before learning the results of serum diagnosis. The lesions of the liver and pancreas due to Ascaridcv are briefly discussed in the chapter on Ascariasis (p. 687). A discussion of the intestinal helminthes from the clinical and therapeutical point of view follows these general considerations. Dibothriocephalus latus. From what is known as to the development of Dibothriocephalus latus, the way by which man is infected is self-evident : infection can only take place through the ingestion of insufficiently cooked fresh-water fish (pike, burbot, perch, grayling and vendace) ; what degree of temperature is necessary to kill the larval forms is still unknown. Dibothriocephalus latus lives in the small intestine of man, alone or in some numbers, frequently also together with Taenia solium. The proglottides are passed always united in large pieces, the ova are deposited through the uterine pore, while the worm is still in the intestine, so that they are easily found in the faeces. The proglottides are so characteristic that they cannot be. confused with those of other species. In reference to whether age or sex is spared by Z). latus, it is not possible to make any definite statement, especially so far as the endemic area is concerned, whether a person resides n it continuously or visits it, so long as his habit of life is in accordance with those of the country. Bendix^ certainly emphasizes the fact that early childhood is as a rule immune : his case was that of a child, aged 4^ years. * Guiart, Compt. rend. Soc. de BioL, Paris, March i6, 1901. 2 Blanchard, A^-ch. d. Par., 1901. ^ Vivaldi and Tonello, Gaz. d. Osp., October 29, 1905. 4 Leuckart, quoted by Kahane. s Blanchard, Acad, de Med., Ocloler 18, 1904. fi Bendix, Verein f. inncre Med., Berlin, June 16, 1902, SUPPLEMENT 659 Sparganum mansonl. According to our present knowledge (Miyake^) the disease occurs almost exclusively in China and japan. On the main island it occurs in all districts, though rarely under observation. It is especially frequent in the neighbourhood of Kioto and Osaka; these places are very near together, and between them there is mutually active intercourse, so that taken together they may be regarded as one district infested by this worm disease. As regards localization in the body, there appears to exist a certain predisposition for definite regions, for instance, the eye and genito-urinary tract. In some cases the parasite manifested the peculiarity of wandering about the body and of appearing at certain favourite points (muse, quadriceps femoris) (Hashimoto^). Most patients complain more or less of the onset of attacks of pain and of sensitiveness to pressure. In those cases in which the patients evacuated the worm during micturition, the symptoms were variable ; sometimes there was tenesmus of the bladder, sometimes pains in the inguinal region, sometimes haematuria. None of these troubles is characteristic of the disease, and does no more than represent the symptoms that follow a mechanical irritation that any kind of foreign body may produce. Besides the onset of attacks of pain, swelling of the regions affected, if superficial, may often be recognized, when a superficial diffuse soft tumour can be felt which often gives pseudo-fluctuation. Some- times a peculiar crackling can be detected internally, as in the making of a snowball. During the further course an abscess not infrequently forms around the worm. When the situation of the worm is super- ficial, *' an inflammatory tumour with a tendency to migrate" is stated by Omi^ to be an important diagnostic sign. That, however, is not always the case, as the observation made by Inoye"^ shows. It would be better to add to this sign the onset of paroxysmal pain and the temporary change in volume of the tumour. When once the parasite is removed, the wound heals just as satisfactorily as any other fresh wounds made at operation. Dipylidium caninum {Tcenia ciiaunerina). This species belongs to parasites of rare occurrence. Up to the year 1905 Bollinger^ collected thirty-six cases from the literature, twenty-nine of which were children and seven adults. Since then ' Miyake, Mitleil. aus d. Grenzgebieie, 1904, xiii. - Hashimoto, quoted by Miyake. ^ Omi, Jji-Shinshi^ Tokio, 1898. ^ Inoye, ibid.^ 1897. ^ Bollinger, Deutsch. Arch.f. klin. Med.j 1905, Ixxxiv. 66o THE ANIMAL PARASITES OF MAN some further cases have come to liglit, so that the number now observed amounts to ninety, and among them only eight adults. The youngest child was 6 weeks old (KohF), in which the first pro- glottides were passed when the child was 40 days old. This pre- ponderating occurrence in children is clearly connected with the close intercourse between children and dogs, and also cats. Bollinger believes that D. canimim in reality occurs more frequently in adults than has hitherto been supposed. In addition, it must be mentioned that this species is quite unknown to many physicians, and is occasionally confused with Tcenia soliutn. One notices almost daily a large quantity of cucumber-seed-like bodies, reddish or whitish- grey, about I cm. long and 2 mm. broad, discharged with the stools. Lindblad'^ remarks that these bodies have lively movements, that they perish rapidly in fresh water, and become white and smooth. These Cestodes, in isolated cases, are parasitic in the intestine in large numbers. Sonnenschein^ expelled four fragments in the case of a boy, aged 4 months ; Asam^ three fragments in the case of a child, aged 19 months ; and Zschokke'^ as many as five or six in that of a boy, aged 4. They do not always produce such striking symptoms as occurred in Pollak's case.^ In other cases gastro-intestinal dis- turbances with or without fever (Kriiger^), emaciation (Zschokke), or even nervous symptoms of central origin in the form of con- vulsions (Brandt^) have been observed. From the nature and mode of infection children must be kept from close contact with dogs and cats as much as possible lo ensure prophylaxis. The appropriate treatment, as it mainly affects children, deserves special mention, whilst the expulsion of the remaining Cestodes may be described in this connection. Among the drugs one may mention flor. kousso I'D grm., pulpa tamar. depur. 2 grm., syrup of sugar 50-0 grm., one- third to be taken every hour (Lindblad). Kamala appears to have no effect, although Huber^ recommends it in small doses according to age from 0*5 to 30 grm. He warns against Filix mas preparations, which otherwise, even in early childhood, vuider careful dosage gives the best results. Young children are given i*o to 2*0 grm. extr. fil. maris, with mint syrup or raspberry syrup 30*0 grm., in the morning twice an hour by the mouth, or i-q grm. extr. fil. maris is mixed with syrup ^ Kohl, Munch, med. Wochenschr., 1904. - Lindblad, Hygiea, xlv. ^ Sonnenschein, Milnch. med. IVochenschr., 1 903. ^ Asam, Afilnch. med. Wochensch}-.^ 1903. ^ Zs~hokke, Cenfralbl. f. Bakt., 1905. " PoUak, Wien. klin. Wochenschr., 1907. ■^ Kriiger, St. Petersb. nied. IVochenschr., 1887. » Brandt, Cenfralbl, f. BakL, 1889. •" Huber, suppl. to Asam, Munch, med. IVochenschr.^ 1903. SUPPLEMENT 66l of mint, and given by means of a stomach tube (Rosenberg^). A few hours afterwards a mild laxative may be taken — one to two table- spoonfuls of aqueous tincture of rhubarb (Asam) — or an enema may be given. In a case reported by Sonnenschein decoction of pome- granate root had no effect, as it was vomited up. Hymenolepis nana. This species, very rare in Central and Northern Europe, inhabits the small intestine, especially of children ; it burrows very deeply into the mucosa. Not uncommonly several thousand have been found in one case (Nicolo,^ E. Stoerk and HaendeP). It is remarkable that these Cestodes have been found so frequently post inoiiem and after vermifuges given for other reasons. Thus the clinical symptoms must often be very indefinite (Stoerk and Haendel), so that one may assume that only a slight percentage of cases of Hynienolepis liana come under observation and are published as such. On the other hand, it is certainly conceivable that with the large number of parasites that frequently occur in one individual a whole series of symptoms, in part quite severe, are capable of being produced. These are partly symptoms of intestinal catarrh, consisting of abdominal pains, constipation, alternating with attacks of diarrhoea, perverse appetite,' and boulimia, abdominal pains of a cramp-like nature, followed by emaciation, headache, sleeplessness, pallor, lassitude, and in part nervous symptoms — epileptiform attacks without loss of consciousness, weakness of memory, melancholia, irregular febrile attacks (Lutz^). Possibly, too, Hymenolepis nana infects the urinary organs, producing true chyluria (Predtetschensky^). Stoerk and Haendel are inclined to think that this species, unlike other Cestod.es parasitic in man and domestic animals, needs no intermediate host for its development, and that the larval forms (cysticercoid) live in the same host as the adults. The diagnosis is based on the demonstration of ova in the stools. As far as expulsion of this Cestode is concerned, santonin, kamala, kousso flowers and thymol appear to have no effect of importance ; whilst extract of male fern, recommended by Grassi^ as a result of his considerable and successful experience, has been given, with the result that the worms really are expelled, and that after the treatment neither worms nor ova are any longer demonstrable in the stools of patients. In his cases of chyluria Predtetschensky prescribed ol. 1 Rosenberg, Ges. f. innere Med., February i6, 1904. ^ Nicolo, Gaz. d. Osp., 1904. ^ Stoerk, E., and Haendel, Wien. klin. IVockenschr., 1907, xxix. * Lu'z, Centralbl.f. Bakl., 1894. ■' Predtetschensky, Zeitschr. f. klin. Med., xl. " Grassi, Centralbl. f. Bak:., 1887. 662 THE ANIMAL PARASITES OF MAN terebinth. 20 drops three times daily for a fortnight, then acid, galhc. 0*5 grm. three times a day for two days, then i*o grm. three times a day ; the urine became clear, but whether permanent cure resulted remained doubtful. Hymcnolcpis diminiita, H, lanceolata, Davainea asiaiica, and D. madagascarensis possess no actual clinical interest ; with regard to the latter it need only be pointed out that Bordier^ in studying a case of chyluria found this species in the kidneys of a person in Madagascar. Taenia solium. Tceiiia solium inhabits the small intestine of man ; single proglottides or whole worms may get into the abdominal cavity and the bladder through fistulae, and penetrating the abdominal wall escape outwards or become discharged with the urine. Symptoms of intestinal stenosis are certainly very rare, as in the case recorded by Steinhaus^ of a child, aged 9, the stenosis ceasing after the expulsion of the segments. The usual position of the worm in the small intestine is with the head closely adherent to the mucosa and the proglottides lying along the intestine ; from time to time portions are discharged with the faeces per rectum. Its position can also be reversed, and the proglottides in the gut become thus discharged by vomiting. The diagnosis depends upon the proglottides being generally discharged in pieces in the stools, or eventually an examination for eggs. Larval infection (Cysticercus celhdoscv) occurs also in man through auto-infection or through food. Cysticercus cellidosce of the skin and subcutaneous tissue occurs very seldom singly ; as a rule they are found in hundreds and thousands in the same individual. They occur in different parts of the body, especially on the flexor surfaces of the extremities (generally symmetrically), small globular swellings, the size of a pea or a hazel nut, smooth, of a tough cartilaginous consistence, fairly movable under the skin, in the muscles less so. They never degenerate or cause the surrounding skin to lose its colour. It is an interesting fact that in the case described by Possell^ nodules on the face, namely in the neighbourhood of the left cheek and behind the left ear, reformed. The following are, according to Posselt, characteristic for cutaneous tumours due to cysticerci : (i) the position in the subcutaneous connective tissue (and almost always simultaneously in the muscles) ; (2) the approximately equal size and regularly / Bordier, quoted by Predtetschensky, loc. cit., p. 95. ^ Steinhaus, Deutsch. med. Wochenschr.^ 1903. 8 Posselt, Wien. klin, IVochenschr.y 1899. SUPPLEMENT 663 rounded oval form ; (3) the peculiar density, almost reminding one of cartilage in its hardness and the sensation of tightly distended thick-walled bladders; (4) proportionately slight mobility; (5) with painlessness, absence of any cutaneous reaction (hyperaemia or swelling of the skin or pigmentation). The very gradual appear- ance generally of the tumours supports the diagnosis, and in addition to this evidence we may emphasize the preponderating liability of the upper part of the body to attack and the symmetrical arrangement of the nodules. Cutaneous and muscular cysticerci cause the most varied symptoms, sensory disturbances, abnormal sensations, depression and a feeling of weariness whenever the diseased parts are moved, weakness in the lower extremities, pains in the course of the sciatic nerve, in addition to those which simulate cramp in the calves, numbness in the hands, pains upon their being moved. In the case of a cysticercus situated in the elbow-joint, painful dragging sensation in the course of the ulnar nerve persisted. In other cases the arm was almost paralysed, or it could not be completely extended ; stiffness and bending of the little finger were noticed. Cysticerci of the gluteal muscle cause trouble upon sitting and upon defaecation. Remittent unilateral headaches were present in the case of a cysti- cercus of the region of the right eyebrow ; pains of a neuralgic character radiated from the diseased temporal region. The cysts may be inflamed and may suppurate ; this especially happens in the case of solitary cutaneous and muscle cysticerci. The best treatment consistS'in puncture of the cysts with a Pravaz syrmge and subsequent injection of a drop of i per cent, sublimate solution. Tincture of iodine has similarly been proposed (Wolff^). Frangenheim^ re- commends early extirpation (this, however, only in the case of solitary cysts). Pelagutti^ believes that in his case diminution in the size of the cysts was obtained by the use of anthelminthic remedies continued over a long period combined with potassium iodide and calcium salts (internally). Cysticercus is very rarely found in the tongue ; there the w^orms generally lie in front of the sulcus terminalis, corresponding to the middle of the tongue, according to Glas.^ In the case recorded by Gaetano^ (a boy, aged 10) there was a nodule on the left side of the tongue which grew very rapidly till it reached the size of a nut ; it was embedded in the muscle and covered over by normal mucosa. Cysticerci are just as rare in the pleurae, in the lungs, in the intestinal submucosa, in the submucosa of the small ' Wolff, " Lesser's Encyclop. d. Haut- u. Geschlechtskrankh.," 1900. ^ Frangenheim, Volkni. klin. Vortr., No. 424. ^ Pelagutti, Giorn. ital. delle nial. vener., 1900. ^ Glas, IVien. kliit. IVochenschr.y 1905. ^ Gaetano, Gioin. int. delle Set. med,^ 1904. 664 THE ANIMAL PARASITES OF MAN intestine, in the mesenteric glands, in the liver, pancreas, spleen and kidneys, in the mamma, in the heart, in the bones and in the great vessels (Huber^). Cysticercus of the eye deserves special mention; in rare cases the cysticercus has been met with in the subcutaneous cellular tissue of the eyelid, once in the muscle bundles of the musculus orbicularis. Subconjunctival cysts are found chiefly in youthful individuals. Their position is most varied, generally in the neighbourhood of the inner angle of the eye. Dilated vessels pass right over the cysts, which are generally movable, together with the base they rest upon, producing a spherical protrusion. The head of the worm can sometimes be seen shining through as a whitish speck. The only symptoms are those of a slight irritation of the connective tissue and some difficulty in closing the lid ; larger cysts dislocate the globe. The diagnosis has the rapid growth of the cystic tumour to support it ; there is the possibility of its being mistaken for a foreign body (Kaldrovils^). After division of the connective tissue capsule extraction is easily performed. It is most rare for the cysticercus to occur in the orbit. Suppuration of the cyst may have serious con- sequences for the eye. It is only exceptionally that the cysticerci gain access to the anterior chamber of the eye. Subretinal cysticerci or those localized in the vitreous are more frequent. Upon examination with the ophthalmoscope there is seen in the vitreous a bluish bladder with a smooth surface. The head is seen as a white patch, and the circle of hooks and the suckers also come into view, also the frequent movements which the head and neck make in the vitreous. Operation generally yields good results ; in rare instances the globe is atrophied and must be enucleated. Formerly cysticerci in the brain were met with in fair frequency, but the number of such cases has generally decreased of late years in a remarkable way, in correspondence with the diminution of cysticerci, which is to be attributed to compulsory meat inspection. Whilst, for example, the post-mortem records of the Pathological Institute in Berlin before the year 1875 showed 20 per cent, cysticerci affecting the brain, this number declined later to 16*3 per cent., and of late years has fallen to i per cent. (Orth^). Nevertheless even now cysticercus still plays no inconsiderable part in the etiology of cerebral diseases. For example, in the clinic of de Amicis at Naples, among seven cases of cysticerci of the skin, they were found four times also in the brain (Sipari*). Cysticerci may occur in the dura mater, arachnoid, pia mater, choroid plexus, the surface of the cerebral * Huber, ** Bibliographic der klin. Helminthologie," 1891, pt. 2. - Kaldrovils, Wien. med. Wochenschr.^ ig02. 2 Orth, Berl. med. Ges., June 29, 1904. * Sipari, " Angelo Trani Neapel," 1900. SUPPLEMENT 665 hemisphere, the medullary substance, the ventricles, the aqueduct, the corpus striatum, corpora quadrigemina, the pineal gland, the pons, the cerebellum, the olfactory trigone, the bulb, the medulla oblongata, and the olive. They are most frequently found in the cortical sub- stance and in the ventricles ; the frequency of the latter situation may be explained by the flow of the fluid (Henneberg^). The severity of the symptoms is not always in proportion to the number of cysticerci. Cases have been known in which ten, twenty and forty cysticerci have been found (Hagen-Thorn^), and yet the clinical symptoms have been remarkably slight. On the other hand, solitary cysts may both run a course completely without symptoms and also cause the severest symptoms when located in specially important parts of the brain (crus, pons, central convolutions). In the case mentioned by Jacobson^ the invasion of the brain by cysticerci was immense ; the largest cyst was found in the cerebral cortex. The chief symptoms of cysticercus of the brain substance consist in the onset of cortical epilepsy, which sometimes runs a very pernicious course, frequently with psychical disturbances, whilst paralyses are absent. Perhaps, too, the localization of pain, spontaneous and on pressure, corresponding with the points observed on the cranium, is of importance. Cysticerci may also change their position in the brain ; patients who had earlier suffered from epileptiform convulsions later showed intra-ocular cysticerci after the cerebral symptoms had completely disappeared. Treatment can only be surgical ; v. Bergmann* operated in two cases with well-marked improvement. Parasites in the ventricles are especially dangerous, more especially so when free in the ventricles, and so capable of giving rise to the danger of sudden closure of the foramen of Majendie (Simmonds,^ Verse^). Stern^ states the symptoms of cysticercus in the fourth ventricle to be the following: general cerebral pressure symptoms (headache, vertigo, vomiting, somnolence, congested disc caused by internal hydrocephalus) ; in addition, there are symptoms which point to disease of the hind-brain — pain and stiffness in the neck, vertigo and cerebellar ataxy, violent and persistent vomiting, slowness of pulse ; and lastly those rare but certain symptoms of a lesion of the bulb, such as diabetes, respiratory disturbances and paralysis of cerebral nerves, especially of the abducens. These are far less marked than the general symptoms of cerebral pressure. One characteristic is the remarkable alternation between severe general ' Ilenneberg, Berl. kliii. Wochenschr.y 1906, xxxii. ■■^ Hagen-Thorn, abstract by Posselt. ^ Jacobson, Berl. klin. IVochenschr., igo6. ^ V. Bergmann, quoted by Fiangenheim, loc. cit., p, 470. •'■ Simmonds, Milnch. med. Wochenschr.^ 1907, xxvii. * Verse, Milnch. med. IVochenschr., 1907, xi. ' Stern, Zeitschr. f, klin. Med., Ixi. 666 THE ANIMAL PARASITES OF MAN symptoms and periods of complete sense of well-being ; in this way a functional nervous affection may be simulated (Jolasse^). Brun's symptom (in the widest sense, sudden onset of violent cerebral symptoms upon change of head-posture) is a specially characteristic sign of free cysticercus in the fourth ventricle ; the disease generally terminates with sudden death from cessation of the heart's action. Defects in motor power, convulsions, implication of other nerves, are rare and unessential complications (Hartmann^). Carefully carried out, lumbar puncture may possess some diagnostic and therapeutic value. Treatment is purely symptomatic, or eventually Neisser's ventricle puncture may be considered. At the base of the brain the cysticerci, as a rule, assume that form which is designated as C. racemosns, and consists of rows of delicate grape- like bladders in groups, sometimes also markedly branched, but generally sterile, which develop in the meshes of the soft meninges and may envelop the nerves and vessels of the base of the brain. Such tumours bring about hydrocephalus and chronic leptomeningitis, which must be regarded as the causes of the clinical disturbances (cysticercus meningitis), attacks of loss of conscious- ness, dementia and apathy, dulness and confusion and headaches. In the case recorded by Meyer^ symptoms which resembled paralysis agitans were noteworthy, and defects in speech in the case recorded by Dursf* (C. racemosus in the region of the left Sylvian fossa). According to Markwald^ C. racemosus of the fourth ventricle is said to represent a characteristic clinical picture : violent headaches, attacks of vertigo followed very soon by deep coma and death in a few days. Treatment in Cysticercus racemosus is ineffectual. In the diagnosis of cerebral cysticerci in general the recognition of multiple cysticerci in the skin and muscle and of the tapeworm is of importance. In cases of cerebral diseases in which cysticerci may be a possible cause, Remmert*^ recommends that the skin of the whole body should be palpated. Cysticercus in the spinal cord and in the vertebral column is occasionally observed ; as a rule, other organs, above all the brain and its membranes, are simultaneously affected. Here, too, the cysticercus occurs in two forms— sometimes the cysts are roundish or oval, solitary or multiple, and at other times Cysticercus raceuwsus occurs. ^ Jolasse, Mibu/i. med. Wochenschr., 1896. - Ilartmann, Wien. klin. Wochenschr., 1902. ^ Meyer, Detitsch. tried. Wochenschr., 1906. * Durst, Lieen. viestnik, 1902. ^ Markwald, Milnch. med. Wochenschr.^ 1895. ^ Remmert, "Dissert. Berlin," 1893. SUi'PLEMENT 667 Taenia saginata. Occurs in the small intestine of man. It is characteristic of the habit of life of this parasite that once it has become mature its proglottides are dropped off daily in increasing numbers because its growth is extraordinarily rapid. The joints are discharged generally spontaneously during the whole day without a stool. An extraordinarily unpleasant sensation is produced by the damp, cool joints slipping down into one's lower garments and over one's legs when walking ; women especially, in whom the proglottides slip through their petticoats on to their legs, complain bitterly of this troublesome symptom. Another unpleasant symptom is superadded in the shape of the proglottides tickling the rectum, and this excites irritable people to the last degree. Different species of tapeworms are not mutually exclusive. B. latns and T. solium frequently occur side by side, so also T. sollnm and T. saginata — for instance, in a butcher's assistant w-e once expelled twelve T. solium and one T. saginata at the same time. The greatest number of Taenia3 which have been observed at one time amounted to forty T. solium (Kleefeld^). Even though the cysticercus of T. saginata is not, as in the case of T. solium, particularly dangerous to man, a parasite, nevertheless, which requires so much nutrient material during its rapid growth, and thereby sets up manifold disturbances in the general condition of health, ought to be expelled as rapidly and thoroughly as possible. Tapeworms are found not uncommonly with other intestinal parasites, such as Ascaris, Oxyuris, Trichocephalus or Ancylostoma. Prunac^ described a case in which a woman passed a Taenia through the anus while she vomited a Fasciola hepatica. The symptomatology of these three large species of Cestodes^ Dibothriocephalus latns, Tcenia solimn, and T. saginata, may very well be summarized together, as, apart from some peculiarities, the clinical symptoms, especially so far as their localization in the intestine is concerned, are practically the same for all three species. In a large number of cases the hosts have no suspicion whatever that they are harbouring a tapeworm ; they feel quite well and free from any disquieting symptoms whatever, and only become aware of the fact that they are the carriers of a tapeworm when the discharge of the segments takes place; on the other hand, it is often difficult to rid people of the idea that they are harbouring a Taenia (Kiichenmeister calls such Taenia imaginata) ; usually it is undigested fibrous shreds of beefsteak which are regarded by the patients as proglottides of taeniae. ' Kleefeld, see Seifert loc. cit. ■^ Prunac, see Eichhorst, ** Handb. d. spez. Path. u. Therap.," ii, p. 281. 42 668 THE ANIMAL PARASITES OF MAN In a large number of cases, disturbances of the intestinal tract set in, e.g., sense of pressure in the abdomen, which sometimes becomes constant on one and the same side, or sometimes changes, now at the umbilicus and again at the epigastrium ; here and there colicky pains are present. Derangements of appetite and digestion are frequently complained of ; the most frequent are the sensations of morbid hunger or irregular appetite, nausea and vomiting. Thus, at the Third Congress of Internal Medicine, Senator recorded a case in which there were symptoms of nervous dyspepsia, cured after a successful vermifuge. There is either constipation or diarrhoea, so that many of such patients are brought for treatment with the diagnosis of " chronic intestinal catarrh " and correspondingly treated. As to the treatment of toxic action of the Taeniae when such arises, see the special section on the subject (bothriocephalus anaemia, p. 644). The frequent disturbances of the general condition, so-called reflex phenomena, so far as the action of toxic substances is not in question, may be explained by the fact of their occurrence in specially sensitive individuals who are affected by such phenomena. The proof that a diseased condition is produced by a tapeworm will be forthcoming with some degree of certainty if the symptoms cease immediately after the removal of the parasites. As a whole series of troubles, which certainly have nothing to do with them, are erroneously ascribed to the tapeworm, as is frequently assumed, one will do well to be somewhat critical in this respect. The treatment is of a threefold nature : prophylactic, symptomatic and radical. Under any circumstances, the best prophylaxis is that which consists in only eating the flesh of those animals in which any of the three larval forms occur (pig, cattle, salmon, pike, burbot, etc.) so prepared that the larval forms have been destroyed and the food thus rendered innocuous. For domestic and public use the rule prescribed by Kiichenmeister is under all circumstances most easily understood, namely to roast or boil till the flesh appear greyish- white and sufficiently done by reason of the coagulation of the albumen and decolorization of the blood. The general prophylaxis simply concerns the tapeworm carriers trying to limit as far as possible the further extension of the parasites in the animal world by carefully rendering the expelled segments and worms harmless (pouring sulphuric acid over the faeces and burning the worms) and also by strictly adhering to official regulations. The official system of meat inspection in this respect has been of immense service, and much can still be done by means of thorough official control over cleanliness in abattoirs and butchers' shops. Galli-Valerio^ very ' Galli-Valerio, Therap. Monatsh. , 1900. SUPPLEMENT 669 rightly desires the aboHtion of the custom of manuring fruit-plants such as strawberries, vegetables and salad with the contents of privies, and would extend the use of privies in the country. Symptomatic treatment consists, in the case of those Taeniae which resist radical attempts at expulsion, of repeated use of drugs injurious to the worm as soon as ever new proglottides are formed, or in special cases, as in the case of persons weakened by diseases or operations, or frail old people, or patients with severe heart failure, gastric or intestinal carcinoma, or in pregnancy, in effecting the expulsion of a large chain of proglottides by the mildest measures possible. Radical treatment of the Taenia is not always equally easy in all three species, even when the means used are the same ; the easiest to expel is T. solituHy then D. latus, and the most difficult T. saginata. That as yet no certain cure exists for Cestodes is clear from the large number of drugs recommended from time to time, and the increase of bungling treatment in this respect ; in addition, there is no depart- ment in which there is so much quackery as in vermifuges. The treatment proper should always be preceded by thorough preparatory treatment, the purpose of which is to render the gut as empty as possible once for all, and on the other hand to put the worms them- selves into a diseased condition. How far the host himself has been made ill by such preliminary cures (herring, pickle, garlic, onions, preserved strawberries), many a person who has had to do with such things can recount. In the opinion of Fischer^ strict preparatory treatment appears to favour the development of toxic substances, or else it disposes to vomiting ; as a rule it causes the patient far more discomfort than the treatment itself. In recent times far less weight is attached to these preparatory treatments than to carefully prepared and correctly dosed drugs ; the preparation is generally limited to relieving the intestine in a simple way, the day before the treatment, of the densest faecal masses, by a simple aperient or water enema. We recommend the following, which has always proved itself to be the best and simplest remedy against T. saginata. The pati-crnt takes early in the evening before the treatment nothing but a plate of soup or a glass of milk, and then takes a laxative (electuar. lenit or infus. sennae compos, or an enema), so that later in the evening one to two stools are passed. In this connection we fail to agree with Grawitz^ and Boas,^ who consider that at least preliminary evacua- tion of the intestines can be dispensed with. On the following ' Fischer, Stockholm, Nordin and Josephson, 1904. '^ Grawitz, Munch, med. Wochenschr.^ 1899. ^ Boas, Dtutsch. med. Wochenschr.^ i839> 670 THE ANIMAL PARASITES OF MAN morning the patient should take a cup of black coffee or tea without anything else, and half an hour later the vermifuge. The best drug is extract, filicis maris aether., which also forms the main constituent of most of the secret remedies recommended for tapeworms. Earlier mishaps with this preparation had their origin principally in insufficient dosage. Also, in addition to correct dosage, extract, filic. maris needs very careful preparation if satisfactory results are to be attained. If preparations wnth the trade mark '' Helfenberg " or "Wohnar" are not used, but the male fern extract has been prepared by a chemist, one must make certain that the roots of the Aspidium filix-mas have been collected in May or October, and only green sappy specimens selected, and that the attached paleae have been separated, that they have been broken up small and ether poured over them with a little spirits of wine while quite fresh. The w^hole mass is to be kept in a cool place, but not too closely covered. If at any time a certain quantity is to be used, it is taken out, the ether carefully distilled in a retort till the extract has a suitable fluid consistency. Fischer attaches great importance to the direction in the Pharmacopoeia being exactly follow^ed, to the effect that the extract is to be carefully stirred before prescribing, as the active sub- stances undergo partial crystallization if kept for any length of time and sink to the bottom, so that the preparation has a different strength and toxicity in different layers. Of this extract 10 to 12 to 15 grm. are to be taken in gelatine capsules within half an hour. We consider it unjustifiable to give greater doses than 15 grm. to adults, as many cases are known in which to some extent severe toxic symptoms have followed, such as headache, sensation of giddiness, dyspnoea and cyanosis, yellow^ vision (xanthopsia), delirium, stupor, the most severe cramps in the extremities, rapidly fatal trismus and tetanus. The most serious are defects of vision of various kinds, which may end in amblyopia and amaurosis, with permanent blindness. A complete collection of toxicological literature up to the year 1903 is to be found in Marx's^ Dissertation. Since that time further instances of such intoxications have been made known. NageP observed them only in severe cases. O. Meyer^ lays special stress on the bad prognosis of the disturbances of vision evoked by poisoning \vith extract, filicis maris. Studt^ has seen two cases of optic neuritis, one with circumscribed, the other with diffuse retinal oedema. Uhthoff'' has only seen one case ; in that reported by Noiszewski^ the toxic retinitis w^as cured; in ViereckV case bilateral concentric limitation ' Marx, •• Diss. WUrzburg," 1903, 2 Nagel, Deutsch. med. Wochenschr., 1903. ^ Meyer, O., i5y Liesen. ^ Donati, Ann. Univ. de Mid. et Chir., Milano, 1875. '" Cerchez, Clinica, 1891, iv. " Wagner, Dentsch. med. Wochenschr., 1902. '-' Kabot, Soc. de Sci. 7n6d,de Lyon, September 9, 1904. '•' Negresco, Soc. de Med. legale^ November 9, 1903. 692 / THE ANIMAL PARASITES OF MAN The route by which Ascaridcv obtain access to the urinary passages must remain undecided. Schliiter*' treated a woman, aged 60, with retention of urine. Upon catheterization the hinder end of an Ascaris hung out from the catheter opening ; the anterior end was fixed in tlie tube and the hunen was obstructed. Perliaps in the female sex Ascaridcv travel from the gut into the vulva and from there into the bladder, as they have already been observed in the vagina, where they cause troublesome symptoms (pruritus pudendi). The diagnosis of ascariasis is not in general difficult ; now and then the worms are discharged spontaneously; if not, the ova, which cannot be mistaken, can easily be detected in the faeces upon micro- scopical examination. Epstein's'^ method — namely, on every occasion to obtain fresh material for examination — is much to be recommended. This consists in introducing a Nelaton's catheter into the rectum with a rotatory motion and then drawing it out. A small portion of faeces forced into the catheter opening is more than sufficient to demon- strate the presence of ova of the parasites upon microscopical examination of a preparation. In spite of all pressure on the part of relatives, treatment directly against Ascaridiv should not be carried out until the diagnosis is certain. As regards prophylaxis, much can be done l')y not throwing the worms, when expelled, on to the dung-hill or into the privy, but straightway into the fire. Metschnikoff"^ has issued a warning .against the consumption of unboiled or badly washed vegetables, salad, strawberries, etc., and also against drinking polluted water. For the expulsion of the worms flores cinae were formerly con- sidered the most useful means ; now, however, santonic lactone — santonin — which is prepared from them, is almost universally preferred. By many, especially in practising among children, flores cinae are still recommended in the form of Stork's worm electuary (consisting of flores cinae, rad. jalapae, valerian and oxymel simplex). Guermonprez^ recommends them because he thinks that santonin only excites the worms and consequently causes unpleasant sym- ptoms. Besides, in the form of the above-mentioned electuary, flores cinae can also be given several times daily with raspberry jelly up to 0-5 grm. to 2 grm. (children and adults). Santonin is prescribed either in single doses from 0*03 to 0-05 to o-i grm. with sugar in the form of powder, or else in oily solution. When given in the latter form the absorption of the santonin in the ' Schliiter, Munch, med. IVochenschr., 1902. - Epstein, see Seifert, " Lehrb. d. Kinderkrankh.," p. 273. =* Metschnikoff, Gaz. hebd. de Med. et Chir., igoi. ^ Guermonprez, see Seifert, Detitsch. med. Zeitg., 1885. SUPPLEMENT 693 stomach is excluded and the whole quantity introduced is thus enabled to reach the worms in the intestinal canal. Kiichenmeister^ has already recommended combination of santonin with ol. ricini. Lewin,^ however, states that ol. morrhuae, ol. olivarum, ol. cocos and ol. cinae can also be taken. In prescribing santonin in oily solu- tion Henoch^ also prefers the combination with ol. ricini. According to Lewin's direction the prescription would run as follows : — ^ Santonin ... ... ... ... ... 0*2 grm. 01. ricini. ... ... ... ... ... 20*0 grm. 01. cinae selh. ... ... ... ... gtt. iv. M.,d.s. S., one tablespoonful to be taken iwo to three times. If the patients should manifest a repugnance to castor oil, Starke's ricinus paste may be selected: — ^ Santonin ... ... ... ... ... o*2grm. 01. ricini ... ... ... ... ... 20'0 grm. 01. cinoe seih. ... ... .. ... gtt. iv. Sacch. albi. ... ... ... ... q.s. Pasta mollis. S., to be used for two days. If necessary the first-mentioned mixture might be given in gelatine capsules. Small children should be given 0*025 grm. santonin in warm olive oil slightly sweetened with sugar (a teaspoonful) in the morning ; if in the course of the forenoon specimens of Ascaris escape, a second dose should follow in the afternoon about two hours after the meal. Older children should be given santonin in combination with castor oil or calomel : — ^ Santonini o*oi to o'02 to 0*03 grm. Calomelan ... ... ... ... ... 0*025 grm. Sacch. albi o*5gim. M.f.p. D. tal. dos. X. S., one powder about six, seven, and eight o'clock on three consecutive days. As santonin causes slight toxic symptoms such as urticaria, vomiting, retention of urine, headache, vertigo, yellow vision (xanthopsia), it is in every case advisable to follow with a laxative to expel the drug from the body as speedily as possible. The urine is coloured yellow from one to two days and assumes a scarlet red colour upon the addition of alkalis ; this, however, soon disappears, while it persists in the case of rhubarb and senna. In the place of santonin iodoform in the form of a powder mixed ' Kiichenmeister, loc. cil. - Lew in, see Sti'ett, Deut^ch. vied. Zeitg.^ 1885. ^ Henoch, idem. 694 THE ANIMAL PARASITES OF MAN with bicarbonate of soda is given by Schidlowsky^ in doses up to o*oi to o'o6 grm. three times daily, and a dose of castor oil on the day after the iodoform is given. Thymol in addition to thymol enemas may be tried, in doses up to 0*5 to 2-0 grm. per diem (Calderone,^ Hausmann-^), also /3-naphthol up to o'45 grm. three times daily (Du Bois^), and — ^j^; Benzo-naphthi.l 2-ogrm. Semin cinae... .. ... ... ... logrni. Sacch. albi o^grm. M.jf.p. Divide in pait. seq. xxii. S., three lo five powders daily. (Ferran^), tilmaron oil i'd to 2-0 to 3*0 grm. in gelatine capsules^ according to age (Bodenstein^). Bnining^, ^ recommends the so-called American worm-seed oil, derived from a plant native to the United States, Cheiiopodiuin anfhdminticinn, Gray. It is given in emulsion (ol. chenopodii anthelm. ico grm., vitelli ovi unius, ol. amygd., gi. arab. pulver. aa io*o grm., aq. destill. 200 grm. ; f. emulsio) up to 0*25 to o*5 grm. three times daily at one to two-hourly intervals, or as a pure oil from 8 to 15 drops in sugar and water ; to be followed an hour after the last dose by oleum ricini or pulvis curellae. If no action takes place by the afternoon, a laxative should again be given. The treatment frequently must be repeated the next day. Thelen° appears- to have had good results from this drug. Corsican moss (mousse de Corse), kamala, Artemisia absinthium, valerian, semen sabadillae, have all been supplanted by santonin and at most are used as adjuvants for the latter. Oxyuris vermicularis (Oxyuriasis). Oxytiridce do not remain at rest in the gut, but leave it, generally at night time, to migrate around the anus, into the gluteal folds, and in females into the vulva and vagina and still higher up, giving rise in these different sites to a whole series of irritative symptoms. In the rectum, also, Oxyuridce give rise to such symptoms, which are mani- fested in the form of catarrhal inflammation ; numerous chronic intestinal catarrhs are thus explained. The frequent coincidence of haemorrhoidal troubles with Oxyuridce may be attributed to the fact that the veins of the rectum participate in those changes which have ' Schidlowsky, see Seifert. - Calderone, idon. ^ Hausmann, St. Peter sh. vied. Wochenschr., 1900. ^ Du Bois, see Lenhartz in " Penzoldt-Siintzing's Handbuch," p. 619. ^ Ferran, idejn, ^ Bodenstein, Wien. med. Presse, 1906. ' Bruning, Med. Klin., 1906. ** IJemy Deutsch. med. Woe hens chr., 1907. ^ Thelen, *' Diss. Rostock," 1907. SUPPLEMENT 695 been described as occurring in the intestinal mucosa. Oxyuridcv may also give rise to prolapse of the anus, either by the tenesmus they bring about having such a prolapse as its direct sequel, or the proctitis that supervenes constituting a further etiological factor for its occurrence (Ungar'). Anal fistulae which still fiu-ther increase the trouble, and even rectal listulac;, appear to be capable of onset in consequence of the irritation of the mucosa brought about by Oxy- jiridce (Trendelenburg"^). The conditions recorded by von Wagener^ and Ruffer^ appear to be of interest. At the post-mortem on a child, aged 5, the former found fifteen to twenty quite minute nodules on some Peyer's patches, and in several of these OxynrUhv were found upon microscopical examination between the calcareous concretions within the patches. He presumes that the parasites penetrated the follicular ulcers, and after healing of the latter that they died and became calcified. In the case of a man who died from cirrhosis of the liver, Ruffer found in the rectum, at a distance of about 6 in. from the anal orifice, several tumours covered by the intestinal mucosa, the smallest of which was the size of a pin's head and the largest that of a w^alnut. The tumours looked like calculi overgrown by connective tissue ; under the microscope, countless Oxyuridcv ova were found in their interior. The symptoms of irritation set up by these migrations from the intestine are troublesome to the last degree ; the pruritus thereby induced is often unendurable ; as this irritation from itching comes on with especial severity during the night, the night's rest is grievously interfered with ; many attacks of night terrors appear to be occasioned by these worms. But the general condition suffers as well ; the children become pallid and affected with nervous excitability. Through the act of scratching the irritated parts the ova of the parasites may be conveyed by contaminated fingers directly into the oral or nasal cavities, certainly also into the oral cavity by the contamination of food (auto-infection). In the case of boys the sexual organs may be excited sympathetically through irritation of the sacral nerves of the rectum ; girls may be induced to practise onanism in consequence of the entrance of the worms into the vulva. As a result of the itching irritation which the scratching gives rise to, and of the irritation due to the parasites migrating to the area surrounding the anus, congestion and inflammatory symptoms may arir^e in the peri-anal and perineal regions (weeping eczema, Seifert),^ ' Ungar, see Seifert, " Lehrbucli der Kinderkrankh.," p. 246. - Trendelenburg, see Seifert, f'de/n. 3 von Wagener, Deutsch. Arch. f. Jdhi. Med., Ixxxi. 4 Rufifer, Brit. Med.Journ., 1 90 1. * Seifert, '* Lehrb. d. Kinderkrankh.," and Lesser's " Encyklop. d. Haut-u. Geschlechts- krankh.," p. 373. 696 THE ANIMAL PARASI lES OF MAN and tliese do not abate till after the removal of the oxyuriasis. Some authors speak of an oxyuriasis cutanea (Majoclii^), in the more limited sense of a dermatitis intertriginoides. So far five such cases have been recorded, one each by Szerlecky/ Michelson,^ Majochi/ Barbagallo'' and Vignolo-Lutati.^ Szerlecky's case was that of a young woman with intertrigo over the thighs (the skin was covered as if with leather) ; Michelson's case was tiiat of a boy, aged 13, with intertrigo on the skin of the genito-crural fold, of the scrotum and of the thigh ; Majochi's was that of a man, aged 38, with the same localization ; Barbagallo's case was that of a boy, aged 14, in vviiom the dermatitis extended to the hypogastrium (rhagades on the scrotum) ; and Vignolo-Lutati's case was that of a man, aged 24, with intertrigo of the peri-anal and perineal I'egion, of the scrotum and the inner side of the thigh. On leaving the gut, Oxyuridai frequently migrate to the stomach, to the oesophagus, to the mouth, to the nasopharyngeal cavity, and into the nose (Zarniko') (the localization in the nose has been referred to as associated with the possibility of auto-infection — sec p. 695 as to the development of embryos from the ova in the moist nasal mucosa). Still the occurrence of Oxyiiridcv in the nose is among the greatest of rarities. Chiari^ records the case of a girl, aged 14, who suffered from pains at the root of the nose and in the left side of the forehead; female specimens of Oxyurlsveniiicularis were evacuated from her nose on several occasions. A similar case is recorded by Hartmann^ ; it was that of a girl, aged 13, with epileptiform convulsions and psychic disturbances ; numerous Oxyurides frequently escaped from her nose. With their departure the symptoms of irritation of the central nervous system also dis- appeared. Rheins^^ records a case, that of a woman, in which a specimen of Oxyurls vennicularis was discharged from the right nostril during the act of sneezing. Proskauer^^ found in the nose of a woman, aged 30, a conglomerate of from fifteen to twenty very small worms which proved to be Oxyuris embryos. The diagnosis of oxyuriasis is not difficult to make, as the trouble- some sensations in the anus and about the genitals necessarily suggest the presence of Oxvnrldcv. As a rule the small white worms are ' Majochi, Boll. d. Sci. vied. d. Bologna, 1893. - Szerlecky, Jouni. Ann. Med. praf., Paris, 1874. ■' Michelson, Bej-l. klin. IVochenschr., 1877, xxxiii. * Majochi, loc. cit. ^ Barbagallo, Gaz. d. Osp., November 16, 1900. •^ Vignolo-Lutali, Atch.f. De>-7ti., Ixxxvii, pt. I. '^ Zarniko, " Die Krankh. d. Nase, u.s.w," S. Karger, Berlin, 1905. " Chiari, " Erfahr. auf d. Gebiete der Hals-u. Naseiikrankh.," Wien, 1887. ® Harlmann, Natti'foischerversainmi., Koln, 1889. '" Rheins, " Der prakt. Arzt.," 1893. " Proskauer, Zeituhr.f. Ohienheilk., 1891. SUPPLliMliNT 697 seen crawling about over recently evacuated faeces, or the ova are found upon microscopical examination of soiled matter adhering to the anus, or in scrapings removed with the spatula from the surface of the skin (in the case of oxyuriasis cutanea). Prophylaxis has to be directed to infection with Oxyurides generally, on the one hand, and, on the other, to the possibility of auto-infection. With reference to the first-mentioned point, Metsch- nikoff's^ directions should be borne in mind, to the effect that badly washed vegetables, salad, etc., ought not to be eaten (vegetables to be rinsed with boiling water), and also that the members of the family of the diseased individual should be examined for Oxyitridoe and eventually be treated (Heller^). With regard to the second point, one has to observe strict cleanliness in general (Barbagallo^ found ova of the parasites in the layer of dirt under the finger-nails). Treatment of oxyuriasis must be of a twofold nature ; first, medicinal, the administration per os of vermicidal drugs in combina- tion with purgatives ; and secondly, local treatment of the gut by means of enemata, suppositories and high injections. Following the method prescribed by Ungar,* pulv. glycyrrhizae co. is first given in the case of smaller children, castor oil or calomel m that of those older, in order to evacuate the intestine, and four times daily on two days following one another a dose of naphthalin, not du'ectly after meal-time, but as far as possible in the interval between two meals, and at the same time the ingestion of fatty or oily nutriment is as far as possible to be avoided. After eight days this treatment should be repeated, and under certain circumstances once again after a further interval of a fortnight. The dose varies between 0-05 and 01 grm. (children of i year old), 0"i toO'2grm. (children of 2 to 3 years old) and 0*2 to 0*4 grm. (children of 4 to 10 years old). Dornbluth^ employs the same medicament in a form only slightly modified from Ungar's method, Barbagallo- gives internally only a purgative (decoct, sennae cum natr. sulfur). Thymol, santonin, kousso, kamala or valerian may be tried instead of naphthalin. For enemata the following are employed : naphthalin in a solution of i in 50, ol. olivar. or thymol o*i in 200 aq. destill., diluted solutions of lysol, menthol in ^ per cent, oily solution, salicylate of soda in watery solution, decoctum tannaceti with santonin, with the addition -of some drops of ol. terebinth. (Barbagallo). Decoctions of garlic, infusion of valerian, sulphur water (sublimate is to be avoided), aq. calcariae, ol. olivanim ' Metschnikoff, Med. Klin., 1907, xlii, p. 1284. - Heller, Deutsch. Arch. f. klin. Med., Ixxvii. * Barbagallo, loc. cit. ^ Ungar, see Seifert, " Lehrb. d. Kinderkrankh," • Dornbluth, Arzil. Zenlral-Anzeiger, 1903. *5 Barbagallo, loc. cit. 6g\< THE ANIMAL PARASITIiS OF MAN cainphoratum (Vignolo-Lutati). Santonin o'l grm. is the best to employ for suppositories. For high injections, large quantities of plain water are employed (2 to 4 litres), or soapy water (0*2 to 0*5 per cent, solution of sapo medicatus, Heller,^ StilP), J per cent, salicylic acid solution or liq. alum. acet. (one tablespoonful to a litre of water, Dornbliith^), or gujanosol (2 to 3 to 4 to 5 per cent, solution, Rahn^). The employ- ment of benzine for such high injections is not advisable according to the experience of Senger,-^ owing to the symptoms of poisoning after the external application of benzine, at least not in the case of young children. That diseases of the intestine which are accompanied by frequent thin fluid evacuations may lead to recovery from oxyuriasis has frequently been observed by us in the case of young children who have suffered from dysentery (Seifert^). Inunctions of cod-liver oil appear to be very valuable in the treatment of oxyuriasis (Szerlecky, Vignolo-Lutati), whilst those with mercurial ointment may easily increase the inflammatory symptoms. The luxury recommended by Esser,^ that patients every evening before going to sleep should have the female Oxyuridcv picked from the anal fold in the knee- elbow position is one which is certainly only in the power of a few people to carry into execution. An essay has been published by Hippius and Lewinson [Dsztlsch. vied. Wochensckr., 1907, xliii.) in which the relationship of Oxytcrida to appendicitis is considered and the treat- ment of oxyuriasis is discussed. The instructive case recorded appears to show that germs through Oxyicrida gain access to the tissue of the appendix, and, indeed, are carried in by them. In view of this more recent communication as to the part whicii intestinal parasites play in the etiology of appendicitis, it seemed to me [O. S.] to be worth whi e to interrogate my surgical colleagues as to this point. About 2,000 appendicectomies have been jointly per- formed by Drs. Burkhardt, Enderlen, Pretzfelder, Riedinger, Rosenberger and Siber, and in not one of these cases could entozoa be found to be a possible cause of the appendicitis. Such figures without doubt speak in favour of the fact that even if in individual cases entozoa might come into reckoning as a possible cause, such an etiological factor must be classed among the greatest of rarities. My colleague, Dr. Ries, who practised for ten years in Mexico, informed me that there practically speaking every Indian without exception harboured para- sites of the most varied kind, and that in spite of the very extensive professional standing he enjoyed among these people he never had under observation among them a single case of appendicitis. As far as the observation of the authors in question as to the treatment of oxyuriasis is concerned, it must be energetically directed to the employment of local measures for the intestine ; they maintain that the use of enemata would be irrational, and that it is -astonishing that this method has been able to maintain its standing down to the present day. ' Heller, loc. cii. - Still, Br a. Med. Journ., 1899. ^ Dornbltith, loc. cit. * Rahn, Miinch. med. Wochenschr., 1 905. * Senger, Berl. klin. Wochenschr., 1907, xxxviii. ^ Seifert, Deutsch. med. Zeilg., 1885. "'■ Esser, Schweiz. Korrespondejizbl . , 1 893- SUPPLEMENT 699 HIRUDINEI (Leeches). The only one of the leeches that comes under consideration from the clinical point of view is Limnatis nilotica [Hcemopsis sanguisitga), which obtains access to the mouth with drinking water, and becomes lodged, even in the case of man, in the pharynx, larynx, trachea, oesophagus and nose. Amongst the causes of severe haemorrhage from the pharynx Jurasz^ mentions the occurrence of leeches in that region : in Northern Europe this must be accounted one of the greatest of rarities, whilst at all times in southern countries, such as South Italy, Spain, Greece, Algiers, Tunis and Egypt, it appears to have been more frequent. Even the physicians of antiquity had much to say about it. Upon the occurrence of blood-stained expectoration, Hippocrates recommends the oral cavity to be examined to see whether a leech is not present in it. Galen speaks of haematemesis due to the presence of leeches in the pharynx and stomach. Similar mention is found in the writings of Celsus,, Asclepiades, Scribonius Largus, Dioscorides, Aetius, Oribasius, Paulus Aegineta and others. In recent times, CortiaF has published observations relating to this subject which he had the opportunity of making in Constantine. Palazzolo^ also in Sicily found leeches in two cases in the pharynx, in one case on the posterior wall, in the other in the crypt over the left tonsil. According to Roset,* leeches adhere by preference behind the uvula, simulating haematemesis and haemoptysis, and the persistent haemor- rhages they give rise to may lead to severe anaemia. Leeches are found in still greater frequency in the larynx than in the pharyngeal cavity. Huber^ records several observations of this kind in his historical and therapeutical study. In the case of a man, aged 64, Ramon de la Sota y Lastra® observed a leech on the nodulus epiglottidis ; this was removed with the forceps. In the case recorded by Photiades,^ a leech had remained adherent to the vocal cord for more than twenty-two days. Maissurianz^ records two such cases : in one the leech had remained in the sinus morgagni for three weeks,, in the other in the same place for ten days. The case recorded by * Jurasz, Heymann's " Handb. d. Laryng. u. Rhinol.," 1899, ii, - Cortial, Union mid.^ 1886. ^ Palazzolo, Bull. del. vial. deW orecchioy etc, 1895. * Roset, Rev. d. Cienc. niM. de Barcelona, 1 907, ii. * Huber, Deutsch: Arch. f. klin. Med., xlvii. 6 Ramon de la Sota y Lastra, Rev. vied, de Sevilla, 1883. ■^ Photiades, Int. Zentralbl. f. Laryng., 1884. ^ Maissurianz, St. Petersb. vied. Wochenschr., 1883. 44 700 THE ANIMAL PARASITES OF MAN Schmolitschew^ is an interesting one ; it was that of a woman who for four days had suffered from violent haem^optysis, the cause of which was a leech that was fixed on the laryngeal wall of the epiglottis close above the vocal cords. In his case (that of a soldier), Godet^ was forced to perform thyrotomy to remove the leech from the larynx. Ficano^ removed a live leech wdth the forceps from the lower laryngeal cavity in a man, aged 30. Massei* reports a similar case. The case reported by Winternitz and Karbinski^ was that of a peasant girl, aged 16, who suffered from coughing, hoarseness, and blood-stained expectoration ; a leech had lodged on the root of the epiglottis. Aubert^ removed a leech from the larynx of a woman after the performance of tracheotomy. Seifert^ reports three cases : in the first the leech had become fixed to the left vocal cord, in the second it was found in the lower laryngeal cavity, and in the third on the border of the left ligamentum aryepiglotticum. Leone^ has published the case of a leech in the larynx, Martin^ two cases with the leech lodged in the lower laryngeal cavity, Be'rthoud^^ a similar case, Palazzolo^^ two such cases, Panzat^^ one case (lower laryngeal cavity). Moucharinski^^ reports a case in which the leech had stayed more than twenty days in the larynx. Martin^* easily removed a leech from the posterior portion of the vocal cord with the forceps. Vieus and Nepeon^^ record a case of a leech in the larynx. It is quite exceptional for leeches to gain access to the trachea ; cases of this kind have been recorded by Aubert,^^ Vicano,^^ Ridola^^ and Tapin^^ (the leech was firmly fixed to the bifurcation and caused coughing, haemoptysis and attacks of asphyxia ; it was easily removed by the aid of a tracheal tube). Now and then leeches are found in the nose. Lusitanus^^ relates the case of a man who suffered from severe headaches. A medical man ordered the application of a leech to the anterior portion of the nostril. Owing to the carelessness of the ' Schmolitschew, Wratsch, 1884. 2 Godet, Arch, de Med. et Pharm. milit., 1887. » Ficano, Rev. de Laryng., 1890. * Massei, Int. Journ. of Laryng., 1890. ^ Winternitz and Karbinski, Prag. vied. Wochenschr.^ 1890. « Aubert, Echo med., 1891. '^ Seifert, Rev. de Laryng., 1893. ^ Leone, Boll. del. mal. delP orecchio, etc., 1892. ® Martin, Arch, de Med. et Pharm. milit., 1891. *o Berlhoud, ibid., 1893. " Palazzolo, Boll. del. mal. delt orecchio, 1895. '2 Panzat, Arch, de Mid. et Pharm. milit., 1896. " Moucharinski, Wratsch, 1896. 1^ Martin, Rev. barcelon de enf. de oido, 1906. '^ Vieus and Nepeon, Monatsschr. f. Ohrenheilk., 1884. *6 Aubert, Echo med., October 12, 1891. '■^ Vicano, Boll. del. mal. deW orecchio, etc., 1892, ix. '^ Ridola, Arch. ital. di Laryng., 1894, ii. '^ Tapin, Siglo med., March 16, 1907. ^ Lusitanus, see Seifert in Ileymann's " Handb.," p. 599. SUPPLEMENT 701 surgeon the leech crawled right into the nose ; it was impossible to extract the leech or to kill it, and it produced a severe haemorrhage which led to the death of the patient within two days. In a case recorded by Sinclair/ a leech, Ha'tnopsis sangiiisuga, gained access to the nose of a boy, aged 3 ; it remained there a fortnight ; it caused frequent attacks of epistaxis and in the end it was removed by means of forceps. Condorelli-Francaviglia^ records a case in which severe epistaxis was caused by a leech which had probably entered the anterior portion of the left nostril by way of the pharynx and become tightly fixed there. It was seen by posterior rhinoscopy, and was removed from in front by means of slightly curved forceps. Sota y Lastra^ mentions the occurrence of leeches in the nose, and Keng^ reports the case of nasal obstruction from a leech. The removal of leeches is effected by means of injections or by the direct sprinkling of salt or acid solutions on their bodies, which brings about their detachment. When possible a previous attempt should be made to seize them with forceps so as to make their immediate extraction possible. The species of Haemadipsa (Looss*^) live in tropical regions in moist places on the ground or in the jungle. They climb bushes and even trees with astonishing rapidity upon the approach of larger animals and also of man (whom they clearly recognize from the vibration of the ground caused by footsteps). From thence they let themselves fall on their victims to suck their blood. Their bites are generally painless, and of themselves not dangerous, but if they are unusually numerous they rapidly accumulate on the body in large numbers and give rise to marked debility and, if the wound become infected, to severe complications and even death. On the other hand, under careful treatment the wounds heal easily and fairly rapidly. Firm leather and firmly adhering clothes afford no certain pro- tection against the attacks of these leeches, as they know how to force themselves with extraordinary rapidity through the narrowest interstices betw^een the clothes and thus gain access to the skin. When they have sucked their fill — and this may take several hours to accomplish — they fall off of themselves. To effect an earlier removal drops of irritative or corrosive fluids are employed (salt solutions, acids, etc.). Tearing away the leech by force should be avoided, as in this way portions of the leech's body may be left behind in the wound and inflammation be set up. 1 Sinclair, BriL Med. Journ., June 20, 1885, i. ■^ Condorelli- Franca viglia, Spallangini, 1892. '^ Sota y Lastra, Rev. vied, de Sevilla, 1887. * Keng, Scot. Med. and Surg. Journ.^ October, 1899. « Looss, " Handb. d. Tropenkrankh.," v. Mense, i, p. 194. 702 THE ANIMAL PARASITES OF MAN ARTHROPODA. Leptus autumnalis (Grass, Harvest, or Gooseberry Mite^). In the hot season of the year, that is, during the months of July and August, it is noticed that those people who stray amongst syringa bushes or who pick gooseberries or kidney beans are attacked by the Leptus autninnalis. On the uncovered parts of the body there appear numerous red spots and papules, which itch and burn smartly. The itching does not commence diffusely, as in the case of scabies (MacLennars^), but is Hmited to the particular points where the parasite is situated. There are especial outbreaks of itching in the morning, arising perhaps from the hatching of ova in the host after lying in the warmth of the bed.^ Leptus fre- quently provokes general erythema, eczematization or severe feverish urticaria, which in France is known by the narne of fievre de grain (Megnieu, Besnier^). If the individual efflorescences be carefully examined, there will be noticed almost without exception a minute boss towards the centre, noticeable by its yellowish-red colour. If an attempt is made to remove it w^ith the point of a needle or to scrape it off the surface, one can often perceive, even with the naked eye, a small reddish creature moving actively about. The treatment of these very troublesome symptoms consists in warm baths with soapy lavages, also lavages with alcohol, spirit salmiac (G. P.), 5 per cent, carbol or creolin solution, diluted vinegar, benzine, emulsions of balsam of Peru, rubbing in sulphur ointment (Sandwith'^) ; ointments of creosote or eucalyptus are recommended. Other grass and grain mites also occasionally penetrate the skin of man and produce transi- tory but sometimes very severe eruptions, urticaria and eczema papulosum, as Geber^ and subsequent to him Josai' have reported of the barley mite. In sensitive individuals the skin becomes bright red, to a greater or less extent their temperature is raised and fre- quently slight febrile affections are present. If the inflammatory ' There is no reason for calling this the gooseberry mite. It is rarely found on this fruit. The gooseberry mite is Bryolia pi-etiosa. - MacLennars, Lancet, 1905. ' [This cannot be the case, as Leptus aututnnalis is the larval form of Trombidiiun hola- sericeum. — F. V. T.] ^ Sack, "Handb. d. Hautkrankh.," v. Mracek, 1907. * Sand with, Lancet, 1905. ^ Geber, " Ilandbuch d. Hautkrankh.," in v. Ziemssen's " Handbuch d. spez. Pathol, u. Therap.," 1884, xiv. ^ Josai. SUPPLEMENT 703 skin symptoms have reached their culminating point after three or four days and no fresh comphcations arise, they only remain for a short while, the effects of scratching and pigment spots being left. Kedani, Akaneesch (The Japanese River or Inundation Disease). This disease is only known in Japan, and is limited to the neighbourhood of some great rivers on the west coast. The people mostly attacked are those who cut the hemp harvest in the infected localities, occasionally those who transport it or come into contact with it (Looss^). The disease is frequently manifested in the form of indefinite disturbances of the general condition ; it commences generally on the sixth day after the presumed infection with rigors, headaches, feeling of weakness, sw^elling of the lymphatic glands in the loin or in the arm-pits ; in the periphery a black dry scab is formed. In addition there is an intense conjunctivitis, and added to symptoms of fever an exanthema resembling measles that lasts from four to seven days. There is frequent delirium and difHculty of hearing which persist for a long while. Obstinate constipation is a striking symptom. At the end of a fortnight, earlier in slighter cases, the fever commences to abate and a rapid convalescence sets in. In pregnant women abortion with fatal issue is frequent. With regard to prophylaxis, Baelz^ recommends as rapid a cultivation of the soil as possible, which has led to a speedy disappearance of the disease . in districts where it was once dreaded. Treatment is symptomatic. Japanese do not tolerate antipyretic drugs as well as Europeans. Dermanyssus gallinae (avium). During the day the resort of bird mites is in the droppings and in the woodwork, etc., of cages in which canaries, crossbills and parrots are kept ; in the crevices of doors, in the chinks between the board planks of bedsteads, so that at night they may seek some domestic animal to suck the blood and so satisfy their hunger. It is by no means rare for young animals, chickens and unfledged pigeons, etc., to perish in consequence of the great loss of blood. This nocturnal habit of life explains why no mites can be found during the day in spite of the most careful examination of the human body, to which they may be transmitted. On the uncovered parts of the body they not only cause severe irritation, but also severe diffuse itching erythema and eczema. Thorough disinfection of the ' Looss, '• Handbuch d. Tropenkrankh.," v. Mense, p. 195. - Baelz, Virchow's Archiv, Ixxviii. 704 THE ANIMAL PARASITES OF MAN cages by hot solution of caustic potash, in addition, sprinkHng over with tar, red carbohc acid or petroleum, thoroughly powdering over the birds with flores pyrethrae, washing with water containing oleum anisi, washing the walls, doors and bedsteads with soap, disinfection of the mattresses, linen and clothes, will protect against further infection. In the case of man the disease needs no special treatment, as the eruptions generally disappear after some days. Heinecke^ recommends lavages with i per cent, carbolic acid solution. \_Vide also p. 492 in body of this work. — F. V. T.] [Dermanyssus hirundinis, Hermann, is identical with this species. By far the best treatment is with paraffin or kerosene oil applied to the places where they pass the day. — F. V. T.] Ixodes reduvius (ricinus). The female is occasionally transmitted to the human skin, and bores its proboscis deep into it and sucks itself full of blood. At sensitive points of the cutaneous surface — for example over the skin of the penis — a feeling of severe pain is produced. Buy's^ observations as to the geographical distribution of the Ixodina show that in all lands in which cattle, horses, sheep and dogs exist, Ixodince are to be found. Recent observations show that the Ixodince play an important part in the transmission of Hsemosporidia {vide body of work, pp. 493, 494). Sprinkling with oil, vaseline, benzine, ether, petroleum, naphtha, turpentine (Jelgenum^), will easily lead to the removal of the parasite ; if the body is torn away with violence and the proboscis is left sticking in the skin, the presence of the latter will give rise to inflammation and suppuration. Sarcoptes scabiei (Scabies). The disease produced by Sarcoptes scabiei shows itself in poly- morphous areas, such as accompany eczema, and are produced on the one hand by the Sarcoptes alone and on the other hand by the scratching with the nails. The localization of both kinds of efflorescences is different from those which are produced by the Sarcoptes ; they occur as papules, vesicles, pustules and mite-tracks, and their usual situation is between the fingers, on the ulnar border of the hand, on the wrist, on the palm of the hand, on the anterior border of the axilla, on the penis and at the base of the thorax. The excoriations are situated on the forearm, over the thigh, over the abdomen, and may be distributed in greater or less degree over the ' Heinecke, Milnch. med. Wochenschr., 1901. - Buy, " Histoire naturelle et medicale des Ixodes," "These de Lyon," 1906. ^ Jelgenum, Med. Weekbladv. Noord- en Ztiid-Nederland, 1901, i, No. 24. SUPPLEMENT 705 whole body ; the back and the face only remain free. The symptoms consist in violent itching, the onset of which specially takes place at night. The mite-tracks are fine curving lines, curved like a, u, c, or s, which appear as if they had been scratched with a fine needle. Upon closer examination with the magnifying glass one sees in their course sm^ll openings. These openings, in persons who keep themselves clean, are scarcely coloured ; but in patients whose occupations necessitate their being associated with coloured or dirty substances, they are dark. The length of the tracks varies from some millimetres to 1 J to 2 cm. They are at the one end, where the Sarcoptes is embedded in the epidermis, widened like a funnel and slightly exfoliated. The track at this point is sharply defined ; the mite shows through the epidermis as a yellowish round point. In the course of the track there develop papulae, vesicles or pustules, which raise the level of the track. The intensity of these inflammatory appearances depends upon the susceptibility of the human individual and upon the capability of the reaction of the skin. There are people in whom scarcely any inflammatory symptoms make their appearance ; on the other hand there are some, especially children and lymphatic individuals^ in whom severe impetiginous ecthymatous pustules, together with their sequelae, are set up. The results produced by scratching consist in papules, which usually bear a small scab of blood, and are arranged in the form of striae, in eczematous surfaces, weeping or sanguineous scabs, vesicles, pustules, etc. The complications that set in are frequently urticaria and even furuncles, lymphangitis and inflammation of the glands, which now and then is followed by the formation of abscesses in the glands. The duration of the disease is unlimited ; when untreated it leads to a form of rare occurrence, that of scabies norvegica^ ; in this the collection of crusts and scales, in which a quantity of dead mites, larvae and ova are present, may become colossal. The symptoms of scabies abate in the presence of intercurrent acute diseases and reappear after the malady is over. The fact has for long contributed to the idea of scabies being regarded as a disease capable of being ^* driven in" upon the internal organs and forming metastases. The diagnosis is rendered certain upon the discovery of a track. Traces of scratching on the extremities and on the abdomen, papular or pustular efflorescences between the fingers, toes, in the neighbour- hood of the wrist, of the elbow, on the anterior border of the arm-pit, on the tuber ischii, in the girdle region, and especially the presence of * [This is produced by a distinct species, vide pp. 519-20. — F. V. T.] 706 THE ANIMAL PARASITES OF MAN disintegrated tracts over the penis (prepuce and glans), will allow of the diagnosis being made. Certain occupational eczemas (grocers, lime- workers, maltsters, bakers and others), also prurigo, must be borne in mind when diagnosing this disease. The prognosis is always a favourable one. Even after such a long duration and after such severe symptoms the disease may completely clear up. There are, however, frequently left behind post- scabious inflammatous and pruriginous conditions which only yield after protracted treatment. Scabiophilia, which persists in certain patients for a long time after the scabies has been cured, must here be mentioned. In the treatment of scabies four points must be kept in view, (i) The mites and the ova must be killed by the treatment ; (2) the treatment must have regard to the intensity of the inflammatory symptoms ; (3) the clothes (body-linen) of the patients must be disinfected ; the bed-linen, the beds and the bedsteads must be cleansed ; (4) when a person suffers from scabies his entourage must be examined, and all diseased conditions treated in the same way as under (3). The treatment (i) should be preceded by a bath wath thorough soap ablution, and when the inflammatory symptoms are not too severe, with green soap. After the bath the skin is dried and the scabies remedy proper applied in warmth. Sulphur preparations receive first consideration ; among such Vlemingkz's mixture occupies a prominent position ; this is rubbed in for half an hour by means of a strong camel-hair brush, to be followed by another bath and powder applications after drying. Repeat this method for three days one after the other, or for two days, and a third time eight days later. The latter method is worthy of recommendation as the ov^a, which perhaps resist the parasiticide action, have by this time developed into larvae, and the latter can then be destroyed with certainty. The remaining sulphur preparations, which are specially employed in the form of ointments, are more complex, as the ointment should remain on the skin. Helmerisch's and Wilkinson's ointments are the kinds specially employed. Nagelschmidt^ recommends thiopinol as a very suitable sulphur preparation in the form of baths or as a 10 or 5 per cent, ointment in the following way : Upon his reception the patient is given a thiopinol bath, in which he remains for thirty minutes. Immediately afterwards 30 to 40 grm. 10 per cent, thiopinol vaseline is carefully rubbed in. The rubbing is repeated daily, and the treatment is concluded on the second to fourth day with a second thiopinol bath. Thiopinol produces no more irritation than the ordinary sulphur ointments ; it is, however, much more penetrative and more capable of absorption. ' Nagelschmidt, Med. Klin., 1907, xxxv. SUPPLEMENT 707 We frequently make use of Kaposi's naphthol ointment, as it renders the skin supple, causes proportionately little irritation, and has but little smell. Treatment with balsam of Peru is certainly expensive, but in the slighter attacks it is relatively the simplest. We give the patient a bath, have him thoroughly dried and rub in 30 to 40 to 50 grm. balsam of Peru carefully and evenly all over, wrap him in a covering of wool, and make him rest in bed for twelve to fifteen hours, to be followed by a bath with careful cleansing with soap ; this treatment need rarely be repeated. The balsam of Peru can be applied undiluted for the rubbings or mixed with ung. glycerini, or resorbin or glycerine in equal parts. [Norman Walker uses balsam of Peru J oz. dissolved in rectified spirit ; to be painted on with a brush.] — ]. P. S. The manufacturers name the undiluted product of the active constituent of balsam of Peru, benzoic acid benzyl-ester, Peruscabin. For the treatment of scabies it is recom- mended by Sachs^ that it should only be administered when mixed with ricinus oil, under the name of Peru oil, in applications repeated three times within thirty-six hours. Sack"2 also considers Peru oil a non-irritant, effectual, pleasant, inodorous and non-staining drug. But he only allows the applica- tions to be used every twelve hours for three to four consecutive days (altogether 200 to 300 grm. of Peru oil are requisite), and after the sixth or seventh rubbing a bath should be taken with the use of Dutch soap. Juliusberg^ considers this treatment specially suited for private practice. Another modern drug is epicarin (/3-oxy- naphthyl-ortho-oxy-meta-tolyol acid) ; this is applied in 10 to 20 per cent, ointments (Pfeiffenberger^), epicarin 7*0 grm., cretae alb. 2*0 grm., vasel. flavi 30*0 grm., lanolin i5"o grm., axungia poric. 45*0 grm. (Rille^) ; epicarin 15-0 grm., sapon. virid. 5*0 grm., axung. poric. 100*0 grm., cretae alb. ico grm. (Kraus^) ; for children, epicarin 5*0 grm., lanolin 90*0 grm., ol. olivar. lo'o grm. (Kaposi'). Siebert^ lays stress upon the odourlessness and colourlessness of epicarm ointment as a strong reason for its use, and points out that it is a harmless drug, the action of which is certain. Endermol (salicylic acid ointment) has a destructive action on the mites even in a O'X per cent, ointment (Wolters,^ Demitsch^^) ; it is, however, very expensive and not wholly free from danger ; and the same applies to nicotiana soap (Taenzer,^' Schumann^^j. ' Sachs, Deutsche med. Wochenschr.^ 1900. '^ Sack, " Handb. d. Hautkrankh.," v. Mracek. '^ Juliusberg, Therap. Monatsh., 1901. ^ Pfeiffenberger, Klin, therap. Wochenschr., 1900. '' Rille, " Die Heilkunde," 1900. ^ Kraus, Allg. wien. med. Zeit.^ 1900. '^ Kaposi, Wien. nied. Wochenschr.^ 1 900. •* Siebert, Munch, med. Wochenschr.^ 1900. '-^ Wolters, Therap. Monatsh., 1898. '° Demitsch, Wratsch, 1905, iv. " Taenzer, Monatsh. f. prakt. Derm., xxi. ^'^ Schumann, Allg. med. Central-Zeitg., 1901. yoS THE ANIMAL PARASITES OF MAN To give an account in detail of the drugs and methods— old and new — used in the treatment of scabies would far outrun the limits of this work. Demodex folliculorum. It is not yet certain whether the Demodex folliculorum is capable of developing pathological conditions in man. VeieP assumes that the hair follicle mite has no connection either with the formation of comedones or even with sebaceous gland disease. Kaposi^ con- siders that they cause no disease in man and cannot be regarded as a cause of acne. Saalfeld^ clearly adheres to the same standpoint, similarly so Jessner,* who, when discussing comedones, makes no mention of acne of hair follicle mites. WeyF and Geber^ adhere to the opinion that the presence of a Demodex in man in contradis- tinction to its presence in animals possesses absolutely no pathogenic influence. On the other hand de Amicis,' Majochi,^ and Dubreuilh'^ report single cases of pronounced circumscribed clear brown pigmentations which they attribute to Demodex follicidorum. In all these cases, moreover, as regards localization the affection had a certain resemblance to pityriasis versicolor ; nevertheless, in the scales separated off with the scalpel no fungi were found, but on the other hand Demodices in moderate quantity. In his earlier cases Majochi has seen the Demodex in the secretion from meibomian glands and had claimed it to be the excitant of chalazion and, as Mibelli^^ did,, considered it to be the cause of some diseases of the eyelids. Ivers^^ found the parasite in 69 per cent, of normal borders of the eyelids, and attributes a pathological signification to it. Hiinsche^^ and Mulder^^ arrive at the same conclusions ; in the light of their investiga- tions the Demodex is found as a constant accessory — certainly not in the meibomian glands, as it is limited only to the internal part of the hair follicle. Lewandowsky^* considers that it can hardly be demon- strated at present that the same parasite which in individual specimens causes no symptoms is capable of producing pathological conditions when markedly increased in numbers. Treatment is by the removal of the comedones, above all, by their mechanical removal by pressure with a watch-key and with the 1 Veiel, V. Ziemssen's " Handb. d. spez. Path. u. Therap.," 1884, xiv. 2 Kaposi, " Path. u. Therap. d. Hautkrankh.," 1899. 3 Saalfeld, Lesser's " Encyclop. d. Haut- u. Geschlechtskrankh.," 1900. * Jessner, *' Kompend. d. Hautkrankh.," 1906, 3rd ed. *, ^ Weyl and Geber, v. Ziemssen's " Handb. d. spez. Path. u. Therap.," 1884, xiv. ■^ de Amicis, quoted by Lewandowsky. *^ Majochi, Centralbl.f. Bakt., xxv. ^ Dubreuilh, La Prat. Derm., Paris, 1901. '° Mibelli, quoted by Lewandowsky. " Ivers, ibid. '■^ Hiinsche, MiXnch. fried. Wochenschr., 1900, xiv. '3 Mulder, Weekbl. v. het Nederl. Itjdschr. v. Geneesk., 1889. '* Lewandowsky, Deutsch. med. Wochenschr. ^ 1907, xx. SUPPLEMENT 709 various comedo-compressors, and by subsequent cleansing of the skin with ether, benzine or spirit. If the eyehds should be affected with blepharitis due to the presence of Demodex in large numbers, epilation and administration of a parasiticide is recommended. Dennodex folliculorunn canis. Transmission from dog to man is in any case very rare, and by man}^ its occurrence is generally doubted. Nevertheless Gruby^ and Remak^ claim that it is transmissible — an opinion which has also been shared by Neumann^ and Ziirn.* The latter saw in the case of a married couple who had the care of mangy dogs the onset of diseased areas on their hands and feet, which were like those on the dogs and contained the same parasites. A. Babes^ also reports several observations which go to show that persons who, to some extent, have been shown to have been in contact with mange-stricken dogs have been attacked by a scabies- like eruption localized over the thorax, abdomen, back and extremi- ties ; large numbers of Demodices were found in the follicular pustules. Lewandowsky^ reports one case — that of an Italian workman, who suffered from an outbreak on the face, like impetigo ; there was crust formation and at the edge of the crusts the epidermis appeared like a narrow row or border of vesicles. A small portion of the covering of the row of vesicles was lifted off, and after slight warming examined in 40 per cent, liquor potassae. In this a large number of animal parasites of the Demodex group were found, and without doubt Demodex folliciiloruni canis alone. Hiinsche^ assumes that Demodex folliculorum penetrates into the tissues and produces abscesses. Treatment first consisted in dusting with zinc amyl powder, but after four days there was no change. After the regular use of xero- form as a powder application, the affection cleared up within fourteen days. INSECTA. Pediculus capitis (Pediculus capitis) (Head Louse). We find Pediculus capitis in very young children and in others more growm up to be the incessant and frequent cause of impeti- ginous crust-forming eczemas. It is more frequent in girls than in ' Gruby, quoted by Lewandowsky. ^ Remak, idid. ^ Neumann, z'di'd. "* Ziirn, idtd. •' Babes, ibid. ° Lewandowsky, Deutsch. med. Wachenschr.^ ^907, xx. ■^ Hunsche, Milnch. med. JVochenschr., 1900, xlv. 7IO THE ANIMAL PARASITES OF MAN boys. In families it is endemic, in schools epidemic, but it also occurs in fair frequency in female adults (servant maids, waitresses) who may pay little attention to bodily cleanliness. The puncture of the parasites sets up a severe irritation, which leads to violent scratching. The consequences of this are the formation of nodules and pustules, crusts and "weeping" patches; the hairs become felted and the final clinical picture is that of plica polonica. The conditions of irritation which are produced by these parasites and then by the scratchings of the impetiginous, and frequently the very severe suppurative processes of the hair-bed, lead to swellings in the neck and sometimes even to glandular suppurations. The eczematous processes not infrequently extend over the face, the neck and the thorax. Blepharitis and conjunctivitis may be due to Pediculns capitis. The means of infection are often very remarkable. Transmission from one individual to another certainly often occurs, but infection may take place in railway carriages and in other ways. A case under the observation of a colleague in Frankfort is a most remarkable one : he diagnosed pediculosis as the cause of a head eczema occurring among the children of one of the best families there. The infection took place through dolls adorned with human hair, in which the presence of nits could be demonstrated. The diagnosis of Pediculns capitis is not difficult to make when the hairs and hairy scalp are carefully examined for nits and living parasites. In better families it is a good plan to point out the corpora delicti to their possessors and to make them aware of the possible sources of infection. As regards treatment, lotions of sabadill vinegar are recom- mended ; in slighter cases these are] quite sufficient. In severe cases cure will not result unless dressings of petroleum, naphthol ointment (5 to 10 per cent.) and balsam of Peru be applied. In the case of plica polonica, the hair must be cut quite short (even in adults) so as to control matting of the hair. To get rid of nits from hair that is not matted, careful combing and washing with strongly alkaline fluids or with hot vinegar is suitable. Pedlculus vestimenti (Clothes Louse). The clothes louse attacks adults by preference, and with especial frequency old and emaciated persons. It lives in the clothes, but derives its nourishment from the body. At the moment at which the clothes louse inserts its proboscis into the skin the person experi- ences a slight sting, which, however, at once ceases to hurt. If the body of the louse is sucked full of blood it falls off and the individual SUPPLEMENT 711 has rest from it for a time. A wheal develops around the haemor- rhagic area of the bitten spot and itches severely. The itching goes on until the eruption is scratched all over. This is followed by crust formation. When many parasites are present the itching reflexes become more severe, and the patients scratch themselves considerably and make long marks at those places where the Pediculi have been. The localization of the scratching effects is characteristic, correspond- ing with folds between portions of clothing (regions between the shoulder-blades, wrist and neck). If the condition lasts for a month, the scratching effects extend over the whole body, and secondary efflorescences become associated with it, such as pustules, ulcers and eczemas. Intermediate between this we find cicatrices and pigmenta- tion, the latter under certain circumstances extending over the whole body. Sulla, Herod, Cardinal Dupet, Philip II, and others are said to have died from louse disease. That even at present many human beings are exposed to the danger of being devoured by lice is a fact that we have had the opportunity of observing on several occasions. Only to record one instance, a man, aged 65, was received into our clinic some time ago in an absolutely neglected condition (he had been slaying for some weeks in a stable, lying on a wretched bed). The whole of the surface of his body was covered with countless furuncles, of greater and less size, which had partly become changed into undermined ulcers. Over the ulcers and beneath their undermined edges Pediculi were swarming. Phthirius ingulnalis {Pediculus pubis) (Crab Louse). The transmission of these parasites generally takes place during coitus, and therefore they especially occur in the pubes. It is possible also that transmission is effected through dirty clothes and bed-linen and privy seats. ^ Starting from the pubes the animals crawl out over the other parts of the body provided with hairs to the abdominal wall and the thorax (so far as these parts are furnished with thick hair) to the arm-pits, the beard, the eyebrows ; not, however, to the hair of the head, or rarely so ; among our numerous cases we have never met with an example of the crab louse attackmg the hair of the head. The irritation produced by the crab louse is extraordinarily severe, especially during the night, as the warmth of the bed incites the lice to active sucking. In consequence of the violent scratching indulged * [A case of infection through a dirty station privy in Switzerland came to my knowledge in 1899, and numbers of pediculi were found there. — F. V. T.] 712 THE ANIMAL PARASITES OF MAN in, eczemas are set up at the points attacked, and these often spread to the neighbouring parts not covered with hair. Of special interest is the onset of maculcC caeruleae (taches bleues) in some persons affected with crab Hce (people disposed to sweating seem to be peculiarly liable to these). They consist in pale blue patches of various size and shape, varying from that of a hemp-seed to that of a lentil, and again to that of a nail in size and form. These are found over the cutaneous surface of the abdomen, thorax and thigh, and are often only seen by a good lateral illumination. Duguet^ considers that the condition is a toxic erythema, that it is set up, on the occasion of the bite of the parasite penetrating the skin, by the poisonous substance derived from it. Oppenheim^ considers that it is a colouring substance that is formed in the salivary glands of the parasites, and which penetrates the skin when the insects bite, and thus forms the maculae caeruleae. We have on several occasions emulated the experiment of Duguet (trituration in a mortar of crab lice freshly taken from the human body and inoculating the mass thus obtained beneath the skin), and have similarly been enabled to produce the maculae caeruleae experimentally, but we have certainly been unable to determine which of the hypotheses is the correct one, the toxic erythema or the colouring substance inhibition theory. The diagnosis of phthirasis is very easy, for either the sexually mature parasites or the nits are found on the hairs. As regards treatment, grey ointment is regarded as a generally useful application ; it gives rise, however, to a slight eczema of the genitals, espejially in males, when injudiciously used. Geber^ recommends petroleum or balsam of Peru, Oppenheim* a i per cent, sublimate solution for lotions, or a mixture of equal parts of petroleum and benzine when the sublimate cannot be borne. The use of a 5 per cent, ointment with hydrarg. oxid. flavum is worth consider- ing in treatment of pediculosis of the eyebrows and eyelashes. The simplest method of treatment, and one with a radical effect, is that by sulphuric ether recommended by Thomer.^ It certainly produces a sharp burning sensation, but the living parasites and nits are destroyed in one sitting. We prefer ether lotions as a rule, and we thoroughly rub the affected parts with a pad of wadding well soaked with the ether. The dead parasites and the nits fall on to what lies * Duguet, Annal. de Derm., II Ser., i. - Oppenheim, "Handb. d. Hauikrankh.," v. Mracek, 1907. ^ Geber, see Seifert, Lesser's " Encyclop.," p. 387. ^ Oppenheim, loc. cit. ^ Thomer, see Seifert, Lesser's " Encyclop.," p. 387. SUPPLEMENT 713 beneath when the rubbing is done thoroughly, and the burning sen- sation caused by the ether only lasts a few minutes. Cimex (Acanthia) lectularia^ {Ciinex lectiilariiis) (Bed Bug). The puncture in the skin made by the bed bug gives rise to an extraordinary amount of severe itching and a burning sensation, and Avhen the skin is sensitive wheals of remarkable size {urticaria ex cimicibus). These eruptions that cause such severe itching are scratched by those attacked, till very soon blood begins to flow, and this generally leads to the formation of a dried crust of blood at the point of eruption. The diagnosis is not always easy, as urticaria arising in other ways frequently leads to similar vigorous scratching and formation of crusts of dried blood. Men who have some experience in this matter (for example, commercial travellers), when they are attacked by severe itching at night, are in the habit of striking a light and searching in their bed and body-linen for the bugs, in order to be able to hand over the corpora delicti to the landlord if need be. The assumption that the bugs in the East play an actual part in the propagation of tuberculosis and bubonic plague has been proved by investigations made by NuttalF to be at least very exaggerated if not wholly without foundation. Further investigations may decide how far the bugs participate in the transmission of kala-azar, as is believed by Rogers to take place. The bed bugs must be exterminated by spraying the chinks and joints in the boards with petroleum and benzine, pulling up the carpets and cleansing the bedsteads. For the treatment of the bite itself the methods recommended as an antidote against insects' stings in general are suitable : 2 per cent, carbol vaseline (Rosen bach^), thymol dissolved in spirit (i in 50^), aethrol or deci-aethrol, form- cethrol (manufactured by Dr. Nordlinger, Florsheim a. /M.), formoP (formol 15 parts, xylol 5 parts, acetone 44 parts, Canada balsam i part), with the aid of a pad of wadding placed over the part bitten, lavages with vinegar, citron juice and spirit of salmiac. 1 Vide genus Cimex, p. 534. ■^ Nuttall, see Sack " Handb.," v. Mracek, p. 290, •^ Rosenbach, Therap. Alonafsh., 1903. ^ Leipzig, med. Monatsh.^ 1907, vi. '' Chemist and Druggist, August 25, 1906. yi^ THE ANIMAL PARASITES OF MAN Pulex irritans (Human Flea). The bite of the flea produces a shght discharge of blood about the size of a pin's head, which rapidly becomes surrounded with a circular area similar to a patch of roseola. The redness fades away after a longer or shorter while (several hours), whilst the discharge of blood is to be seen for one or two days longer. In dirty people the whole body mav be covered with such discharges of blood. Individuals with very delicate, sensitive skin, especially small children, show true wheal formation at the site of the bite. In certain cases there develops from one such single bite an urticaria that extends over a large part of the body. The manner by which an irritating sub- stance is introduced into the skin upon biting by the bed bug and also by the flea is clear. The bite is followed by a feeling of itching, which is liable to rob nervous persons of their sleep. Sensitive individuals are upset even by the fleas moving over the surface of the skin during their rest at night. Treatment consists in extreme cleanliness, capture of the parasites^ sprinkling the body and bed-linen with insect powders. The fleas are difficult to remove from barracks, schools and hospitals. Dermatophilus (Sarcopsylla) penetrans (Sand Flea). The fertilized females penetrate into the skin with their heads, and here they swell, in consequence of the numerous and growing eggs and larvae, to a white ball the size of a small pea, on which the head is recognizable only as a small brown point. In this way a small brown tumour arises, over which, at the commencement, the skin is not reddened ; after some days, however, it becomes inflamed; in the centre of it a small opening is seen. If the parasite is not extracted the skin that lies over it becomes destroyed by suppuration, and thus becomes removed. At the commencement the part affected itches, with increasing inflammation ; the symptoms of irritation become more severe and may amount to actual pain. If the small suppurative processes be neglected, inflammation and gangrenous and septic processes may arise. The region of the body sought out by preference by the sand flea is the sole of the foot, the toes, under the free ends of the nails and the digito-plantoid folds — more rarely the scrotum, thigh and other parts are attacked (Scheube^). The number of parasites found on one person may amount to several hundreds. Scheube, "Die Krankh. d. warmer Lander," 1896. SUPPLEMENT 715 Treatment consists in the removal of the parasites from the skin with a needle or a small sharp knife and the application of a bandage. Rubbing the feet with copaiba or Peru balsam, sprinkling them with insect powder, or washing them with bay rum (Berger^) acts as a prophylactic or removes the irritation of the skin produced by the parasites. Myiasis. Under the name of myiasis we designate the complex symptoms which parasitic dipterous larvae give rise to in man (Braun), and we conceive under the term myiasis externa (dermatosa s. cutanea) all lesions of the human integument caused by fly larvae and of the cavities covered with mucosa therewith connected, such as the external auditory meatus, the oro-nasal cavity, the urethra and vagina. The occurrence of dipterous larvae in the digestive tract is named myiasis intestinalia or interna. Myiasis externa. The larvae of a species of fly belonging to the Muscidce, Liicilia inacellaria,^ are found in relative frequency in the nose, especially in America and India.^ Riley* has stated that the screw-worm of Central America and of the United States is nothing else than the larva of Liicilia inacellaria, and also that the Brazilian fly named '' berna " may be no other than Lncilia macellaria. Their offspring may set up inflammatory disturbances in the soft tissues of man. This fly has a wide distribution, from the Argentine Republic to Canada, also in the British portions of the East Indies, where the disease is named " peenash." This word is derived from the Sanskrit, and is said to be a collective name for all diseases of the nose. Lahory^ states that w^ithin a period of nine years ninety-one cases of " peenash '^ occurred in Allyghar, two of these ending fatally. Liicilia macellaria is not at all timid but bold, like the house-flies and blue-bottles, its relatives. It not only lives at no great distance from human dwellings, and forces its way into villas and country houses, but even attacks its victims without awaking them from their sleep. Although this species shows a certain preference for nasal cavities affected with catarrh or pus (v. Frantzius^), and also the external auditory meatus, ' Berger, Therap. Monalsh., April, 1907. - {Chrysomyia macellaria, p. 587. — F. V. T.] ^ [C. macellaria, Fabricius, the screw- worm fly, is found in tropical America and the West Indies. The genus is restricted to America. The species from India is a Pycnosoma. — F. V. T.] ^ Riley, American Naturalist, 1883, xvii. •^ Lahory, Edin. Med. lourn., 1856. 6 V. Frantzius, Virchow's Archiv, 1868, xliii. 45 7l6 THE ANIMAL PARASITES OF MAN as well as ulcerated or wounded parts of the body, and even badly ulcerated skin carcinoma (Lutz^), it is not a rare thing for it to penetrate into one of the above-mentioned cavities rapidly to deposit its eggs, without these parts having been previously affected. The report also of Conil,*^ in which these flics bear the name of Calliphora anthropophaga,^ is an interesting one. Probably it was the same species of Muscid in the cases of myiasis nasi observed by von Tengemann, Delasiauve,* Weber,^ Mankiewicz,^ and Kirschmann.^ In the case recorded by Prima,^ and in that recorded by Britton,^ the issue was a fatal one ; in the latter the larvae escaped through the pharynx and nose ; the hyoid bone and the soft parts of the palate were destroyed, the speech and power of swallowing 'were hindered. At the post-mortetn extensive destruction of the internal nose was found, so that the nasal bones could only be kept in their position by the aid of the external skin. Even during life 227 larvae escaped. Similar destructive processes were found in the case communicated by Richardson. ^^ In two cases reported by Schmidt^^ 300 and 350 larvae were respectively removed from the nose, and the patients recovered. Wolinz^^ found his patient had lost consciousness, and that in the pus filling up the entrances to the nose numerous larvae were moving; recovery followed. In the case communicated by Adler,^^ more than 150 larvae escaped from the nose of an old man. Curran^* states that people suffering from ^' peenash " frequently die from meningitis. The cases reported by Pierre'^ related to the forms of severe myiasis frequently to be observed in Guiana. In a patient who was suffering from typhus ( ? typhoid), Douglas^^ found the conjunctival sacs full of larvae ; in two other individuals the nasal cavities were attacked. The case observed by Summa^^ was that of a man, aged 28, who suffered from nasal obstruction, foetor, epistaxis and pain in the nose. Out of seven of the cases occurring at Fort Clark, U.S.A., and in its ' Lu*z, see Joseph, Deutsch. med. Zeitg., 1885. * Conil, Annal. de Science nat. zooL, 1878. • [This fly belongs to the genus Cordylobia, and is peculiar to Africa, C. anthropophaga^ or the tumbri fly, is, when a larva, a subcutaneous parasite of man and animals. — F. V. T.] ^ Delasiauve, Gerhardt's "Handb. d. Kinderkrankh.," 1878, iii. 2 Weber, Mexique Rec. d. Mhn. de Med. milit.^ 1867. ^ Mankiewicz, Virchow's Archtv, 1868, xliv. ' Kirschmann, Wien. med. Wochenschr., 1881. ^ Prima, ** These de Paris," 1881. ^ Britton, Cambridge, Massachusetts, 1883. '" Richardson, Medical Monthly, 1883. " Schmidt, Texas Med. Journ., 1887. '•- Wolinz, Wratsch, 1884. i3 Adler, Med. Record, 1885. '* Curran, Med. Press and Circ, 1887. '=* Pierre, "These de Paris," 1888. '^ Douglas, Kansas City Med. Index, 1890. " Summa, St. Louis, 1889. SUPPLEMENT 717 neighbourhood, six ended fatally ; in all these cases KimbalP dia- gnosed ozaena ; attracted by the strong odour the flies forced their way into the noses of the patients when asleep and there deposited their ova. In a case reported by Carriere^ an abscess of the nasal septum was produced by the larvae of flies ; Chiodi^ reports seven cases of myiasis due to Lucilia macellaria ; among these was a case of rhinitis myiatica, in which a cerebral abscess leading to a fatal termination developed, being produced by the migration of a larva into the brain. Among the three cases of Lesbini^ was that of a girl, aged 16, with 250 larvae in the diseased nasal cavity. Quintano^ observed larvae beneath the eyelids in one case. It is possible that the cases of Cesare^ and Calamida'' w^ere those of myiasis nasi due to Lucilia macellaria. The larvae are also found in the nasal accessory sinuses, as is seen from the cases reported by De Saulle^ (frontal sinus), Delasiauve^ (frontal sinus), MacGregor^^ (antrum of Highmore), and Bordenave^^ (antrum of Highmore). If a survey is made of the literature of the cases described of myiasis nasi produced by Lucilia macellaria^^ the following information is forthcoming : In Europe this form of the disease is of very rare occurrence, whilst in America and India^^ it is frequent. Persons suffering from ozaena are rendered the most liable to danger as the penetrating odour entices the flies in tropical countries with intense frequency, so much so that v. Frantzius does not consider this myiasis as an independent disease, but as a complication of ozaena of frequent occurrence in warm countries. The infection is so far of interest in its nature, in that it only takes place during the day. The fly is on the wing only by day when the sun is shining, and consequently only deposits its eggs at this time. Therefore persons suffering from ozaena are principally exposed to the danger of being pursued by the flies when they succumb to sleep during the mid-day hours in the open or in dwellings that are not closed up. Headache is the symptom which most troubles the patients. It extends over the whole cranium and persists uninterruptedly, with ' Kimball, New York Med. Journ., 1893. 2 Carriere, Gaz. hebd. de Med. et de Chir., 1898, xciv. 3 Chiodi, La Argent. Med., March i, 1905. ^ Lesbini, ibid. •^ Quintano, *' Cronic oftalm. de Cadiz," 1878. 6 Cesare, Arch. ital. di Otol., April, 1903. ■^ Calamida, Giorn. d. A\ Accad. de Med. di Torino, September, 1903. « De Saulh, Gaz. des Hdp., Paris, 1857. » Delasiauve, Gaz. hebd. de Med., Paris, 1885. ^0 MacGregor, Arch. gin. de Med., No. 1,031. " Bordenave, ** Deuxieme Mem. presente a I'Acad. de Chir.," v, p. 387. " [And the other species, of course, must be included here. — F. V. T.] '^ [Concerning Europe and India, macellaria does not occur. — F. V. T.] 7l8 THE ANIMAL PARASITES OF MAN more or less severe periods. Violent headaches in the frontal and buccal regions are almost always present in this complaint; they are experienced either only on one side or on both simultaneously ; sometimes the pain is extended to the lower jaw and region of the neck, following the whole extent of the trigeminal nerve. The inflammation of the nasal mucosa produced by the penetration into it of the larvae extends right into the frontal sinus and antrum. Simultaneously the patients, at the height of their trouble, suffer from persistent sleeplessness and severe vertigo, so that they reel and cannot walk straight ; excessive sneezing always sets in at the com- mencement. The larvae immediately spread over the nasal mucosa to seek a place suitable to feed, and irritate the nasal mucous membrane by the tickling sensation they produce. Later the patients frequently sneeze when the maggots move to and fro. One very characteristic symptom consists in the peculiar swelling of the face, which is extended either over the whole or only one half of it, and may alternate with attacks of erysipelas (Brokaw^). The discharge from the nose is of special diagnostic value. It consists of a blood-stained serous matter or blood-stained fluid, which is perpetually trickling from one or both nostrils. The larvae especially choose the anterior portions of the nasal cavity, where they can be seen lying in groups together at the base of the choanae. The consequence of this is that the soft palate becomes intensely swollen, and this in turn makes swallowang very difficult ; speech is impeded, and the voice acquires a nasal intonation. Symptoms of fever become more or less pronounced according to the number of larvae present, and according to the nature and constitution of the individual. The appetite is in abeyance throughout the whole dura- tion of the illness, and sometimes there is the onset of slight attacks of diarrhoea. If the larvae are not removed in good time there follows excessive destruction of the interior of the nose and of the turbinals ; and the whole nasal framework undergoes disintegration, frequently, too, the velum palati, so that the larvae come into sight in the oral cavity. Individuals thus severely attacked succumb through exhaustion, symptoms of meningitis (cerebral abscess) or septicaemia (Prima"). Twenty-one out of thirty-eight cases recorded (collected) by Maillard^ died. The method of prophylaxis is self-evident from what has been stated. On bright summer days neither the healthy nor those suffering ' Brokaw, see Seifert, in Heymann's " Handb.," p. 595. 2 Prima, "These de Paris," 1881. ^ Maillard, " These de Montpellier," 1870. SUPPLEMENT 719 from diseases of the nose should sleep during the day-time in the open or in public habitations : sufferers from nasal diseases should pay special attention to this. Treatment consists in the removal of the larvae ; this, however, is not always easy. With regard to the methods which have proved to be effectual in the destruction of living larvae and their expulsion from the nose, strongly smelling and easily diluted fluids come first, such as alcohol, eau-de-Cologne, and ether, which should kill the creatures when injected into the nostrils. The earlier physicians, such as Salzmann,^ Honold,^ and Henkel,^ have seen good results from the use of these methods, whilst Mankiewicz'' and Goldstein^ obtained no results whatever. Kimball's^ careful investigations have shown that a decoc- tion of bitter herbs recommended by Behrends'' (tansy, wormwood) have just as little effect as the tobacco decoction employed by Boerhave^ and Kilgour.-* The sternutatories employed by the older physicians are entirely neglected. Delasiauve'^ experienced good results from the inhalation of the smoke of paper cigarettes, which were soaked with a solution of 2-0 pot. arsenic in 30*0 distilled water. Whilst, according to Kimball, balsam of Peru had no effect on the larvae, Mankiewicz succeeded in removing the larvae from the nose with the help of that drug. Turpentine steam or mixtures of turpentine employed by Indian physicians have not been very effectual according to Moore,^^ Kimball and Goldstein. Success has been attained in some cases by the use of insufflations of calomel (Roura,^^ Cerna,^^ Schmidt^"*) or of iodoform (PascaP^). Joseph^^ recommends con- centrated alum solution being sniffed up into the nose as very effectual. Sublimate and carbol solutions do not appear to be very successful (Kimball, Moore, Goldstein), whilst benzine inhalations (Pierre^^) have shown better results. Scheppegrell^^ strongly recommends injections of oil which kifl the larvae, while it is perfectly harmless to the nasal mucosa. Cesare^^ employed nasal lavages with solutions of salicylate of soda with good results, and Calamida^^ lavages with physiological saline solution. Bresgen^^ recommends the nose being ' Salzmann, see Tiedemann, Mannheim, 1844. "^ Honold, ibid. 3 Henkel, ibid. ^ Mankiewicz, Virchov/s Archiv, 1868, xliv. * Goldstein, New York Med. yourn.y 1892. 6 Kimball, ibid., 1893. ' Behrends, see Tiedemann. ^ Boerhave, ibid. ^ Kilgour, ibid. ^^ Ddtasiauve, loc. cit. '1 Moore, Chicago Med. Times, 1893. ^^ Roura, Gaz. di San. inilit., 1884. •3 Cerna, New York Med. J own., 1893. '^ Schmidt, Texas Courier, 1884. '5 Pascal, Arch. d. Med. milit., 1895. '^ Joseph, Deutsch. med. Zeitg., 1885. '7 Pierre, "These de Paris," 1888. '» Scheppegrell, New York Med. Journ., 1898. '^ Cesare, loc. cit. ^o Calamida, loc. cit. 2' Bresgen, Eulenburg's " Real. Encyclopadie," third edition. 720 THE ANIMAL PARASITES OF MAN cocainized and the larvae being removed with a pincette. Roorda- Smit^ cocainized the nose, then insufflated calomel and plugged the nose with a gauze tampon dusted with calomel. After two hours fifty-six larvae crawled out along the plug. Continuation of the treatment resulted in a complete cure. Injections of chloroform water (Jourdran^) or chloroform inhala- tions, or injections of pure chloroform into the nose, have proved the most effectual (Goldstein,^ Osborn,^ Jourdran, Durham,^ Jennings,^ Kimball/ Mackenzie,^ Oatmann/^ Zarniko,^^ Antony,^' Folkes^^). Camphorated carbolic solutions are very well spoken of : Grayson^^ states that these kill the larvae immediately. Some authors have removed the larvae with forceps (Goldstein^'^), others with pincettes ; thus Brokaw extracted 200 fragments with the forceps, Pascal eighty fragments with the pincettes, and Wolinz^^ also appears to have removed the larvae with forceps. Greater operative measures than these do not appear to have been undertaken in latter days ; yet Morgagni^*^ states that the army surgeon, Caesar Mogatus, at Bologna, first trephined the frontal sinus and then extracted a '' worm " from it. Larvae of other Muscidce have come under observation much more rarely (ChevaP' [larvae of Galleria melloiiella^^], Bond,^^ DumesniP^ [larvae of PiopJiila casei]). Species of the genus Scolopendra {Myriapoda), which all shun the light and seek their food during the night — which consists of animal and vegetable substances — frequently make their way into the nasal cavities of people when asleep. They are found not only in the nose, but in the accessory cavities. In the chapter on the '* Parasites of the Nose "^^ we have collected striking ' Roorda-Smit, Deutsch. vied. Wochenschr., ico6. ^ Jourdran, Aich. de Med. nav., 1895. ^ Goldstein, Neiv York Med. Journ., 1892. ^ Osborn, Daniel's Mid. Journ.^ 1891. 5 Durham, Chicago Med. Times, 1893. ^ Jennings, Kansas City Med. Index, 1890. "^ Kimball, New York Med. Journ. ^ 1893. ^ Mackenzie, " Diseases of the Nose and Throat." ^ Oatmann, Med. Mirror, February, 1894. ^° Zarniko, " Lehrb, d. Krankh. d. Nase." ' Antony, Bull, Soc. vied, des Hop. de Paris, 1903. '2 Folkes, New York Med. Record, 1907. '3 Grayson, St. Louis Med. and Surg, yonrn., 1891. 1* Goldstein, N'ew York Med. Journ., 1892. '^ Wolinz, Wralsch, 1884. '^ Morgagni, see Tiedemnnn. '^ Cheval, Journ. de Med. et de Chir., 1893, '^ [This is the larva of a moth. — F. V. T.] '* Bond, ////. Zeniralbl. f. Laryng., 1896. 2° Dumesnil, see Friedreich, " Die Krankh. d. Nase," 1858. ^' Seifert, j^^ Heymann's "Handb." SUPPLEMENT 72 I instances, but we have omitted to mention the observation made by Bertrand^ (Scolopendra in sinus maxillaris) and that made by Berg- mann^ (Scolopendra in sinus frontahs). In the same chapter some remarks are made as to the occurrence in the nose of earwigs, cater- pillars, scorpions and termites, as well as of animals which have not been identified. The larvae that develop in the auditory meatus penetrate the membrana tympani, destroy the middle ear and may produce menin- gitis and intracranial suppurations. In one case Vesescu^ extracted seven living larvae from the ear with the aid of a thin pair of pincettes. Kohler^ recommends the infusion of drops of ol. terebinth, to destroy the larvae, Ouintano^ the insufflation of the following powder : Oxid, hydrarg. rubr., sulfur., aa I'o grm., pulv. gi. arab. 8*0 grm. ; Lesbini^ recommends tincture of iodine. In the case reported by Henneberg'' the larvae were those of Lucilia ca'sar. Eye affections due to Lucilia iiiacellaria are very uncommon ; the literature relating to the lesions of the eye produced by the larvae of flies has been collected in Kayser's^ work. In the cases under the observation of Schultz-Zeyden'** both the eyes of a female tramp were destroyed, and quantities of larvae were also found in the nasal fossae and in the ears. The Lucilia is found relatively seldom on the cutaneous surface. Henneberg's^° case was that of a neglected girl, aged 20, in whom countless larvae (L. ccesar) were found in a plica polonica ; after the plica polonica had been removed the scalp was found to be covered with a large quantity of ulcers which swarmed with larvae, large and small. The skin of the trunk was also much macerated and covered w^ith larvae. Death resulted from sepsis ; Westenhoffer" remarks on this case that a lesion of the head from which the patient had suffered previously and the perpetual state of intoxication in which she was had probably given rise to the lodgment of the fly larvae. Whether the communications made by Munk^^ Qf maggots in the mouth relate to Lucilia I do not know. Vesescu,^^ in one case with extensive ' Bertrand, Soc. med. de Bologne, 1839. 2 Bergmann, Koi-respondenzbl. d. deiilsch. Ges. f. Psych., Neuwied, 1 859. * Vesescu, Riv. sliintelor vied., February, 1906. * Kohler, Monatsschr. f. Ohretiheilk., 1885. * Quintano, see Seifert, loc. cit. ^ Lesbini, La Argent. Med., 1905. ■^ Henneberg, Berl. med. Ges,, February 18, 1903. ^ Kayser, Kliu. Monatsbl. f. Augenheilk., 1905. ^ Schultz-Zeyden, ^^r/. i/z'«. Wochenschr., 1906. ^^ Henneberg, Berl. med. Ges., February 18, 1903. ^' Westenhoffer, Verein f. innere Med., Berlin, May 7, 1906. ^2 Munk, Wien. med. Presse, xxi. *^ Vesescu, loc. cit. 72 2 THE ANIMAL PARASITES OF MAN ulceration and deep fistulae in the skin, removed 176 larv^e with the pincette. In Roorda-Smit's^ case there were two ulcers in the neck of a girl, aged 17, and larvae appeared at their base. After dusting with calomel and the appUcation of a bandage the next day fifty-two dead or half-dead larvae came to light. Recovery took place. Lesbini/ in the case of an old lady, saw numerous larvae in an ulcer of the leg she was suffering from. Hector's^ case appears to have been one of myiasis cutanea provoked by Lucilia. The first exact observations of myiasis cutanea from SarcopJiaga magnifica are due to Wohlfahrt,* in whose honour Portschinsky'"' named this species of fly S. wohlfahrti. Portschinsky ascertained that S. ivohlfahrii was not confined to man as its sole host, but that several of our domestic animals, such as cattle, horses, pigs, dogs and geese, w^ere visited. In these animals small wounds serve to entice the flies and to supply them with a suitable site for the deposition of their eggs. The oral armature of the young larvae renders it easy for them to penetrate not only the mucosa and cutaneous surface but also intact places in the submucous connective tissue. In many localities more than half the herds have proved to be infected by the flies. The fly only frequents open spaces and never enters human dwellings, and is so timid that it approaches man only during sleep ; infection, therefore, takes place only out of doors, in summer, in clear, warm weather, and only in such individuals as sleep in the open air. Individuals are most exposed to risk who suffer from catarrhs or inflammations, combined with purulent secretions of the nasal cavity (ozaena), or otorrhcea, or ulcers in any parts of the body accessible to the female fly. The frequency and intensity of the infection will be in inverse proportion to the advance in civilization of the inhabitants, their idea of cleanliness, their having timely medical aid and the chances of their being rapidly attended to. On that account the majority of cases of myiasis (Sarcophaga) are reported from Russia. The literature of this kind of myiasis nasalis is not very extensive ; in addition to Wohlfahrt, Portschinsky and Joseph,* there is a communication by Gerstacker,^ who found fifteen adult larvae of 5. wohlfahrti in the nasal cavity of one man. The larvae transmitted from Ordruf by Dr. Thomas to Low,^ in Vienna, which were discharged from the nose of a woman, ' Roorda-Smit, Deutsche med. Wockenschr.^ 1906. ^ Lesbini, loc. cit. ' Hector, Lancet^ 1902. * Wohlfahrt, " De vermibus per nares excretis," Norimbergae, 1770. * Portschinsky, "Norae Soc. entomolog. Rossicae," 1875. * Joseph, Deutsch. vied. Zeitg., 1885. ' Gerstacker, "Sitzungsberichte d. Ges. f. naturf. Freunde in Berlin," 1875. " Low, Wien. med. Wochenschr,, 1883, xxxi. SUPPLEMENT 723 aged 71, suffering from czaena, were recognized by the well-known dipterologist Braun as belonging to S. wohlfahrti. Among the cases reported by Joseph, one only affected the nose; it was that of a peasant girl, aged 11, who had suffered from ozaena; she had travelled on the open road and had there gone to sleep. Severe symptoms set in and death followed under delirium. In making the post-mortem it was found that the interior of the nose was extensively destroyed by larvae of S. wohlfahrti. Powell found Sarcophaga larvae in two persons who had slept in the open air ; the larvae were killed by injections of chloroform and sublimate. Destruction of the eyes by S. wohlfahrti has only been observed in a few cases ; it is reported by Cloquet^ that, in the case of a ragman who had lain some time in the fields, both eyes were pierced by larvae. On the outer skin the larvae of S. wohlfahrti have been found more than once in inflammatory or festering areas. Freund^ demonstrated that from a live year old child, which had suffered for some time from an impetiginous eczema of the skin of the head, from two suppurating abscess cavities which extended to the periosteum, which was already affected, twenty-one living larvae were taken ; rapid healing took place under antiseptic bandaging. The small treatise by Balzer and Schimpff^ contains two new observations on myiasis externa ; in the one case an ulcer on a man's foot was full of larvae, in the other case thehead of a woman showed numerous larvae without the skin of the head being destroyed. Brandt's'* observation is interesting, for he found such larvae in the gums of a sick person. The impression which one obtains of the active movement of larvae on wounds is a strange and at the same time uncanny one. One finds that the larvae to obtain protection against the drying of the surface of the abscess almost incessantly burrow with their heads, first contracting and then expanding the body, which rises and falls, and keeping the tail upwards. Owing to these movements producing irritation, increase of inflammation may ultimately arise, causing erysipelas and cellulitis. The treatment of myiasis nasalis caused by Sarcophaga is the same as in myiasis caused by Lucilia, and in the other places where found it is merely a question of the removal of the larvae and the subsequent proper treatment of the surface of the abscess. In Northern Nigeria * Cloquet, see Schultz-Zehden, loc. cit. "^ Freund, Ges. f. innere Med. in Wien, December 5, 1901 ; and Wien. med. Wochenschr . ^ 1910, li. 3 Balzer and Schimpff, Annal. de Derm, et de Syph.y 1902. < Brandt, Wratsch, 1888. 724 THE ANIMAL PARASITES OF MAN Lelean^ found Anchmeromyia depressa to be the cause of myiasis externa.^ The occurrence of Oestiid larv^ in a human being is very rare, at least up till now myiasis oestrosa has been very seldom observed in man in Europe. Whilst the hosts of the Muscidct comprise a considerable number of warm-blooded animals, on which the larvae develop, each species of the Oestridce appears, on the other hand, to have a definite host or some definite hosts of the class Mammalia. No species of Oestrid is peculiar to man. Although in America, as well as in Europe, Oestrus lioniiuis was spoken of up to the middle of the last century, no such species exists. But in both hemispheres, in America much more often than in Europe, Oestrid larvae have been found in man. In Florida, Mexico, New Granada, Argentina, Brazil, Costa Rica and other districts, and especially where large herds of cattle are kept, myiasis oestrosa has been observed in shepherds, huntsmen and amongst the rural population. The larvae of Hypodenna hovls, according to the obser- vations of Goudot,^ occur as a parasite in man. Poilroux^ found larvae of cavicolous Oestridce in the nose of a man, aged 55. Amongst the species of warble flies, whose larvae are parasites in domestic animals and game in Europe, reliable observers have found larvae of two kinds, Hypodenna bovis and Hypoderma diana, also in man.^ The larvae of H. bovis have very seldom been observed in the nose. The . case quoted by Kirschmann,*^ which was that of a peasant woman, aged 50, who was suffering from ozaena, and in which violent attacks of sneezing, epistaxis, pain in the forehead, and swelling of the face were observed, is, according to Low^ and Joseph,^ not an Oestrid ; Muscid larvae were evidently the cause. By the injection of diluted iron chloride solution seventy-nine larvae were removed from the nose. In the case reported by Kazoux^ the species of larva is not definitely known — at least, v. Frantzius^^ did not consider them Oestrid larvae. Joseph does not definitely say that Oestrid larvae were the cause of a case which he quotes. He was sent a number of uninjured larvae of Oestrus ovis ready to pupate, which ' Lelean, Bril. Med. Journ.^ 1904. 2 [Numerous instances of attacks by Auchmeromyia are known and referred to under that genus, pp. 593-4' The species referred to here is not depressa. Walker. — F. V. T.] •'' Goudot, Annul, d. Set. itat., 1845. ^ Poilroux, Joiirn de Mid., Chir., etc., 1809. ^ \Hypodertna linearis is frequently confused with H. bovis. — F. V. T.] 6 Kirschmann, Wien. med. Wochenschr., 1881. ■^ Low, Wien, med. Wochenschr., 1882. ^ Joseph, Deiitsch. vied, Zeitg.^ 1885. ^ Razoux, Journ. de Mid, Chir., etc., 1 758. *° V. Frantzius, Virchow's Archiv, 1868, xliii. SUPPLEMENT 725 were said to have been expelled, during violent sneezing, from the nose of a peasant woman who had suffered for six months from continuous frontal headache and chronic nasal catarrh. The Oestrides prefer to use the surfaces of wounds on the skin of man to lay their eggs, which develop into larvae ; but they often use their ovipositors^ to make a fresh wound. In this case there arise in the skin, and particularly in the subcutaneous connective tissue of the neck, in the region of the shoulder, as wxU as in other parts of the body painful, furuncle-like inflammations which are known under the name of gad-fly boils. These boils may become the size of pigeons' eggs ; if several are together, they appear to form a connected tumour. Each tumour is elastic and somewhat movable, and has an orifice through which the larva breathes and discharges its excreta. At times these turn to festers and gangrenous disintegrations, which may even cause the loss of a limb. Wilms^ had the opportunity a few years ago of observing a case of myiasis dermatosa oestrosa in Leipzig. The fistula which led to the larva was slit open and the larva extracted. As a notable characteristic of myiasis oestrosa Joseph states that the larva3 grow very slowly. The flight time of the Oestridcv is the hot summer months. Adams^ observed on the Isthmus of Panama a number of cases of a skin disease which is caused by the larvae of Dermatobia noxialis {Gtisano-peliido-Muche). The larvae penetrate not only the skin but also the mucous membrane of the pharynx and larynx, and from there proceed through the tissue to the subcutaneous cellular tissue. The infection seems to result from bathing. The study of ^' thimni," a human myiasis caused by Oestrus ovis, by Ed. and Et. Sergent,^ deals more with the zoology and with the geographical distribution of this insect in North Africa than with the clinical appearances of myiasis. [This paper deals with matters of great interest, with important facts. — F. V. T.] The treatment consists in the removal of the larvae (from the nose) ; in Brazil it is the custom to drop tobacco juice into the boil in order to kill the larvae (Strauch^). One is only justified in speaking of myiasis intestinalis when there is no doubt that living fly maggots or flies themselves can be proved to have been found in the fresh contents of the stomach or intestine ^ [The O. strides appear to lay their ova on the hair of animals. They do not puncture the skin.-F. V. T.] - Wilms, Deutsch. med. Wochensclu:, 1897. 8 Adams, [our7t. Atner. Med. Asscc, 1904. ^ Ed. and Et. Sergent, Annal. de VInst. Pasteur, 1907. ^ Strauch,y^//;v/. of Cut. Dis., 1906. 726 THE ANIMAL PARASITES OF MAN (Schlesinger and Weichselbaum^). In the discussion of myiasis intestinalis we give the evidence of Schlesinger and Weichselbaum, as well as that of Wirsing,^ to which must be added a number of other investigations. In a great number of acute cases apparently only the stomach was affected, there being no signs in the intestine. In these cases sudden illness is noticed, colic, sometimes unbearable pains in the region of the stomach, pyrosis, vomiting or continuous intense inclination to vomit, occasionally even with the mixture of blood. Frequently a general feeling of malaise, twinges of pain in the muscles, and attacks of giddiness were notified, very rarely fever. Generally all the symptoms disappeared in a short lime when the larvae had been removed by an act of vomiting or by washing out the stomach. It is well to note that in the history of many cases the pains preceding the expulsion of the larvae are stated to be extremely violent. Acute myiasis of the intestinal canal frequently runs a course without special symptoms and is only an accidental condition ; one . has, however, in such cases to guard against errors. The faeces may be deposited in vessels or places where fly larvae are in great numbers, or a subsequent infection of the faeces with the eggs or larvae of flies may have taken place. Only when the inspection of the excrement immediately following defaecation proves the presence of living larvae, and when there were certainly no fly larvae in the vessel previously, can one speak of the passing of fly larvae from the intestine. More frequent than the cases showing no special symptoms are those with pronounced disturbances in the intestinal passage, obstruction or diarrhoea (also constipation and diarrhoea alternately), violent and sometimes agonizing abdominal pains (Pottiez^), which preceded the evacuation of the larvae and subsided after their removal. General symptoms, like weakness, languor, transitory vague pains, loss of appetite, sickness, rarely fever, giddiness, attacks of faintness, epileptic attacks (Krause*) are observed. In a few cases blood and pus have been noticed in the evacuation of the bowels. In the cases of chronic myiasis of the intestine the aspect of the disease is dominated by the complex symptom of colitis mucosa. The following features are noticeable, namely, the intermittent passing of blood, the influence over the expulsion of the larvae of mechanical procedure (massaging of the abdomen), the duration of the process for several years, the sometimes enormous number of 1 Schlesinger and Weichselbaum, Wien. klin. Wochenschr., 1902, i. 2 Wirsing, Zeituhr.f. klin. Med., 1906, Ix. 3 Pottiez, Bull, de V Acad, royale de Med. de Belgique, xv. ^ Krause, Detiisch. med. Wochenschr., 1886, xvii. SUPPLEMENT 727 insects contained in the dejecta. Another cHnically important factor is the passing of the larvae in batches. While for some time no larvcC may appear in the stools, they may suddenly be ejected in great numbers, either because the conditions of feeding are not suitable, or because medicaments remove them from the intestine. The haemorrhage is ascribed by Schlesinger and Weichselbaum directly to lesions of the mucous membrane caused by the larvae ; in the case reported by these writers there were found shreds of tissue as well as pus in the stool. The pains occurring spontaneously in the abdomen are at times influenced by position and attitude of the body, often they were more violent after rest and after evacuation of the bowels ; often they were continuous, but in that case less intense ; pressure on the abdomen is generally little felt. The condition of the blood was in two cases (Pasquale^ and Schlesinger and Weichselbaum) a marked chlorotic one. The state of nutrition seems almost always to suffer with prolongation of the disease, but in Peiper's^ cases this was not so. The condition of the appetite was in some instances good, in others very bad. A frequent symptom is headache of a migraine-like character and neuralgic pains in different parts. Schlesinger and Weichselbaum's case shows that there are forms of myiasis intestinalis which, after prolonged sickness, lead to death, and that in consequence of the formation of intestinal abscesses stricture of the intestine may arise from the subsequent formation of a scar. The question of the mode of infection is interesting ; in this mouth, nose and anus must be considered. The most frequent way is certainly by means of food on which flies have laid their eggs, or which is permeated with young maggots. This may be raw (especially grated) meat, cheese, fruit, salad, milk, cabbage, cold farinaceous foods, raspberries. When the stomach is affected, when the gastric juice has lost acidity and power of digestion, the larvae will be able to stay and develop more easily. According to Csokor,^ if the eggs get into the gastro-intestinal canal of man with the food, the delicate stages of the young larvae would certainly not surviv^e the action of the gastric juice. Salzmann^ assumed that the invasion occasionally occurred through the rectum, the larvae creeping into the anus while the person is asleep. Wirsing accepts this method of infection for two of his cases, where it was a question of the infection of an infant. Salzmann* reports a case where the maggots of Anthomyia^ scalaris 1 Pasquale, Centralbl.f. Bakt., 1891. 2 Peiper, " Fliegenlarv. als gelegentl. Parasiten d. Menschen," Berlin, 1900. 3 Csokor, Wieti. klin. Wochenschr., 1901, p. 129. 4 Salzmann, Wiirttemberg. med. Korrespondenzbl. 1883, liii. 3 [This is presumably Homalomyia [FaTinia) scalaris, — F. V. T.] 728 THE ANIMAL PARASITES OF MAN were passed in great numbers from the urethra of an old man. The patient had been catheterized on account of urethral stricture and was probably infected with eggs or larvae at the same time. The diagnosis of the affection is easy and sure, if Hving larva' are found in the contents of the stomach or in the stools, and if contami- nation is out of the question. The number of different species of flies whose larva3 are found in myiasis intestinalis is considerable. The larvae of species of Anthomyia (.4. canicularis,^ A. scalaris, etc.), of Sarcophaga carnaria and 6\ niagnifica and of Mnsca vomitoria^ are especially observed. The prognosis is certainly generally favourable, but must be made with some reserve in chronic cases, in view of the observations of Schlesinger and Weichselbaum (intestinal stenosis). The treatment must aim at removing the larvae as soon as possible from the digestive canal. In cases of myiasis of the stomach, a thorough washing out of the stomach (Joseph,^ Staniek*) is to be preferred to emetics used with success in individual instances ; perhaps it would be advisable to add menthol or thymol to the mixture. In myiasis of the intestine internal remedies and local treatment of the intestine must be considered. So far santonin seems to have proved to be the best remedy. In some cases extract, filicis maris, calomel, semina cucurbitae, naphthalene o'l to 0*5 (Peiper^), infus. of Persian insect powder (5 in 200), mineral waters, Carlsbad water, seem to have had good results. P'or irrigation of the rectum, weak solutions of argentum nitricum, tannin, thymol, gelatine, ol. ricini, naphthalene may be used. Wirsing administered an aperient (Rurella compound liquorice powder) and a soap enema after the passing of the first larvae. The principal thing is the prophylaxis, which must include the careful protection of articles of food, on which flies may lay their eggs (protection by glass dishes, tulle or fine wire nets). Fruit should not be eaten before being washed or rubbed with a cloth. ^ [This fly, common in houses, is known as Homalomyia canicularis, and the next belongs to the same genus. — F. V. T.] 2 [This fly belongs to the genus Calliphora, not Musca.—Y. V. T.] 5 Joseph, Deiitsch. vied. Zei(g., 1885 and 1887. ^ Staniek, see Schlesinger and Weichselbaum, p. 47. « Peiper, " Fliegenlarv. als gelegentl. Parasiten d. Menschen," Berlin, 1900. SUPPLEMENT 729 Gastricolous Oestridae (Creeping Disease). Syn. : Creeping eruptio7i; Larva migrans ; H autmaulwurf ; Dermatomyiasis linearis migrans oestrosa ; Hy ponomoderma ; Dermatitis linearis migrans ; Linea migrans; Epidermiditis linearis migrans Wolossatik j Kriechkrankheit ; Haut- kratzschorf ; Myiase hypodermique. Under the name ^' creeping disease," R. J. Lee^ has recorded a peculiar affection of the skin in a three year old girl, which appeared first in the form of pale red, thread-like irregular protuberances, which seemed partly to become entwined on the right malleolus and had spread without causing special disturbances to the abdomen. Dickinson, Fox and Duckworth^ reported, in connection with this, that they observed a growth of this red line of about i in. per diem. Since then a number of similar cases have been reported which, without doubt, were cases of larvae creeping under the skin. Crocker^ saw such a case in a two year old girl, the progress of the red line varying in one night between 4 and 7J in. In Europe the first case was observed in Vienna, by v. Neumann and Rille,"^ also in a two year old girl. V. Samson-Himmelstjerna,^ Sokoloff,*^ Rawnitzky' found larvae at the end of the tract, which had been recorded as larvae of Gastro- philus by Cholodowsky.^ According to Blanchard {Arch. f. Par., 190 1 ) the larvae were those of Hypoderma hovis. How these larvae get into the skin has not yet been definitely ascertained ; v. Samson is of the opinion that they usually obtain access to man as larvae, Stelwagon^ believes that the infection generally occurs in a seaside watering place; a patient of EhrmannV^ fell ill when he returned from the manoeuvres, where he had lain for some time on the ground. Here and there it is reported that the eruption was preceded for a longer or shorter time by lesions of the skin (incised wounds, furuncles, slight excoriations, V. Harlingen"). Twice it has been suggested that perhaps the parasites might come from vineyard snails (Crocker, Lenglet and Delaunay'^), and it is » R. J. Lee,/<7Mm. Clin. Soc: Lond., November 27, 1874. 2 Dickinson, Fox and Duckworth, ibid.^ 1875. 3 Crocker, " Diseases of the Skin," 1893 ; ** Atlas of the Diseases of the Skin." * V. Neumann and Rille, Wten. klin. Wochenschr. , 1895 ; Dermatologenkongr., Graz, 1895. * V. Samson-Himmelstjerna, Wratsch, 1895; Arch.f. Derm. u. Syph., i897- « Sokoloff, Wratsch, 1896. '^ Rawnitzky, Derm. Zeitschr., v, p. 704. « Cholodowsky, Wratsch, 1896. ' Stelwagon, Journ. Cut. Dis., xxii, 8. '» Ehrmann, Wien. derm. Ges., November 17, 1897. " V. Harlingen, Amer. Journ. of Med. Set., 1902. 12 Lenglet and Delaunay, Annal. de Derm, et de Syph., 1904. 730 THE ANIMAL PARASITEvS OF MAN pointed out by v. Samson that in Russia the infection of peasants who work in the fields was specially frequent. It is noticeable how frequently the affection begins on uncovered parts of the body (face, hands, arms) ; but that fact, on the whole, is not in conflict with the statement (Kengsep') that the disease makes its first appearance over the nates, because children often sit on the ground and play with that part of their body uncovered. A case observed by us was that of an elderly lady who did not do this and was properly clothed, yet showed the typical lines of creeping disease on the nates, and asserted again and again that she had the feeling as if a worm were creeping under her skin. The disease occurs in children as well as adults, so that age, sex and calling offer no determining point etiologically. The clinical symptoms of the disease consist in the sudden appearance of itching and burning ; if the cause is looked for one perceives a red line, raised but little above th^^ surface of the skin, with irregular curves, never branched, but often entwined, broadening more or less rapidly at one end (i to 15 cm. in twenty-four hours). The larva can be seen sometimes with a lens under pressure of the skin as a dark spot ; formations of pus, such as other larvae produce, are not noticed ; now and again there is a formation of little vesicles {Hamburger,^ v. Harlingen,^ Bruno,^ Ehrmann,^ Brodier and Fouquet,*^ Rawnitzky''). It may happen that the parasite burrows through a small region of the skin with many close curves for some time; on the other hand, observations exist where it covered large tracts in a short time. The itching and smarting cease in the place left by the larva, so that the patients even in the shortest tract can point out at which end the larva is, even if they have not watched the lengthening of the tract. Very rarely the larva invades the mucous membrane of the mouth, the nose, and the conjunctiva, proceeding from thence to the external cutaneous area. The localization of the affection is very varied ; the primary seat has been observed on the glutei muscles (Lee, Kengsep, Morris,^ Rille, Seifert) and their surroundings (Stelwagon, Hamburger, Bruno), on the lower extremities (Stelwagon, Lenglet and Delaunay, Hutchins, Moorhead, Lee, Crocker, Schmid,^ v. Harlingen), on the ' Kengsep, Derm. Centralbl.y 1906, vii. 2 Hamburger, y<7«/-«. of Cut. Dis.^ 1904. '^ V. Harlingen, loc. cit. ^ Bruno, v. Rille and Riecke, " Handb. d. Hautkrankh. v. Mracek." ^ Ehrmann, loc. cit. ^ Brodier and Fouquet, Bull, de la Soc.frauf. d. Derm., 1904. ' Rawnitzky, loc. cit. ^ Morris, Brit. /onrn. Derm., 1896. " Schmid, Verein der Aerzte in Steiermark, February 12, 1900. SUPPLEMENT 731 upper extremities (Samson, Meade and Freeman, Hutchins, Sokoloff, V. Harlingen, Brodier and Fouquet, Shelmire,^ Stelwagon), on the face (Sokoloff, Moorhead, Kumberg,^ Rawnitzky, Crocker, Boas^), on the neck (Sokoloff), and on the body (Ehrmann, Brodier and Fouquet, Kaposi,^ Topsent*''). The duration of the affection varies very much ; it varies between a few hours and some years*^ ; several times a spontaneous recovery has been reported. The diagnosis of the disease is not at all difficult owing to its peculiar appearance. The treatment can only consist in the removal or killing of the larvae, since one cannot rely on spontaneous recovery, even if it has occurred in some cases. If one should succeed in locating the larva as a black spot at the end of the tract, its removal by means of a needle is the simplest method (Quortrup and Boas^). In some instances a cure has been successfully accomplished by excision of the active end of the tract (v. Neumann and Rille, Schmid). In opposition to this method, which not all patients will allow, the method practised by Arab women (Rille and Riecke^) of killing the worm with red hot needles is quite rational. Shelmire^ used the electrolytic needle for the destruction of the maggots, Stelwagon^^ made use of cataphoresis, by means of which he applied a sub- limate solution, afterwards cauterizing with a drop of nitric acid, as excision was refused. Crocker" and v. Harlingen^^ injected small quantities of carbolic acid; Moorhead^^ by a single freezing of the skin with ethyl chloride, attained a definite cessation of the attack at the active end. Hutchins^^ in one case made use of hypo- dermic injection of a few drops of solution of cocaine and afterwards of I to 2 drops of chloroform ; in a second case of repeated applica- tions of tincture of iodide, as Lenglet and Delaunay^^ did. ' Shelm'ne, /ourn. Cut. Dis., 1905. 2 Kumberg, St. Peter sb. med. Wochenschr., 1898. ' Boas, Monatsh.f. prakt. Derm., 1907, xliv. ^ Kaposi, Wien. klin. Wochenschr.^ 1898. ^ Topsent, Arch, de Par., 1901. 6 [This is extremely unlikely, as the bots of Hypoderma only live for nine or ten months at the most !— F. V. T.] '^ Quortrup and Boas, Hospitalstid., 1907. ** Rille and Riecke, " Handb. d. Hautkrankh.," v. Mracek, 1907, iv. ^ Shelmire, loc. cit. " Stelwagon, loc. cit. 1^ Crocker, loc. cit. " V. Harlingen, loc. cit. '^ Moorhead, Texas Med. News, 1906. " Hutchins, y<7«rw. Cut. Dis., 1906. ^•^ Lenglet and Delaunay, loc. cit. 46 732 THE ANIMAL PARASITES OP^ MAN V. Harlingen' allayed the affection in his first case by rubbing in sapo viridis and tar, in Kensep's^ case the cure seems to have been accomplished by an ointment containing resorcin, in Meade and Freeman's'^ case by a 20 per cent, ichthyol paste. In our case we made exclusive use of Lassar's paste ; within four weeks a cure resulted, probably spontaneously, since one cannot ascribe any essential effect to this paste. * V. Harlingen, loc. cit. 2 Kensep, loc. cit. ^ Meade and Freeman, Brit. Jourjt. Deriu.y October, 1906. APPENDIX ON PROTOZOOLOGY 733 APPENDIX ON PROTOZOOLOGY, Comprising Notes on Recent Researches, Formulae of some Culture Media, and Brief Notes on General Protozoological Technique. BY H. B. FANTHAM, M.A., D.Sc. I.— NOTES ON RECENT RESEARCHES. Since the foregoing section on Protozoology was sent to press, certain interesting observations and results have been published. Brief notes on such, and some references thereto, are now added. It is necessary, however, to remark that sometimes it is impossible to give a precise or rigid definition to a genus of Protozoa, owing to differences of opinion, to differences regarding nomenclature or to incompleteness of knowledge. Such a lack of definition, while incon- venient for the time being, is not unhopeful, as it directs attention to the necessity for further work, which is inevitable in such a relatively new and wide subject as protozoology. Thus, it may be noted in illustration that Minchin, in 1912, in his text-book regarding the genus Efitaniceha writes : '^ The entozoic amoebae are commonly placed in a distinct genus, Entamoeha, distinguished from the free- living forms by little, however, except their habitat and the general (but not invariable) absence of a contractile vacuole." DiflPerences between Entamoeba histolytica and E. coli. — In continuation of the remarks on pp. 34 and 40, it may be added that Lugol's solution (iodine in aqueous potassium iodide solution) in fresh specimens shows by brownish staining the presence of glycogen in the vacuoles of Entaniceba coli. Such a reaction is rarely or never given by E. histolytica. Phagedaenic Amoebae. — Carini and others record cases in which the skin around an operation wound in connection with liver abscess became gangrenous. Amoebae, possibly Entamoeba histolytica, were found therein and may have been responsible for the gangreno- phagedaenic action. Endamoeba ginglvalis (see pp. 43, 44). — Smith and Barrett,^ after * Joui'TU of Parasitol., i, p. 159. 734 I^HE ANIMAL PARASITES OF MAN analysing the early literature, state (June, 191 5) that Endamceba gingivalis, Gros, 1849, is the correct name for the following organisms : E. buccalisy Prowazek, 1904 (see p. 43) ; Amoeba gingivalis, Gros, 1849; Aniceba buccalis, Steinberg, 1862, and Amoeba dentalis, Grassi, 1879. They conclude that E. gingivalis is the causal agent of pyorrhoea alveolaris, and that this disease responds to treatment with emetine. Entamoeba kartulisi (see p. 44), synonym E. maxillaris, Kartulis, is considered to be E. gingivalis. Smith and Barrett adopt the generic name Endamoeba, Leidy, 1879 {see footnote on p. 31, also p. 34). Leidy worked on Endamoeba blatta^. Cralgia and Cralgiasis {see p. 45). — Barlow^ (May, 191 5) found Craigia {Paramoeba) hominis in cases of chronic diarrhoea and mild dysentery in Honduras. He also described a new species of Craigia under the name of C. migrans. Fifty-six cases were studied, five of which were due to Craigia hominis, the remainder to C. migrans. In C. migrans, each flagellate, on attaining full development, becomes an amoeba without dividing. Each amoeba encysts and produces a number of flagellates which are somewhat like cercomonads. On the other hand, in C. hominis the flagellate form produces, by longi- tudinal fission, several generations of flagellates before entering upon the amoebic stage. The cysts of C. migrans contain fewer ^' swarmers " (flagellul^) than those of C. hominis, but the ''swarmers" are some- what larger, namely, 5 jjl instead of 3 /^ in diameter. Further, there is no accessory nuclear body in C. migrans, but its flagellum stains more deeply than that of C. hominis and has a peculiar banded iippearance. Human Trichomoniasis {see pp. 52-56).— Lynch^ (April and May, 1915), working in Charleston, seems to favour the view that the trichomonads found in the vagina, urethra, mouth, lungs and alimentary tract are one and the same organism, and that these flagellates may further excite already existing inflammatory condi- tions. He gives detailed histories of cases of {a) infection of the vagina and gums, and (6) intestinal infection manifested as intermit- tent attacks of diarrhoea. The flagellates were found in catarrhal vaginal discharge, in blood-stained scrapings from the gums (together with Endamceba buccalis), and in stools after a purge of magnesium sulphate. The parasites were tetratrichomonads {see footnote, p. 53), that is, each possessed four flagella anteriorly as well as an undulating membrane. Lynch successfully infected rabbits from the cases and from cultures of the parasite. Encysted trichomonads were seen in a Amer. Journ. Trop. Dis. and Prevent. Aled.^ ii, p. 68o. /hid., p. 627 ; New York Med. y^^^rw.. May I, 1915, ci, p. APPENDIX ON PROTOZOOLOGY 735 patient's stools, in rabbits infected therefrom and in cultures. The culture medium used was bouillon acidified with about 0*05 per cent, acetic acid and the cultures were maintained at 30° C. Trichomonads occur in the digestive tracts, for example, the caeca of rats and mice (fig. 422). In man allied flagellates can occur in similar situations, as well as in other parts of the intestine. Fig. 422. — Trichomonas from csecum and gut of rat: «, nucleus; bl, blepharoplast ; y?, flagella ; ax^ axostyle ; ///, undulating membrane ; b, line of attachment of undulating membrane to the body. X 2,000 approx. (Original.) Other trichomonad-like organisms have been recently described from the faeces of man, more particularly from cases of chronic dysentery in the tropics. Derrieu and Raynaud^ (July» 1914)^ working in Algeria, found a flagellate possessing five free flagella anteriorly and an undulating membrane apparently lateral. They named the parasite Hexamastix ardin-delteiliy but the generic name Hexaniastix is pre-occupied. Chatterjee^ (January, 1915), working in India, found probably the same flagellate and called it Pentatrichomonas bengalensis. Chllomastix (Tetramitus) mesnili (see p. 57). — Alexeieff^ (1914) now places the parasite originally called Macrostoma mesnili, by Wenyon (1910), in the genus Chilomastix, Alexeieff. The differential characters of the genera Tetramitus and Chilomastix are not especially well marked. According to Alexeieff, Tetramitus is characterized by four unequal flagella (which he figures anteriorly), a ventral cytostome in the form of a linear cleft and a pulsatile vacuole in front of the anterior nucleus. Chilomastix, according to the same author, has Bull. Soc. Path. Exot., vii, p. 571. Ind. Med. Gaz., 1, p. 5. Zool. Anzeiger, xliv, pp. 203, 206 ; and ibid.^ xxxix, p. 678. 736 THE ANIMAL PARASITES OF MAN three forwardly directed flagella and a fourth backwardly directed one in the cytostome, which is well developed (fig. 423). Some authors consider that the fourth flagellum forms the edge of an undulating membrane in the cytostome. Diagrams of Chilomastix niesnili are given in fig. 423. Fig. 423. — Chilomastix ( 1 etramitus) mesnili. a, b, c, flagellate forms ; dy rounded or encysted form. X 2,500. (Original.) Giardla (Lamblia) intestinalis {see p. 57). — Alexeiefif^ (1914) con- siders that Lamblia intestinalis, h^Lmbl, should be placed in the genus Giardla, Kunstler, 1882. Bipartition occurs in the encysted state. The axostyles persist in the quadrinucleate cyst. Cercomonas hominis {see p. 61). — This parasite is considered by some authors to be of a doubtful nature, as it is thought to have been mistaken for deformed or incompletely observed Trichomonas or Chilo- mastix or even Lamblia. Wenyon^ (1910) described Cercomonas longicauda from cultures of human faeces. It is considered that the genus is very confused, and the author points out that the tail flagellum has been overlooked. He considers that the genus Cercomonas should include flagellates with an anterior blunt end from which arises a single long flagellum, and a posterior tapering end also with a flagellum, which can be traced ' ZooL Anzeiger, xliv, p. 210. - Quart. Journ. Micros. Sci., Iv, p. 241. APPENDIX ON PROTOZOOLOGY 737 over the surface of the body towards the insertion of the anterior flagellum. Another species, Ccrcojiionas parva, has been found in cultures of human faeces by Hartmann and Chagas^ (1910). It has a somewhat different structure. Furthur researches are necessary on the organisms variously referred to the genus Cerconwnas. Transmlssive Phase of Trypanosomes in Vertebrates. — In addition to the general remarks on the morphology of trypanosomes set forth on pp. 70 to 72, it may be noted that Woodcock^ (November, 1 9 14) states that, in certain cases, there is a definite transmissive phase of a trypanosome in its vertebrate host. He quotes the work of Minchin and himself on T. noctiice of the little owl, in which the transmissive form is spindle-shaped and occurs in the bird's peripheral blood during the early summer months {see p. 69). A similar phase occurs in T. Jringlllarum, and Robertson^ has found that the short, stumpy form of T. gauibiense is its transmissive phase in vertebrates. Trypanosoma lewisi {see p. 88). — Brown (1914-15) has published some interesting results on the potential pathogenicity of T. lewisi, Blepharoplastless Trypanosomes {see p. loi). — Laveran^ (April, 19 1 5) suggested a practical use of strains of blepharoplastless trypano- somes produced by the action of drugs. He finds that tryposafrol will also produce such strains, and remarks on blepharoplastless strains of T. evansi and T. brucei, which in the former case can undergo 450 passages without reversion, and in the latter 273 passages. He states that if it is desired to inoculate surra or nagana to Capridae or Bovidae in order to produce immunity, use should be made of the blepharoplastless races of the respective trypanosomes, which races are a little less virulent than the corresponding normal ones. Also, the immunity which follows from an infection due to blepharoplastless T. evansi or T. brucei is only a little less coifiplete than that following infections from either of the respective normal strains. The Experimental Introduction of certain Insect Flagellates into various Vertebrates, and Its bearing on the Evolution of Leishmaniasis. — In continuation of the remarks on pp. 103, 104, and 112, further researches have been conducted on the introduction into vertebrates of flagellates normally parasitic in insects. The vertebrates became infected by inoculation with the flagellates or by ' A/em. Inst. Oswalao Cruz, ii, p. 67. ^Arch.f. Protistenk.^ xxxv, p. 197. ^ Proc. Roy. Soc, B, Ixxxv, p. 527. 4 C. R. Acad. Set., clx, p. 543. 738 THE ANIMAL PARASITES OF MAN being fed on insects containing the protozoa. Fantham and Porter^ (June, 1915) published the following results. Flagellates from sanguivorous and non-sanguivorous insects were used, and cold- blooded as well as w-arm-blooded vertebrates as hosts. The intro- duced protozoa were pathogenic to the mammals, but not markedly so to the cold-blooded vertebrates. Herpetomonas jaaihiiu, H. stratio- myice^ H. pediaili, and Crithidia gerridis (parasitic in certain water- bugs) proved pathogenic to mice. A puppy was infected by way of the digestive tract with H. ctenocephali. Frogs became infected with H. jaciilum and with C. gerridis^ toads and grass snakes w'lih. H. jacnlum, lizards with C. gerridis, and sticklebacks with //. jaadiim. Second and third passages of some of the parasites were obtained. The protozoa, whether Herpetofuonas or Crithidia, were present in the vertebrate hosts in either the non-flagellate or the flagellate form, or usually both. They were more abundant in the internal organs of the hosts, more particularly in the liver, spleen and bone-marrow'. In all experiments in which C. gerridis was used the parasite invariably retained the crithidial facies in the vertebrate host. No transition to a trypanosome was ever seen. Infections in adult animals were not so heavy as in the young ones, and the parasites were more virulent in young hosts, as is the case with Mediterranean kala-azar in children. The mode of infection of the vertebrate in Nature seems to be contaminative, either by its food or through an already existing abrasion or puncture on the surface of its body. Cases in which the flagellate-infected insects have been allowed to suck the blood of vertebrates have proved negative up to the present. In areas where leishmaniases are endemic, an examination should be made of all insects and other invertebrates likely to come into contact with men or dogs, or rats and mice (see below), in order to ascertain if these invertebrates harbour herpetomonads. Preventive measures should be directed against such invertebrates, especially arthropods. Further, it is likely that certain vertebrates, such as reptiles and amphibia (especially those that are insectivorous), may serve as reservoirs of leishmaniases, or, as they should preferably be termed, herpeto- moniases. From such reservoirs the herpetomonads may reach man by the agency of ectoparasites or flies, especially such as are sanguivorous. That vertebrates in Nature can harbour herpetomonads in their blood has been shown by the work of Button and Todd (1903) on the herpetomonads of Gambian mice, while the recently published ' Proc. Camb, Philosoph. Soc, xviii, p. 137 ; and ^nna/s Trop. Med. and ParasitoL, ix, P- 335- APPENDIX ON PROTOZOOLOGY 739 investigations of Fantham and Porter^ (June, 1915) on natural herpe- tomonads in the blood of mice in England have shown that these rodents may be a natural reservoir of herpetomoniasis. The origin of the infection of mice is to be sought in a flagellate of an ecto- parasite of the mouse, very probably Herpetomonas pattoni parasitic in various fleas, which protozoon can adapt itself to life in the blood of mice. Herpetomonads were also found naturally in the blood of birds by Sergent (1907). Recently, Fantham and Porter have successfully infected birds with herpetomonads experimentally. The significance of insect flagellates in relation to the evolution of disease has recently been set forth by Fantham^ (June, 1915). The deductions to be made from the occurrence of a herpetomonad stage in Leishmania, especially in L. tropica, in man himself, and of flagel- late stages of the so-called Histoplasma capsiilainm in man are fully discussed and correlated. It is pointed out that flagellosis of plants (see p. 104) may possibly be connected with leishmaniasis. The evolution of Lcishmanla from flagellates of invertebrates is thus traced and the way again indicated for preventive measures against leishmaniasis, as first set forth by Dodds Price and Rogers. Franchini and Mantovani (March, 191 5) have successfully infected rats and mice by inoculation or by feeding with Herpeto- monas muscat domestical obtained from flies and from cultures. It is of great interest to note that the recent observations of Ed. and Et. Sergent, Lemaire and Senevet^ (19 14) have demonstrated the presence of a herpetomonad flagellate in cultures of the blood and organs of geckos obtained from areas in Algeria in which Oriental sore, due to L. tropica, is prevalent. Phlebotomiis flies, which may harbour a natural herpetomonad, feed on the geckos and on men. Hence animals like geckos may possibly act as reservoirs of leishmaniasis. Lindsay'^ (1914) writes that the parasite of dermo- mucosal leishmaniasis in Paraguay is believed by native sufferers to be conserved in rattlesnakes, and spread by ticks or flies {Simutium) feeding on the reptiles and transferring the parasite to man. The Transmission of Spirochaeta duttoni (see p. 116). — It is probable that Ornitlwdorus savignyi acts as the transmitting agent of S. c/////o;// in places like Somaliland (Drake-Brockman, 1915).'' Spirochaeta bronchialis {see p. 122). — The morphology and life- history of S. broncliialis have been investigated by Fantham^ (July, 1915). From researches conducted in the Anglo-Egyptian Sudan, he ' Parasitology, viii, p, 128. - Annals Trop. Med. and FarasiloL, ix, p. 335. =* Bull. Soc. Path. Exot., vii, p. 577. 4 Trans. Soc. Trop. Med. and Hyg., vii, p. 259. ^ Ibid., viii, p. 201. ^ Annals Trop. Med. and Parasitol., ix, p. 391. 740 THE ANIMAL PARASITES OF MAN found that S. bronchialis is an organism presenting marked poly- morphism, a feature that has only been determined by the examination of numerous preparations from the deeper bronchial regions of various patients. S. bronchialis varies in length from 5 ft to 27 fi, and its breadth is about 0-2/A to o-6yLt. These variations are due to the processes of growth and division. Many of the parasites measure either 14 yu- to 16 />t long, or 7 /A to 9yLt, the latter resulting from transverse division of the former. The ends show much variation in form, but approach the acuminate type on the whole. The discrepancies in dimensions given by the very few previous workers on the subject are probably the result of the measurement of a limited number of parasites. All such sizes can be found on some occasion during the progress of the disease, when a larger number of spirochsetes is examined. The movements of S. bronchialis are active, but of relatively short duration, when it is removed from the body. The number of coils of the spirochaete is rather an index of its rapidity of motion than a fixed characteristic of the species. The motile phase of S. bronchialis is succeeded by one of granule formation, the granules or coccoid bodies serving as a resting stage from which new spirochaetes are produced. The formation of coccoid bodies and reproduction of spirochaetes from them can be observed in life. S. bronchialis is a species distinct from the spirochaetes occurring in the mouth. It differs from them in morphology, pathogenicity and in staining reactions. It is not a developmental form of any bacterium, and is an entity in itself. The passage from man to man is effected most probably by means of spirochaetes, and especially coccoid bodies, that leave the body in the spray with expired air and byway of the nasal secretions. Owing to the fragility and short life of S. bronchialis extracorporeally, the resistant coccoid bodies in air, in dried sputum and dust, and possibly also on the bodies of flies and other insects, are probably instrumental in inducing attacks of bronchial spirochaetosis in human beings, espe- cially those having a lowered bodily resistance, such as occurs after a chill. Lurie (December, 1915), has described a case from Serbia. The Spirochaetes of the Human Mouth {see p. 122). — Two species of spirochaetes were recorded as occurring in the human mouth about forty or fifty years ago. These are Spirochccta buccalis, Steinberg (often ascribed to Cohn, 1875), and S. dentinni, Miller (often attributed to Koch, 1877). The most recent work on S. dentinm and S. bnccalis is that of Fantham^ (July, 1915), who observed the parasites ascribed to Cohn * Annals 'Irop. Med. and ParasitoL, ix, p. 402. APPENDIX OX PROTOZOOLOGY 74I and to Koch, these being the two common spirochcetes seen in the mouths of natives of the Sudan and of Europeans in England, as well as the forms described and cultivated by recent investigators. Some of the mouth spirochaetes are not very active, but there is marked corkscrew and boring movement, and they are flexible. Tangles or tomenta of these mouth spirochetes are common. Internal structure is seen with some difficulty, but in some specimens it can be determined, and chromatin granules are then seen. Muhlens (1907) figured stained specimens of S. hiiccalis and S. dentiiim, in which chromatin-coloured granules were distributed along the bodies of the organisms. S. deiitiiim has tapering ends, and varies in length from 4 //. to lOyLt. S. dentitun is rather like Treponema pallidum, and has been placed by some workers— for example, Dobell— in the genus Treponema. It has already been mentioned, on p. 128, that Noguchi cultivated three species of Treponema from the human mouth — namely, T. macro- denthun, T. microdentium, and T. mucosum, but they cannot be easily distinguished morphologically, and so may appear to be biological varieties of S. dentium. S. huccalls has somewhat rounded or bluntly acuminate ends and varies in length from g fi to 22 /jl. A slight membrane or crest may sometimes be observed. S. huccalls was found to be the pre- dominant spirochaste in the mouths of eight natives examined by Fantham in the Anglo-Egyptian Sudan. S. huccalls and S. dentium take up stains well and with relative ease. Intracellular stages of the parasites are uncommon. Multiplica- tion by binary fission has also been observed. Coccoid bodies or granule stages of the mouth spirochaetes are formed, but appear to be relatively few in number. J. G. and D. Thomson^ {1914) have written an interesting paper on various spirochaetes occurring in the alimentary tract of man and of some of the lower animals. They have also given a useful list of references, and the work of some of the earlier authors is discussed in the paper. With regard to the general morphology of spirochaetes, it may be noted that the so-called axial fibre of Zuelzer is acknowledged to be homologous with the membrane or crista of molluscan spirochaetes. Coccidia in Cattle. — Regarding the remarks on coccidiosis or ** red dysentery " in cattle on p. 147, it may be added that Schultz^ (July, 1915) has found the malady among cattle in the Philippine Islands. He states that some irregular or atypical cases of apparent rinderpest are really due to coccidia. As has been pointed out by Proc. Roy. Soc. Med.^ vii, pt. i, p. 47. " Jotirn. Infect. Dis., xvii, p. 95. 742 THE ANIMAL PARASITES OF MAN Montgomery, rinderpest can be transmitted by blood inoculation, while coccidiosis cannot be so transmitted, but may be diagnosed by the microscope. These differences should be remembered as the two diseases are often found to be associated and are difHcult to separate clinically. Coccidia have also been found in Australian cattle. The Haemosporidia. — It is likely that this order {see p. 151) may be soon abolished. MesniP (April, 19 15) considers that the grouping of the three families, Plasmodiidae (or Haemamcebidae), Haemogreg- arinidae and Piroplasmidae in the order Haemosporidia is no longer possible, because of the coccidian nature of the Haemogregarines {see p. 154). The Coccidia are divisible into the Adeleidea and the Eimeridea (see p. 141). The Haemogregarinidae are allied to the former, and the Plasmodiidae to the latter. The Piroplasmidae, until more is known of their life-cycle in the invertebrate host, cannot be more definitely placed. The Leucocytozoa of Birds. — Regarding the statement, on p. 153, that Laveran and FYanca consider that avian leucocytozoa may inhabit red blood cells, it may be added that Franga''^ (April, 1915) remarks that the action of the parasites on the red cells is very rapid and very intense. The host cells become so altered that it is difficult to -recognize their true nature. He used very young birds in his researches. Two shapes of host cell are considered, namely, those with fusiform prolongations, and those which are rounded and with- out such prolongations (see p. 153). The movements and form of the Leucocytozoa determine the shape of the host cell, as was pointed out by Fantham^ in 1910. Schizogony of these parasites has been seen by Franga (191 5) and by Coles (1914), in addition to Fantham (1910), and to Moldovan (1913), mentioned on p. 153. Schizogony may also take place in the lungs of the host. The genus Leucocyiozodn, established by Ziemann in 1898, belongs to the family Haemamoebidae. II.— FORMUL/E OF SOME CULTURE MEDIA. (i) Culture Media for growing Amoebae.— There has been much discussion as to whether the true parasitic Entamcebcv or Endamcehce can be grown on culture media [see p. 42). Undoubtedly certain free-living amoebae can be so grown, and it is considered that some of the earlier researches on the so-called artificial growth of the dysenteric amoebae were really due to contaminations with free-living forms. The following media are worthy of note : — ' Bid I. Soc. Path. Exot., viii, p. 241. - Ibid., p. 229. 3 Pj.^^^ 2:001. Soc. Lond., 1910, p. 694 APPENDIX ON PROTOZOOLOGY 743 Musgrave and Clegg in 1904 devised a culture medium for amoebae. The organisms grown by them were probably not dysen- teric amoebae, as was thought, but free-living forms. Phillips^ (1915) •gives a slightly modified formula of Musgrave and Clegg's medium, thus : — Agar-agar ... ... ... ... 2*5 grm. Sodium chloride ... ... ... ... 0*05 ,, Liebig's beef extract ... ... ... 0*05 ,, Normal sodium hydroxide ... ... ... 2*0 c.c. Distilled water ... ... ... ... 100*0 ,, Without clarifying, sterilize at 7 kilograms pressure per square centi- metre for about three-quarters of an hour. It should be neutral to phenolphthalein. Anna W. Williams" (191 1) described a medium consisting of fresh tissue spread on agar plates for the culture of amoebae. There are three stages in the procedure : (i) obtaining living amoebae free from other living organisms ; (2) obtaining sterile tissue ; and (3) making successive transplants of amoebae and tissue, and showing that every transplant is free from other living organisms. Each step requires many controls. The essentials of the method may now be given. Remove aseptically and rapidly the tissue required, such as brain, liver, kidney, or spleen, from a freshly killed animal (guinea-pig, rabbit, or dog). Put each tissue on a separate agar plate. Cut the selected tissue into tiny pieces, and spread them over freshly made agar plates. Place these plates in a thermostat at 36° C. for tw^enty- four hours to insure sterility. Add the broken up tissue to the amoebae, free from bacteria, and maintain the cultures in thermostats, some at 36° C, and some at 20° C. to 24° C. Emulsions of liver and brain in sterile neutral glycerine may also be used. The freshly removed tissue serves as food for the amoebae. The cultural amoebae mentioned on p. 42 were grown on such media or modifications thereof. One modified medium actually used was brain tissue, to which blood was added from day to day, and an easily assimilable bacterium (one of the influenza group of bacilli) was present, which did not overgrow the medium at a temperature of 38° C. Different conditions of food and of temperature produced morphological variations in the cultural amoebae. Couret and ]. Walker^ (1913) state that they have cultivated five varieties of intestinal amoebae, the associated bacteria having been previously separated. They used a medium consisting of agar to ' *' Amoebiasis and the Dysenteries," p. 8. - Journ. Med. Research, xxv, p. 263 ; and Proc. Soc. Exper. Biol, and Med., viii, p. 56. 3 Tourn. Exper. Med., xviii, p. 252. 744 THE ANIMAL PARASITES OF MAN which sterile autolysed tissue had been added. The sterile tissue, such as brain or liver, was kept in a sterile thermostat at a temperature of 40° C. for ten to twenty days. The surface of the agar should be broken up before use, and the medium must not be too acid (not over I '5 per cent.). They consider that autolysed tissue is necessary for the growth of Entamoebae, and that naturally associated bacteria aid growth by autolysing the tissues. (2) Culture Media for the growth of Protozoa parasitic in the Blood. — MacNeal and Novy,^ in 1903, used a mixture of blood and agar for the cultivation of trypanosomes such as T. lewisi and T. brucei. They employed varying proportions of the blood and agar, a medium consisting of two parts of deiibrinated rabbit's blood mixed with one part of agar being useful. The trypanosomes grew in the water of condensation. Some of the authors' earlier formulae contained different proportions of blood and agar w^ith a little peptone, while one of these media contained meat extract, agar, peptone, salt and sodium carbonate. The temperature, like the proportion of blood and agar, varied with the trypanosome investigated, but the optimum was 25° C. Mathis^ (1906) somewhat simplified the technique of Novy and MacNeal. He collected the blood of a suitable animal, such as rabbit, cow or dog, strict asepsis not being essential. The blood was defibrinated in the ordinary way. One part of blood was added to two parts of agar at 50° C. The mixture was sterilized several times by heating to 75° C. or 100° C. Slopes were made and the water of condensation was inoculated with a little blood containing the trypanosomes. Blood may be obtained from a superficial vein or from the heart. Novy-MacNeal-Nicolle or N. N-. N. Medium. — In 1908 C. Nicolle^ brought forward a modification of the Novy-MacNeal (N.N.) medium. The formula is as follows : — Agar ... ... ;.. ... ... I4grni. Sea salt ... ... ... ... ... 6 ,, Water ... ... ... ... ... 900 ,, Apparently pure sodium chloride can be substituted equally well for sea salt. The mixture is placed in tubes and sterilized in an autoclave. To each tube one-third of its volume of rabbit blood, taken by aseptic puncture of the heart, is added. The salt agar is kept liquid at 45° C. to 50° C. and the blood is added to the mixture. The culture medium so prepared is maintained for five days at 37° C, and then for a few days at room temperature. ' See Sleeping Sickness Bulletin (1909), i, No. 8, p. 287. 2 C. R. Soc. Biol, Ixi, p. 550. 3 c^ ^, ^^^^ ^^i^ (,jj1^^ p_ g^^. APPENDIX ON PROTOZOOLOGY 745 This medium was devised for the cultivation of Leishmania {see p. 106), but trypanosomes may also be grown thereon. Subsequently, NicoUe recommended the use of citrated rat's blood heated to 45° C. for half an hour, instead of defibrinated rabbit's blood. On such a medium, J. G. Thomson and Sinton^ (19 12) succeeded in growing Trypanosoma ganihiense and T. rhodesieiise (see pp. 76, 83). Noguchi's media for the cultivation of Spirochaetes and Trepo- nemata are described on pp. 123, 125. Hata's modification is discussed on p. 126. Bass's glucose-blood medium for the cultivation of malarial parasites is described on pp. 170-172. It has also been used successfully for the cultivation of Piroplasnia or Babesia {see p. 172). III.— BRIEF NOTES ON GENERAL PROTOZOOLOGICAL TECHNIQUE. The object of this book is to give accounts of the structure and life-histories of the numerous parasitic organisms that affect man more particularly. It is, therefore, inappropriate to devote much space to a consideration of technique, regarding which many volumes have already been written. Methods of procedure are largely matters of opinion, and the technique that gives brilliant results when used by one investigator may be a complete failure in the hands of another. In the present appendix, brief notes regarding certain relatively simple methods only can be given, because the number of fixatives in use is very great ; there are also large numbers of stains as well as many modifications of them, while the methods of apply- ing both fixatives and stains are, perhaps, still more numerous. There are so many, in fact, that confusion frequently arises from the multiplicity of choice presented to the worker. Those desiring more information on the subject of technique are advised to consult the treatises of Bolles Lee^ and of Langeron.-^ Fresh Material. {a) Simple Examination. Fluid Substances, such as Blood and Sputum. — A small quantity of the substance to be examined is taken on a sterile platinum loop and transferred to a perfectly clean glass slide. A clean cover-slip is gently lowered on to the drop, air bubbles being avoided. The preparation ^ Annals Trop. Med. and ParasitoL, vi, p. 331. * " The Microtomist's Vade Mecum " (7th edition, 1913). London : J. and A. Churchill. ^ '* Precis de Microscopic" (1913). Paris : Masson et Cie. 746 . THE ANIMAL PARASITES OF MAN is luted with vaseline or paraffin and examined first with a low power and then with a high power objective. The light is cut down by partly closing the diaphragm of the substage of the microscope. Skin Ulcers and Similar Sores. — Scrapings are made from the edge of the sore, mixed with sterile physiological salt solution, and prepared and examined as above. Fceces. — A small portion of faeces, or flakes of mucus (which may be blood-stained) from the same, is removed on a sterile platinum loop, spread out thinly after dilution, if necessary, with physiological salt solution on a slide, covered and examined as before. Alternatively, hanging drop preparations of blood, ulcerative tissue, or faeces, appropriately diluted if necessary with sodium citrate or physiological salt solution, may be made on a cover-slip, which is inverted over a slide with a well in it. The cover-slip is then luted and examined. For the elucidation of the developmental processes of such organisms as trypanosomes, spirochaetes and piroplasms, fresh preparations may be often kept under observation longer by the use of a thermostat, maintained at or near blood heat, in which the microscope is inserted. (b) Intra vitaui Staining of fresh Preparations. intra vitani staining is of service on some occasions, more particularly for the study of the nucleus and other chromatoid substances of the living organism. Two methods are in common use. In the first case, the stain, employed usually in very dilute solution, is mixed wdth the medium containing the organism. The latter takes up some of the stain, the amount of coloration depending on the organism concerned and on the stain employed. The commoner intra vitam stains are pure, medicinal (zinc-free) methylene blue and neutral red, used in aqueous solutions. A solution of methylene blue of i per 1,000 of water may be tried, while neutral red in the proportion of i per 3,000 parts of water has proved of service. The second method of vital colouring consists in placing a drop of I per cent, solution of methylene blue on a slide or cover-slip, slightly spreading it, and allowing it to dry. The living organism is then placed in a drop of saline on the prepared slide or cover-slip, which is then mounted and examined under the microscope. Progressive staining of the organism occurs and its internal structure can be seen. A similar procedure may be followed for neutral red. Intra vitam staining is useful for relatively large and easily deformed protozoa such as ciliates, as well as for amoebae and flagellata of the gut. When examining very actively motile organisms, it is sometimes APPENDIX ON PROTOZOOLOGY 747 useful to endeavour to restrict their movements by adding a little gum or gelatine to the medium. (c) Examination by aid of the Paraboloid Condenser, The use of one of the dark-ground illuminators (so-called ultra- microscopes) is of service for the detection of minute living organisms or of organisms present in small numbers only. The forms of paraboloid condenser manufactured by the firms of Zeiss and Leitz can be recommended. For details of their methods of employment, reference should be made to the leaflets of the firms supplying the said instruments. By the use of the paraboloid condenser, the finer details of certain stages of life-cycles, such as the formation of granules in spirochaetes and treponemata, can be observed more readily than by using the ordinary substage of the microscope. The use of the paraboloid condenser for the detection of small numbers of living organisms renders it of value for rapid diagnostic purposes. Stained Material. Fuller accounts of the technique of fixed and stained material will be found in Bolles Lee and in Langeron, already mentioned. Thin Films. — For the examination of blood-inhabiting Protozoa, it is necessary to make first thin films or smears of blood. There are many ways of doing this, and opinions differ as to their respective merits. A simple method is to take a straight surgical needle about 2 in. long, the eye of which has been removed, and a clean glass slide. The patieiit's skin is pricked, and when the bead of blood reaches the size of a small pin's head, the slide is applied to the surface of the blood, about J in. from the far (left-hand) end of the slide. The shaft of the needle is laid across the drop of blood, which spreads between the slide and the needle. The latter is drawn evenly along the slide towards the right. The film is dried by waving it in the air. The film should possess a straight edge parallel with that of the slide and should be as uniform and thin as possible. Another glass slide may be used as a spreader, or a cover-slip or thin glass rod may be employed. Thick Films. — These are of service in detecting malarial paiasites or trypanosomes, especially when the parasites are few. The method of Ross, or a modification thereof, has been much used. A small drop of fresh blood is spread evenly and quickly with a needle-point over a square area somewhat less than that of an ordinary square cover- glass. The blood is allowed to dry. The film is then carefully dehaemoglobinized in water in which there is a trace of acetic acid. The dehaemoglobinizing fluid is then carefully drained off and the 47 748 THE ANIMAL PARASITES OF MAN film again dried. It is fixed in absolute alcohol and stained with Romanowsky's solution. A cubic millimetre of blood divided into quarters may be thus dehaemoglobinized and stained. The parasites in such a cubic millimetre of blood may be counted. Such a proced- ure was followed by R. Ross and D. Thomson/ in determining the periodic variation of the numbers of trypanosomes in the blood of a patient, as mentioned and figured on pp. 78 and 79. For cytologica details of various Protozoa, thin film preparations on cover-slips or slides are often useful. Cover-slip preparations are preferable, unless the organisms under investigation are extremely scanty. The medium containing the organisms, such as blood, lymph, intestinal contents, sputum, scrapings of ulcers, and urine, is spread thinly, either alone or diluted with a little physiological salt solution, on the cover-slip. Fixation while still wet is necessary. Various methods are employed. Fixatives. — A useful procedure is to fix the wet film by exposure to 4 per cent, osmic acid vapour for ten to thirty seconds, then place in absolute alcohol for five minutes to harden. Grade down from absolute alcohol through 90 per cent., 70 per cent., 50 per cent., and 30 per cent, alcohols to water. Stain wet with a suitable stain such as haematoxylin, and gradually dehydrate by grading through the necessary strengths of alcohol, clear in xylol or other oily clearing medium and mount in Canada balsam. Other fixatives may be employed, such as are also useful for fixing pieces of tissue for sectioning. Films or smears on cover-slips while still wet are floated on the surface of the fixative in a watch glass. Some good fixatives of wide application are : — Schaudinn's Fluid. — This consists of a mixture of Saturated aqueous solution of corrosive sublimate ... ... 2 volumes Absolute alcohol ... ... .., ... ... i volume Two modifications of Schaudinn's formula may be found useful. A saturated solution of corrosive sublimate in physiological salt solution may be substituted for the aqueous one, and the addition of a few drops of glacial acetic acid to either of the preceding mixtures may be made. Some workers prefer to use hot fixatives, raised to a temperature of about 50^" C. Fixation by corrosive sublimate solutions must be followed by thorough removal of the mercury salt by washing repeatedly in 30 per cent, alcohol or with iodine-alcohol. ' Froc. Roy. Soc.^ B, Ixxxii, p. 411. APPENDIX ON PROTOZOOLOGY 749 Bouin's Fluid, or modifications thereof, is also very useful for wet fixation. Bouin's picro-formol solution consists of : — Saturated aqueous solution of picric acid ... 30 volumes Formalin, 40 per cent. ... ... ... 10 ,, Acetic acid, glacial ... ... ... 2 ,, The best-known modification is one due to Duboscq and Brasil, and often known as Bouin-Duboscq Fluid. Its formula is as follows : — Alcohol, 80 per cent. ... ... ... 150 c.c. Formalin, 40 per cent. ... ... ... 60 ,, Acetic acid, glacial ... .., ... 15 ,, Picric acid ... ... ... ... i grm. Thorough washing of the smear or cover-slip preparation with 70 per cent, alcohol until the yellow colour disappears is necessary to remove excess of fixative. Other fixatives, which may be of use, more especially for fixing small pieces of tissue for sectioning, are the solutions of Flemming (chromo-aceto-osmic acids) and of Zenker (sublimate-bichromate- acetic, with sodium sulphate). Regarding the time of fixation, there is much difference of opinion. Usually, exposure to or contact with the fixative for five minutes is sufficient in the case of films or smears. Material for sections should be cut into small cubic pieces, of a thickness of about 5 mm. (J in.). One or two hours should be sufficient time for the fixation of such pieces of tissue, though some, as Langeron, prefer a longer time of fixation. On the other hand, Gustav Mann' recom- mends a short fixation period. The excess of fixative should be thoroughly washed out of the tissue in the manner appropriate to the particular fixative used. If it is desired to keep the tissue for some time before sectioning and staining, it should be transferred to 70 per cent, alcohol. When fluid fixatives are employed, large quantities of the fixing media are necessary. The volume of the fixative should be at least ten to twenty times that of the object, and the latter should be sus- pended in the middle of the fixative. The tissue should be fixed as soon as possible after the death of the host. For sectioning tissue parasitized by Protozoa, embedding in parafifin is generally recommended. Microtome sections should not, if possible, exceed 5 //, in thickness. Details of special procedures must be sought in larger works. Staining. — Here, as with fixatives, much choice is presented. The various modifications of the Romanowsky stain have aided greatly in 1 << Physiological Histology," 1902, Clarendon Press, Oxford. 750 THE ANIMAL PARASITES OF MAN the detection of various Protozoa parasitic in the blood. Such stains, however, leave something to be desired in the revealing of finer cytological details. Other stains, more especially the haematoxylins, must be employed for cytological purposes. FormulcC of some of the principal Romanowsky and ha^matoxylin stains may now be given. The underlying principle of the Romanowsky Stain is the reaction between alkaline methylene blue and eosm, forming the so-called eosinate of methylene blue which stains chromatin purplish-red. A solution of medicinal methylene blue after having been subjected to the action of an alkali, such as sodium carbonate, becomes partly converted into certain derivatives, the chief of which are methylene azure and methylene violet. These substances are also present in matured polychrome methylene blue. The formula of a slightly modified Romanoivsky Stain which gives excellent results is given below : — Two stock solutions are required — Solution A. — Methylene blue, pure medicinal ... ... i*o grm. Sodium carbonate ... ... ... ... 0*5 ,, W^ater ... ... ... ... ... loo-o c.c. Keep in a warm incubator for two or three days, until the solution is distinctly purple in colour. It improves with age. Solution B. — Eosin, water soluble, extra B. A. ... ... 10 grm. Water ... ... ... ... ... 1,000 'O c.c. This solution must be kept in the dark, in dark-tinted (amber-coloured) bottles, as unfortu- nately it is decolorized by light. Before use each stock solution must be diluted. Thus, make up 5 c.c. of each stock solution to 100 c.c. by adding distilled water. For staining, i volume of solution A is added to 2 or 3 volumes of solution B. Mix thoroughly by shaking, pour the mixture over the film, previously fixed in absolute alcohol, and stain for ten to fifteen minutes. Wash carefully in running w^ater, then dry. The cytoplasm of a protozoan parasite will be stained blue, the chromatin purplish- red and vacuoles or very tenuous protoplasm wall remain colourless. The exact proportions of solutions A and B, which must be mixed together, should be determined by experiment. Freshly mixed stain must be used on each occasion. Leishman's Stain is the precipitate resulting from the interaction of alkaline methylene blue and eosin. The washed and dried precipitate is collected and dissolved in pure methyl alcohol, which acts as a fixative ; 0*015 grm. of Leishman powder may be dissolved in 10 c.c. of methyl alcohol for staniing films. The film is covered with the solution for one minute, twice the volume of water is then added and mixed with the stain on the slide. The staining is then appendix: on protozoology 751 continued for five to ten minutes, and the film is finally washed with water. Gienisii's Stain. — This should be procured ready made. Azure II is a mixture of methylene azure and methylene blue. (Methylene azure is sometimes known as Giemsa's Azure I.) The formula given by Giemsa himself in 19 12 is : — Azure Il-eosin ... ... ... ... 3*0 grm. Azure II ... ... ... ... ... 0'8 ,, Glycerine, pure ... ... ... ... 125*0 ,, Methyl alcohol, pure ... ... ... 375"0 ,, The film is first fixed in absolute alcohol. The proportion of stain usually used is one drop of stain to i c.c. of water. Stain for about ten minutes and then wash in water. The details of the application of the Giemsa stain to films fixed wet and to sections must be sought in larger works on technique. These works should also be consulted for information regarding the use of Pappenheim's Panchrome mixture. There are numerous formulae of stains containing ripened Hama- toxylin or its essential principle, Hceniatein. A mordant is necessary, one of the alums being usually employed. The mordant may be included as an ingredient in the staining mixture, or it may be used separately as in the case of the so-called iron-haematoxylins, wlierein ferric ammonium alum is used separately and is followed by staining with haematoxylin or haematein. A few of these stains of general application may now be mentioned. Delafield's {or Grenachers) Hcematoxylin, Hgematoxylin crystals ... ... ... 4 gi^m. Absolute alcohol ... ... ... ... 25 c.c. Saturated aqueous solution of ammonia-alum ... 400 ,, Mix these ingredients, and leave exposed to light and air for three to four days. Filter and add — Glycerine ... ... ... ... ... 100 c.c. Methyl alcohol ... ... ... ... ico ,, Allow the mixture to stand until the colour is sufficiently deep, then filter and place in a stoppered botde. The solution should be allowed to ripen for at least two months before use. Dilute aqueous solutions of the stain are of service for films and for sections. A trace of acetic acid may be added at the moment of use, for sharp differentiation. Ehrlich's acid haematoxylin, Mayer's ha^malum, and Mayer's glych^malum are also useful. Their formulae will be found in larger works. 75^ THE ANIMAL PARASITES OF MAN The chief Iron-H cematoxylin Stain is that devised by Heidenhain. Unfortunately the procedure involved is a long one, and various modifications have been made to obviate this disadvantage. Hsema- tein maybe used instead of ripened haematoxylin. One efficacious modification of Heidenhain's stain is that of Rosenbusch. The smear or tissue, after fixation, must be graded downwards through the alcohols to water. Mordant for one and a half hours in a 3J- per cent, aqueous solution of ferric ammonium sulphate. Stain for about three minutes in i per cent, solution of ripe haematoxylin or haematein in absolute or 96 per cent, alcohol, to which a drop of saturated aqueous sokition of lithium carbonate, sufficient to produce a wine-red colour, has been added. Differentiate under the microscope with a very dilute solution of the ferric ammonium sulphate. Wash, gradually dehydrate, clear and mount in balsam. It must be remarked that iron-haematoxylin is a regressive stain, hence great care must be exercised in differentiating with the iron alum. Gentian Violet. — A i per cent, alcoholic solution of gentian violet, or of methyl violet, or of crystal violet, will be found useful for stain- ing spirochaetes. Methyl Green. — This substance is considered to be a chromatin stain, for either fresh or perhaps recently fixed tissues. A concen- trated aqueous solution contains about i per cent, of the stain. This should be added to a i per cent, solution of acetic acid. It may be used for demonstrating the nuclei of ciliates. In conclusion it is essential to remember that the actual magnifica- tion of figures of Protozoa should be given, and not merely the combination of objective and ocular that has been used, for unless the tube-length and distance of the drawing board from the ocular be also given, it is not possible to compute the magnification from such information. Drawings should always be made with the aid of a camera lucida, drawing prism or other form of projection apparatus. APPENDIX ON TREMATODA AND NEMATODA 753 APPENDIX ON TREMATODA AND NEMATODA. BY J. W. W. STEPHENS, M.D., B.C., D.P.H. TREMATODA. Artyfechinostomum sufrartyfex, Clayton Lane, 19 15. — Leiper thinks this may be the same as Echhiostoma jualayamnn, Leiper, 1 91 1, which species Odhner assigns to the genus Euparyphium. Metagonimus (Yokogawa) yokogawai occurs in dogs in Shanghai. Encysted cercariae probably in the perch. Opisthorchis sp. — Skin covered with spines. Gut forks almost reach end of body. (Esophagus two to three times length of pharynx. Ovary multilobed. Ovary and testes in posterior fourth of body. Vitellaria end opposite the ovary. Distinguished from 0. felineus by presence of spines and lobed ovary ; from 0. pseudo- felineus and 0. noverca by the lobed ovary, and by the fact that the yolk glands do not extend as far as the anterior testis. It agrees with Poirier's description of 0. viverrini in the Indian civet cat, but whether this species has spines on the cuticle is not known. Habitat. — Man in Chiengmai (Malay States). Fifteen per cent, of prisoners in the jail showed the ova of this species in their faeces. Schistosome cercariae. Schistosome cercariae belong to the furcocercous division of the Distomata cercariae. Distomata cercariae. Body without a floating membrane. Tail absent, or if present not cleft to the base. Mouth anterior, gut forked. Oral sucker present. Ventral sucker near middle of body. Eyes generally absent. Group Fercocercous cercariae, Cercariae single (not in colonies). Tail forked at its end. Family. Schistosomidae. Pharynx absent. 754 THE ANIMAL PARASITES OF MAN Cercaria bilharzia, Leiper, 1915. Pigment spots (eyes) anterior to ventral sucker absent, cuticular keel on forks of tail absent. In Bullinus sp. and Planorbis hoissyi in Egypt, (?) in Physopsis africanuy South Africa. Adult form, Schistosoma hcematohium. Cercaria bilharziella, Leiper, 1915. Cuticular keel on tail forks present. Pigment spots (eyes) in front of ventral sucker present. In Planorbis hoissyi and P. mareoticus, and in Melanin sp. Adult form(?). For characters of numerous other cercariae which occur in fresh water molluscs see " Die Susswasserfauna Deutschlands," Max Liihe, H. 17 (Gustav Fischer, Jena, 1909). The characters of Cercaria japonica of S. japonicuni in the mollusc Katayama nosophora and of C. mansoni have still to be defined. Schistosoma mansoni, Sambon, 1907. The evidence appears to be strong that terminal-spined eggs are not found in the West Indies, and that therefore the lateral-spined eggs found in faeces there belong probably to S. mansoni. If this be true, then the egg described by Stephens and Christophers in man in India probably also belongs to another species of Schistosome. NEMATODA. Ancylostomiasis. — Treatment: (i) Oleum chenopodii (U.S. P.), dose iri X to irt xv on a lump of sugar, three doses at two-hourly intervals, preceded and followed by a purge. It is cheap, not unpleasant to take, and non-toxic. Effective also against Ascaris lumbricoides. (2) Milk of the higueron Ficns laurifolia. A spoonful in milk, three times daily for three days followed by a purge. Described as a harmless but very sucessfu form of treatment. Ground-itch. — Completely cured in a few days by a 3 per cent, solution of salicylic acid in ethyl alcohol. Apply for five minutes twice daily. Ascaris lumbricoides can be kept alive for twelve days in Kronecker's solution ; NaHO 0069 grammes, normal saline 1,000 c.c. Eggs are laid and develop in about a fortnight at ordinary room temperature. At 70"^ C. they are readily killed. APPENDIX ON TREMATODA AND NEMATODA 755 Filariasis. — Dutcher and Whitmarsh have cultivated from the blood and from the exudation fluids of cases of filariasis (elephanti- asis, lymphangitis, etc.), in about sixteen cases, a bacillus resembling B. subtil is. Controls were negative. They propose the name Bacillus lymphangiticus for this organism, and they believe it to be the cause of the diseases grouped under the designation " filariasis." Oncocerca volvulus. — Unsheathed embryos (indistinguishable from those taken from the uterus of this worm) have been found in lymph- atic glands and in the blood (if considerable pressure is used so as to squeeze out lymph at the time of taking the linger blood, otherwise none occurs in the specimens). The measurements in dried films are : Nerve ring 237 per cent, of length ; Gi cell 69-6 per cent. ; end of last tail cell 96-3 per cent; total length 274-3 /n. Strongyloides stercoralis. — Pathology : They occur in the wall of the intestine and may be associated with ulceration. They also occur in lymphatics and blood-vessels. 756 THE ANIMAL PARASITES OF MAN BIBLIOGRAPHY. [In the following pages the letters C. f. B., P. u. Inf. are used to indicate the Centralhlatt fur Bakteriologie_, Pathologie und Infektions-Krankheiten.'] (A) PROTOZOA (pp. 25 to 210, 617 to 637, and y^z to 742). [This list affiles to the earlier literature only. More recent references are given as footnotes in the text.'] (a) General. BtJTSCHLl, O. Protozoa in Bronn's Klass. u. Ordn. d. Tierreichs, Leipz., 1880- 1889. Calkins, G. N. The Protozoa, Columbia Univ. Biol. Ser., vi. New York, iqoi. Delage, Y., and E. Herouard. Traite de Zool. Concr., i, La cellule et le& protozoaires, Paris, i8q6. Farmer, J. B., J. J. Lister, E. A. Minchin and S. J. Hickson. Protozoa, in A Treatise on Zoology, edited by E. Ray Lankester, London, 1903, i, 2. Lang, A. Lehrb. d. vergl. Anatomic d. wirbellos. Tiere, 2. Aufl., 2. Lief,. Protozoa, Jena, 1901. (b) Pathogenic Protozoa in General. Doflein, F. Die Protozoen als Parasiten u. Krankheitserreger, Jena, 1901 ;. Lehrbuch der Protozoenkunde, 1912. Doflein, F., and S. v. Prowazek. Die pathog. Protoz. mit Ausnahme d. Hamospor., in Handb. d. path. Mikroorganism. ; issued by W, Kolle and A. Wassermann, 11. and 12. Lief, Jena, 1903. Kastner, p. Die tierpathogenen Protozoen, Berlin, 1906. KISSKALT, K., and M. Hartmann. Praktikum der Bakteriologie und Proto- zoologie, Jena, 1907. LtJHE, M. Die im Blute schmarotzenden Protozoen und ihre nachsten Ver- wandten, in Handb. d. Tropenkrankh., issued by C. Mense, Leipz., 1906,, iii. Pfeiffer, L. Die Protozoen als Krankheitserreger, 2. Aufl., Jena, 1891 ; Supple- ment, Jena, 1895. Roos, E. Die im menschl. Darm vork. Protozoen u. ihre Bedeutg., Med. Klinik, 1905, i, p. 1328. SCHNElDEMtJHL, G. Die Protozoen als Krankheitserreger der Menschen und der Haustiere, Leipz., 1898. SlEVERS, R. Zur Kenntn. d. Verbreitg. d. Darmparas. d. Mensch, Helsingfors, 1905, Festschrift f. Palmen. Ward, H. B. Protozoa, Wood's Ref. Handbook of the Med Sci., 1904, viii. Class I — Sarcodina (pp. 29 to 50). Order. Amoebina (p. 29). Entamoeha coli; Entamceba histolytica (pp. 32 to 41, 618 to 620, and y2>3). Blanchard, R. Traite de Zool. medic, 1885, i, Paris, p. 15. — Maladies paras., 1895, p. 658. Bowmann, M. H. Dysentery in the Philippines, Journ. Trop. Med., 1901, iv,. p. 4^0. BIBLIOGRAPHY 757 Bunting, C. H. Hsematogenous Amoebic Abscess of the Lung, Arch. f. Schiffs- u. Tropenhyg., iqo6, x, p. ^'i- Calandruccio. Anim. par. dell' uomo in Sicilia, Atti Accad. Gioen., iv, iSgo^ ii, p. 95. Casagrandi, O., and P. Barbagallo. SulP amoeba coli, Boll. Accad. Gioen. sci,. nat., Catania, 1895. Entamooba hofninis s. Ainosba coli Losch, Annal. d'Igiene sperim., 1897^ vii, I. Celli, a., and R. FiOCCA. Beitr. z. Amoebenforsch., ii, C. f. B. u. Par., 1894,. xvi, p. 329; Ric. int. alia biol. d. Amoebe, Bull. Accad. med. Roma, 1894- 95, xxi, p. 285; abstracted in C. f. B., P. u. Inf., 1897, i, xxi, p. 290. Councilman, W. P., and H. A. Lafleur. Amoebic Dysentery, Johns Hopkins Hosp. Rep., 1891, ii, p. 395. Craig, C. F. Etiology and Pathology of Amoebic Infection of the Intestines- and Liver, Intern. Clin., Philad., 1905 (14), iv, p. 242. Cunningham, D. Seventh Ann. Rep. San. Comm. of India, Calcutta, 1871. — Unters. iib. d. Verb, mikrosk. Organ, z. Cholera in Indien, Zeitschr. f. Biol.,. 1872, viii, p. 251; Quart. Journ. Micros. Sci., 1881 {2), xxi, p. 234. Grassi, B. Dei protozoi par. e spec, di quelli che sono nell' uomo, Gazz. med. ital.-lomb., 1879 (8), i, p. 445; Int. ad ale. prot. endop., Atti soc. ital. sci. nat., 1882, xxiv, p. i; Morf. e sist. di ale. prot. par., Atti Ace. Lincei. Rendic. {4), iv, i, p. 5; Signif. patol. d. prot. par. dell' uomo, ibid., p. 83. Gross, A. Beobacht. iib. Amoebenenteritis, Arch. f. klin. Med., 1903, Ixxvi,. p. 429. Harris, H. F. Amoebic Dysentery, Amer. Journ. of Med. Sci., April, 1898. — Experimentell bei Hunden erzeugte Dysent., Arch. f. path. Anat., 1901, clxvi, p. 66; On the Alterations Produced in the Large Intestine of Dogs by the Amoeba colij Philadelphia, 1901. Hoppe-Seyler, G. Ub. Erkrankung des Wurmfortsatzes bei chron. Amoeben- enteritis, Miinch, med. Wochenschr., 1904, No. 15. Jaeger, H. Die in Ostpreuss. heim. Ruhr eine Amoebendysent., C. f. B., P. u. Inf., 1902, i Abt. Orig., xxxi, p. 551. — Erwiderg. a. d. Bemerk. Shigas, ibid.^ 1902, xxxii, p. 865. JANOWSKI, W. Zur Atiol. d. Dys., C. f. B., P. u. Inf., 1897, i, xxi, pp. 88, 151,. 194, 234. JtJRGENS. Zur Kenntn. d. Darm-Amoeb. u. d. Amoeben-Enteritis, Veroff. a. d. Geb. d. Milit.-Sanitatswes., Berl., 1902, Heft 20, p. iii. Kartulis. tJber Riesenamoeben ( .?) bei chron. Darmentdg. d. Agypt, Virch.. Arch. f. Path., 1885, xcix, p. 145; Zur Atiol. d. Dysent. in Agypt, C. f. B. u. Par., 1887, V, p. 745; Ub. trop. Leberabsc. u. ihr Verb. z. Dysent., Virch. Arch. f. Path., 1889, cxviii, p. 97; Einiges iib. d. Path. d. Dysenterie-Amceb., C. f. B. u. Par., 1891, ix, p. 365; Article : Dysenteric- in Spec. Path. u. Ther. v. H. Nothnagel, Wien, 1896, v, 3. — Gehirnabscesse nach dysent. Leberabsc, C. f. B., P. u. Inf., 1904, i Orig., xxxvii, p. 527. Koch, R., and G. Gaffky. Bericht iib. d. Tatigkeit d. z. Erforschg. d. Cholera ents. Kommiss., Arb. a. d. kais. Gesundheitsamt, 1887, iii. Kovacs, F. Beob. u. Vers. iib. d. sog. Amoeben-Dys., Zeitschr. f. Heilkde., 1892, xiii, p: 509. Kruse, W., and Pasquale. Eine Exped. nach Agypt, Deutsch. med. Wochen- schr., 1893, No. 15, p. 354; No. 16, p. 368. Untersuch. iib. Dys. u. Leberabsc, Zeitschr. f. Hyg., 1894, xvi, p. i. ■^58 THE ANIMAL PARASITES OF MAN Lambl. Aus d. Franz-Joseph-Kinderspit. in Prag, i860, i, p. 362. Iesage, a. Culture de Tamibe de la dysenteric des pays chauds, Ann. Inst. Pasteur, 1905, xviii, p. 9; 1905, xix, p. 8. Lewis. Sixth Ann. Rep. San. Comm. of India, Calcutta, 1870. LOSCH, F. Massenh. Entw. v. Amoeben im Dickd., Virch. Arch. f. Path., 1875, Ixv, p. 196. JMarchoux. Note sur la dysenteric d. pays chauds, C. R. Soc. Biol., Paris, 1899 (11), i, p. 870. MUSGRAVE, W. E., and M. T. Clegg. Amoebae, their Cultivation and Etiological Significance. " Treatment of Intestinal Amoebiasis," in the Trop. Manila, 1904, Bur. of Govern. Lab., Biol. Lab., No. 18. NORMAND. Note sur deux cas de colite parasit., Arch. med. nav., 1879, xxxii, p. 211. Quincke and Roos. tJber Amoebenenteritis, Berl. klin. Wochenschr., 1893, XXX, No. 45, p. 1089. Roos, E. Zur Kenntn. d. Amoebenenteritis, Arch. f. exper. Path. u. Pharm., 1894, xxxiii, p. 389. RUGE, A. Amocbenruhr., Handb. d. Tropenkrankh., issued by C. Mense, 1906, iii, p. I. JSCHAUDINN, Fr. Untcrs. lib. d. Fortpfianz. einig. Rhizopod, Arb. a. d. kais. Gesundheitsamt, 1903, xix, 3, p. 547. SCHUBERG, A. Die paras. Amocb. d. menschl. Darms, C. f. B. u. Par., 1893, xiii, pp. 598, 654, 701. Shiga, K. Bemerk. zu Jaegers Die in Ostpreuss. einh. Ruhr cine Amoebendys., C. f. B., P. u. Inf., 1902, i Abt. Orig., xxxii, p. 352. Strong, R. P., and W. E. Musgrave. Report on the Etiology of the Dysentery of Manila, Rept. Surg. -Gen. of the Army to the Secretary of War, Wash- ington, 1900, p. 251. Verdun. Sur quelq. caract. specif, de I'amibe de la dysenteric et des absces trop. du foie, C. R. Soc. Biol., 1904, Ivi^ p. 183. WOOLLEY, P. G., and W. E. Musgrave. The Pathology of Intestinal Amcebia- sis, Dept. of the Int. Bureau of Government Lab., No. 32, Manila, 1905, Journ. Amer. Med. Assoc, Chicago, 1905, xlv, p. 1371. Entamoeba huccalis (pp. 43 and 620). Leyden, E. v., and W. Loewenthal. Entam. hucc. Prow, bei eincm Fall von Carcinom d. Mundbod, Charite-Ann., Berl., 1905, xxix; Berl. klin. Wochenschr., 1905, xlii. No. 7, p. 187. Prowazek, S. Entamceba buccalis n. sp., Arb. a. d. kais. Gesundheitsamt, Berl., 1904, xxi, i, p. 42. Tietze, Al. Ein Protozoenbef. i. ein. erkrankt. Parotis., Mitt. Grenzgeb. Med. u. Chirurg., 1905, xiv, p. 302. Entatnceba undulans (p. 43). •Castellani, a. Dysentery in Ceylon, Journ. of the Ceylon Branch of the Brit. Med. Assoc, 1904. — Observations on some Protozoa found in Human Faeces, C. f. B., P. u. Inf., 1905, i Abt. Orig., xxxviii, p. 67. BIBLIOGRAPHY 759 Entamceba kartulisi (pp. 44 and 734). DOFLEIN, F. Die Protoz. als Paras, u. Krankheitserreg., Jena, 1901, p. 30. Flexner. Amoebse in an Abscess of the Jaw, Johns Hopkins Hosp. Bull., 1892^ No. XXV ; abstracted in C. f. B., P. u. Inf., 1893, xiv, p. 288. KarTULIS. Ub. pathog. Protoz. b. Mensch., Zeitschr. f. Hyg., 1893, xiii, p. 9. — trber Amoebenosteomyelitis d. Unterkief, C. f. B., P. u. Inf., 1903, i Abt.^ Ref. xxxiii, p. 471. Amceba gingivalisj A. buc calls and A. dentalis (pp. 44 and 733). Grassi, B. Gazz. med, ital.-lomb., 1879 (8), i, No. 45, p. 445. Gros, G. Fragm. d'helm. et de phys. micros., Bull. soc. Imp. d. natur. de Moscou, 1849, ij 2, p. 555. Steinberg. In Zeitschr. f. neuere Med, (Russ.), issued by Walter in Kiew,. 1862, Nos. 21-24. Craigia [Parajnccba) honiinis (pp. 45 and 734). Cr.aIG, Ch. F. a new Intestinal Parasite of Man : Paramceba hominis, Amer. Journ. Med. Sci., 1906, N.S. cxxxii, Philad. and New York, p. 214. SCHAUDINN, Fr. tJber den Zeugungskreis von Paramoeba eilhardi n. g. n. sp.^ Sitzgsber. Kgl. Pr. Akad. d. Wiss., Berlin, Phys.-math. CI., 1896, No. 2. Entamoeba -pulnionalis (p. 45). Artault, St. Flore et faune d. cav. pulm.. Arch, de paras., 1898, i, p. 275. Blanc, L. Sur une Amibe viv. accid. dans le poumon du mouton, Ann. Soc. Linn. Lyon, 1899 (2), xlv, p. 529. Amceba uro genitalis (pp. 45, 46). Baelz, E. Ub. einige neue Paras, d. Mensch., Berl. klin. Wochenschr., 1883^ p. 237. Jeffries. Present, of a Specimen of Urine containing Amoebae. Med. Rcc.^ New York, 1904, xlvi, p. 356. JiJRGENS. In Deutsche med. Wochenschr., 1892, p. 454. KaRTULIS. Pathog. Prot. b. Mensch, Zeitschr. f. Hyg., 1893, xiii, p. 2, Anm. 2. PoSNER, C. Ub. Amoeben im Harn, Berl, klin, Wochenschr., 1893, xxx,. No. 28, p. 674. WlJNHOFF, J. A. Over amoeburie, Nederl. Tijdschr. v. Geneeskde., 1895^ p. 107. Ajncuba iniurai (p, 46). IjiMA, J. On a New Rhizopod Parasite of Man, Ann, zool. japon., 1898, ii, 3^ p. 85; abstracted in C, f. B,, P, u. Inf., 1899 (i), xxv, p. 885. MlURA, K, Amoebenfund i, d, Punktionsfiiiss. bei Tumoren d, Peritonealh., Mitt. med. Facult. d. kais. Jap. Univ., Tokyo, 1901, v, p. i. Chlamydo-phrys and Leydenia (pp. 47 to 50). ClENKOWSKI, L. Ub. einige Rhizop. u. verwandte Organismen, Arch. f. mikr. Anat., 1876, xii, p. 39. LauENSTEIN, C. Ub. ein. Fund von Leyd. gemniif., Deutsche med, Wochen- schr., 1897, xxiii, p, 733' Leyden, E. v., and F. Schaudinn. Leyd. genimif,, ein neuer i. d, Ascites- Fliissigk. d. leb, Mensch. gefund. amoebenahnl. Rhizop., Sitzgsb. kgl. Preuss. Akad, d, Wiss,, Berlin, 1896, xxxix, p. 951. y6o THE ANIMAL PARASITES OF MAN Leyden, E. v. Zur Atiol. d. Carcin., Zeitschr. f. klin. Med., igoi, xliii, p. 4. SCHAUDINN, Fr. Untersuch. iib. d. Fortpflanz. einig. Rhizopod., Arb. a. d. kais. Gesundheitsamt, 1Q03, xix, 3, p. 560. Schneider, A. Beitr. z. Naturgesch. d. Infus., Miillers Arch, f. Anat., Phys. u. wiss. Med., Jahrg. 1854, p. 191. Class 11. — Mastigophora (pp. 50 to 128, 620 to 633, and 734 to 741). BiLAND, J. Beitr. z. Frage d. Pathog. d. Flagellat., Deutsch. Arch. f. klin. Med., 1905, Ixxxvi, p. 274. Blochmann, F. Mikrosk. Tierw. d. Siisswassers, 2. Aufl., 1895. Kent, W. S. Manual of the Infusoria, London, 1880-81. Prowazek, S. Flagellatenstudien, Arch. f. Protistenkde., 1903, ii, p. 195. Senn, G. Flagellata in Engler und Prantl^ Die natiirlich. Pflanzenfam., Lief 202, 203, Leipzig, 1900. Stein, F. v. Der Organismus der Infus., iii, Der Org. d, Flagellaten, Leipzig, 1878. Trichomonas vaginalis (pp. 52, 53, and 734). Baatz, P. Trich. vag. in der weibL Harnblase, Monatsber. f. UroL, 1902, vii, 8. Blochmann, F. Bemerk. iiber einige FlagelL, Z. f. wiss. ZooL, 1884, xl, p. 42. Dock, G. 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Atti della societa per gli studi della malaria, Roma, 1899-1906, i-vii. BiGNAMi, A. Das Tropenfieber u. die Sommer-Herbstfieber d. gemass. Klimate, C. f. B., P. u. Inf., 1898, (i), xxiv, p. 650. 774 " THE ANIMAL PARASITES OF MAN BiGNAMi and Bastanielli. Osserv. nelle febbre malar, estivo-autunn., Rif. med., 1890, p. 1334- Studi suir inf. mal., Bull. R. Accad. med., Roma, 1893-94, xx. Sulla strutt. dei par. mal. e in specie dei gameti d. par. est.-aut., Atti Soc. stud. d. mal., 1899, i. Celli, a., and F. Sanfelice. Uber d. Paras, d. roth. Blutk. im Menschen u. in Thieren, Fortschr. d. Med., 1891, pp. 499, 541; Ann. istit. d'igiene sperim., Roma, 1891, N.S. i. Danilewski, B. Zur Parasit. d. Blutes, Biol. Centralbl., 1885-86, v, p. 529. — La parasitologie comp. du sang, Charkow, 1889. — Sur les microb. d' infect, malar, aigue et chron. chez les oiseaux et chez rhomme, Ann. Inst. Pasteur, 1890, p. 753; 1891, p. 758. — tJber den Polymitus malariae, C. f. B. u. Par., 1891, ix, p. 397. 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Feletti. Uber d. Paras, d. Malaria, C. f. B. u. P., 1890, vii, pp. 396, 340; Malariapares. in d. Vogeln, ibid.^ 1891, iv, pp. 403, 429, 461; Weiteres zur Malariafrage, ibid.^ 1891, x, pp. 449, 481, 517. Jancs6, N. Zur Frage d. Inf. d. Anopheles claviger mit Mal. -Paras, b. nied. Temp., C. f. B., P. u. Inf., 1904, i Abt. Orig., xxxvi, p. 624; Der Einfi. d. Temp. a. d. geschl. Generationsentw. der Mal. -Paras, u. auf d. exper. Mal.-Erkr., ibid., 1905, xxxviii, p. 650. Koch, R., and R. Pfeiffer. Beitrage z. Protozoenforsch. : I, Die Coccidien- krankh. d. Kaninchen, Berlin, 1892. Arztl. Beob. i. d. Tropen, Verhandl. d. D. Kol.-Ges. Abt. Charlottenburg- Berlin, 1897-98, Heft 7, p. 280. Die Malaria in Deutsch-Ostafrika, Arb. kais. Gesundheitsamt, 1898, xiv, p. 292. — — Reiseberichte liber Rinderpest . . . trop. Malaria . . . Berlin, 1898. Ergebn. d. wiss. Exped. nach Italien z. Erf. d. Malaria, Deutsche med. Wochensch., 1899, p. 69, [Zwei] Berichte iib. d. Tatigkeit d. 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Protistenkunde., 1904, iv, p. i. Vaney, C., and A. Conte. Sur une nouv. microsp., Pleisto-phora mirandellaej paras, de I'ovaire ^^ Alhtirnns mirandella Blanch., C. R. Acad. Sci., Paris, 1901, cxxxiii, p. 644. Order. Actinomyxidia (p. 187). Some earlier literature will be found quoted in : — Caullery, M., and F. Mesnil. Recherches sur les Actinomyxidies, I, Arch. f. Protistenkde., 1905, vi, 3, p. 272. Order. Sarcosporidia) (pp. 187 to 194). Bertram, A. Beitr. zur Kenntn. d. Sarcosp., Zool. Jahrb., 1892, v, p. 581. Blanchard, R. Sur un nouv. type d. Sarcospor., C. R. Acad. Sci., Paris, 1885, c, p. 1599- — Note sur les Sarcospor. et sur un ess. d. classif. d. ces sporoz.. Bull. Soc. Zool. France, 1885, x, p. 244. DaMmann, C. Psorosp.-Krankh. beim Schaf., Arch, f path. Anat., 1867, xli, p. 283. Eecke, J. VAN. Sarcosporidien, Geneesk. Tijdschr. v. Nederl. -Indie, 1892, xxxii; Jaarsverl. Labor, path. An. en Bact. te Weltevreden (1892), Batavia, 1893. Foret, p. Observ. rel. au devel. de la cuticle chez le Sarcocystis tenella, Arch. d'Anat. Micr., 1903, vi, p. 86; C. R. Soc. Biol., Paris, 1903, Iv, p. 1054- Hessling, v. Histol. Mittheil., Zeitschr. f.. wiss. Zool., 1854, v, p. 189. Koch, M. uber Sarcosporidien, Verh. V. intern. Zool. Congr., Berlin, Jena, 1902, p. 674. — Die experimentelle Ubertrag. d. Miescherschen Schlauche, Berl. klin. Wochenschr., 1904, li, p. 321. KORTE, W. E. de. On the Presence of Sarcosporidia in the Thigh Muscle of Macaciis rhesus, Journ. of Hyg., Cambridge, 1905, v, p. 451. Laveran, a., and F. Mesnil. Morph. d. sarcospor., C. R. Soc. Biol., 1899 (x), vi, p. 245. 49 780 THE ANIMAL PARASITES OF MAN Leisering and Winkler. Psorosp.-Krankh. beim Schaf., Ber. lib. Veterin.- Wesen, Konigr., Sachsen, 1865; Arch. f. path. Anat., 1865, xxxvii, p. 431. Manz, W. Beitr. z. Kenntn. d. Miescherschen Schlauche, Arch. mikr. Anat., 1867, iii, p. 345- MiESCHER, F. t)ber eigent. Schlauche in d. Musk, einer Hausmaus, Ber. iiber die Verh. d. naturf. Ges., Basel, 1843, v, p. iq8; Reprinted in Verb. d. V. internat. Zool.-Congr., Berlin, Jena, igo2, p. 679. PiANA, G. P. Fasi evol. d. Sarcosp., La clinica veter., 1896, p. 145; C. f. B., P. u. Inf. (i;, XX, p. 39. Pluymers, L. Des sarcosp. et de leur role dans la pathog. d. myositis. Arch. Med. exp. et d'Anat. pathoL, 1896, p. 761; C. f. B., P. u. Inf. (i), xxii, p. 245- Rainey, G. Structure and Development of Cysticercus Cells as found in the Muscles of the Pig, Phil. Trans. Roy. Soc, 1858, cxlvii, p. iii. RiECK, V. Sporozoen als Krankheitserreger, Deutsche Zeitschr. f. Thiermed. u. vergl. Path., 1889, xiv, p. 75. RiEVEL and Behrens. Beitr. zur Kenntn. d. Sarcosp. und deren Enzyme, C. f. B., P. u. Inf., 1903, i Abt. Orig., xxxv, p. 341. RiVOLTA. Dei paras, veget., Torino, 1873; Giorn. an., fis. e pat. d. anim., 1874, vi, p. 25. ScHNElDEMtJHL, G. tJber Sarcosporidien, Thiermed. Vortr., Leipzig, 1897, iii, p. II. SiEBOLD, C. Th. V. Zusatz [zu Hessling. histol. Mittheil.], Zeitschr. f. wiss. Zool., 1854, V, p. 199. SlEDAMGROTZKY, O. Psorosp. in d. Musk. d. Pferde, Wochenschr. f. Thier- heilkde. u. Viehz., 1872, xvi, p. 97. Smith, Th. The Production of Sarcosporidia in the Mouse by Feeding Infected Muscle Tissue, Journ. Exp. Med., Baltimore, 1902, vi, p. i. — Further Observations on the Transmission of Sarcocystis maris by Feeding, Journ. Med. Res., 1905, xiii, p. 429. Stiles, Ch. W. Notes on Parasites : 18, Presence of Sarcosporidia in Birds, U.S. Dept. of Agric, Bur. of An. Ind., Bull. 3, 1893, p. 79. Sarcosforidia observed in Man (pp. 193, 194). Baraban and St. Remy. Sur un cas d. tub. psorosp. obs. chez I'homme, C. R. Soc. Biol., Paris, 1894 (x), i, p. 201. Le parasitisme d. sarcosp. chez Phomme, Bibliogr. anat., 1894, p. 79. Braun, M. Zum Vork. d. Sarcosporid. b. Menschen, C. f. B. u. Par., 1895 (i), xviii, p. 13. Kartulis. tJber pathog. Protoz. b. Menschen, Zeitschr, f. Hyg., 1893, xiii, p. i. LiNDEMANN. Ober d. hygien. Bedeutung d. Gregarinen, Deutsche Zeitschr. f. Staatsarzneikde., 1868. Rosenberg. Ein Befund von Psorosp. im Herzmuskel d. Mensch., Zeitschr. f. Hyg., 1892, xi, p. 435. VuiLLEMlN, P. Le Sarcocystis tenella, paras, de I'homme, C. R. Acad. Sci., Paris, 1902, cxxxiv, p. 11 52. Order. Haplosporidia (pp. 194 to 197). Caullery, M., and F. Mesnil. Rech. sur les Haplosporidies, Arch, de Zool. exp., 1905, ser. iv, iv, p. loi, in which a good bibliography is given. BIBLIOGRAPHY 781 MiNCHiN, E. A., and H. B. Fantham. Rhino s'poridium hinealyi n. g., n. sp., a New Sporozoon from the Mucous Membrane of the Septum Nasi of Man, Quart. Journ. Micros. Sci., 1905, xlix, p. 521. Class lY — Infusoria (pp. ig8 to 210). BOtschli, O. Studien liber ... die Conjugation d. Infusorien, Abh. d. Senckenb. naturf. Ges., 1876, x. Ehrenberg, Ch. G. Die Infusionsthierchen als vollkommene Organismen, Leipz., 1838. GuiART, J. Sur un nouv. infus. paras, de Phomme, C. R. Soc. Biol., Paris, 1003, iv; p. 245. Hertwig, R. Uber die Conjugation d. Infusorien, Abh. kgl. bayer. Akad. d. Wiss., 1889, iij Kl., xvii. Kent, Sav. A Manual of the Infusoria, London, 1880-1882. Maupas, E. Rech. exper. sur la multipl. des Infusoires cilies, Arch. Zool. exp., 1888 (2), vi. — Le rajeunissement karyogamique chez les Cilies, ihid.^ 1889, vii. Stein, Fr. v. Der Organismus der Infusionsthiere, Leipz., 1 859-1 867. Balantidium coli (pp. 200 to 204 and 637). ASKANAZY, M. Pathog. Bedtg. d. Bal, coli, Wien. med. Wochenschr., 1903, liii, p. 127; Verb. d. D. path. Ges., v (1902), Berlin, 1903, p. 224. Casagrandi, O., and P. Barbagallo. Bal. coli s. Parani. coli_, Catania, 1896, 8vo. Collmann, B. Fiinf Falle von Bal. coli im Darm d. Mensch., In. -Diss., Kgsbg., Pr., 1900. Ehrnroth, E. Z. Frage der Pathogenitat d. Bal. coli, Zeitschr. f. klin. Med., 1903, xlix, p. 321. Grassi, B. Signif. patol. d. prot. par. dell' uomo, Atti Accad. Lincei, Rendic, 1888 (4), iv, Sem. i, p. 86. Janowski, W. Ein Fall von Bal. coli im Stuhl, Zeitschr. f. klin. Med., 1897, xxxii, p. 415. (With copious literature compiled by Shegalow, Solowjew and Klimenko.) Klimenko, W. Beitr. z. Pathol, d. Bal. coli, Beitr. z. path. Anat. u. allg. Path., 1903, xxxiii, p. 281. KOSLOWSKI, J. J. Zur Lehre v. d. Infus., die als Paras, im Verdauungskan. d. Mensch. vork.. Arch. f. Verdauungskrankh., 1905, xi, p. 31. Kossler, K. Ein Fall von Balantidium-Colixis, Wien. med. Wochenschr., 1906, Ivi, p. 522. Maggiora, a. Microsk. u. bacter. Beob. wahrend einer epid. dysent. Dick- darmentzdg., C. f. B. u. Par., 1892, xi, p. 181. Malmsten, p. H. Infusorien als Intestinalthiere b. Mensch., Arch. f. path. Anat., 1857, xii, p. 302. Nagel. Ob. ein. Fall v. Infusorienenteritis, Miinch. med. Wochenschr., 1905, No. 44- Shegalow, J. P. Ein Fall von Bal. coli bei einem 5 jahr. Madchen, Jahrb. f. Kinderhlkde., 1899, xlix, p. 425. SiEVERS, R. tJber Bal. coli im menschl. Darm u. dessen Vork. in Schwed. u. Finland, Arch. f. Verdauungskrankh., 1900, v. Abstracted in C. f. B., P. u. Inf., 1900 (i), xxviii, p. 328. 7^2 THE ANIMAL PARASITES OF MAN SlEVERS, R. Zur Kenntn. d. Verbreit. v. Darmparas. d. Menschen in Finland, Helsingfors, 1905; P>stschr. f. Palmen, No. 10. SOLOWJEW. Bal. coli als Erreger chron. Durchfalle, C. f. B., P. u. Inf., iqoi (i), xxix, pp. 821, 849. [Solowjew's additional communication that appeared in " Wratsch," 190I5 Nos. 12 and 14, as well as in the " Russki Wratsch," 1902, No. 14, has been translated into German by Klimenko (1. c.).] Stokvis, B. J. Paramaecium in sputa, Nederl. Tijdschr. v. Geneeskde., 1884 (2), XX. Strong, R. P., and W. E. Musgrave. Preliminary Note of a Case of Infection with Balantid.him colij Bull. Johns Hopkins Hosp., Baltimore, 1901, xii, p. 31. The Clinical and Pathological Significance of Balantidhim coli, Dept. of Int. Bureau, Govt. Labor. Biol., Manila, No. 26, 1905, p. i. Wlajeff, G. Zur Frage d. Atiol. u. Behandlg. d. Dysenterie, Wracebraja Gaseta, Kemmern, 1905, xii, p. 913; abstracted in C. f. B., P. u. Inf., 1906, i, Ref. xxxvii, p. 757. WoiT, O. Drei neue Falle von Bal. coli i. menschl. Darm., Deutsch. Arch, f. klin. Med., 1898, Ix, p. 363. Balantidium niiniitmn (pp. 204 and 637). Jakoby, M., and F. Schaudinn. Ub. zwei neue Infus. i. Darm. d. Mensch., C. f. B., P. u. Inf., 1899 (i), XXV, p. 487. SCHULZ. Colfoda cucullus im Darm d. Mensch., Berl. klin. Wochenschr., 1899,. No. 16, p. 353. Nyctothenis (pp. 204 to 206 and 637). Castellani, a. Observations on some Protozoa found in Human Faeces, C. f. B., P. u. Inf., 1905, i Abt. Orig., xxxviii, p. 66. Jakoby, M., and F. Schaudinn. tJber zwei Infus. i. Darm d. Mensch., ibid.^ 1899 (i), xxv, p. 487. Krause, P. tJb. Infus. im Typhusstuhle nebst Beschreibg. einer bisher noch nicht beob. Art. {Balantiditan giganteic7n), Deutsch. Arch, f, klin. Med.,. 1906, Ixxxvi, p. 442. Chlamydozoa (pp. 207 to 210). Bosc, F. J. Les malad. bryocytiques (malad. a protozoaires), II, La maladie vaccinale {Plasmodium vaccines), C. f. B., P. u. Inf., i Grig., xxxvi, p. 630; xxxvii, pp. 39, 195. — Les malad. bryocyt., Ill, La variole et son parasite [Plasmodium variolae), ibid., xxxix, pp. 36, 129, 247, 389, 594. Calkins, G. N. The Life-history of Cytoryctes variola, Journ. Med. Research, Boston, 1904, xi, p. 136. Councilman, Magrath, Brinckenhoff, Tyzzer, Southard, Thompson, Ban- croft and Calkins. Studies on the Pathology and on the Etiology of Variola and of Vaccinae, Journ. Med. Research, Boston, 1904, xi, i, 1904. Gorini, C. tJber die bei der mit Vaccine ausgef. Hornhautimpf. vorkomm. Zelleinschlusse, C. f. B., P. u. Inf., 1900, i, Abt. xxviii, pp. 233, 589;. 1902, i Orig., xxii, p. in. BIBLIOGRAPHY 783 GUARNIERI, G. Ric. sulla patogenesi ed etiol. delT inf. vacc. e variolosa, Arch, sci. med., Torino, 1892, xvi. — Ulteriori ric. sulla etiol. e sulla patog. della inf. vacc, Clinica moderna, Firenze, 1897, iii. Ht)CKEL. Die Vaccinekorperchen, Beitr. z. pathol. Anat. u. z. allg. Path., Supp. II, 1898. LOEFF, A. VAN DER, in Weekbl. van het Nederl. Tijdschr. v. Geneeskde., 1886, No. 46. Mt)HLENS, P., and M. Hartmann^ Zur Kenntnis d. Vaccineerregers, C. f. B., P. u. Inf., 1906, i Orig., xxxxi, pp. 41, 203, 338, 435. Proscher, F. Uber d. kiinstl. Ziichtung eines " unsichtbaren " Mikroorgan. aus der Vaccine, C. f. B., P. u. Inf., 1906, i Orig., xl, 3, p. ZZI- Prowazek, S. Unters. iib. d. Vaccine, I, Arb. a. d. kais. Gesundheitsamt, 1905, xxii, p. 535. — Unters. iib. d. Vaccine, II, ihid.j 1906, xxiii, p. 525. Salmon, P. Rech. sur Pinfect. dans la vaccine et la variole, Annal. Inst. Pasteur, 1897, xi, No. 4. Schulze, F. E. Cytorrhyctes hiis Siegel, Berl. klin. Wochenschr., 1905, No. 21. SCHULZE, W. Impfungen mit Luesmaterial an Kaninchenaugen, Klin. Monatsbl f. Augenheilkde., 1905, xliii. — Das Verhalten der Cytorrhyctes luis in der mit Syphilis geimpften Kanin cheniris, Beitr. z. path. Anat. u. z. allg. Path., 1906, xxxix, p. 180. SlEGEL, J. Zur Kritik der bisherigen Cytorrhyctesarbeiten, C. f. B., P. u. Inf., 1906, i Orig., xlii, pp. 128, 225, 321, 480. Wasielewski, v. Beitr. z. Kenntnis d. Vaccineerregers, Zeitschr. f. Hyg., 1901, xxxviii, p. 212. (B) PLATYHELMINTHES (pp. 211 to 359, 638 to 698 and 753 to 755). Class II — Trematodes (pp. 212 to 282, 638 to 644, 753, and 754). \^M.B. — The literature, which is very comprehensive, has, up to the year 1892, been quoted and critically examined in Braun's monograph on the Trema- todes : Bd. iv, Abth. i, of Bronn's " Klass. u. Ord. d. Thierreichs," Leipz. Of works that have appeared later it is not possible to do more than enumerate the following.] Bettendorf, H. Musculatur u. Sinneszell. d. Tremat., Zool. Jahrb. Anat., 1897, X, p. 307. Blochmann, F. Die Epithelfrage bei Cestoden u. Trematoden, Hamburg, 1896. Braun, M. Arten d. Gattg. Clinostomuvi, Zool. Jahrb., 1900, Syst. xiv, p. i. — Trematoden d. Chelonier, Mitt. zool. Mus. Berlin, 1901, ii, p. i. . — Trematoden d. Chiroptera, Annal. K. k. naturh. Hofmus., Wien, 1900, xv, p. 217. — ZurKenntn. d. Tremat. d. Saugeth., Zool. Jahrb., 1901, Syst. xiv, p. 311- — Fascioliden d. Vogel, ihii., 1902, xvi, p. i. Brugge, G. Zur Kenntn. d. Excretionsgefasssyst. d. Cestoden u. Tremat., Zool. Jahrb. Anat., 1902, xvi, p. 208. FiSCHOEDER, F. Die Paramphistomiden d. Saugeth., Zool. Jahrb., 1903, Syst. xvii, p. 485- Gronkowski, C. v. Zum feineren Bau d. Tremat., Poln. Arch. f. biol. u. med. Wiss., 1902, i. Hein, W. Zur Epithelfrage d. Tremat., Zeitschr. f. wiss. Zool., 1904, Ixxvii, p. 546. 784 ^i^HE ANIMAL PARASITES OF MAN Looss, A. Die Distomen unserer Fische und Frosche, Stuttg., 1894; Bibl. zooL, xvi. - Rech. faune paras, de I'Egypte, I, M^m. Inst. 6gypt., 1896, iii, p. i. -- Weit. Beitr. z. Kenntn. d. Tremat.-Fauna Agypt, Zool. Jahrb., 1900, Syst. xii, p. 521. — Uber neue u. bekannte Tremat. aus Seeschildkroten, ibid.^ 1902, xvi, p. 411. Maclaren, W. Beitr. z. Kenntn. einig. Tremat., Jen. Zeitschr. f. Naturw., 1903, xxxviii, p. 573. MONTICELLI, F. S. Stud, tremat. Endopar., I, Zool. Jahrb., 1893, Suppl. iii. ROEWER, C. F. Beitr. z. Histogenese v. Cercariaeum helicis, Jen. Zeitschr. f. Naturw., 1906, xli, p. 185. ScHUBMANN, W. Eibildung u. Embryonalentw. v. Fasciola he-patica^ Zool. Jahrb. Anat., 1905, xxi, p. 571. ZlEGLER, H. E, Das Ectoderm d. Plathelminthen, Verh, D. zool. Ges., 1905, p. 35. Watsonius watsoni (pp. 234, 235). CONYNGHAM, H. F. A New Trematode of Man, Brit. Med. Journ., 1904, ii, p. 663; Lancet, 1904, ii, p. 464. Shipley, A. E. Cladorchis watsoni (Conyngham), a Human Parasite from . Africa, Thompson, Yates and Johnston Lab. Report, Liverpool, 1905, vi, I, p. 129. Gastrodiscus hominis (pp. 236, 237). Giles, G. M. A Report of an Investigation into the Causes of the Disease known in Assam as K^la-azar and Beriberi, Shillong, 1890, p. 125. Leuckart, R. Die Paras, d. Mensch., 2. Aufl., ii, p. 450, where the first dis- covery is reported in greater detail. Lewis, T. R., and McConnel. A New Parasite Affecting Man, Proc. Asiatic Soc, Bengal, 1876, p. 182. Fasciola he-patica (pp. 237 to 244, and 638). Ammon. Klin. Darst. d. Krankh. d. menschl. Auges, Dresden, 1838. BOSSUAT, E. Les helminth, dans le foie, Arch, de Paras., 1902, vi, p. 186. [The author is in error when he writes '* The name Dist. sibiricum originated from M. Braun " !] COE, W. R. Bau des Embryos v. Dist. hef., Zool. Jahrb. Anat., 1896, ix, p. 561. DUFFEK, E. Dist. hef. beim Mensch., Wien. klin. Wochenschr., 1902, p. 772. Gaide, cf. under Clonorchis sinensis (p. 787). Gescheidt And Ammon. Die Entoz. d. Auges, Zeitschr. f. Ophth., 1833, iii, P- 405. Greeff, R. trber d. Vork. v. Wurmern im Auge, Arch. f. Augenheilkde., 1907, Ivi, p. 334- Havet, J. Contrib. k I'^tud. d. syst. nerv. d. Tremat., La Cellule, 1900, xvii, p. 351. Henneguy, L. F. Rech. sur la mode de form, de I'ceuf du Dist. hep.. Arch. d'anat. micr., 1906, ix, p. 47. Khouri, a. Le Halzoun, Arch, de Paras., 1904, ix, i, p. 78. KtJCHENMElSTER, F. On Animal and Vegetable Parasites of the Human Body, translated by E. Lankester, London, 1857. Leuckart, R. Z. Entw. d. Lebereg., Arch. f. Naturg., 1882, i, p. 80. LUTZ, A. Lebensgesch. d. Dist. hep., C. f. B. u. P., xi, p. 783; xiii, p. 320. BIBLIOGRAPHY 785 Malherbe. Progr. med., i8q8, vii. No. 4. Marcinowski, K. Das untere Schlundgangl. von. Dist. hef., Jen. Zeitschr. f. Naturw., xxxvii, 1903, p. 544. NORDMANN, A. V. Mikrograph. Beitr. z. Naturgesch. d. wirbellos. Thiere, Berlin, 1832, ii, p. 9. Pallas, P. S. De infestis viventibus intra viventia. Diss, in., Lugd., Batavia, 1760. Saito, S. Beitr. z. Kenntn. d. geogr. Verbr. d. Dist. he-p., C. f. B., P. u. Inf., 1906, i Orig., xli, p. 822. Schaper. Die Leberegelkrankheit. d. Schafe, Deutsche Zeitschr. f. Tiermed., 1890, xvi, p. I. SOMMER, L. Anat. d. Leberegels, Z. f. w. ZooL, 1880, xxxiv, p. 539. Stieda, L. Beitr. z. Anat. d. Plattw. : I, Arch. f. Anat. u. Phys., 1867, p. 52. — tJber d. angebl, inneren Zusammenhang d. mannl. u. weibl. Org. b. Tremat., ibid., 1 87 1, p. 31. Stiles, C. W. Frogs, Toads and Carp as Eradicators of Fluke Diseases, Ann. Rep. Bur. of Anim. Ind., 1901, Wash., 1902, xviii, p. 220. Thomas, P. The Life-history of the Liver Fluke, Quart. Journ. Micros. Sci., 1883, xxiii, p. 99. \_N.B. — A bibliography of cases has been compiled by Davaine (1877), Leuckart (1889-1894), Moniez (1896), Blanchard (1889), and Huber (1895), in addition to Khouri (1. c.).] Fasciola gigantica (pp. 244, 245). COBBOLD, Th. Sp. Description of a New Trematode Worm {Fasciola gigantica)^ Edin. New Phil. Journ., 1855, N.S. ii, p. 262. — Entozoa, an Introduction to the Study of Helminthes, London, 1864, pi. i. Gouvea, H. de. La distomatose pulm. par la douve du foie, Th^se, Paris, 1895. LOOSS, A. Rech. sur la faune de PEgypte, Mem. Inst, egypt., 1896, iii, p. 2>2>' — Obs.«a prop, d'une note . . . C. f. B., P. u. Inf. (i), 1898, xxiii, p. 459. Railliet, a. Sur une forme partic. de douve h^pat. prov. de Senegal, C. R. Soc. Biol., Paris, 1895, loe ser., ii, p. 338. Fasciolofsis buski (pp. 245, 246, and 638). BUDD, G. On Diseases of the Liver, London, 1852. CoBBOLD, T. Sp. On the Supposed Rarity of . . . Dist. crassunij Journ. Linn. Soc, 1875, xii, p. 285; Obs. on the Large Human Fluke, Veter- inarian, 1876. Giles, G. M., cf. under Gastrodisciis hominis (p. 784). Lankester, E. Manual of Animal and Vegetable Parasites (Kiichenmeister), London, 1857, App. i, B. p. 437. Leidy, J. On Distomum he-patictim, Proc. Acad. Nat. Sci., Philadelphia, 1873, P- 364. Odhner, Th. Fasciolofsis buski , C. f. B., P. u. Inf., i Orig., xxxi, p. 573. Fasciolo-psis rathouisi (pp. 246, 247). PoiRlER, P. Note sur une nouv. esp. de Dist. paras, de I'homme, Arch. Zool. exp., 1887 (2), V, p. 203. 786 THE ANIMAL PARASITES OF MAN Paragoninms ringeri (pp. 249 to 251, 630 and 640). Baelz, E. trber paras. 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Notes on Parasites, No. 26; Dist. (Mesogon.) westermannij Discovery of a Parasite of Man, new to the United States, Vet. Journ., 1894, p. 107. Stiles, C. W., and A. Hassall. Notes on Parasites, No. 50 : A Muscle Fluke in American Swine, XVI Ann. Rep., Bur. of Anim. Industry (1899), Wash., 1900, p. 559. — No. 51, The Lung Fluke in Swine, ibid.^ p. 560. Taniguchi. Ein Fall von Distomum-Y.xlirdin'kving des Gehirns mit dem Sym- ptomenkomplex von Jacksonscher Epilepsie, Arch. f. Psych, u. Nerven- heilk., 1904, xxxviii, No. i. Ward, H. B. Dist. westerm. in den Vereinigten Staaten, C. f. B. u. P., 1894, xiv, p. 362; 1895, xvii, p. 304. Yamagiva, K. Lungendistomenkrankh. in Japan, Arch. f. path. Anat., 1892, cxxvii; Zur Atiologie der Jacksonschen Epilepsie, ibid.., 1890, cxix. 0-phisthorchis felineiis and Metorchis truncatus (pp. 252 to 255, 261 and 262). ASKANAZY, M. Uber Inf. d. Mensch. mit Dist. felin. in Ostpreussen u. ihren Zusammenhang mit Leberkrebs, C. f. B., P. u. Inf. 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[An almost complete collection of the literature relating to Cestodes up to 1895 is to be found in Braun's monograph on the tapeworms in Bronn's '* Klassen und Ordnung des Thierreiches," iv, p. 2 ; of later books the follow- ing may be mentioned.] Bartels, E. Cysticercus fasciolaris, Anat., Beitr. zur Entw. und Umwandl. in Taenia crassicollis^ Zool. Jahrb. Anat., 1902, xvi, p. 511. Blanchard, R. Sur quelq. Cest. monstr., Progr. med., 1894 (2), xx. Blochmann, F. Die Epithelfrage bei Cestoden und Tremat., Hambg., 1896. Boas, J. E. V. Triplotaenia mirabiliSj Zool. Jahrb., 1902, Syst. xvii, p. 329. Brandes, G. Teratol. Cestoden, Ztschr. f. d. ges. Nat., Halle, 1899, p. 105. Bugge, G. Zur Kenntn. der Excretionsgefass-Syst. der Cestoden und Tremat., Zool. Jahrb. Anat., 1902, xvi, p. 177. 79C THE ANIMAL PARASITES OF MAN ChilDj C. M. Abnormality in Moniezia ex-pansa, Biol. Bull. Woods HolL, 1902, iii, pp. 95, 143. CoHN, L. Zur Anat. imd Syst. der Vogelcestoden, Nov. Act. Acad. Caes. Leop.- Carol. Nat. Cur., Halle, 1901, Ixxix, No. 3. Drago, U. Azione sperim. dei succhi diger. sulT involucro della ova di ale. Tenie, Arch, de Paras., 1906, x, p. 321. FUHRMANN, O. Ein getrenntgeschlechtlicher Cestode, Zool. Jahrb., 1904, Syst. XX, p. 131. Grohmann, W. Die Abnormitaten in den Progl. der Cestoden, Inaug. -Dissert., Giessen, 1906. KUNSEMULLER, F. Zur Kenntnis der polycephalen Blasenwiirmer, insbesondere des Coeniirus cerebralis Rud. und des C. serialis Gerv., Zool. Jahrb. Anat., 1903, xviii, p. 507. Lt)HE, M. Zur Anat. und Syst. der Bothrioceph., Verhandl. der Deutsch. zool. Ges., 1899, P- 30. — Review of Braun's Bothr.-Syst., C. f. B., P. u. Inf., 1902, i Orig., xxi, p. 318. Messineo, G. Sul veleno conten. in alcune Tenie delP uomo, Atti Accad. Gioenia sci. nat., Catania, 1901 (4), xiv, No. 6. MiNGAZZiNl, P. Sul vario modo di fissaz. delle Tenie alia parete intest.. Rich. Labor, anat., Roma, 1904, x, p. 5. ROSSLER, P. tJber den fein. Bau der Cysticerken, Zool. Jahrb. Anat., 1902, xvi, p. 423. Saint-Remy, G. Develop, embr. Taenia serrata, Arch, de Paras., 1901, iv, P- 143. Schaaf, H. Zur Kenntn, der Kopfanlage der Cysticerken, insbes. des Cysti- cercus der Taenia solium L., Zool. Jahrb. Anat., 1906, xxii, p. 435. Spengel, J. W. Die Monozootie der Cestoden, Ztschr. f. w. Zool., 1905, Ixxxii, p. 252. Stiles, Ch. W. Revision of Ad. Tapeworm of Hares and Rabbits, Proc. U.S. Nat. Mus., 1896, xix. Stiles, C. W., and A. Hassall. Tapeworms of Poultry, U.S. Dept of Agric, Bur. of Anim. Ind., 1896, Bull. 12. ViGENER, J. tiber dreikant. Bandwiirmer a. d. Fam. d. Taeniiden, Jahrb. nass. Ver. f. Naturke., Wiesb., 1903, p. 115. Zernecke, F. Unters. iiber d. fein. Bau d. Cestod., Inaug.-Diss., Rostock, 1895. Dihothrioce-phalus latiis (pp. 310 to 315, and 658). {a) Anatomy. BOttcher, a. Studien uber den Bau des Bothr. latus, Arch. f. path. Anat., 1864, xxx, p. 97; 1869, xlvii, p. 370. ESCHRICHT, D. F. Anat.-phys. Untersuchung. iiber die Bothrioceph., Nov. Act. Ac. 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The Influence of Temperature upon the Biting of Mosquitoes, Parasitology, 1910, iii, No. 4, p. 478. James, S. P. The Protection of India from Yellow Fever, Indian Journ. of Med. Res., 1913, i, No. 2, pp. 213-217. — Note on the Practicability of Stegomyia Reduction in Indian Seaports, Ind. Journ. Med. Res., 1913, i, No. 2, pp. 258-262. BIBLIOGRAPHY 829 James, S. P., and \V. G. Liston. A Monograph of the Anopheles Mosquitoes of India, Calcutta, 1904, 4to, 30 pi. ; 2nd ed., iqii. King, H. H. Animals Injurious to Man and Animals, Mosquitoes, Fourth Rept. Well. Lab. Trop. Res., iqii, pp. 99-112. — On the Bionomics of the Sandflies of Tokar, Bull. Ent. Res., 1913, iv, pt. i, p. 83. Lahille, F. Sobre un Anopheles, una Stegomyia y la Notation, de las Xerva- duras alares de los Mosquitos, Ann. d. Mus. Nagion. d. Hist, Nat. de Buenos Aires, 1912, xxiii, p. 253-263. Leicester, G. F. ■ The Culicidae of Malaya, Studies from the Inst, of Med. Res., Federated Malay States, 1908, iii. Leon, N. Vorl. Mitt. Uber den Saugriissel der Anopheliden, Zool. Anz., 1Q04, xxvii, p. 730. LiSTON, W. G., and T. G. Akula. A Stegomyia Survey of the City and Island of Bombay, Proc. Gen. Mah Comm., Madras, November, 1912, Simla, 1913, p. 187. Ludlow, C. S. Disease-bearing Mosquitoes of North and Central America, the West Indies, and the Philippine Islands, Bull 4, War Dept., Office Surgeon-General, U.S.A., 1914. — The Philippine Mosquitoes, Psyche, iqii, xviii. No. 4, pp. 125-133. LUTZ, A. Brazilian Simulidae, Mem. d. Inst. Oswaldo Cruz, 1910. — Contribuigao para o estudo das C erato-pogoninas heniatofagas encontradas Mem. Inst. Oswaldo Cruz, 1913, v, pp. 45-72. — Contribuigao para o estudo das C erato-pogoninas heniatofagas encontradas no Brazil, Mem. Inst. Oswaldo Cruz, 1912, pp. 1-33. — Contribuigao para o conhecimento das Ceratopogonina^ do Brazil, Mem. Inst. Oswaldo Cruz, 1914, v, pp. 8i-q8. — Segunda contribuigao para o conhecimento das especies brazileiras do genus Siniidiuin, Mem. do. Inst. Oswaldo Cruz, 1910, ii, p. 213. LuTZ, A., and A. Neiva. Contribuigoes para o biologia des Megarininas, Mem. Inst, Oswaldo Cruz, 1913, v, pp. 129-141. — Contribuigoes para o estudo das Megarhininse, Mem. Inst. Oswaldo Cruz, IQ14, pp. 50-56. — Contribuigao para o conhecimento das especies do genero Phlehotomiis existentes no Brazil, Mem, Inst. Oswaldo Cruz, 1912, iv, pp. 84-95. MacFie, J. W^. S. A Note on the Action of Common Salt on the Larvae of Stegomyia fasciata, Bull. Ent. Res., 1914, iv, pt. 4, pp. 33Q-344- Malloch, J. R. American Black Flies or Buffalo Gnats, U.S. Dept. Agri., Bur. Ent. Tech. Sci., 1914., No, 26. Mansion, J. Les Phlebotomes en Corse, Bull. Soc. Path. Exot,, Paris, 1913, vi. No. 9, pp. 637-641, Mhaskar, K. S. Stegomyia in Karachi, Proc. Gen. Malaria Com., Madras, 1912, Simla, 1913- Mitchell, E, G. Mosquito Life, New York and London, 1907. Newstead, R. Papataci Flies {Phlehotomiis) of the Maltese Islands, Bull. Ent. Res., 191 1, ii, pt. i. — The Papataci Flies [Phlebotomus) of the Maltese Islands, Bull. Ent. Res., IQ11-12, ii, p. 47- — Notes on Phlehotomiis with descriptions of New Species, Bull. Ent. Res,, 1912, iii, p. 361. — Notes on Phlehotomiis with descriptions of New Species, Bull. Ent. Res., 1914, v, pt. 2, pp. 129-136, 83a THE ANIMAL PARASITES OF MAN Newstead, R., andH. F. Carter. On Some New Species of African Mosquitoes [Culicidae], Ann. Trop. Med. and Par., vii, p. 233. Description of a New Genus and three New Species of Anopheline Mos- quito, Ann. Trop. Med. and Par., 1910, iv, p. ^yy. — — On a New Genus of Culicinx from the Amazon Region, Ann. Trop. Med. and Par., 1910, iv, p. 553. Newstead, R., and H. W. Thomas. The Mosquitoes of the Amazon Region, Ann. Trop. Med. and Par., 1910, iv, p. 141. NUTTALL and Shipley. Structure and Biology of Anoi>heleSj Journ. of Hygiene, January, April, and October, iqoi, and January and April, 1903. Studies in Relation to Malaria; II, Structure and Biology of Ano-pheleSy Journ. of Hyg., 1901, i; 1902, ii ; 1903, iii. PeryasSU, a. Os Culicideos do Brazil, Trabalho do Institute de Manguinhos, 1908. Ross, E. H. The Reduction of Domestic Mosquitoes, London, John Murray. RoUBAUD, E. Quelques mots sur les Phlebotomes de I'Afrique occidentale frangaise, Bull. Soc. Path. Exot., I9i3j vi, pp. 126-128. SCHINER, J. R. Fauna austriaca. Die Fliegen (Diptera), Wien, 1860-64. Schwetz, J. Preliminary Notes on the Mosquitoes of Kabinda, Belgian Congo, Ann. Trop. Med. and Par., 1914, viii, p. 163. Stanton, A. T. The Anopheles of Malaya, Bull. Ent. Res., 1913, pt. i, iv, pp. 129-133. — The Anopheles Mosquitoes of Malaya and their Larvae, with some Notes on Malaria-carrying Species, Journ. London School Trop. Med., 1912, ii, pt. I, pp. 3-II- Summers, S. L. M. A New Species of Phlebotomus from South America, Bull. Ent. Res., 1912, iii, p. 209. Taylor, F. H. The Culicida^ of Australia, Trans. Ent. Soc. Lond., March 31, 1914, pp. 683-708. — A Revision of the Culicidae in the Macleay Museum, Sydney, Proc. Linn. Soc, N.S. Wales, 1913, xxxviii, pt. 4, pp. 747-760. — Report of Entomologist, Reprint from Report for Year iqii of the Australian Inst. Trop. Med., 1913, pts. xiii, xiv, pp. 24. — Description of Mosquitoes collected in the Northern Territory during the Expedition, 191 1, Rept. on Health and Disease in the Northern Territory, Bull. No. lA, 1012, p. 25. — Culicidse of Papua, Trans. Ent. Soc. Lond., 1914, pt. i, pp. 185-205. Theobald, F. V. A Monograph of the Mosquitoes of the World, 1901-1911, vols, i-v, and atlas and plates. — New Culicidae from the Sudan, Ann. Trop. Med. and Par., 1913, vii, p. 591. — A New Genus and Two, New Species of Culicidae from the Sudan, Fourth Rept. Wellcome Trop. Res. Lab., 191 1, Vol. B, Gen. Science, pp. 151- 156. — Three New Culicidae from the Transvaal, Entomol., March, 1912. — A New Mosquito from North China, Entomol., June, 1913, p. 179. — A New Mosquito from Samoa, Entomol., January, 1914, p. 36. — Culicidae of the R. Zool. Soc, '' Natura Artis Magistra," Amsterdam Overgt. uit het Tijdsch. v. Ent., 191 1, liv, pp. 233-240. — The Distribution of the Yellow Fever Mosquito [Stegomyia fasciata^ Fabri- cius) and general notes on its Bionomics, First Int. Cong. d'Ent., 191 1, pp. 145-170. BIBLIOGRAPHY 83 1 Theobald, F. V. A New Species of Culicidse, Rev. Zool. africaine, 1912, ii, fas. 1. — Novae Culicidae, 191 1, pt. i, pp. 35. — Culicidae (Percy Sladen Trust Exp. to Indian Ocean in 1905), Trans. Linn. Soc. Lond., 1912 (2nd ser., Zool.), xv, pt. i, pp. 81-94. — Culicidae from New Caledonia and the Loyalty Islands, Nova Cal., Zool., 1913, i-iii, No. 3, p. 163. — A New Genus and Two New Species of Culicidae from the Sudan, Fourth Report Wellcome Trop. Lab., igii, Vol. B, Gen. Sci., pp. 151-156. TOWNSEND, C. H. T. A Phlehotomtis the practically certain Carrier of Verruga, Science, 1913, xxxviii, pp. 194-195. — The Vector of Verruga, Phlehotonms verrucarum, sp. nov., Insecuta Inscitiae Menstruus, Washington, D.C., 1913, i, No. 9, pp. 107-109. Urich, F. W. Mosquitoes of Trinidad, Proc. Agri. Soc. Trini. and Tobago, 1913, xiii. No. 10, pp. 525-530. WiLLCOCKS, F. C A Preliminary Note on the Prevalence of Mosquitoes in Cairo and its Environs, Ann. Trop. Med. and Par., 1909, v, p. 583. The House-fly (p. 586). Bayon, H. Leprosy and House-flies, Ann. Trop. Med. and Par., 191 5, ix, pp. 1-90. BiSHOPP, DORE, and Parman. Notes on Certain Points of the Economic Importance in the Biology of the House-fly, Journ. Eco. Ent., 1914, viii, pp. 54-71. Felt, E. P. Methods of Controlling the House-fly and thu-s Prek-enting the Dissemination of Disease, New York Med. Journ., April 2, 1910. Hewitt, C. G. The Biology of the House-fly in relation to Public Health, Journ. Roy. Inst. Pub. Health, October, 1908. — On the Bionomics of certain Calyptrate Muscidae and their Economic Signi- ficance, with especial reference to Flies inhabiting Houses, Journ. Eco. Biol., 1907, ii, p. 3- — The Structure, Development, and Bionomics of the House-fly, Miisca domestica, II and III, Bionomics, &c., and the Relations to Human Disease, Quart. Journ. Micros. Sci., 1908, and December, 1909, pt. 3, liv. — A Preliminary Account of the Life-history of the Common House-fly [Mtisca do7nestica, L.), Mems. and Proc. Manchester Lit. and Phil. Soc, 1906- 1907, pt. I, Ii. — House-flies and the Public Health, Ottawa Naturalist, May, 1910, xxir, pp. 31-38. — Notes on the Pupation of the House-fly [Musca don^estica) and its mode of Over-wintering, Can. Ent., March, 191 5, xlvii, pp. 73-78. HiNDLE, E. The Flight of the House-fly, Proc. Camb. Phil. Soc, 1914, xvii, pt. 4, pp. 310-313. House-flies, U.S. Dept. Agri., Div. Ent., Circ. No. 71, 1906. Howard, L. O. The House-fly, Disease Carrier, New York, 191 1. Hutchinson, R. H. The Migratory Habit of the House-fly Larvae as indicat ing a Favourable Remedial Measure, Bull. U.S. Dept. Agri., 1914, No. 14. Jepson, F. p. The Breeding of the Common House-fly {Musca domestica) during the Winter Months, Journ. Eco. Biol., 1909, iv, pt. 3. MORRELL, A. W. House-fly Baits and Poisons, Journ. Eco. Ent., 1914, vii. No. 3, pp. 268-273. ^^2 THH ANIMAL PARASITES OF MAN Nevvstead, R. Preliminary Report on the Habits, Life Cycle, and Breeding- places of the Common House-fly {Miisca domestica^ L.) as observed in the City of Liverpool, &c., Health Committee Rept., City Liverpool^ 1907. — - Report on the Habits, Life Cycle, and Breeding-places of the Common House-fly, Liverpool, IQ07, Ann. Trop. Med. and Par., February, iqo8. Pack.ard, a. S. On the Transformation of the Common House-fly, with Notes on Allied Forms, Proc. Boston Soc. Nat. Hist., 1874, xvi, p. 136. Reports of the Local Government Board on Public Healthy &c. New Series, Nos. 5 and 16, 1909. Preliminary Reports on Flies as Carriers of Infec- tion, and further Preliminary Reports (Paper by Copeman, Jepson, Nuttall, Graham Smith, and Austen), containing Bibliography and Abstracts up to date. SURF.ACE, H. A. To keep down House-flies, Zool. Press Bull., Div. ZooL, Dept. Agric, Pennsylvania, 1915, No. 313. Recommends ground phos- phate rock, scattered over manure heaps. Brachycera, &c. (pp. 600 to 612. and 613 to 615). Ashlev-Emile, L. E. Zambesi ulcer, Journ. Trop. IMed., 1905; Arch. f. Schiffs.- u. Tropen-Hyg., 1906, x, p. 164. Austen, E. E. African Blood-sucking Flies other than Mosciuitoes and Tsetse- flies, 1909. — A Monograph of the Tsetse-flies, 1903. — • Supplementary Notes on Tsetse-flies, Brit. Med. Journ., 1904. — A Handbook of the Tsetse-flies, iSqi. — Some Dipterous Insects which cause Myiasis in Man, Trans. Soc. Trop. Med. and Hyg., March, 1910, p. 215. — New African Tabanidae, Bull. Ent. Res., 19 14, iv, pt. 4, pp. 283-300. Bezzi, M. Etudes systematiques sur les Muscides hematophages du genre Lyperosia, Archiv. Parasit., 191 1, xv, pp. 1 10-143. — Die Gattungen der blutsaugenden Musciden, Zeitsch. f. Hymenoj). u. Dipt., 1907, vii, p. 413. Bl.anch.ard, C. Contrib. a I'Etude des Dipteres paras., .4nn. Soc. Ent. France, 1895, Ixv, p. 641. Carter, H. F. Descriptions of Three New African Species of the genus Tabanus, Ann. Trop. Med. and Par., 1914, vi, p. 435. Chevrel, R. Sur le Myiase des Voies urinaires, Arch, de Paras., 1909, xii, p. 369. Cluss, F. Myiasis interna and externa, Inaug.-Diss., Tubingen, 1902. Cockerill, J. D. A. A Dcnnatobia in Guatemala, Entomologist, 1914, xlvii, p. 131. Davy, J. B. Notes on the Habits of Glossina fiisca, Bull. Ent. Res., 1910-1911, i, P- 143- DONITZ. Fine neue afrik. F liege, Cordylobia muriitm, Sitz. Ges. naturf. Frde. Berl., 1905, p. 248. Dubreuilh, W. Les Dipteres cutic. chez I'Homme, Arch. Med. exp. et d'Anat. path., 1894, vi, p. 328. Enderlein, G. Die Respirationsorgane der Gastriden, Sitz. k. Akad. d. Wiss., Wien, Math-nat., 1899, Kl. cviii. Fell, T. E. Notes on Tsetse-flies and on Prophylactic Measures against Sleeping Sickness, &c.. Bull. Ent. Res., 1912, iii, p. 227. Fiske, W. F. The Bionomics of Glossina, Bull. Ent. Res., 1913, iv, pt. 2, p. 95- BIBLIOGRAPHY 833 Francaviglia, M. C. An cora suUa myiasi auricolare, Boll. Sedute Accad. G.ioenia, Catania, 1914, No. 31, pp. 15-23. Fuller, C. The Skin-maggot of Man [Cordylobia anthro-po-phaga)^ Agri. Journ. Union of South Africa, Pretoria, 1914, vii. No. 6, pp. 866-874. Gedoelst, L. Contrib. a I'Etude des larves cuticoles des muscides africaines. Arch, de Paras., 1905, ix, p. 568. Giles, G. M. The A^natomy of the Biting Flies of the genus Stomoxys and Glossina, Journ. Trop. Med., 1906, ix. Grunberg. Ueber Blutsaugende Musciden, Zool. Anz., 1906, xxx, p. 78. Grunberg, K. Afrik. Musciden mit parasit. lebenden Larven, Sitz. Ges. Nat. Frde. Berl., 1903, p. 400. — - Die Blutsaugenden Dipteren. — Ein neuer Fall des Vorkommens. der Larve der Rinderdasselfiiege im menschlichen Auge, Sitz. Ges. Nat. Frde. Berl., 1913, 5 and 6, pp. 298- 304- Hewitt, C. G. Observations on the Feeding Habits of the Stable Fly, Stomoxys calcitrans, L., Trans. Roy. Soc. Canada, 1914, viii, pp. 37-42. HUBER, G., and F. L. Flack. An Unusual Case of Screw-worms in the Nose and Nasal Accessory Sinuses, Journ. Amer. Assoc. Chicago, 1914, Ixiii, No. 26, p. 228. Joseph, G. Uber Fliegen als Schadlinge und Paras, d. Menschen., Dtsche. Med. Ztg., 1885, i, p. ?,1, and 1887, iii, pp. 713-725. Jack, R. W. Observations on the Breeding Haunts of Glossina morsitans. Bull. Ent. Res., 1911-1912, ii, p. 357. King, H. H. Blood-sucking Flies other than Mosquitoes, Fourth Rept. Well. Trop. Res. Lab., 1911, pp. 112-126. — Observations on the Occurrence of Glossina in the Mongolia Province of the Anglo-Egyptian Sudan, Bull. Ent. Res., 1912, iii, p. 89. Kingham, a. Notes on the Preliminary Stages of Glossina morsitans. Bull. Ent. Res., 1911-1912, ii, p. 291.. Le Dantec and Boye. Note sur une myiase observee chez I'homme en Quince frang. (Reun. biol. de Bordeaux), Le Caducee, 1905, v, p. 9; Arch. f. Schiffs- u. Tropen-Hyg., 1906, x, p. 71. Lloyd, L. Notes on Glossina morsitans in the Luangwa Valley, N. Rhodesia, Bull. Ent. Res., 1912, iii, p. 233. Low, F. Uber Myiasis und ihre Erzeuger, Wien. med. Wchschr., 1882, xxii, p. 247, and xxiii, p. 972. LoWNE, B. T. Physiology, Morphology, and Development of the Blow-fly, two vols. LuTZ, A. Tabanidas do Brazil e de alguno Estados visinhos, Mem. Inst. Osw^aldo Cruz, 1913, v, pp. 143-190. — Novas contribuigoes para o con hecmento das Pangoninas e Chrysopinas do Brazil, Mem. Inst. Oswaldo Cruz, 191 1, iii, fas. i, pp. 65-84. LUTZ, A., and A. Neiva. Los Tabanidae do Estado do Rio de Janeiro, Mem. Inst. Oswaldo Cruz, 1914, vi, pp. 69-80. McConnell, R. E. Some Observations on the Larva of Auchmeromyia luteola, Fabr., Bull. Ent. Res., 1913, iv, pt. i, p. 29. — Notes on the Occurrence and Habits of Glossina fusci-pes in Uganda, Bull. Ent. Res., 1912, iii, p. 55. MacFie, J. W. S. Experiments and Observations upon Glossina palfalis, Bull. Ent. Res., 1912, iii, p. 61. Magalhaes, p. de. Subsidio ao estudo das Myiases Rio de Janeiro, 1892. Meigen, J. W. Systemat. Beschr. d. bek. Europ. zweifliigel. Insecten, 1818- i8xpulsion of guinea-worm, 676 Chloroform injections or inhalations in nasal myiasis, 720 Chloromyxiace, 184 Chloromyxum leydigi, trophozoite of, 182 Choanoflagellata, 52 Cholera Investigation Commission, work of, with regard to dysentery, 30 — motions, spirochaetes found in, 122 — spread by house-fly, 586 Cholodkowsky, " wormlet " burrowing into human epidermis, 599 Chorea, case of, cured after expulsion of Taenia, 648 Chorion enveloping ova of nematodes, 371 Chorioptes bovis, 521 — characters, 517 — species found on man, 521 Christophers, on Babesia cants, 176, 177 — on Lencocytogregarina canis, 155 — see Stephens and Christophers Christopherson, J. B., case of espundia, 108, 628 Chrithoptes nionunguiculosiis, 489 Chromidial apparatus of protozoa, 26 Chrysoconops, 577 Chrysomela hoemoptera, gregarine from, 131 Chrysomyia and Pycnosoma, distinguishing features, 588 — (Cotnpsomyia) macellaria (screw-worm fly), 587 larvae of, invasion by, fatal results from, 587 — regions of body invaded by, 587 references to, 587 synonyms, 587 — viridula, characters of, 588 larvae (maggots) of, discharged from nose, 588 Chrysops host of Filaria loa, 601 — dimidiata, 601 — silacea, 601 — transmission of surra by, 601 Chrystia, characters, 562, 568 Chyluria, association of Hymenolepis mada- gascarensis with, 662 — following infection by Eustrongylus gigas, 683 by Hymenolepis nana, 661 — from Filaria bancrofti infection, 677 treatment, 677 — in filariasis, 402 — without lymphatic obstruction, 401 Cigarettes, paper, smoking of, in nasal myiasis, 719 Ciliata, 198 — classification, 199 — macro-nucleus and micro-nucleus of, 198 — morphology of, 198 — peristome of, 198 — reproduction of, 198 Ciliophora, 198 Cimex (Acanthia) lectularia (bed bug), see Cimex lectularius (following) — honeti, host of Trypanosoma criizi, 87 — ciliatus, 537 INDEX 847 Cimex columbarius , 536 bite of, 536 — hirundinis (swallow bug), 537 — lectularius (bed hug), artificial host of Trypanosoma criizi, 87 bite of, treatment, 713 ■ — blood-sucking, 535 — — characters and habits 534 extermination of, 713 infection by, 713 diagnosis, 713 larval stage, 535 ova of, 535 peculiar odour emitted by, 535 — — persistent accompaniment of man by, 535 possible transmission of kala- azar by, 107, 713 transmission of SpirochcBta re- currentis by, 120, 121 — macrocephalus , 536 — rotundatus (tropical bed bug), 535, 536 carrier of virus of polio- myelitis, 536 geographical distribution, 536 points of distinction from C. lectularius , 536 — possible connection with kala-azar, 536 — sp., connection with Oriental sore, 536 CimicidcB, characters, 534 Circulatory system, disturbances of, in ancylostomiasis, 683 Cisterns, screening against mosquitoes, 636 Cladorchiina>, 231, 234 — male and female organs, 234 — morphology, 234 Clark, see Howard and Clark Clasping and clinging organs in permanent parasites, 4 Climates, temperate, prevalence of Ascaris lumbricoides in young children in, 464 Clonorchis endemicus, geographical distri- bution, 260 habitat and hosts of, 259 intermediate host of, first unknown, 261 second, 261 life-history, 261 — — morphology, 259 synonyms, 259 — morphology, 258 — sinensis, geographical distribution, 259 — — habitat and hosts of, 259 infection by, 640 diagnosis, 641 prophylaxis against, 641 — — — syniTotoms, 640, 641 treatment, symptomatic, 641 morphology, 258, 259 organs of, diagram showing, 259 ova of, 259 sites of invasion in body, 640 Cloquet, destruction of eyes by Sarcophaga larvae, 723 Clothes louse, see Pediculus vestimenti Cnidosporidia, 129, 194 Cobbold, Ligula niansoni, 318 Cocaine, application, followed by calomel insufflations, in nasal myiasis, 720 — hypodermic injection in creeping disease, 731 Coccidce (scale insects), 532 Coccidia, copulation in, 137 — experimental infection with, 136 — ova of helminthes mistaken for (foot- note), 137 — pathogenicity of, 136 Coccidia-like organisms in various diseases of man, 150 Coccidiidea, 129, 135, 137 — characters and habitat, 28 — classification of, 141 — gametes of, 137, 139, 140. — hosts of, 137 — life-cycle of, 138-141 — macrogametes, 139, 140 — merozoites of, 138, 139, 140 — microgametes of, 137, 139, 140 — morphology of, 138 — occurrence, 137 — oocysts of, 141 — schizogony in, 138 — sporoblasts and sporocyst of, 141 — sporozoites of, 138, 139, 140 Coccidioides imniitis, 150 — pyogenes, 150 Coccidiomorpha, 129, 151 Coccidiosis. avian, 142-145 — diagnosis of, 742 — human, doubtful cases, 149 hepatic cases, 148 intestinal cases, 148 — in cattle, 147, 741 — in rabbits, intestinal and hepatic, 145, 147 — rinderpest mistaken for, 741 Coccidium jalinum, 150 hosts of, 150 Cochin-China diarrhoea, see Diarrhoea (Cochin-China) Cockchafer, intermediate host of Echino- rhynchus qigas, 477 Cocoons of Hirudinea, 481 Cod-liver oil, inunctions of, in evacuation of Oxyuridm, 698 Coelosporidiuni, 195 Coenurus, definition of, 301 — cerebralis, experimental rearing of tape- worms from, 15 reared experimentally, 15 . section showing cephalic invagina- tions, 304 — scolices in, 303 Coleoptera, characters, 531, 532 — larvae of, accidental parasites, 542 Colitis associated with B alantidiiim coli, 201 with intestinal amoebae, 30 — mucous, complicating intestinal myiasis, 726, 727 Collargol in balantidian dysentery, 637 848 THE ANIMAL PARASITES OF MAN Collargol, irrigation of lower bowel with, in gangrenous dysentery, 619 — rectal administration in billiarziasis, 643 Colon, cysts of, (Esophagostoniiim contained in, 441, 443, 444 — descending, means of access of Schisto- stoma hceynatobium to, 272 Colorada, mite attacking man, 486 Colpoda cucullus, 204 Comedones, removal in infection by Demo- dex folliculorum , 708 Commensals, 6, 20 — nature of, 6 Congo floor maggot, 593, 594 Conjunctiva, dipterous larvae in, 716 — trachoma bodies in infected epithelial cells of, 209 Conjunctivitis due to head louse infection, 710 Connective tissue, subcutaneous invasion by Loa loa, 678 Conorhinus megistus intermediate host of Trypanosoma cruzi, 83, 537 see Triatoma megista — nigrovarius , bite of, 539 — protractus , 539 — renggeri (great black bug of Pampas), blood-sucking, 539 - — rubrofasciatus, bite of, 538 trypanosome inhabiting, 99 — sanguisuga (blood-sucking cone nose), bite of, 537, 538 ova of, 538 — sp. (?), bite of, symptoms following, 538 — variegatus, 539 Conseil, mode of transmission of relapsing fever, 120 Constipation, prevention during malarial attacks, 635 — set up by Ascarides, 657 Copaiva balsam in bilharziasis, 643 Copper, black oxide of, as tapeworm driig, 674 Copra itch, mite causing, 513 — — treatment. 513 Copulation, modes of, in tapeworms, 297 Cordylobia anthropophaga, characters, 592 geographical distribution, 593 hosts of, 592 larvae of, characters, 592 lesions set up by invasion, 592 life-history, 592 references to, 593 — grunbergi, larvae, characters, 591, 592 synonyms, 591 — rodhani, 593 geographical distribution, 593 Coreotrypanosis, 87 Corethra, 565 Corethrina, 565 Coriscus subcoleoptratus, bite of, 540 geographical distribution, 540 synonyms, 540 Cornea, parasitic crustacean (Caligiis cur- tus) invading, 483 Cortical layer of Cestoda, 289 Councilman and Lafieur on nomenclature of amoebae, 31 Couret and Walker, J., culture medium for intestinal amoebae, 743, 744 Couvy, Spirochceta gallinarum, 119 — see Marchoux and Couvy Cover-slip preparations, 748 Cows attacked by Leptus autuninalis, 486 Crab louse, see Phthirius inguinalis Craig, C. F., on Entarnceba histolytica, 37, 41 on Paramoeba (Craigla) hominis, 44, 45 Craigia hominis, 45, 734 — migrans, 734 Craigiasis, 734 — nature of, 734 Crane-fly, Gregarina longa from larva of, 130 • Craw-craw (filaria infection of the skin), 378, 514 Crawley, experimental infection with Sarco- cystis, 192 — movements of gregarines, 131 Creeping disease, cases of, various authors reporting, 729 — — clinical symptoms, 730 — — duration, 731 localization of, 730 mode of origin, 729, 730 synonyms, 729 ■ treatment, 731 methods reported by various authors, 731 — eruptions, 599 Creosote in bronchial spirochaetosis, 633 — mounting agent for flukes, 471 Creplin, discovery of progeny of Diphyllo- botlirium {Bothriocephaliis) ditremnm, 13 — psorosperms, 181 Cristina and Caronia, treatment of infan- tile kala-azar, 627 Crithidia, 67, 103 — inoculation experiments with, 104, 738 — fasciculata, 104 host of, 104 — qerridis, 738 — hosts of, 104 — hyalommm, 104 — melophagia, host of, 104 — morphology, 104 — natural flagellates of Arthropoda, 104 Crustacea, parasitic, change of original fea- tures in, 4 loss of digestive sy^stem in, 3 or free-living, invading man abnor- mally (footnote), 483 Csokor, mode of infection in intestinal myiasis, 727 Ctenocephalus canis, herpetomonad inhabit- ing, 103 transmission of Trypanosoma lewisi by, 88, 90, 92 — distinctive characters, 545 — felis (cat flea), 547 INDEX 849 Ctenocephalus, hosts of, 547 Ctenophthalmus, distinctive characters, 545 Ctenopsylla, 548 — distinctive characters, 545 — musculi, transmission of Trypanosoma lewisi by, 90 Cucumber seeds in intestinal myiasis, 728 Culex and Anopheles, larvae of, position in water compared, 554 — • — ova of, method of depositing com- pared, 554 points of difference between, 551 — characters, 564 — fatigans (common tropical gnat), dis- tinguishing character, 576 transmission of Filarise by, 576 — head of male and female, 549, 556 — human malaria not spread by species of, 158 — larva of, 553 — ova of, 557, 558 — pipiens (common gnat), 575 — — characters, 576 ova of, localities selected for deposi- tion, 553 — species of, development of Plasmodium relicUim in, 170 Ctilicidoe, classification of, 501 — number of species, 552 , — scales of, 560 CulicincB, characters, 563, 571 Culicoides, blooi-sucking habits of, 580 — larvae of, 580 — ornatus, bite of, 581 — possible carrier of germ of Delhi boil, 580 — pupae of, 580 Culture media for amoebae, 742 — — for blood protozoa, 744 Cunningham, discovery of intestinal amoebae, 29 Cutaneous glands, unicellular, of nema- todes, 361 — tumours due to cysticerci, characteristics of, 662, 663 Cyclasterium, 208 Cyelocoelum mutabile, progeny of, dis- covery, 12 Cycloleppteron, characters, 561, 567 Cyclophyllidea, 308 Cyclopidce, characters, 390 Cyclops, characters, 390 — intermediate host of Draciinculus medin- ensis, 388 — virescens, 389 Cyclospora, 141 Cyrtoneura stabulans, larvae of, habitat, 585 Cysticerci, cutaneous and muscular, (sym- ptoms set up by, 663 treatment, 663 — development from oncosphere of Teen iidcB, 303 — early researches on, 282 — experimentally reared from tapeworms, 15 Cysticerci, number of, in relation to species of Taeniae, 16 — origin of, 14 — regions of body site of, 663, 664 — subretinal, 664 — tapeworms experimentally reared from, 15 Cysticercoid, morphology of, diagram show- ing, 301 Cysticercoids, echinococcus-like conditions in, 304 Cysticercus acanthotrias , 336 and C. cellulosoi, 336, 337 — hovis, 340 amount of prevalence in ox, 340, in Prussia and Berlin, 341 , artificial infection of human beings with, 340 rarity in man, 341 — celluloscR, amount of injury inflicted by, depends on situation in body, 8 development to ToBuia solium, 340 infection of skin and subcutaneous tissues, 662 — — and C. acanthotrias, 336, 337 decrease of frequency in pork, how effected, 334 development, time taken for, 334 habitat, 332, 333 — - liosts of, 332 — — how conveyed to man, 334, 335 'u man, 335 I -— long persistence of, 337 i in sheep, 337 organs of body invaded by, 335 sex distribution of invasion by, 335 vitality of, 334 — development of, 301, 302 diagram showing, 303 : — fasciolaris, host of, 338 ' possible means of spread to man, 338 j — forms of, 301 ! — morphology of, diagram showing, 301 ' — avis, 337 1 — papilliform invagination into bladder, 301 — pisiformis, hosts of, 338 in evaginated condition, 304 — ■ racemosus , 335, 336 — tenuicoUis, 337, 338 experimental rearing of tapeworms from, 15 — with developed scolex at bottom of in- vagination, 304 Cystoflagellata, 52 Cysts, intestinal, CEsophagostomum con- tained in, 441, 443, 444 Cytorhyctes, 208 — aphtharum, 208 — cytoplasm of, 209 — luis, 124, 208 — scarlatince, 208 — vaccinice, 208 — variolce, 208 minute granules in, 210 850 THE ANIMAL PARASITES OF MAN Dactylomyia, characters, 562 Daniels, C. W., iridocyclitis in trypanoso- miasis, 623 — — parasitic coleopterous larva (foot- note), 542 treatment of trypanosomiasis, 622 yellow pigment in kidney and liver cells in ancylostomiasis, 647 Danielsia, characters, 564 Danilewsky, discovery of endoglobular para- sites, similar to malarial, in birds, 157 of Leucocytozoa by, 153 Danube, banks of, Simulium columbaschen- sis plague on, 578 Darling, experimental infection with Sarco- cystis muris, 192 — Endotrypanum schaudinni, 99 — Histoplasma capsulatum, 112 — researches on Entamceha tetragena, 38, 40, 41 Darwin, Charles, great black bug of Pam- pas (Conorhinus renggeri), 539 Dauernheim, Ascarides in bile-ducts, 688 Daughter cysts of echiuococcus, 350, 351 mode of origin, 350, 351, 352 Davaine, Cercomonas hominis, 61 — milk cure in expulsion of Strong yloides stercoralis, 675 — mode of development of Ascaris lumbri- coides, 464, 465 Davainea asiatica. 662 morphology, 330 — madagascarensis , association with chyl- uria, 662 cases of human infection by, 330 morphology, 329 — morphology, 329 Davaineidce, 309, 329 Davaineinoi, 309, 329 Davidson, bite of Rasahus biguttatus , 540 Deeks, W. E., treatment of amoebic dysen- tery, 619 Deguy, see Labadie-Lagrave and Deguy Deinocerites, characters, 564 Delanoe, pneumocysts in rats, 90 Delhi boil, see Oriental sore Demarquay, observation of Filaria ban- crofti in man, 390 Demodex folliculorum, 521 affecting eyelids, 708 as cause of chalazion, 708 characters, 522 infection by, treatment, 708 views of different authors respect- ing, 708 synonyms, 522 var. canis, 522 infection by, transmission from dog to man, 709 treatment, 709 Demodicidoe (mites of hair follicles), charac- ters, 522 Dendriomyia, characters, 565 Dengue fever, micrococcus supposed car- rier, 576 Dermacentor, characters of, 497 — occidentalis , bite of, effects, 504 characters and morphology, 504 (wood tick), characters, 504 — reticnlatus, hosts of, 502, 503 transmission of Babesia caballi by, 178 — variabilis, 505 — venustus, characters and morphology, 503 hosts of, wild and domestic, 504 Dermanyssus gallinoe, disinfection methods against, 704 effect on skin of host, 492 infection, symptoms set up by, 703 morphology, 492 synonyms, 492 — hirundinis, disinfection against, 704 skin affections due to, 492 Dermatitis intertriginoides set up by Oxyii- ris vermicularis , 696 Dermatobia cyaniventris, 596, 597 characters, 598 geographical distribution, 598 hosts of, 596 larvae of, characters, 597 local or vernacular names for, 598 see also Mosquito worm — noxialis, 597 larvae of, skin disease caused by, 725 Dermatocentor reticulatus, var. occiden- I talis, carrier of Rocky Mountain spotted I fever, 496 I Dermatodectes, 521 ! Dermatophagoides scheretnetewskyi, 521 Dermatophilus, 543 — ccBcata, 544 — penetrans (jigger, chigoe), characters and morphology, 544 geographical distribution, 544 j — (Sarcopsylla) penetrans (sand flea), en- I trance beneath skin, 714 1 host of, 613 ' — lesions produced by, 714 I — treatment, 715 I — possible carrier of leprosy, 613 Dermo-muscular layer of nematodes, 361 Derrieu and Raynaud, chronic dysentery due to trichomonads, 624 trichomonad-like organism discovered by, 624, 735 Desvoidea, characters, 563 Deve, experimental development of hyatid scolices, 353 Diarrhoea associated with Balantidium coli, 201 with invasion by Balantidium minu- tum, 204 with Lamblia intestinalis , 59, 625 with Trichomonas hominis, 54, 624 with presence of Watsonius watsoni, 235 with Prowazekia asiatica, 65 — blood-stained, in Strongyloides sterco- ralis infection, 674, 675 INDEX 851 Diarrhoea caused by Difdmus tunensis, 57 — chylous, from Filaria bancrofti infec- tion, 678 — (Cochin-China) in cases of infection with Strong yloides stercoralis, 380, 381 — flagellate, 623 — ■ climatic distribution, 623 — infantile, spread by house-fly, 586 — white (or white scour), in fowls, causal agent, 145 DibothriocephalidcB, 308, 309 — morphology, 308 Dibothriocephalince, 308, 309 — morphology, 308 Dibothriocephalus, 309 — cordatus, 315 cephalic end, 315 hosts of, 315 — excretory apparatus, collecting tubes, island formation, 292 — felis, 313 — latus, 310 chains of segments, 311 development, 311 developmental cycle of, 16 disturbances produced by, in man, 314 duration of life, 315 embryophore of, 298 experimental infection of man with, 312 geographical distribution, 313, 314 growth of, 306 — — habitat in man, 658 head of transverse section, 311 hosts of, 313 human parasite invading animals, 7 in muscles of trunk of burbot, 313 intermediate host of, 255 means of transmission to man and other hosts, 314 mode of infection, 658 morphology, 310 ova of Fasciola hepatica to be dis- tinguished from, 242 ovum of, development, 312 parasitic association with Tcsnia solium, 658 percentage in sufferers from tape- worms in various localities, 314 plerocercoids of, 313 habitat and host, 311 how destroyed, 315 inhabiting fish, 314 proglottids of, average number found daily, 312 proglottis, fairly mature, stained pre- paration, 311 prophylaxis against, 668 supposed origin of, 11 — - — symptoms produced by infection by, 667, 668 synonyms, 310 topographical anatomy, transverse section through proglottis showing, 296 — morphology, 309 — parvus, habitat, 316 Dibothriocephalus parvus, how distin- guished from D. latus and D. cordatus, 316 morphology, 316 — plerocercoid of, 300 — synonyms, 309 Dicrocoeliidce, morphology, 232, 265 Dicrocceliuni dendriticuni, intermediate host unknown, 267 morphology, 266 organs of, diagram showing, 265 ova and miracidia, 266 synonyms, 266 — lanceolatum, incidental human parasite, 7 Diesing, first record of case of Metastrongy- lus apri in man, 433 Difdmus tunensis, cause of diarrhoea, 57 characters, 57 Difflugia enchelys, 47 Digenea, morphology, 230 Digestive system, loss of, in parasites, 3 Dinoflagellata, 52 Dioctophyme, 431 — gigas, hosts and habitat in body, 431 — ■ — in man, source of, 431 infection by, 431 morphology, 431 ova of, 432 synonyms, 431 Dioctophymidoe, 431 — characters, 375 Dionis de Carrieres, liver-fluke in right hypochondriac region, 244 Diphyllobothrium (Bothriocephalus) ditre- mum, discovery of progeny of, 13 Diplodiscus subclavatus, hosts of, 6 Diplogonoporus, morphology, 316 — grandis, egg of, 298 morphology, 316 ventral view of genitalia of left side, 317 of portion of strobila, 317 Diptera, bibliography, 612 — biting-mouthed and other noxious car- riers of disease, 600 — characters, 531, 532 — digestive tract of, inhabited by Herpe- tomonads, 102 — larvae of, in man, references to, 599 Dipylidiidoe, 309, 320 Dipylidium, morphology, 320 — synonyms, 320 — caninum, 320 confused with Taenia solium, 660 cysticercoids of, 322 hosts of, 322 dogs and cats infected with, through skin parasites, 323 embryo of, development, 321 expulsion of, drugs suitable for, 660 hosts of, 6, 7, 322 invasion by, effect on central nervous system, 649 morphology, 320 oncosphere of, 299 oncospheres of, animals selected as hosts for development, 299 852 THE ANIMAL PARASITES OF MAN Dipylidium caninum, prevalence of infec- tion by, in children, 659, 660 proglottids of, 322 proglottis of, central portion, 321 region of human body inhabited by, 660 rostellum of, 289 Dirksen, effect on anaemia of expulsion of Tosnia solium, 648 Dirofilaria immitis, 417 — magalhaesi, morphology, 417 — morphology, 416 — repens, 417 Discophora, see Hirudinea Disporea in Myxosporidia, 182, 184 Distoma, opercula of ova of, discovery, 12 — echinatum, redia of, in later stage, 227 Distomata, morphology, 231 — cercarice, 753 Distomum ophthalmohium, 244 Doeveren, van, on transmission of intestinal worms, 11 Doflein, Coccidiomorpha, 129, 151 — Cnidosporidia, 129, 194 — Entamceba kartulisi, 44 — Trypanosoma equiperdum, 97 Dog and cat, parasites common to, 6 — blood of, life-cycle of Babesia (Piro- plasma) canis in, 175 — DipylidiuTYi canimim parasite in, 322 — fleas, natural flagellates in, 111, 112 transmission of canine kala-azar by, 103 see also Ctenocephahis canis and Pulex serraticeps — host of Dibothriocephalus latus, 313, 315 of Paragonimus kellicotii, 250 — intestine of, Isospora bigemina parasitic in, 149 — mange, transmission to man, 523 — rearing of Tcenia echinococcns in, 356 — to dog, transmission of leucocytogre- garine from, by tick, 155 — transmission of infection with Demodex folliculorum canis to man, 709 Dogs, contraction of surra by, 9G — dermal infection with larvae of Ancylo- stomum duodenale, 455 — echinococci in, 346 — experiments with, to prove transmission of infantile kala-azar by fleas. 111 — infected with Dipylidium caninum through skin parasites, 323 — infection with infantile leishmaniasis, 110 experimental, 110 natural, 110 — Leishmania tropica in, 108 — malignant jaundice in, carrier of, 493 cause of, 177 — nagana fatal to, 94 — percentage infected with Tcenia echino- coccns in various cities and countries (footnote), 345 — prevention of echinococcus infection by, 346 Dogs, segregation of, as preventive against Oriental sore, 628 — skin diseases in, due to young nema- todes, 378 Doliocystidce, 135 Domestic animals, species of Sarcoptes transmissible from, to man, 520 trypanosomes deleterious or lethal to, 69, 70 Dormouse, inoculation of Trypanosoma lewisi into, 90 Dornbliith, method of evacuation of Oxyu- ridoe, 697 Dorr, tapeworm drug recommended by, 674 Dourine, anaemia and paralysis in, 97 — periods or stages of, 97 — trypanosome causing, 97 Dovecots, habitat of Argas reflexus, 506 Dracontiasis, disorder named by Galen, 386 Draciinculidce, 385 — characters, 374 Dracunculus medinensis (Guinea worm), anterior extremity, 387 expulsion by extraction, 676 — methods and drugs for, 676 female, transverse section of, 388 — — infection by, prophylaxis against, 676 — symptoms and lesions following, 389 intermediate host, 388 larvae of, 388 life-history, 386, 388 methods of extraction from body, 389 morphology, 386 period of development in man, 389 regions of body inhabited by, 386 synonyms, 386 viviparous nematode, 371 — morphology, 385 Dromedaries, " mbori " in, 96 Drone fly, 583, 584 Drosophila melanogaster, larvae of, charac- ters and habitat, 584 effects produced by ingestion of, 584 Drosophilidoe, characters and habitat, 584 Drouillard, favourable effects of expulsion of Ascaridoe, 649 Drugs, reaction of spirochsetes to, 115 Dubini, Filaria (?) conjunctivae in man, 405 Dubreuilh, infection with Demodex folti- culorum, 708 Ducks, destruction of mosquito larvae by, 636 — how infected by Echinostoma echinatum, 226 Dufour, creation of name Gregarina by, 129 Duguet, maculae caeruleae (tdches bleues) due to infection by crab louse, 712 Dujardin, development of Taeniae, 14 — psorosperms, 181 Duke, anterior station of trypanosomes in Glossince, 101 — Trypanosoma gambiense in antelope, 76 Dum-dum fever, 105 INDEX 853 Dumesnil, larvae of Muscidce in nose, 720 Duodenum, flagellate stage of Larnblia in- testinalis found in, 59 — - human, habitat of Ancylostoma duo- denale, 450 — possible invasion by Balantidium 7ninu- tuvi, 204 — species of Trichostrongylus inhabiting, 435, 436 Durham, specimens of Leptus (bete rouge), 486 Dutton, Trypanosoma gamhiense, 68 — and Todd, herpetomonads in mice, 738 researches on Spirochoeta duttoni, 116, 117 Duval, liver-flukes in veins, 243 Dysentery, amoeba as causal agent, 30, 34 — experiments made to prove, 30, 40, 618 — amoebic, acute, symptoms, 618 carriers, 618 chronic, 618 experimental production, 618 gangrenous, treatment, 619 — — incubation period long, 618 latent, 618 leading to liver abscess, 618 preventive measures, 620 relief of griping and straining, 618, 619 — ■ — treatment by emetine hydrochloride, 618, 619 by ipecacuanha, 619 by preliminary administration of castor oil, 619 of liver abscess, 620 — — — surgical, 619 — association of Lamhlia intestinalis with, 59, 625 of Noc's entamoeba vd^ith, 41 of Trichomonas with, 56, 624 — bacillary, 31 Entamoeba coli present in cases of, 38 — bacillus, discovery of, 31 — balantidian or ciliate, 201, 202, 637 prophylaxis, 637 ■ — symptoms, 637 treatment, 637 — discoveries of Cholera Investigation Commission with regard to, 30 — discovery of intestinal amoebae in cases of, 30 — flagellate, 623 diet in, 625 geographical distribution, 624 prognosis, 625 prophylaxis, 625 protection of food supply against fsecal contamination in, 625 treatment, 624, 625 — followed by recovery from oxyuriasis, 698 — production by injection of amoebae into cats, 35 — red, geographical distribution, 147, 741 in cattle, cause of, 147 — so-called, due to mites, 512 Dysentery spread by house-fly, 586 — treatment by bismuth subnitrate, 619 Dysodius lunatus, bite of, 541 Ear, abscess of, liver-fluke in, 244 — parasites in, 615 see also Myiasis, auricular Ears of hosts infested by Ornithodorus m6g- nini rendered painful, 510 Earth-eating in connection with Trichuris trichiura infection, 679 East Coast fever in cattle, cause of, 178, 179 pathogenic agent, 174 Eau de Cologne, application in nasal myia- sis, 719 Echidnophaga, 543 Echinococcus, brood capsules, 349 and scolices, mode of formation, 348, 349 transformation into daughter cysts, 351, 353 — cysticus jertilis, 350 — cysts causing urticaria, 651 — daughter cysts, 350 — definition of, 301 — fluid, chemistry of, 353 — frequency of infection by, of various organs in slaughtered animals, 347 — hominis in liver, incised fibrous capsule and wall showing daughter cysts, 351 — in dogs, oxen, sheep and pigs, 346 — in man, age incidence, 355 death at various stages of develop- ment, 356 — — geographical distribution, 354, 355 organs of body invaded by, 355 percentage of prevalence in Central Europe, 354 secondary, 356 sex incidence, 355 — infection, prevention of spread by dogs, 346 — m,ultilocularis (alveolar colloid), develop- ment, 357, 358 feeding experiments with Taenia from, 358 booklets of, 359 in liver of ox, 357 in man, early disintegration, 357 geographical distribution, 358 invasion, results of, 358 _ _ _ site of, 358 morphology of, 356 reasons for distinction from hydatid or unilocular, 357, 358 — of liver rupturing into abdominal cavity, 652 — rate of growth, 354 — rich in glycogen, 348 — scolex, 349 development in rabbits, 353 854 THE ANIMAL PARASITES OF MAN Echinococcus scolex iu jirocess of vesicular metamorphosis, section through, 351 invaginated, section through, 350 transformation into daughter cyst, 352 — scolices in, 303, 349 — serum diagnosis of, 359 — structure and development, 347 — veterinorum, 350 brood capsules and scolices, 350 hooklets of, 355 Echinorhynchus, anatomy, 475 — excretory organs, 475, 476 — "floating ovaries" of, 150 — gigas, hosts of, 477 incidental human parasite, 7 intermediate hosts, 477 morphology, 477 — hominis, 478 — moniliformis , hosts, habitat and inter- mediate host of, 478 — nervous system, 475 — ova of, 477 — protrusor proboscidis, 475 — receptaculum proboscidis, 475 — retractor proboscidis, 475 receptaculi, 475 — sexual organs, 476 Echinostoma, cercarise of, 225 — echinatum, method of infection of duck.3 and geese by, 226 — ilocanum, habitat, 268 morphology, 267 organs of, diagram showing, 268 — m,alayanum, habitat, 269 morphology, 268, 269 — morphology, 267 EchinostomidxB, 267 — morphology, 233 EchinostominoB, 267 — morphology, 233 Ectoparasites, 1 — permanent, changes in, 3 Ectoplasm of protozoa, 25, 26 substances deposited in, 26 Ectoschiza, 135 Eczema due to head louse infection, 709, 710 — following infection by crab louse, 712 — occupational, diagnosis from scabies, 706 — peri-anal and perineal, set up by migra- tions of Oxyuris vermicularis, 695 — purulent, following infection by ancylo- stomes, 684 — resulting from clothes louse infection, 711 — set up by infection with Dermanyssus gallince, 703 by Leptus autiunnalis , 702 Ehrenberg, Spirochceta plicafilis, 114 Ehrlich's acid hsematoxylin, 751 Eimer, researches on coccidia, 136 Eimeria, 142 — avium, causal agent of white diarrhoea or white scour in fowls, and blaclfhead in turkeys, 145 cause of fatal epizootics among game birds and poultry, 142 Eimeria avium, infection by, method of, 145 life-cycle of, 142-145 period, 144, 145 phases, 142-144 merozoites of, 143 microgametes and macrogametes of, 143, 144 — — relation to E. stiedoi, 145 sporozoites of, 143 trophozoites of, 143 — falciformis, 136 — (Coccidium) schuhergi, life-cycle of, 138- 141 — hominis, 150 bodies described as, 150 — stiedce, ascribed cause of " red dysen- tery " in cattle, 147 host of, 7, 145 oocysts of, 142, 146 parasitic in rabbit and occasionally in man, 145, 148 schizogony, 147 — — effects, 147 synonyms, 145 see also Coccidiosis — synonyms, 142 Eimeridea, 141, 742 Electrolytic needle, application in creeping disease, 731 Elephantiasis arabum from Filaria han- crofti infection, sites of body affected by, 677 symptoms, 677 treatment, 677 — scroti in filariasis, 402 Ellermann, " rhizopods in poliomyelitis acuta," 46 Elmassian, discovery of Entamoeba minuta, 42 of trypanosomes in " mal de caderas," 68 Embryophore of tapeworms, 298 Emetine and vaccine treatment combined iu pyorrhoea alveolaris, 620 — hydrochloride in flagellate dysentery, 625 — — in treatment of amoebic dysentery, 618, 619 — in oral endamoebiasis, 620 Emily, expulsion of guinea worm, 676 — method of extraction of Dracnnculus medinensis , 389 Endamoeba, 31, 34, 734 — see also Entamoeba Endamoebiasis, oral, 620 treatment, 620 Endermol, application in scabies, 707 Endoparasites, 1 — intermediate generations invading inter- mediate hosts, 5 hosts of, 5 — young of, leaving host or organ of host inhabited by parents, 5 Endophlebitis set up by Schistosoma hcema- tobium, 274, 275 Endoplasm of protozoa, 25, 26 substances deposited in, 26 INDEX 855 Endoschiza, 135 Endotoxins in Trypanosoma equiperdiim, 98 Endotrypanurn schaudinni, 99 Enemata, in evacuation of Oxyuridce, 697 Entamreba, 31, 734 — africana, see under Entamoeba tetragena — bticcalis, 43, 734 association with cancer of oral cavity, 43 with dental caries, 43 — with pyorrhoea alveolaris, 43, 734 characters, 43 possible relation of E. pulmonalis to, 45 — biitschlii, 34 — coli, 32, 618, 733 characters, 33 — — cysts of, 33 in normal faeces, 33 encystment process, 33 — cytological changes during, 33 how distinguished from E. histolytica, 34, 40, 733 life-cycle of, 32, 33 non-pathogenic and non-culturable, 618 parasite of human intestine, 618 may be present in bacillary dysen- tery, 38 schizogony of, 32, 33 so-called autogamy of, 34 — — sporogony of, 32, 33 — gingivalis, 733 synonyms, 734 — hartmanni, 34 — histolytica, 32, 34, 45, 618, 733 causal agent of amoebic dysentery, 35 changes in intestine produced by, 35 — — ■ characters, 34, 35 cysts of, permanent, injection pro- ducing infection, 37 dysentery following experimental in- fection with, 618 how distinguished from E. coli, 34, 40, 733 present in large intestine, 38 — — producing liver abscess, 35, 620 sporulation of (so-called), 37 and E. coli, differences between, 34, 40, 733 mixed infection, 38 and E. tetragena, identity of, 38, 40, 41 — hominis, 42 — kartulisi, 44, 734 association with dental caries, 44 — maxillaris , 734 — minuta, 42 relation to E. histolytica, 40, 42 — mortinatalium, 45 — nipponica, 42 — Noc's, 41 association with liver abscess and dysentery, 41 — phagocytoides, 42 — poleki, 34 Entamoeba pulmonalis, 45 relation with E. buccalis, 45 — tetragena, 38 characters of, 39 described as E. africana by Hart- mann, 38 found in amoebic dysentery, 38 infection by, 40 multiplication of, 39 nucleus of, 39 ' part of life-cycle of E. histolytica, 38 reproduction of, 39 trophozoites, 39, 40 and E. histolytica identical, 38, 40, 41 — tropicalis, 41 — undulans, 43, 44 characters of, 43 probable flagellate nature of, 44 — williamsi, 34 Entamoebse of vertebrates, 34 — question of cultivation on artificial media, 42, 742 Enteritis, hsemorrhagic, in Strongyloides stercoralis infection, 674 — produced by injection of amoebae into cats, 36, 37 — verminosa in children, 688 Entozoa, 1 — derivation of, 21 Enyaliopsis durandi, bite of, 542 — petersi, 542 — species of, producing ulcers, 542 Eosinophilia in ancylostomiasis, 647 — in bilharziasis, 641 — in hydatid disease, 652 Epicarin, application in scabies, 707 Epidermis, human, excavation of tunnels in, by Sarcoptes scabiei, var. hominis, 519 — " wormlet " burrowing into, 599 Epistaxis, association with presence of Lin- guatula in nasal cavity, 526, 527 — leeches in nose causing, 701 Epithelium of nematodes, 360 Epizoa, 1 EproboscidcB, see Pupipara Epstein, experimental infection with Ascaris lumbricoides , 465 — transmission of trichomonad infection, 56 Epstein's method of diagnosis of ascariasis, 692 Equines, baleri in, causal agent, 95 — biliary fever in, cause of, 177 Erdmann, experimental infection with Sarcosporidia, 192 Erythema, autumn, set up by Leptut autum^nalis, 485, 486 — following bite of Argas reflexus, 506 — set up by infection with Dermanyssus gallince, 703 by Leptus autumnalis , 702 i Eschatocephalus, characters of, 497 I — hosts and habitat of, 497 Escomel, dysentery due to Trichomonas, I 56, 624 856 THE ANIMAL PARASITES OF MAN Escomel, treatment of esjjundia, 629 of lamblial diarrhoea, 625 Espundia, 108, 628 — course of, 629 — geographical distribution, 628 — pathology of, 628 — prophylactic measures against, 629 — transmission of, 629 — treatment, 629 Ether, application in nasal myiasis, 719 — lotions in crab louse infection, 712 — sulphuric, in ^ crab louse infection, 712 Ethyl chloride, freezing by, in creeping disease, 731 Eucalyptus oil in expulsion of ancylostomes, 687 Euflagellata, 52 — characters of, 52 Eugregarinea, schizogony absent in, 134 EuLyes amoena, 542 Euphorbia, infection by Herpetomonas davidi, 104 Eupodidoe, characters, 491 Euquinine in malaria, 635 Europe, Central, percentage of prevalence of echinococcus in man in, 354 Eustrongylus gigas, infection by, site of, 681 symptoms, 682 Evans, discovery of trypanosomes in blood of horses with " surra " disease, 67 — see also Steel and Evans Excretory apparatus of cestodes, 291 — or segmental organs of Hirudinea, 481 — organs of Echinorhynchus, 475 of nematodes, 366, 367 Eye, cysticerci in, 335, 664 — cysticercus of, 664 diagnosis from foreign body, 664 — diseases due to Liicilia Tnacellaria, 721 — human, infection with filaria, 406 — — nematodes observed in, 412 — invasion by Loa loa, 678 — paragonimiasis of, 639 Eyeball, bodies found in, probably young liver-flukes, 244 Eyebrows and eyelashes, pediculosis of, treatment, 712 Eyelid, iipper, extraction of larva of Hypo- derma hovis from, 596 Eyelids, Demodex folliculorum affecting, 708 Eyes, destruction of, by Sarcophaga wohl- fahrti, 723 — loss of, in parasites, 3 Fabre, prophylaxis against ancylostomiasis, 685 Face, swelling of, in nasal myiasis, 718 Faeces, amoebse found in, 30, 34, 47, 48 — asexual multiplication of Chlamydophrys enchelys in, 47 — examination for protozoa, 746 — human, fresh, Strongyloides stercoralis larva from, 382 Faeces, larvae of Uomalomyia canicularis found in, 584 of Piophila casei found in, 583 — males of Oxyuris vermicularis rarely met with in, 468 — normal, cysts of Enta1no^ba coli present in, 33 — preservation of ova of flukes in, 472 — N^orticella in, 206 Fanapapea intestinalis identical with Tetramitus mesnili, 57 Fantham, H. B., appendix on protozoology (recent researches, formulae of culture media, general protozoological tech- nique), 733 avian coccidiosis, 145 pathogenic spirochaetes, 119 classification of Haplosporidia, 195 of Schizogregarines, 135 experimental infection with Spiro- chceta duttoni, 117 granule phase of spirochaetes, 120 Herpetomonas ctenocephali, 103 pedictili, 103 latent forms of trypanosomes, 73, 74, 77 molluscan spirochaetes breaking up into granules, 119 — — morphology and life-cycle of Eimeria avium, 142-145 and life-history of Spirochoeta hronchialis, 739 nuclear phenomena of Babesia hovis, 176 Protozoa, 25 recent work on spirochaetes of human mouth, 740, 741 Rhinos poridium kinealyi, 195-197 schizogony in Leucocytozoon lovati, 153 significance of insect flagellates in relation to disease, 104, 739 Theileria parva, 179 — — Trypanosoma rhodesiense, 69, 76 ■ and Porter, experimental introduction of insect flagellates into vertebrates, 104, 112, 738 inoculation experiments with Her- petomonas jaciilum-, 104, 738 — natural herpetomonads in mice, 739 researches on Nosema apis, 185 ■ — ^ on Spirochceta. duttoni, 116 ' — — and Thomson, J. G., cultivation of Babesia canis, 172 periodic cyclical variation of trypanosomes in blood, 78 see also Stephens and Fantham Fasciola, 237 — gigantica, distribution, 244 habitat, 244 invading lung, 245 morphology, 244 — section illustrating, 243 synonyms, 244 — hepatica, 237, 238, 239, 638 INDEX 857 Fasciola hepatica, cercaria of, 228 encysted, 228 development of, 226 fixtition method, 471 geographical distribution, 238 half transverse section through, 214 hosts of, 6, 7, 238 incidental human parasite, 7 intermediate host, 240, 241 invading and infecting pharynx, 242 regions of body other than liver, 243 life-history, 241 — — method of infection of sheep by, 226 miracidium of, 223 morphology, 237 sections illustrating, 238, 239 ova of, to be distinguished from those of Dibothriocephalus latus, 242 ovum of, 223 from liver of sheep, 240 redia, in early stage, 227 — — synonyms, 237 — morphology, 237 Fascioliasis, prevention and treatment, G38 — symptoms of, 638 — see also Liver-fluke disease Fasciolidce, 237 — morphology. 231 Fasciolince, 237 — morphology, 231 FasciolopsincB, 245 — morphology, 231 Fasciolopsis buski, fixation method, 471 geographical distribution, 246 habitat, 246, 638 morphology, 245, 246 symptoms set up by invasion by, 638 — fiillehorni, cirrus sac, 247 habitat, 249 ' morphology, 247 — ventral aspect showing, 248 — goddardi, morphology and geographical distribution, 247 — rathouisi, geographical distribution, 247 habitat, 247 morphology, 246 — morphology, 245 Feltinella, characters, 561, 567 Females, greater prevalence of head louse infection among, 709, 710 — rarity of bilharziasis in, 643 Fern root, new species, effects as vermi- fuge, 673 Fibrin, clots of, Dioctophyme gigas in man traced to, 431 Ficalbia, characters, 565 Fievre de grain, 702 Fiji, intermediate host of filaria in, 575 — manifestations of filariasis in, 402 Filaria associated with phthisis, 408 — hancrofti, anatomy of, diagrams show- ing, 391 discoveries relating to, 390 diseases following infection by, 398i 400 Filaria hancrofti, embryos, 392 female, characters, 392 geographical distribution, 392, 403 habitat In body, 392 ■ infection by, diseases resulting from, 676, 677 larvae of, absence from blood in those suffering from filarial disease, 400 distribution in body, 392, 393 method of concentration, 395 of preservation, 395 morphology, determination by fixa- tion and staining methods, 395, 396 — periodicity of, in peripheral blood, 393, 394 separation of red corpuscles from. of Tseniorhynchus carry- 395 species ing, 577 — — — structure, 396 life-history, 398 male, characters, 392 mosquitoes acting as hosts of, 398 ova of, 392 synonyms, 390 transmission of, 576 — (?) conjunctiva, 404, 405 — — morphology, 404 normal hosts of, 406 sites of infection in man, 405 synonyms, 404 — demarquayi, 403, 404 geographical distribution, 403 morphology, 403 — infection of skin by, 378 — intermediate host of, in Fiji, 575 — (?) kilimarce, 407 — loa, host of, 601 — medinensis, antiquity of knowledge con- cerning, 386 — morphology, 390 - — oculi humani, association with cataract, 406 — perstans, carrier of, 508 — ( ?) romanorum orientalis, morphology, 407 — (?) sp. (?), 407 — taniguchi, 404 Filariasis, cultivation of bacillus from cases of, 755 — prevalence proportionate to prevalence of Mikrofilaria hancrofti in blood, 400 Filariidce, 390 — characters, 374 Filariinoe, 390 Filmaron, administration in expulsion of ancylostomes, 687 — as vermicide, 672 — oil, dosage of, 672 effects of, 672 in expulsion of Ascarides, 694 ''Filterable viruses," 207 Filtration experiments with CMamydozoon granules, 209 Finlaya, characters, 564 Finsen, echinococcus cysts causing urti- caria, 651 858 THE ANIMAL PARASITES OF MAN Finucane, lymphangitic symptoms ia chil- dren from Filaria hancrofti infection, 676 Fischer, effects of new species of fern root as a vermifuge, 672, 673 — retinal haemorrhages in ancyloatome anaemia, 646 Fish, destruction of mosquito larvae by, 636 — disease of, due to invasion by Myxo- sporidia, 182, 184 — eating of raw or badly cooked, favours transmission of Dibotliriocephalus latus in man, 315 — fresh-water, second intermediate host of Clonorchis endemicus, 261 proved by feeding ex- periments, 261 — intermediate host of Dihothriocephalus latus, 255 of Opisthorchis felineus, 255 — parasitic ciliate destructive to, 206 — plerocercoids of Dibothriocephalus latus inhabiting, 314 — trypanoplasms in, 68 Fistulas, anal and rectal, set up by migra- tion of Oxyuris vermicularis, 695 — formation of, through migration of para- sites, 9 — urethral, arising from bilharziasis, 642 ■ — treatment, 644 Fixation, time of, 749 Fixatives, 748, 749 — for wet films, 748 — hot, 748 Flagella may occur among rhizopods, 52 — of Flagellata, 50, 51 Flagellata, 50 — characters of, 28, 50, 51 — classification, 52 — formation of colonies of individuals, 51 — habitat, 28, 52 — multiplication of, 51 — non-flagellate stages, 51 — nuclear apparatus of, 51 — post-flagellate and pre-flagellate stages, 52 Flagellates, aggregation rosettes of, 51 — dysentery in children due to, 56, 624 — in blood of horses, diseases associated with, 67, 68 — natural, in dog fleas, 112 of invertebrates, evolution of Leish- lYiania from, 739 —^ parasitic, in insects, experimental intro- duction into vertebrates, 104, 112, 737, 738 relation to evolution of leishmaniasis, 112, 737, 738 in relation to evolution of disease, 739 Flagellosis of plants, possible connection with leishmaniasis, 104, 739 Flat worms, see Platyhelminthes Flea, human, see Pulcx irritans Fleas acting as intermediate hosts, 543 — blood-suckers, 543 — carriers of plague, 543 ]'^leas, cocoons of, 543 — fed on infected rat, percentage infected with trypanosomes, 93 -- herpetomonads in gut of, 103 — larvae of, 543 — life-cycle of Trypanosoma lewisi in, 88, 90 — ■ method of controlling, during experi- ments, 93 — ova of, 543 — jjossible transmission of kala-azar by. 111 Flemming's solution, 749 Flesh of animals containing larvae of tape- worms must be thoroughly cooked be- fore eating, 668 Flesh-fly, see Sarcophaga carnosa Flexner, Entamoeba kartulisi, 44 Flies, larvae of different species of, found in intestinal myiasis, 728 Flukes, clearing and mounting agents, 471 — differentiation methods, 471 — fixation methods, 471 — miracidia of, discovery, 12 — ova of, preservation in faeces, urine, bile, 472 transference to glycerine, 472 — preservation and examination of, 471 — staining methods, 471 — see also Trematoda Foetus, Trypanosoma cruzi in, 88 Foley, transmission of relapsing fever, 120, 121 Food, transmission of trichomonad infec- tion by, 56 Foods, decomposing, inhabited by and nutriment of TyroglyphidcB, 511 Foot, sole of, hepatic flukes found in swell- ing on, 243 Foot-and-mouth disease, 207, 208 Foraminifera, 27, 47 — characters and habitat, 27 Forde and Dutton, discovery of human try- panosomes, 68 Foreign body, diagnosis of cysticercus of eye from, 664 Formalin, fixation of cestodes by, 472 Fowler, method of administering male fern to children, 671, 672 Fowler's solution in sleeping sickness, 623 Fowls, fatal effects of Spirochoeta galli- narum on, 119 Fox, host of Dibothriocephalus latus, 313 Framboesia tropica, see Yaws Fran9a, action of leucocytozoa on red cells, 153, 742 — genera of Piroplasmidce, 174 Francaviglia, auricular myiasis, 615 Franchini, experimental infection of verte- brates with herpetomonads, 103, 104, 112, 739 — Hcemocystozoon brasiliense, 104 Frese, O., rhabdites found in gastric fluid obtained by lavage, 378 Freund, sarcophaga larvae from abscess cavities, 723 Frog, rectum and bladder of, ciliates para- sitic in, 207 INDEX 859 Frontal sinus, invasion by Ancylostoma duodenale, 683 scolopendra in, 721 Fruit pickers affected by Leptus autumna- lis (footnote), 485 Fiilleborn, cultivation of larval forms of Ancylostoma and Strongyloides, 474 Furcocercous cercarise, 753 Fiirst, cases of Ascarides invading- larynx and trachea, 691 Gabel, diarrhoea due to Difdmus tunensis, 624 — Difdmus tunensis, 57, 624 Gad-flies, 600 — see also TabanidcB Gaetano, cysticercus of tongue, 663 Galen, disorder named " dracontiasis " by, 386 Gall-bladder, Schistosoma hcematohium, eggs of, in, 274 Gall sickness in cattle, cause of, 98, 180, 611 Galleria melonella, larvae of, in nose, 720 Galli-Vallerio, bothriocephalus anaemia, 646 — infection by trichomonads, 56 — Oxyuris and Trichocephalus infection in relation to appendicitis, 654 Galyl in syphilis, 632 — ■ in trypanosomiasis, 622 Gamasidce (coleopterous or insect mites), characters, 491 hosts and prey of, 491 Gambia horse sickness, cause of, 100 Game infested by Dermacentor reticulatus, 503 Game-birds, fatal epizootics among, due to Eimeria avium, 142 Gametes of Coccidiidea, 137, 139, 140 — of gregarines, 132, 133 — of malarial parasites, 162 Gametocytes of Coccidia, 140 — of gregarines, 132, 133 — of malarial parasites, 162 Gametogony, in Eimeria avium, 143 Gam,marus pulex occasionally parasitic in man, 483 Garlic and saline injections in Trichuris trichiura infection, 680 GastrodisciidcB, 236 — morphology, 231 Gastrodiscoides, how distinguished from Gastrodiscus, 236 Gastrodiscus, 236 — mgyptiacus, hosts of, 237 — ho minis, 236 genital pore, 236 geographical distribution, 237 ■ habitat, 237 morphology, 236 ova, 237 testes, 236 — male and female genitalia, 236 — morphology, 236 Gastrophilus, 599 Gastrophilus equi, 599 — hcemorrhoidalis , 599 — larvae of, in stomach and intestine, 599 — nasalis, 599 — pecorum, 599 Geber, treatment of crab louse infection, 712 Gecko, blood and organs of, herpetomonad flagellate in cultures from, 739 Gedoelst, Cordylobia rodhani, 593 Geese, how infected with Echinostom,a echinatum, 226 Genital apparatus of Cestoda, 293-296 Genitalia as means of distinguishing species of Glossina, 604 Genitals, external female, invaded by Oxyuris vermicularis, 467 Genser, von, Ascaris infection in relation to appendicitis, 653 Gentian violet, 752 Gerbillus indicus, 154 Gerlach, stages of liver-fluke disease in sheep, 240 Germany, districts of, j3ercentage of occur- rence of echinococcus in man in, 354 — trichinosis in, epidemics of, 423, 429 — — prophylaxis against, 429 Giard, microscopical investigations of con- jugation in gregarines, 130 Giardia (Lamblia) intestinalis, 736 Giemsa's stain, 751 formula of, 751 Giesker, liver-fluke in sole of foot, 243 Gilesia, characters, 564 Gingivitis, nematode larvae in periosteum of upper jaw associated with, 378 — see also Entamoeba gingivalis, 733; and E. buccalis, 43 Girard, effects of Trichocephalus infection, 651 — Trichocephalus infection in relation to appendicitis, 653 Glas, cysticercus of tongue, 663 Glossina austenii, characters, 605 — brevipalpis group, characters, 606, 607 — caliginea, characters, 605 — characters, 603, 604 — development of trypanosomes in, 101 — fusca, characters, 606 — — group, characters, 606 — fuscipleuris, characters, 606 — habitat of species, 605 — ■ larvae of, 604 — longipalpi.-i, characters, 606 head of, 664 — longipennis , characters, 607 — medicorum, characters, 607 — morsitans , characters, 606 development of Trypanosoma brucei in, 94 development of Trypanosoma rhode- siense in, percentage of fly becoming infective, 82 developmental cycle of Trypanosoma rhodesiense in, 81, 82 — — geographical distribution, 608 group, characters, 605 method of reproduction, 604 54 86o HE ANIMAL PARASITES OF MAN Glossina morsitans, race submorsitans, 609 transmission of nagana (tsetse-fly disease) by, 93 of Trypanosoma rhodesiense by, 69, 81, 608 — nigrofiisca, characters, 606 ■ — pallicera, characters, 605 — pallidipes , antenna of, 604 — — characters, 606 — palpalis, 100 blood-sucking, 607 carrier of sleeping" sickness, 605, 607 — — characters, 605 development of Trypanosoma gam- biense in, 74, 75 geographical distribution, 607 group, characters, 605 larval and pupal stages, 608 — — proportion becoming infected, 608 salivary glands of, invasion by Try- panosoma gavibiense, 75 transmission of sleeping sickness in- fection by, 68 — puparia of, 604, 605 — special means of distinguishing species, 604 — species of, artificial infection with human trypanosome, 60.5 — spread of trypanosome diseases by, 603 — tabaniformis, characters, 606 — tachinoides, characters, 605 Glycerine, mounting agent for flukes, 472 — transference of ova of flukes to, 472 Glycerophosphates and arsenic in bronchial spirochaetosis, 633 Glychaemalum, Mayer's, 751 Glyciphagi, differentiation from Tyro- glyphi, 513 — buski, 513 — cursor, 513 — domesticus , cause of grocer's itch, 513 — hippopodes, 513 — prunorum, 513 Glycogen, echinococcus rich in, 348 Gnathobdellida, 481 Gnathostoma hispidum, hosts of, 385 — morphology, 384 — siamense, infection by, associated with tumour of breast, 385 morphology, 384 — sp., hosts of, 385 — spinigeriim, 385 hosts of, 385 morphology, 385 Gnathostotnidce, 384 — characters, 374 Gnats, see Culex Goebel, bilharziasis, 641, 642 Goeldia, characters, 565 Golden beetle, intermediate host of Echino- rhynchiis gigas, 478 Goldmann, male fern extract in expulsion of Strongyloides stercoralis , 675 — sebirol as vermicide, 672 — taeniol in ancylostomiasis, 686 Goldschmidt, excretory apparatus oi Ascaris lumbricoides , 367 — formation of ova of trematodes, 223 Golgi, description of asexual cycle in blood in case of quartan parasite, 157 Gonder, relation of infantile kala-azar to Oriental sore, 108, 109 — strain of Trypanosoma lewisi losing resistance to arseno-phenyl-glycin, 03 — Theileria parva, 178 Gordiidce, 375 — characters of, 479 — larvae of, 479 Gordius aquations, 479 — chilensis, 479 — pustulosus, 479 — species invading man, 479 — tolosanus , 479 — tricuspidatus , 479 — varius, 479 — villoti, 479 — violaceus, 479 Grabhamia, characters, 564, 576 — do7'salis, 576 — geographical distribution, "576 — sollicitans, geographical distribution, 576 Graffe, escape of ascarides from inguinal tumour, 656 Granate root as vermifuge, 673 Granuloma inguinale, spirochsete associated with, 122 Grass, harvest or gooseberry mite, see Lep- tus autuynnalis Grassi, Cercomonas and Trichomonas, 54 — development of cestodes without inter- mediate host, 17 of Trichuris trichiura, 420 — discovery of amabse in stools, 30 — experimental self-infection with Oxyuris vermicularis, 469 — expulsion of Hymenolepis nana, 661 — Hymenolepis nana, 324 — larval stage of Hymenolepis diminuta, 327 — mosquitoes in relation to human malaria, 158 — on Entamoeba coli, 32, 33 — self-infection with Ascaris lumbi'icoides, 465 Great black bug of Pampas, see Conorhinus renggeri Gregarina blattarum, 135 — longa from larva of crane-fly, 130 — munieri, from Chrysomela hcemoptera, 131 — ovata, host of, 135 Gregarines, ectoplasm of, 131 — endoplasm of, 131 — gametes of, 132, 133 — gametocytes of, 132, 133 — mode of infection, 134 — monocystid, 130 hosts of, 130 — morphology of, 130 — movements of, 131 — myonemes of, 130, 131 INDEX 8^1 Gregarines, polycystid, 130, 131 protomerite, deutomerite and epi- merite of, 131 — resistant spores of, purpose of, 134. — spore-production, 132, 133 — sporocyst of, 134 — sporozoites of, 132, 133 — syzygy of, 132 — trophozoites of, 132, 133 — zygotes of, 132, 133 Gregarinida, characters and habitat, 28, 130 — classification, 134 — history of discoveries relating to, 129 Grey ointment in crab louse infection, 712 Grocer's itch, cause of, 513 Ground-itch, skin affection set up by in- vasion of larvae of Ancylostomum duo- denale, 455 — treatment, 754 Guarnieri's bodies, 207, 209 Gubler, case of human hepatic coccidiosis, 148 Guermonprez, method of expulsion of ascarides, 692 Guinea-pigs, experimental infection with Sarcocystis muris, 192 — natural occurrence of Paraplasma in, 180 Guinea worm, see Dracunculus medinensis Gurley on Myxosporidia, 182, 183 H. H^MADIPSA, 482 — blood-sucking pest in tropics, 482 Haemagogus, characters, 565 Hsemalum, Mayer's, 751 Haemamoeba, 151, 742 Haemaphysalis, characters of, 497 — leachi (dog tick), carrier of malignant jaundice in dogs, 493 transmitting agent of Babesia canis, 177 — ^punctata, characters and morphology, 502 — — hosts of, 503 synonyms, 502, 503 Haematein, essential principle of haema- toxylin, 751 — solutions of, in staining flukes, 471 Hoematobia irritans, 610 Hoematopinus spinulosus, 88 effect on strain of Trypanosoma lewisi being passed through, 93 Haematoxylin, Delafield's (or Grenacher's), 751 — Ehrlich's acid, 751 Haematuria in bilharziasis, 641 Haementaria, 482 — officinalis used medicinally, 482 Hmmocvstozoon brasiHense, 104 Haemoglobinuria, infectious, in cattle, cause of, 177 Hmmogregarina balfouri (jaculi), 154 — gerbilii, 154 Uoemogregarina marceaui, 154 — nobrei, 154 — schaudinni, var. africana, 154 Haemogregarines, characters of, 154, 742 — hosts of, 153 — in red blood corpuscles, 154 — leucocytic, 154 — transmission of, 153 — variation in size and appearance of, 154 Haemolysis, cure, after expulsion of Ascaridce, 649 Hcemonchus contortus, diseases due to in- vasion by, 437, 438 geographical distribution, 437 habitat and hosts of, 437 life-history, 438 morphology, 436, 437 symptoms caused by invasion mis taken for those of ancylostomiasis, 438 — • morphology, 436 Hcemoproteus (Halteridium) columbce, in- sects transmitting, 151, 612 life-cycle of, 152 danilewskyi, 152 Haemorrhoidal veins. Schistosoma hoem,a- tobium in, 273 superior, plexus formed by, in rectum, 272 Haemorrhoids set up by migrations of Oxyuris vermicularis , 694 H- vulus, 418 Labbe, copulation in Coccidia, 137 Lacompte, nematodes in human eye, 412 LoRlaps echidniniis, Leishmau granules in, 493 — stabularis, 493 Lafleur, see Councilman and Laflenr Lagocheilascaris, characters, 466 — minor, host of, 467 lesions set up by, 467 — — morphology, 467 Lakes, mosquitoes depositing ova in, 553 Lama, possible carrier of leprosy, 613 Lambkin's mercury cream in treatment of syphilis prevailing in Uganda, 632 Lambl, discovery of human intestinal amoebae, 29 Lamblia intestinalis , 57, 625, 736 association with diarrhoea, 59, 60, 625 treatment, 625 — — characters, 57 flagella of, 57, 58 — — hosts of, 59 — — infection with, 60 nuclear apparatus, 58 site in intestine of flagellate and cystic stages, 59 synonyms, 57, 736 Lankester, liver-fluke in abscess of ear, 244 — Sarcocystis, 193 Lankesterella, 154 Larva migrans, 599 Larvae, dipterous, in conjunctiva, 716 — — in nasal accessory sinuses, 717 in nose in enormous numbers, 716, 717 — in wounds, movement of, 723 INDEX 867 Larvae in wounds, see also under Names of Parasites Larvicides, use in campaign against mos- quitoes, 636 Larynx, ascarides invading, 691 — leeches in, 699, 700 La Spada, echinococcus of liver rupturing into abdominal cavity, 652 Lasioconops, characters, 564 Lassar's paste, application in creeping disease, 732 Laurer's canal of trematodes, 221, 222 Laveran, A., classification of trypanosomes, 71 cross-immunity experiments with Try- panosoma rhodesiense and T. hrucei, 80, 94 — with trypanosomes, 80 discovery of true malarial parasites by, 157 — — latent forms of trypanosomes, 74 on Leucocytozoa, 153, 742 Trypanosoma pecaudi, 95 and Franchini, inoculation experi- ments with Crithidia fasciculata, 104 with Herpetomonas cteno- cephali, 103 — — with H. pattoni, 103 and Mesnil, isolation of sarcocystin, 191 — — — on the spore of Sarcocystis tenella, 193 " Trypanosomes et Trypano- somiases," 617 and Thiroux, treatment of sleeping sickness, 623 Laverania, characters, 164, 569 — nialarice (Plasmodium falciparum), crescents of, 162, 167, 168 sites of development, 169 — — cultivation of, clumping in, 172 — cultures of, number of spores produced, 172 development, duration of, 167 distinctive characters of, 169 — — invasion of spleen by, 168 — — — — merozoites, number of, 168 number in one red blood cor- puscle, 167 oocysts of, in stomach of Ano- pheles, 163 ookinete of, in stomach of Anopheles maculipennis, 162 parasite of malignant tertian or sub-tertian fever, 167 and quotidian malaria, 167, 633 — ^ pathological effects, 634 question of varieties or sub- species, 167 " signet-ring " stage, 167 sporozoites, 169 stages of development in intes- tine of Anopheles maculipennis, 162 — — synonyms, 167 trophozoites of, 167, 168 Leo, R. J., creeping disease, 729 Leeches in upper air passages, 699, 700 cases reported by various authors, 699, 700, 701 mention among ancient writers, 699, 700 — invading body, means of riddance, 701 — see also Hirudinea, Rhyncobdellidos Leeuwenhoek, opposition to theory of spon- taneous generation, 10 Leger, L., classification of Coccidiidea, 141, 142 — genus Crithidia, 104 — researches on Coccidia, 137 Leger, M., proportion of population in Tonkin infected with Glonorchis en- demicus, 260 Leger, M. and A., proposed classification of Leucocytozoa, 153 Leichtenstern, bothriocephalus anaemia, 646 — toxic symptoms following thymol ad- ministration, 686 Leidy, genus Endammha, 31, 34, 734 Leignathus sylviarum, 493 Leiper, R. T., Gastrodiscoides, 236 host of Filaria loa, 601 — — identity of (Esophagostomum hrumpti with (Es. apiostomum, 444 report of Bilharzia Mission under, 277 Leipzig, frequency of infection of various organs of animals with echinococcus slaughtered at, 347 Leiseri ng, percentage of rats infected with Trichinella, 427 Leishman, Sir W. B., experimental re- searches on infection with Spirochceta duttoni, 117, 118 on parasite of Indian kala-azar, 105 treatment of Indian kala-azar, 626 Leishman-Donovan body, see Leishmania donovani Leishman granules in Laelaps echidninus, 493 Leishmania, 67, 104 ' — donovani, 105 cause of Indian kala-azar, 105, 626 cultivation methods, 106 inoculation experiments with, 107 localization of infection, 105 morphology, 105, 106 possible mode of transmission, 107 — evolution from flagellates of inverte- brates, 739 — infantum., cause of infantile kala-azar, 105, 109, 627 cultivation methods, 109 immunity to, 112 in dogs, 110 — — inoculation, 110 animals suitable for, 110 probable transmitter. 111 — probable origin of, 103, 739 — tropica, 105, 107 cause of Oriental sore, 105, 107, 627 i cultivation methods, 108 868 THE ANIMAL PARASITES OF MAN Leishmania tropica, hosts of, 108 inoculation, experimental, 108 possible transmitters, 108 synonyms, 107 Leishmaniasis, cutaneous, 107 — dermo-mucosal, supposed mode of trans- mission in Paraguay, 739 — evolution of, relation of experimental in- troduction of insect flagellates into ver- tebrates on, 737, 738, 739 — experimental production in white mice, 103 — geographical distribution, 105, 107, 109 — infantile, see Kala-azar, infantile — naso-oral, see Espundia — possible reservoirs, 738, 739 — treatment, G26-629 Leishman's stain, 750 Lemaire, herpetomonad flagellate in cul- tures of blood and organs of gecko, 739 Lenhartz, bothriocephalus anaemia, 646 Lentospora cerebralis, 184 Lepidoptera, characters, 531, 532 Leprosy, possible carrier of, 579, 613 Leptidce, 603 — blood-sucking species, 603 — characters, 603 Leptis scolopacea, 603 — strigosa, 603 Leptodera, life-history of, 19 — appendiculata, occasional parasite, 7 — pellio, facultative parasitism of, 8 Leptomonas, 102 — biitschlii, 102 Leptotheca, 184 Leptiis autumnalis (grass, harvest or goose- berry mite), animals attacked by, 486 skin irritation set up by, 702 habitat of, 485 hosts of, 485, 486 — ■ — nut and fruit pickers affected by, 485 skin affection set up by, 485, 486 so-called proboscis of, 485, 486 — geographical distribution of species, 486 — undescribed species of, 486 Lesbini, dipterous larvae in nose in enor- mous numbers, 717 Letulle, pathological changes in rectum due to Schistosoma hosmatohium, 274 Leuckart, E.., advances in helminthology due to, 15, 16 attempt at self-infection with Ascaris lumhricoides, 464, 465 change of host in parasites, 20, 21 classes of parasites, 1 development of Acanthocephala and TAnguatulida, 17 , of alveolar echinococcus, 357, 358 of nematodes, 17 —of Trichinella spiralis, 423 distinction between Cercomonas and Trichomonas, 54 experimental self-infection with Oxyuris vermiciilaris , 469 facultative parasitism, 7 Leuckart, K., feeding experiments with T(Bnia saginata, 340 with Trichinellae, 423 — — growth of echinococcus, 354 — — heterogony in Strongyloides stei^- coralis, 381 — — method of infection with Trichuris ovis, 420 migration of oncospheres, 302 name of Coccidia first given by, 135 Trichocephalus in association with cholera, 658 and Thomas, P., life-history of liver-. fluke, 241 Leucocytogregarina, 154 — canis, life-cycle diagram, 155 transmission from dog to dog by tick, 155 Leucocytogregarines, 154 Leucocytosis in bilharziasis, 642 Leucocytozoa, action of, on red blood cells, 153, 742 — classification proposed, 153 — hosts of, 153 — morphology of, 153 — schizogony of, 153, 742 Leucocytozoon type of Iloimosporidia, 152 — lovati, schizogony in, 153 — ziemanni, schizogony in, 153 Leucomaines, effects on living organisms, 9 Levaditi, cultivation of spinal ganglia of rabid monkeys, 210 Lewandowsky, infection with Demodex fol- liculorum, 708 Lewin, expulsion of Ascarides, 693 Lewis, finding of intestinal amoebae, 29 — studies of filariasis, 391 Leydenia, 49 — gemmipara, 49 in ascites, 49 association with possible ascites and malignant growth in abdomen, 49, 50 characters of, 49 — — cytoplasm containing blood corpuscles, 50 pseudopodia of, joining several in- dividuals, 49 Leydig, psorosperms, 181 Lice, Ilerpetomonads in gut of, 103 — transmission of relapsing fever by, 120 — wingless, owing to parasitic life, 3 — see also Pedicididce Lieberkiihn, investigations of Coccidia, 135 of gregarines, 130 — psorosperms, 181 Liesen, Ascaris in peritoneal cavity, 656^ Ligula, excretory apparatus, collecting tubes, island formation, 292 — plerocercoid of, 300 Limatus, characters, 565 Limnaeus, species other than L. triincatulus intermediate hosts of Fasciola hepatica, 242 — truncatulus, amount of ova deposited by, 242 geographical distribution, 241 INDEX 869 Limnoeus truncatidus, intermediate host of Fasciola hepatica (?), 240, 241 hosts of liver-fluke, 240, 241 Limnatis, characters, 482 — nilotica, characters, 482 geographical distribution, 482 habitat, 482 only leech of clinical importance as parasite, 699, 701 synonyms, 482 Lindblad, Dipylidium caninum, 660 Lindner, G., peritrichal Infusoria (stalkless Vorticella), 206 Lindsay, possible mode of -transmission of dermo-mucosal leishmaniasis to man in Paraguay, 739 Linguatula, 523, 524 — rhinaria, characters and morphology, 524 development and life-history of, 524, 525, 526 larvae of, 524, 525 occurrence at autopsies, 526 organs of body invaded by, 524, 525, 526 ova of, 524, 525 parasitic in nasal cavity of animals and man, 523, 524 synonyms, 524 — serrata, hosts of, 527 synonyms (footnote), 527 Linguatulida, development of, 17 Linguatulidoe, blood-sucking, 523 — change of original features in, 4 — characters and morphology, 523 — hosts of, 523 — larvae of, 523 — nature of, 2 — relation to Arachnoidea, 19 — separation from Helminthes, 2 Lini, escape of Ascarides from umbilicus, 656 Linnaeus, discoveries as to origin of Hel- minthes, 10, 11 — so-called dysentery infection due to mites, 512 Lipari, cysticerci of brain, 664 Lipuria in bilharziasis, 641 Lithocystis, endoplasm of, contents, 131 Liver, abscess of, association of Entamoeba histolytica with, 35 of Noc's entamoeba with, 41 caused by invasion of Ascaridoe, 690 due to amoebic dysentery, treatment, 620 set up by amoebae, 35 — and bile-ducts, habitat of Clonorchis endemicus, 259, 260 — and kidney cells, yellow pigment in, in ancylostomiasis, 647 — and portal vein. Schistosoma hcRmato- hium, most easily found post mortem in, 273 — coccidiosis of, in man, cases, 148 — encystment of Porocephalus constrictus in, 526, 527 — female Ascarides depositing ova in, 689 Liver, human, eggs of Schistosoma japoni- cum, showing " spines " and " hoods " at opposite pole, 279 — invasion by larvae of Linguatula rhinaria, 525, 526 — migration of oncospheres from intestine to, 302 — pathological changes associated with in- vasion by Opisthorchis felineus, 253 — — — in, due to ova of Schistosoma japo7iicum, 281 set up by Clonorchis endemicus , 260 Liver-fluke, supposed origin of, 10 — see also Fasciola hepatica — disease, diagnosis, 242 in man, 242 in sheep, 238 ravages caused by, 238, 239 stages of, 240, 241 pathological anatomy, 241 symptoms, 239 Liverpool School of Tropical Medicine, ex- pedition to investigate trypanosome in- fections, 68 Lizards, haemogregarines from, 154 Loa, morphology, 409, 411 — • loa, duration of life of, 414 early historical accounts of, 412 geographical distribution, 414 larvae of, in blood, 412, 414 periodicity, 413 structure, 412 lesions produced through invasion by, 413, 414 life-history, 414 morphology, 409, 411 ova of, 410 • sites of body invaded by, 412, 678 synonyms, 409 Lobaczewski, prophylaxis against body, head and clothes lice, 615 Lobker, cause of ancylostome anaemia, 648 Locusts injurious to man, 542 Looss, infection by Ancylostoma duodenale through skin, 683 — origin of lateral-spined eggs of Schisto- soma hcematohium,, 273 — prevalence of Heterophyes heterophyes, 264 — skin affections set up by invasion of larvae of Anciflostoma duodenale, 455 — symptoms of lymphangitis from Filaria bancrofti infection, 676 — toxic action of ancylostomes, 647 — Trichostrongylus instabilis in man, 435 Lophius piscatorius, 186 Lophoscelomyia, characters, 562, 568 Losch, discovery of intestinal amoebae in case of dysentery, 29, 30, 32 Loschia, 34 Lota vulgaris, see Burbot Lounsbury, life-cycle of Amblyomma hebrceum, 495 Louse disease, historical instances of death from, 711 870 THE ANIMAL PARASITES OF MAN Low, personal experiments with regard to malaria infection^ 158 — treatment of Oriental sore, 628 Lucilia argyrocephala cause of myiasis in French West Africa, G14 — - ccesar, 588 — macellaria, larvae of, causing eye diseases, 721 ■ — in nose, 715, 716 see also Myiasis, nasal on cutaneous surface, 721, 722 penetrating auditory meatus, 721 — nobilis, larvae (maggots) of, discharge from auditory meatus, 588 — sericata, 588 Lumbricosis, typhoid, 650 Lumbricus, Monocystis agilis from seminal vessels of, 130, 132 — teres, see Ascaris lumhricoides, 464 Lund's larva, characters, 593 Lung, abscess of, set up by amoebae, 35 — amoebae found in, 45 — Balantidium coli occurring in, 202 — bilharziasis of, 642, 643 — gangrene of, possible occurrence of Cercomonads in, 62 — invasion by Fasciola gigantica, 245 by Paragoninius ringeri, 251 by Schistosoma hmmatohium, 274 — Trichomonads found in, 56 Lung-fluke disease, geographical distribu- tion, 639 prognosis, 640 symptoms, 639 treatment, 640 Lussana, toxic theory of ancylostome anaemia, 646 Liitz, ascarides in pulmonary artery, 656 — Ceratopogoninoe described by, 580 — experimental infection with Ascaris hrm- hricoides, 465 — favourable effects of expulsion of Ascaridm, 649 — perforative peritonitis due to Ascaris, 656 Lyctocoris campestris, bite of, 541 characters, 541 Lygceidce, characters, 541 Lymphangitis from Filaria bancrofti infec- tion, symptoms, 676 — in filariasis, 401 Lymphatic glands, enlarged, in filariasis, 402 — vessels and glands, destruction without lymphatic obstruction, 401 distribution and connections of, 400, 401 — vessels, invasion by Onchocerca volvulus, 418, 419 Lymphatics, Strongyloides stercoralis in, 755 Lynch, human trichomoniasis, 734 Lynchia, transmitting Haltcridium , 151 Lyperosia, differentiation from Stomoxys, 610 — exiqua, life-history, 610 — irrifans, var. weisii, 610 Macaciis sinicus, inoculation with Leish- mania donovani, 107 MacCallum, " exflagellation," 152 — investigations of Proteosoma and Hal- tcridium in birds, 158 McDonagh, J. E. R., life-cycle of organism of syphilis, 124 MacFadyean and Stockman, Babesia diver- gens, 177 Macfie, Trypanosoma nigeriense, 76 — and Gallagher, treatment of sleeping sickness, 622 Mackenzie, periodicity of larvae of Filaria bancrofti in peripheral blood, 393 Mackie, suggested transmission of relapsing fever by lice, 120 — treatment of Indian kala-azar, 623 Macleayia, characters, 563 MacNeal, see Novy and MacNeal Macrostoma niesnili, 57, 735 Maculae caerulae (taches bleties) due to in- fection by crab louse, 712 Maggots, see under Names of Parasites and Regions of Body — in nose, see Myiasis, nasal Magnesium sulphate in flagellate dysen- tery, 625 Maillard, fatal cases of nasal myiasis, 718 Majochi, case of intertrigo set up by Oxyuris vermicularis , 696 — infection with Demodex folliculorum, 708 Mai de caderas in horses, trypanosomes associated with, 68 trypanosome causing, 96 Malaria, acute, 156 — atypical forms, 634 — campaign against, commencement and progress of, 158 — chronic, 156 — development of parasites of, 159 — diagnosis (pathognomonic signs), 635 — geographical distribution, 155 — historical, 157 — in birds spread by Culex, 158 — - in man, 155 — latent, in children of natives, 158 — masked, 156 — parasites of, 164-170, 633 asexual generation, cultivation in vitro, 170 copulation, 160, 161, 162 exflagellation (footnote), 162 — — gametocytes of, 160, 161, 162 human, development, 159 occurs only in Anopheles, 158, 159 differential characters, 171 species of, 164, 633 — see also Laverania malarioe, Plasmodium malaria?, Plasmodium re- lictnm, Plasmodium vivax macrogametes of, 160, 161, 162 — — merozoites of, 159, 160 — — methods of detecting, 747 microgametes of, 160, 161, 162 movements, discovery of, 157 INDEX 871 Malaria, parasites of, not transmissible to mammals, 159 ookinetes, 160, 161, 163 schizogony of, 161, 172 sporozoites of, 159, 160, 164 penetration of red blood cor- puscles by, 159, 160 sporulation, 160, 161, 163 — pig-mentation of organs, 165 (footnote), 634 — prevention of constipation during, 635 — preventive measures against mosquitoes, 635, 636 against parasite in man, 635, 636 by quinine administration, 636 — prophylaxis, 636, 637 — relief of symptoms, 635 — symptoms, 156, 633 — synonyms, 155, 633 - — tertian, malignant, paroxysms of, 634 1 — treatment by quinine, 635 Malarial fever, quartan, 156 duplex or triplex, appearance of, 167 parasite of, 166 malignant or sub-tertian, parasite of, 167 pernicious symptoms, explanation, 172 quotidian, 156 rhythmical, course of, 155 symptoms, 155, 633 tertian, 156 ■ — simple or spring, parasite of, 164 — — typical, clinical features, 633, 634 Male fern, administration to children, 671, 672 emulsion, injection of, 671 ethereal extract best vermifuge for TcBTiia saginata, 670 dosage and method of adminis- tration, 670, 671 ■ extract of, expulsion of Hymenolepis nana by, 661 in bilharziasis, 643 in expulsion of ancylostomes, 686 — in intestinal myiasis, 728 poisoning, 670, 671 — antidotes to and remedies for, 671 bad effects on vision, 670 Malignant malarial parasites, sporulation, influence of temperature on, 163 stages of, 163 Mallory's bodies, 208 Mamma, tumours of, association of Diocto- phyme gigas with, 431 Mammals, human malarial parasites not transmissible to, 159 — leucocytogregarines in, 154, 155 — r; d blood corpuscles of, Babesia para- sitic in, 154 M:rT, incidental parasites of, 7 — infection with animal trypanosome, 96 — parasites found only in, 6 Mange, see Dog mange Mangold, feeding experiments with Taenia from multilocular echiriococcus, 358 Manguinhosia, characters, 562, 568, 569 Manson, Sir Patrick, development of Paragonimas ringeri, 251 discovery of Sparganum 7nansoni, 317 infection of skin by Filaria per- stans, 378 on Spirochaeta carteri, 631 pathognomonic signs of malaria, 635 — prophylaxis against ancylosto- mia,sis, 685 researches on malaria, 158, 635 studies of filariasis, 391 treatment of Indian kala-azar, 626 ^ — of Oriental sore, 628 Manson, P. T., infected with malaria by infected mosquitoes, 158 Manson ia, 577 Manson's method of administration of atoxyl in sleeping sickness, 622 Manteufel, immunity of Ornithodorus mou- bata against infection with Spirochceta duttoni, 119 Marchiafava, discovery of movements in malarial parasites, 157 Marchoux, amoebic abscesses in liver of experimental cats, 35 — Spirochceta gallinarum, 119 — and Couvy, Leishman granules in Loelaps echidninus, 493 Mareo, Helminthiasis meningitiformis , 649 Margaropus annulatus australis, hosts of, 505 pest to man in Jamaica in larval stage, 505 — characters of, 497 — microplus , 505 Marx, male fern administration, 671 — toxic action of male fern, 670 Marzinovsky, prophylaxis against Pedicu- lus vestimenti, 616 Mastigophora, 28, 50, 760 — aggregation rosettes of, 51 — characters and habitat, 28 Mathis, carriers of Entamoeba histolytica, 40 — diarrhoea due to Lamblia intestinalis , 625 — Lamblia intestinalis , 59, 60 — modification of Novy-MacNeal medium, 744 Maurer's dots, 168, 170, 171 Maxillary sinus, Scolopendra in, 721 Mayer's glychsemalum, 751 — hsemalum, 751 Mbori in dromedaries, 06 Measles, 207 Meat inspection, decrease of cysticerci in pork effected by, 334 Meatus, auditory, larvae of Anthom,yia plu- vialis found in, 584 Rhinosporidium in, 196 maggots of Lucilia nobilis in, 588 872 THE ANIMAL PARASITES OF MAN Meatus, auditory, synonyms, 438 Mecistocirrus, habitat, 438 — morphology, 438 — fordi, morphology, 438, 439 • synonyms, 438 Medullary layer of Cestoda, 289 Megarhinince, characters, 563, 570 Megarhinus, characters, 563, 570 Megnin, development of cestodes, 16 Mehlis, discovery of progeny of Distoma, Typhloccelum flavum-, and Cathoemasia hians, 12 Mehlis' gland secretion in trematodes, 223 Melanoconion, distinguishing characters, 564, 576 — atratus, characters, 576 geographical distribution, 576 Melanolestes abdominalis, 540 — morio, geographical distribution, 540 synonyms, 540 Mello-Leitao, flagellate dysentery in chil- dren, 56, 624 Melnikow-Raswedenkow, development of alveolar echinococcus, 357, 358 Melophagus ovinus (sheep ked), bite of, 611 Crithidia inhabiting, 104 Meningitis, fatal, peenash terminating in, 716 — symptoms of, due to Ascaridce infection, 649 — terminating nasal myiasis fatally, 718 Mense, expulsion of Guinea worm, 676 Mercier, nematodes in human eye, 412 Mercury, benzoate of, in infantile kala- azar, 627 — cream (Lambkin's) in syphilis prevailing in Uganda, 632 — in expulsion of Strongyloides stercoralis , 675 Mermis, 469 — hominis oris, 469 Mermithidce, 469 — characters, 375 Merogregarina, 135 Merogony, 185 Meront, 185 Merozoites of Coccidiidea, 138, 139, 140 — of malarial parasites, 161 Mesenteric vein, superior, tributary of por- tal vein, 272 Mesnil, on Actinomyxidia, 187 — on Hsemosporidia, 742 — on Haplosporidia, 194 — and Ringenbach, cross-immunity ex- periments with trypanosomes, 80 trypanolytic reactions, 80 — see also Laveran and Mesnil Messineo, effects of experimental injection of extracts of Taenia, 648 Metagonimus, 264 — (Yokogawa yokogawai, 264, 753 geographical distribution, 265 habitat, 265 host and intermediate host, 265 life-history, 265 morphology, 264 Metastrongylinae, characters, 432 Metastrongylus, morphology, 432 — apri, hosts of, 433 in man, cases recorded, 433 invasion of air-passages by, 433 morphology, 432 synonyms, 432 Methyl green, 752 Methylene blue in bilharziasis, 643 in flagellate diarrhoea, 625 Metorchiince, 261 — morphology, 232 Metorchis, 261 — conjunctus, organs of, diagram showing, 258 — truncatus , habitat and hosts of, 262 morphology, 261, 262 organs of, diagram showing, 262 Metschnikoff, intestinal parasites in relation to appendicitis, 652, 653 — prophylaxis against oxyuriasis, 697 Meyer, disturbances of vision in male fern poisoning, 670 Mibelli, infection with Demodex follicu- lorum, 708 Mice, experimental infection with herpeto- monads, 103, 104, 112, 737, 738, 739 with Sarcocystis muris, 191 — with SpirochoBta duttoni, 117 — natural herpetomonads in, 738, 739 — occasionally hosts of Hymenolepis diminuta, 326 — Sarcosporidia in, 187 — spherical contracted forms of Tricho- monas intestinalis in, 56 Michelson, case of intertrigo set up by Oxyuris vermicularis , 696 Microgametes of Coccidiidea, 137, 139, 140 Microscope, use of, discoveries of parasites from, 10 Microsporidia, 129, 184 — characters and habitat, 28 — morphology of, 185 — various pathogenic members, 186 Midges, see Chironomidce, Ceratopogoninoe, Psychodidce Miescher's tubes, 187, 188 Mikrofilaria hancrofti, prevalence in blood, prevalence of filarial diseases propor- tionate to,, 400 and Mikroloa lea, distinction be- tween, 398 — diurna, larvae of Loa loa, 412 presence in blood, 412, 414 — perstans, morphology, 416 and M. diurna, simultaneous presence in blood, 414 — philippinensis , 407 — powelli, 407 Microfilariae, periodic, 393, 394 Milk cure in expulsion of Strongyloides stercoralis, 675 Milton, bilharzial vaginitis, 643 — treatment of bilharziasis, 643 Mimomvia, characters, 565 INDEX 873 Miiicliiii on genus Entamceha, 733 — researches on Trypanosoma lewisi, 89-92 — see also Nicoll and Minchin Mineliin and Fantham, on Rhinosporidium kinealyi, 195, 196, 197 Minchin and Woodcock on Trypanosoma noctuoB, 737 Mineral waters in intestinal myiasis, 728 Miners, prophylaxis against ancylostomiasis in, 684 Mines infected with ancylostomes, disinfec- tion of, 685 Miracidia of digenetic trematodes, mor- phology of, 226, 227 Miracidium, germ cells of, 227 Mitchell, treatment of Oriental sore, 628 Mites attacking man, geographical distri- bution of species, 486 — case of so-called dysentery said to be due to, 512 — living endoparasitically in animals and birds, 491 — see also Acarina — see also Arachnoidea Mochlonyx, 565 Moiriez, species of Chorioptes found on man, 521 Moldovan, schizogony in Leucocytozoon ziemanni, 153 Molluscs, spirochsetes in, 114 — fresh-water, round Cairo, cercariae of bilharzia type in, 277 Molluscum contagiosum, 207, 208 Monas pyophila, 62 characters of, 62 Mondiere, perforation of appendix by Ascaris, 655 Monera, 26 Moniez, Aleurobius (Tyroglyphus) farinoB, 511 — on derivation of entozoa, 21 Monkeys, dysentery in, associated with presence of (Esophagostomum apio- stomum, 444 — experimental infection with Spirochmta duttoni, 117 — inoculation experiments with yaws upon, 128 — rabid, spinal ganglia of, cultivation, 210 — Trypanosoma simicB virulent to, 100 Monocystis, hosts of, 134, 135 — agilis from seminal vessels of Lumbri- cus, 130, 132 life-cycle of, 132, 133 Monogenea, canalis vitello-intestinalis, 222 — ova of, deposition, 223, 224 — post-embryonic development in, 224 Monostomum lentis, 244 Monothalamia (testaceous amoebae), charac- ters of, 47 Montgomery, transmission of rinderpest. 742 Moore and Breinl, latent bodies of Try- panosoma gainbiense, 77 Moosbrugger, earth-eating in connection with Trichuris trichiura infection, 679 — trichocephalus anaemia, 651 Moriggia, Glyciphagus cursor, 513 Morkowitin, Oxyuris infection in relation to appendicitis, 653 Morphia, injection of, in relief of griping and straining in amoebic dysentery, 618, 619 Morsasca, trichocephalus anaemia, 651 Moscato, chyluria following infection by Eustrongylus gigas, 682 Mosquito nets, use of, 636 — worm in Trinidad, 598 how destroyed, 598 Mosquitoes, abdomen, 550 — acting as hosts of Filaria bancrofti, 398 — alimentary canal, 550, 551 — anatomical remarks on, 548 — antennae of, 548 — aquatic in larval and pupal stages, 555 — breeding places of, 553, 557 — campaign against, in prevention of malaria, 636 — copulation of, 553 — distinguishing features of Chironomidce (midges) from, 579 — females alone blood-suckers, 552 fertilized in autumn, hibernation of, 555 — first development of malarial parasite in, traced in Plasmodium relictum, 170 — labrum, labium, and hypopharynx, 548, 549 — larvae, food of, 557 living in salt water, 557 position assumed in water, 557 — length of egg, larval and pupal life, 555 — maxillae and mandibles, 548, 549 — ova of, 558 float on water, 559 — • — localities for deposition of, 553 — proboscis of, 548 — pupae of, 558 — spread of malaria in birds by, 158 — systematic remarks on, 548 — typical structure of, diagram showing, 558 — ubiquitous existence of, 555 — see also Culicidm Moth-like appearance of PsychodidcB, 581 Mott, F. W., association of Treponema with general paralysis, 125 Moty, Oxyuris infection in relation to appendicitis, 653 Mouqui, mite attacking man, 486 Mouth, human, cultivation of species of Treponema from, 741 spirochsetes in, 122, 740 — • infection with Oxyuris vermicularis solely through, 469 — maggots in, 721 Mucidus, characters, 563, 571 Mulder, infection with Demodex follicu- lorum, 708 Mules, murrina in, trypanosome causing, 98 — "surra" in, 95 Miiller, D., echinococcus cysts causing urticaria, 651 874 THE ANIMAL PARASITES OF MAN Miiller, J., discovery of Myxosporidia, 181 — O. F., discovery of and views as to cercarise, 12 of origin of tapeworms by, 11 Miiller's psorosperms, 135 Murrina in mules, trypanosome causing", 98 Musca domestica (common house-fly), 586 breeding grounds, destruction of, 586 characters, 585, 586 destruction of, methods, 586 diseases spread by, 586 hibernation as puparia, 586 larvae (maggots) of, characters, 586 life-cycle of, 586 ova of, places where deposited, 586 pupa of, 586 — pattoni, 611 MuscidoB, 584 — African, larvae of, 590 causing myiasis in man (footnote), 590 — blood-sucking, 603 — larvae of, other than Lucilia, in nose, 720 Muscles, encystment of Trichinella spiralis in, 425 — invasion by Trichinella spiralis, 424, 425 — of nematodes, 361 — sarcosporidia in, 191 Muscular system of Hiriidinea, 480 Musgrave, on human intestinal amoebae, 31 — and Clegg's culture media for amoebae, 743 Mussels, fresh-water, spirochaetes of, 114 Mutualists, nature of, 6 Myiasis, 715 — auricular, 615 treatment, 615 — dermatosa oestrosa, 725 — due to Sarcophaga, 589, 590 — externa, 715 methods of treatment recommended by various authors, 719, 720 rare situations of, 723 — gastric, treatment, 728 — human, occurring in mountains of Cen- tral Sahara, 598 — in French West Africa, cause of, 614 — intestinal, 725, 726 chronic, 726 complicated by mucous colitis, 726, 727 diagnosis, 728 irrigation of rectum in, 728 larvae of different species of flies found I in, 728 ! modes of infection, 727 i views of various authors on, 727 prognosis, 728 prophylaxis, 728 symptoms, 726 treatment, 728 and cutaneous, fly causing, 585 — larvae of African Muscidoe causing (footnote), 590 — nasal, 715 cases of, authors reporting, 716, 717 Myiasis, nasal, connection with ozaena, 717, 722, 723 discharge from nose in, 718 due to Sarcophaga, treatment, 723 fatal termination of, 718 from Sarcophaga wohlfahrti, 722, 723 maggots of flies setting up, 588 prophylaxis against, 718 symptoms, 717, 718 treatment, 719 see also Peenash — cestrosa, geographical distribution, 724 prevalent among rural population, 724 rare in man, 724 treatment, 725 Myriapoda parasitic in intestine and nose of man, 483 MyxidiidcB, 184 Myxidium lieberkiihni, 182 Myxobolidce, 184 Myxobolus cyprini, 184 — pfeifferi, 184 — cause of barbel disease, 184 spore formation, 183 — neurohius, 184 — schematic representation of spore of, 182 Myxoedematous form of Brazilian trypano- somiasis, 88 Myxosporidia, 129, 181 — authors adding to knowledge of, 182 describing species causing diseases in fishes, 182 — characters and habitat, 28, 182 — free forms of, 182 — introduction of term of, by Biitschli, 181 — invasion by, causing disease in fishes, 182, 184 — mode of infection, 184 — multinucleate trophozoite of, 182 — plasmotomy, 182 — spore formation, 182, 183 — tissue parasites, 182 Myzomyia, characters, 561, 567 — funesta, breeding places of, 557 Myzorhynchella, characters, 561, 568 Myzorhynchus, characters, 562, 568 Nabarro, on sleeping sickness, 68 — on Spirochmta duttoni, 116 Nagana (tsetse-fly disease), agent of trans- mission, 93 — fatal to horses, asses and dogs, 94 — prevalent among and generally fatal to cattle, 93, 94 --- treatment by arsenic, 94 — trypanosomes in blood of horses suffer- ing from, 68 Nagel, chloroform and syrup of senna in expulsion of ancylostomes, 686 — filmaron in expulsion of ancylostomes, 687 Nagelschmidt, treatment of scabies, 706 INDEX 875 Naphthalene iu evacuation of Oxyuridce, 697 — in intestinal myiasis, 728 Naphthol ointment, dressings of, in head louse infection, 710 (Kaposi's), application in scabies, 707 Nasal cavity, deposition of ova of Oestrus ovis in, 598 see also Myiasis, nasal Ling-uatula parasitic in, 523, 524, 526, 527 — polypus, Bhinosporidium causing, 195- 197 Nason, Ascaris in appendix, causing in- testinal obstruction, 654 Nasse, investigations of Coccidia, 135 Natal, larva of, characters, 591 Nattan-Larrier, cross-immunity experiments with trypanosomes, 80 — Tetramitus mesnili, 57, 624 Natural flagellates of insects, 103, 104, 107, 112, 739 Naunyn, mode of formation of daughter cysts of echinococcus, 352 Neave, S., ulcers set up by invasion by larvae of Cordylobia anthropophaga, 592 Necator, 447 — americanus, 450 geographical distribution, 459 habitat, 459 — — morphology, 457, 458 organs compared with those of Ancylo- stoma duodenale, 458 — characters, 457 — exilidens, characters, 459 habitat, 459 Negri, experimental infection with Sarco- cystis muris, 192 — on Neuroryctes, 208 Negri's bodies, 208, 209 Neligan, Leishmania tropica in dogs, 108 Nemathelminthes, 360 Nematoda, see Nematodes Nematode larvae in blood in cases of pru- ritus, 378 in periosteum of upper jaw in case of gingivitis, 378 Nematodes, anatomy of, 360 — bursa copulatrix of males, 370 — chorion enveloping ova, 371 — classification of, 374 — clearing of, 473 — cutaneous glands, unicellular, 361 — cuticle of, 360 — cutis of, 361 — dermo-muscular layer of, 361 — development of, 17, 371 ' — embryos, 372 — encapsuled forms of, 17 — epithelium of, 360 — excretory canals, anterior, 367 organs, 366, 367 special, lacking in certain genera, 367 — — pore and duct, 367 vesicle, 367 — fixation of, 473 — oflandular stomach of, 363 Nematodes, gubernaculum of male genital apparatus, 369 — hatched from eggs of Sphcerularia, 5 — heterogony in, 372 — hind gut, 363 — ■ infection by, 644 — intestinal caeca, 364 canal, 363 — "isolation tissue," 362, 363, 364 — life spent in intermediate and final host, 18 — marine, ventral gland of (so-called), 367 — mounting head of, 473 — muscles of, 361 — nervous system, 364-366 — observed in human eye, 412 in man, 376 — oesophageal glands, 364 — oesophagus of, 363 — organs of sense lacking in parasitic species, 366 — ova of, 371 conveyance to definite host with inter- mediate host, 373 without intermediate host, 372 detection, 473 developmental capacity, 371, 372 — ovejector, 368 — oviduct, 368 — parasitic and free-living, connection, 20 — preservation and examination of, 473 — rolling of, 473 — seminal receptacle, 368 — sexual organs, 367 female, 367, 368 diagram of, 368 male, 369 diagram of, 368 — small, detection of, 473 — spicules of male genital apparatus, 369 — staining of, 473 — testis of, 369 — " tuft-like " or " phagocytic " organs, 362 — viviparous species, 371 — young, skin diseases due to, in dogs, 378 Nematodirus, habitat, 438 — morphology, 438 Neocellia, characters, 562, 569 Neomyzomyia, characters, 561, 567 Neopsylla, distinctive characters, 545 Neosalvarsan in syphilis, 632 — in yaws, 632 Neosporidia, 129, 181 — characters, 28, 129, 181 Nephrophages sanguinarius, characters and morphology, 490 presence in urine, 490 Nervous system of Cestoda, 289, 290 of Echinorhynchus, 475 of Hirudinea, 481 of Insecta, 530 of nematodes, 364-366 central, effect of Dipylidium caninum on, 649 Neumann, mosquitoes transmitting Plasmo- dium relictum, 170 876 THE ANIMAL PARASITES OF MAN Neumann, podophyllin in expulsion of ancylostomes, 687 — synopsis of genus Ornithodorus, 508 — jbable of species of Argas, 505 Neuritis, optic, following male fern poison- ing, 670 Neuroptera, characters, 531 Neuroryctes, 208 — hydrophobioe, 208 minute granules in, 210 Neurosporidium, 195 — cephalodisci, 195 Newstead, Amblyomma cayennense, 501 — life-cycle of Phlebotomus, 582 ■ — Margaropus anmUatus aiistralis, 505 — means of separating sjDecies of Glossina, 604 Niaibi, mite attacking man, 486 Nicoll, development of cestodes without intermediate host, 17 ■ — and Minchin, cysticercoids in rat fleas, 327, 328 Nicolle, immunity experiments with Leish- mania infantum and L. tropica, 112 — and others, transmission of relapsing fever by lice, 120, 121 Nicollia, 174 — quadrigemina, 174 Nicotiana soap, application in scabies, 707 Nits, methods of getting rid of, from hair, 710 Nitzsch, views as to cercariae, 12 Noc, cultivation of species of amoeba by, 41 — on Lamblia intestinalis , 60, 625 Noguchi, cultivation of parasite of rabies, 210 of Treponema from human mouth, 128, 741 of Treponema pallidum, 125 — method of cultivation of spirochsetes, 123 — Spirochoeta phagedenis, 122 — Treponema calligyrum, 126 — and Cohen, cultivation of so-called trachoma bodies, 210 — and Moore, association of Treponema with general paralysis, 125 Noller, development of Trypanosoma lewisi in dog flea (Ctenocephalus canis), 90, 92 — method of controlling fleas during ex- periments, 93 Nordmann, von, discovery of miracidia of flukes, 12 Normand, association of amoebae with colitis, 30 of Strongyloides stercoralis with diar- rhoea, 380 Norway itch (scabies norvegica), 520 Nose, ascarides in, 690 — dipterous larvae in, in enormous num- bers, 716, 717 — discharge from, in nasal myiasis, 718 — human, Myriapoda parasitic in, 483 — larvae of Hypoderma bovis in, 724 of Lucilia macellaria in, 715, 716 see also Myiasis, nasal of Oxyuris vermicularis in, 469 — leeches in, 700, 701 Nose, leeches in, causing epistaxis, 701 — maggots in, 588 — Oxyiiridoi migrating into, 695, 696 Nosema apis, 184 - — — life-cycle of, 185 pansporoblast and sporoblast >f, 185 planont of, 185 — bombycis, 184 spores of, 186 Notoedres cati, 521 — cuniculi, 521 — notoedres, 521 Novy and MacNeal, artificial cultivation of trypanosomes, 69 Novy -MacNeal medium, 744 Mathis's modification, 744 Novy-MacNeal-Nicolle medium, best for cul- tivation of Leishmania infantum, 109 for cultivation of Leishmania tropica, 108 formula, 744 Nut-pickers affected by Leptus aiitumnalis (footnote), 485 Nuttall, Spirochoita marchouxi, 119 — Piroplasmidce, 174 Nuttall and Hadwen, trypan-blue in treat- ment of piroplasmosis, 178 — and others, nuclear phenomena of Babesia canis, 176 Theileria parva, 179 Nuttallia, characters, 174 — equi, cause of equine piroplasmosis, 174, 178 life-cycle in red blood corpuscles, 173 — herpestidis , 174 Nycteribiidce, 611 Nyctotherus, 204 Nyctotherus africanus, 206 — faba, 205 — — morphology, 205 — giganteus, 205, 206 morphology, 205 Nyssorhynchus, characters, 562, 569 0. Occiput, abscess of, liver-fluke in, 243 Ochindundu, bite of, 541 — characters of, 541 Ochromyia anthropophaga , larvae of, charac- ters (footnote), 590, 591 — — — hosts of, 590 (Edema following bite of Argas reflexus,o06 Oerley, induction of facultative parasitism of Bhabditis pellio, 377 (Esophageal glands of nematodes, 364 CEsophagostomeoe, characters, 439 (Esophagostomum, morphology, 441 — apiostomum, habitat and host of, 444 morphology, 444 — brumpti, habitat, 441 morphology, 441 — stephanostomum, habitat, 444 var. thomasi, morphology, 442, 443, 444 (Esophagus of Hirudinea, 480 — of nematodes, 363 INDEX 877 CEsophagus, trichomonads in, 00 Oestrijoe (wabble flies), boils produced by, 725 — cavicolous, 598 — cutaneous, 595 — flight time of, 725 — gastricolous, 599 infection by, 729 see also Creeping disease — hosts of, 594 — larvae of, occurrence in man rare, 724 — method of depositing ova on skin of man, 725 Oestrus {Cephalomyia) oris, 598 geographical distribution, 598 ova of, deposition in nasal cavity, 598 Oil, injections of, in nasal myiasis, 719 Ointment, application in scabies, 706 Oken, views as to origin of cercariae, 12 Oleum chenopodii in ancylostomiasis, 754 Oligosporulea, 195 Oligotricha, 29 Oliver, artificial infection of human beings M^ith Cysticercus bovis, 340 Omentum, abscess of, with Ascaris ova in pus, 657 Onii, diagnostic sign, of presence of Spar- ganum mansoni in body, 659 Onchocerca, 417 — volvulus, 417 — — distribution in West Africa, 419 — - — invading lymphatic vessels, 418, 419 invasion in man associated with formation of tumours, 418 — — measurements, 755 morphology, 417, 418 Onchocercinw, 417 Oncospheres (embryos) of tapeworms, 298, 299 certain species of animals neces- sary for, 299 development into plerocercoid, 300 further develojjment must take place in suitable animals, 299 — migration in body, 302 O'Neil, filaria infection of skin, 378 Onions, Anguillulina putrefaciens living in, 379 Oocysts of Cuccidiidea, 141 — of malarial parasites, 163 Opalina, 198, 207 — ranarum, 207 Ophryocystis, 135 Op isthorchiidcB , morphology, 232 Opisthorchiinw , 252 Opisthorchis, 252 — felineus, development, 254 — — geographical distribution, 252 hosts of, 252 intermediate, 254 mode of infection by, 254 morphology, 252 synonyms, 252 — pseudo felineus, anatomy of, diagram illustrating, 254 Opisthorchis sp., habitat, 753 morphology, 753 Opisthotonos, disappearance after exjiulsion of Ascaridce, 649 Oppenheim, maculae caerulae (taches bleues) due to infection by crab louse, 7l2 — treatment of crab louse infection, 712 Oppila^ao, synonym of Brazilian trypaiioso- miasis, 87 Oral cavity, cancer of, association of Ent- amoeba buccalis with, 43 trichomonads in, 55, 56 Orbit, cysticercus of, 664 Orbital cavity, Pycnosoma maggots invad- ing, 588 Orchitis from Filaria bancrojti infection, 677 Oribates sp., 489 Oriental sore, cause of, 107, 627 experimental production, 109 geographical distribution, 108 germ of, possible carrier, 580 immunity to, procured by inocula- tion, 108 objection to name, 107 occurrence in dogs, 108 parasite producing, 107, 627 pathology of, 627 — — preventive measures, 628 relation of infantile kala-azar to, shown experimentally, 109 sites of occurrence on body, 108 transmission of, bugs possibly con- nected with, 108, 536 treatment, 628 Ornithodorus coriaceus, geographical distri- bution, 509 — megnini, characters, 510 ears of hosts infested by, 510 geographical distribution, 510 hosts of, 510 — moubata, carrier of African tick fever^ 116, 496 of Filaria perstans, 508 of spirochaete of relapsing fevei*, 508 geographical distribution, 509 immunity against infection with Spirochceta duttoni, 119 length of life apart from host, 495 Malpighian secretion passed by, sig- nificance, 117 transmission of Spirochceta duttoni by, 116 — savignyi, 509 geographical distribution, 509 transmitting Spirochceta duttoni, 739 — synopsis of genus, 508 — talaje, 119 bite of, 509 geographical distribution, 509 — tholozani, geographical distribution, 51Q — turicata, 119 bite of, effects, 509 Omithomyia lagopodis, bite of, 611 Orthoptera, characters, 531 878 THE ANIMAL PARASITES OF MAN Otter, Brazilian, liost of Paragonimus rudis, 251 Ova, transmission of intestinal worms by, 11 — see also under Names of parasites Owen, Trichina spiralis, 423 Ox, liver of, Echinococeus multilocularis in, 357 — Sarcocystis hlanchardi from, 190 — gad fly (Tabanus hovinus), 601 Oxazine producing blepharoplastless try- panosomes, 101 Oxen, amount of prevalence of Cysticercus bovis in, 340, 341 — echinococci in, 346 — how infected with Para7nphistomum cervi, 226 Oxygen necessary in cultivation of spiro- chaetes, 123 Oxyuriasis, diagnosis, 696 — dysentery followed by recovery from, 698 — in children, 695 treatment, 697, 698 — prophylaxis against, 697 — treatment by drugs and purgatives, 697 local, 697 Oxyuridce, 467 — migrating into nose, 695, 696 — morphology, 375 — relationship to appendicitis, 698 Oxyuris, 467 — ambigua, 469 — compar, 469 — curvula, 469 — encapsuled in female pelvis, 657 — in appendix, 654, 655 — infection in relation to appendicitis, 653 — invading peritoneal cavity, 657 — lacks intermediate host, 21 — mastigodes, 469 — poculum, 469 — tenuicauda, 469 — toxic action of, 651 vermicularis, association with appendi- citis and typhlitis, 467 development, 468 direct, 469 experimental self-infection with, 469 habitat, 467 infection by, 694 with, mode of, 469 . — larvae of, found in nose, 469 life-history of, 467, 468, 469 males rarely -met with in faeces, 468 migration from intestine, lesions and symptoms of irritation set up by, 694, 695 of, in and from intestinal tract, 467 morphology, 467 ova of, where deposited, 467 supposed origin of, 11 Oyster, spirochaete of, 114 Ozaena, connection of nasal myiasis with, 717, 722, 723 P. Page, case of escape, of ascarides from abdominal operation wound, 654, 655 Paget, observation of encapsuled Trichi- nellae, 423 Pallas, on transmission of intestinal worms, 11 Panama, larvicide used at, in campaign against mosquitoes, 636 — Canal, Stegomyia fasciata source of danger to, 574 Pani-ghao, skin affection set up by pene- tration of larvae of Ancylostomum duo- denale, 455 Panoplites, 577 Pansporoblast, 183, 186 Papataci fever, carrying agent of, 582 Pappenheim's panchrome mixture, 751 Paraboloid condenser, 747 Paraffin, embedding in, for sectioning tissue parasitized by protozoa, 749 Paragonimiasis, 639 — affecting regions other than lung, 639 — prophylaxis, 640 — see also Lung-fluke disease Paragonimus, morphology, 249 — compactus, host of, 251 — kellicotti, hosts of, 250 spines of, 251 — ringeri (lung-fluke), 639 development, 251 diseases caused by, 251 habitat, 251 internal organs, diagram illustrating, 250 morphology, 249, 250 sites of body in which found post ■mortem, 639 spines of, 251 synonyms, 249 — rudis, host of, 251 — westermannii, host of, 250 — — morphology, diagram illustrating, 250 spines of, 251 Paraguay, supposed mode of transmission of dermo-mucosal leishmaniasis in, 739 Paralysis due to tick bites, 613 geographical distribution, 613 — of dourine, 97 Paramoeba, 44 — hominis, 45, 734 characters of, 45 now called Craigia hominis, 45, 734 ParamphistomidoB, 231, 234 ParamphistomincB, 231 Paramphistomum cervi, method of infection of oxen by, 226 Paraplasma, 180 — doubt as to organismal nature, 180 — occurs naturally in guinea-pigs, 180 — flavigenum possibly associated with yel- low fever, 180 morphology, 180 — subflavigenum, 180 Parasites, definition, 1 INDEX 87$ Parasites, derivation of, 19 — diagnosis of presence of, 10 — discoveries from use of microscope, 10 — great fertility of, 5 — hereditary transmission of, 19 — human, Opisthorchis felineus most fre- quently found at autopsies at Tomsk, 253 — incidental, 6 human, 7 — influence on host, 8 — invading many hosts, 6 — limited to closely related hosts, 6 to one species of host, 6 — migrations in host, injuries set up by, 9 — movements of, disorders set up by, 9 — occasional (temporary), 1 — origin of, 10 — permanent, bodily changes in, 3 clasping and clinging organs in, 4 classes of, 2 hermaphroditism in, 4 loss of organs in, 3 (stationary), 1, 2 — transference from one host to another, 7 Parasitic life, advantages of, 20 Parasitism, facultative, 7 Pariah dogs, liver of, habitat of Paropis- thorchis caninus, 257 North-west Provinces, India, percent- age infected with Paropisth orchis cani- nus, 257 Paropisthorchis, 255 — caninus, genital pore, 255 habitat, 257 morphology, 255 seminal vesicle, 257 synonyms, 255 uterine coils, 257 vitellaria, 255 Partridges, Plasmodium relictum cause of fatal disease in, 170 Pasquale, see Kruse and Pasquale Pasteur, L., researches on silkworm disease, 184 Patterson, maggots of Pycnosoma removed from orbital cavity, 588 Patton, genus Crithidia, 104 — Herpetomonas muscce doniesticce , 102 — Piroplasma gihsoni, 177 — places Leishman-Donovan body in genus Herpetomonas, 107 — probable transmission of Leishmania, 107, 108 — and Cragg, life-history of Lyperosia exigua, 610 Peacock, observation of encapsuled Trichi- nellae, 423 Pebrine bodies or Nosema bombycis of Arthropoda, 184 Pediculida> (lice), characters, 532 Pediculoides ventricosus, effects on man, 469 — — morphology, 489 — shape of preg-nant female, 489 — — synonyms, 489 Pediculis capitis (head louse), characters and morphology, 532, 533 Pediculis_ capitis, geographical distribu- tion, 533 habitat, 533 infection by, 709 — causing eczema, 709, 710 diagnosis, 710 greater prevalence among females, 709, 710 remarkable instances, 710 resulting in blepharitis and con- junctivitis, 710 in plica polonica, 710 treatment, 710 mouth parts of, 533 ova of, 533 prophylaxis against, 615, 616 — vestimenti (clothes louse), characters, 533 habitat, 533 infection by, 710 lesions and symptoms following, 711 pest among soldiers during cam- paigns, 533 prophylaxis against, 615, 616 transmission of relapsing fever by, 120, 630 Peenash (nasal myiasis), 588, 715 — ending in fatal meningitis, 716 Peiper, cause of ancylostome anaemia, 648 Pelagutti, treatment of cutaneous and mus- cular cysticerci, 663 Pelletierinum as vermifuge, 673 Pelvic and abdominal organs, blood-supply of, as illustrating distribution of Schis- tosoma haematobium in body, 272 Pelvis, female, Oxyuris encapsuled in, 657 Pentastoma armillatus, hosts of, 528 — denticulatum, former name of larval stage of Linguatula rhinaria, 525, 526 — moniliformis, hosts and habitat of, 528 synonyms (footnote), 528 PentastomidoB, references to, 528 Pentateuch, " fiery serpents " mentioned in, probable identification, 386 Pentatrichomonas bengalensis, 624, 735 Pereira, case of chorea cured after expul- sion of Taenia, 648 Perinseum, tumours of, association of Dioc- tophyme gigas with, 431 Peritoneal cavity, Ascaris in, 656 Oxyuris invading, 657 Peritonitis, perforative, due to Ascaris, 656 Peritricha, 29, 200 Perroncito, artificial infection of human beings with Cysticercus bovis, 340 — infection with Lamblia intestinalis , 60 Persia, importation of African tick fever into, 613 Persian insect powder infusion in intes- tinal myiasis, 728 Peru oil, application in scabies, 707 Petrie, treatment of bilharziasis, 643 Petroleum as larvicide in campaign against mosquitoes, 636 — dressings of, in head louse infection, 710 — in crab louse infection, 712 — and benzine in crab louse infection, 712 88o THE ANIMAL PARASITES OF MAN Pfeiffer, L., pathogenicity of Coccidia, 136 Pfeiffer, R., Coccidia, 136 Pharynx, ascarides invading, 691 — invasion and infection in man by Fasciola hepatica, 242 — leeches in, 699 — of Hirudinea, 480 Philaematomyia, position of genus, 611 — insignis, 611 Philippine Islands, experiments on amoebae in, 618 Philips, eucalyptus oil in expulsion of ancylostomes, 687 Phillips, L. P., on Musgrave and Clegg's medium, 743 treatment of balantidian dysentery, 637 Phlehotomince, characters, 581 Phlebotomus, blood-sucking, 581 — characters, 581 — duboscii, 582 — intermedins, 582 — geographical distribution, 581 of species, 582 — larvae of, habitat, 582 — longipalpis, 582 — papatacii, 581, 582 — squamiventris , 582 Phonergates hicoloripes, 541 Phoniomyia, characters, 565 Phora rufipes, 589 larvae (maggots) of, habitat, 583 Phoridce, characters, 582 Phthiriasis, agents of, 533 Phthirius inguinalis (crab louse), charac- ters, 534 habitat, 534 infection by, diagnosis, 712 how eifected, 711 lesions and symptoms following, 711, 712 sites of body affected, 711 treatment, 712 rapid reproduction of, 534 — pubis, prophylaxis against, 616 Phthisis, filaria associated with, 408 Physaloptera, habitat and hosts of species, 460 — caucasica, morphology, 461 — mordens, geographical distribution, 461 habitat and host, 461 morphology, 461, 462 — morphology, 460 Physalopteridce, 375, 460 Phytoparasites, 1 Pierantoni, Agamofilaria labialis, 407 Pig concerned in transmission of Balanti- dium coli, 202 — development of Trichinella spiralis in, 426, 427 — domestic, normal host of Cysticercus cel- luloscp, 332 — echinococci in, 346 — geoQ-raphical distribution of Tmnia solium corresponds with that of, 334 — host of Paragonimus kellicotti, 250 Pig, intestine of, Fasciolopsis buski in, 246 — Metastrongylus apri in, 433 — organs infected with echinococcus, per- centage of frequency, 347 — rectum of, Balantidium coli present in, 202 — Sarcocystis miescheriana in, 190 — Sarcosporidia in, 187 — trichinous, proportion to healthy, in Prussia, 429, 430 — Trypanosoma simios virulent to, 100 Pigeon lofts inhabited by Argas rejlexus, 506 Piophila casei, characters, 583 larvae of, found in faeces, 583 in nose, 720 Piroplasma, 172, 173, 174 — • see Babesia — gibsoni, 111 — hosts of, 173, 174 PiroplasmidoB, 172, 742 — genera of, 174 Piroplasmosis, treatment of, 178 by trypan-blue, 178 symptoms of, 178 transmission by ticks from recovered to uninfected animals, 178 Placobdella, 482 — catenigera, geographical distribution, 482 Plague, fleas carriers of, 543, 547 Planont, 185 Plants, flagellosis of, possible connection with leishmaniasis, 739 Plasmodium, 151, 742 — falciparum, see Laverania malarias — malaricB, development in red corpuscles of man, asexual stage, 166 distinctive characters, 167 lesions set up by, not marked, 634 parasite of quartan malaria, 166, 633 pigment granules of, 166, 167 schizogony of, 166 synonyms, 166 trophozoites of, differ from those of tertian parasite, 166 — or haemamoeba type of Hoemosporidia includes malarial parasites of man and birds, 151 — relictum, first development of malarial parasite in mosquito traced in, 170 hosts of, 170 mosquitoes transmitting, 170 stages in life-history, 170 synonyms, 170 — species, differential table of, 171 — tenue, 170 — vivax, agent of simple tertian malarial fever, 164, 633 cultivation of, clumping not observed in, 172 number of spores produced, 172 — — development in red blood corpuscles of man, 160, 164, 165 — — — of ''Polymitus," 160, 165 time occupied by, 165 distinctive characters of, 166 INDEX 88l Plasniociium rivax, lesions set up by, uot marked, 634 life-cycle of, 160, 164 merozoites of, 165 migration, 165 — — micro- and macrogametocytes of, 165 — 23igment granules, 165 small variety, 166 — — •' stippling," 165 synonyms, 164 Platylielminthes (flat worms), 211 — central nervous system of, 211 — classification, 212 — - definition, 211 — diseases caused by, 638 — excretory apparatus, 211 — hermapliroditic, 211 — integument of body of, 211 — method of reproduction, 211 — morphology, 211 Plerocercoid, definition of, 301 Plerocercus, definition of, 301 Pleurae, invasion by Faragonimus ringeri, 251 Plica polonica due to head louse infection, 709, 710 treatment, 710 Plimmer, H. G., treatment of sleeping sick- ness with antimony, 623 and Bradford, Sir J. Eose, Trypano- soma brucei, 93 Pliny, Ascaris lumhricoides known to, 464 Pneumocystis carinii, 90 Pneumocysts in rats, 90 Pocock, geographical distribution of Orni- thodorus mouhata, 508, 509 Podophyllin in expulsion of ancylostomes, 687 Polar capsule, 181, 183, 184, 186 — filament, 183, 184, 186 Polecat, intestine of, Isospora higernina parasitic in, 149 Poliomyelitis acuta, possible rhizopods in, 46 — carrier of, 610 — epidemic, insects transmitting, 612 — virus of, 536 carried by house-fly, 586 Pollack, invasion by Loa loa, 678 Polymastigina, 52 Polymitus of Plasmodium vivax, 160, 165 — form of malarial parasites (footnote), 162 Polypus, nasal, caused by Rhinospori- dium- kinealyi, 195, 196 Polysporea, 182, 184 Polysporulea, 195 Polystoynum integerrimum, organs of, 218 Ponds, mosquitoes depositing ova in, 553 Pork, cysticerci in, cause of decrease, 334 — eating of, cause of trichinosis, 423 means of infecting man with cysti- cerci, 334 — inspection of, in prophylaxis against trichinosis, 429 Porocephalus, 523 — armillatus , 527 synonyms (footnote), 528 Porocephalus constrictus, characters, 526 hosts of, 526, 527 organs of body invaded by, 526, 527 synonyms, 526 Port Natal sickness (Cape ailment), 488 Portal vein and liver. Schistosoma hcema- tobium most easily found post mortem in, 273 and vena cava, communication be- tween, how formed, 272 tributaries of, as illustrating distri- bution of Schistosoma hcematobium in body, 272 Porter, A., Crithidia pulicis. 111 generic differences among insect flagellates, 103 (fig. 49) Herpetomonas muscm domesticre, 102 Leucocytogregarina, 154 Theileria parva, 179 — see also Fantham and Porter Portschinsky, deposition of ova of Oestrus oris, 598 — method of destroying Tabanidce, 601 Posner, case of amoebae in urine, 46 Posselt, cutaneous tumours due to cysti- cerci, 662 — reasons for distinction of multilocular from hydatid or unilocular echino- coccus, 358 Post-flagellate stage in herpetomonads, 103 in Crithidia, 104 Potassium iodide in treatment of cutaneous and muscular cysticerci, 663 — permanganate, application in Oriental sore, 628 Pou d'agouti, mite attacking man, 486 Poultry, fatal epizootics among, due to Eimeria avium, 142 Poultrymen attacked with Dermanyssus gallince, 493 Poupee-Desportes, Guinea worm infection, 676 Powell, method of destruction of Sarco- phaga larvae, 723 Predtetschensky, expulsion of Hymenolepis nana, 661, 662 Pre-flagellate stage in herpetomonads, 103 in Crithidia, 104 Price, Dodds, method of prevention of Indian kala-azar, 627, 739 Prima, fatal case of myiasis externa, 716 Privies, disinfection of, as prophylactic against ancylostomiasis, 685 Proflagellata, 115 Proskauer, case of Oxyuridm in nose, 696 Prostomata, 230 Protargol in balantidian dysentery, 637 Proteid destruction in ancylostomiasis, 647 — metabolism in anaemia, 645 Proteosoma, spread of malaria in birds by, 158 Protista defined, 29 — spirochsetes classed among, 115 Protomonadina, 52, 60 — classification, 60, 61 882 THE ANIMAL PARASITES OF MAN Protozoa, 25, 756 — alternation of generations in, 27 — blood-inhabiting, examination of, 747 — characters, 25 — chromodial apparatus of, 26 — classification, 27 — clinical and therapeutical notes relating to, 617 — cytological details, method of examin- ing, 748 — definition of, 25 — digestive apparatus, 26 — ectoplasm and endoplasm of, 25, 26 — encystment of, 27 — examination, methods for, 745, 746 — food of, 26 — genera of, precise definition sometimes impossible, 733 — hereditary transmission of (footnote), 19 — intra vitam staining of fresh prepara- tions, 746 — nucleus of, 26, 27 — organellae, 29 — parasitic in blood, culture media Ltr, 744 — propagation of, 27 — sectioning tissue parasitized by, 749 — or bacteria, question whether spiro- chsetes to be classed among, 115 Protozoology, notes on technique, 745-752 — recent researches in, 733 Prowazek, balantidian dysentery, 637 — Chlamydozoa, 207 — Entamceha biltschlii, &c., 34 buccalis, 43 — Herpetomonas muscce domesticm, 102 — lamblial diarrhoea, 625 — variety of Trichomonas intestinalis in- habiting oral cavity, 56 — and Aragao, filtration experiments with chlamydozoal granules, 209 Prowazekia, characters of, 63 — asiatica, 65 — cruzi, characters, 66 — javanensis, characters, 66 — parva, 66 — urinaria, 63 characters, 63 flagellate stage, 64 in cultures associated with bacteria, 65 synonyms, 63, 64 — weinbergi, characters, 66 Prowazek's bodies, 208 Pruner, Porocephalus constrictus, 526, 527 Pruritus ani due to escape of ascarides, 688 set up by migration of Oxyuris ver- rnicularis, 695 — nematode larvae in blood associated with, 378 Prussia, oxen infected with Cysticercus bovis in, 341 — percentage of pigs infected with cysti- cerci in, 334 — proportion of trichinous to healthy pigs in, 429, 430 Pseudo-helminthes, 8 Pseudomeningitis due to Ascaridoe infec- tion, 649, 650 Pseudo-myxoedematous form of Brazilian trypanosomiasis, 88 Pseudonavicellae, 129, 1.30 — amoeboid germs in, 130 Pseudoneuroptera, characters, 531 Pseudo-parasites, 6, 8 Pseudophyllidea, morphology, 308 Pseudotaeniorhynchus, 576 — characters, 564 Psorophora, characters, 563, 571 — ovum of, 557, 558 Psoroptes, characters, 517 Psorospermia of Arthi'opoda , 184 Psorosperms (Myxosporidia), discovery of, 181 — egg-shaped, former name for Coccidia, 135 Psychodidce (owl midges), moth-like appear- ance of, 581 PsychodinoR, characters, 581 Pterocephalus, host of, 135 Pterygota, classification, 531 Pulex, distinctive characters, 545 — irritans (human flea), bite of, effects, 714 treatment, 714 carrier of plague bacillus, 543 characters, 545 larva of, 546 raaj transmit Trypanosoma lewisi, 92 — pallipes, 548 — serraticeps (dog flea), 546 Pulicidw (true fleas), characters, 543 — classification of genera, 545 Pulmonary artery, ascarides in, 656 Pumpkin seeds as vermifuge, 673 Pupipara or Eproboscidce, blood-sucking, 611 Purgatives for expulsion of ascarides, 693 — in arrest of development of trichinosis, 681 Pustules arising from clothes louse infec- tion, 711 Putnam, Oxyuris in appendix, 654 Pycnosoma, characters of, 588 — and Chrysomyia, distinguishing fea- tures, 588 — putorium, spread of amoebic dysentery by, 614 Pyelitis following invasion by Eustrongylus gig as, 682 Pygiopsylla, distinctive characters, 545 Pyorrhoea alveolaris, association of Ent- amoeba buccalis with, 43, 734 of species of Treponema with, 128 treatment, 620 Pyretophorus, characters, 561, 567 Pyronin producing blepharoplastless try- panosomes (T. brucei), 101 Quincke and Eoos, species of amoebae named by, 31 Quinine, administration as preventive against malaria, 636 INDEX 883 Quinine, administration in malaria, 635 dosage, 635 methods of administration, 635 time for, 635 treatment by, 635 — in Indian kala-azar, 626 — lotion, irrigation of lower bowel with, in gangrenous dysentery, 619 Rabbit, development of hydatid scolices in, 353 — host of Eimeria stiedce, 145 — intestinal coccidiosis in, 145, 147 - — intestine of, section infected by Eimeria stiedce, 145 — kidney of, use in cultivation of Tre- ponema pallidum, 126 — liver of, section through nodule infected by Eimeria stiedce, 147 — Sarcosporidia in, 187 Eabies, parasite of, cultivation, 210 Eadiolaria, characters and habitat, 28 Radium treatment of Oriental sore, 628 Railliet, method of infection with Trichuris depressiuscula, 420 Rainey's corpuscles, 189 Rain-water barrels, mosquitoes depositing ova in, 553, 557 Ramstedt, Oxyuris infection in relation to appendicitis, 653 Ranken, treatment of sleeping sickness with antimony, 623 Basahus bigiittatus, bite of, 540 — — geographical distribution, 540 — — synonyms, 540 Rat attacked by Dermatophilus (Sarco- psylla) penetrans, 613 — blood of, transference of Trypanosoma brucei from, to blood of snake, 102 — blood parasite, see Trypanosoma lewisi — gut and caecum of. Trichomonas from, 735 — infection with Trichinella, method of, 427 with Trichinella spiralis in slaughter- houses and knackers' yards, 427 with Trypanosoma lewisi, mode of, 92, 93 — muscles of, invaded by Trichinella spiralis, 425 — normal host of Trichinella spiralis, 427 — pneumocysts in, 90 — sewer and black, hosts of Hymenolepis diminuta, 326 — flea (Ceratophyllus fasciatus), cysticer- coid of Hymenolepis diminuta found in, 327, 328 host of rat trypanosome, 88, 90, 543 larval stages of Hymenolepis murina occurring in, 17 see also Ceratophyllus fasciatus — Trypanosoma lewisi in, 88 Rectum, administration of quinine by, in malaria, 635 — bilharziasis of, 642 treatment, 644 — irrigation of, in intestinal myiasis, 728 — means of access of Schistosoma hasma- tobium to, 272 — pathological changes in, due to Schisto- soma hcematobium, 274, 275 — plexus formed in, by superior haemor- rhoidal veins, 272 Redi, origin of flesh maggots, 10 Redise of trematodes, 225, 226, 227, 228 Reduviidce, bites of, 537 — characters of, 537 — geographical distribution, 537 Reduvius personatus, bite of, sometimes fatal, 539 geographical distribution, 539 Red-water fever, European, in cattle, cause of, 177 Reighardia, 523 Relapsing fever, 120, 629 African, cause of, 116, 630 incubation period, 630 prophylaxis, 631 symptoms, 630, 631 — treatment, 631 American, 630 Asiatic, mortality from, 631 prophylactic measures, 631 symptoms, 631 treatment, 631 complications, 630 East African, cause of, 122 European, agent of, 122, 629 incubation period, 630 prophylaxis, 630 symptoms, 630 treatment, 630 Indian, cause of, 122 North African, prophylactic measures, 631 symptoms, 631 — treatment, 631 and Egyptian, cause of, 122, 631 — — prophylactic measures, 630 spirochaetes causing, 115, 120, 122, 508 transmission by lice, 120, 630 by ticks, 117, 630, 631 treatment, 630 Remak, investigations of Coccidia, 135 Reptiles, haemogregarines in, 153, 154 — Sarcosporidia in, 187 Resorcin ointment, application in creeping disease, 732 Respiration, organs of, in Insecta, 530 Retinal haemorrhages in ancylostome anaemia, 646 Reyher, bothriocephalus anaemia, 644, 645 Rhabdites in gastric fluid obtained by lavage, 378 — mellio, presence in vagina, 377 — niellyi, 378 mode of infection in man, 378 884 THE ANIMAL PARASITES OF MAN Bhahditis pellio, induction of facultative parasitism, 377 morphology, 377 synonyms, 377 Ehabdonema, alternation of parasitic and free-living generations, 20 — life-history of, 19 — propagation of, parasitic generation during free life, 18 Rheins, case of Oxyuridce in nose, 696 Rhinosporidium, 195 — hosts of, 197 — in conjunctival polypus, 197 — in external auditory meatus, 196 — in horses, 197 — in nasal polypus, 195 — in papilloma of penis, 197 — kinealyi (or seeberi), 195, 197 causal agent of a nasal polypus, 195, 196 cysts of, 196 geographical distribution, 195, 196 pansporoblasts of, 196 trophozoites of, 196 tumours produced by, 197 Rhipicentor, characters of, 497 BhipicephalcB , characters of, 496, 497 Rhipicephalus, characters of, 497 — species of, transmitting Theileria parva, 179 — annulatus, carrier of Texas fever in cattle, 494 ■ moulting of, 496 — appendiculatus and B. simus, carriers of Rhodesian fever in cattle, 494 — bursa, trasmitting agent of Babesia bovis, 177 — sanguineus, geographical distribution, 505 hosts of, 505 synonyms, 505 — — transmission of leucocytogregarine from dog to dog by, 155 transmitting agent of Babesia canis, 177 Bhizoglyphii, characters and habitat, 514 Bhizoglyphus parasiticus , characters, 514, 515 skin disease produced by, 514 Rhizopods, flagella occurring among, 52 — possible association with poliomyelitis acuta, 47 Rhodesian fever in cattle, carriers of, 494 Bhodinus prolixus, bite of, 541 geographical distribution, 542 Bhyncobdellidce, 482 Rhyncobothrium, scolices of, 305 Bhyncota, see Hemiptera Bicinidm classed among mutualists, 6 Riley, see Walsh and Biley Rinderpest and coccidiosis, 741 — method of transmission, 742 River fever set up by kedani mite in Japan, 487 Rivolta, experimental infection with Cocci- dia, 136 — on Sarcocystis lindemanni, 193 Robertson, Miss, development of Trypano- soma gambiense, 74, 75 in Glossina palpalis, 74, 75 forms of Trypanosoma gambiense, 73, 737 Rocky Mountain spotted fdver, carrier of, 496 tick fever, carrier of, 503 mortality, 504 Rodenwaldt, distribution of larvae of Filaria immitis in body, 393 — periodicity of larvae of Filaria bancrofti in peripheral blood, 393 Rogers, Sir L., agent transmitting kala- azar, 713 cultivation of Leishmania dono- vani, 105, 106 places Leishmann-Donovan body in genus Herpetomonas, 107 transmission of surra by Chrysops, 601 treatment of amoebic dysentery, 618 treatment of Indian kala-azar, 626 of pyorrhoea alveolaris, 620 Rokitansky, perforation of intestine by Ascaris, 655 Romani, agglutinating haemolytic action of serum of ancylostome patients, 648 Romanowsky stain, 749 slightly modified, formula of, 750 underlying principle of, 750 Roos, presence of cercomonads in gan- grenous lung, 62 — and Harris, penetration of intestinal blood-vessels by amoebae, 36 Rosenquist, proteid metabolism in anaemia, 645 Ross, E. H., Treponema pallidum, 124 Ross, Sir Ronald, campaign against mos- quitoes in prevention of malaria, 636 development of malarial parasite in mosquito traced in Plasmodium relic- turn by, 171 discovery of transmission of malarial parasites by mosquito, 158 " Prevention of Malaria," 617, 633 relapses in malarial fever, 161, 162 trichomonads and cercomonads, 56 and Thomson, D., cyclical varia- tion of trypanosomes in blood, 78 — method of determining number of trypanosomes in blood, 747, 748 Rossia, characters, 568 Rossi^lla, morphology, 174 — rossi, 174 Rostellum of Cestoda, 289 Rothschild, classification of genera of Puli- cidcp, 545 Roubaud, cause of myiasis in French West Africa, 614 — life-histo-v of Cordylobia nnthropo- vhaga, 614 — Pycnosoma putorium, 614 INDEX 885 Eovelli, larval stage of Hymenolepis diminuta, 327 Row, experimental production of Oriental sore, 109 — treatment of Oriental sore, 628 Rudolphi, origin of helminthes, 12 E-uffer, lesions produced by Oxyuris vermi- cularis, 695 Buncliiomyia, characters, 565 S. Sabadill vinegar, lotions of, in head louse infection, 710 Sabatier on change of hosts, 21 Sabethes, characters, 565 Sabethoides, characters, 565 Sachs, treatment of scabies, 707 Sack, treatment of scabies, 707 Sahara, Central, human myiasis occurring in mountains of, 598 St. Artault, Trichomonas pulmonalis, 56 Saline solution, physiological, lavages of, in myiasis, 719 Salol as tapeworm drug, 674 Salt water, mosquito larvae living in, 557 Salvarsan in Asiatic relapsing fever, 631 — in North African relapsing fever, 631 — in Oriental sore, 628 — in relapsing fever, 630 — in tropical syphilis, 632 — in trypanosomiasis, 623 — in yaws, 632 dosage, 632 Salzmann, mode of infection in intestinal myiasis, 727 Sambon, L. W., Linguatula serrata, 527 personal experiments with regard to malarial infection, 158 Samelsohn, retinal haemorrhages in ancylo- stome anaemia, 646 Sandal oil in chyluria from Filaria han- crofti infection, 677 Sand flea, see Dermatophilus (Sarcopsylla) penetrans — flies, 577 and fever due to them in North China, 613 biting in Hampshire, 579 Sandflies, haunts of, 613 see also Siynulium Sandler, trichocephalus anaemia, 651 Sandwith, F. M., toxic symptoms following thymol administration, 686 treatment of bilharziasis, 643 Santonin in bilharziasis, 643 — in expulsion of ascarides, 692 ^ of OxifuridcB, 697 — in intestinal myiasis, 728 Sapo viridis and tar, application in creep- ing disease, 732 Sarcocystin, isolation of, 191 Sarcocystis hertrami, 193 — blanchardi, 193 from ox, 190 Sarcocystis colii, 193 spore, 193 — hueti, 193 — lindemanni, 193 — miescheriana, 188, 193 from pig, 190 — muris, 193 deadly to host, 191 experimental infection with, 191, 192 gymnospores of, 191 spore of, site of sarcocystin, 192 — of muscles, 191 — pansporoblasts of, 189 — recognition from other foreign bodies, 188 — spores of, 189 — tenella, 193 from sheep, 190 — — spores of, 191 toxin isolated fronl, 191 Sarcodina, 27, 29 — characters and habitat, 27 Sarcoid globules in miracidium of Schis- tosoma hcBmatobium, 276 Sarcophaga carnosa (flesh fly), characters, 589 larvae of, 589 regions of human body invaded by, 589 viviparous, 589 — chrysostoma, 590 — hcematodes, 589 — hoBniorrhoidalis , 589 — magnifica, geographical distribution , 589 larvae of, regions of human body in- vaded by, 589 references to, 589 — plinthopyga, probably concerned in dis- semination of yaws, 590 — ruficomis, 589 — wohlfahrti, larvae of, method of destroy- ing, 723 unusual situations of, 723 nasal myiasis from, 722, 723 Sarcopsyllidoe, characters, 543 Sarcoptes, characters, 517 — species transmissible from domestic animals to man, 520 — auchenii, 520 — canis, 520 — dromedarii , 520 — equi, 520 — leonis, 520 — minor, hosts of, 520 — ovis, 520 — scahiei, characters, 518 infection by, disease produced by, 704 see also Scabies synonyms, 518 crnstosce, 519 var. hominis, 519 development of, 519 excavation of tunnels in human epidermis by, 517, 519 ■ transmission of, natural and artificial, 519 886 THE ANIMAL PARASITES OF MAN Sarcoptes suis, 520 — vulpis, 520 Sarcoptidce (itch mites), characters, 516 — development, stages in male and female, 517 — hosts of, 516 — rate of breeding, 517 Sarcoptince, 517, 518 Sarcosporidia, 129, 187, 193 — chambers of, 189 — characters and habitat, 28, 188 — experimental transmission, 191, 192 — fatal to sheep, 188 — hosts of, 187 — in man, 193 — morphology, 188 — muscles affected, 188 Sarcosporidiosis, possible percentage of animals affected by, 191 Scabies, 704 — diagnosis, 705 from occupational eczema, 706 — mite tracks of, 705 — prognosis, 706 — symptoms of, 705 — treatment, 706, 707 — norvegica (Norway itch), 520, 705 Scabiophilia, 706 Scarlet fever, cell inclusions in, 208 Schaudinn, classification of intestinal amoebae, 31 — cytological changes during encystment process of Entamoeba coli, 33 — infection by trichomonads, 56 — intensity of infection with Entamoeba coli, 33 — on Leydenia genimipara, 49 — on Paranioeba hominis, 44 — penetration of red blood corpuscles by sporozoites of tertian parasite, 159 — relapses in malarial fever, 161 — researches on Coccidia, 137, 138, 139 on Entamoeba histolytica, 34, 37 — Treponema pallidum, 114 Schaudinn's fluid, 748 Scheube, lung-fluke disease, 639 Schewiakoff, movements of gregarines, 131 Schiller, Ascaris and Oxyuris infection in relation to appendicitis, 653 — effects of trichocephalus infection, 651 Schistocephalus, pleroceroid of, 300 Schistosoma, morphology, 269 — synonyms, 269 — cercariae, 753 — hmmatobium, distribution in body, 272 endoiihlebitis set up by, 274, 275 female, diameter, 273, 274 morphology, 271 genitalia, 276 geographical distribution, 276 in caval system, 274 in gall-bladder, 274 in hsemorrhoidal veins, 273 in lungs, 274 Schistosoma hcematobium in vesico-pros- tatic plexus, 273, 274 infection hj, 641 see also Bilharziasis male, anterior end, diagram showing organs, 271 carrying female in canalis gynae- cophorus, 270 diameter, 273 morphology, 270 and female in copula, transverse section through, 271 means of access to descending colon, rectum, anal canal, bladder and caval system, 272 miracidium of, sarcoid globules in,. 276 most easily found post mortem in portal vein and liver, 273 ova in utero, diameter, 273 lateral spined, 273 origin of, 273 ovum of, 277 pathological changes in rectum and bladder due to, 274, 275 synonyms, 270 — japonicum, 277 anterior end with testes, posterior end with point of union of caeca, 278 — — female, morphology, 278 from dog, 280 egg from faeces, 280 uterine egg, 280 habitat, 280 liver showing eggs in the intra- and interlobular connective tissue, 282 male, morphology, 277 — ^ and female in copula, 279 — — mode of infection by, 279 ova of, 278 from human liver, showing "spines" and "hoods" at opposite pole, 279 sites in which found in body, 282" — mansoni, 754 Schistosomidce, 269, 753 [ — morphology, 233 I Schizocystis, 135 j Schizogony absent in Eugregarinea, 134 i — in Coccidiidea, 138 — in Leucociitozoa, 153 — of malarial parasites, 161, 172 Schizogregarinea, 135 Schizotrypanum cruzi, 83 Schleip, blood examination in diagnosis of I trichinosis, 681 j Schlesinger, intestinal myiasis, 727 Schliiter, haemorrhagic enteritis from Strongyloides infection, 674 Schmidt, larvae in nose in enormous num- bers, 716 — Trichomonas pulmonalis , 56 Schneider, A., on Coccidia, 137 on Eimeria, 142 on gregarines, 130 INDEX 887 Schuberg, copulation in Coccidia, 137 — immunity of Ornithodorus moubata against infection with Spirochceta dut- toni, 119 Schiiffner, peculiar fever resembling typhoid, 613 Schiiffner's dots, 165, 166, 171 Schultz, on Coccidia in cattle, 741 Schupfer, typhoid lumbricosis, 650 Schwankhaus, Ascaris infection in relation to appendicitis, 653 Schweriner itch following infection by ancylostomes, 684 Schwetz, life-history of Auchmeromyia luteola, 614 Scolex of tapeworms, 300, 303, 304 morphology, 304 Scolopendra in maxillary and frontal sinuses, 721 Screw worm, Indian, see Pycnosoma fly, see Chrysomyia (Compsomyia) macellaria Scutomyia, characters, 563 Seal, host of Dibothriocephalus latus, 315 Sebirol as vermicide, 672 Seeber, Ehinosporidium described by, 197 Sehrt, abscess of omentum with Ascaris ova in pus, 657 Seidelin, association of Paraplasma bodies with yellow fever, 180 Seifert, blood-stained diarrhoea from Strongyloides stercoralis infection, 674 Selenidium, 135 Sellards, see Walker and Sellards Senevet, herpetomonad flagellate in cul- tures of blood and organs of geckos, 739 Senna, syrup of, in expulsion of ancylo- stomes, 686 Sense, organs of, lacking in parasitic nematodes, 366 Sepsidce, characters, 583 — larvae (maggots) of, 583 Septicaemia terminating nasal myiasis fatally, 718 Sergent, transmission of relapsing fever, 120, 121 Sergent, Ed. and Et., herpetomonad flagel- late in cultures of blood and organs of gecko, 739 ''thymni," 725 Sergent, E. and L., deposition of ova of Oestrus ovis, 598 transmission of trypanosomes by species of Tabanus, 601 Sergent and Gillot, treatment of North African relapsing fever, 631 , Sergentella hominis, 210 Serous fluid, bodies resembling amoebae found in, 46 Serum diagnosis of echinococcus, 359 complement deviation, 359 precipitin reaction, 359 — human, action on Trypanosoma rhode- siense, 80 — immune, action on Trypanosoma rhode- siense, 80 Setaria, habitat, 407 — morphology, 407 — equina, hosts and habitat of, 408 morphology, 408 synonyms, 408 — (Filaria) hcemorrhagica, 408 — labiata papillosa, 408 Sexual organs, irritative effects on, set up by migrations of Oxyuris vermicularis, 695 of Echinorhynchus, 476 of Hirudinea, 481 of Insecta, 530 of nematodes, 367, 368, 369 Sheep, baleri in, causal agent, 95 — " carceag " in, cause of, 177 — Cysticercus cellulosoe in, 337 — echinococci in, 346 — heart-water fever in, carrier of, 493 — how infected by Fasciola hepatica, 226 — liver-fluke disease in, death from apo- plexy in first period, 240 period of anaemia, 240 of immigration, 240 — of migration of flukes, 241 of wasting, 240 ravages caused by, 238 — organs infected with echinococcus, per- centage of frequency, 347 — Sarcosporidia fatal to, 188 — section of Sarcocystis tenella from, 190 Sheep-ked, see Melophagus ovinus Shell gland secretion in trematodes, 223 Shiga, discovery of dysentery bacillus, 31 — species of amoebae distinguished by, 31 Shipley, A. E., prophylaxis against clothes lice, 616 Sick, cases of ascarides in bile-ducts, 688 Siebert, application of epicarin in scabies, 707 Siebold, v., development of Taeniae, 14 — explanation of bladder worms, 14 — feeding experiments with Tcenia echino- coccus, 356 — investigations of Gregarinida, 129 — observation of Pseudonavicellae, 129 — psorosperms, 181 — views as to development of Helminthes, 13 Siedlecki, researches on Coccidia, 137 Siegel, Cytorhyctes, 208 — Cytorhyctes luis, 124, 208 Silcock, case of human hepatic coccidiosis, 148 Silkworm disease, " gelbsucht," 207 Nosema cause of, 184 Silvanus surinamensis , characters and habitat, 543 Silver tick, see Amblyomm,a cayennense Simond, researches on Coccidia, 137 Simulidce, 577 Simulium, bite of species of, 578, 579 — characters, 577 — larvae of, 578 — life-cycle of, 578 888 THE ANIMAL PARASITES OF MAN Simulium, wing of, 579 — buissoni, possible connection with spread of leprosy, 579 — columoascnensis , geograjjhical distribu- tion, 578 — damnosum, geographical distribution, 578 — griseicoLLis, geographical distribution, 579 — latipes, 579 — meridionale, possible carrier of chicken cholera, 579 — occidentalis, 579 — wellmanni, 579 Sinton, culture of trypanosome forms of T. gamhiense, 76 of T. rhodesiense, 83 — Prowazekia urinaria, 64, 65 Siphunciilata, 532 — see also Pediculidoi Skin affections caused by cereal mites, 489 due to Dermanyssiis hirundinis , 492 set up by Leptus autiimnalis, reme- dies against, 702 by Trombidium tlalsahuate, 486 — disease caused by larvae of Dermatobia noxialis, 725 due to young nematodes in dogs, 378 produced by Bhizoglyphus parasi- ticus, 514 — diseases set up by penetration of larvae of Ancylostoma duodenale, 455 various names for, 455 — filaria infection of, 378 — infection by Ancylostoma duodenale through, 683 by larvae of Ancylostoma duodenale, 454, 455 — lesions due to Sparganum mansoni, 318 — mole, 599 — parasites of dogs and cats infecting them with Dipylidium caninum, 323 — surface of, larvae on, 721, 722 Skusea, characters, 563 Slaughter-houses, infection of rats with Trichinella in, 427 Sleeping sickness, 68, 69, 72, 76, 620 association of trypanosomes with, 68 cerebral stage, 621 due to Trypanosoma gambiense, 68, 72, 620 rhodesiense, 69, 76, 620 symptoms, 622 febrile or glandular stage, 621 incubation period, 621 investigation of, 68 parasites producing, 72, 76, 605 pathology of, 621 preventive measures, 623 Rhodesian, daily number of trypano- somes in blood from case of, 79 transmission of, 68, 605, 607, 608 experimental (with apes), 68 treatment by arsenic and arsenical preparations, 622, 623 by atoxyl, 622 Sleeping sickness, treatment by tartar emetic, 622 must be commenced in early stages to be effective, 622 Sleeping Sickness Bureau Bulletin, founda- tion of, 69 Sloth, blood of, inhabited by Endotry- parium schaudinni, 99 Smith, Theobald, experimental infection of mice with Sarcocystis maris, 191 — and Barrett, Endamoeba, 734 Endamoeba gingivaiis, 733 treatment of oral endamoebiasis, 620 Smith and Kilborne, 174, 176, 177 — and Weidman, Entamoeba mortinatalium , 45 Smithia, morphology, 174 — microti, 174 — talpcB, 174 Snake, blood of, transference of Trypano- soma brucei to, from blood of rat, 102 Soamin in sleeping sickness, 623 Soda, bicarbonate, with iodoform in expul- sion of ascarides, 694 — salicylate of, lavages of, in nasal mvia- sis, 719 Soldiers, Pediculus vestimenti pest among, during campaigns, 533 — prophylaxis against clothes lice among, 616 Solium, derivation of specific term (foot- note), 331 Souma in bovines and equines, causal agent, 100 Space parasites, 20 Spaniopsis tabaniformis , 614 Sparganum, 317 — mansoni, 317 cephalic end, 318 diagnostic signs of j^resence, 659 discovery of, 317 geographical distribution, 659 habitat in body of man, 659 migration in body, 318 plerocercoid of, 318 skin lesions due to, 318 ; symptoms set up by invasion, 059 synonyms, 317 transverse section of, 318 — proliferum, 318 acne-like condition set up by, 318 geographical distribution, 320 mode of infection, 320 morphology, 319 synonyms, 318 Spengel, Filaria (?) kilimaroe, 407 Spermatozoa of Trematoda, no essential difference in structure from those of other animals, 222 Sphcerularia, nematodes hatched from eggs of, 5 Spiders, see Arachnoidea , 483 Spinal ganglia of rabid monkeys, cultiva- tion, 210 Spinning mites, see Tetranychidcs Spirochaeta, 115 INDEX 889 Spirochoeta aboriginalis, association with granuloma inguinale, 122 — acuminata, 122, 128 — anodontce, 114 — anserinu, 119, 122 — balbiunii, 114 — berberu, agent of North African and Egyptian relapsing fever, 122 — bvonchialis, 122, 632, 739 mode of infection, 740 morjihology and life-history, 739, 740 — buccalis, 122 morphology, 741 — cavtcri, agent of Indian relapsing fever, 122 — dentium, 122, 128 morphology, 741 — duttoni, 116 agent transmitting, 116 cause of African relapsing fever, 116, 630 cultivation of, 123 geographical distribution, 116, 119 infection by, experimental, 117 summary of methods and re- sults, 118, 119 — — — ■ immunity of Ornithodorus mou- bata against, 119 transmission of, 739 — eurygyraia, 122 — galiinarum, 119, 122 agent of transmission, 119 appearance in hsemocoelic fluid of Argas yersicus, 119 • cultivation of, 123 fatal to fowls, 119 — gigantea, 114 — granulosa, 116 — hachaizce in cholera motions, 122 — laverani, small size of, 122 — marchouxi, see Spirochceta galiinarum — muris, 122 — novyi, agent of North American relaps- ing fever, 122 cultivation of, 123 — obermeieri, see Spirochceta recurrentis — obtusa, 122, 128 — ovina, 122 — phagedenis, 122 — 2}licatilis, 114 — recurrentis , 120 agents of transmission, 120 cause of European relapsing fever, 120, 122 ■ cultivation of, 123 incubation period, 630 morphology, 120 — ref ring ens, 122, 128 association with Treponema jyalUdum., 122 — rossii, agent of East African relapsing fever, 122 cultivation of, 123 — schaudinni, agent of ulcus tropicum, 122 — stenogyrata, 122 — theileri, 122 Spirochceta vincenti, 122 Spirochcetacea, 115 Spirochsetes, 114 — blood inhabiting, 116 — classed among Frotista, 29, 115 — cultivation 01, 123 presence of oxygen necessary for, 123 — granule phase of, 120 — hosts of, 114 — in alimentary tract, 741 — in human mouth, 122, 740 — in voinited matter, 122 — mode of division, 115 — molluscan, breaking up into granules, 119 — morphology and morphological varia- tion, 114, 115 — of human mouth, recent Avork on, 740, 741 — of relapsing fever, periodic increase and decrease in blood, 115 — reaction to drugs, 115 — systematic position, 115 Spirochcetoidea, 115 Spirochsetoses, 629 — bronchial, diseases for which mistaken, 632 treatment, 633 — relapsing fever, 629 — syphilis, 632 — yaws, 632 Spiroschaudinnia, 115 Spleen, development of crescents of tertian malignant parasite in, 169 — enlargement due to ova of Schistosoma japonicum, 282 in malaria, 634 — pigmentation of, following malaria (footnote), 165 Splenic blood, citrated, cultivation of Leishmania donovani in, 106 — vein, tributary of portal vein, 272 Splenomegaly, association of Histoplasma capsulatum. with, 112 of Toxoplasma pyrogenes with, 113 — infantile (kala-azar), 109, 627 Spontaneous generation, theory of, 10 early opposition to, 10 Sporoblasts of Coccidiidea , 141 — of malarial parasites, 163 — of Myxosporidia, 183 Sporocyst, germ balls of, 227 — of Coccidiidea, 141 — of gregarines, 134 — of trematodes, 225, 227 Sporogony, 144, 186 Sporozoa, 28, 128 — characters and habitat, 28 — classification, 129 — hosts of, 129 — relation to Protozoa, 19 I Sporozoites of Coccidiidea, 138, 139, 140 i — of gregarines, 132, 133 ' — of malarial parasites, 159 j Stained material, examination of, 747 ' Staining, 749 890 THE ANIMAL PARASITES OF MAN Stallion's disease (dourine), trypanosomes in blood of horses with, 68 Stannus, species of EnyaLiopsis producing ulcers, 542 — and Yorke, observation of Trypanosoma rhodesiense in animals inoculated from case of sleeping sickness, 78 Staphylocystis, 304 Staubli, blood examination in diagnosis of trichinosis, 681 Steel and Evans, experimental transmission of Trypanosoma evansi, 67 Steenstrup, discovery of method of multi- plication of Heiminthes, 13 Stegomyia, breeding of, prevention, 636 — characters, 563, 571 — ovum of, 557, 558 — transmission of yellow fever by, 555 — albipes, characters, 572 — albocephala, characters, 573 — albolateralis , characters, 573 — albomarginata, characters, 573 — amesii, characters, 573 — argenteomaculata, characters, 572 — argenteopunctata, characters, 573 — assamensis, characters, 573 — auriostriata, characters, 573 — crassipes, characters, 573 — dubia, characters, 573 — fasciata, biting hours of, 574 breeding of, 574 carrier of yellow fever, 574 characters, 572, 574 development of Plasmodium relictum in, 171 — — distinguishing characters of S. scu- tellaris from, 575 domesticated species, 574 food of, 574 geographical distribution, 574 larvae of, habitat, 574 ova of, 574 possible host of Leishmania tropica, 108 source of danger to Panama Canal, 574 supposed intermediate host of para- site of Bagdad sore, 575 transportation of, 574 — gelebinensis, characters, 572 — grantii, characters, 573 — lilii, characters, 572 — mediopunctata, characters, 573 — minuta, characters, 573 — minutissima, characters, 572 — nigeria, characters, 572 — poweri, characters, 572 — pseudonigeria, characters, 572 — pseudonivea, characters, 573 — pseudoscutellaris , 394, 575 characters, 572 intermediate host of filaria in Fiji, 575 — punctolateralis , characters, 573 — scutellaris , characters, 572 distinguishing character from S. fas- ciata, 575 Stegomyia simpsoni, characters, 572 — terreus, characters, 573 — tripunctata, characters, 573 — W-alba, characters, 572 — wellmannii, characters, 572 Stein, interrelation of pseudonavicellae and gregarines, 129 Stein, v., classification of Infusoria, 199 — discovery of meal worm in bladder worm, 303 Steinhaus, intestinal stenosis following in- fection by Tcenia solium, 662 Stempell, on Nosema bombycis, 184 Stephens, J. W. W., appendix on Trema- toda and Nematoda, 753 Nemathelminthes, 360 Plasmodium tenue, 170 Platyhelminthes or flat worms, 211, 638 and Christophers, Maurer's dots, 168 relapses and latent infection of malaria, 158 and Fantham, length of Trypanosoma gambiense, 73 Trypanosoma rhodesiense, 69, 76 Stern, symptoms of cysticercus in fourth ventricle, 665 Stethomyia, characters, 561, 567 Stiles, C. W., infection with Lamblia in- testinalis, 60 prophylaxis against flagellate diar- rhoea, 625 Stillborn child, problematical " monocystid gregarines " from lung tissue of, 150 Stitt, alkaloid of quinine in malaria, 635 — paroxysms of malignant tertian fever, 634 Stock, bilharziasis, 641 — treatment of bilharziasis, 644 Stokvis, Balantidium coli occurring in lung, 202 Stomach, cancer of, Lamblia intestinalis in, 59, 60 — fluid from, obtained by lavage, rhabditea found in, 378 — larvae of Gastrophilus inhabiting, 599 — trichomonads in, 55 — Tristrongylus instabilis in, 435 — wall, fibrous thickenings in, produced by species of Gnathostoma, 385 Stomoxys, characters, 609 — differentiation of Lyperosia from, 610 — disease carrier, 603 — species of, 610 — calcitrans (stinging or stable fly), 609 diseases transmitted by, 610 ova, larval and pupal stages, 609 transmission of epidemic polio- myelitis by, 612 Stools, larvae of Blaps mortisaga in, 542 — method of discovering head of tape- worms in, 674 StreblidcB (bat parasites), 611 Strepsiptera, characters, 531 Strong and Musgrave, species of amoebae distinguished by, 31 INDEX 891 Strung ylidce, 375, 432 — free life of young stages, 20 Strongyloides, European, free-living gene- ration generally absent in, 383 — fulleborni, 384 — intestinalis , geogra2)hieal distribution, 384 ■ — larvae of, cultivation, 474 — - life-history of, 19 — morphology, 379 — stercoralis , free-living form, morphology of, 381 generation, female, 382 ■ habitat in body, 755 — — heterogony of, 381 infection by, diagnosis, 675, 676 diarrhoea associated with, 381 expulsive treatment, 675 — pathological significance, 674 — prophylaxis against, 675 symptoms, 674, 675 larva from fresh human faeces, 382 — mature filariform, 383 mode of development, 373 occurrence in man, 384 — — parasitic generation, morphology, 381 ova, 381, 382 synonyms, 380 and Ancijlo stoma duodenale, larvae of, diiit'erences between, 451 — synonyms, 379 — toxic action of, 651 — vivipara, 384 Strongylopkiimata, 208 Stuelp, amaurosis following male fern poisoning, 671 Stuertz, chyluria following infection by Eustrongylus gigas, 682 Stylorhynchus, host of, 135 — oblongatus, gametes of, morphological differentiation, 133, 134 Stypticin in bilharziasis, 643 Sublimate, corrosive, saturated aqueous, fixation of cestodes by, 472 solutions, fixation by, 748 — solution in crab louse infection, 712 injection in expulsion of Guinea worm, 676 into cutaneous and muscular cysti- cerci, 663 Suckers of Cestoda, 289 Sucking worms, see Tr-etnatoda Suctoria, 29, 198 — characters and habitat, 29 Sulphur, flowers of, projjhylactic against clothes lice, 616 — preparations, application in scabies, 706 Sump bunches, skin affection set up by t penetration of larvae of Ancylostoma duodenale, 455 Surra, animals among which prevailing, 95 — causal agent of, 95 — geographical distribution, 95 — transmission by Chrysops, 601 by Stomoxys, 96, 610 by Tabanus sp., 96 56 I Surra, trypanosomes in blood of horses with, 67 Swallow bug, see Cimcx hirundinis Swammerdam, discoveries of origin of para- sites, 10 Swamps, drainage of, in prevention of malaria, 636 Sweden, ox Avarble fly (Hypodernia bovis) attacking man in, 596 Swellengrebel and Strickland, on Trypano- soma lewisi, 92 Symbiosis, 6 Symmers, bilharziasis of lung, 642, 643 Symphoromyia, characters, 603 Syngamece, characters, 459 Syngamus, 459 — habitat and hosts of species, 459 — kingi, habitat and host, 460 morphology, 459, 460 — trachcalis, bursa of, 461 Syphilis, inoculation with, j^roducing no immunity to yaws, 128 — non-immunity to, produced by inocula- tion with yaws, 128 — parasite of, 114, 124, 125, 632 — tertiary eruptions of, Treponemata diffi- cult to find in, 125 — treatment, 632 Syrphidoz, rat-tailed larvae of, characters and habitat, 583, 584 ^J^J^J of gregarines, 132 Szerlicky, case of intertrigo set up by Oxyuris vermicularis, 696 Tabanid^ (gdd flies), characters, 600, 601 — diseases transmitted by, 601 — larvae, 600 — method of destruction, 601 — ova, 600 — pupae, 600 Tabanus, species of, transmitting trypano- somes, 96, 601 — bovinus (ox gad fly), 601 Taenia, 331 — africana, mature segment of, 342 morphology, 342 oncosphere of, 299 — — proglottis and head of, 343 — bremneri, morphology, 337 — capensis, 339 — coenurus, nervous system, head and part of neck showing, 291 — confusa, mature and gravid segments, 344 morphology, 343 — crassicollis, anatomy of, longitudinal section showing, 290 cysticercus of, 338 host of, 6 — derivation of name (footnote), 331 — echinococcus, hooklets of, 355, 359 hosts of, 345 morphology, 344 organs of, 345 892 tHE ANIMAL PARASITES OE MAN Tcenia echinococcus, percentage of dogs infected with, in various cities and countries (footnote), 345 — — rearing of, in dog, 356 synonyms, 344- see also Echinococcus — expulsion of, resulting in cure of chorea, 648 — extracts of, experimental injection, effects, 648 — lata (Dibothriocephalus latus), supposed origin of, 11 — lophosorna, 339 — marginata, 337 cysticercus of, 338 hooks of, 338 hosts of, 338 — oncospheres of species of, animals selected as hosts for development, 299 migration from intestine through blood-vessels to liver, 302 — saginata, cysticercus of, 340 expulsion of, best method for, 669, 67^ — — frequency in man, 341 • genitalia, proglottis showing, 293 geographical distribution, 341 habitat in man, 667 host of, 6 malformations, 339 morphology, 339 parasitic in man in association with other tapeworms, 667 proglottids of, feeding experiments with, 340 prophylaxis against, 668 race incidence of infection, 340 symptoms produced by infection by, 667, 668 synonyms, 338 uterine e^g, 298 — serrata, cysticercus of, 338 hooks of, 338 host of, 338 — solium, 339 carriers of, 335 diagnosis of presence in body, 662 Dipylidium caninum confused with, 680 expulsion from body, effect on anaemia, 648 measures for, must be thorough, 336 geographical distribution, 334 corresponds with that of domes- tic pig, 334 ■ habitat in body of man, 662 head of, 332 host of, 6 in man, mode of infection, 335 larval infection, 662 see also Cysticercus celluloses malformations of, 332 modes of transmission, 336 morphology, 331 parasitic association with Dibothrio- cephalus latus, 658 Tcenia solium, proglottids, 332 prophylaxis against, 668 symptoms produced by infection by, 667, 668 synonyms, 331 — species of, respective times required for development of cysticercus from date of infection, 304 various, respective time required for growth, 306 Taeniae, development of, 14 — infection by, treatment, symptomatic, 669 — nervous system of, 290 — oncospheres of, 14 — species of, in relation to cystic forms, 16 Tceniidce, 331 — egg-shell substance, 297 — eggs of, 297 — morphology, 309 — oncospheres of, development of cysti- cerci from, 303 — rostellum, 289 of, ring encircling, 291, 292 Taeniol, administration in ancylostomiasis, 686 — effects of, 672 Taeniorhynchus, 576, 577 — africana, 577 — annulipes, 577 — australiensis , 577 — characters, 564 — major, 577 — ova of, 557, 558, 577 — titillans, 577 carrier of larvae of Filaria bancrofti, 577 — uniformis , bll carrier of larvae of Filaria bancrofti, 577 Tallqvist, experimental bothriocephalus anaemia, 646 Tamne or thimni of Kabyles, 598 Taniguchi, paragonimiasis of brain, 639 Tapeworms, adult, length of life, 307 — biology, 307 — caudal vesicle, 300 — cysticerci experimentally reared from, 15 — development of, 297 embryonal, 298 — embryophore, 298 — experimental rearing of, 15 from cysticerci, 15 — expulsion by preliminary aperients, 669 by vermifuges, 669 — found in association with other intes- tinal parasites, 667 — individuality of, early researches as to, 283 — infection by, symptomatic treatment, 669 — injury inflicted by, depends on number in host, 9 — larvae of, sexual maturity must take ! place in terminal host, 304 — larval stages, development, 298-301 modes of, 300 INDEX 893 Tapeworms, metamorphosis of, 15 of larva into, 305 — method of discovering- head in stools, 674 — oncospheres (embryos) of, 298, 299 transformation into bladder worms, 303 — origin of, discovery, 11 early researches as to, 283 — ova of, 297 consistency, 297 — plerocercoid of, 300 — scolex of, 300, 303, 304 Tar and sapo viridis, application in creep- ing disease, 732 Tarsonemidce, characters of, 488 Tarsonemus intectus, 489 — uncinatiis , 489 Tartar emetic in espundia, 629 in Indian kala-azar, 626 in infantile kala-azar, 627 in Oriental sore, 628 in sleeping sickness, 622 Taschenberg, Silvanus surinamensis , 542 Taylor, treatment of bronchial spirochaeto- sis, 633 Technique, protozoological, fixed and stained material, 747 fresh material, 745 notes on, 745-752 Teeth, carious, spirochaete associated with, 122 Teichomyza fusca, larvae of, habitat, 584 Teissier, mercury in expulsion of Strongy- loides stercoralis, 675 Telosporidia, 28, 129 — characters, 28, 129 Temnocephalid(B, habitat and habits of, 20 Terebinthine oil in chyluria from Filaria bancrofti infection, 677 Ternidens, characters, 439 — deminutus, 439, 440 habitat, 441 Tersesthes, 581 Testis, enlarged, in filariasis, 401 — of Ancylostoma duodenale, 449 Tetramitus, 57 — and Chilomastix, differential characters, 735, 736 — how differing from Trichomonas, 57 — mesnili, causal agent of colitis, 57 Fanapapea intestinalis identical with, 57 habitat, 57 — — synonyms, 57 Tetranychidoe (spinning mites), characters of, 488 Tetranychus, 488 — molestissirnmus, geographical distribu- tion, 488 itching produced by, 488 — telarius, var. russeolus, effects produced by, 488 Tetratrichomonas, 53 (footnote), 734 Texas fever in cattle, carriers of, 177, 494 causal agent, 173, 177 Theiler, 178, 180 Theileria, 174, 178 — annulata, 180 , — characters of, 174 — mutans, 180 — parva, 178, 179 agents of transmission, 179 Koch's blue bodies in, 179 life-cycle in tick, 179 morphology of, 178 pathogenic agent of East Coast fever in cattle, 174, 178 — stordii, 180 Thelohan on Myxosporidia, 182, 183 Thelohania contejeani, 186 Theobald, F. V., Arthropoda (jointed limbed animals), 483 Theobaldia, 575 — annulata, bite of, 575 characters, 575 domestic form, 575 geographical distribution, 575 larvae of, habitat, 575 — characters, 564 — spathipalpis , bite of, 575 characters, 575 geographical distribution, 575 Theobaldinella, 575 Thiarsol in infantile kala-azar, 627 Thiopinol, application in scabies, 706 Thomas, W., introduction of atoxyl in try- panosomiasis, 622 Thomer, treatment of crab louse infection, 712 Thomson, D., sites of development of crescents of tertian malignant parasite, 169 see also Ross, Sir R., and Thomson, D. Thomson, J. D., researches on Trypanosoma lewisi, 89, 90, 92 Thomson, J. G., and Fantham, cultivation of Babesia (Piroplasma) canis by Bass's method, 172 nuclear phenomena of Babesia canis in cultures, 176 see also Fantham and Thomson, J. G. Thomson, J. G., and Sinton, culture of Trypanosoma rhodesiense, 82, 83 culture of trypanosome forms of T. gambiense, 76 medium employed by, for growth of Trypanosoma gambiense and T. rhodesiense, 745 and Thomson, D., methods of cultiva- tion of malarial parasites, 171, 172 number of merozoites of malig- nant tertian parasite, 168 spirochaetes in alimentary tract, 741 Thornhill, toxic symptoms following thymol administration, 686 " Thymni " or tamne of Kabyles, 598, 725 Thymol, administration of, in expulsion of ancylostomes, 685, 686 mode of, 685, 686 of ascarides, 694 894 THE ANIMAL PARASITES OF MAN Thymol, administration of, in expulsion of Oxynridop, 697 of Strongyloides siercoralis, 075 in flapfellate dysentery, 624 in Trichiiris trichiiira infection, 679, 680 followed by benzene enemata, 680 ■ toxic symptoms following, 686 — enemata in arrest of trichinosis, 681 in expulsion of ascarides, 694 Thymoluria, 686 Thymotol, administration in ancylosto- miasis, 686 Thyroiditis, parasitic, 87 — see also Trypanosomiasis , Brazilian Thysanoptera, characters, 531 Tick, stag-es of life-cycle of Babesia canis and B. hovis in, 176, 177 — bites, paralysis due to, 613 — or relapsing- fever, 116, 630 African, carrier of, 116, 496, 630 • importation into Persia, 613 pathogenic agent, 116 — paralysis, cause of, 504 Ticks, transmission of piroplasmosis by, from recovered to uninfected animals, 178 Tiger, host of Paragonimus westermannii; 250 Tinea rotunda, see Ascaris lumhricoides Toad, rectum and urinary bladder of, Opalina parasitic in, 207 Todd, on leucocytogregarines in birds, 154 — tick paralysis, 613 — see also Button and Todd Tommasi-Crudeli, early researches on malaria, 156 Tomsk, Opisthorchis felineus, human para- site most frequently found at autopsies at, 253 Tongue, cysticercus of, 663 Townsend, Simulium occidentaJis , 579 Toxascaris, characters, 465 — limhata, morphology, 466 ovum of, 466 synonyms, 466 Toxoplasma, 112 — hosts of, 113 — pyrogenes. association with splenome- galy, 113 Toxorhynchites, characters, 563, 570 Trachea, ascarides invading, 691 Trachoma bodies in infected epithelial cells of conjunctiva. 209 (fig. 119) so-called, cultivation, 210 Trematoda. endoparasitic life spent in in- termediate and final host, 18 — relation to Turhellaria, 19 Trematodes (sucking worms or flukes), 212 — ao-e attained by, 230 — alimentary canal, 217 — asexual generations, 224, 225 — cercariae (larval stages), 225, 227, 228 — cirrus sac, 221 — copulation iii, 222 -. — cross, 22? Trematodes, development, 12, 222 embryonic and post-embryonic, 224 final, conditions necessary for, 225 — developmental cycle, 229 — digenetic, adult, animals harbouring, how infected, 226 development, 224, 226 miracidia of, 226 — endoparasitic, biology of, 229 hosts and habitat of, 229 — excretory bladder, 219 system, 219 terminal flame cell, 219 — food of, 218 — found in man, classification, 230 — genital pore, 222 — intestine of, variation in, 217 — investing layer of, 213 — Laurer's canal of, 221, 222 — metraterm of, 221, 222 — miracidia of, 223, 224 — morphology of, 212 - movements of, 216 - muscular system of, 214 - nervous system of, 216 — organs of sense, 216 — origin of, 12 of parasitism in, 20 — ova of, deposition, 223 formation, 223 — parenchyma of, 213 muscles of, 214 — redise of, 225, 226, 227, 228 — salivary glands, 217 — sexual organs, 220 deviation from typical position (footnote), 222 female, 220, 221 male, 220 — shell gland secretion in, 223 — sporocyst of, 225, 227 — suckers of, 213, 214 — and turbellaria, genetic relationship be- tween, 20 Treponema, 114, 115, 123 — calliqyrum, 126 — cultivation of species from human mouth, 128, 741 — macro d entium , 128 — microdentium , 128 — morphology, 124 — mucosum, 128 — pallidum, 114 causal agent of syphilis, 124 ^ — - cultivation of, method, 125 difficult to find in tertiary eruptions of syphilis. 125 granule fcrmation, 124, 125, 127 morphological and pathogenic varia- tions, 126 morphology, 124, 125 Svirochoeta refringens associated with, 122 - — • — synonyms, 124 — pertenue, ciiltivation, 128 granule formation, 127 INDEX 895 Treponema pertenue, mode of infection, 128 morphology, 127 pathogenic agent of yaws, 114, 127 reasons for considering specific cause of yaws, 128 — species of, association with pyorrhoea alveolaris, 128 Treutler, filaria associated with phthisis, 408 — parasite, probably liver-fluke, in vein, 243 Triaenophorus, excretory vessels, island formation, 292 — plerocercoid of, 300 Triatoma megista, discovery of Trypano- soma cruzi in, 83, 84 jihases of development of Trypano- soma cruzi in, 87 preventive measures against, 623 Triboulet, Ascaris infection in relation to appendicitis, 653 Trichina spiralis, 423 Trichinella, 421 — development in definite host, 18 — spiralis, 421 development of, 373 history of, 423, 424 geographical distribution not in correspondence with occurrence of trichinosis in man, 427, 428 hosts of, 6 in man, percentage of invasion according to nationalities determined by post-mortem examination, 428 infection by, 680 distribution in body after, 424 — see also Trichinosis invasion and encystment in muscles, 424, 425 mammals in which developed experi- mentally, 421 infected by, in order of frequency, 421 inhabited by, 421 morphology, 421 normal hosts of, 427 — — symptoms produced by, in periods of invasion, dissemination and encystment, 424, 425 viviparoiis nematode, 371 Trichinellse, development in encysted con- dition, 427 — encysted, in man and other mammals, early observations of, 423 — fatal case of infection by, 423 - — feeding experiments with, 423 Trichinellida?, 419 — characters, 375 Trichinellince, 421 Trichinosis, amount of prevalence in North America, 428 — diagnosis, 681 -by blood examination, 681 — epidemics of, 423 in Germany, 423, 429 r — geographical distribution, 428 Trichinosis in man, geographical distribu- tion of Trichinella spiralis not in cor- respondence with occurrence of, 427, 428 — prophylaxis against, 429, 431 — symptoms of, 424, 425, 680 — treatment, before and after development, 681 Trichocephali in appendix, 655 Trichocephalus anaemia, 651 — infection by, effects of, 651 in relation to appendicitis, 653 — lacks intermediate host, 21 Trichomonads, habitat in body, 55, 735 — question of cysts of, 56 Trichomonas, 52 — characters of, 52 — diarrhoea due to, 57, 624, 734 — from gut and caecum of rat, 735 — hominis same as T. intestinalis, 54 — intestinalis, 45, 54 — — • axostyle of, 55 characters of, 55 flagella of, 55 relation to T. vaginalis, 54 spherical contracted forms in mice, 56 transmission, modes of, 56 — points of difference of Tetramitus from, 57 — regions of body other than intestine in which found, 55, 56 — vaginalis, 52, 760 characters of, 52, 53 flagella of, 53 nucleus of, 53 — — presence in urethra of male, 53 Trichomoniasis, human, recent researches in, 734 — oral, treatment, 625 — vaginal, treatment, 625 Trichopalpus, 603 — larvae, characters and habitat, 603 — ohscurus, 603 Trichoptera, characters, 531 Trichosoma crassicaudum, female parasitic, 4 habitat of, 4 Trichostrongylince , characters and habitat, 433 Trichostrongylus, morphology, 434 — instabilis, habitat, 435 hosts of, 435 ^ in man, cases recorded, 435 morphology, 434 — probolurus , habitat, 435 hosts of, 435 morphology, 435 — vitrinus, hosts of, 436 morphology, 435, 436 Trichotrachelida-, oesophagus of, 363 — unicellular cutaneous glands of, 361 Trichurince, 419 Trichuris, morphology, 419 — alcocki, 421 — cameli, 421 — campanula, 421 — crenata, 421 infection with, 420 896 THE ANIMAL PARASITES OF MAN Trichuris depressiuscula, 420, 421 infection with, 420 — discolor, 421 — giraffcB, 421 — globidosa, 421 — nodosus, 421 — avis, 421 infection with, 420 — trichiura, habitat in man, 420 infection with, sources, 679 symptoms, 679 treatment, 679 mammals inhabited by, 421 mode of attachment to wall of in- testine, 679 morphology, 419 ova, development of, 420 embryo-containing, 420 infection by, 420 parasitic in large intestine, 678 percentage found at autopsies, 420 synonyms, 419 — unquiculata, 421 Trinidad, mosquito worm in, 598 Triodontophorus, bursal formula (foot- note), 439 Troglotremidcs, 249 — morphology, 232 Trombidiidas, characters, 485 Trombidium, 485 — fuliginosum, 486 — gymnopterosum, 486 — serraticeps, 486 — tlalsahuate, skin affections set up by, 486 Trophozoites of Coccidia, 140, 143 — of Entamoeha tetragena, 39, 40 — of gregarines, 132 — of malarial parasites, 159 — of Microsporidia, 185 — of Myxosporidia, 182 Tropical Diseases Bureau Bulletin, founda- tion of, 69 Tropical sore, see Oriental sore Trouessart, Histioqaster (entomophagus ?) spermaticus , 515 Trypan-blue treatment of piroplasmosis, 178 dosage for dogs, horses and cattle, 178 Trypanophis, 63 Trvpanoplasma, characters of, 63 — hosts of, 63 T: ypanoplasms in iish, 68 Trypanosoma, 67 — americanum, 69 — hoylei, 99 experimental infection with, 99 host of, 99 — hriicei. 93, 94 and T . rhodesiense, question of dis- til- ction or identity, 80, 83, 94 blepharoplastless strains, 101, 737 cause of nagana ("tsetse-fly disease), 93 development in Glossina morsitans, 94 drug resistance of, 101 innocuous to big ffame, 70 morphology and life-history in verte- brate host, 94 Trypanosoma hrucei, nucleus, blepharoplast and flagellum of, 70 posterior nuclei in, 83 strain from Uganda, 95 from Zululand, 94, 95 — caprce, monomorphic, 100 — cazalboui, causal agent of " souma," 100 monomorphic, 100 — characters, 67 — congolense, agents of transmission, 100 cause of Gambia horse sickness, 100 geographical distribution, 100 monomorphic, 100 probable synonyms, 100 — cruzi, 83 crithidial forms, 86 culture, 87 geographical distribution, 83, 84 hosts of, 85, 86, 87 in foetus, 88 invertebrate host of, 83, 84, 537 life-history in invertebrate host, 80 in vertebrate host, 84 modes of multiplication (" sexual " and asexual), 85, 86 microgametes and macrogametes, 85 morphology, 84 possible reservoir of, 87 schizogony of, 84, 85, 86 — dimorphon, 100 — equi, 83, 98 — equinum, cause of " mal de caderas," 96 morphology, 96 transmission of, 97 — equiperdum, 97 cause of " dourine " or stallion disease, 97 endotoxins in, 98 morphology, 98 posterior nuclei in, 83 progress of disease, 97 — evansi, blepharoplastless strains, 737 causal agent of surra, 95 morphology, 95, 96 possible case in man, 96 synonyms, 95 transmission of, 95 experimental, 67 variety causing " mbori " in drome- daries, 96 — fringillarum, 737 — gambiense, 68, 72 cause of sleeping sickness, 68, 605 cultivation of, medium used for, 745 cultures of trypan osome forms of, 76 development in Glossina palpalis, 74, 75 effect of serum reactions on, 80 immunization against, does not pro- tect against infection by T. rhodesiense, 80 in antelope, 76 innocuous to big game, 70 invasion of salivary glands of Glos- sina palpalis, 75 latent forms of, 77 INDEX 897 Trypanosoma gambiense, morphology, 72 in circulating blood, 73 serum from animals infected Avith, no effect on T. rhodesiense, 80 trypanolytic for, 80 synonyms, 72 - — hippicum, agents transmitting, 99 ■ cause of " murrina " in mules, 98 morphology, 98 — lewisi, crithidial forms, 91 development in rectum of rat flea, 91, 93 inoculation experiments, 90 life-cycle in invertebrate host, 90, 91 in vertebrate host, 88, 89 morphology, 88 multiplication rosettes, 71 potential pathogenicity, 737 — — rosette forms, 89, 90 strain of, losing resistance to arseno- phenyl-glycin, how effected, 93 — — transference from blood of rat to blood of snake, 102 transmission of, 88 — nanum, 100 — nigeriense, 76 — noctiicB, 69, 737 — pecaudi, 95 causal agent of baleri in sheep and equines, 95 posterior nuclei in, 83 — pecorum, 100 — rhodesiense, 69, 76 and T . brucei, question of distinction or identity, 80, 83, 94 animal reactions, 78 cause of Rhodesiau sleeping sickness, 69, 76, 605 cultivation of, 83 medium used for, 745 developmental cycle in Glossina mor- sitans, 81 effect of serum reactions on, 80 Glossina morsitans transmitting, 608 immunization against T. gambiense does prevent infection by, experiments proving, 80 latent or resting forms of, 77, 78 morphology, 76, 77 non-pathogenic to antelopes, 70 partial immunity against, 81 pathogenic to man and laboratory animals, 70 posterior nuclei in, 83 reservoir of,' 81 resistant to atoxyl, 78 — — serum from animals infected with T. gambiense has no effect on, 80 transmission of, 69, 81 climatic factors affecting, 81 virulence of, compared with that of T. gambiense, 78 — simioB, virulent to monkeys and pigs, 100 — theileri, 98, 611 — — geographical distribution, 98 morphology, 98 Trypanosoma ugandce, 95 — uniforme, hosts of, 101 monomorphic, 101 — vivax, fatal to cattle, 99 — — monomorphic, 99 transmission of, 100 Trypanosome, animal, infection of human being with, 96 — diseases spread by Glossina, 603 — human, 68, 69 ■ artificial infection of species of Glossina with, 605 — infections, Liverpool School of Tropical Medicine Expedition sent to investi- gate, 68 Trypanosomes, adaptation of, 101 — artificial cultivation, 69 — blepharoplastless, 101, 737 — classification, 71, 72 — deleterious or fatal to domestic animals, 69 — general note on development in Glos- sina, 101 — hosts of, 67, 68, 69 — immunity to, in antelope, 69 — in blood, cultures aid in detection of, 69 ~ — cyclical variation, 78 — — daily number from case of Rhodesian sleeping sickness, 79 method of determining number, 748 multiplication, 71 periodicity, 69 seasonal variation, 69 — in cerebrospinal fluid from cases of sleeping sickness, 68 — latent forms, non-flagellate, from in- ternal organs of vertebrates, 73, 74, 77 — monomorphic, 99 — morphology of, 70 — nuclei of, 70 — pathogenic to man and domestic animals, 70 — percentage of fleas fed on infected rat becoming infected with, 93 — polymorphism, 72 — posterior nuclei in, 83 — resting stages, 72 — transmission from one vertebrate host to another, 72 — transmissive stage in vertebrates, 737 — transmitted experimentally by Stomoxys, 610 — undulating membrane, 71 Trypanosomiasis, African, see Sleeping sickness — Brazilian, acute, 87 chronic, varieties of, 87, 88 clinical features, 87 hereditary transmission, 88 histopathology, 88 suggested treatment, 623 synonyms, 87 — cryptic, 69 TrypanosomidcB, 61 — characters, 66 — genera of, 67 898 THE ANIMAL PARASITES OF IMAN Tryposafrol, producing blepharoplastless trypanosomes, 737 Tsetse-fly, see Glossina morsitans — disease, see Nagana Tuberculosis, see Cestode tuberculosis "Tuft-like" or "phagocytic" organs of nematodes, 362 Tumours, subcutaneous, associated with in- vasion by Onchocerca volvulus, 418 Turbellaria, parasitic, 2 — relation of Trematoda and Cestoda to, 19 — and trematodes, genetic relationship be- tween, 20 Turkeys, blackhead in, 145 Turpentine in flagellate diarrhoea, 624 — in nasal myiasis, 719 — oil of, in bilharziasis, 643 Tydens molestus, habitat, 491 host-tormenting, 491 Tylenchus putrefaciens , 379 Typhlitis, association of Oxyuris vermicu- laris with, 467 Typhlocoelum flavum, progeny of, discovery, 12 Typhoid fever, helminthes as predisposing factor of, 657 peculiar fever resembling, 613 spread by house-fly, 586 symptoms of, in lumbricosis, 650 — vaccine in bilharziasis, 644 Typhus, possibly due to a chlamydozoon, 207 Tyroglyphi, differentiation of Glyciphagi from, 513 TyroglyphidcB, characters, 511 — habitat and food of, 511 Tyroglyphus longior, 512 characters of, 512 habitat, 512 — minor, var. castellani, cause of copra itch, 513 — siro, 512 — — characters of, 511 U. Uganda, strain of Trypanosoma brucei from, 95 — syphilis in, treatment, 632 TJhlenhuth (and others), endotoxins in Try- panosoma equiperdum, 98 Ulcers arising from clothes louse infection, 711 — and boils due to invasion by Cordylobia anthropophaga, 592 — examination for protozoa, 746 — production by species of Enyaliopsis , 542 Ulcus tropicum, agent of, 122 Umbilicus, ascarides escaping from, 656 Unger, treatment of oxyuriases, 697 Uranotsenia, characters, 565 Urethra, fistulae of, arising from bilhar- ziasis, 642 treatment, 644 — larvae of Homalomyia canicularis found in, 585 Urethra, maggots passed from, 728 — male, presence of Trichomonas vagina- lis in, 53 Urinary apparatus, symptoms of bilharzia- sis mainly centred in, 641 — passages, invasion by ascarides, 692 Urine, amoebae found in, 45, 46 — human, aphides said to have been passed in (footnote), 532 — occurrence of Anguillula aceti in, 379 — presence of Nephrophages sanguinarius in, 490 — preservation of ova of flukes in, 472 Urosporidium, 194 — fuliginosum, 195 Urotropine in bilharziasis, 643 Urticaria, echinococcus cysts causing, 651, 652 — set up by Leptus autumnalis, 702 Uterus, cervix, polypoid tumour of, with Schistosoma infection, 643 Uzara in flasrellate diarrhoea, 625 Vaccine and emetine treatment combined in pyorrhoea alveolaris, 620 Vaccinia, cell inclusions in, 207, 208 Vagina atrophied in Acole'inoB, 297 — presence of Bhabditis pellio in, 377 Vaginitis, acute, due to Schistosoma infec- tion, 643 Vanilli.smus, so-called, cause of, 512 Varicose glands in filariasis, 402 Variola, cell inclusions in, 207, 208 Vegetable food, raw, avoidance of, in pro- phylaxis against Oxyuriasis, 697 — matter, decomjoosing, Tyroglyphida; in, 511 larvae of Homalomyia canicularis found in, 585 Veins, liver-flukes found in, 243 Vena cava and portal vein, communication between, how formed, 272 Verallina, characters, 565 Vermifuges, 669-675 Vertebrates, entamoebse of, 34 — experimental introduction of insect flagellates into, 104, 112, 737, 738 — internal organs of, latent forms of try- panosomes from, 73, 74 — multiplication of trypanosomes in blood of, 71 — spirochaetes in, 116, 122 Vesicles, formation of, in creeping disease, 730 Vesico-prostatic plexus. Schistosoma haema- tobium in, 273, 274 Vianna, histopathology of Brazilian try- panosomiasis, 88 — treatment of espundia, 629 Viereck, discovery of Entamceba tetragena by, 38 Vignolo-Lutari, case of intertrigo set up by Oxyuris vermicularis, 696 INDEX 899 Villot, larvae of Goirliidoe, 479 Vineg^ar, Anguilliila aceti found in, 379 — see also Sabadill vinegar Virchow, R., development of Trichinella spiralis, 423 doubtful case of human coccidiosis, 149 Echinococcus multilocularis , 356 Vital, liver-fluke in vein, 243 Vleming-kz's mixture, application in scabies, 706 Vog-t, C, on the Helminthes, 3 Vomited matter, spirochaetes in, 122 Vorticella in faeces, 206 W. Wagener, von, lesions produced by Oxyuris vermicularis , 695 ' — life-history of Oxyuris vermicularis , 467 Waldenburg-, experimental infection vs^ith Coccidia, 136 Walker and Sellards, experiments with dysenteric amoebae, 618 Walker, E. L., balantidiasis, 203 on Entamoeba histolytica, 40 | prevention and treatment of balanti- dian dysentery, 637 j Walker, Norman, treatment of scabies, 707 ' Walrus, host of Dibothriocephalus cordatus, \ 315 . ' Walsh and Riley, Rasahus biguttatus, 540 Beduvius personatus , 540 Warble uieo {uesiriace), liosts of, 594 j Warburg, extract of male fern in expulsion I of ancylostomes, 687 i Wasielewski, Hoemoproteus (Halteridium) \ danilewskyi, var, falconis, 152 [ Water, eggs of mosquitoes float on, 559 ! — filtered and boiled, as prophylactic ag-ainst bilharziasis, 644 — infected, avoidance of, in prophylaxis against Guinea worm infection, 676 — larvae of Stegomyia fasciata occur in all collections of, 574 — mature larvae of Ancylostoma duodenale capable of living- in, 454 j — receptacles, screening against mosqui- i toes, 636 j — stagnant, mosquitoes depositing ova in, j 553, 557 — transmission of trichomonad infection j by, 56, 624 j — weeds harbouring mosquito larvae, des- : truction of, 636 I Watercress, passage of larvae of Syrphidce into human beings through eating, 584 Watsonius, 234 — watsoni, 234, 235 — — diarrhoea in host associated with, 235 female organs, 235 habitat, 235 male organs, 234 morphology, 234 ova, 235 57 Watsonius watsoni, synonyms, 234 Weichselbaum, intestinal myiasis, 726, 727 Weidman, see Smith and Weidman Welland, Ascaris sp., 465 Wellmann, the ochindundu, 541 Wendelstadt and Fellmer, trypanosomes, mutation experiments with, 102 Wenyou, C. M., connection of Cimex sp. with Oriental sore, 536 possible host of Leishmania tropica, 108 on Entamoeba histolytica, 40, 41 on genus Cercomonas, 736 — spherical contracted forms of Tricho- monas intestinalis , 56 — supposed intermediate host of parasite of Bagdad sore, 575 — Tetramitus mesnili, 57 — transmission experiments with Trypano- soma lewisi, 92, 93 Werbitzki, blepharoplastless trypanosomes, 101 Wheler, Dermacentor reticulatus, 502 — length of life of Ixodes plumbeus (dog tick) apart from host, 495 — life-history of Ixodes reduvius, 494 Whip worm, see Trichuris trichiura White mice, experimental production of disease like leishmaniasis in, 103 infection with Herpetomonas cteno- cephali and H. pattoni, 103 — scour in fowls, causal agent, 145 Whitfield, A., and Hobday, F., transmis- sion of dog mange to man, 523 Whittles, nematode larvae in periosteum of upper jaw in case of gingivitis, 378 Wiggins, locust injurious to man, 542 Wijnhoff, cases of amoebae in urine, 46 Wild game, Trypanosoma rhodesiense pre- sent in, 81 Wilkinson's ointment, application in scabies, 706 Williams, Anna W., culture media for amoebae, 743 on cultural amoebae, 42 Williams, H. XJ., invasion of human beings by Trichinella according to nationali- ties, 428 Wilms, myiasis oestrosa dermatosa, 725 Winogradoff, post-mortem discoveries of Opisthorchis felineus, 252, 253 Wirsing, mode of infection of intestinal myiasis, 727 Wohlfahrt, myiasis cutanea from Sarco- phaga, 722 Wolff, treatment of cutaneous and muscu- lar cysticerci, 663 Woodcock, transmissive phase of trypano- somes, 737 Wood tick, see Dermacentor occidentalis Worm abscesses, formation of, 9 — electuary (Stork's) in expulsion of ascarides, 692 — seed oil in expulsion of ascarides, 694 " Wormlet " burrowing into human epider- mis, 599 900 THE ANIMAL PARASITES OF MAN Worms, intestinal, hereditary transmission of, former belief in, 11 of lower animals represent young stages, 21 spontaneous generation, belief in, 12 transmission by ova, discovery of, 11 Wounds, larvae in, movements of, 723 Wright, Bhinosporidium kinealyi, 197 Wurtz and Cleri, invasion by Loa loa, 678 Wyeomyia, characters, 565 Xenopsylla, distinctive characters, 545 — host of cysticercoids of Hymenolepis murina and H. nana, 328 — brasiliensis, 547 — cheopis, 546 carrier of plague bacillus, 543, 547 host of Trypanosoma lewisi, 92 Xeroform, application in Demodex follicu- lorum canis infection, 709 Xyphorhyncus firmus, 131 Y. Yaws, climatic distribution, 632 — inoculation with, experimental, 127 producing no immunity to syphilis, 128 — non-immunity to, produced by inocula- tion with syphilis, 128 — pathogenic agent of, 114, 127, 128, 632 — prophylaxis, 632 — species of Sarcophaga concerned in dis- semination of, 590 — stages of, 632 — treatment, 632 Yellow fever, mosquito carrier of, 574 Paraplasma fiavigenum said to be associated with, 180 transmission by Stegomyia, 555 — pigment in kidney and liver cells in ancylostomiasis, 647 Yorke and Blacklock, classification of try- jjanosomes, 72 — see also Blacklock and Yorke — see also Stannus and Yorke Z. Zarniko, case of Oxyuridoe in nose, 696 Zeder, special class of cysticerci established by, 282 Zeller, Echinococcus multilocularis , 356 Zenker, development of Trichinella spiralis, 423 — fatal case of infection by Trichinellae, 423 — Linguatula serrata, 527 Zenker's solution, 749 Ziemann, infection by Loa loa, 678 — varieties or sub-species of malignant ter- tian parasite, 167 Zinn, blood-stained diarrhoea from Strongy- loides stercoralis infection, 674 — extract of male fern in expulsion of ancylostomes, 687 Zooparasites, 1 Zschokke, experimental infection of man with Dibothriocephalus latus, 312 — Rhinosporidium in horses, 197 Zuelzer, on spirochsetes, 114, 741 Zululand, strain of Trypanosoma brucei from, 94 Ziirn, case of transmission of infection by Demodex folliculorum canis to man, 709 Zygotes of Coccidia, 141, 144 — of gregarines, 132, 133 ^' • ? THIS BOOK IS DUE ON THE LAST DATE STAMPED BELOW BOO ARE a REN. IIVIMEDIAIfc KfcV,ALL Mir FED 1 1 REC'D MAR ZZVTi EO'O \ ^«DV ftflAYl 1976 ER K. LIBRARY, UNIVERSITY OF CALIFORNIA, DAVIS Book Slip-Series 458 U 71201 Fantham, H. B. Animal parasites of man. QL757 F3 LIBRARY UNIVERSITY OF CALIFORNIA DAVIS