r rrd- Issued October 10, 1910. DEPARTMENT OF AGRICULTURE, BUREAU OF ANIMAL INDUSTRY.— Bulletin 125, Part 1. A. D. MELVIN, Chief of Bureau. THE GID PARASITE AND ALLIED SPECIES OF THE CESTODE GENUS MULTICEPS. I. HISTORICAL REVIEW. BY MAURICE C. HALL, Junior Zoologist, Zoological Division. WASHINGTON : GOVERNMENT PRINTING OFFICE. 1910. THE BUREAU OF ANIMAL INDUSTRY. Chief: A. D. Melvin. Assistant Chief: A. M. Farrington. Chief Cleric: Charles C. Carroll. Animal Husbandry Division: George M. Rommel, chief. Biochemic Division: M. Dorset, chief. Dairy Division: B. H. Rawl, chief. Inspection Division: Rice P. Steddom, chief; Morris Wooden, R. A. Ramsay, and Albert E. Behnke, associate chiefs. Pathological Division: John R. Mohler, chief. Quarantine Division: Richard W. Hickman, chief. Zoological Division: B. H. Ransom, chief. Experiment Station: E. C. Schroeder, superintendent. Editor: James M. Pickens. zoological division. Chief: B. H. Ransom. Assistant Zoologist: Albert Hassall. Junior Zoologists: Harry W. Graybill, Maurice C. Hall, Howard Crawley, and WiNTHROP D. Foster. Issued October 10, 1910. U. S. DEPARTMENT OF AGRICULTURE, BUREAU OF ANIMAL INDUSTRY.— Bulletin 125, Part 1. A. D. MF.LVIN, Chief of Bureau. THE GID PARASITE AND ALLIED SPECIES OF THE CESTODE GENUS MULTICEPS. I. HISTORICAL REVIEW. BY MAURICE C. HALL, 'f Junior Zoologist, Zoological Division. WASHINGTON: GOVERNMENT PRINTING OFFICE. 1910 LETTER OF TRANSMITTAL. U. S. Department of Agriculture, Bureau of Animal Industry, Washington, D. C, June 16, 1910. Sir: I have the honor to transmit herewith, and to recommend for pubhcation as a bulletin, the accompanying manuscript entitled "The Gid Parasite and Allied Species of the Cestode Genus Multiceps. Part 1. Historical review," by Maurice C. Hall, of the Zoological Division of this Bureau. Mr. Hall has been making a most comprehensive study of gid, and his investigations will furnish an important contribution to our knowl- edge of this deadly disease of sheep, which has only in recent years been recognized as established in the United States, the first definite evidence of its presence as an enzootic having been published in 1905 in Bureau of Animal Industry Bulletin 66. It is intended to publish later, as succeeding parts of the present bulletin, the results of Mr. Hall's investigations, now in progress, con- cerning the morphology and life histories of the parasites in question, as well as the symptomatology, treatment, prophylaxis, etc., of gid. Respectfully, A. D. Melvin, Chief of Bureau . Hon. James Wilson, Secretary of Agriculture. OCT 25 1910 CONTENTS. Page. Introduction 5 Multiceps multiceps 6 Historical sketch 6 Gid in the United States 16 Gid in Canada 29 The hosts and occurrences of the larval Multiceps multiceps 30 The occurrences of the adult Multiceps multiceps 41 Economic importance of gid 42 Alleged causes of gid 46 Names applied to gid and giddy animals 47 Common names of the gid parasite 49 Synonymy 50 Multiceps serialis 56 Historical sketch 56 The hosts and occurrences of the larval Multiceps serialis 58 The occurrences of the adult Multiceps serialis 63 Economic importance 64 Synonymy 65 Multiceps lemuris 66 Historical sketch y. 66 Synonymy 66 Multiceps polytiiberculosus 67 Historical sketch 67 Synonymy 67 Multiceps spalacis 67 Historical sketch 67 Synonymy 67 Cysticercus hotryoides 68 Historical sketch 68 Synonymy 68 Acephalocystis ovis tragelaphi 68 Historical sketch 68 Synonymy '. 68 ILLUSTRATION. Fig. 1. — Map of Montana, showing distribution of gid in sheep 25 3 THE GID PARASITE AND ALLIED SPECIES OF THE CESTODE GENUS MULTICEPS. PART I. HISTORICAL REVIEW. INTRODUCTION. Coenurus is the name comraonly applied to a larval cestode group of considerable importance to helminthologists from a historical and scientific standpoint, for it was with one of its species, commonly referred to as Ccenurus cerehralis, that Steenstrup's theory of the alternation of generations was first completely demonstrated for cestodes by Kiichenmeister, who, in 1853, produced the adult cestode or tapeworm in the primary host by feeding the larval form to the dog, and produced the larval cestode or bladderworm in the secondary host by feeding the eggs of the adult tapeworm to the sheep. This work of Kiichenmeister's and that of Von Siebold along the same line is taken by Braun (1894a), '^ in his classic work on cestodes, as marking the beginning of the fourth and latest period in helminthology, dating from 1851. This same species, C. cerehralis, is of considerable economic interest to veterinarians and stock raisers, and especially to sheepmen, as being the cause of the disease commonly known among English- speaking people as gid. In spite of the fact that the disease caused by this parasite, as well as something of its nature, was probably known in the fourth and fifth centuries B. C, and that the parasite itself was observed at least as early as 1634 A. D., its parasitic nature known since 1780, and its life history known for over half a century, there are still some mistaken popular ideas about it, and also some errors, disagreements, and uncer- tainties in the writings of scientists as to the specific identity of this and various other forms of coenurus that have been described from different hosts, and also as to the correctness with which certain par- « Bibliographic citations refer, wherever possible, to Stiles and Hassall's (1902-19 — ) Index-Catalogue of Medical and Veterinary Zoology, Authors, Bureau of Animal Indus- try Bulletin 39, United States Department of Agriculture. References not in Bul- letin 39 are indicated by the use of Greek letters and will be covered in a supplemental bibliography, to be published later. 5 6 THE GID PAEASITE AND ALLIED SPECIES. asites are listed from certain hosts. The writer has endeavored to correct some of these errors in this paper, and it is proposed in a series of papers to give a comprehensive account of the cestodes having a coenurus larva. The first form to be considered is the brain bladderworm of sheep, usually known as Ccenurus cerebralis, but which, as will be shown, should be known by the name Multice])s multiceps, proposed here for the first time. In this article the word "coenurus " will not usually be capitalized ; it will be used merely as the name of a larval stage, hke the words " cysticercus," "cercaria," "leptocephalus," etc. It is not entitled to be used as a generic or subgeneric name, owing to the pri- ority of Multiceps, but as it is still much more commonly used in this way than Multiceps, and as reference must be constantly made to quotations where it is used in combination with some specific name, especially in the form Ccenurus cerehralis, it will often be clearer to use this form instead of the correct one. MULTICEPS MULTICEPS. HISTORICAL SKETCH. Braun (1894a) makes his first period in helminthology cover the work of antiquity and the middle ages up to 1600, and in the litera- ture of this period, relatively barren from a scientific standpoint, almost no references are to be found that can be construed as refer- ring to gid. However, a disease like gid, involving, as it does, a deli- cate arrangement of alternating hosts, must have existed long before primitive man passed from the hunting to the pastoral stage. It is not the sort of disease to arise by rapid facultative adjustment or out-of-hand adaptation. The very fact that gid exists to-day is proof enough in a disease of this sort that it existed thousands of 3^ears ago. Undoubtedly, in the days when the ancestral dog pur- sued the wild sheep, the nice adaptation of a brain parasite that would interfere with muscular activity and blunt the sense perceptions, making flight and escape difficult, must have furnished a striking example of a life habit well calculated to perpetuate a parasite, but it could scarcely have been more satisfactory than the new arrange- ment introduced by man when he domesticated the sheep and put its former enemy, the dog, in charge of it to run over its pastures as a constant companion and to eat the discarded heads and diseased brains of giddy sheep — an enemy still. A prolonged search of ancient literature would no doubt show some references which might readily be taken as descriptions of gid. The symptoms are so striking that pastoral peoples, like the Arabs, Jews, and Greeks, must have noted and described them; but finding such references involves a tedious search and more time than can profitably be spent on the work. HISTOKICAL SKETCH OF MULTICEPS MULTICEPS. 7 One such reference occurs in Kuhn's edition of Hippocrates (1825^), who is beUeved to have lived 460 to 375 B. C. The follow- ing is quoted from Adams's translation of Hippocrates (1886ar), describing excess of fluids on the brain in epilepsy: This you may ascertain in particular, from beasts of the fiock [i. e. , sheep] which are seized with this disease, and more especially goats, for they are most frequently attacked with it. If you will cut open the head you will find the brain humid, full of sweat, and having a bad smell. It is, of course, impossible to make a positive statement of fact on anything less than complete and accurate observations. Obviously there was no one in the time of Hippocrates who could be expected to make and record such observations in a case of gid, and existing editions of Hippocrates are open to the suspicion of having in them observations not properly referable to Hippocrates. Hence we can not say certainly that Hippocrates actually saw cases of gid, but on the strength of the reference given, agreeing as it does with the certainty that gid among sheep must have existed for ages, it is fair to state that Hippocrates probably saw cases of gid four or five centuries before the Christian era. The fact that the brain of sheep was found full of fluid points, among other things, to hydrocephaly, which may follow the invasion of the gid parasite, according to Miiller (1877a), or to the gid parasite itself. Gid probablj'- was not rare in those days when sheep were everywhere tended by dogs and the prophylaxis of the disease was undreamed of. The "bad smell" may have been due to delay in post-mortem examination, to hydro- cephalus purulentus as a sequel of gid, or it may easily have been noted in the coenurus vesicle, as my own observations show that the coenurus fluid serves as an excellent medium for decomposition bacteria, the odor of the fluid in a graduate becoming intolerable in twenty-four hours at ordinary room temperature. Guetebruch (1766nr), according to Kuchenmeister (1880a), states in an article on gid that when perforation of the skull occurs, as it sometimes does in gid, the brain decomposes and becomes purulent, the brain and bone marrow turning to water and becoming putrid. The writer has never seen such a case, but it is evident that if the perforation of the skull were followed by perforation of the skin as well, it would afford entrance to bacteria, with possibly a result similar to the one given. Finally, the fact that these post-mortem findings are given for sheep suffering from "the sacred disease," a term covering epilepsy and other brain disorders, would indicate the possibility of gid, as the symptoms of nervous disturbances are very marked in this disease. Adams, the translator of Hippocrates from whom the foregoing quotation is taken, and himself a physician, refers to the lines quoted as follows : It is well known that this is also the case with sheep, and that they are subject to the disease called the sturdy [i. e., gid], which is indisputably a sort of epilepsy. 8 THE GID PARASITE AND ALLIED SPECIES. In the somewhat limited literature on helminthology for the period from 1600 to 1800, Braun's (1894a) second period, the gid parasite figures to a proportionally large and increasing extent. The citations from this period are given rather fully, as they are in works which are not readily available to many. In the first part of the nineteenth century, Braun's (1894a) third period, there are numerous references to gid, and since 1850 and the work of Kiichenmeister, which was done soon after, not a year' has passed in which few to many notes on the brain bladderworm, its adult tapeworm, or its effects, have not appeared. This increase in the amount of literature is perhaps concomitant with an increase in number and distribution of sheep and cases of gid, as well as with increasing knowledge of the parasite. In general the large amount of literature is due to the attractive combination of scientific and eco- nomic interest which has induced many persons to publish notes on the disease and its parasite from one or both standpoints. The early notes on coenurus deal only with Ccenurus cerehralis (= MnUiceps multiceps) and especially with the disease caused by it. It was nearly two hundred years after Scultetus (1672a) had seen the first unmistakable case of gid that I have found recorded, before the first coenurus which we may regard as other than C. cerehralis was noted by De Blainville (1828a). Scultetus saw his case in 1634. The first available note published during Braun's (1894a) second period of helminthology dealing with C. cerehralis is that of Rolfinck (1656a) who, in a work on medical anatomy, writes of vesicles full of water and humor in the third ventricle of sheep as the cause of a vertigo. This may be safely accepted as a reference to C. cerehralis. The description is in general terms just the one a casual observer would give of this parasite, as witness the statement of a correspondent to the veterinary editor of a periodical (Vet. Ed. Amer. Shepherd's Bulletin 1903;-) to the effect that he found in a sheep's head "a bag of water which burst and ran out when I pressed upon it." The next available article on the subject of gid published during this period is that of Wepfer (1658a). The part relating to C. cerehralis gives at this early date notes on the characteristic symptoms of the disease, its pathology, and the morphology of the water bladder. The disease is further recorded as a frequent cause of death in cattle, and the peasants are credited with a form of operation involving percussion and surprisingly good for that date. Heusinger (1853ar) quotes from a work of Bartholinus (lQQ7a), not available to me, a statement of a species of frenzy and vertigo which in 1661 attacked horses, cattle, and sheep, and notes that worms were found in the heads of the animals attacked. These cases may have included, and very likely did include, cases of gid. The next available article dealing with C. cerehralis is that of Scultetus (1672a), who in a Latin treatise on surgery gives the HISTOKICAL SKETCH OF MULTICEPS MULTICEPS. 9 description of a case seen at the earliest date at which we have found a case recorded. The following is quoted from an English translation of the same work (Scultetus, 1674a) : Observation X. Of a Vertigo in a Sheep, proceeding from an Abscess in the Brain. In the Year 1634, December the 24th. Being in the shop of Nicolas Kite « he made mention of his sheep, among which one was troubled with a Vertigo, or Giddiness, the Germans call it Wirbling: this Disease one who dealt in sheep affirmed to be inci- dent to the fairest of the Flock; that hereby their whole Brain would be turned into Water and then they would fall down dead on a sudden. The Chyrurgion therefore commanded that one of those sheep which was weakened with this Giddiness, and turning around, should be killed, and sent me the head. Scultetus found nothing in the ventricle. Afterward I lifted up the organs of smelling * * * and on the left -side, between the Brain and the Pia mater, I found an abscess, like the Bladder of a Fish, full of very clear water * * * I wondered that * * * the sheep should not labour under an Apoplexy, or a Palsy, rather than a Vertigo. In 1645 Scultetus lost a sheep by the same disease, and in the work just noted writes: I dissected the Head * * * and presently on the left-side as it were of the backward part of the Head, under the Dura Mater, I found a Bag of the thickness of a Fisches Blader, filled with Water, and little Worms, such as are bred in Cheese; for it began to putrefie at the bottom. This Coated Tumour being bigger than a Hens Egg, had so insinuated itself into the substance of the Brain, that it did somewhat press upon the third Ventricle. This Sheep, as the Shepheard reports, turned herself round about towards the night b all that day she dyed. That gid was not uncommon in the seventeenth century is clear from the fact that Rolfinck (1656a), writing of vertigo, refers to it as occasionally (nonnumquam) caused b}^ sacs of water on the brain m sheep. Wepfer (165Sa) notes it as a serious and common disease of cattle in Switzerland. In the account of Scultetus (1674a) it appears that a sheep dealer recognizes the disease as one common enough in Germany at that time to have a colloquial name, "Wirbling." The next reference to gid is by Wepfer (1681a) and is identical with the one already given, being in a later edition of the original work of 1658. Ktichenmeister (1880a) refers to an article by Brunner (1694a'), not available to me, and quotes from it a statement to the effect that Brunner had dissected the head of a giddy calf, "vituli vertiginosi," and in the cerebral substance had found three hydatids the size of pigeon eggs and full of limpid fluid. Kiichenmeister takes this to refer to Coenurus cerehralis, which it obviously does. a The original Latin text reads "in tonstrino Nicolai Reutte." The translator has translated not only the text but also the proper names, rendering the German name Reutte by its English equivalent, Kite. ^This last statement should read " towards the right," the Latin word here being "dextram." 51674°— Bull. 125, pt. 1—10 2 10 THE GID PAEASITE AND ALLIED SPECIES. The next reference is in Wepfer (1724a). The first two parts of this article consist of the two parts making up the edition of 1658. With these is incorporated a third part. The same references to gid occur in the parts already published and referred to above. In the new part is a new reference to hydatids in the brain of cattle as being commonly believed to be the cause of the vertigo accompanying them. He has seen the peasants perforate the skull and extract these in operations and has also seen the hydatids demonstrated post-mortem. Hoffberg (1759a), in a dissertation on Oervus tarandus, first presented in 1754, writes under the heading of diseases of this animal, of a vertigo or "Ringsjuka" causing the reindeer to turn in circles. Braun (1894a) takes this as a reference to Ccenurus cere- hralis, which is a i)erfectly reasonable assumption. The presence of the parasite in the reindeer, however, is unsupported by post- mortem evidence in this reference, and, so far as I am aware, such evidence is lacking in any subsequent writings. The occurrence of the gid parasite in the reindeer must therefore be considered doubt- ful. It seems the more doubtful in that Brehm (1877cv) states that reindeer are attacked by the larva of a gadfly, specified by Moniez (1880a) as CepJienomya trompe, which penetrates from the nasal cavity to the brain, causing a fatal "Drehkrankheit" or gid, and it may have been this disease, apparently a common one, which Hoffberg saw. Kiichenmeister (1880a) quotes from a treatise on diseases of sheep by Guetebruck (1766ci'), already noted as not available. In this treatise it is stated that the disease attacks lambs and yearlings, but not old sheep; that some are born with it; that a water bladder forms on the brain and may penetrate the skull; that when the disease has not gone too far the flesh may be used and the head and feet thrown away [very bad advice], but if the disease has gone too far the entire carcass should be done away with. As a method of treatment he gives venesection on the temple and nose. Stier (1776a) has an article on gid, of which only the review was seen by me, the original (Stier, 1775a) not being available. The article takes up a long list of supposed causes of gid and rejects them, the water bladder in the head being held guilty of causing the trouble. Stier also draws a careful distinction between actual gid due to C. cerehralis and simulated gid due to the presence of (Estrus larva* in the nostrils, the latter presenting the symptoms most com- monly mistaken for gid. According to footnotes in Bloch (1780a), Hastfer (1776a') and Ranstler (1776(t) have published references to gid, but these are not available. Bloch states that they attributed gid to the bladder on HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 11 the brain, and that Ranstler was the first to notice the small bodies on the bladder and surmised that worms arose from them. Accordino; to Braun (1894a) and others, the cestode nature of the water bladder found in the brain of giddy sheep was first pointed out by Leske (1780a) and by Goeze (1780a), independently. These references are not available to me. Braun notes that Goeze recog- nized the cestode heads and considered them as the embryos of the bladderworms which are found in the omentum and liver of sheep and swine. He also notes that Leske found Tsenia multiceps ( = Ccenurus cerebralis) , recognizing the characteristic hooks and suckers. Kiichenmeister (1880a) quotes part of Leske's article showing that Leske made a very careful study of the morphology and pathology of the parasite. He noted the heads invaginate and evaginate through the bladder wall. From the presence of so many of these heads, he observes that we may consider the animal as many tapeworms attached to a common bladder, or as one tape- worm with many heads. Hence it would be appropriate to call it the many-headed tapeworm, so he names it Taenia multice'ps. This last is important, as it establishes the fact that the correct specific name of the gid parasite is multice'ps. The preceding note from Braun (1894a) confirms the correctness of Kiichenmeister's (1880a) quotation, and in addition Mr. Sherborn has very kindly verified the reference in the library of the British Museum. It appears from evidence to be considered later that Leske's work antedates that of Goeze in the same year. Were it otherwise, Goeze's article need not be considered, as, according to Braun's synopsis, he regarded the heads of the parasite as the embryos of the bladder- worms found in the omentum of sheep and swine, and hence pre- sumably proposed no new name for the brain parasite, as there would be no reason for it under the circumstances or a proper appli- cation for the name had he done so. In a discussion of the synonymy of this parasite, Stiles and Steven- son (1905a) accept as the specific name the one proposed by Bloch (1780a). Bloch makes the genus Vermis vesicularis for the bladder- worms, and divides these into three species, of which Vermis vesicu- laris sociaUs is the brain bladderworm of sheep. But though this article of Bloch's bears the same date as those of Leske and Goeze, viz, 1780, Leske's article is nevertheless older, and the name pro- posed by him is therefore entitled to priority. This is evident from Bloch's own article, which shows that Bloch had read Leske's article of, the same year. Bloch states that Ranstler first noticed the small bodies on the bladder walls and surmised that worms arose from them, but that Leske and Goeze observed that these bodies were actually bladderworms. He states that Leske has described them 12 THE GID PARASITE AND ALLIED SPECIES. very completel}'^ and figured one accurately. Bloch very signifi- cantly adds that Leske numbered the parasites among the tape- worms, "Bandwiirmer," where, according to Bloch, they can not properly be reckoned, for reasons already given by him. Tt is evident from the last statement that Bloch had not overlooked Leske's Tsenia multiceps and that he believed he was correcting an error by proposing the name Vermis vesicularis socialis. However, subsequent work on cestode life history has shown the invalidity of all classifications which place vesicular worms in a group apart from the strobila forms and has justified Leske's judgment in uniting them. Unfortunately for Leske's name, Rudolphi (1810a) did not list it as a synonym of Cmnurus cerehralis, although he listed Leske's article in his bibliography. For this reason Leske's name has been very generally overlooked, as research in nomenclature has com- monly gone back through Rudolphi to the names quoted by him. Stiles and Stevenson (1905a) do not give Leske's name, Txnia mul- ticeps, in their table of synonymy, and in selecting the oldest name available to them have overlooked the rather obscure references to Leske's unavailable article. On calling Doctor Stiles's attention to the omission he pointed out to me that Sherborn (1902a) refers to Leske (1780a) with the comment "No n. spp." I wrote Mr. Sherborn, asking him to verify this reference, which he very kindly did. In a personal communication he quotes substantially the part quoted by Ktichenmeister (1880a), and states that he over- looked the name in his former reading. Mr. Sherborn was also good enough to supply copies of Leske's illustrations. These show very close observation. Following the independent discoveries by Goeze and Leske of the cestode nature of the water bladder from the brain of giddy sheep, there arose some controversy as to which of them was entitled to priority. According to Braun (1894a), Boerner (1780a) published an article discussing this point and holding Goeze as the discoverer. Subsequently, Goeze (1782a) repudiated Boerner's article, deploring the misunderstanding between himself and Leske. He states that he has explained the situation in a previous publication, the date of which is not given and which is unavailable to me. Leske's priority is C(mceded by Rudolphi (1808a) and by Davaine (1860a). The matter of priority here is apparently not concerned in the nomen- clature, and what honor lies in priority of discovery belongs to Leske, so far as the available evidence shows. Goeze (1782a) divides his genus ^^ Taenia, Bandwurm," into two main classes as he calls them — Tsenia visceralis, the visceral tape- worms, and Tsenia intestinalis, the intestinal tapeworms. Under the former he lists, among other species, ' ' Taenia vesicularis cerebrina" HISTOEICAL SKETCH OF MULTICEPS MULTICEPS. 13 from the brain of giddy sheep, Multice/ps, the many-headed, with many heads and bodies in a common bladder. And later on he states that from the numerous heads one may call the parasite "Vielkopf (Multiceps).'' From the above, Stiles and Stevenson (1905a) have taken the generic name Multiceps. The generic name used by Bloch (1780a) is evidently unavailable, being composed of two words and there- fore contrary to Article 8 of the International Code of Zoological Nomenclature, as given by Stiles (1905y): "A generic name must consist of a single word, simple or compound." Rudolphi (1809a) rejected Bloch's Vermis vesicularis as incon- gruous and unsystematic. Sherborn (1902a) is in error in listing Vermis Bloch 1782 as a generic name. The combination Vermis vesicularis is always used, whether with or without various specific names attached. As heretofore shown (p. 11), the earliest specific name of the parasite is that of Leske (1780a) as given in the name Tsenia multi- ceps. If the parasite in question is to be removed from the genus Tsenia, then the new combination must use the earliest available generic or subgeneric name, and since Goeze's (1782a) use of the scientific name Multiceps is evidently generic or subgeneric in intent, being clearly used to distinguish the many-headed gid parasite from the single-headed cysticercus forms, it is necessary to use it in the new name. The tendency for some time, and certainly a desirable tendency, has been to break up the large and heterogeneous group of animals formerly listed in the genus Tsenia, and to restrict the use of this name. The present situation has already been stated by Stiles (1905y) as follows: Most authors recognize that Taenia is to be divided into the subgenera Tsenia, Multi- ceps (i. e. Ccenurus), and Echinococcus. Some authors, however, incline to recognize these subgenera as of full generic rank. It seems advisable to restrict the generic name Tsenia to those forms which have a cysticercus stage in the life history. These alone make up a large group with a fairly close similarity in the adult and larval stages. To retain in this already large genus forms having a ccenurus or echinococcus larva seems unnecessary and undesirable. Long ago Leuckart (1886d) wrote: The Cosnwus * * * is related to the Cysticercus as a compound to a simple animal — a sufficient reason for systematic zoologists to separate them. Generic rank is accorded to particular groups of species which in the course of evolution have attained distinctive characteristics, and I see no reason for withholding sucli rank from forms in which these distinctive characteristics occur in the larva instead of the adult. This point is of especial importance in a case of this sort 14 THE GID PAEASITE AND ALLIED SPECIES. where the animal is found in the Larval stage in the great majority of cases, the adult being seldom seen or recognized. This view is in accord with that of Stiles and Stevenson (1905a), from whom the following is quoted: Opinions may differ as to whether this group [Multiceps] should be given generic or subgeneric rank. Personally we see no serious argument against recognizing a distinct genus on basis of the "larval" stage. Adopting, then, the genus Multiceps Goeze, 1782a, and the species multiceps Leske, 1780a, as the oldest available names, the correct technical name of the gid parasite is Multiceps multiceps (Leske, 1780a), Hall, 1910/9. From 1782 to 1800, the latter date marking the beginning of Braun's (1894a) third period in helminthology, numerous observa- tions were made on gid, most of them merely confirming the previous work of Leske, Goeze, and Bloch, or adding minor points of more or less importance and interest. By 1800 the gid disease had been observed certainly for over a centurv and a half and very likely for twenty-two centuries, its parasite had been named, described, and figured, and had a fairly large number of synonyms in addition to its correct name, the symptoms and pathology of the disease had been given, together with the symptoms of diseases simulating gid, and methods of operation had been used which only lacked aseptic pre- cautions to make them equivalent to good modern methods, and which were as good, perhaps, as most methods now in actual use. There remained, then, the work of finding out the life history and basing on that a rational prophylaxis. As a matter of fact the dis- covery of this life history by Kiichenmeister and Von Siebold marks the beginning of the fourth and last period in helminthology. The contributions of the third period to the subject of gid are largely WTong and unnecessary theories of causation as well as unsatisfac- tory methods of treatment. In addition, the large amount of litera- ture in this period lists the parasite from several new hosts, often erroneously, and adds considerably to the synonyms by which it is known. During this period new records of the disease show a widening geographical distribution, and unsatisfactory and unsub- stantiated statements of its presence in the United States begin to appear as early as 1809. The essential contributions in the literature of this period have been covered in tables and discussions to be given later, and the important events marking the modern period of helmin- thology may next be considered. Von Siebold (1844a) proposed as an explanation of the true nature of bladderworms that they were cestode embryos which in attaining a new host had gone astray, ending as encysted, incompletely devel- oped forms. Thus Cysticercus fasciolaris of the mouse was held to be such an incomplete sexless modification of Tsenia crassicollis of the HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 15 cat. He ventured to predict that in time the various tapeworms would be identified in their relation to certain cysticercus, coenurus, and echinococcus forms. Dujardin (1845a) advanced a similar theory, and this view or modifications of it became popular m scientific circles durmg the five or six years following Von Siebold's publication. It required the experimental work of Von Siebold and Kiichenmeister in 1851 and 1852 to complete this half truth. In the meantime the advo- cates of spontaneous generation lost ground to those who urged that the bladderworms were altered, degenerate cestodes or were incom- pletely developed embryonal forms. A prominent champion of the last theory, Kiichenmeister (1851e), finally published a note stating that he had produced Tsenia cras- sicipes [= T. crassiceps] of the fox by feeding Cysticercus pisiformis. A little later (Kiichenmeister, 1851d) he corrected this statement, changing his identification of the adult worm to T. serrata. This marks the beginning of the modern use of the now general experi- mental feeding methods of determining life histories. It remained for Von Siebold (1852a), the supporter of the theory of hydropic degeneration of bladderworms, to furnish additional proof that his theory was wrong, for this same year he produced the adult cestode from the gid bladderworm. The following year Kiichenmeister (1853e) succeeded in experi- mentally demonstrating, for the first time, the entire life history of a cestode. He fed Coenurus cerehralis to a dog and produced a tape- worm which he called Txnia ccenurus. He then fed the gravid pro- glottids of this tapew^orm to a sheep, and produced in it the early stages of the ccenurus in the brain. From this experiment Kiichenmeister concludes that sheep are infected in pasture by dogs dropping proglottids. Other animals, he thinks, may also harbor the tapeworm, and he claims this would certainly be true of wolves in Hungary and Poland. This statement is evidently mere assertion, as it is not verified by the record of such a finding either at the time or subsequently. At this date no de- scription of T. ccenurus had been published and its anatomy had not been studied. Indeed, the following year Von Siebold (1854b) states that he finds the adult of Coenurus cerehralis to be Txnia serrata. While the occurrence of T. ccenurus in the wolf is a proba- bility, it is nothmg more, so far as all available records show. On the evidence at hand Kiichenmeister formulated a set of rules for the prophylaxis of gid which is practically complete. It is as follows : 1. Feed dry food the year round and do not pasture. 2. Once or twice a year, purge the sheep and dogs in some inclosed place to get rid of tapeworms, and burn the feces. 16 THE OID PARASITE AND ALLIED SPECIES. 3. Do not, as is usually done, throw the heads of giddy sheep to the dogs, or, as Klichennieister after investigation finds to be done, throw the brain to the dogs before cooking the heads. Where there are wolves one must also bury or burn the intestines of those that are killed, and not throw them away to infect the fields. Such a program is not altogether practicable or necessary, but it only needs trifling amendment to bring it down to date. Had it been adhered to only as regards keeping dogs free from tapeworms and heads of giddy sheep away from carnivora for the last half cen- tury, gid would probably have been a rare disease by this, for it is really one of the most readily preventable of diseases. The next year Kiichenmeister's work was confirmed by Von Beneden (1854cv and 1854/?), Eschricht (1854CV), Gurlt — according to Kiichenmeister (1854n') — Haubner (1854c and 1854d), Leuckart (1854c), and Roll (1854^'), all of whom produced gid in sheep by feeding proglottids of Tsenia ccenurus sent them by Ktichenmeister. As a result of these experiments and others performed soon after, the important phases of the life history of the gid tapeworm were determined. It was found that the disease began with an invasion period during which the embryos were migrating through the body. Then followed an interval of apparent recovery, during which the growth of the bladdery vesicle was going on, to the point where the heads became developed and exsertile. Here the third and final stage of gid occurred, the characteristic symptoms, corresponding to particular locations of the parasite, becoming more aggravated with the increase in growth and number of heads until death occurred. Subsequent work has added to our knowledge of the morphology of the gid parasite, of the symptoms, pathology, and simulation of the disease, and of the need of avoiding bacterial infection in opera- tion. It has added numerous synonyms to the nomenclature, and recorded, correctly or incorrectly, new hosts and new areas of infec- tion, among the latter the United States. No essential points have been added to our knowledge of the life history of the parasite or the prophylaxis of the disease. GID IN THE UNITED STATES. The history of gid in the United States is, to a remarkable extent, a matter of conjecture. So far as I have been able to discover, the first claim of its occurrence here was made a century ago by Liv- ingston (1809ci'). His claim is based on very unsatisfactory evidence. The following is a rather full quotation of the case : The staggers or dizziness, which is also known by various other names, has occurred in three instances in my flock, and always attacked lambs under one year. * * * They were taken very suddenly * * * by a species of convulsion, in which the neck was twisted to one side; they lost the use of their legs; when raised they would GID IN THE UNITED STATES. 17 attempt to follow the flock, but turned round and fell; in a few days they were inca- pable even of standing, of moving their heads or any of their limbs. As they were very valuable sheep, I paid particular attention to them; grass and grain were given them, which they would readily eat, though they could not move any part but their jaws. In this state they lay a week without motion, except of their eyes and mouth. * * * In about ten days they could stand without support, but fell when they attempted to walk. * * * At intervals they would get better * * * but they were always found laying in some part of the field as if they were dead. * * * In the course of about six weeks they so far recovered as to be able to join the flock; one of them * * * received a blow * * * that killed him; the other two recov- ered, but very slowly; and even at the end of eight months they bore evident marks of their complaint. This disorder is found, upon dissection, to be owing to a bag containing water within the skull. * * * It may * * * be justly considered as incurable by the doctor, but not, as I have shown, by the nurse. * * * But a sheep must be extremely valuable to pay for three months' constant attention. It seems unlikely that the above cases were gid. Their occurrence in lambs fits in with the theory of gid, and the general symptoms, though not typical, might have been gid. On the other hand, the alternation between periods of normal activity and entire collapse does not look like gid, and the gradual betterment over a period of eight months runs counter to the clinical history of the disease. Moreover, leaving out the case of the lamb that was killed while recovering, the per cent of recoveries was 100. Some writers have claimed a spontaneous recovery in 2 per cent of all cases, but the writer knows of no evidence showing that any cases ever recover when the formation of the bladder is once under way, and a degen- eration of the parasite in its earher stages, indicated by the brain concretions according to Spinola (1858b), would not give a long period of slow recovery. Moreover, the three scattering cases given would indicate a sporadic infection, not to be expected in the case of gid. Doctor Mohler of tliis Bureau suggests a meningitis as the particular disease simulating gid in this instance, a theory which seems to fit the case very well. The lack of post-mortem evidence is unfortu- nate, as even typical cases of gid may be simulated by other things. Cole (1847a:), in a book published in Boston, discussing "Sturdy, or Water in the Head," states: A writer on this subject says that he knew a shepherd in Europe that saved nearly all on which he operated in this manner [by trocar], while he himself lost nearly all on which he operated. This sentence suggests that the writer referred to had operated outside of Europe and most Ukely in the United States, but this is, of course, mere speculation. Later, a competent scientist, Leidy (1856a and 1856b) records Coenurus cerehralis in a list of parasites "observed by the author," but does not state whether it was collected in the United States. McClure (1870^'), writing from the United States, says that he has known as many as five coenuri to occur in the brain of sheep. He 51674°— Bull. 125, pt 1—10 3 18 THE GID PARASITE AND ALLIED SPECIES. does not specify that this observation was made in the United States, however, or that the disease occurs here. Verrill (1870d), writing of gidj says: "In this country [United States] the disease is far more common than most persons suppose." Unfortunately, he cites no Hterature and no cases in support of this statement, and a request for further information has not been answered. Tellor (1879a) says: "Hydatid in the brain, or turnsick, although reported from New York and other States, is a curiosity rather than a scourge." He does not claim to have seen the disease. . Crutchfield (ISSOo'), of Hamilton County, Tenn., says: I have lost a few sheep by "staggers," "turnsick," etc., properly Hydatid on the brain, by allowing the sheep to range upon low, wet, spongy lands. By removing them at once the disease ceased. The evidence here is not sufficient to enable one to pass judgment on the case. There is no statement of symptoms or autopsy find- ings, and the cessation of the disease on removing the sheep from low, wet ground might or might not have followed in the case of gid. Hence this case must remain uncertain. Killebrew (1880<:tr), writing from the same State, Tennessee, in the same year does not claim to have seen the disease, but Stewart (1880a), writing from New York, says of Ccenurus cerehralis: "The presence of this parasite has been discovered * * * in numerous sheep in this country." Stewart's statement is not convincing, but in connection with other things it shows a belief on the part of men interested in the sheep business that gid existed in this country. Later events indicate that their belief and their statements to that effect are quite as likely to have been based on fact as to have been unfounded. Wernicke (1886a) records C. cerehralis from sheep in Buenos Aires. He believes it imported from Europe and states that it is a source of worry to breeders. It seems altogether likely that if gid had been imported to South America from Europe by 1886, it had probably been imported to the United States from the same source even earlier. In this connection, Powers (1887«') writes from New York the fol- lowing year concerning gid : I have never seen a case of this, knowing it to be such, nor have I seen an American shepherd who has met with it. It was probably imported from England, and it seems to prevail chiefly in the Eastern States. * * * i made many autopsies of sheep * ■* * for the bladder or cyst of this parasite, but I never found one. When the case is long drawn out, the bladder or tumour on the brain by constant pressure on the skull, absorbs it to such a degree that a finger pressed on the spot discovers a soft spot in the plate of the bone, or the latter even bulges out in a protuberance. * * * Twice I have seen this phenomenon in my own flocks and in rude fashion lanced them, thereby saving the sheep. GID IN THE UNITED STATES. 19 There is an evident contradiction between the statement that the writer has never seen gid and that he lias operated on his sheep for it. How easy it would be to import a case of gid may be surmised from Rabe's (lS89a) case in a gazelle imported from South Africa fourteen days before death. There is also the possibility of import- ing the adult worm in some of the numerous dogs which have been imported to this country. Professor Law, in a personal communica- tion, writes under date of July 2, 1909: Owing to its rapid development in the lamb it is less likely to be imported in the condition of larva, but among the many imported dogs the Tsenia must have been often imported. All things considered, the likelihood of importing the disease via the dog is perhaps as great as that of importing it in the sheep, but I would not consider the latter less likely. Rabe's case and others to be considered later show this. Moreover, a possible four to six months is not a very rapid development of disease in these days of rapid transit. An outbreak of gid attributed by Doctor Law and by Taylor and Boynton (1910a) to imported dogs is discussed later in this paper. The writer has collected evidence in Montana indicating that the gid parasite has been imported in dogs in some instances and the disease spread by the sale or gift of these dogs and their offspring. Nearly twenty years ago, Curtice (1890c) writes of larval cestodes in sheep : ' ' Tsenia marginata is more common in the United States, and T. coenurus next." He hazards the guess that in the West wolves, coyotes, and foxes may harbor the parasite. In a personal communication Doctor Curtice writes of the above under date of July 26, 1909: "I have never seen T. ccenurus. I must have made statement on information by reading." In another article Curtice (1892g) has the following: The tapeworms identified as T. coenurus were found but once in Colorado. The species may have been one arising from rabbit cysticerci and wrongly identified. The specimens were taken from a sheep dog. They are now in the bureau collection. I have examined these specimens (Nos. 2839 and 2840), and while they are not in good condition it is still possible to determine the essential things. They are not T. ccenurus, so far as the material furnishes data on the subject. To mention two evident differences, the eggs are decidedly oval, and the handle of the large hook is of an entirely different shape. About the year 1895 the subject of gid in the United States begins to receive notice in sheepmen's periodicals. Thus we find gid diag- nosed by the veterinary editor of one paper (Vet. Ed. Amer, Sheep Breeder, 1895fi') in a case where correspondents from an unspecified locality give a history of staggering to the right in an imported 20 THE GID PAEASITE AND ALLIED SPECIES. Slu-opshire ewe. The animal became unable to rise and was killed. On post-mortem examination a third of a teacupful of water ran out of the head. We are obliged to concur in the diagnosis given and consider that the disease was very likely imported with the sheep. Later in the same year the same diagnosis is given by this editor (Vet. Ed. Amer. Sheep Breeder, 1895/3) in a second case from an unspecified locality, with the characteristic symptoms of giddiness or turning, followed by death. Another case is diagnosed as gid on the same symptoms two years later (Vet. Ed. Amer. Sheep Breeder, 1897^). Sommer (1896c) did not find T. coenurus in an examination of fifty dogs at Washington, D. C. The adult tapeworm, T. ccEuurus, was reported from Nebraska by Ward (1896b), but Stiles (1898a) on an examination of the head of the specimen pronounced it T. serialis. Doctor Stiles tells me that he based tliis identification on the bifid guard of the small hook, an inadequate diagnostic character, as the corresponding guard of T. coenurus is also bifid. (See Reinitz, 1885a, and Ransom, 1905d.) On the other hand, the larva and adult of T. serialis are known to occur in Nebraska, which makes it likely that Stiles was correct. Ward (1897b) agrees with Stiles that it was T. serialis. Knowles (1897<:v) writes as follows: As numbers of inquiries come to this office relative to gid, or staggers, or so-called tumsick in sheep, I * * * append a well-written description, etc., of this dis- ease by Doctor Curtis. [This should be Curtice.] Doctor Knowles tells the writer that he saw his first cases of gid in Montana during the year that the above was written, 1897. Stiles (1898a), writing from this laboratory, says of Cosnurus cere- hralis: Fortunately it does not seem to be prevalent in this country. * * * It has been impossible for the writer to find any possible evidence of the existence of the gid bladderworm in this country, yet in view of the importations from Europe of sheep and dogs it is difficult to believe that we are entirely free from this parasite. In a footnote he says: One extremely doubtful case has been reported to us from Minnesota of its occur- rence under the skin of a horse. This latter case has not been examined by the bureau, but I would suggest that Txnia serialis is common in America, and consider- ing the tissue in which this parasite was found, it is not at all improbable that the Minnesota case was one of Coenurus serialis ( Taenia serialis) rather than C. cerebralis. Railliet's (1893a) earlier note of this case is based on correspond- ence. As this case stands we may choose between considering it as the first and only case of O. serialis in the horse and in its normal loca- tion, or regarding it as one of several cases of C. cerebralis in the horse, occurring in a location in which it has been reported twice GID IN THE UNITED STATES. 21 from the sheep. The case is too doubtful to pass judgment on, and the report may have been an error in the first place. Wallace (1900a) diagnoses a case for a correspondent from Iowa as gid in sheep. The symptoms are suspicious, but not clearly gid. Shaw (1901a), writing of the sheep industry of Minnesota, says that gid "has not been markedly prevalent in Minnesota." In a personal communication dated July 27, 1909, Professor Shaw writes: I have seen cases which I supposed to be gid in sheep, but I have never seen the parasite itself * * *, Dr. H. M. Reynolds, veterinarian of our [Minnesota] station * * * tells me that his experience is similar to mine. He has not yet seen the parasite. The veterinary ecHtor formerly referred to (Vet. Ed. Amer. Sheep Breeder, 1901;- and 1901^) diagnoses a case as gid in reply to two correspondents from Montana who describe the symptoms and post- mortem findings of their sheep. The diagnosis is unmistakably cor- rect. He states (1901o) that gid is "fortunately not very common except in the native sheep of the plains." Strictly speaking, the only native sheep in America are the Bighorn sheep, Ovis montana, of the mountains, never reported as subjects of gid. The reference is per- haps to native-bred sheep. The diseased sheep in this case came from Colorado, and the editor states : It [C. cerebralis] is especially common in Colorado, where 70 per cent of sheep examined by Doctor Curtice were infested by it. It is unquestionably quite as com- mon in all the western country from Mexico as far north as the animals mentioned [foxes, wolves, and coyotes] exist. It has already been noted that Doctor Curtice says that he has never seen T. cmnurus. Finally the editor states that he has recently operated on seven shee]) for gid. This is the first record of what appears to be a clear case of the finding of the parasite in the United States. On attempt- ing to secure further information about these cases it was learned that the veterinary editor in question was deceased. In another sheep-breeders' periodical (Vet. Ed. Amer. Shepherd's Buhetin, 1902