THE HEAD-CAPJULB 
 
 or 
 
 COLEOFTEPR 
 
 by 
 
 Fanner ^att^Fwaith Stickh^y 
 
 B 
 
. r\ ■ 
 
THE HEAD-CAPSULE OF COLEOPTERA 
 
 BY 
 
 
 FENNER SATTERTHWAITE STICKNEY 
 B. S. University of California, 1916 
 M. S. University of Illinois, 1918 
 
 THESIS 
 
 Submitted in Partial Fulfillment of the Requirements for the 
 
 Degree of 
 
 DOCTOR OF PHILOSOPHY 
 
 IN ENTOMOLOGY 
 
 IN 
 
 THE GRADUATE SCHOOL 
 
 OF THE 
 
 UNIVERSITY OF ILLINOIS 
 
 1921 
 

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TABLii OF CONTENTS 
 
 I. Introduction page 
 
 II. Acknowledgments 
 
 III, Materials 
 
 IV. Head-capsule 
 
 Epicranial suture 
 
 Vertex 
 
 Occipital suture 
 
 Occiput 
 
 Compound eyes 
 
 Oculata 
 
 Supratentorina 
 
 Ocelli 
 
 Antennaria 
 
 Antacoria 
 
 Antacava 
 
 Pretentorina 
 
 Front 
 
 Clypens 
 
 Postclypens 
 
 Clypealia 
 
 Mandibular i a 
 
 Preclypens 
 
 Lab rum 
 
 Occipital foramen 
 
 Submentum 
 
 Me tatentorina 
 
 Maxillaria 
 
 1 
 
 3 
 
 <7 
 
 12 
 
 14 
 
 19 
 
 20 
 
 21 
 
 21 
 
 22 
 
 22 
 
 23 
 
 23 
 
 24 
 
 24 
 
 24 
 
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 26 
 
 27 
 
 27 
 
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Cervix 33 
 
 Cervical sclerites 34 
 
 Odontoidea 35 
 
 Postgena 35 
 
 Grula 39 
 
 Spicraniiim 41 
 
 Paracoila 41 
 
 Postcoila 42 
 
 Precoila 42 
 
 Tentorium 42 
 
 Pre tentorium 44 
 
 Metatentorium 46 
 
 Corpotentorium 47 
 
 Laminatentorium 49 
 
 Supratentorium 49 
 
 V. Some Phylogenetic Considerations 50 
 
 71 . Summary 61 
 
 7II. Bibliography 65 
 
 VIII. Plates I - XX7II 
 IX. Vita 
 
Digitized by the Internet Archive 
 in 2016 
 
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I IITTHODUGTION 
 
 LeConte and Horn pulDlished in 1883 a classification of the Cole- 
 optera that has stood the wear of time remarkably well. Since then, 
 however, a number of new classifications have been proposed: lameere 
 (1900 and 1903), Ganglbauer (1892-1904), Handlirsch (1906-1908), 
 
 Kolbe (1901, 1908, and 1911), Sharp (1909), and Gahan (1911), all of 
 which differ more or less seriously in one way or another, and show, 
 for one thing, the need of further comparative morphological data, 
 
 / which is-, of course, indispensable to the building of any thorough 
 classification. leng's recent catalogue (1920) also emphasizes this 
 need. 
 
 A review of the literature seems to show but few studies based 
 
 of 
 
 on the comparative morphology of a comprehensive series^coleopterous 
 families. A number of European workers have published comparative 
 studies of the wings of Goleoptera, the most recent being by d’Orchy- 
 mont (1920). Sharp and 1'lu.ir (1912) and Lluir (1918) have published 
 the results of their inve stigations on the male genital tube in Gole- 
 optera. Various internal structures have been discussed from time 
 to time by a number of workers. Narrower in scope is the work of 
 d'Orchymont (1916) on the classification of the Hydrophiloidea, based 
 on a study of both the adult and the larva. Hyslop (1917), Bbving 
 and Gharaplain (1920), Graighead (1920), and Gage (1920), have pub- 
 lished papers on the comparative morphology of various families, 
 based on a study of the larvae. There are probably other comparative 
 papers more or less extensive in scope, but I have not been able to 
 find any such literature based on a study of the head-capsule. The 
 comparative morphology of the head-capsule of some other orders. 
 
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 however, has been investigated: Peterson (1915) on the Thysanoptera , 
 Peterson (1916) on the Diptera, Yuasa (1920) on the Orthoptera, and 
 Hoke (1922) on the Plecoptera. These simply draw attention to the 
 need of such an investigation of the head-capsule of Coleoptera. 
 
 With the "broader vision in mind of a more satisfactory and nat- 
 ural classification of the Coleoptera, the following study on the 
 comparative morphology of the head-capsule is offered. This study 
 does not aim "by any means to exhaust the su"bject. There have "been 
 too few species investigated in each family to justify the making of 
 any sweeping statements. This study can simply point out character- 
 istic conditions of structures as found in the different species of 
 the families studied, revealing, therefore, inharmonies, and perhaps 
 suggesting improvments on the present arrangement of the classifica- 
 tion. 
 
 In order to reach a correct estimate of the degree of speciali- 
 zation of the various parts of the head-capsule, an hypothetical 
 f yp6 . representing a supposed primitive condition, has "been construe* 
 ed. The structure of this hypothetical type is based on the struc- 
 ture of the head-capsule of generalized insects and of generalized 
 adult and larval Coleoptera. Hach structure has been treated separ- 
 ately, starting from the hypothetical type. The submentum has been 
 included in this study because of its bearing on certain develop- 
 mental processes. All statements made refer to the species listed 
 under ’’materials'’ only. The material studied was soaked in a 10^ 
 solution of potassium hydroxide until clarified, then washed in dis- 
 tilled vrater to remove the hydroxide, and preserved in 70% alcohol. 
 All dissections were made under a binocular microscope in 70% 
 

 
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3 
 
 alcohol in Syracuse watch glasses, 
 
 II. ACKUOWIEDGMEIITS 
 
 This study was pursued under the supervision of Professor Alex. D. 
 MacGillivray, to v;hom I am under the deepest obligations for all that 
 his supervision has meant to me in the way of helpful suggestions and 
 real inspiration, I must further thank him for permission to use his 
 unpublished morphological nomenclature. I am also greatly indebted 
 to Professor S. A. Forbes for suggestions and for furnishing a large 
 number of species from the collections of the Illinois State Natural 
 History Survey and from the collections of the University of Illinois 
 I am further greatly indebted to Professor E. F. Wickham of the Iowa 
 State University, who supplied me with a considerable number of spe- 
 cies belonging to rare families; to Mr, E, A, Schwarz and the author- 
 ities of the United States National Museum for representatives of 
 seven very rare families from the collections of the. Museum; to Mr. 
 
 W. S, Blatchley of Indianapolis for many very rare species; and to 
 Dr. Edwin C, 7an Dyke of the University of California for a specimen 
 of a species of Othnius. Of the many courtesies that Dr. Ghas. P, 
 Alexander of the Illinois State Natural History Survey has shown me 
 I am duly appreciative. Finally, to Mrs. Elizabeth Stickney, who has 
 helped me greatly in the preparation of the drawings, I am under deep 
 obligations, 
 
 III. MATERIALS 
 
 has been made to make this study as comprehensive as 
 possible, including not only a v/ide series of families, but also a 
 representation of the different subgroups within the families. Of the 
 eighty-one families , exclusive of the Strepsiptera, listed by leConte 
 3.nd Horn, representatives of all but one are listed in this study. 
 
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4 
 
 Leng in his recent catalogue lists one hundred and nine families. Of 
 these ninety-four families have been studied representing one hundred 
 and fifty-five species, of which one hundred and thirty have been 
 figured. The fundamental structure of the head is, except in a few 
 cases, practically similar for the two sexes. The sex has therefore 
 been disregarded, except in the case of the brentid, Eupsalis rainuta, 
 the male of which has a long slender snout, as contrasted with the 
 large broad snout of the female. The latter has been figured. 
 
 A number of attempts were made to arrange the figures in a lin- 
 ear series leading from the generalized to the specialized forms. All 
 attempts proved unsatisfactory. No matter what structure or condi- 
 tion of a structure was used, the structure showed itself to be un- 
 stable within narrov; limits and therefore could not be relied upon to 
 illustrate a definite line of development. However, the meagre re- 
 sults obtained in trying to arrange the drawings in a linear series 
 emphasized an important fact: that the various families of Coleoptera 
 and even the subgroups within the families, have developed along many 
 lines. For this study, the arrangement finally decided on, including 
 the species, is that adopted by Leng. This arrangement will be valu- 
 able, in so far as the head-capsule is concerned, in showing the need 
 for further morphological work towards the improvement of our classi- 
 fication of the Coleoptera. Owning to the number of drawings present- 
 ed in this study it was deemed more practical to omit detailed de- 
 scriptions. The salient features, only, of the various structures 
 are discussed. The following list is arranged according to Leng's 
 catalogue, and includes only those species studied:-- 
 
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5 
 
 GOLEOPTSHA 
 Suborder Adephaga 
 
 Caraboidea. 
 
 1 . Cicindelidae . 
 
 Megacephalini , Tetracha Carolina 37! ^ 
 
 Gicindelini . Cicindela formosa (Sigs. 3,US,Z4?^37Z) 
 
 2 . Carabidae . 
 
 Garabinae.-- Galosoma calidum .4, ^ZC>,ZS"0^3 73) 
 
 Harpalinae . Harpalus erraticus (Figs ) 
 
 Harpalus caliginosus 
 
 3. Amphi zoidae . Amphizoa lecontei (Figs. ) 
 
 4,0mophronidae . Omophroa americanum (Figs ) 
 
 5 .Haliplidae . Peltodytes 12-punctatus .7 /Z3 76> ) 
 
 6 . Dytiscidae , 
 
 Golymbetinae . Goptotomus longulus 
 Pytiscinae . Pytiscus hybrides 
 
 Cybistrinae . Gybister fimbriolatus (Figs . ) 
 
 Gyrinoidea. 
 
 7 . Gyrinidae . Pineutes assimilis , 7^ /3^Z3~S^37^) 
 
 Suborder Polyphaga 
 
 Hydrophiloidea, 
 
 8 . Hydrophilidae . 
 
 Hydroscaphinae . Hydroscapha natans (Figs. ) 
 
 Hydrophilinae . 
 
 Hydrophilini . Hydrous triangularis (Figs./<^ /wJ/ ^ :5"^ 37^ ) 
 
 Hydrophilus obtusatus JZ^ J3Z ZS/^^O) 
 
 Silphoidea. 
 
 9 .Platypsyllidae . ilatypsyllus castoris (Figs. 
 
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 lO.Leptinidae . Leptinus testaceus ^ S / ) 
 
 11 .Silphidae . T- Necrophorus Carolina i^ig^ • 
 
 12 . Scydmaenidae . — Connophron fossiger (Figs . ) 
 13.0rthoperidae . Molamba lunata (Figs. /6>^ ^ ) 
 Staphylinoidea. 
 
 14,Staphylinidae . 
 
 Steninae.-- iStenus flavicornis (Figs. 
 
 Paederinae . Gastrolobium bicolor (Pigs.^^JJ, / 
 
 Staphylininae . Creophilus villosus ) 
 
 Tachyporinae , i'achinus fimbriatus (Figs . J3/) 
 Aleocharinae . 
 
 Myrmedoniini . Aleodorus bilobata 
 
 Xenodusa cava 
 
 Aleocharini . Aleochara Ista (Jigs , /^/4Z^ 
 
 15. Pselaphidae . Pilopius lacustris (Figs. ) 
 
 16 . Ptilidae . Limulodus paradoxus 67^3^9) 
 
 17 . Sphaeriidae . Sphaerius politus (Figs ) 
 
 le.Scaphidiidae . -- Scaphidium quadriguttatum (Figs . >2^ ) 
 19 .Histeridae . 
 
 Hololeptinae . Hololepta fossularis 
 Histrinae.-- Hister merdarius (5‘igs 
 Saprinini.-- 2aprinus lugens 
 Oantharoidea, 
 
 20 .Lycidae . Calopteron terminale {Y\g^ . Z7^ /49^yZ7(r 
 21 .Lampyridae . 
 
 Lucidotini . Lucidota corrusca 
 
 Photini.-- Photinus pyralis 3 
 
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 23.Cantharidae . 
 
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 Cantharini . Podahrus rugulosus, Podahrus tomentosus, 
 Cantharis hilineatus (Pig, 
 
 24 .Ivlelyridae . — Gollops nigriceps (Pigs ^ ) 
 
 25 . Cleridae . Trichodes nuttali Z77^fB) 
 
 26 . Corynetidae . 3ecrobia rufipes (Pigs ) 
 lymexyloidea. 
 
 27 . Lymexylidae . Hylecoetus luguhris (Pigs. ) 
 
 28. Micromal thidae Micromalt hi dae dehilis (Pigs. ) 
 
 Gupesoidea. 
 
 29 .Cupesidae . Oupes concolor Z7f^ ^^0 ) 
 
 Mordelloidea. 
 
 SO.Gephaloidae . Gephaloon lepturides ) 
 
 Sl.Oedemeridae . 
 
 Nacerdini . Hacerda melanura (Pigs US;7S7y2S/, 
 hsclerihi.-- Asclera ruficollis 
 32 .Mordellidae . Tomoxia hidentata (Pigs ) 
 
 33. Phipiphoridae . — Macro siagon dirnidiatum * Z6,/Sf 
 
 34. Meloidae . 
 
 Meloinae.-- Ppicauta marginata ,3-^^ 7^0^ 
 
 Zonitinae.-- Zonitis atripennis 
 35 .jiuryste thidae Puryste thus dehilis (Pigs .3^ ) 
 
 36 .Othniidae . Othnius sp. (Fig". ) 
 
 57 .Pythidae . x^ytho americanus 
 
 38 .ryrochroidae . Neopyrochroa flahellata (Pigs . 
 
 39 .Pedilidae . Macratria murina (Pigs. 
 
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 iilateroidea. 
 
 41 .Hhipiceridae . Sandalus niger (Figs . ) 
 
 42 .iilateridae . 
 
 Pyrophorinae , — Alaus ooulatus (iigs. 
 
 Elaterinae . Melanotus communis 
 Cardiophorinae . Gardiophorus gagates 
 
 43. Eucnemidae . Tharops iruficornis ) 
 
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 Chrysobothrini . Chrysobothri s femorata 
 Agrilini.-- Agrilus ruiicollis 
 Dryopoidea. 
 
 46. Psephenidae r--Psephenus lecontei (Figs. 
 
 47 . Dryopidae . Eelichus striatus (Figs. 
 
 48 . Elmidae . Stenelmis sinuata (Figs . /74 
 
 49 . Heteroceridae . Heterocerus undatus (Figs.3'^/Z^ 4/f ) 
 
 50. Georyssidae . Georyssus calif ornicus 7 99^^^) 
 
 Bascilloidea. 
 
 bl.Eucinetidae . — Eucinetus morio (Figs . ^ 4-Z/) 
 52,Gyphonidae . Gyphon ruficollis /JS 
 
 Byrrhoidea. 
 
 SS.Bermestidae . Dermestes lardarius 
 54.3yrrhidae . — Byrrhus americanus CFlgs , 
 
 55.Nosodendridae . Nosodendron unicolor (Figs 
 

 
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 56 .Rhysodidae . Rhy&odes americanus 
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 57 .Ostomidae . — Tene5roides sinuatus 
 
 58 . Nitidulidae . — 
 
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 SO.Cucujidae . 
 
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 Dacnini . Megalodacne fasciata (Pigs. ^ i7/>2 
 
 62 . Derodontidae . — Derodontus maculatus (Pigs . ) 
 
 63. Gryptophagidae . Anchicera ephippiata (Pigs . ) 
 
 64. Byturidae . Byturus unicolor /f/, ) 
 
 65. Myoetophagidae.-- Mycetophagus punctatus (Pigs . 
 
 66. Colydiidae . — Philotherraus glabriculus (Pigs . %?/"^ ^88) 
 
 67 .Lathridiidae . Melanophthalma angularis (Pigs .7-^/>^.r ) 
 66 .Myoetaeidae . Phymaphora pulchella iligs . // /f6. d/f, ^C) 
 
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 70. Phalacridae . — Phalacrus politus 
 
 71 . Coccinellidae . 
 
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 7E.Alleculidae . — f seudocistela brevis (Figs . 
 
 73. Tenebrioni dae , 
 
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 Diaperinae. — Diaperis maculata 
 
 74 . Lagriidae . Arthroraacra aenea (Fi gs . 7^ 20SI OZ 
 
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 76 .Melandrjridae . Penthe obliquata (Figs . ) 
 
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 77 .Ptinidae . Ptinus brunneus (Figs . 
 
 78 .Anobiidae . Sitodrepa panicea (Fi^s . 
 
 79 .Bostrichidae . Boetrichus bicornis (Figs . ) 
 
 SO.Lyctidae. — Lyctus planicollis (Figs. 
 
 81. Sphindidae. — Sphindus americanus (Figs . 
 
 82. Cisidae . Plesiocis cribrum (Pigs . 
 
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 83,Scarabaeidae . 
 
 Aphodiinae. — Aphodius firaetarius (Pigs.fi^7/^ 
 
 Melolonthinae . Bichelonyx elongata (Figs. ^3,^ -7.7^) 
 
 Putelinae.-- Pelidnota punctata (Figs . ^70) 
 Dynastinae . Btrategus julianus (Figs . 7^/ ) 
 
 Cetoniinae. — Osmoderma ereraicola (Figs. ) 
 
 84. Trogidae. — Troz suberosus (Figs. 9i, z/9, 
 
 85.1ucanidae . Psudolucanus dama .?7 ZZC ) 
 
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 86. Passalidae. — Passalus cornuta (Figs 
 Cerambycoidea. 
 
 87 . Cerambycidae . 
 
 Prioninae . 
 
 Parandrini . Parandra brunnea 
 
 Prionini. — Derobrachus brunneus JC0,ZJ.s^^ J ) 
 
 Cerambycinae . Glycobiiis speciosus . /6/^ ) 
 
 ^Lamiinae. — Tetraopes tetraophthalmus (Pigs . ) 
 
 88 .Chrysoinelidae . 
 
 Donaciinae. — Donacia piscatrix (Figs ^7/) 
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 Griocerinae . Griocerus asparagi (Figs ^7^7 ) 
 
 Gryptocepbalinae . Gryptacephalus quadruplex 
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 Ghrysomelinae . Leptinotarsa decimlineata (Figs 
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 89 .Mylabridae . — Pachymerus arthriticus J/Z,Z3C>p^?^ "f S'O ) 
 
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 Brentoidea. 
 
 90. Brentidae . — Eupsalis minuta .jCfZZZ 3^0^ 4 ) 
 
 Gurculi onoidea . 
 
 91. Belidae. — Ithycerus noveboracensis 
 
 9S.Platystomiidae . Eurymycter fasciatns (Figs ) 
 
 93 . Gur culi oni dae . 
 
 xihynohitinae . xiiiynchi tes bicolor (Fi^s 
 
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12 
 
 Attelabinae . Attelabus analis JZC -3^4 d>S) 
 
 Allocorhyninae . Allocorhynus slossoni (Figs. ) 
 
 Otiorhynchinae . Epicaerus nubricatus J/6 ) 
 
 Gurculioninae . Lixns maoer 
 
 Thecesterninae . ^beoesternus Immeralis (Figs. 
 
 Calendrinae . Sphenophorus aequalis (Figs ./2/ ) 
 Scolytoidea. 
 
 94 .Ecolytidae . 
 
 Ecolytinae . i^colytus quadrispinosus ,/ZZ.^Z4:TjC:SJ‘^fo) 
 
 Eylesininae . Dendroctonus valens 3^9^^?/) 
 
 IV. HEAD-CAPSUIS. 
 
 There exists a distinct homogeneity in the general character of 
 the structure of the head-capsule of Coleopters. Its uniform strong 
 chitinization is typical, bo is the spacious area occupied by the ! 
 mouth-parts, producing a relatively broad cephalic end. Especially 
 characteristic is the wide space between the occipital foramen and 
 the submentum. Then, there is that indescribable similarity of struc j- 
 ture, even between groups widely separated, that can best be appre- 
 ciated from a thorough knowledge of the morphology. As an illustra- 
 tion, there is little superficial resemblance between Harpalus (Figs. 
 ^^ZS(J74) and Phalacrus (Figs , either in external or internal 
 morphology. Their distinct differences are merely due to two diver- 
 gent lines of development. The structures of the dorsal surface of 
 Phalacrus have become highly specialized, whereas those of Harpalus 
 are relatively generalized. On the other hand, on the ventral sur- 
 face the metatentorina has remained in a relatively primitive condi- 
 tion in Phalacrus, whereas in Harpalus its position is highly special- 
 ized. The internal structures of Phalacrus are rudimentary or lack- 
 

 
 
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13 
 
 ing, wiiile in Harpalus they are in a v/ell-developed primitive state. 
 Considering the degree of generalization of each species, there can 
 hardly be any question that Harpalus is the more generalized. The 
 above comparative description simply illustrates roughly the problems 
 of complexity of development that are encountered. The two distinct- 
 ive kinds of development as shown above for Harpalus and Phalacrus, 
 that of specialization of the dorsal surface and that of separate 
 specialization of the ventral surface, do not in the least necessari- 
 ly parallel one another in the Goleoptera. Indeed, these two lines 
 of development are predominantly divergent. In the majority of spe- 
 cies, the development is trending towards the obliteration of sutures 
 and consequent consolidation of sclerites, and towards the develop- 
 ment of a compactness of form of the sclerites that do not consoli- 
 date. The general trend towards a cephalization of migratory struc- 
 tures is a part of this process, too, as well as the development of 
 a stronger chitinize-tion of the head-capsule as a 'whole. The entire 
 phenomenon appears to be for purposes of strengthening the head. Be- 
 sides Phalacrus typical examples are Tenebroides (Fig.^’f) , Clischro- 
 chilus (Fig. 6/), Megalodacne (Fig. ^5), and all the Scarabaeoidea. 
 
 On the other hand these processes have lagged behind on the ventral 
 surf ace. in Phalacrus and others. It is true that the hypothetical 
 type (Fig.^y/) shows a marked cephalization of the submentum, with 
 obliteration of sutures resulting from this migration. Yet the pre- 
 dominant condition of the heads studied shows only a certain degree 
 of removal from the primitive type. Stronger chitinization has 
 probably kept pace with the cephalization of the dorsal surface, as 
 well as the growing compactness of such a structure as the submentum. 
 But the very significant structure of the ventral surface, the meta- 
 
 
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 11 
 
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14 
 
 ina, only shows a certain degree of removal from the primitive con- 
 dition in most of the heads. 
 
 Developmental processes such as are discussed above can be best 
 worked out through a comparative study of a large series of forms. 
 
 In fact the determination of the homologies of some structures enter- 
 ing into these developmental processes, as for example the various 
 changes in the epicranial suture, and the determination of the nature 
 and line of development of the area between the occipital foramen and 
 the submen turn, would probably be most difficult without this compara- 
 tive study. The chief value of this study lies in all probability 
 in the determination of homologies, to the end of understanding the 
 lines of development present. The homology existing between the var- 
 ious structures of the head-cai^sule of Coleoptera and other orders of 
 insects, particularly the generalized orders, seems to work out satis ■ 
 factorily. From such an homology the hypothetical type was construct • 
 ed without much difficulty. In general appearance it is oblong and 
 rather flattened dorso-ventrally. Such a form is fairly character- 
 istic of generalized insects and of the more generalized Coleoptera. 
 The mouth may be considered as directed cephalad. Such a direction 
 is representative of the vast majority of the heads, and for purposes 
 of convenience, at least, the following discussion considers the head 
 as extending cephalad. Some possible exceptions in which the head 
 appears to be directed ventrad are found in Calopteron (Fig. /■^^), Ma- 
 crosiagon ('Fig.Jff) , Tharops (Fig./7<^) , Throscus (Fig,/^f) , Byrrhus (Fip: 
 yfO) , and a few others. It should not be forgotten, however, that in 
 primitive insects the mouth is directed ventrad, and the occipital 
 foremen is on one side instead of at the opposite end. 
 
 Epicranial suture . -- The line of closure of the head in the em- 
 
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15 
 
 bryo is represented by the epicranial suture. The complete epicranial 
 suture is typical of generalized insects. V/here it is present in Co- 
 leoptera, this denotes a generalized condition. The primitive form 
 of the epicranial suture is that of a deep inverted Y, with the ce- 
 phalic ends of the arms near the lateral border of the labrum. The 
 hypothetical head is represented as having a complete epicranial su- 
 ture. The epicranial stem extends to a transverse line drawn through 
 the middle of the compound eyes. Branching here, the epicranial arras 
 continue to the margin of the head cephalad of the compound eyes. A 
 complete epicranial suture is not of general occurence in the Coleop- 
 tera. It is practically complete in Hydrous (Pig.^ ) and Hydrophilus 
 (Fig./Z), very distinct and sharply invaginated in both, particularly 
 so in the former, and characteristic in form. Each arm reaches the 
 margin of the head almost immediately cephalad of a compound eye, and 
 the arms are not as generalized in position as they are in Epicaiuta,^ 
 where they are quite distinct. The only other occurrence of a com- 
 plete epicranial suture is in Chelymorpha (Fig.//^) , Here the arras 
 meet the stem farther caudad than in the other genera named. The arms 
 in this last genus are distinctly curved, as contrasted with the more 
 or less straight arms in the above-mentioned genera. 
 
 The epicranial arms or some portion of them are present in all 
 Coleoptera, except possibly in Galopteron (Fig..^7)and Photinus (Fig.^^) 
 Dne or more species of every superfamily of the Adephaga and Poly- 
 phaga, except the Elateroidea, Byrrhoidea , Hhysodoidea , and Hhyncho- 
 phora, have the arms complete. In the Caraboidea they are prominent 
 as nearly straight sutures across the head, as in Tetracha (Fig.^), 
 Cicindela (Fig.J ), Calosoma (Fig. ^ ) , and Harpalus (Fig.^ ). Their 
 most generalized condition in the Adephaga is found in Omophron (Fig^F 
 

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16 
 
 in which they extend from the meson at a sharp angle. . Hep resentative 
 species of other superfamilies that have the arms complete are; Ne- 
 crophorus (Fig./;j)^ Tachinus (Fig.//), Chauliognathus (Fii?.j/)), Cupes 
 (Fig.J^), Gephaloon (Fig.j/), Uotoxus (Fig.fCg.), Heterocerus (Fig.^3), 
 Eucinetus (Fig.ii?), Mycetophagus (Fig.^/-) , Tenehrio (Fig.^^), Bostri- 
 chus ,Fig.^(^), Aphodius . (Fig.^iO), and nearly all the Gerarnhycoidea. 
 Species having parts of the epicranial stem preserved are not very 
 common. In Omophrom (Fig. 5), Tachinus (Fig.//), Penthe (Fig.^^), 
 and a number of the Gerarnhycoidea. parts of the cephalic end can be 
 identified; in Omophron (Fig. ^) , Phengodes (Fig.^^) , Gupes (Fig.J(^), 
 Sitodrepa (Fig. , Blepharida (i'ig.//<^) , and a number of the Hhyn- 
 chophora, parts of the caudal end are present. Ghalcophora (Fig. -^7) 
 Tetraopes (Fig./(PZ are peculiar in possessing practically all of the 
 stem but little of the arms. The arms in Ghalcophora are as short as 
 in any other species studied. Parts of the arms are present in every 
 degree of length from nearly meeting on the meson, as in Nosodendron 
 (Fig. .5*/) , to almost complete disappearance as in Ghalcophora (Fig.7f^) 
 and Hhysodes (Fig.//^). They also shov/ varying degrees of disappear- 
 ance and invagination, from the deep^ distinct invaginations of such 
 forms as Dineutes (Fig. ^ , Necrophorus (Fi?./^)^ Tachinus (Fi g./7 ) , 
 leterocerus (Fig.;T,/)^ and Arthromacra (Fig. 7/), to the faint or slen- 
 ier and shallow or not at all invaginated sutures characteristic of 
 bhe bcarabaeoidea. The character of the invagination associated with 
 the epicranial arms is not as simple as may be thought. In Harpalus 
 , the epicranial arms extend from the meson along the 
 5dge of the invagination to the pretentorinae , f rom which they extend 
 .0 the bottom of the invagination, curve late: ad, and continue to the 
 mrgin of the head. The course of the epicranial arms can be better 
 
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17 
 
 understood from Omophron (Fig. 5’), a related genus, which has re- 
 tained the cephalic portion of the epicranial stem. The line of the 
 invagination appears to be and often is considered to be simply the 
 clypeal suture. If a specimen of Harpalus is soaked for a long time 
 in potassium hydroxide the invagination can be opened and the various 
 structures in this region studied advantageously. The invagination 
 when opened (Fig./^f) will be seen to assume a deep wide wedge-shaped 
 form, extending entirely across the dorsal surface of the head. The 
 pretentorinae are located on the external dorsal surface just caudad 
 of the cephalic margin of the invagination. Altnougn the epicranial 
 arms are not in evidence anywhere between the meson and the preten- 
 torinae, it is assumed that the cephalic marginal ridge must repre- 
 sent them, since the pretentorinae are not only located caudad of 
 the invagination, but the arms are quite distinct, extending from 
 the pretentorinae to the bottom of the invagination, in which they 
 then curve laterad and continue to the margin of the head. From a 
 cross-section (Fig.^*^) it will be seen that the pretentorium expands 
 cephalad in characteristic form from the pretentorina along the epi- 
 cranial arm to the bottom of the invagination. In Galosoma (Fig.-^ ) 
 the same condition of this region is found as in Harpalus. In Omo- 
 phron (Fig. .5’) the epicranial arms are distinct between the preten- 
 torinae, extending from the meson along the cephalic border of the 
 invagination. From figure/^*^ it will be observed that the cephalic 
 border of the invagination in Harpalus is along the imaginary line 
 of the f ronto-clypeal suture. This border may represent the cephal- 
 ic limit of the front. The invagination, then, in Harpalus and Ga- 
 lOEoma includes the entire front. The line of the invagination in- 
 stead of being solely a part of the epicranial suture is in fact 
 ■■ 
 
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 A 
 
 
18 
 
 compound in nature, representing the approximation of the caudal bor- 
 ders of the front and post-clypeus , and that part of it between the 
 pretentorinae may be termed the clypofrons. . Laterad of the pre- 
 tentorinae to the margin of the head the line of the invagination is 
 readily seen to be an approximation of a part of the vertex with the 
 caud&l border of the postclypeus and cannot be included in the clypo 
 frons . Due to the more primitive position of the epicranial arras in 
 Omophron the invagination in this genus contains only a part of the 
 front, hence the line of the invagination between the pretentorinae 
 is simple in nature. In Tachinus (lig.//) the epicranial stem ex- 
 tends distinctly into the invagination, the arms continuing in the 
 same to the margin of the head. As should be expected, the preten- 
 torinae are within the invagination. The line of the invagination 
 in Tachinus is then of a different character from that of either Har- 
 palus or Omophron. In Tachinus, it has nothing whatever to do with 
 the epicranial suture nor with any other suture, being throughout 
 the approximation of parts of the external dorsal surface of the ver- 
 tex and the front. It is obvious from the above discussion of three 
 types of the invagination associated with the epicranial arms that 
 the dorsal surface of the head-capsule in Coleoptera must be studied 
 most carefully before a correct interpretation of the parts can be 
 made. This is most true in the case of any invagination that may be 
 present. The latter may not be readily observed when the head-wall 
 is strongly and darkly chitinized, necessitating treatment of such 
 specimens before the parts can be clearly made out. In some genera 
 such as Derraestes (Pig.5^) , Scolytus (Fig./^Xi » and Dendroctonus (Fig. 
 Il%) all external trace of the line of the invagination may be lost. 
 
 In such cases a true understanding of the parts can only be gained 
 

 t' 
 
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19 
 
 from an examination of the ental surface of the head. But in special • 
 ized forms the ental indication of the invagination may also he ef- 
 faced. 
 
 The epicranial suture can always he located from the determina- 
 tion of the position of the pre tentorina . The letter is always close • 
 ly associated with the epicranial suture, being present either in or 
 just off of the suture, in which case the pretentorina resembles a 
 sort of pocket. There is usually little difficulty experienced in 
 locating the suture. The cephalic ends of the arms are the most per- 
 sistent parts of it, being present when the remainder of the suture 
 cannot be identified. Interesting examples are found in most Hhyn- 
 chophora, where the remnants of the epicranial arms are represented 
 by short furrows located at the cephalic and of the snout. The epi- 
 cranial arms are typically structures of the dorsal aspect, but with 
 the shifting and modification of other parts of the head may be con- 
 fined to the lateral aspect as in Lineutes (?io-./^;f) , Helichus (Fig./^ 
 Adalia (Fig.^^) , or to the ventral aspect as in Gybister (Fig.^^) , 
 Hydrous (Fig.^^) , and Phalacrus (Fig.J^) . 
 
 Vertex. -- That part of the head-capsule not embraced by the 
 three primary sclerites cephalad of the epicranial arms, the occiput, 
 and the postgena, constitutes the vertex. Its extent is determined 
 by the form and size of the three above-mentioned areas. For in- 
 stance, in those species with much reduced epicranial arms, as in 
 Creophilus (Fig./^), Limulodes (Fig.^/), Adalia (Fig.//), and Phalac- 
 rus (Fig./>^), the extent of the vertex is correspondingly increased. 
 
 In the Phynchophora , as represented by such species as lixus (Fig./<:j) 
 and Bphenophorus (Fig./i/) , it is very extensive, including practical- 
 ly all (of the snout of the dorsal and lateral surfaces. 
 
 The area 
 

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20 
 
 located on the lateral aspect cephalad of the compound eyes, between 
 the latter and the epicranial suture, is the gena, a part of the ver- 
 tex. The limits of the gena are not definite. The prominent ridge 
 in many genera, dorso-raesad of each gena and antacoria, is the so- 
 called frontal ridge, that extends in the general direction from the 
 epicranial arras to the mesal margin of the compound eyes. The frontal 
 ridge is prominent in Harpalus (f ig. ^ ) , Ilecrophorus (Fig. 43). Tri- 
 chodes (Fig.x3^) , Pyrochroa (Fig.^^), Dermestes (Fig.-3X, and many 
 others. 
 
 Occipital Suture .-- In generalized insects the occipital suture 
 is confined to the ventral or caudal aspect, beginning near the lat- 
 eral margin of the postcoila and extending around the caudal or dor- 
 sal margin of the occipital foramen. In the Goleoptera this suture 
 arises lateral of the postcoila, extends cephalad for a considerable 
 distance, then curves abruptly lateral, extending onto the dorsal as- 
 pect of the head, where it joins the suture of the other side at the 
 meson. The genus Cicindela possesses the most generalized condition 
 of this suture found in the Goleoptera. The cephalic end of the su- 
 ture is modified into a ridge. This ridge is considered a later de- 
 velopment, and is not shown in the hypothetical type. It unquestion- 
 ably represents a part of the occipital suture, and can be identified 
 in practically all the Goleoptera, as in such widely separated groups 
 as Molamba (Fig.j^/) , nacerda (Fig.^//) , and Byturus (Fig.J/57. The oc- 
 cipital suture separates the vertex from the occiput and the postgena 
 Only the Garaboidea seem to possess with certainty an unmodified oc- 
 cipital suture. In Gicindela (Fig. 3 ) it is complete, and nearly so 
 in T.etracha (Fig.Z- ) but very faint in great part. In Galosoma (Fig. 
 ZS^t the unmodified suture begins farther caudal and is more distinct. 
 

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El 
 
 In Omophron (Fig.llJ) two short, characteristically curved, lateral 
 ridges no doubt represent remnants of the occipital suture. The ridge 
 across the lateral aspect in Peltodytes (Pig./2^) may also represent 
 this suture. In Cybister (Fig.Jf^ it is probably represented by the 
 crescent-shaped suture on the ventral aspect. Hidges or furrows ap- 
 pearing in the same general location in other species, such as Aleo- 
 chara , Throscus (i'ig.ZfJ) , Cyphon (Fig, JO/) , and Aphodius (Fig 
 
 337 >, may possibly be horaologized as occipital sutures. In most cases! 
 these ridges seem to be merely to mark the limits to which the head 
 is telescoped in. the prothorax. 
 
 Occiput . -- In those species possessing an occipital suture the 
 occiput is recognized as a distinct area. It includes the region be- 
 tween the occipital foramen and the occipital suture as far as the 
 postgena, appearing as a sort of broad band across the dorsal aspect, 
 divided by the epicranial stem and indistinguishably fused on the lat 
 eral aspect with the postgenae. Examples of a well-marked occiput 
 are present in Tetracha (Fig.>^), Gicindela (Fig. J ) , Oalosoma (Fig. 
 
 ■^ ) , and Earpalus (Fig. ^). In Omophron (Fig./^/) the occipital su- 
 ture is so short that the limits of the occiput cannot be definitely 
 determined. In those species not possessing a recognizable unmodi- 
 fied portion of the occipital suture, the limits of the occiput can 
 only be judged accordingly. Even in generalized insects the occiput 
 is nearly always fused with the pos-tgenae, and is so represented in 
 the hypothetical type. 
 
 Compound Eyes. There is a great similarity in the form and lo- 
 cation of the compound eyes. The general form is oval. They are lo- 
 cated near the middle of the lateral margin of the head. Such a form 
 and location is given in the hypothetical type. There are a number 
 

 
 
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 Yt Sft^'^ -f' '■ ^ : U' -rt.t' >c, aoiiop 
 
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 rtrt - A-C» , .Jw_a» » -; • • “^^ilflfvjlr' ‘•**'‘"'*'*( 
 
 • ■ If'y: “*. f ^ -**'#V 
 
 V . \« 
 
 f ^ i-'n iid^jfto 
 
 i/ri 'f v'X©»,x<'' iJi .Xi«»v4*-.|ti f' iw34. 
 
 o’, ... '’ .1 
 
 *<< 
 
 •Mcr 
 
 ,f. ' 
 
 h . . 
 
 33 i 
 
22 
 
 of interesting variations in form from tJie normal type. Dineutes 
 CEi.g,J3^ and Tetraopes (Fig.y<9Z) Jaeve four complete eyes. This phe- 
 nomenon is produced by a projection of part of the vertex. into the 
 eye that in time completely separates the t7i,^o halves. The line of 
 closure between the projection and the opposite side is indicated by 
 a distinct line--the exoculata. The beginning of such a projection 
 is shown in Cephaloon (Fig./cT4) , Fpicanta (Fig./^O) end many others. 
 
 In Cistela (Fig.^/)/) and Osmoderna {Fig.Z/3) the projection extends 
 more than half-way across the eye. In Throscus (Fig./^/) the projec- 
 tionneariy separates the two halves. The eyes of Peltodytes fl'ig./ ) 
 Fhotinus (Fig,Z9) and Stenus (Fig./^) are very large. Unusual forms 
 of the eyes are found in Hypophagus (Fig.^^), where they are very 
 long and narrow, nearly meeting on the dorso-meson; in Gryptocephalus 
 (Fig./<J^) , where they are prominent, crescent-shaped, and extend w'ell 
 caudad on the dorsal surface; and in Aphodius where they 
 
 are relatively small and square-like. The eyes of Limulodes (Fip-./^j) 
 are transparent and almost invisible; tnose of Leptinus (Fig./,?j) are 
 completely wanting. 
 
 Oculata. The oculata is present only on the inside periphery 
 
 of the eye as a broad ring-like shelf. It is considered of little 
 importance in this study. Its general size is indicated by the dot- 
 ted area within the eyes of Cicindela (Fig./Z5j , Dineutes (Fip.'./J,5)’, 
 PassaiuE (Fig.,2,2/) , and a few others. In forms with divided eyes the 
 two sides of an oculata are pressed togetner forming an exoculata. 
 
 Supratentorina. -- The supratentorinae represent the point of at- 
 tachment on the head-capsule of the supratentorium. They are not 
 thought to be primary invaginations and may probably represent no 
 more than depressions. In the Goleoptera the supratentorinae are not 
 

 
 ■ .h.' , 
 
 ^1 - *• 
 
 ’"} ii 
 
 *,THfw "j"'*' ’■ '1 _ 
 
 4 - 4 >uiW-jCt'' 4 »i 4 tae *«# s..ji 4 eiiaiirt tof : 
 
 ■ - _ m. -'® - to* ' 
 
 Ef-| .X 8 .> if**/' •ri'jt,/ 'U* T"' ^l-‘o ^oa^iv aoowi«>« 
 
 ’ ■ ' '.■ J'c. - i - ■■'.!- 3 .▼tap n" 
 
 €'*i‘ 
 
 ,.' 56 '. iP«rf ^i 5 f cUf f 
 
 ™ 'i ‘ » 
 
 3-, A ■■-•’f 
 
 
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 ifini *'‘je<-, cx^'T xo 'dwa' . ^.i' ■ 
 
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 nj ./!J St . •?•;!> 5^w' n.-?ii(f; ,9^« 
 
 
 
 [35^™ 
 
 ., *'K‘‘ - ■’ 
 
 
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 411^- , V;.' aw ' I L *u 4^8 ;fc3»r'YTifii I-isixo?^ eaJ xfo. 
 
 ^ •» 
 
 . 9Sf ilr r ■ 
 
 XlaajiTtiBri/isi^x 'rx^j 
 
 «:Jt 
 
 * V/.\.'i-^ acttx * ‘gel t*: ■» ;=^ 
 
 
 
 ] 
 
 
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 tx^fjt}tnq '^liUt i ^anajo.. „>yso cl a^t 
 
 ' >•) 
 
 ;.M? ie fMn >£l<iM-s ii ^r; . •’per' * . -H-* ofii t® 
 
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 jb ►fJi "iti i' if-.’ • t' i« « t b 
 
 h Uu. 
 
 
 
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 1 .1fcTA. ,iP» (jMireniC . 'E. A.-in.l, ^tnt4JWf^‘^a 9»V “''4* itm boi 
 
 ■■>U< 40 , « .v.ftf:x* fii ... 1 *ir --•»_# «2 ,!»s.^‘i*» »'ot*<a«'i 
 
 • ^ 
 
 . : ■,-»t»'’i|| ^^‘XXi^djD i*^ ^ i Oifir^i 
 
 . - : t r: a ▼ii? to »n d^i o-t'fiva ♦JiJ ><r j 
 
 .•> .r iAi-i wwtJM^is '^'LftjhiX j ad :i ^ 
 
 nAJihli .Bar,Ud^f^ab 4 j 4 ar/;««Ci^,; 
 
 • .-< #pi«pip 
 
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 4 
 
E3 
 
 prominent ,, as tne pretentorinae and me tatentorinae often are. They 
 are situated on the dorsal surface of the vertex. Their presence is 
 not general, occurring commonly only in tne Staphylinoidea . Outside 
 of tnis superfamily the supratentorinae are found only in Phyconomus 
 (Pig. and Philothermus (Pig./^). The supratentoria are usually 
 attached to the inner membrane of the body-wall, hut unless an actual 
 mark of their presence is indicated on the external surface the supra- 
 tentorinae are not considered as present. In generalized insects tne 
 latter are generally present. Their presence in tne Coleoptera 
 should indicate a primitive state. 
 
 Ocelli . -- In all the Coleoptera examined, no indication of any 
 ocelli has been observed. 
 
 Antennaria. -- In generalized insects there is a ring-like scle- 
 rite surrounding the periphery of each antacoria. In Coleoptera this 
 sclerite is present, but it is distinguished from the head-capsule by 
 a ridge only. On most heads it is considerably reduced in size, about 
 all that can be seen of it externally being its projection, the anta- 
 coila, upon which the scepe of the antenna articulates. On the other 
 hand, in Sandalus (Pig./J/) , Derobrachis (Pig.Zi^) , Tetraopes (Pig.^ 25 ), 
 Leptinotarsa (Pig.>2j/) and Anoplitis (Pig.;L3^ ,the antennaria is quite 
 prominent. The most generalized position of the antennaria is consi- 
 dered to be on the gena cephalad of the eyes, notwi thstandinpr that in 
 generalized insects the antennaria is quite frequently found distinct- 
 ly between the eyes. Embryology, however, shows that the antennae are 
 
 / 
 
 postoral in origin. Furthermore, in coleopterous larvae each anten- 
 laria is located cephalad of a compound eye. Such a position in cole- 
 Dpterous adults should denote the more generalized condition. The an- 
 tennaria is very unstable in position. There is hardly a superfamily 
 
vUl r-'wW** o'* •'* 
 
 >!■■.» 'V 'n ' ' i,K' 
 
 .« ‘ :- t a. . ^ 1 -^ ^a tr.- 
 
 
 . ) 
 
 ,f -.j- fc'. ^,as.■«aW 
 
 tooi: "«'■ rti 'va# V- noaisjCtt 
 
 ’*-tc“uco-./.'i. al ■;.1» bat«fr--*«- 0 Sai-iOW«dM^ e^^^»o|- 
 
 islt^. A itol’ijoi , ■'«?' - '■ ( tV-S « ' “■ 
 
 q.-T, -itiJ i-’inriftl’^’B ct :ttOt«i|tt<q, 
 
 t urflJ-. 'tet. aV • .jIhis's W «■' J.pa. ^^aiseJaaH 
 
 i*t\ >'y< 
 
 j-(*f»f,«*sO oil <- ' aiss'i '*. 
 
 . •, .« H-3 •rijai'it'a-i 
 
 '^k r-»pott!-.3.i.''i 0.1 ,6V.-te^ii- 7^.^ 
 
 ai;’’’V: ".’■ .»ov‘-:c?'^ .b#^‘ jkwi •ii'Ifo-.S- 
 
 ^ • - • a 
 
 •'' ^ f, 
 
 .■i’ 
 
 1 
 
 ,,. , -.vJrqroiB; ,M .looocfoo^Bt'afc 
 
 V;«»-«i-»i^vt.- a*' j.v- jsiwt uoM»oa»j.f.at6 i ij- ‘f;; *^4'**^*? 
 
 L|»''vs^ii> 'ct ^aoif .%li»diio^ia i'*| 
 
 U .i,oi4')'?-'.m .77 
 
 .rj B«1 ' Ti-/ | 
 
 a^iifr '.• a».'7ooW 
 
 Misir 1 !« 
 
 iJ .l.•..•^ ..X'.. ■-, ..a ?■■: ;< J-xj‘'ra|!j6 J6e»-«Ma‘* -^If.-W fi«-.»tj 
 
 :■ j: t im-- <i I ' ;ti ^ ^ -«D 
 
 .-• I*<*lii'-u ■■*'f.!i';M(^5 .itvvH S .•i»,0- .^'*>'2 *«»''i-» taewlOfl ^■ 
 
 0 0.^ cjlBi'.'tn .■.’ssuti .aw, '--'I'^''* * 3s%' »* “*;*f ' 
 
 W .■ : ...rs^fjiisea '^O* '♦loaVt W'.xaV> 
 
 * ' ' - ._ ^ _ 0 .V ai 0« K ir’ iV 
 
 ndT It I ' 
 
 AJ Iff* •i jrlTi^^i*-' 
 
 ' Ak 
 
24 
 
 in which it does not appear in both the generalized position and else 
 where. In the Scarahaeoidea and Gerambycoidea, though the position 
 of the antennaria varies within certain limits, yet it shows a char- 
 acteristic location. In the former it is either on the lateral or 
 ventral aspects, while in the latter it occurs only on the dorsal as- 
 pect. In no other large groups does the antennaria appear so con- 
 stant in position. In Galopteron (Fig.J/) and Phengodes (Pig.^^) it 
 is exceptionally large; in Dineutes and Alobates (Fig.24/2) it 
 
 is exceptionally small. 
 
 Antacoria . -- The membrane attaching the antenna to the head-cap- 
 sule is the antacoria. In removing the antennae the antacoria is 
 often torn, and as it plays no significant part in this study no at- 
 tempt was made to represent it in every case. The antacoria varies 
 
 in size depending upon the size and shape of the scape. It is indi- 
 cated in a number of figures by the stippled area; Omophron (Pig./.^7) , 
 IJecrophorus (Fig./^) , Galopteron (Fig.//^) , Phengodes (Pig./;r<5) , and 
 Ghanliognathus (Fig./^^). 
 
 Antacava . -- The depression in the vertex, usually in the gena, 
 within which the antennaria and antacoria are situated is the anta- 
 cava. It is always present, so far as is known, and is developed 
 into a deep socket in Dineutes (Fig./%?.fJ, Gonnophron (Fig./3^) , Sco- 
 lytus (Fig. 2“#^, and Dendro c tonus(Fig . 
 
 Pretentorina . -- The points of invagination of the pretentoria 
 on the head-capsule are the pre tentorinae . They are always located 
 along the epicranial suture in the Goleoptera. In generalized in- 
 sects and the more generalized Goleoptera, they are situated on the 
 
 lateral margin of the head. They are therefore represented in this 
 position on the hypothetical type. Ihe pretentorinae have been 
 

 -ifSl-nstXtXi ! 
 
 ■y^. 
 
 ra 
 
 ki'-' 
 
 •Vi 
 
 a' : 
 
 wn ' * ¥ i 
 
 •^^6 likk^ R ^J* -:,r ^ 1 I* 
 
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 nt .'srrt£f« n - 
 
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 1 ^ •* * * ** 
 
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 I j((. ^*’ B!# 
 
 tTf^- .to 9l J# 
 
 iu .ijo 4 ftiCsJttfirji 
 
 I i'^n^tl' eat 4 ;;!. -^iirC » 4 .t 
 
 ' ‘ «.i ^ '* V ~ 
 
 ■^ * .' ‘.=' 41 ' • » ^ U 
 
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 ^ ^ *“ j 
 
 r.- . .. ..VVfSfl- ■ •;:J> . % u«n<5 1 J ifi ^ 
 
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 -i» ^-1* ' 
 
 1-12 r-ft «niu* dll' oa £(S# ,n' 5 *it aafiro , 
 
 i ’ * ■ T ^ .,B ^ ^ . . . ; , . ->v 
 
 ^rt-T^'V -- f- / 4* ■?.*'# \»4t , 8tJ^ .L -• aiF^Sri acun , t^a^\ 
 
 i;^ ' ■ ?• ?S rklf-' 
 
 CtjiHti - Ia %4^MAh. 'J' .X^ ^P^$f3ir> /rlS af»<Tuj 3fti5^5■8^^ *>Jtiit-UJt 
 
 * ,<«j5. "I -.1 
 
 
 f. H 4 r^;i#;^!rO $ii£ 'ritr->isi .^'t« lo »- 
 
 i' .,• ^ •• ‘ i. -V, ? ^‘ ■ 
 
 r ■'• ^ '^- ¥ ^ V . . jm 
 
 , ?(<rcsf i*’' ) '. n^a- Ainel-jlr:? , !^\.3 ?ti:'* iU^ocfaotiDtJ!?^ 
 
 “‘‘'I' , ^ ' . .-.Yifrf ' .■' • 'i.r--j 
 
 w-.(¥^S*tV 
 
 •r" . ' . »»«>i rej ;ii J30*i*’4az<i(5i‘ r. dt *. r^ .-^ ^r g c- .ai jg^v 
 
 ■ , r^ *■ 
 
 ,: i.t b! I Ir 9T/?. L iiooAJfit; ^•v^®.al't(lA^^4KJ‘■R J 
 
 ‘■* - •■ 
 
 C<3. xjel (fft *tDra-Jen c^^.-^Xr ’-•' 
 
 ■-; ' ' ' • ■ * j . r.’J 
 
 -■ - 
 
 in’' , hC.%_ , 
 
 
 ^ ■• * fi— ‘ d .1C, 
 
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 , '' ’ I i 
 
 V. •*’ ^6 n»?iiat^*j ^•.’- i»a4 .. : -ar triViiaM _ft.‘ &r-» q1086 ‘ 
 
 
 
 : T: t 
 
 II “C !" 
 
 T Bite ; vutn - *'■ **X4 Cii . . *h|X^ Xeo/ ) M ^ s^ 
 
 
 
 
 .1 
 
S5 
 
 identified in every species studied except possibly Calopteron (Fig. 
 n) and Piiotinus (Fig.^Jf). In tiie latter they are represented by 
 depressions caudad of the eyes. Their position along the epicranial 
 suture varies greatly. In widely separated groups they may be prim- 
 itively located, as illustrated in such diverse forms as Oraophron 
 (Fig.:5'j, llecrophorus (Fig./^ j , bcaphidium (Fig..J3) , Ghauliognathus 
 (Fig.,?/)), Cephaloon (Fig.J/), Nacerda , Aiaue (Fir-.y-f) , Phy- 
 
 conomus (Fig.^3;, and Glycobius (Fig./<9/) . The position of the pre- 
 tentorina evidently cannot possess any important significance in 
 every instance, yet its position may be characteristic sometimes. In 
 the Scarabaeoidea it is never on the dorsal surface; in the Ceramby- 
 coidea it is always on the dorsal surface. In this respect, the pre- 
 tentorinae and the antennariae behave similarly. As a matter of fact 
 they are usually associated together, but there are some striking ex- 
 ceptions. The antennsriae ol Llacrosiagon (Fig.^^j are located well 
 caudad of the cephalic margin of the eyes, while the pretentorinae 
 are situated at the ventro-lateral margin of the head(± ig./ff) . The 
 
 opposite condition is found in Phenolia (1 ig./<^/j . The antennariae in 
 
 located 
 
 the vast majority of cases are/^caudad of the pretentorinae. 
 
 The pre tentorinae are the great landmarks of the head-capsule. 
 
 On their location the determination of the presence and position of 
 the epicranial suture is often dependent, and, consequently , the ho- 
 mologies oi large areas of the head-capsule. A case in point is that 
 of the xdiync hophora,in which the pre tentorinae are located near the 
 cephalic end of the snout, on the dorso-laterai margin. The epicra- 
 
 nial suture is reduced to the very short cephalic ends of the epicra- 
 nial arms, and though we cannot, therefore, indicate with precision 
 the cephalic limits of the vertex, its approximate limits can be 
 

 
 
 ■ » 
 
 
 
 1^,'' .*’''^ '*.i < ■ ’.J'i- " ■?■ ■'■'.^\^IH 'W 
 
 r.0i/!«o^ ^lr't^ ,. .ij,> 
 
 fc’’ ■''>■' ■- 
 
 . S;6 ^lO, t'S*^**^ i ^{Fi? I 
 
 |- ■ ■' w ^ ' TT *'"• \^ ' *-■ 
 
 A%l^^iy^) 45!/x£iilix£t) 
 
 ', '’t%-^ .ywiiJtWA , f’^^|||j||f?^ ‘<v ': ■ 
 
 A<,^n 1'/^ J •*•:>) . V^V:x*Vf . ' i/J-0^iC|;‘f«i^ ,j awraon 
 
 
 ^ 5'-?jr"' :4'C ’. ~*^ ' '\lJP-- • 'B 
 
 T ‘ ''! ■ *'* '- '^ i " a .»> •■^rf' •: 
 
 « .jw ^ . - . Mk ,i4 4 -. ■ * ^iL • ^ ■ij> . r^' ^ ^ Am^ i *■ ^ .^. v* A ^ -4A T. . A 1^ 
 
 X^jU4,gip .fi ;^!J^i--'X -C n.'^':»q 
 
 ^ ■*"' - jf V ' ' _ ' ^. _ -#»*' ,- ^ ■ ■ ^- y * 
 
 ^ -fil .oo^*t'i/r- £^-yj;''L<irh 
 
 , -^' 
 
 : - :.■ ^iSt '# ■ “’ I- 
 
 W- • J '- %: ' ' . -.1 if 'f '‘■JVia' V '»-■ , 
 
 ^ ■ ' ■■ H >'’ .' 
 
 ““^- ■ *x' f »V :.f ' ■" /■■ ‘ " ' ■* 
 
 
 V-/- '* *' ^- . ■ •' ' - Hfcj ^ ‘I 
 
 4 ?j.-’ -• £ To. »J ,v^-. 
 
 ■" ■■|’■■■^- - ' ••■?>*' * ' | ■ '-"r -if ~ i 
 
 <USrt . 
 
 -**. ■ X.l: 
 
 . -» 1 _ •• » * • .^ , _ • 
 
 P4 i if(^' - t^X’vu --f‘*‘ . f/c’.ifu I I xuj - V* *1/ 
 
 it n *’ . ‘T, , ' ^ ".(■'_ 
 
 ^ ■!'■ ■- . .;.*--■( •( :>df^j5-j*£ '4 -aiC^ :v u-iy^Cr 'tO' Jxl»;-*H g>fit 
 
 1 ,.«..-^c'j.. '^yKM *ij^ll^ ' "'* X-? ia ^ rr' c’'' 
 
 ( i;> ^ IM V».t : -it-Ua.; '' • -^'jt,e41-' fi.6 ‘ 
 
 •f»l^#:.:^-^i TH?J3I>' ‘'‘ i; >' , -y.lt , ' 
 
 i£( %i ;fili. i 3^., ^♦S vv ^ ^'il.t- r3 &j[^«x,£r‘ !%:£' - '»7130-X^I 
 
 ifflf ,evn ;-v?*<ni #t;» a.''' lii, » q;^ 
 
 t 
 
 w'-^ . 7i*^f» ‘11^ . .’ ■;. f**w V ■ j »~* ^-<- - i^'’ */ ••*' nr . •-'ViiJs'W 
 
 j - t ttl\. • tii'i t3C ^tis^ X*4lX<^v4^t 4^fiW| 
 
 ftf, i r i m: iH43lt. 419#!^^ ♦* • ' '®'i' ■-'» • * ' H ^.‘'' 
 
 
 yf 7itas^t 
 
 3 J -' i - f A«» iJiJta 
 
 ,<J* ] 
 
 • -it' 
 
 ,1 u ialaf ' 
 
 Ik 
 
 
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 /fi •.Mx'i' 5Xidl 
 
 fg . ■ %'H 
 
 
26 
 
 judged., wnicn would show the vertex to occupy nearly all the dorsal 
 and lateral aspects of the snout. An exception to the above-mention- 
 ed position of the pre tentorinae is found in the Scolytidae (Pigs.^-^f) 
 In which the pre tentorinae are located in the antacava. 
 
 Front . ~~ The sclerite embraced by the epicranial arms is the 
 front. In the hypotnetical type (Pig./ ) its caudal and lateral lim- 
 its are the epicranial arms. Its cephalic limit is indistinguishable , 
 since the front is fused with the postclypens. The approximate line 
 of fusion is indicated by a dotted line. There is no indication in 
 any head of a fronto-clypeal suture. The size of the front depends 
 upon the position and direction of the epicranial arms. In those spe- 
 cies possessing tne inverted Y type of arms, the front assumes con- 
 siderable proportions, but where the arms have been forced into a 
 more or less straight line across the nead, the invagination associ- 
 ated with the epicranial arms includes practically the entire front. 
 As mentioned previously, the epicranial arms extend in this manner a- 
 cross the head in a wide series of families. The front must hence as- 
 sume this form. In just as wide a series of families, the epicranial 
 arms are in process of disappearing. In such cases, the caudal lim- 
 its of the front can only be judged approximately. In great reduc- 
 tion of the arms, as represented by Phalacrus (Pig.J.^/) and Macrosia- 
 gon (Pig./:^), the vertex, the front and the postclypeus are indis- 
 tinguishably fused into one area. 
 
 Clypeus . -- In all species the clypeus is divided into two dis- 
 tinct sclerites, the postclypeus and the preclj^eus. This condition 
 is not present so far as I hnov; in the more generalized insects such 
 as the Orthoptera and Plecoptera. In some ITeuroptera, however, the 
 preclypeas is a large characteristicscleri te , quite similar to what has 
 
‘ f ^ \ ^ * ;^»' ' i - t * . K' ■ 
 
 
 
 ■** ’■*. 
 
 
 ' ■ * ?' 
 
 11^* 
 
 ■Ti* ‘ _ ‘ . €i 
 
 
 ‘^at '7i *^*tili|V^.i Cotst^f tt7 
 
 Kii ’ 
 
 
 ! '\tl eit - . 3i>riJU :• - 
 
 M. fJ 12Z ' '*■' in. 1 v» i, .^4£i ‘■C,i$:- l»i~ 
 
 -V ' -^ 'i,' ' 
 
 *^fj'^:U^a'Jt^>iibfiJk ei .ijf4.tr s; i ;*si^ do^ s^t 
 
 
 HB 
 
 
 (.«'.w,^i ...I-- '<, ..IV T«' 
 
 ^ - a* I >V - .-, ; ■' 2 
 
 
 ’.’’il^* 'S-j? I - » 
 
 s 
 
 :.a Ir V 1 ' •■; -Dr. i 1 6 n o ^ . 'ttot^sr r'‘ 
 
 "i/ • ' -■>■ •' 
 
 
 '«■ 
 
 ■^lU' ,3ni'i::< !v ■jq%^'^'iA^‘0’riArtii c^it &£tf^9^nna^\^0do 
 
 “ i^r ■- 
 
 W. ' ' * .. , ' ' ' '■' * • «i^, pi 
 
 (^ti-i iiaU frral'^i(t ’imis^xr,^4f\U 7iV.iU%tf »sfrjL ioj«rtc««4 ' 
 
 : , ■ - .. .-’ '.^ V .' ^ ' V-;'. '■' ■> % ' j ' 
 
 Va-i. ;'x-l:-' sii(fAP s : 
 
 ' T ^ V -V ■ -n 
 
 ^ - ■ ■ ■'^- y 
 
 <1*: JWDfr -ei?*^ * 
 
 li^lit*' ' f i*7iv " 
 
 19 
 
 rf K’ 
 
 4g V 
 
 
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 ' ■ - . - - . 1>2, t:;.idx '^6 ^s^i.Z9&: .jfh’iWiM ;isf^ 
 
 ‘■-•■.f ^ f'X>i;>''- Bj, uv}' xstff. ix2 ^ .fliiol 
 
 [ £ fl>v ‘‘QAis 4ii .- « iti 
 
 T ^ * - ■ T-' “ 
 
 ■ - ’ p t’n *xmr: * t • ^ ttjjno* 
 
 ~3l ^ 
 
 
 'S» 
 
 ■'" -i •- ^ *.■ 
 
 »% : 
 
 
 
 ' ^ 
 
 *?■ I-J 
 
 t 'iotnoii i j 
 
 viU /tS.*' rff' .-X.f)*- 
 
 *j04' ■^^.^:X ^6'- . 
 
 j.ii. , v • * »' j 
 
 \A Dij:; oitUi j 
 
 JiJ 
 
 '<• #> 
 
 f .' ^ fJin .'yjKJ' ■ t.J atl#0 . '.^i.OSl t , ^:- 't X'lfll Vft 
 
 
 pltfsiAt I -T.'.' ‘ : Ivt Q ' 
 
 r>5.‘'.J . • i«»¥ .' f-a i.:! 
 
 • »? 'jftp 
 
 
 « ;« 
 
 ^ r f Tt 
 
 
 f , ^4 Vc ^ lit >cri.4Af ,e^,Ni‘j..'X a 
 
 m iw aa r: " T"- 
 
 X -^lit 
 
 v(^£00t 
 
 
27 
 
 "been designated as tne preclypeiis in the Goleoptera. It may possibly 
 represent the extraordinary development and differentiation of the la 
 bracoria, but its size, shape and form would militate against such an 
 assumption. It seems much easier to believe that this area is a true 
 sclerite, and in this discussion it will be so considered. 
 
 Postclypeus. -- The broad cephalic part of the area between the 
 epicranial arms in the hypothetical type (Pig. / J represents the post 
 clypeus. In generalized Goleoptera it is of considerable size if the 
 dorsal surface has retained a generalized form. The shape, form, and 
 size of the postclypeus is correlated with the position and extent of 
 the epicranial arms, which has already been discussed. In highly 
 specialized forms like the Scarabaeoidea (Pigs .7^ , the postclypeus 
 may be even more extensive. Among the Staphylinidae , the postclypeus 
 may be very large in Tachinus (Pig.//) and Aleochara (Pig./r) , and 
 very small in Grsophilus (Pig.//). In Ghalcophora (Pig.^ff) the ceph- 
 alic and of the vertex is located on the ventral aspect of the head, 
 and the postclypeus is reduced to hardly more than a line. Among the 
 Gerambycoidea the postclypeus is generally very large. In Hhyncho- 
 phora it is quite reduced in size, except possibly in the Scolytidae. 
 
 Glypealia. — The caudo-ls teral projection or lobe of the post- 
 clypens is the clypealia. In Orthopters and Plecoptera the clypealia 
 is not separated from the remainder of the postcljrpens . In the larvae 
 of Gorydalis it is a very prominent sclerite. The clypealia in the Go- 
 leoptera is often separated from the postclypeus proper by a distinct 
 furrow or suture. It is quite loosely attached to the postclypeus in 
 the Gicindelidae (Pigs , the Garabidae (Figs ./^<^/i^) , 
 
 many of the Gerambycoidea i'^lgsJ^/o//C8) , and others. Difficulty is 
 often experienced in removing the mandibles from the head without 
 
- 
 
 
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28 
 
 detaching the clypeaiia. The close resemblance between the Ileuropterg 
 and the Coleoptera in other respects would lead one to believe that 
 this similar structure in the two orders must be homologous. The pre- 
 sence of this furrow in the Coleoptera is wide-spread, as a glance at 
 the figures will show. It can probably show little signiiicance as ar 
 indication of primitiveness. It must, though, have been present in 
 the primitive Coleoptera, and is hence shown in the hypothetical type, 
 Mandibularia. — In most of the Orthoptera a small triangular ares 
 is present, extending from the precoila to the cephalic end of the oc- 
 cipital suture. This sclerite is known as the mandibularia, IJo such 
 area has been located in the Coleoptera, 
 
 Preclypeus, — The dorsal surface of the larvae of Corydalis is 
 very generalized. On this surface there is a prominent submembranous 
 sclerite between the postclypeus and the labrum, the preclypeus. Such 
 a sclerite, very similar in size, form, texture, and position, is pre- 
 sent in Tachinus (Fig.//), Arthromacra (Fig,//), Trichodes (Fig.y;^,) , 
 and Glycobius (Fig./^/) . This sclerite is considered the preclype^. 
 Figure is a longitudinal section of the dorsal aspect of the head 
 of Harpalus, and shows the characteristic position of the preclypeus. 
 It is always present in the Coleoptera, though often considerably re- 
 duced in size. The preclypeus is always membranous except in ihotinus 
 (Fig.^^), where it is chitinized and the labrurn is membranous. Be- 
 sides the forms mentioned above, the preclypeus is large and prominent 
 in Necrophorus (Fig./J), Connophron (Fig./^), oiiacratria (Fig.-ji/) , Phi- 
 lothermus (Fig./2? ) , Hippodamia (Fig.^ ) , and many others. Very fre- 
 quently the cephalic end ol the postclypeus is infolded, thus carrying 
 the preclypeus and the labrum witn it. In such cases the preclypeus 
 cannot be seen irom the dorsal aspect. The preclypeus , , no matter now 
 

 
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29 
 
 deeply it is infolded, is except in a few ceses, sharply differentiat- 
 ed from the postelypeus and the labrum. In Glisehrochilus - 
 
 and G haul iogna thus the postelypeus and the lahrura were in 
 
 such close approximation that the preclypeus could not be observed un- 
 
 til the two above-mentioned sclerites were separated, and this was 
 possible only after long soaking in potassium hydroxide. In most of 
 the ^-ihynchophora, due to the fusion or absence of the lab rum, the pre 
 clypeus could not be identified. The preclypeus, hov/ever, was promi- 
 nent in Attelabus fPig./^^?j and somewhat reduced in Thecesternus (Fi.?. 
 and Spicaerus (Fig..?^^) , 
 
 Lab rum.-- The broad prominent sclerite attached to the cephalic 
 end of the clypeus in generalized insects is the labrum. In Goleop- 
 tera possessing other generalized structures, the labrum is typically 
 of the same general form. The labrum is so shown in the hypothetical 
 type. In position the hypothetical labrum should be, with the preclyp 
 eus, in accordance with their condition in generalized insects, on the 
 same general level with the remainder of the dorsal surface. The gen- 
 eralized form and position of the labrum is present in every super- 
 family except the Slateroidea, Dryopoidea, Hhysodoidea, Scarabaeoidea , 
 and the xihynchophora in which the postelypeus has been infolded, thus 
 forcing the labrum onto the ventral aspect. All the superfamilies 
 aontaining species with the labrum generalized, contain about as many 
 with it in various degrees of specialization, as to form, size, tex- 
 ture and position. The labrum in Gonnophron (Fig./^ ) , Photinus (Fig. 
 Z^) , Phiiothermus' (Fig./y ) , Melanophthalmus (Fig./^) , Eurymycter (Fig 
 //S) t and others, are large and membranous. The labrum of Aphodius 
 (iig.3«?7) is large but very thin and delicate. The labrum of Scolytus 
 (Fif.Jt;^ and Lendroctonus (Fig.^<$^ are probably membranous. In the 
 
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30 
 
 latter, a signif icant-looking slightly chitinized structure is locatec 
 in the memhrane within the mouth that may represent the lab rum. In 
 Thecesternus flig.^^/} there is a membranous area closely joined to the 
 postclypeus that probaoly represents the labrum. In iiupsalis (Fig.3^^) 
 lixus (I’ig.Ji^), Sphenophorus (Fig.^^y) , and hhynchites (Fig.J^z) , there 
 is an area within the mouth, bounded by furrov/s, that may be the la- 
 brurn. Such are particularly suggestive in view of the 'fact that in 
 the same location and lying flat against the postclypeus a very thin 
 but a relatively large and well chitinized labrum was found in Bpicae- 
 rus (Fig.3if) and Attelabus (Fig.J^jj. The labrum in Tharops (Fig.2?4 » | 
 Hosodendron {Fig.^bfj , Phyconomus (Fig.J^^) , and Perobrachus (Fig.j^^i 
 is considerably reduced in size.. It is present in every species, ex- 
 cept possibly the rhynchophorous genera named above, where it is al- 
 ways said to be wanting. 
 
 Occipital foramen. --The prominent opening in the caudal part of 
 the head is known as the occipital foramen. It is generally very 
 large, but in some of theLIordelloidea (Figs and in Connophron 
 (Fig.476), Cupes (Fig./:T;>T, and Bhysodes (Fig./^^j , is reduced in dimen- 
 sions, due to the constriction of the caudal end of the head. In Ca- 
 lopteron (Fig./^^; , Photinus (Fig./^^^j , Alaus (Fig./^^) , and Tetraopes 
 (Fig.^,^, the occipital foramen is extraordinarily large. 
 
 Submentum. - - In order to understand clearly the developmental 
 processes that take place on the ventral surface, it is necess-ry to 
 consider a sclerite, belonging to the moutir-par ts , the submentum. In 
 generalized insects the submentum is not only adjacent to but is one 
 of the covering parts of the occipital foramen, iiuch a position is 
 not found in the Goleoptera. Here, it is always located cephalad of 
 the occipital foramen with a distinct area between the two. In the 
 

 
 
 
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31 
 
 vast majority of heads this area is very wide. It is considered as 
 having been present in primitive forms, and is so sho?m on the hypo- 
 thetical type. The submentum in generalized insects is a large quad- 
 rangular movable plate. Many genera of Coleoptera show a similar 
 size, form , and mobility, as in Leptinus (Fig.2^<y, llecrophorus (Fig. 
 ZS‘f) , the btaphylinoidea (Figs .2^3^ Z65T, Nacerda (Fig.z^/) , Pyrochroa (Fig 
 , Alaus (Fig.^f/), the Dryopoidea (Fl^s , the Dascilloidea 
 
 (Fig. ) , and Byrrhus (Fig.^^^) . In these three characters, and the 
 additional one of position in respect to the paracoila, which in gen- 
 eralized insects is normally found beneath the submentum, th^t of the 
 Adephaga seems to be the most primitive. The innumerable sizes and 
 forms assumed by this structure throughout the entire series of fami- 
 lies can best be judged by glancing at the figures. It is extraordi- 
 narily large in Ahysodes (Fig.3^;.5r. 
 
 Metatentorina. — The invaginations on the external surface of the 
 head-capsule of the metatentoria are the metatentorinae . In the Or- 
 thoptera the metatentorinae are located along the cephalo-lateral 
 border of the occipital foramen as invaginations between the maxill- 
 ariae and the postgenae. They are not in any way associated with the 
 submentum in generalized insects or in the Coleoptera. I'ne same rel- 
 ative position of the metatentorinae is maintained in the Plecoptera. 
 In a number of Coleoptera, as in Helichus (Fig.2f^), Stenelmis (Fig.^^y' 
 Heterocerus (Fig.^'fy) , and even in the platystomid, Eurymycter (Fig.^^/' 
 this same generalized position of the metatentorinae is found. In a 
 number of Coleoptera the me tatentorinae are situated considerably 
 cephalad of the occipital foramen. The question might be raised as 
 to whether the metatentorinae that are so located could possibly be 
 more generalized in position than those situated adjacent to the 
 
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 r ' -/ ■.• ^- V T' ''1 
 
 ^ * tJXiO.f ' *'•■ ■ iXittVCJ rfX. -■-•'^♦ 0 < ^'iOjsJ 
 
32 
 
 occipital foramen. In every instance in which the raetatentorinae are 
 located cephalad of the occipital foramen, a suture connects the met- 
 atentorinae with the foramen. In only a few cases does the suture ex- 
 tend much farther cephalad than the me tatentorinae . This suture in 
 the vast majority of heads studied does not extend cephalad. It is 
 readily seen how the me tatentorinae might he drawn cephalad and as a 
 result a suture he formed marking their line of migration. In such a 
 process one would naturally not expect to find a suture located ceph- 
 alad of the me tatentorinae , and in the cases in which the suture does 
 extend so it is easy to understand that tne force of the cephalic 
 pull might have been communicated to this region, producing in conse- 
 quence a suture or invagination. Due to the shape of the head, it is 
 most difficult to believe that tne me tatentorinae could be drawn caud- 
 ad, and if they were so drawn, it would seem that in this process 
 there would be formed a suture cephalad of the mets tentorinae , mark- 
 ing the line of migration. In this discussion tne generalized posi- 
 tion of the me tatentorinae will he considered as that of their general 
 ized position in more primitive insects, at or near the occipital for- 
 amen. 
 
 In the development of the coleopterous head the metatentorinae 
 have Shown a tendency to migrate cephalad. The cephalic migration of 
 of the metatentorinae and the ventral migration of the pretentorinae 
 and other structures, were no douht due to the same force, the result 
 being a closer approximation of parts, which naturally supplied in- 
 creased firmness to the head's mechanics of operation. The Dryopoidea 
 show the most generalized position of the metatentorinae. lenera of 
 this superfamily have been mentioned above. The Elateroidea 
 probably possess the next most generalized metatentorinae, such as in 
 
.Vfc •,>uj^'sie £>^\ ‘ ? jpi •Vty.-i'i' ttl •»: v*« ** nfTo^xtetiJ 
 
 O! ' ' ' ' '^ ' '"* 
 
 % ..uUiMsi at^o.1. *' ri4:j0i.is« -^a^ 
 
 ftl .tvoi:#-i-:i3.iiB io i'all xld^i ' d<f t/.Xf«»'t- 
 
 ^%;^7u3 « .' j i«^X<l /oti i*X«Oir B30«0*I(J 
 
 tny ai ;^ttm4trnS to BalA 
 
 ■ 1} ► . ■' ' . ■ . X 
 
 ^ 7, :>j ..-!0^'/x et-t:; v*' v ^f(5iir'^XX-tfq. 
 
 :c »,' « Qg<f j^*>if)l';-- t ' ‘^ftf.tjitj a MCeiT^ 
 
 ^ift^o-xcr fiii f »t --id, OX^C •? 
 
 i>i/ .-?'?f '3.ir »:'iiL «fi^ Ti* i’ e 
 
 xt^it ^n •1‘Tf «cf>f 
 
 ^■^ili^Z^^iyao V«s > ;**oq Xl^sij 
 
 - Oit'^ M, X-Co a^oOM OXrjdj 
 
 *•’5'^ t» MMiHilA U.T ©*>*u:« •'« t,<>| 
 
 kfXt't-T 1^'*' aG'9® -J#5f'’ ffj^it«ypt> f'‘P ,*‘-lt;l<i-* .1 >< ‘1,4*1 TO 
 
 I ' ' ' - " !■ l 
 
 ’ ^-sU . 7,H.‘!ai4a * in «QX-4# -. *S t- ^:si© 
 
 ■•!f 4 *■ ' -^ _, ' I 
 
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 t ' • ■ i » ’ ] 
 
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 % t dflXJ vfdil^ 
 
 fliS &. 
 
33 
 
 Sandalus and Alans The Cucu;)oidea shov; the meta- 
 
 tentorinae just a little removed from the occipital foramen, as in 
 Kegalodacne fFig.J/ij , Anchicera (Fig.,?y^) , Philothermus (Fig.^//) , and 
 others. In the genera of other superfamilies, as l!losodendron (Fig.J<^' 
 Anoplitis (Fig.^;9^ , and Tetraopes (Fig.?^^) , almost the same degree of 
 primitiveness is shown. The large superfamily Mordelloidea show the 
 me tatentorinae to have migrated to a'bout half the distance between the 
 occipital foramen and the submentum. This condition is fairly consis- 
 tent througn the group. In the Bostrichoidea, the Scarabaeoidea , and 
 the Cerambycoidea , the me tatentorinae show considerable variation in 
 position. This is shown by a comparison of their position in Bostri- 
 chus (Fig.J^j^ and Bphindus (Fig.:?^^. In the families to which Tetra- 
 cha (Fig.2-y<^ , Calosoma (Fig.2;J>) , Cybister (Fig.^cJ^ , Dineutes (Fig^c?"^, 
 Hecrophorus (Fig.2;r^) , and Glicobius (Fiv.jy/) belong, the metatentori- 
 nae have aavanced very far cephalad, near to . the submentum. In Pho- 
 tinus (Fig, ^7^ and Ghauliognathus (Fig.iT^) they are located on or 
 quite near the paracoila. 
 
 Maxi liar ia . -- There is a narrow plate surrounding the lateral anc 
 caudal margins of the occipital ioramen in some generalized insects, 
 between which and tne postgenae the me tatentorinae are invaginated. 
 This plate has disappeared in the Goleoptera. 
 
 Gervix. -- Tne structure connecting the head-capsule with tne pro- 
 thorax is called the cervix. It is normally composed of membrane, 
 and a number of cervical sclerites. The size of the cervix depends 
 upon tne size of tne occipital foramen, and tne degree of mobility of 
 the head. In the lampyroidea and some other forms the cervix is very 
 large. The cervix in fihysodes (Fig, 30-^) is composed of tough fibrous 
 membrane, quite different in structure from the normal cervix. In 
 
W?: ■ i 
 
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 lit ’C • v**? • »i-. r ‘'^ .^?"4 
 
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34 
 
 nearly ail of the Haynchophora the cervix is heavily supported hy 
 strong tendons attached at its cephalic end. These tendons take care 
 of the added strain on the cervix due to the elongation of the snout. 
 There are more or less small cervical tendons appearing occasionally 
 tnroughout the whole series of families studied. In Ilolamba (Fig.^^/) 
 the cervix is invaginated within the prothorax, doubling upon itself. 
 The cervix in Bostrichus (iig.^3^ doubles back upon the head-capsule, 
 which is produced into a round projection. 
 
 Cervical sclerites. -- The most prominent and persistent cervical 
 sclerites are the pleural cervical sclerites, the cervepisternrun and 
 the cerve.pimeron. The former is usually the larger of the two, and 
 articulates at its cephalic end either against an odontoidea or sim- 
 ply against the undifferentiated area surrounded by the occipital 
 foramen. The latter usually extends in a dij'lerent direction from 
 the iormer, and articulates at its cephalic end with the cervepister- 
 num and at its caudal end with the prothorax. In Gantharis (Fiec.7/5)', 
 Macratria (Fig.i’?^ , Psephenus {Pig.7^, and others there is a single 
 large sclerite present. In the Adephaga, in Leptinus (Fig , Hy- 
 porphagus (Fig. 77^ , Phyconumus (Fis.yo^, Pseudocisteia (Fig. ^7^, and 
 many others, there is a single small subcircular sclerite present. 
 Both of these types probably represent the cervepisternum. In a very 
 large number of genera cervical sclerites are always wanting. They 
 are poorly or not at all developed in the Cerambycoidea. None of the 
 Nhynchophora studied possess a cervical sclerite except Furymycter 
 (Fig.J^/) , in which it is very small. The ventral cervical sclerite 
 is the cervisternum. Nacerda (Fig. 7?/), Gucujus (Fig.^//)) , Hister (Fig 
 Tenebrio (Fig.^i^) and a few others, possess two small small cer- 
 visterna, while Tomoxia (Fi^.7?^ possesses a long narrow one. The 
 

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35 
 
 cervis ternum is on tne wnole of infrequent occurrence. i"iie dorsal 
 cervical sclerite is called tne cervinotum. It occurs even less fre- 
 quently tnan tne cervis ternum. Hydrous (Pig.//) possesses a subquad- 
 rangular distinctly chitinized cervinotum. In Aleocnara (Pig./57 tne 
 cervinotum is divided into two distinct ligntly cnitinized subtrian- 
 gular sclerites. Two much larger square-like scierites are situated 
 caudad of tnese. All of tne Hydropnilidae and Scarabaeidae possess 
 setaceous caudo-lateral sclerites, wnicn do not seem to be present in 
 otner genera studied. 
 
 Odontoidea. -- Owing to the strong chi tinize.tion of the head and 
 the close fit ol the nead in the prothorax, there is little need for 
 a special prbcess or projection on the head for the articulation of 
 the cervepisternum. Such a process is called an odontoidea. The few 
 instances in which it does occur are in Scaphidium (Pig.J?/j , Pucine- 
 tus (Pig.-^^j, Gyphon (Pig.f^^^ , Byrrhus (Pig.^^y) , Hosodendron (Pig.y-p^)" 
 Phyconomus (Pig.yj^j) and Arthromacra (Pig.-^^) . The latter genus, it 
 is interesting to note, possesses no cervepisternum. 
 
 Postgena. -- That part of the head-capsule on the caudal surface, 
 mesad of the occipital suture and ventrad of the occiput, in general- 
 ized orthopterous insects is the postgena. The occiput is considered 
 as extending to near the middle of the dorso-ventral length of the 
 occipital foramen, thus limiting the dorsal extent of the postgenae. 
 The latter are widely separated from each other by the occipital for- 
 amen. ihe lateral parts of the large subraentum distinctly cover the 
 mesal parts of the postgenae. In the Coleoptera the occipital suture 
 separates this region from the remainder of the head-capsule, as in 
 generalized insects, and the occiput is also indis tinguishably fused 
 with the postgenae. But the postgenae, instead of being widely sep- 
 

 
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arated and their mesal parts being covered by the lateral parts of 
 the submentum are directly connected by the broad area located be- 
 tween the occipital foramen and the submentnm (Fig. / ) ; and the sub- 
 mentum is attached to the mesal portion of the cephalic margin of the 
 postgenae. This area is indistinguishably ftised with and undifferen- 
 tiated from the postgenae in the hypothetical and the more general-' 
 ized types of ventral surface. 
 
 In generalized insects the submentum is joined to the cervix by 
 the undifferentiated part of the cervix, the cervicoria. In the ceph 
 alic migration of the submentum, a part or all of the cervicoria must 
 have been drav/n in behind. The postgenae alone bordered the lateral 
 margins of the submentum and the cervicoria. A possible line of de- 
 velopment may have been the chi tinization of the cervicoria and the 
 disappearance of the sutures separating it from the postgenae. The 
 broad area between the occipital foramen and the submentum would in 
 such a case be a modification of part or all of the cervicoria. It 
 is much easier to believe, instead, that because of the membranous 
 condition of the cervicoria, no strong resistance was offered to an 
 approach of the mesal margins of the chitinous postgenae, which would 
 migrate thus to serve as a firm foundational support for the submen- 
 tum, as well as to strengthen the solidity of the head-capsule as a 
 whole. The mesal margins would in time approach so close as to press 
 the two sides of the cervicoria into a line. Fusion of the mesal mar 
 gins of the postgenae would finally take place, and the cervicoria 
 would be divided into two parts. One part would be attached to the 
 submentum entirely separeted from that portion of the cervicoria bor- 
 dering the occipital foramen, and would be finally reduced to a su- 
 ture. The other part of the cervicoria would remain as a portion of 
 
f ■ 
 
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 ;-fdir^i ant t^:\-\,ii':)( iWAset^Jtr' tr •■'cylr|J^t^ i^-^ l'ti|^ •-^M’ 
 
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 ^ " I ■ ■■ ~ ^ ' 4 ' ■ ' ! 
 
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 #iti *1^' 7 'Ji -* 4^ • j -.li .4‘ ti .. utc^dystj 
 
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37 
 
 the cervix. Whether the mesal margins of the postgenae would ap- 
 proach the meson as broad surfaces, or as narrow ones and later elon- 
 gatejit would be difficult to state with certainty, though the latter 
 would probably happen. In the primitive coleopterous head it is not 
 unlikely that the head was much shorter, and that the area between 
 the occipital foramen and the submentum was correspondingly shorter, 
 so that there would not be the necessity for a broad fusion of the 
 mesal margin of the postgenae. So far as observed there has been re- 
 tained in no coleopterous head any marks of the line of fusion of the 
 postgenae . 
 
 Abundant evidence is found for the conclusion that the broad a- 
 rea between the occipital foramen and the submentum must be a part 
 of the postgenae. A similar condition as in the Coleoptera is pre- 
 sent in many families of insects. In many Eymenoptera the mesal mar- 
 gins of the postgenae are fused between the occipital foramen and the 
 articulation of the labium. That this is the true interpretation of 
 this structure is shown by the fact that there is often a suture on 
 the meson showing the lime of fusion, and is further substantiated by 
 the fact that this area bears on its ventral margin the paracoilae. 
 
 In all cyclorrhaphous and orthorrhaphous jJiptera the area ventrad of 
 the occipital foramen is a continuous chitinized piece, similar to 
 that found in the Goleoptera, and is derived from the fusion of the 
 mesal margins of the postgenae. 
 
 It may be well to add here that the expression '’cephalic migra- 
 tion" as applied to the changed location of such a structure as the 
 submentum, may be only relatively correct in its suggestion. Very 
 probably the occipital foramen in Coleoptera has migrated dorsad, at 
 least it has assumed this typical position, ouch a migration in 
 

 
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 :> '•‘WiDT?’ '. I»A*^i-' t!*’ '!•. i'. r:-? t sC hpa^'X- ^i{i ^o^'imAj* fl ] 
 
 
 1 
 
 ■V 
 
 
 m. 
 
 noij&tkY ^».I/iRlft 
 
 
 \4 
 
 \ 
 
 * Ik<* ■“ 
 
 >i '4 l,•i^i^ 4 ^xt 7 ;>^‘‘ ' ftjtW vf H*v 
 
 J 
 
 ■ •* 6 - fj,© f ^ I' J 
 
 * , . . • ll« -W , ' I-. 
 
 .■♦j-^aM'ix <!Ji ad V "T- ■***''• - ■ "V v' ,iMnt#*ifi'f»'‘ 
 
 f-< .' '•<•>;' ‘'ur <: I .-JiV^aitt-* 'Xii aaawi^ Lr'J.iPM- * x f 
 
 1 ' '*■' ’ ' • u’’ *• 
 
 9 t aoi> •'^jWi *6 -W Tii,U> 
 
 
38 
 
 itself would produce an elongation of the ventral surface, with a con • 
 sequent production of the hroad area between the occipital foramen 
 and the submentum. Whether the force exerted on the head was direct- 
 ed more towards the cephalization of the submentum or towards the dor • 
 salization of the occipital foramen it would be hard to saj. ine con ■ 
 dition resulting would be in either case approximately the same. For 
 purposes of convenience and simplicity of language , the expression 
 "cephalic migration" is used in this discussion. 
 
 This broad area betm^een the occipital foramen and the submentum 
 is sometimes designated as the gula. Usually, though, the term gula 
 is used to indicate a sclerite, on the meson of the ventral aspect, 
 separated from the remainder of the head-capsule by distinct subpara- 
 llel sutures,, the gular sutures. The terra gula cannot satisfactorily 
 be appoied to both an indefinite region and a distinct sclerite. The 
 gula is considered in the latter sense in this discussion. For the 
 present the area between the occipital foramen and the submentum will 
 simply be designated as the raesal parts of the postgenae. 
 
 The cephalic end of a postgena is usually modified into a rather 
 flat area sloping towards the meson, upon which are situated a para- 
 coila ^nd a postcoila. This flat area is distinguished from the re- 
 mainder of the postgena by a distinct ridge, which usually extends in 
 a deep curve from near the lateral margin of a postcoila to near the 
 caudo-lateral margin of the submentum. Sometimes this ridge is di- 
 rected towards the occipital foramen, as in Leptinus (Fi.pr.Zf?) , Gan- 
 tharis (Fig.^7f) , Ifacrosiagon (Fig.W?^) , Tharops (Fig.^fz) , Psephenus 
 (Fig.^7^ , Helichus (Fig.Wf^) , and others. Attention has already been 
 called to the fact that the cephalic portion of this ridge represents 
 the cephalic end of the occipital suture. This modified part is 
 
II I ’ ii x i ii 'n i nrrrTft>t'i i i' rn p "'lTiTT^'~’ ~nr; 
 
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39 
 
 reduced in size or indistinguishably fused with the remainder of the 
 head, in Photinus (Fig./7i) , Fucinetus (Fig.3^^) , Alobates (Fig.^i^") , 
 Tenehrio (Fig.?>Z6) , Hyporphagus (Fig. 32 f) , most of the Scarahaeoidea, 
 Sphenophorus (Fig.?^?) , Fupsalis (Fig.;?<io), and Lixus (Fig.j(i<^). 
 
 Gula . -- There seems to he considerable confusion as to what con- 
 stitutes the gula, Some investigators consider it as simply the cen- 
 tral portion of the ventral surface of the head-capsule. Others re- 
 strict it to the distinct sclerite appearing on the meson of the ven- 
 tral surface of certain insects, including the Goleoptera. Since 
 this sclerite is distinctive and characteristic it needs a name. The 
 term gula is quite satisfactory to apply to it. The origin of the 
 gula also rests in considerable confusion, Comstock and Kochi (1902) 
 consider it the sternum of the cervical segment, and to strengthen 
 their argument they cite the situation in Gorydalis, in v/hich they 
 state that the sternellum of the cervical segment is retained back of 
 the gula, and they figure it as the small quadrangular sclerite imme- 
 diately caudad of the gula. They do not mention the very much larger 
 rectangular cervical sclerite caudad of their '’sternellum”. The lar- 
 ger sclerite is suggestive of the sternellum, and the smaller one 
 
 would then represent their "sternum”. It is generally stated that 
 
 the 
 
 the gula is derived from^chi tinization of the neck membrane or cervix 
 In the discussion preceding, dealing with the metatentorinae and the 
 postgenae, I haveattempted to show the impossibility of such a deri- 
 vation. In many orders of insects structures on the caudal aspect of 
 the head have been designated as the gula. My interpretation would 
 prevent the recognition of the presence of the gula in any insects 
 other than the Goleoptera, Keuroptera, Trichoptera, and possibly the 
 Mecoptera. 
 
' 'T. " . 
 
 
 i - ■ . V'^'i ’^l '"' ■ 
 
 b- iil'K^^ ^i> ‘ T.\tiX^e^,k2/!^l -in vifX/t £_ 
 
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 ■ ■ .: - - _ia . *- 
 
 is ^ '4., 
 
 
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 li ‘ _ ' ’v. « . .,;_ • , > 
 
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 ■* ^ - ' - **:• ■•> 1 hvi:^- 'tO: t: . n.-rxX^ ■ ■ 'I^itXt>9 ieM>X©0 
 
 .’/« -/•.;, A’ 
 
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 .« ■ 
 
 
 ^«. ' UAH -<WBJi~0JACNrf^?p|fY4;r..t •4i/-,f :^6tL»al ‘ •ioV’T-^ AAiJia'oaJX" cl'.j 
 
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 f ‘ ^ .’^©’^.•1 7 yXo I £|!T» 4X>d. Jl X»>?JU!KilU^ 6 tS 
 
 fc Hi. om el ^Xmi^ txtd to •#'«’:■ ^ U IfeO' Boi^lAr^oili. ©«iX /a#v©7a- 
 
 r I- . i 
 
 I 
 
 
 ^jpTo^^^ooelff I 
 
 
40 
 
 practically 
 
 In some Coleoptera there is^no gula (Figs ^ 7 ), whereas in 
 
 others there is a complete gala (i'igs,Z^/^ ). Between these ex- 
 
 treme types there are found in the Coleoptera all gradations of a gu- 
 la in process of formation (Figs. 2 W,^^<^^ ). buch a range of vari- 
 
 ation in a structure simply emphasizes the importance of a compara- 
 tive study of a large series of forms. The question as to which is 
 the more generalized condition, the presence of a complete gula or 
 the absence of one, has been discussed previously, with the verdict 
 in favor of the latter condition. It would then be impossible , other 
 reasons not being considered, for the gula to be the sternum of the 
 cervical segment. The gula is the sclerite formed by the migration 
 of the me tatentorinae from the occipital foramen towards the submen- 
 tum, and must be derived from the postgenae. In this migration of 
 the me tatentorinae sutures are produced which are knovm as the gular 
 sutures. The area between the gular sutures is the gula. It is evi- 
 dent, then, that the presence snd extent of the gula depend upon the 
 position of the me tatentorinae and the sutures which they produce. 
 
 The gula is complete in all the Adephaga except Peltodytes , 
 
 in the Silphidae, the Staphylinid&e , the Sphaeriidae, the first four 
 families of the Cantharoidea, and the ;^carabaeidae , except Pseudolu- 
 canus (Fig.jy,?) . Isolated examples in other families of a complete 
 gula are Scaphidium (Fig.^^^ , Helichus (Fig.^^'^) , Cyphon (Fig.y<?/) , Byr - 
 rhus (Fig. 30 ^) , and Leptinotarsa (Fig.3-5*-^ . The gular sutures nearly 
 meet in Gastrolobium (Fig.^^^) and Creophilus (Fig.^^ , and are entire- 
 ly confluent in Necrophorus ^ Bcaphidium , Hister . 
 
 (Fig.77i0) , Phengodes (Fig.i/.^) , Ghauliognathus (Fig.y;^) , and in all the 
 Phynchophora except Furymycter (Fig.J^/) , in which genus the gular su- 
 tures are very short and widely separated. The gula in cases of con- 
 

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 ,t * . 7 I !i t; 4 T ft i ilft. tfjs^, f r/:1 
 
 
41 
 
 fluency of the middle portion of the gular sutures on the meson is 
 simply invaginated. The gula is complete hut extremely short in Gal- 
 opteron (Fig. 27 /) and Photinus (Fig.2/z.) . The cephalic ends of the gu- 
 lar sutures meet on the meson near the occipital foramen in Philother - 
 mus (Fig. ^ 77 ) and Anoplitis (Fig.^jy/) . The gula is partially membran- 
 ous in xTosodendron (Fig.3<?f) , bitodrepa (Fig.J^z) and Plesiocis (Fig, 3 ?ii , 
 Part of it is deeply invaginated transversely in Scaphidium (Fig.^^7) , 
 Chauliognathus (Fig./c57) , Helichus (Fig,^^^) , btenelmis (Fig.y^?’^ , and 
 others . 
 
 Api cranium. -- Ivlost of the sclerites of the head-capsule are so 
 closely united in some Coleoptera as to appear as a single piece. 
 Straus -Pur ckheim (1828) so regarded this region in the head-capsule 
 of the I.iay-heetle , and named it the epicranium. Used in the sense of 
 indicating a closely united area, the term seems satisfactory and may 
 often prove convenient. It can then he only a relative term, includ- 
 ing at times no more than the paired sclerites of the head, vvhen the 
 epicranial suture is well developed. Occasionally, the preclypeus 
 and the lahrum are closely joined to the head-capsule. In such cases 
 the epicranium would embrace these structures, also. 
 
 Paracoila . -- The small rather concave projections at the caudo- 
 mesal margins of the postgenae against which the maxillae articulate 
 are the paracoilae. They are usually easily identified, as in Molam- 
 ha (Pig. 26/) , Pytho (Pig.2^<i) , Heterocerus ( Fi g , and. Phymaphora I 
 (Pig.^/f') . They are usually chitinized, hut are membranous in a con- 
 siderable number of genera. The paracoilae are insignif icant in size 
 in G-astrolobium (Fig, 2^^) , all of the Hlateroidea, Jistela (Fig. 32/) , 
 and Tenebrio (Fig. 32^) . In generalized insects the paracoilae are sit- 
 uated beneath the submentum. They are found in a similar position in 
 

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42 
 
 tiie Adephaga (Figs , in Hydrous (Fig.^5i^) , and a few others. 
 
 They are also always located on the postgenae in generalized insects. 
 This position is of great importance in determining the identity of 
 the postgenae in specialized insects. 
 
 Postcoila . -- At the cephalic end of a postgena is always found a 
 distinct crescent-shaped acetabulum-- the postcoila, against which 
 the postartis of the mandible articulates. Throughout the genera the 
 postcoilae exhibit some degree of difference in exact position upon 
 the postgenae, in size, and in degree of shallowness. Their form and 
 position are indicated in Galosoma (Fig.i,?i?) , bphaerius (Fisr.^^g’) , IJo- 
 toxus (Fig.7^^) , and Derobrachus (Fig. 3 ^^) . In generalized insects the 
 postcoila are also always located on the postgenae, and their posi- 
 tion is of equal importance with that of the paracoilae in determin- 
 ing the identity of the postgenae. 
 
 Precoila. -- The rounded condyle on the ventral surface of each 
 clypealia, against which the preartis of the mandible articulates is 
 the precoila. It is usually crescent-shaped, but sometimes, as in 
 Peltodytes (Fig.i^.^) , Cybister (Fig.i5f), Greophilus (Pig.^^ 5 ^ , Endomy- 
 chus (Fig.J^c) , Dichelonyx (Fig.jjr) , and others, the form is spher- 
 ical. The precoilae are large in Macrasiagon (Fig.^^'y) , Epicauta (Fig 
 ) , Tharops (Fig.^fz) , Ghauliognathus (Fig.p//) , Bostrichus (Fig. 3 ,^?), 
 and others, and small in Dineutes (FigvZ^^, Photinus (Fig.27il, Heli- 
 chus (Fig.Jf^) , Derodontus (Fig. 3 / 3 ) , and particularly so in Psephenus 
 (Flg,Jff), T he precoilae can readily be recognized by their character 
 istic form, oince they are always located on the caudo-lateral lobe 
 of the postclypeus, or the clypealia of certain insects, they are a 
 great aid in determining the limits of the postclypeu.s . 
 
 Tentorium. -- The endoskeleton of the head is known as the ten- 
 
\ 
 
 rr! 
 
 4? 
 
 ■/fl^o -. -^Re^fqsubA -©.I.** 
 
 sr-^biauiteit- 
 
 rw* 
 
 ••y-f 
 
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43 
 
 toriura. It is composed of two anterior and two posterior arms or pro 
 Sections, and their various modifications. The anterior arms arise 
 from the pretentorinae , and are knovv'n as the pretentoria. The pos- 
 terior arms arise from the me tatentorinae , and are known as the meta- 
 tentoria. In generalized insects one end of the pretentoria expand 
 along and are continuous with the epicranial suture. liach extends 
 in a caudo-mesal direction, and narrov/s for a short distance, then 
 expands along its mesal margin until a fusion is formed with the pre- 
 tentorium of the other side, producing the cephalic bridge or lami- 
 natentorium. The pretentoria separate and then fuse again farther 
 caudad with each me ta tentorium. The metatentoria extend cephalo-me- 
 sad a very short distance, their mesal margins expand and completely 
 fuse on the meson, producing the caudal bridge, or corpotentorium. 
 
 The fusion of the pretentoria and the metatentoria is supposed to 
 take place along the cephalic margin of the corpotentorium. The dor- 
 sal pro;iections arising from the lateral margins of the pretentoria 
 and extending toward and attached to the dorsal wall of the head, are 
 the super tentoria. The ring-like plate surrounding the inside peri- 
 phery of the occipital foramen is indistinguishably fused with and is 
 a part of the metatentoria. The tentoria, as a whole, are distinctly 
 chitinized and v^ell-developed. The typical condition of the tentor- 
 ium in generalized insects is practically duplicated among the Goleo^ 
 tera. The hypothetical type (I'ig.37b) has been constructed with this 
 similarity in mind. The greatest difficulty experienced was in de- 
 ciding upon the primitive type of laminatentorium, whether it should 
 be represented as complete or incomplete, that is, whether the two 
 sider of the laminatentorium fuse on the meson or not. Many Coleop- 
 tera that in other respects are quite generalized do not show a com- 
 

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44 
 
 plete laminatentorium, as Tetraclia (Pig. 37/) , Omophron (Pip:.J73l , Di- 
 neutes (Fig. ^73) , leptinus (Fig.J^/) , Tachinus (Fig.j^ 7 ) , and Stenelmis 
 (Fig.y/g') . The hjrpothetical laminatentorium is represented as nearly- 
 meeting on the meson. At least, such a condition is thought to he 
 not far removed from that which actually existed in the primitive Go-- 
 leoptera. liraulodes (Fig.3B'7) , liuryste this (Fig./^<i) , Pytho (Fig.^^7) , 
 Philothermus (Fig.^^^) , Melanophthalma (Fig.f.??) , Hyporphagus (Fig.-^x<? 
 bphindus (Fig.T'r^) , and others, possess practically no trace of a lam-. 
 inatentorium, hut well-developed supratentoria. The latter are fair., 
 ly well developed in Photinus (Fig.3?f) , Gollops (Fig..;^/) , and Alaus 
 (Fig.f/^ , hut there is neither a laminatentorium nor a corpotentor- 
 ium present. There is no trace of any one of the three above-men- 
 tioned structures in Tharops (Fig.-j^/j) and Throscus (Fig,^/-^). Phalac- 
 rus (Fig.y^^) and Furymycter (Fig.f'^'^) possess only the rudiments of 
 the pretentoria and metatentoria. The tentoria of the Geramhycoidea 
 (Pigs. ) are very delicate and membranous. The pretentoria, 
 
 the metatentoria, and the corpotentoria are always complete, while 
 the laminatentorium end supratentoria are always either rudimentary 
 or absent except in Pachymerus (Fig.-#^^/?) , in which the supretentoria 
 are present. Other genera showing completely membranous tentoria are 
 numerous. Among these are Gupes (Fig./^?/!?) , Auryste thus (Fig.^^'<5?^) , Hh. .. 
 sodes (Fig.-^^6) , Languria (Fig../^Z) , and Pseudoci stela (Fig. . 
 
 Pretentorium. -- A generalized condition of the pre tentori-um, 
 characterized by a strong chitinization as a whole, and possessing a 
 broad flaring cephalic end, is found in the Adephaga (Pigs,37'5^ ) , 
 
 Leptinus (Fig.3^/) , Hecrophorus (Fig.Ji^Z,' , the Staphylinidae (Figs.^yj' 
 
 ) , Georyssus (Fig.y^^) , liucinetus (Pig.^..2/) , Dermestes {Fig.^.?^) , 
 
 and Lerod ontus (Fig.y^^). There are all degrees n1 Present 
 
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45 
 
 from the most generalised pretentoria to those very delicate raemhran" 
 ous ones represented hy such forms as Heterocerus (Fig.^/^), Fndomy- 
 chus (Fig.yf*/) , -Pseudocistela (Fig.y^^, Hyporphagus (Fig.-^cf^yj , the ma- 
 jority of the Geramhycoidea, and the xihynchophora. A very prevalent 
 type , possessing a distinctly chitinized cephalic end and a membran- 
 ous caudal portion sharply separated from the former, is represented 
 in iilpicauta (Fig.-^'^.^T, Pytho (Fig. , Chalcophora (Fig.y/^T, Alobates 
 (Fig.^f^), Bostrichus (Fig./j'/) , Diabrotica (Fig./7Sl , Bendroctonus 
 , and most of the Dcarabaeoidea. ihidimentary pretentoria 
 are found in a number of genera, as Calopteron (Fig. 3^?) , Tenebroides 
 (Fig. ^^ 7 ), Phalacrus (Fig.^^z] , and perhaps all of the Phynchophora , 
 except Bendroctonus . The pretentoria of the Ahynchophora are for the 
 most part very delicate and fragile. The difiiculty of making a dis- 
 
 section shov/ing the pretentoria intact is intensified by the close 
 in 
 
 packing with^the snout of the greatly developed tendons of the mouth-. 
 parts and the strongly chitinized pharynx. In no instance, except 
 in Bendroctonus (Fig.-/7/) was a pretentorium preserved intact. In 
 bcolytus (Fig.-^fo) the pre tentorium is evidently rudimentary, but in 
 none of the other genera is there a clear indication of such being 
 the case, owing to the presence of frayed ends, suggesting that not 
 all of the pretentorium has been seen. The presence in the rdiyncho- 
 phora (Figs . ) of a suture in the right position for an epi- 
 
 cranial suture, an invagination within the suture that suggests the 
 pre tentorina , and a projection arising from the invagination, alT: 
 furnish evidence that this projection is probably the cephalic por- 
 tion of the pre tentorium. The pretentorium of Passalus is extraordi- 
 narily developed. The cephalic part extends as a slender bar dorso- 
 caudad, nearly half way to the occipital foramen, then bends sudden- 
 

 
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46 
 
 ly and extends as a huge straight arm tov;ard the corpo tentorium. The 
 tremendous development of the caudal part is due to the need of a 
 strong support for the dorsal surface, which hears a prominent horn 
 used in fighting. 
 
 Me ta tentorium. -- The form and size of the metatentorium is quite 
 decidedly indicated by the position of the metatentorinae . Those 
 genera possessing primitive me tatentorinae are very apt to possess 
 the primitive type of metatentorium, one that is short and simple, 
 as Limulodes (?ig.;jj'^), Sphaerius (Fig.jf^i^) , Anchicera (Fig.f^.?;>1 , PhilO" 
 thermus (Pig.-j^.?^') , Melanophthalma (Pig.yy^) , and Sphindus (Pig. . 
 
 It is significant to note that all of these genera are very small in 
 size. The cephalic migration of the metatentorinae is due to a sim- 
 ilar movement of the metatentoria. In those genera in which the met., 
 atentorinae have migrated from their primitive position near the oc- 
 cipital foramen, the metatentoria are found more or less deeply in- 
 vaginated along the gular sutures, the sutures being the products of 
 these invaginations. In most genera the metatentoria advance but 
 little or not at all farther cephalad than the metatentorinae , but 
 there are some exceptions, in which the metatentoria taper down grad- 
 ually, as in Helichus (Pig.^/7), Gyphon (Pig./^i) , Lyctus (Pig.-jJ^.^*4 , anc 
 all of the Scarabaeoidea, except Pseudolucanus (Pig.y/^ . In those 
 genera in which the gular sutures are confluent on the meson, the a:u-. 
 la itself is simply invaginated, becoming a part of the metatentor- 
 ium. The same type of development has taken place in Necrophorus 
 (Pig,3i*-j) , bcaphidium (Pig.^?/) , Hister (Pig.^^^.cJ , Phengodes (Pig.jf,}^ , 
 Chauliognathus (Pig.39’t) , and Georyssus (Pig.-f^^?) , as in the Hhyncho- 
 
 phora. In ilecrophoinis , Phengodes, Chauliognathus , Aupsalis (Pig./^-^) , 
 and Thecesternus (Pig.^J?/) , the line of fusion of the invaginations 
 
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47 
 
 of the two sides has disappeared. In Ghauliognathus the invagination 
 is greatly reduced, and in Phengodes nothing remains hut a mere line , 
 These two latter genera seem to show a greater specialization of the 
 gular region than any other genera studied. A correspondingly deep- 
 er invagination of the ring-like plate surrounding the inside peri- 
 phery of the occipital foramen has occurred with that of the gula. 
 
 The whole phenomenon appears to he due to an especially strong ceph- 
 alic pull on the raetatentoria. This can he readily understood in 
 the case of the Ahynchophora , in wnich the elongation of the snout 
 would encourage this result. A second force may play a part here, 
 that of the narrowing of the snout, which might assist in the enfold- 
 ment of the gula. 
 
 A large number of genera, scattered throughout the series of 
 families, possess prominent projections along the mesal margins of 
 the metatentoria , caudad of the corpotentorium, as those of Cyhister 
 (Fig.J/7) , Necrohia (Pig.J?’^) , Grlischrochilus (Fig.f^f) , Phyconomus 
 (Fig.y'y^) , and Boros (Fig.y^,^ . Some of these projections have dis- 
 tinct tendons attached to them, as in Scaphidium (Fig.Jv) , Parandra 
 (Fi g,y^ 7 ) , Derohrachus (Fig.7-^J^ , Bonacia (Fig.^;7) , and Criocerus (Fi?, 
 7/^). Prominent lateral projections are present in Lyctus (Fig.7.r;5)- , 
 Osmoderma (Fig.y^i) , Parandra, Glycohius (Fig.-j^d?) , Ahynchites (Fig. 
 
 , Epicaerus (Fig.^^4) , and Thecesternus (Fig.^^*;^ . Passalus (Fig. 
 is peculiar in the possession of a large well-chi tinized second- 
 ary bridge arising from the mesal margin of the metatentoria. 
 
 Corpotentorium. -- Owing to the simplicity of form of a struc- 
 ture like the corpotentorium, hut little change is indicated in it 
 through most of the genera. !£'he hypothetical type shows this struc- 
 ture to he a rather narrow simple hand (Fig.J7^) . Such is fairly 
 

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48 
 
 characteristic of the vast majority of forms. The corpotentorium is 
 sometimes very hroad, as in Ptinus (Fig.y^^) , Bostrichus (Pig./^^) , 
 Parandra (Fig.^/) , Beraorachus (Fig./<^^1 , Glycohius (Fig.^<^f) , and Eup- 
 salis (Fig,//^). In contrast are many that are quite slender and 
 arched, as Bineutes (Fig.J/^) , Btenus (Fig.^irf), Hister (Fig.3?>2}, Beo- 
 ryssus (Fig.-^^^) , and Mycetophagus (Fig.^^^/) . The form of these lattei- 
 has evidently resulted from the narrowing of the space between the 
 metatentoria. An exceptionally large number of genera have only a 
 rudimentary corpotentorium, or none at all, as in all of the Lampy- 
 roidea except Trichides (Fig.;??^^ and Becrobia (Fig.^^f) , Epicauta (Fig. 
 
 , Macrosiagon (Fig,/^f) , all of the nlateroidea, Psephenus (Fig.-//^! 
 idiysodes (Fig.y^<^) , Phalacrus y Hippodamia (Fig.-^/^) , and Ar- 
 
 thromacra . The reasons for the loss of the corpotentorium 
 
 are not always evident, though in most cases , either the arms of th« 
 tentoria have expanded and approximately met on the meson, as in Hhy-. 
 sodes and Plesiocis (Fig.-ji^oy) , or the arras are directed towards the 
 meson and meet there, as in Gonnophron (Fig.j^,?) and all of the Elat- 
 eroidea, or the pharynx rests snugly between the tentorial arms, as 
 is so perfectly found in Ghauliognathus , where the pharynx is wedged 
 so tightly between them that the whole seem like one piece, all of 
 which conditions supply firmness to the tentorial arras and obviate 
 the necessity for a corpotentorium. Of the hhynchophora , Euryraycter 
 (Fig.-^^^) , Epicaerus (Fig.f/^), and Lixus (Fig./^^ possess no corpoten-. 
 torium. Scolytus (Fig.y^<jj) possesses a very rudimentary one. The 
 disappearance of the corpotentorium can best be explained in the caso 
 of the Hhynchophora by the fusion of the metatentoria into one solid , 
 plate that needs no added support. The corpotentorium of most of th(i 
 Gerambycoidea is a delicate membranous structure. A cbmmom modifi- 
 
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49 
 
 cation of the corpotentoriura is the mesal projections on its cephal- 
 ic "border, as in lachinus (Fig. 3^7) , Cephaloon (Fig.-j?^/) , Tomoxia (Fig 
 
 Phenolia (Fig.^^^?), and Philothernms (Fig./^s) . 
 
 Laminatentorium. -- A structure of the tentorium that shows per- 
 haps a greater varia"bility in form than any other is the laminaten- 
 torium, which appears in a great array of shapes and sizes, from the 
 forming of a perfect and "broad bridge to total disappearance. .'fhen 
 the two sides of the laminatentorium meet on the meson, the mesal 
 margins of the laminatentorium very frequently are "bent ventrad, pro. 
 ducing a projection. Examples of this development are seen in Galo- 
 soma (Fig. 37^) , Gyhister (Fig.J?/) , E"ecrophorus (Fig.j^iv , Aleochara 
 (Fig.3?Sf) , Heterocerus (Fig.^^/^) , Dermestes (Fig.-^^^i, and Endomychus 
 (Fig.-^^^S') . Genera that show the laminatentorium as hardly more than 
 touching on the meson are Peltodytes (Fig.3/<^>) , Scaphidium (Fi?.3V), 
 Pyrochroa (Fig.*7^<?<^ , Ilotoxus (Fig.f'/^) , Psephenus (Fig.f/i) , Gucujus 
 (Fig.yj/), Penthe (Fig./^y), and most of the 3cara"baeidae . Ihe reduc- 
 tion of the laminatentorium takes place so gradually that it is hard 
 to tell when it has completely disappeared. There seems to "be no in 
 dication of it in Limulodes (Fig.J^^) , Gollops (Fig..?^/) , Pseudociste- 
 la (Fig.f.j?3l , Hyporphagus (Fig.y;>u) , Ephindus (Fig.y^'^) , and others. 
 
 In Tomoxia (Fig./<?^) , Pytho (Fig.y<97) , Alaus (Fig.-j*’/^ , Tharops (Fig.^/^ 
 Throscus (Fig..//f) , and others there is no laminatentorium. The pre- 
 tentorial arms curve towards the meson and serve a similar purpose. 
 The laminatentorium of Tachinus and Epics uta is in the form 
 
 of a lobe. That of Psephenus (Fig. -7/^), Pichelonyx ( Fig. -737) , and Pel 
 idnota (Fig.TV^) possesses two long sharp cephalic projections. 
 
 Supratentorium. -- Arising from the dorsal surface of each pre- 
 tentorium cephalad of the laminatentorium and usually extending 
 

 
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50 
 
 towards the dorsal surface of the head is a projection of variable 
 form, the supratentorium. Its generalized condition would show a ra- 
 ther distinctly chitinized structure, with an expanded ventral end 
 that gradually narrows, then expands flat-like against the ental sur-. 
 face of the head-wall. This kind of structure is found in a very 
 large number of genera, as all of the Adephaga except Peltodytes (Pi,-:. 
 374 ,), the Hydrophilidae , Leptinus (Fig.ja/) , all of the titaphylinoidea 
 except Aleochara (Pig.^^S^) and Eister (Fig.^j-i) , Pytho , Eoso- 
 
 dendron (Fig.^-^^) , languria (Fig.-jJ^ji) , Megalodacne {Fig.y?}) , Derobra- 
 chus (Fig.-j^^s^) , and others. The supratentorium is a surprisingly per- 
 sistent structure considering the large number of genera that posses:; 
 it in a more or less rudimentary state. The broken ends of the ten- 
 toria of Aupsalis (Fig.y^^J , Lixus {Fig.-z^^sr) , and bphenophorus 
 are expanded, which expansion may include a part of the supratentor- 
 ia. This seems reasonable to believe when a rudimentary one is evi- 
 dently present in Ahynchites (Fig.^.^^) and Attelabus (Fig.-^f^ . Sphen- 
 ophorus is peculiar in having the lateral margins of the tentoria 
 fused to the oculata. The head-capsule of this genus is suddenly 
 constricted at this point, which probably placed the tentorium and 
 head-wall in contact, a fusion finally resulting. The supratentor- 
 ia of Cucujus (Fig. -^3/) and Passalus are stout structures, but ex- 
 tremely short. In the former, this condition is due to the flatness 
 of the head, in the latter, to the unusually close proximity of a 
 part of the pretentoria to the dorsal surface. 
 
 7. SOLIS PHYLOOSKSTIC COIJSIDEEATIOES 
 Y/ith the hypothetical type of coleopterous head in mind, it is 
 possible to note the lines of development that have taken place with' 
 in the various groups. Structures have developed away from the prim- 
 

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51 
 
 itive condition with varying degrees of rapidity, and sometimes in 
 different directions. It is difficult from this study to decide on 
 the amount of importance to attach to the change that may take place 
 in a particular structure, and to average the degree of primitive- 
 ness as a whole of the development of the structures characteristic 
 of a group, and to indicate with precision the place in the primi- 
 tive scale of each group. As previously mentioned, owing to the 
 wide field covered in this investigation it was impossible to find 
 time to study a large representation of genera within each family. 
 Such a study should help one considerably in reaching clearer con- 
 clusions as to the arrangement of the genera in the primitive scale. 
 Using this study as a basis, it is quite possible, however, to dis- 
 cuss the relative importance of the structural changes exhibited by 
 the different groups, and to suggest possible inconsistencies in the 
 present arrangement of certain genera. 
 
 Adephaga.- The characteristic possession of the occipital suture 
 of complete epicranial arms, of a generalized position of the anten-^ 
 nariae and of the pretentorinae , of a generalized form of labrurr and 
 of tentorium, would force such families as the Cicindelidae , Gar-bi- 
 dae , Amphizoidae and Oraophronidae into a primitive place in the phy- 
 logenetic scale, in spite of their specialized ventral surface. Uo 
 other large group shows as many primitive characters. The genera of 
 the above-mentioned families show a great similarity in all their 
 structures and must be closely related. Of these families Omophron 
 possibly possesses the most generalized dorsal surface, though the 
 occipital suture in this genus is not nearly complete, and does not 
 show on this surface. Omophron can hardljr hold an intermediate po~ 
 
 sition between the Garabidae and the Haliulidae. for the latter 
 

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 4 JLet^Ep , to -•■aoDEXiflETTi^ taE 8 aT;is. 
 
 43 " 
 
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 "!l, 
 
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 rS 1 ^ ~ I ' *T ' M ^ 
 
 l^pc . 4 S»»t,TU‘'J ,T^ tp e^<iE^nJt ,®i ^3 0 i^'aria^px| i 
 
 ft^• ecK' ea Evrorit qaat^^^TaX ’xarf^cj'.^ 
 
 rx^ aX 'vcti i/' f jjBia * i n^df , aiiiXbiHt t^tsolf a^em-aroda 1 
 
 tOYr 'ioa^a seXX tm>'l fet^<s^TO cc« 
 
 * A r y n 
 
 tidt ff*>a 6 dt AfitofT i^£fJ '^‘fsaas^Jop 
 
 fZ'ii ‘ *off J>n» , op^i'i<ii j'Qn e f a tii^ aA s>**.M^.tt ,Catlqioo 
 
 -O'’ «£Kt#16'‘*«Tfr.*,| { tx^p ''(KfziSxfomO *T '^oct'j*"r *rr ‘ cjo wosfaFj 
 
 I V.’S ■ '«S 
 
52 
 
 latter family, as represented by Peltodytes, shows little similarity 
 with any other family of the Adephaga. The form of the head is dis- 
 tinctly specialized, being globular, with enormous eyes, and the 
 mouth-parts and the very small lab rum are directed distinctly ven- 
 tral. The occipital suture is probably lacking, the epicranial su- 
 ture is shorter than in any other genus of the Adephaga. On the ven< 
 tral surface, the metatentorinae extend only halfway between the oc- 
 cipital foramen and the submentura, and the gular sutures extend no 
 farther cephalad, whereas in all the other genera of the Adephaga 
 the metatentorinae are close to the submentum and complete gular su- 
 tures are present, finally, the tentorium of Peltodytes is not so 
 well developed, and the supratentoria are small and do not reach the 
 ental surface of the head wall. In all other genera of the Adephaga 
 the tentorium is large and strongly chitinized in all its parts. The 
 Pytiscidae and Gyrinidae are very similar fundamentally , and differ 
 from other adephagous families in the position on the ventral aspect 
 of such structures as the antennariae and the pretentorinae , and by 
 a rudimentary or absent occipital suture. The instability of the 
 epicranial suture is well illustrated by the case of the two closely 
 related genera, Dytiscus and Cybister, the former possessing com- 
 plete, distinct arms, and a considerable part of the caudal end of 
 the stem, the latter incomplete and rather indistinct arms and no 
 part of the stem. The Pytiscidae and the Gyrinidae are undoubtedly 
 more specialized than the first four families of the Adephaga. 
 
 Hydrophiloidea. -The form and texture of the head of the Hydro - 
 philidae is quite similar to that of the Pytiscidae and Gyrinidae. 
 The heads of all these families are broad and short and strongly chi 
 latinized, ihe antennariae and pretentorina e of the Hydrophilidae 
 

 ill ,. V, ',W ^:.. ',• ^ . ■*'■'%'' • k ‘ '' 
 
 ,X.tiriori:fcl.€ ;4a \^tvraX ley^al |1 
 
 , ■ ,■ ?,•• . ,1 ■>.•/-. ;2. ■ ,q_ -J 
 
 t^iit i>s:r> ?• r^o# 
 
 ■ 
 
 :4fctco»'i-i4'.?i' ^tii- 4 »’ ■' 'i 4i..'i^t^int i--eoo wf$ ^'•.Jb.»t-1 
 
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 ^ ' ■* -1. ..' , '' '4?1 
 
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 ■ f. J'ft A. tXe i hfl.i <»^oX^o8aX ,£4p;^a et;# 
 
 xX^iaJ (<t;oei!ir o«X2tiXl;^0 •rff laas « ,«^t3 eP,P- 
 
 . xi'‘ xm'HlioftX xaaJ J'atiX eo* oj*jU f LsJtcs<ia-^9’i:0^ 
 
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 M 
 
 lnu- 3 :Oj#^ 5 II oX X' fta0XXi4< 
 
 Jt--*’ Xir»' <>1*' ri>* ' afteorf 
 
 
53 
 
 have migrated ventrad, as in the other two families. The metatentor- 
 inae, the gala, and the tentorium are almost identical in form and 
 condition, also. This superfamily is evidently very closely related 
 to the Dytiscidae and Gyrinidae, 
 
 Silphoidea. -- The genera of the Silphoidea show many inconsis- 
 tencies. The dorsal and ventral surfaces and the endo skeleton of no 
 two genera are nearly alike, llecrophorus possesses the most general- 
 ized dorsal surface, which is very like that of the first four fami- 
 lies of the Adephaga. On the other hand, this genus possesses the 
 most specialized ventral surface present in this superfamily. The 
 gular sutures are complete, and for more than half their extent meet 
 on the meson. In contrast, there are practically no gular sutures 
 in Gonnophron, a scydmaenid, and Molamha, a corylophid. iJo greater 
 extremes of this surface can be found within the limits of any other 
 superfamily. The tentorium exhibits just as great extremes in devel ■ 
 opment. That of leptinus is quite generalized; that of Necrophorus 
 well developed but specialized, owing to the invagination of the 
 gula; that of Gonnophron with no corpotentorium nor laminatentorium 
 and rudimentary supratentoria ; that of Molamba the most rudimentary 
 tentorium of any genus included within this study, No clearer illus 
 tration of the difficulties of indicating relationship can be shcAai 
 than that exhibited by the families of the Silphoidea. 
 
 Staphylinoidea, -- The various genera of the Staphylinidae show 
 considerable homogeneity, Tachinus departs the most from the charac. 
 teristic condition, in possessing a much more generalized dorsal sur- 
 face than the other members of this family. The Staphylinidae poss- 
 ess a tentorium and a ventral surface similar on the whole to that o: 
 the Adephaga and the fiydrophiloidea . Tachinus, through the added 
 

 
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 i; tne ftot ai :N66tf^'ei* »at.,.»e'ij 
 
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 ,, .oal* .BOWiir-OO t 
 t'B.R 4^^ off 
 
 Sf ^ ^ : /:sj.X*SSf3il - ., ■' ■H- y . » 
 
 ii-Bfs7i5B# oeti^ 7U.A% art? .- ^»:-c. i ^j8 
 
 ©r It <6^ 1 1 as £4 S'loJi ^n± .aelqae;^ 
 
 ■ * * i . - 3, ♦ . - » ^ " , ' ' . 
 
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 4>70'6-nl '^2,trV 
 
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 „ Uif? -io a.tX' , 6Ma.XWi'ee-!l9 ‘Jorf fristfs-ireS Xia* 
 
 
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 fsi©i n "U'^^.'rc> oi ^’JrKv . %*' ^0 aiai^cjffiat 
 
 iTf’ tiitfi • e ^ aitt , i,i^ ' ii-o i * ixr/© li>ii.X 'tOg, «i«;l f ■ 10 - pol^slt 
 
 , '■ .S? ; ^ ' ^ ' ’ T 
 
 
 I, »v ^^B^i <“• V fJti I 
 
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 0'(> &*€"•’ \M9Reii0mid 
 
 a ^ 
 
 k^ifs X^*rl;'ft I BTOB iipjCT- ikni^«^i&jp80<^ Hi 
 
 [-13170 ,v.I^.;:^ ei/lJ U- i-.fctJtM'm-iprtfO aadi’ ©oai 
 
 U. 'X« ax ffailjr t."’ y/«»J£t‘l'» fc>^«r ■»' 9 'Ca' 0 >a»''‘« * 4 * Wi’OJnojf; e- 98 » 
 
 
 
 f*<JP 
 
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 '■-:J ras «* .fqi^AA 9^ 
 
54 
 
 similarity of its dorsal surface, would seem to bear a particularly 
 close relationship to these groups. The Sphaeriidae, as represented 
 by Sphaerius, and the Ptilidae, as represented by Limulodes, possess 
 a primitive condition of the metatentorinae , while Scaphidium and 
 Hister possess confluent gular sutures. The Pselaphidae, as repre- 
 sented by Pilopius, in form of head and character of the ventral sur 
 face and tentorium shows a close relationship to Gonnophron. The 
 irregularities of structure displayed by the families of this group 
 are nearly as great as those displayed by the families of the Sil- 
 phoidea. Both of these superfamilies probably need considerable re- 
 vising. 
 
 Gantharoidea . -- All of the genera of the Gantharoidea possess 
 complete epicranial arms and dorsal position of the antennariae , and 
 lack a corpotentorium, except the Gleridae and the Corynetidae, rep- 
 resented by Trichodes and llecrobia respectively. All possess a com- 
 plete gula except the latter genera and Gollops, the representative 
 of the Melyridae. This superfamily can be divided into three sub- ^ 
 groups. One group will include the Lycidae and Lampyridae, charac- 
 terized by a globular head, short, broad gula, and absence of the pre 
 tentorinae . A second group will include the Phengodidae and the Gan 
 tharidae, characterized by a longer and flatter head and longer gula 
 The third group will include the Gleridae and the Gorynetidae, poss- 
 essing short incomplete arms, incomplete gular sutures, and a well— 
 developed corpotentorium. Gollops does not seem to fit in very well 
 anywhere, though this genus, through the form of its dorsal surface 
 and tentorium may possibly lean towards the second group. Gantharis 
 of the second group, is peculiar in possessing a normal gula, where- 
 
 as in the other forms the ^ula is rudimentary, though in all other 
 

 •'f' 
 
 
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 ,i\s;f n 6 ^tCf 4 '^, 
 
 
 ■'.'i . 
 
 
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 * Wii;' 3itx;V 9 
 
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 aP . 
 
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 Xi'> a 1 tit oj Liftaft Xoh at»a& aiioXXoC \^ia£rXxc>itii©;ro^t5P, AftqoXaTaa 
 
 1 
 
 Xj|fcXoii '|i4 i £C /uKOt t(Sf* <^1aJpo.^:4’ , «X<C^-..:/l3yc4l .aieri^pca 
 
 
 
 p.. ofif/ , roj^^ls'/a'icn ts uX 
 
 
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55 
 
 respects this Renus is like the other memhers of this group. The 
 first and second groups are no doubt closely related. The latter is 
 probably the more generalized, possessing a dorsal surface resemblin 
 to a marked extent the dorsal surface of the Carabidae. The gula, 
 in so far as it is complete, also resembles the condition of this 
 structure in the families of the Adephaga, and particularly in IJecro 
 phorus, of the Silphoidea. It has, however, developed much farther, 
 even, than in Necrophorus, being not only invaginated, but reduced 
 to a more or less rudimentary state. The first group probably e- 
 volved from the second by the change in the shape of the head, due 
 to the enormous development of the compound eyes and the loss of 
 much of the gular region, which has probably fused with the cervix. 
 Parallel with these changes, the pretentoria shifted their position 
 so that they could better support the globular head, and came to as- 
 sume the direction ordinarily assumed by the supratentoria of other 
 genera. The third group would appear to be more closely related to 
 the Mordelloidea than to this superfamily. The condition of the epi 
 cranial suture, the position of the antennariae, pretentorinae , and 
 metatentorinae , the degree of development of the gula and tentorium, 
 would all favor this assumption, 
 
 Cupesoidea. -- The representative of the Gupesidae studied does 
 not show any particular peculiarities of structure of the head that 
 would entitle it to be placed in a separate superfamily. It can be 
 very readily included with the Ifordelloidea, for approximately the 
 same reasons as the Cleridae and Corynetidae, 
 
 Mordelloidea. -- The genera of the Mordelloidea show considerabl 
 homogeneity. Most of them possess heads that are elongate, with a 
 
 generalized dorsal surface and r. largft postclypeus and labrum, both 
 
35 
 
 
 
 ‘ -■ ■-•••■ .. T- . ®. , " ' '^ ■•■ 
 
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 Li tb&M «d!t %|'(^/&X'E'*i -;iWaX3* ^dxr43X: oij (^qsfd^^^o.i^a :. i:a^&^-in 
 
 I. . ■ - -.'1 ■; ,- ■ 
 
 ■ jyifc/aMktte-i rr?i«'i^;;« f#aioi» o ’^rri^as&GQq ,Jb^sifa'£4cr^ 
 
 bjfT . r^el/Jd^^iaO ei?/ iO iUj«*ioA adi: « ot 
 
 6:0 
 
 airffl' Tia LGi^fuTvO^y oPla , sJi xa#; o« ni 
 
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 / 
 
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 at^^loXXoib^oil %T5 -a aoep 
 
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56 
 
 on the same general level with the rest of the dorsal surface. The 
 epicranial suture, however, shows considerable instability, sometime! 
 showing complete arms and stem, as in Epicauta, while in others the 
 epicranial suture may be extremely reduced, as in Macrosiagon, If 
 these two genera are at all related, the condition of the epicranial 
 suture can hardly throw any light on the degree of relationship. On 
 the other hand, the metatentorinae are stable in position, and are 
 located almost uniformly about half-way between the occipital fora- 
 men and the submentum. Another characteristic feature is the fact i 
 that most of the genera possess heads that are distinctly constrict- 
 ed at their caudal ends to form a neck. The Oederaeridae, the Gepha- 
 loidae, the Pyrochroidae , the Pedilidae, the Anthicidae,, and possib 
 ly the Gupesidae and Lleloidae, seem particularly closely related, 
 through the possession of similarly shaped heads, whose caudal ends 
 are distinctly constricted, of a generalized dorsal surface, of com- 
 pound eyes that are usually more or less emarginated, of a ventral 
 surface whose structures are practically in a similar condition, and 
 of a tentorium that is alike throughout, with the exception of the 
 ileloidae, in which the corpo tentorium is absent. Though the dorsal 
 surface of the Ivlordelloidea resembles to a considerable extent that 
 of such families as the Garabidae and Gantharidae , yet this surface 
 differs so widely in closely related groups that we can hardly place 
 too much importance on the resemblance here. 7/hat seems to be far 
 more important is the totally different condition of the ventral sur 
 faces in the Mordelloidea and the Garabidae, for instance. In the 
 former the gular sutures and the metatentorinae extend uniformly 
 half way between the occipital foramen and the submentum; in the Gar 
 abidae the gular sutures extend the whole distance between these 
 
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 ( k 4 » 9 V/ aquoi^ ftftJ-ai®! mi 09 aadlt/^ 
 
 PJ! 
 
 trt r.rf odt .»704 eoiJ 4 X<fWi-Pi ®'4 t-rii 4 c 604 X 6 iToq[fri rfooaj^/doi 
 
 r'^j- ? ' ' .■ ’.,» H i 
 
 Wi- arli ±6 :ioXttbadt 6rfi tX cioah * 
 
 f , ,» ’ i 
 
 '.-•Ki , • .nOi;»*^/:.: , 6 m.r)i(fat 40 toXX^^V^ ioi aaoB^I^ 
 
 
 J5n«»iX» «t»xi;iafl "iB-lATA ,xacrt0' 
 
 kz.iQ ^(U til injriiRpfiiditJ xfiX l^J-^qXvuo 4:?i, u^eteia^r 
 
57 
 
 parts, and the metatentorinae nearly to the suhmentura. The Mordell- 
 oidea and such groups as the Adephaga and Hydrophiloidea do not ap- 
 pear to be nearly related, 
 
 Elateroidea. -- The members of this superfamily are homogeneous 
 with the exception of the Buprestidae. The pre tentorinae and the an > 
 tennariae are located on the dorsal surface, and the epicranial arms 
 extend no farther caudad than the pre tentorinae . On the ventral as- 
 pect, the metatentorinae extend but little cephalad of the occipital 
 foramen, and the gular sutures no farther. The corpo tentorium is 
 lacking in all. In the Buprestidae, on the other hand, the preten- 
 torinae and antennariae have migrated distinctly ventrad, and are nc: 
 visible at all from the dorsal surface. The gular sutures extend 
 the whole distance between the occipital foramen and the submentum, 
 and there is a distinct corpotentorium. The Buprestidae are hardly 
 closely related to the rest of the Elateroidea. 
 
 Dryopoidea. -- With the exception of the Oeoryssidae, the member: 
 of the Dryopoidea appear to be fairly homogeneous. The chief charac- 
 teristics are very prominent widely separated metatentorinae, locat- 
 ed very near the occipital foramen, and a very broad submentum. The 
 ventral surface of the Creoryssidae , as represented by G-eoryssus, is 
 totally different. In this family the gular sutures extend nearly 
 to the submentum, and are confluent through practically their entire 
 extent. The metatentorinae cannot be definitely located. Georyssus 
 would appear to be related to Hister, both through the condition of 
 the gula, of the tentorium, and of the dorsal surface. The general I 
 appearance of the ventral surface in the Dryopoidea is much like tha’i 
 of the Elateroidea, and in other characteristics they are not unlike i 
 These two superfamilies appear to be related. : 
 

 
 ■ ' , ■ » 
 
 
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 m ■ " ’'■■ - -- ■ * 
 
 Wfj CstS't^'4,,4ii;^ 'aO . . ^ at% £%oiAe>f ix^tii' iiBlsra^ on hiwSx9 
 
 
 
 ?»,rfs^c pX0AQ0,o>»£^|’i.^ ; 'a'-iftf 
 
 ntaJ: 0 0/r ttsifcxrJtira ' 0/*> hojo^ .iiwD^’xolt 
 
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 ,ntiSi\OBidii^ tao-id xiiii a Lne^jABrnB^Q^ W aBI. «e£j ^10v t« 
 
 „.. ® £ *•■ . , '■' * ' ■- 
 »Ji *^.*^rr }«\;%ooC JiBUit^ oniBi BA , 0»i ft 1*0^1 1* «{> erfl 'Vo ftOAlii/8, Xa;^1c©v 
 
 /)xi3ljr8 i0iir>iA ,1*4.0 '? %XXa1o1 
 
 * " ■ 'i *- « w ^ I . ' AJ 
 
 * * A < ^S 
 
 (■©li’iis Tl5ftl jt/X‘>ollo.j^iq i‘aQ.uAaOQ ons bp.n ,$it)a^daB Bsit ol 
 
 s*j&0T^o<>?» x^.9£xrtlsh rtf 5ocuxeo ©BOlT(ola0l-l&«^ ,laotx0 
 
 .,, I S) j4 - I ' 
 
 ^ nollilififii' Pds n^acjriJ’ tf 5 ,ocT ApSnlBt fid ci fiXxipw, 
 
 «i#M30u 4f#t!T .Bod^i^xso-Xa(S;i(JS •^tiS lo ,aXtf3 ©tfl 
 
 ni)^ Hi #iiV T»8#i7?:ff4i i8il4»>?V 0tfl lo '®bflaiA0O4Bl 
 
 
 4^iXmr tfco oto «olr tiXTolca*4#^id larflo r.l ,i 'r^loi*' taX^ 0tfl toi 
 
 V. • 'i® •( 
 
 .h*3»!in drf l>t t»*fil»^ ' a.uH '■'•i' *«•' 
 
 
58 
 
 Dascilloidea , The two representatives of two of the families 
 of the Dascilloidea studied are similar in structure. There are no 
 outstanding features that would warrant placing them in a separate 
 superfamily. They are negative enough in their characteristics to 
 fit in, perhaps, in a number of other superfamilies, such as the Mor-. 
 delloidea, Cucujoidea, and Tenebrionoidea . 
 
 Byrrhoidea. -- The representatives of the Dermestidae and the 
 Byrrhidae studied, Dermestes and Byrrhus, do not seem to differ de- 
 cidedly from one another, but the ITosodendronidae , represented by No- 
 sodendron, may fit in better in some other group, such as the Dryo- 
 poidea. They resemble the members of this latter group in the posi- 
 tion of the metatentorinae and in tne form of the submentum. In Do- 
 sodendron, the me tatentorinae are almost adjacent to the occipital 
 foramen, with a membranous area included between them. That part of 
 the metatentorium surrounding the periphery of the occipital foramen 
 projects deep into the head. The supratentoria are well developed. 
 Hone of these characteristics are possessed by the other two genera i 
 studied, 
 
 i^hysodoidea. -- The representative of this family studied, Hhy- 
 sodes, shows distinct peculiarities in the structure of its head, 
 such as an extremely thick chitinous head-wall, a fibrous condition 
 of the cervix, an arrow-shaped head, enormously developed submentum, 
 and very small mouth-parts. Its relationships are not at all clear. 
 
 Gucu.joidea. -- The members of the Gucujoidea show on the whole a ; 
 short broad head-capsule, a distinct ventral migration of the anten- 
 nariae and pre tentorinae , the degeneration of the epicranial arras, | 
 and a slight migration of the metatentorinae from the occipital for- l 
 amen. There is some variation in the dorsal aspect. The genera ’ 
 
X 
 
 
 j^iatbna^ 
 
 :^=CK 
 
 
 iCTl 
 
 •$lS, y^ 
 
 T , - 4 :»fei ^a c asM Q i^ 
 
 ■ 
 
 *«i? owt fo owt ftd^- > c 6 UX loflBCt 
 
 
 3 S. 
 
 ' ' V - ^'^-} '^ '' '^ - .' Jir: 
 
 ? On 0 ‘ta^#^- 9 flf 5 r ..^Mj/Oinr>a sfl ex^ ij^viXa'ja ‘ft-^liioXXIoeaCC oiii lo 
 
 I! 
 
 -xol. fiinX liOiHk' , 56 ilia^t'T 9 q;x/s xeflJ'Q U> iecfe»x>"/» Ai 
 
 ■ ■ i , ^ .3?: ' J 
 
 1 ^- , ^ .«j 6 f»ioaoXxtfoAat f>aa ^oiJio^'i)wrf>x /0 ,^fl«i&ioiIe‘ 
 
 ,, ii r ^ 
 
 Adit- *tAa o 4 T ??drld^t 5 tai>fta 3 .^si ^kt^" r-^ ,nahicftx^£ ' 
 
 i 
 
 Q.J-axtfla!ii e iat TirjisaXq .'aaxtaw XXiJow ^ailt 
 
 0 ^ .’I &}^jtor^0 s 
 
 •«a «, 1 MJ \^’if^ .\,XX£ 3 <txaqa-e 
 ti - : ^ \ ff- ® 
 
 
 rJ*i 
 
 w 
 
 )-06 xo'i'iXfr o t to*>ed SToo ofr fen^' e'^ViVVioCr ,folAcrj» 
 
 lO^ JSotcpee^qox i'ecl’ , © 2 ^‘t -iad Oao "^Xiefc^o 
 
 odt - 2 i? if})«« ^qiio-iy Otti'.B 4 i' x&i^ocf ct ^ac ,itoxBAef?oa 
 
 
 ax Ov'oxjf •reiisX pXrtjt- to stotfroo® ecLt oXi^pesotr '♦«aAl’o 
 
 ■ >s ■ ' • ■ • ■ ■ i ' ' ‘ ■ ’ " ' -■ • 
 
 Vi- 
 
 -ct! t\% .iw^ta^-AcfiTB aiiit tVinxo*'. m" 1? ai An»t oRaixod'tt-a-J'B^ftm ©aJ- to too it 
 
 ’TdLU<iiAP 6 ‘Vril <yf tooo-'£M taqmlB p-ia o^irlx-ot'aad'B^aai 
 
 
 jto t'ri*<^ ‘ ;piU7 ,at»5nt a«®wlorf rei)frryrr| ^exa‘'aao£iBXtJajft:’ a atlw r.^®«:*xo^ 
 
 «• 
 
 «5®itS0l: ,X:f.t/r^ivoo io 3alAajjOtXij«Vi?Jltdtn»tatj>©-ddt 
 
 *• ■*• -5 .'-•■* ‘ i y 
 
 . pa('oX«» 7 ftf , IX^# ftT*tria'i’'sC'';/ar/t.axqi/o td* or.t oXctjI qoafc fe^oatoxe 
 
 ‘-f ii^ ^ • _i /,* 
 
 f !« 7 ©aox, '^.T '■^Irl-lo oai vd fear’s ^koq »tJ» ■foilaXtatoaxaio arsiafij 50 0 ac 3 
 
 “ > '** t-j''‘ 
 
 1 ' ' :S ^ * ■'*■' '’" '■ ' ' 
 
 „-j " ' ■ ^ 1 
 
 “iSi- i v.XX«^t alsXt la evitetaaeetq^x o ioto»j |i( j^^ 
 
 s<^ t tt »*str^rrTt« etf;; ci i^iai 'jroa^ ^oallaJ antoAe ,«aX'oa 
 
 ,t»eXiinf» 
 
 FiaraXii^oou. ««ox-^F‘lt , iXrvi'-i>a«fil eiioa ti Iffo ;ie cm' as iJoi:mI'‘j 
 
 ,^'”lncr:r‘0tt^ <f-qpl 3 Y«B» ^XStto®‘Tt>a^ .jbr^ari fcaq.»a*i-wciT 7 a m' .xilvteo; tAt to 
 
 rat»> ir© ta i t v-*^ »qi*,aaoirtX&‘i- ai:I XJLii^ ^xev A'ha ^.i 
 
 .Tfl 
 
 eXo 'w " <t pO to'J^ Eof'oojtfO la 9 io<Jtoa® BttX » ~ » a e 1 c- '. 
 
 -n ,T 
 
 «. ^ • • V ^ 
 
 fc«t ,xo 40 i- iAx 3 :ofr 7 toitZlRiX M , AXixaqaa-BPOjl oaoid Ixortfe 
 
 iDlA/itUrj/j^^ ju(l :o^ac^ii©^£rf^8/»fc o41 ,9aalxotx»al»xq ^itr aaii 
 
 ! 
 
 aJ^ 
 
 -lOt iftflqJioDC' ct©*t 1 ^aaJt'o^ © qJ^ ta ao'Xl'vx^Xfir j-^lXa’a tn 
 
59 
 
 belonging to the Llyoetopha^idae , the Myce taeidae , the Melanophthal- 
 midf^e, the Endomychidae , and perhaps those of a few other families, 
 possess dorsal surfaces that strongly resemble the dorsal surface of 
 the Mordelloidea. In Derodontus and Philothermus , a colydiid, the 
 antennariae and the pre tentorinae are both on the dorsal aspect, lo- 
 cated considerably caudad. The ventral aspect of all the members of 
 the group, however, is strikingly similar. The tentorium is fairly 
 uniform throughout the group except in Tenebroides and Phalacrus, in 
 which the pretentoriura has distinctly degenerated, due probably to 
 the heavy chi tinization of the head wall. In the shape of the head 
 and the condition of the metatentorinae and the gular sutures, the 
 Gucujoidea show a similarity to the Mordelloidea, and to the super- 
 family Tenebrionoidea. 
 
 Tenebrionoidea. — There are no particular characteristics of th 
 head-capsule that would separate this superfamily from the typical 
 Mordelloidea. For instance, the dorsal and ventral surfaces and the 
 endoskeleton of Arthromacra, Pseudocistela , and Penthe , are almost 
 exactly like these parts of such genera of the Mordelloidea as Epi- 
 cauta, Gephaloon, and Macratria. The corpotentorium ot Arthromacra 
 and Epicauta is, however, lacking. 
 
 Bostrichoidea. --The structure ol the various parts of the head- 
 capsule of the representatives of the Bostrichoidea varies consider- 
 ably. The epicranial arms are complete in Bostrichus, Lyctus, Plesi 
 ocis, and Sphindus. With the exception of Sphindus these families 
 show a distinct ventral migration of the antennariae ana pretentor- 
 inae. In contrast, Ptinus and Sitodrepa do not possess complete 
 arms, and the antennariae and pre tentorinae are located more dorsad, 
 markedly so in the former. On the ventral surface, there is consid- 
 

 . VS’ ■ » I 
 
 .|r;?' ^ ct.^ sa^cOlod I. 
 
 M'il 
 
 ,srM«B-i*"rlirt#'jik. wS.1: 
 
 ’ iC' ■ *? ' . * *■ 
 
 : 
 
 ’H'S .BosSsss Xfi-»T6ii; *4.+ 'afi<f«^apt %■'«&«' »«»*bo?-: li 
 
 , W, '" ■ - ^' . ' ■ "' ' ., -r ' ■ r 
 
 ' ■ a' ■ ~ -tjjj ,..»?- ' 
 
 ■at t.Q.'%to6- wSa- slKiTiiace ."na 
 
 i 0 l>e^4J $ 
 
 19 
 
 l:.^' 
 
 
 
 3 
 
 .tXiX4^ snT’ ,.'x^ifI;Bi9' ,qm:^ ^at 
 
 a 
 
 S'' 
 
 
 0\fi^ ^/tt/jHan ftj 
 
 ... j i”k ““ 
 
 ,i. • ' • , . - -■ ® 'W ' • ^,, ,.XM', 
 
 W# -''44?. -*y^' s&di ACSt.’^|o>:aolf^ 
 
 •^g. i*« M ■ .?!■-■ , • ■; 
 
 ■ ",*J 
 
 
 y. « w .-O^'l^irfi ,Ay©&^aiX%'£i*^iJ^ '«if ''R 
 
 ^^1.. .... MIM- ■ .*-- ■■ ■■% ,f^'; ' W" ,■>' , a r,' - 
 
 ■ ,'fia , \ 
 
 " .« ft 
 
 1 
 
 a tyte i i tf »q |#/ ■ ■ 
 
 WV, ; . ;u'”V; ^ ^ i 
 
 Ki'fil' fjx;.r^ *>G/i ij» tot ' 
 
 it ' ■' *'■ ' •vt' " ■'- ■ ^ . 'try- ' •».*■' a .3 . » - '^-' »’ '. 
 
 /cojfji4 ^C aot ; 
 
 ’ - ■'’ ' ■" . , t r ' 
 
 ^i-?. ’-'r*' *o ^'Ci4 ^ to sl'xjaq a!>jl+ i>rfxr,_^X#Oexo- >1 
 
 ■ ■ ' . . 'Vr- . ■■ v-. ..' w?i' ■»-■•• 
 
 M, 
 
 a 
 
 w 
 
 ilOJ c cq’io 0 , a0OX^q©0J?» 
 
 ^^Wfi ■ 
 
 u 
 
 ’III .»iiJ- to ftt&q: ysmi'ta'v .>^1 ';0-A. 
 
 fd?^;Pl‘<ir9vX^st£ii»?ioaftt>T: pijrag^o 
 
 
 
 1 
 
 1 
 
 ■i^!Cl’.t;^T' n->dK Vc taXco 
 
 .1 ^ ^ ^ ^ -m ■. 
 
 ^ ~i£y*’4^r\'»'iq r-4j-< •iftjbxxjAT itj^ 't 0 XvijiisiA a w<^*f 
 
 i^c-0 ot XtttjA *i*-^a#*ifi3oe al;> .gaa 
 
 .^Otiioia *iyAOO;- ofe%i^iXi'^oJft».t'lvf Nir*: t»»X,^i444S|||AJr .aai.BffliMii 
 
 i **i ; i«.*v* UAT »j!^^ fti iOa 
 
 fir 1 
 
60 
 
 erable variation in the position of the metatentorinae . The tentor- 
 ium also varies a great deal. This is probably not a very homogen- 
 eous group. Sphindus seems to be more structurally similar to 6phae 
 rius than any other genus studied. 
 
 Scarabaeoidea. -- This is a very homogeneous group, characterize 
 by a heavy chitinization of the head-wall, degeneration of the epi- 
 cranial suture, ventral migration of the antennariae and the preten- 
 torinae, and a complete gula, except in Pseudolucanus , in which the 
 gular sutures extend no more than half the distance between the oc- 
 cipital foramen and the submentum. In the development of the ven- 
 tral surface and perhaps the tentorium, the Scarabaeoidea should be 
 classed with the Adephaga and related groups. 
 
 Gerambycoidea. -- This is also a very homogeneous group, charac- 
 terized by a weah chi tinization of the head wall, a distinct dorsal 
 position of the antennariae and the pre tentorinae , and a more or lesi 
 well-developed epicranial suture. The gular sutures are generally 
 short. The tentorium is delicate and membranous throughout. 
 
 Rhyncho phora . - - The members of this group may or may not poss- 
 ess a snout. They may or may not possess confluent gular sutures. 
 7/hen they do, the condition is the same as in other Goleoptera. They 
 may or may not possess a labrum. There is no definite character of 
 the head-capsule by which the Khynchophora can be separated from 
 other Goleoptera. The affinities of this group are not at all clear 
 
 Gonclusions . -- The condition of the dorsal surface throughout 
 the groups is most variable, and can be relied upon but little to 
 furnish evidence of the degree of relationship. On the ventral sur- 
 face the condition is much more stable, and probably much more reli- 
 able in indicating affinities. The tentorium is less variable than 
 

 .'-•t '.j/J" 
 
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 ifii-: . : ' 
 
 
 
 
 
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 • i '. ' ” ^ 
 
 ' .} 
 
 i Ax»I"ia’. -sXa 
 
 ^ i ~J r ■ 
 
 lU 
 
 
 - ' vv : , ; ^'10 . 
 
 ✓ 
 
 '■i V 
 
 C iii • 
 
 
 a.u 0 •:■ 
 
 
 • 
 
 i ofl '72 s;fil ■ ■: 
 
 ?iU?0 
 
 '•: r X j 
 
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 ^■ .-> 'lx u Xi 
 
 l.-A 
 
 ^Aei»-:’>i1i-, :, y. 
 
 
 
 
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 istifWi'. . 
 
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 -. r> a |»A( 
 
 
 
 vv; ■ - ■ -..Di.-'t ••; f L' *iitn : ' ‘ 
 
 .;':■ i- f '. ■3TC7t 
 
 Pa 
 
 ■ : f'^w 
 
 C 
 
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 .r.» ! rjsmdofe 
 
 i. 
 
 
 
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 ■-,-<■ r> 
 
 
 t.m c.i,.r 5 C;r:r;, 3 <i ^ '■ 
 
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 V Ij I 
 
 ,.: '> , I 
 
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 ^ ■ ■■ f 5 i'‘i ' H 
 
 , f *air ■ mi “to ' i;aT ^-*-■ -.;:-r. j ’itf 
 
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 fltf' ■ --4 ' .'.-.■-'rai' 
 
 i I 
 
 £:V.>i .:.■T^<^..■i i JW in . T * . 
 
 'vvldm^ffi. -C^a' -Vi- ri;ri'io..l -.t 
 
 ■v,^ -X..; 'tc ■ 
 
 K' y n; V, ■'...■ 'i'- '.-’i-' ..:TO«a-.i 
 
 i-**!:-* • *’• 
 
 > .rr; ( 
 
 ; IJ D i o D efi 
 
 •;o' 
 
 " *r ^ *1 
 
 r 4 U V ^ 
 
 ,- 1 i y I ’ ' ■<■'■' i- '.^ d< i. • v '-’ ’ ® 
 
 tli 3 *^:; aaoX ■ 
 
 •rr -,t ©ioc . .. .df^k 
 
 Ji. '•> c'CCiitj ..r iir)i.i% 
 
 ■ I •• w 
 
 ■;.,:/ - 1 ^ ' T :(• 6 ai“i ;.Aii . ’^d'l 
 
 r . -j-’o-; ‘ -C 
 
 .- . r»'-. '-'•/ »; a on* , ' ^.i* . - uti> 
 
 , 4 f fliino ttnX'-Jl i .'■• ?’:'i:- 
 
 V =; i\i>l ■ 
 
 / 
 
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 lOD , 
 
 riai 
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 I 
 
61 
 
 the dorsal surface, and less stable than the ventral surface. Weigh- 
 ing the evidence presented in this study, tv;o large groups of most 
 of the families can he made. The Cupesoidea, the Mordelloidea, the 
 iilateroidea, the Dryopoidea, the Dascilloidea, the Byrrhoidea, the 
 CucuQoidea, the Tenebrionoidea , and the Bostrichoidea should probab- 
 ly be grouped together, while the characteristic families of the 
 Gantharoidea , the Staphylinoidea, and the Silphoidea should perhaps 
 be grouped with the Adephaga and Hydrophiloidea . She Scarabaeoidea 
 may also be included in this latter group. The affinities of the 
 Cerambycoidea and the Rhynchophora are too vague to include either 
 in the two larger groups. 
 
 VI. SraVARY 
 
 1. This investigation deals with the homoloa-y of all the struc- 
 tures of the head-capsule of one or more representatives of ninety- 
 four of the one hundred and nine families of Goleoptera listed by 
 Leng in his recent catalogue. One hundred and fifty-five genera, of 
 which one hundred and thirty have been figured, nearly all represent 
 ing different important subgroups, have been studied. 
 
 2. This serial study has made it possible to identify the same 
 structures in a wide series of forms, and to definitely fix the ho- 
 mology of all the parts of the heed-capsule. 
 
 3. Hypothetical types have been constructed, based on the struc 
 ture of the head-capsule of generalized insects and Goleoptera. These 
 show the Goleoptera to have developed in general a consolidation of 
 sclerites and a heavier chi tinization of the wall of the head, a com 
 pacting of the head as a whole, and an approximation of the dorsal 
 and ventral movable parts at the cephalic end. 
 
 4. The epicrs.nial suture has been identified in all but two 
 
Vi :,r. X 
 
 
 ‘3 
 
 -M 
 
 ■^■11 
 
 
 
 - iif i - ^ 
 
 ■'' 9Ait B^.'. ^*/ 2 ^ /#:«iO Bdi 't^[ 
 
 ' , .>ia : ■ ' p' .'■■ -V 7 
 
 : ^i.- 
 
 SBB±Opr^Mf 
 
 '■•'I'X IS ti 
 
 J.:^ 
 
 .'<? '"', ■ ' • - 
 
 () 4 i- fiG*' , 0 «Jb 4 .iptJjctfQ[] 
 
 ' "f.- ' ' -F* 
 
 
 
 ' ■’ ' 31 ^ * e = * ' "*” ■ 1 
 
 
 iO- j) 0 'Jv f‘te .1 
 
 ' 
 
 s.'iT e/ 4 ^ 
 
 >iS 
 
 I n 
 
 «'fii<:C*7:i> a "il <5i?p'sv^i' ocrf' ®%jn FtoKgodoBrici'^? ■' P«^4ooi^«jmale!S-fr 
 
 
 n^t 
 
 ^a?»' 
 
 ± ■ 
 
 ^ f 
 
 f-. vV - ^H" 
 
 .it'"’ 
 
 
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 . eoaotj "«3 taX _ow|' ' »<^ aJtJ ^ 
 
 
 <: • 
 
 >,ip 
 
 i' 4 Si ■'■^ 
 
 ^r ^ j 3 ^ r® 
 
 t,i 
 
 itr 
 
 •\ 
 
 
 f* 
 
 1 -. 
 
 
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 akvUx' ^to ea^'M oXi;»a(jo4-|f»e^ 6St to 'ee't«4 ;|^.? 
 
 XX-i^'Yi't £)fi ^fe 4 cr. 9 ‘-?e«^ tiif vi>G^ »tx oifiV a^e- 
 
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62 
 
 of the genera studied. It has proved a great aid in determining the 
 limits of neighboring parts. »7hat may appear to he a distinct epi- 
 cranial suture may not even he a suture. It is sometimes distinctly 
 invaginated. Its identity can only he definitely fixed hy determin- 
 ing the location of the pretentorinae , which are always associated 
 with it. 
 
 5. The limits of the vertex are dependent upon the position of 
 the epicranial suture. In the Hhynchophora nearly all of the snout 
 belongs to the vertex. 
 
 6. The unmodified occipital suture has been identified only in 
 the Adephaga. The cephalic end of it on the ventral surface is al- 
 ways represented hy part of a curving ridge, which is present in all 
 hut a few genera. 
 
 7. The supratentorinae have been identified in a few genera, 
 nearly all of which belong to the iitaphylinoidea. 
 
 8. The pre tentorinae are the great landmarks of the head-capsulo 
 and have been identified in all but two genera. They are, in the 
 vast majority of genera, located near the cephalic end of the epi- 
 cranial arms. A definite determination of the pretentorinae cannot 
 always be made without an ental examination of the head. 
 
 9. The size and form of the front is dependent upon the posi- 
 tion of the epicranial arms. In the Cerambycoidea it is large. In 
 many genera, as illustrated by Omophron, Harpalus, and Tachinus, it 
 is partly or wholly invaginated. It may probably be rudimentary or 
 wholly lost in many genera in which the me sal parts of the epicranial; 
 arms have disappeared. 
 
 10. 7/hat has been celled the clypeal suture in such genera as 
 Gicindela and Harpalus is not even a suture, but the line of invagi- 
 

 
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63 
 
 nation of the front. 
 
 11. The clypeus is always divided into the postclypeus and the 
 preelypens. The preclypeus is with one exception always distinctly 
 membranous. It may be as large or larger than the labrxim. 
 
 IS. There ib a distinct clypealia present in the Ooleoptera, 
 and in widely separated groups, such as the Adephaga and the Geram- 
 bycoidea. 
 
 13. The lab rum may be indistinctly determined in both iihyncho- 
 phora and other Ooleoptera. It may also be quite distinct in some 
 Hhynchophora where it is considered to be absent. 
 
 14. The submentum is always located distinctly cephalad of the 
 occipital foramen, with a chitinized area between it and the foramen, 
 
 15. The me tatentorinae may be located on the cephalo-lateral 
 border of the occipital foramen, as in generalized insects, or they 
 may be far cephalad of this location. 
 
 16. All that region between the occipital foramen and the sub- 
 mentum is a part of the postgenae, produced by the fusion on the me- 
 son of the raesal margins of the postgenae. 
 
 17. The gular sutures result fro n the cephalic . migration of the 
 raetatentorinae . 
 
 18. The gula is that area included between the gular sutures, 
 and is, therefore, derived from the postgenae. The majority of the 
 Ooleoptera possess a gula that extends no more than half the distance 
 between the occipital foramen and the submentum. 
 
 j 19. The tentorium of the Ooleoptera is typically quite similar 
 in form and development to that of generalized insects. Frequent 
 nodif ications are loss of chitinization, loss of corpotentorium and 
 laminatentorium. Occasionally the pretentorium may be rudimentary. 
 

 
 
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64 
 
 The functions of the absent parts are assumed by other parts of the 
 tentorium, or by the pharynx, or the head may be so compsct and chi-: 
 tinized that a tentorium is no longer needed. 
 
 20. The cephalic migration of the submentum, and the subsequent 
 formation of an indistinguishable area between it and the occipital 
 foramen is due either to the caudad migration of the occipital for- 
 amen, or to the cephalic pull on the mouth-parts, or to both. The 
 cephalic migration of the metatentoria , and, therefore, the metaten- 
 torinae, with consequent production of the gula, is probably due to 
 the cephalic pull on the tentorium to furnish a firmer support for 
 the muscles and tendons of the mouth-parts. 
 

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65 
 
 VI. BIBIIOGHA.PHY 
 
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qS 
 
 t-'* 
 
 
 .oiaav" 
 
 ir ' 
 
 < V ! 
 
 .MX-av ;X ./^ti,fcfr.*j 11^ .eoex 
 ,e: .A .EIJUMIIi. ,.A ,©IIY08 
 
 .ftViii'ra/C' ^ aisi i^atwA fa 9B7tii*l .Os4x^‘ 
 
 f .I(jt :Yt» ,.aij3< •'' 
 
 u 61 - ^ ,2to^TeBwa 
 
 , JU &h ;^n:rtj;: ?0 'T'-^^xa-Vjit'r cCliiiXoYi 
 
 i''; 
 
 J^. 
 
 f ■' 
 
 V K* ■ .k-i& 
 
 ^ .ilii e ;exx-vi /I 
 
 4 
 
 _ .IHtJCl 5ac .; ,Ji9(y!Q!»KOO 
 
 - I ' '^- .' ■ , ' 
 
 :u.* ,,?4»R .**J ^ ai*A' fc co^eCaila .SO&f^ 
 
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 ▼V 
 
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 t ^ ^ 
 
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 t ^4 
 
 Tip’<I <w»i‘ ^i;**AOiv«^g tl it# MclX«;>ixiut«Xa iX 1#6<|^ K^f.t .&XtJI 
 
 0 ;oOX-I€ ;53 ..*X1 .^<iri ./toA .eiflTaq , jv 
 
 ij 
 
 . ‘ “i .r*J>l .u»’jWqoor«t v©fc 0tlMf3 aOi^#rt#a a*t ,0£I6X 
 
 «X<1 s ;^j1f CS |G*-i :.«0 », 
 
 r * ^ s' ^ 
 
 ?i 
 
 
 |V^ 
 
 .■^r .lo kt . rn . aax 1 1 1 ♦‘Xi OO0 wiit ^ a t - I #4.* - . 0S9 1 
 
 , # 
 
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66 
 
 39S-396; 7 figs. 
 
 GA1JGIBAU3H, I. 
 
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 HYSLOP, J. A. 
 
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 KOLBE, H. J. 
 
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 LAI.IEEEE, A. 
 
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 1903. Nouvelles notes pour la classification des Coleopteres. 
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67 
 
 IE CONTE, J. L., and HORN, G. H. 
 
 1883. Glassification of the Goleoptera of North America. 
 Smithsonian Mis. Coll., No. 507. 
 
 LENG, G. W. 
 
 19E0. Catalogue of the Goleoptera of America, north of Mexico. 
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 MUIR, P. 
 
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 PETERSON, A. 
 
 1915. Morphological studies on the head and mouth-parts of the 
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 1916. The head-capsule and mouth-parts of Diptera. 111. Biol, 
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 RILEY, 11 , A. 
 
 1904. The emhryological development of the skeleton of the head 
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 SCHIOLTE, J. G. 
 
 1861-1683, Le Me tamorphosi Eleutheratorum Ob servationes : Bidrap 
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 pis. Kjobenhaven. 
 
 SHARP, D. 
 
 1909. Insects. Cambridge Nat. Hist., 6: 184-298. 
 
 SHARP, D. , and MUIR, P. 
 
 1912. The comparative anatomy of the male genital tube in Co- 
 leoptera. Trans. Ent. Soc. Lond., 1912: 477-639; pi. 
 42-78. 
 

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 STMUS-DUxiKEjCilLr, H. 3. 
 
 1628. Considerations generales sur I’anatoraie coraparee des ani- 
 raaux articules, auxquelles on a joint I’anatomie descrip- 
 tive du Hanneton vulgaire. P. 51-58; 10 pis. 
 
EXPLAIIATIOH OP PLATE I 
 Dorsal aspect of the head 
 Pig. 1. Hypothetical head. 
 
 '2. Tetracha Carolina. 
 
 3. Gicindela forrnosa. 
 
 4. Galosona calidum. 
 
 5. Oniophron arnericanum. 
 
 6. Harpalus erraticus. 
 
 7. Peltodytes 12-punctatns . 
 
 8. Gyhister f imhriolatus . 
 
 9. Dineutes assimilis. 
 
 10. Molamha lunata. 
 
 11. Hydrous triangularis. 
 
 12. Hydrophilus obtusatus. 
 
 13. Hecrophorus Carolina. 
 
 14. Greophilus villosus. 
 
 15. Aleochara lata. 
 
 16. Gonnophron fossiger. 
 
 17. Tachinus fimbria tus. 
 
 18. Leptinus testaceus. 
 
 19. Stenus flavicornis. 
 
 20. Gastrolobium bicolor. 
 
EXPIANATION OF PLATE II 
 
 Dorsal 
 Pig. 21. 
 22 . 
 
 23. 
 
 24. 
 
 25. 
 
 26 . 
 
 27. 
 
 28. 
 
 29. 
 
 30. 
 
 31. 
 
 32. 
 
 33. 
 
 34. 
 
 35. 
 
 36 . 
 
 37. 
 
 38. 
 
 39. 
 
 40. 
 
 aspect of the head 
 Limulodes paradoxus. 
 
 Sphaerius politus. 
 
 Scaphidiuni quadriguttatum. 
 Hister merdarius. 
 
 Nacerda melanuro. , 
 
 Ivlacrosiagon dimidietum. 
 Calopteron terminale . 
 
 Photinus pyralis. 
 
 Phengodes plumosa. 
 Ghauliognathus pennsylvanicus . 
 Collops nigriceps. 
 
 Trichodes nuttali. 
 
 Necrohia rufipes. 
 
 Epioauta marginata. 
 
 Pytho americanus. 
 
 Gupes concolor. 
 
 Gephaloon lepturides. 
 
 Tomoxia hidentata. 
 
 Eurystethus dehilis. 
 Ueopyrochroa flabellata. 
 
PLATE II 
 
EXPLAKATIOI OP PLATE III 
 
 Dorsal 
 Pig. 41. 
 4E. 
 
 43. 
 
 44. 
 
 45. 
 
 46. 
 
 47. 
 
 48. 
 
 49 . 
 
 50. 
 
 51. 
 
 52. 
 
 53. 
 
 54. 
 
 55. 
 
 56 . 
 
 57. 
 
 58. 
 
 59. 
 
 60. 
 61. 
 62. 
 
 aspect of the head 
 Macratria murina. 
 
 IJotoxus anchora. 
 
 Sandalus niger. 
 
 Alaus ociilatus. 
 
 Throscus chevrolati . 
 
 Thar ops ruficornis. 
 
 Ghalcophora virginiensis . 
 Psephenus lecontei. 
 
 Helichus striatus. 
 
 Eucinetus morio. 
 
 Gyphon ruficollis. 
 
 Stenelmis sinuata. 
 
 Heterocerus undatus. 
 
 Georyssas californicus . 
 Derraestes lardarias. 
 
 Byrrhus americanus. 
 
 Nosodendron unicolor. 
 
 Tihysodes americanus. 
 Tenehroides sinuatus. 
 
 Phenolia grossa. 
 
 Glischrochilus quadriguttatus . 
 Gucujus clavipes. 
 
PLATE III 
 
EXPLAiJATIOIJ Oi" PLATE IV 
 
 Dorsal 
 Fig. 63. 
 
 64. 
 
 65. 
 
 66 . 
 
 67. 
 
 68 . 
 
 69. 
 
 70. 
 
 71. 
 
 72. 
 
 73. 
 
 74. 
 
 75. 
 
 76. 
 
 77. 
 
 78. 
 
 79. 
 
 80. 
 
 81. 
 
 82. 
 
 83. 
 
 aspect of the head 
 Phyconofiius raarihus . 
 Languria raozardi. 
 Megalodacne fasciata. 
 Derodontiis maculatus. 
 Anchicera ephippiata. 
 Byturus unicolor. 
 
 LIyce tophagus punctatus. 
 Philothermus glahriculus. 
 Phymaphora pulchella. 
 Phalacrus politus. 
 Alohates pennsylvanica . 
 Melanophthalma ansrularis. 
 Endoraychus higuttatus. 
 Hippodamia convergens. 
 Arthroraacra aenea. 
 Pseudocistela brevis. 
 Adalia bi-punctata. 
 Tenebrio molitor. 
 
 Boros unicolor. 
 Hyporphagus sp. 
 
 Penthe obliquata. 
 
PLATE 17 
 
t 
 
EXPLAIJATIOII OP PLATE V 
 
 Dorsal aspect of the head 
 
 Pig. 84. 
 
 Ptinus hrunneus. 
 
 85. 
 
 Sitodrepa panidea. 
 
 86. 
 
 Bostrichns hicornis. 
 
 87. 
 
 lyctus planicollis. 
 
 88. 
 
 Plesiocis crihrum. 
 
 89. 
 
 Ephindus americanus. 
 
 90. 
 
 Aphodius fimentarius. 
 
 91. 
 
 Dichelonyx elongata. 
 
 92. 
 
 Pelidnota punctata. 
 
 93. 
 
 Strategus iulianus. 
 
 94. 
 
 Harpalus erraticus, 
 longitudinal cross-section 
 
 95. Osrnoderma erernicola. 
 
 96. Trox suherosus . 
 
 97. Pseudolucanus dama. 
 
 98. Passalus cornuta. 
 
 99. 
 
 Byne ta ferruginea. 
 
 100. 
 
 Derohrachus hrunneus. 
 
 101. 
 
 G-lycohius speciosus. 
 
 102. 
 
 Tetraopes te traophthalmus . 
 
 103. 
 
 Donacia piscatrix. 
 
 104. 
 
 Griocerus asparagi. 
 
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PLATS V 
 
 A. 
 
EXPLANATION OF PLATE VI 
 
 Dorsal 
 Pig. .105. 
 106. 
 
 107. 
 
 108. 
 
 109. 
 
 110 . 
 111 . 
 118. 
 
 113. 
 
 114. 
 
 115. 
 
 116. 
 
 117. 
 
 118. 
 
 119. 
 
 120 . 
 121 . 
 122 . 
 123. 
 
 aspect of the head 
 Parandra hrunnea. 
 
 Crypt ocephalus quadruplex. 
 Chrysochus auratus. 
 Leptinotarsa decimlineata . 
 Eupsalis rainuta. 
 
 Blepharida rhois. 
 
 Anoplitis nervosa. 
 Ghelymorpha argas. 
 Pachymerus arthriticus. 
 Diahrotica 12-punctata. 
 Euryraycter fasciatus. 
 Spicaerns irahricatus. 
 Hhynchites hicolor. 
 Dendroctonus valens. 
 Thecesternus humeralis. 
 Attelahus analis. 
 ophenophorus aequalis 
 iicolytus quadrispinosTis . 
 Lixus macer. 
 
PLATE 71 
 
EXPLAilATIOH OF PLATE VII 
 
 Lateral 
 Pig:. 124. 
 
 125. 
 
 126. 
 
 127. 
 
 128. 
 
 129. 
 
 130. 
 
 131. 
 
 132. 
 
 133. 
 
 134. 
 
 135. 
 
 136. 
 
 137. 
 
 138. 
 
 139. 
 
 140. 
 
 141. 
 
 142. 
 
 143. 
 
 aspect of the heed 
 Tetracha Carolina. 
 Gicindela formosa. 
 Calcsoma calidnm. 
 Omophron americamira. 
 Peltodytes 12-punctatus . 
 Harpalus erraticus. 
 Gybister fimbriolatus . 
 Hydrous triangularis. 
 Hydrophilus obtusatus. 
 Leptinus testaceus. 
 Necrophorus Carolina. 
 Dineutes assirailis. 
 Gonnophron fossiger. 
 Ivlolaraba lunata. 
 otenus flavicornis. 
 Gastrolobium bicolor. 
 Greophilus villosus. 
 Tachinus fimbriatus. 
 Aleochara lata. 
 
 Limulodes paradoxus. 
 
EXPLAIIATIOW OF PLATE 71 1 1 
 
 Lateral aspect of the head 
 Fig. 144. Ephaerius politus. 
 
 145. Scaphidium quadriguttatum. 
 
 146. Ghaiiliognathus pennsylvanicus , 
 cross-section shov/ing invagination 
 of part of the gula. 
 
 147. Hister raerdarius. 
 
 146. Photinus pyralis. 
 
 149. Galopteron terminale . 
 
 150. Phenorodes plnraosa. 
 
 151. Ghauliognathns pennsylvanicns . 
 
 152. Gollops nigriceps. 
 
 153. Trichodes nuttali. 
 
 154. Necrohia rufipes. 
 
 155. Gupes concolor. 
 
 156. Gephaloon lepturides. 
 
 157. Hacerda melanura. 
 
 158. Tomoxia hidentata. 
 
 159. Macrosiagon dirnidiatnrn. 
 
 160. Epicauta marginata. 
 
 161. Eurystethus dehilis. 
 
 162. IJeopyrochroa flahellata. 
 
 163. Pytho americanus, 
 
 164. Harpalus erraticus, 
 
 dorsal aspect, showing invagination 
 associated with epicranial snture. 
 
SXi'LAMTION OF PLATF IX 
 
 lateral 
 Fig. 165. 
 166. 
 167. 
 166. 
 
 169. 
 
 170. 
 
 171. 
 17E. 
 
 173. 
 
 174. 
 
 175. 
 
 176. 
 
 177. 
 176. 
 179. 
 160. 
 161. 
 162. 
 163. 
 
 184. 
 
 185. 
 
 aspect of the head 
 Llacratria nmrina. 
 
 Notoxns anchora. 
 
 Eandalus niger. 
 
 Throscus chevrolati. 
 
 Alaus oculatus. 
 
 Tharops ruficornis. 
 
 Heterocerus -andatus. 
 Ghalcophora virginiensis . 
 Georyssus calif ornicus . 
 Psephenus lecontei. 
 
 Eelichus striatus. 
 
 Etenelmis sinuata. 
 
 Eucinetus raorio. 
 
 Gyphon ruficollis. 
 
 Dermestes lardarias. 
 
 Byrrhus americanus. 
 llosodendron unicolor. 
 
 Hhysodes americanus. 
 Tenehroides sinuatus. 
 
 Phenolia grossa. 
 
 Glischrochilus quadriguttatus . 
 
PLATii IX 
 
EXPLAIIATIOIJ OF PLATE X 
 
 Lateral aspect of the head 
 Fig. 186. 1137-00110111118 marinus. 
 
 187, Guc-ajus clavipes. 
 
 188, Languria mozardi. 
 
 189. Megalodacne fasciata. 
 
 190. Anchicera ephippiata. 
 
 191. 
 
 Byturus unicolor. 
 
 192. 
 
 Mycetophagus punctatus. 
 
 193. 
 
 Derodontus asculatus. 
 
 194. 
 
 Philo thermus glabriculus. 
 
 195. 
 
 ILelanophthalma angularis. 
 
 196. 
 
 Phymaphora pulchella. 
 
 197. 
 
 Sndomychus higuttatus. 
 
 198. 
 
 Phalacrus politus. 
 
 199. 
 
 Hippodamia convererens. 
 
 200. 
 
 Adalia bi-punctata. 
 
 201. 
 
 Pseudocistela brevis. 
 
 202. 
 
 Alobates pennsylvanica. 
 
 203. 
 
 Tenebrio molitor. 
 
 204. 
 
 Boros unicolor. 
 
 E05. Arthromacra aenea. 
 
PLATE X 
 
EXPIAIIATIOII OE PLATE XI 
 Lateral aspect of the head 
 Pig. 206. Hyporphagus sp. 
 
 207. Penthe ohliquata. 
 
 208. Ptinus hrunneus. 
 
 209. Eitodrepa panicea. 
 
 210. Lyctus planicollis. 
 
 211. Bostrichus hicornis. 
 
 212. Sphindus americanus . 
 
 213. Plesiocis crihrum. 
 
 214. Aphodius firaetarius. 
 
 215. Dichelonyx elongata. 
 
 216. Pelidnota punctata. 
 
 217. btrategus julianus. 
 
 218. Osmoderma eremicola. 
 
 219. Trox suherosus. 
 
 220. Pseudolucanus dama. 
 
 221. Passalus cornuta, 
 
 222. Parandra hrunnea. 
 
 223. Lerobrachus brunneus. 
 
 224. Glycobius speciosus. 
 
 225. Tetraopes tetraophthalmus . 
 
PLATE XI 
 
EXPLANATIOH 0? PLATE ZII 
 Lateral aspect of the head 
 Pig. 226. Donacia piscatrix. 
 
 227. Syneta ferruginea. 
 
 228. Criocerus asparagi. 
 
 229. Gryptocephalus quadruplex. 
 
 230. Ghrysochus auratus. 
 
 231. Leptino tarsus decimlineata . 
 
 232. Liahrotiea 12-punctata . 
 
 233. Blepharida rhois. 
 
 234. Anoplitis nervosa. 
 
 235. Ghelymorpha argus. 
 
 236. Pachyraerus arthriticus . 
 
 237. Eupsalis mimita. 
 
 238. Eurymycter fasciatus. 
 
 239. xihynchites hicolor, 
 
 240. Attelabus analis. 
 
 241. Epicaerus imbricatus. 
 
 242. Lixus raacer, 
 
 243. Thecesternus huraeralis. 
 
PLA^^ XII 
 
EXPLAMTIOIT OF PLATE XIII 
 Lateral aspect of the head 
 Fig. 244. Ephenophorus aequalis. 
 
 245. Ecolytus quadri spinosus . 
 
 246. Dendroctonus valens. 
 Ventral aspect of the head 
 
 Fig. 247. Hypothetical head. 
 
 248. Tetracha Carolina. 
 
 249. Gicindela formosa. 
 
 250. Calosoma calidum. 
 
 251. Harpalus erraticus. 
 
 252. Omophron araericanum. 
 
 253. Peltodytes 12-punctatus . 
 
 254. Gybister f imbriolatu-s . 
 
 255. Lineutes assimilis. 
 
 256. Hydrous triangularis . 
 
 257. Hydrophilus obtusatus. 
 
 258. Leptinus testaceus. 
 
 259. Hecrophorus Carolina. 
 
 260. Gonnophron fossiger. 
 
 261. Molamba lunata. 
 
EXPLAHATIOH OF PLATE XI Y 
 Ventral aepect of the head 
 Fig. 262, Stenus flavicornis. 
 
 263. Gastrolo'bium hicolor, 
 
 264. Greophilus villosus. 
 
 265. Tachinus fimbriatus, 
 
 266. Aleochara lata. 
 
 267. Limulodes paradoxus. 
 
 268. Sphaerius politus, 
 
 269. Scaphidium quadri gut ta turn. 
 
 270. Hister merdarius. 
 
 271. Calopteron terminale. 
 
 272. Photinus pyralis, v 
 
 273. Phengodes plumosa. 
 
 274. Chauliognathus pennsylvanicus . 
 
 275. Gantharis bilineatus. 
 
 276. Gollops nigriceps. 
 
 277. Trichodes nuttali, 
 
 278. Necrobia rufipes. 
 
 279. Gupes concolor. 
 
iiXPLAlIATIOlJ OF PIATS X7 
 
 Ventral aspect of the head 
 
 Fig. 280. 
 
 Cephaloon lepturides. 
 
 281. 
 
 ITacerda melanura. 
 
 282. 
 
 Tomoxia hidentata. 
 
 283. 
 
 IvI[acrosiagon dimidiat-ara. 
 
 284. 
 
 Epicauta raarginata . 
 
 E85. Eurystethus dehilis. 
 
 286. 
 
 Pytho americanus. 
 
 287. 
 
 IJeopyrochroa flahellata. 
 
 288. 
 
 Macratria raurina. 
 
 289. 
 
 Sandalus niger. 
 
 290. 
 
 Notoxus anchora. 
 
 291. 
 
 Alans ocnlatiis. 
 
 292. 
 
 Tharops ruficornis. 
 
 293. 
 
 xhroscus chevrolati. 
 
 294. 
 
 Ghalcophora virginiensis . 
 
 295. 
 
 Psephenns lecontei. 
 
 296 
 
 Helichus striatus 
 
PLATE X¥ 
 
TOi UBS*® 
 
 on«' 
 
EXPIAIJATIOH OF PLATS X7I 
 
 '/entral aspect of the head 
 
 Fig. 297. 
 
 Stenelrais sinuata. 
 
 298. 
 
 Heterocerus undatus . 
 
 299. 
 
 Oeoryssus calif ornicus . 
 
 300. Encinetus morio. 
 
 301. 
 
 Cyphon ruficollis. 
 
 302. 
 
 Dermestes lardarius . 
 
 303. 
 
 Byrrhus americanis. 
 
 304. 
 
 Nosodendron unicolor. 
 
 305. 
 
 Ehysodes araericanura. 
 
 306. 
 
 Tenebroides sinuatus. 
 
 307. 
 
 Phenolia grossa. 
 
 308. 
 
 Olischrochilus quadriguttatus . 
 
 309. 
 
 Phyconomus marinus. 
 
 310. 
 
 Cucujus clavipes. 
 
 311. 
 
 Languria raozardi. 
 
 312. 
 
 Megalodacne fasciata. 
 
 313. 
 
 Derodontus maculatus. 
 
 314. 
 
 Anchicera ephippiata. 
 
 315. Byturus uni color. 
 

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EXPLANATIOIJ OF PLATE Till 
 
 Ventral aspect of the head 
 
 Pig. 316. 
 
 Kycetophagus punctatas. 
 
 317. 
 
 Philothermus glahriculus 
 
 318. 
 
 Melanophthalma angularis 
 
 319. Phymaphora pulchella. 
 
 320. 
 
 Endomychus higuttat-as . 
 
 321. 
 
 Phalacrus politus. 
 
 322. 
 
 Hippodaraia convergens. 
 
 323. 
 
 Adalia hi -punctata. 
 
 324. 
 
 Pseudocistela hrevis. 
 
 325. 
 
 Alobates pennsylvanica. 
 
 326. 
 
 Tenebrio molitor. 
 
 327. 
 
 Boros unicolor. 
 
 328. 
 
 Arthromacra aenea. 
 
 329. 
 
 Hyporphagus sp. 
 
 330. 
 
 Penthe obliquata. 
 
 331. 
 
 Ptinus brunneus. 
 
 332. 
 
 Sitodrepa panicea. 
 
 333. 
 
 Bostrichus bicornis. 
 
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PLATii XVll 
 
EXPLAIIATION OP PLATE XVIII 
 
 Ventral aspect of the head 
 Pig. 334. Lyctus planicollis. 
 
 335. Ephindus americanus. 
 
 336. Plesiocis crihiurn. 
 
 337. Aphodius f ime tarius . 
 
 338. Lichelonyic elongata. 
 
 339. Pelidnota punctata. 
 
 340. Etrategus julianus. 
 
 341. Osraoderma eremicola. 
 
 34S. Trox suberosus. 
 
 343. Pseudolucanus dama. 
 
 344. Passalus cornuta. 
 
 345. Parandra brunnea. 
 
 346. Derobrachus brunneus. 
 
 347. Olycobius speciosus. 
 
 348. Tetraopes te traophthalmus . 
 
 349. Donacia piscatrix. 
 
 350. Eyneta ferroginea. 
 
 351. Criocerus asparagi . 
 
 352. Gryptocephalus quadruplex. 
 
PLATS XVIII 
 
EXPLANATION OF PLATE XIX 
 Ventral aspect of the head 
 Fig. 353. Ghrysochus anratiis. 
 
 354. Leptinotarsa decimlineata . 
 
 355. Diahrotica lE-panctata. 
 
 356. Blepharida rhois. 
 
 357. Anoplitis nervosa. 
 
 358. Chelymorpha argus. 
 
 359. Pachymerus arthriticns. 
 
 360. Enpsalis minuta. 
 
 361. Euryraycter fasciatus. 
 
 36E. Ehynchites hicolor. 
 
 363. Attelabus analis. 
 
 364. Epicaerus imhricatus. 
 
 365. Scolytus quadrispinosus . 
 
 366. Lixus macer, 
 
 367. Thecesternns h-ameralis. 
 
 368. Ephenophorus aequalis. 
 
 369. Lendroctonns valens. 
 
PLATE XIX 
 
PI ATE XX 
 
 374 
 
EXPLANATION OP PLATE XXI 
 
 Endoskeleton of the head 
 
 Pig. 387. 
 
 Tachinus fimhriatus. 
 
 388. 
 
 Aleochara lata. 
 
 38S. 
 
 Li.'nulodes paradoxus. 
 
 390. 
 
 Ephaerius politus. 
 
 391. 
 
 Ecaphidium quadriguttatum. 
 
 39E. 
 
 Hister merdarius. 
 
 393. 
 
 Galopteron terrainale. 
 
 394. 
 
 Photinus pyralis. 
 
 395. 
 
 Phengodes plumosa. 
 
 396 . 
 
 Chauliognathus pennsylvanicus , 
 
 397. 
 
 Collops nigrioeps. 
 
 398. 
 
 Trichodes nuttali. 
 
 399. 
 
 Necrohia rufipes. 
 
 400. 
 
 Gupes concolor. 
 
 401. 
 
 Gephaloon lepturides. 
 
 402. 
 
 Nacerda raelarmra. 
 
 403, Tomoxia hidentata. 
 
PLATE XXII 
 
EXPLAMTIOII OF PLATE XXIII 
 
 Endoskeleton of the head 
 
 Fig. 419. 
 
 Heterocerus undatus. 
 
 420. 
 
 Oeoryssus californicus . 
 
 421. 
 
 Eucinetus morio. 
 
 422. 
 
 Cyphon ruficollis. 
 
 423. 
 
 Derraestes lardarius. 
 
 424. 
 
 Byrrhus americanus . 
 
 425. 
 
 llosodendron unicolor. 
 
 426. 
 
 Ehysodes americanus. 
 
 427. 
 
 Tenebroides sinuata. 
 
 428. 
 
 Phenolia grossa. 
 
 429. 
 
 Glischrochilus quadriguttatus . 
 
 430. 
 
 Phyconomus marinus. 
 
 431. 
 
 Cucujus clavipes. 
 
 432. 
 
 Languria raozardi. 
 
 433. 
 
 Megalodacne fasciatus. 
 
 434. 
 
 Lerodontus maculatus. 
 
 435 
 
 Anchicera ephippiata 
 
PLATE XXIII 
 
EXPLAHATIOil OF PLATE XXIV 
 
 Endoskeleton of the head 
 Fig. 436. Byturus unicolor. 
 
 437. Mycetophagus punctatus. 
 
 438. Philo the rraus glahriculus. 
 
 439. llelanophthalma angularis. 
 
 440. Phyraaphora pulchella. 
 
 441. Endomychus higuttatus. 
 44E. Phalacrus politus. 
 
 443. Hippodamia convergens. 
 
 444. Adalia hi -punctata. 
 
 445. Pseudocistela hrevis. 
 
 446. Alohates pennsylvanica. 
 
 447. Tenehrio molitor. 
 
 448. Boros unicolor. 
 
 449. Arthromacra aenea. 
 
 450. Hyporphag:us sp. 
 
 451. Penthe ohliquata. 
 
EXPLAIIATIOII OF PI ATE XXV 
 
 Endoskeleton of the head 
 Fig. 45E. Ptinus hrunneus. 
 
 453. Eitodrepa panicea. 
 
 454. Bostrichus bicornis. 
 
 455. lyctus planicollis. 
 
 456. Sphindas americanus. 
 
 457. Plesiocis cribruna. 
 
 458. Aphodius firnetarins. 
 
 459. Dichelonyx elongata. 
 
 460. Pelidnota punctata. 
 
 461. Btrategus julianus. 
 
 462. Osmoderma erimicola. 
 
 463. Trox suberosus. 
 
 464. Pseudolucanus dama. 
 
 465. Passalus cornutus. 
 
 466. Passalus cornutus, 
 
 lateral aspect, cross-section. 
 
 
EXPIANATIOiJ OF PLATE XXVI 
 
 Endoskeleton of the head 
 Fig. 467. Parandra brunnea. 
 
 468. Derobrachus brunneus. 
 
 469. Glycobius speciosus. 
 
 470. Tetraopes tetraophthalmus . 
 
 471. Donacia piscatrix. 
 
 472. Syneta ferruginea. 
 
 473. Criocerus asparagi. 
 
 474. Gryptocephalus quadruplex. 
 
 475. Blepharida rhois. 
 
 476. Ghrysochiis auratus. 
 
 477. Leptinus testaceus. 
 
 478. Diabrotica 12-punctata. 
 
 479. Anoplitis nervosa. 
 
 480. Pachymerus arthriticus. 
 
 481. Ghelyraorpha argus. 
 
SXPIAHATION OF PLATE XX7II 
 
 Endoskeleton of the head 
 Fig, 482. Eupsalis ininuta. 
 
 483. Eurymycter fasciat-us. 
 
 484. Hhynchites hicolor. 
 
 485. Attelahus analis. 
 
 486. Epicaerus irahricatus. 
 
 487. Thecesternus humeralis. 
 
 488. Lixiis macer, 
 
 489. Sphenophorus aequalis. 
 
 490. Ecolytus quadrispinosus . 
 
 491. Lendroctonus valens. 
 
V 
 
VITA 
 
 The author of this thesis, Fenner Satterthwaite Stickney, was 
 horn in Wilson, North Carolina, August 6, 1892, He attended the 
 public schools of Wilson, until May 1909. The following year he 
 spent on a farm in Beaufort County, North Carolina. During the sum- 
 mer of 1910, he moved to the Hawaiian Islands, and entered in Sep- 
 tember of that year the McKinley High School of Honolulu, from which 
 he graduated in June 1912. From September 1912 until June 1914, he 
 studied agriculture, including a course in Entomology, at the Col- 
 lege, now the University, of Hawaii. In August 1914, he registered 
 in the College of Agriculture of the University of California; he 
 majored in Entomology, and received the degree of Bachelor of Sci- 
 ence in Agriculture from that University in May 1916. The year 1916 
 -1917 he spent on a farm in Brunsv/ick County, North Carolina, en- 
 gaged in practical agriculture. He attended the Summer Session of 
 1917 at the University of Illinois, specializing in economic Ento- 
 mology. During the year 1917-18, he was registered in the Graduate i 
 School of the University of Illinois, from which he received the de- 
 gree of Master of Science in Entomology in June 1918, June 1, 1918, 
 he was inducted into the United States Army and ordered to Camp 
 Dodge, lov/a, vrhere he was engaged in sanitary entomological work un- 
 til August 1918, when he was sent overseas. After six months in 
 France as a member of Sanitary Squad #70, he was returned to the U- 
 nited States, and received his discharge in Honolulu in June 1919. 
 From September 1919 to June 1921 he was a Fellow in Entomology at 
 the University of Illinois. During the summer of 1920 he assisted 
 in extension entomological work at the North Dakota Ag'ri cultural 
 College . 
 
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EXPLAMTIOil OF FLATE XX7III 
 
 .493. Amphizoa lecontei, dorsal aspect, 
 
 493. Amphizoa lecontei, lateral aspect. 
 
 494. Amphizoa lecontei, ventral aspect. 
 
 495. Amphizoa lecontei, endoshele ton. 
 
 496. Pilopius lacustris, dorsal aspect. 
 
 497. Pilopius lacustris, lateral aspect 
 
 498. Pilopius lacustris, ventral aspect 
 
 499. Pilopius lacustris, endoskele ton. 
 
 500. Othnius sp., dorsal aspect. 
 
 501. Othnius sp., lateral aspect. 
 
 502. Othnius sp., ventral aspect. 
 
 503. Othnius sp . , endoskele ton. 
 

 
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