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THE UNIVERSITY OF ILLINOIS LIBRARY NATURAL HISTORY SURVEY 5705 ILL v7cop.4 ILLINOIS BIOLOGICAL MONOGRAPHS Vol. VII October, 1922 No. 4 Editorial Committee Stephen Alfred Forbes William Trelease Henry Baldwin Ward Published under the Auspices of the Graduate School by the University or Illinois Press Copyright, 1923 by the University or Illinois Distributed February 9, 1923 A CLASSIFICATION OF THE LARVAE OF THE TENTHREDINOIDEA WITH FOURTEEN PLATES BY HACHIRO YUASA Contributions from the Entomological Laboratories of the University of Illinois No. 69 THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENTOMOLOGY IN THE GRADUATE SCHOOL OF THE UNIVERSITY OF ILLINOIS 1920 TABLE OF CONTENTS I. Introduction 7 II. Morphology 14 III. Taxonomy 35 Superfamily Tenthredinoidea 37 Families of Tenthredinoidea 38 Family Xyelidae 39 Family Blasticotomidae 42 Family Tenthredinidae 42 Subfamilies of Tenthredinidae 43 Subfamily Diprioninae 45 Neodiprion Rohwer 46 Species of Neodiprion 47 Monoctenus Dahlbom 48 Diprion Schrank 49 49 Subfamily Emphytinae , Genera of Emphytinae 50 Subfamily Selandriinae 51 Genera of Selandriinae 52 Thrinax Konow , 52 Species of Thrinax 53 Strongylogaster Dahlbom 53 Species of Strongylogaster 53 Selandria Leach 54 Subfamily Dolerinae 54 Dolerus Jurine 55 Species of Dolerus 56 Subfamily Phyllotominae 57 Tribe Phyllotomini 57 Genera of Phyllotomini 58 Endelomyia Ashmead 58 Caliroa Costa 58 Species of Caliroa 59 Tribe Phlebatrophini 59 Phlebatrophia MacGillivray 60 Subfamily Tenthredininae 60 Genera of Tenthredininae 61 Macrophya Dahlbom 62 Species of Macrophya 63 Subfamily Cimbicinae 64 Genera of Cimbicinae 65 Cimbex Olivier 65 Trichiosoma Leach 65 Abia Leach 65 Species of Abia 66 Subfamily Hoplocampinae 66 Genera of Hoplocampinae 67 Hemichroa Stephens 67 Marlattia Ashmead 68 Caulocampus Rohwer 68 Subfamily Dineurinae 69 Subfamily Cladiinae : 70 Genera of Cladiinae 71 Trichiocampus Hartig 71 Species of Trichiocampus 72 Priophorus Dahlbom 73 Species of Priophorus 73 Cladius Rossi 74 Subfamily Nematinae 74 Genera of Nematinae 75 Diphadnus Hartig 76 Pnstiphora Latreille 77 Species of Pristiphora 77 Micronematus Konow 79 Lygaeonematus Konow 81 Pachynematus Konow 81 Species of Pachynematus 82 Nematus Panzer 82 Species of Nematus 83 Croesus Leach 83 Amauronematus Konow 84 Species of Amauronematus 84 Pteronidea Rohwer 85 Species of Pteronidea 85 Pontania Costa 88 Species of Pontania 88 Subfamily Blennocampinae 91 Genera of Blennocampinae 92 Tomostethus Konow 92 Blennocampa Hartig 93 Erythraspiaes Ashmead 94 Monophadnus Hartig 94 Hypergyricus MacGillivray 94 Species of Hypergyricus 94 Monophadnoides Ashmead 95 Isodyctium Ashmead 95 Subfamily Fenusinae 96 Genera of Fenusinae 96 Kaliofenusa MacGillivray 97 Fenusa Leach 97 Subfamily Scolioneurinae 98 Metallus Forbes ;. ... 98 Species of Metallus 99 Subfamily Hylotominae 99 Hylotoma Latreille 100 Species of Hylotoma 100 Subfamily Schizocerinae 101 Schizocerus Lepeletier 102 Subfamily Acordulecerinae 103 Acordulecera Say 103 Species of Acordulecera 103 Family Pamphiliidae 104 Species of Pamphiliidae 105 Family Cephidae 108 Genera of Cephidae ...;•.. 109 Janus Stephens 110 Species of Janus 110 Adirus Konow 110 Trachelus Jurine Ill Cephus Latreille ... Ill Species of Cephus Ill Hartigia Schiodte 112 Family Xiphydriidae 112 Xiphydria Fallen 113 Family Siricidae . 114 Tremex Jurine 115 Family Megalodontidae 116 Family Oryssidae 117 Oryssus Latreille 118 IV. Phylogeny 120 V. Summary 133 VI. Bibliography 135 VII. Explanation of Plates 141 VIII. Index 169 325J LARVAE OF THE TENTHREDJNOIDEA—YUASA I. INTRODUCTION That the cardinal principle of modern taxonomy is based on the funda- mental facts of evolution and that the essential problem of classification is the phylogenetic relationship of organisms need no argument. In order to ascertain genetic affinities, it is not sufficient to investigate the morphological characters alone, but all other attributes, physiological and biological, must be considered. It is also evident that the immature stages of organisms should receive as thoro consideration as the adult if taxonomy of insects is to attain that degree of comparative perfection obtained in the classification of other organisms. Systematic entomologists, dealing as they do with animals of such diversity and complexity morphologically and biologically, have from early times recognized, at least to some extent, the taxonomic significance and value of the developmental stages of insects, but the practical difficul- ties in obtaining necessary materials, accurately determined and adequate in quantity and range, have made progress in this phase of insect taxonomy very tardy. A good start, however, has been made by recent workers as was pointed out by Brues (1919), and their results vindicate both the possibilitity and practicability of such investigations. There is, moreover, an urgent demand for such studies from economic entomologists, who are constantly confronted by the problem of identifying the immature stages of economic species. The present study is an attempt to deal with the larvae of the Ten- thredinoidea from the standpoint of synoptic and, to some extent, genetic classification. The systematic significance of the morphological characters will be discussed in part two; the taxonomic treatment of the families, subfamilies, genera, and species will constitute part three; and, as full a discussion of the phylogenetic relationship of the families as is possible with the data at hand, will form part four. No one appreciates the inade- quacy of this study, both in thoroughness and comprehensiveness, more than the author, but it is hoped that he has opened a way for those who will advance our knowledge of this highly interesting group of insects to a more satisfactory condition in the future. The taxonomic literature dealing with the adults of the Tenthredinoidea is extensive. The historical development of the subject is interesting to students of this group of insects but a detailed account is out of place here. However, a brief statement of the history of the group is desirable. 8 ILLINOIS BIOLOGICAL MONOGRAPHS [326 Linnaeus in the fourth edition of the Systema Naturae (1744) estab- lished the order Hymenoptera under the name of Gymnoptera and applied to the order its present designation in the first edition of the Fauna Suecica. The name Piezata was proposed by Fabricius (1775) for the order, but this name never came into general use. Latreille (1796), following Lin- naeus, divided the order into two sections, Terebrantia and Aculeata. The first section included two groups, Phytophaga, which comprises the Tenthredinoidea, and the Entomophaga or parasitic Hymenoptera. The Ditrocha and Monotrocha of Hartig (1837) correspond approximately with the two sections of Latreille. Gerstaecker clearly recognized the Tenthredinoidea as a unique compact group and proposed in 1867 to divide the order Hymenoptera into two suborders. He used the name Symphyta for the Tenthredinoidea and Apocrita for the remainder of the order. The term Symphyta thus antedates Konow's (1890) subordinal name Chalastogastra. Various terms have been proposed for this group of Hymenoptera and the following are coextensive with the superfamily name Tenthredinoidea as used in the present paper: Phytophaga, Ses- siliventres, Securifera, Serrifera, Symphyta, and Chalastograstra. Rohwer and Cushman (1917) proposed a third suborder of Hymenoptera, Idiogas- tra, for the family Oryssidae and placed it between the Chalastogastra and Clistogastra of Konow. Early students of the Tenthredinoidea divided the superfamily into two groups, Phyllophaga for the Tenthredinidae or "Tenthredo" of Linnaeus and Xyllophaga for the Siricidae or "Urocerus" of Geoffroy. With the exception of Stephens (1835) and Andre" (1879), who recognized the additional families Xiphydriidae and Cephidae, respectively, besides the two families mentioned above, the old system was followed for many years. With the progress in studies of the world fauna of this group of insects, modern writers h<ave proposed many elaborate schemes of classification. Konow in 1890 suggested one family and three subfamilies and Dalla Torre (1894) catalogued one family divided into eighteen subfamilies, while Ashmead (1898) proposed fifteen families and twenty-seven subfamilies. Enslin (1911) criticized Konow's three divisions as unnatural and proposed four families, Oryssidae, Siricidae, Cephidae, and Tenth- redinidae, thus reverting to a considerable extent to the scheme of the old school as represented by Cameron (1882) and others. The recent and more important systems are those proposed by Konow (1905), MacGilliv- ray (1906), and Rohwer (1911). These systems, when compared, show a great discrepancy in the number and rank of the groups which formerly constituted the family Tenthredinidae, as is indicated graphically in Plate XIV. MacGillivray, whose classification is based on a thoro-going phylogenetic study of the wings, is of the opinion that the large complex of genera obtained in this family are readily separable into a number of 3271 LARVAE OF THE TENTHREDINOIDEA—YUASA 9 definite groups on structural differences, and that they are best dealt with by considering them simply as subfamilies. In general the systems of Konow and Rohwer are, with the exceptions noted below, more in accordance with each other in their essential features than either one of them is with that of MacGillivray. A comparison of these three

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