THE UNIVERSITY OF ILLINOIS LIBRARY rL_L_ 5 - V. c_op- 2 CD 00 < ' sturn this book on or before the itest Date stamped below. A arge is made on all overdue oks. University of Illinois Library UL28: .952 &i;g4 1952 %Po S IQ";^ 'APR 1 1953 DFn 7 W54 '•> d ^r-. ''/./'ji. Lit]—H41 Digitized by tine Internet Arciiive in 2011 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/genericclassific15ross ILLINOIS BIOLOGICAL MONOGRAPHS Vol. XV No. 2 Published by the University of Illinois Under the Auspices of the Graduate School Ukbana, Illinois 1937 EDITORIAL COMMITTEE John Theodore Buchholz Fred Wilbur Tanner Harley Jones Van Cleave UNIVERSITY OF ILLINOIS 1000—7-37—11700 ,. PRESS A GENERIC CLASSIFICATION OF THE NEARCTIC SAWFLIES (HYMENOPTERA, SYMPHYTA) WITH SEVENTEEN PLATES BY Herbert H. Ross Contribution No. 188 from the Entomological Laboratories of the University of Illinois, in Cooperation with the Illinois State Natural History Survey CONTENTS Introduction 7 Methods 7 Materials 8 Morphology 9 Head and Appendages 9 Thorax and Appendages 22 Abdomen and Appendages 29 Phylogeny 33 The Superfamilies of Sawflies 33 Family Groupings 34 Hypothesis of Genealogy .... 35 Larval Characters 45 - Biology 46 Summary of Phylogeny 48 Taxonomy 50 Superfamily Tenthredinoidea 51 Superfamily Megalodontoidea 106 Superfamily Siricoidea 110 Superfamily Cephoidea 114 Bibliography 117 Plates 127 Index 162 ACKNOWLEDGMENT This monograph is an elaboration of a thesis sub- mitted in partial fulfillment for the degree of Doctor of Philosophy in Entomology in the Graduate School of the University of Illinois in 1933. The work was done under the direction of Dr. W. P. Hayes, to whom I am indebted for unceasing help throughout the investigation. I am indebted also to Dr. R. Malaise, of Stockholm, Sweden, to Mr. Takeuchi, of Kyoto, Japan, and to Mr. O. Conde, of Riga, Latvia, for much valuable Eurasian material; and to Mr. R. B. Benson, of the British Museum, for a great deal of assistance and information on the genotypes of many genera. A great aid in my work has been the frequent dis- cussion of many points regarding morphology, tax- onomy, nomenclature, and phylogeny with Dr. H. J. Van Cleave, of the University of Illinois, and with Dr. T. H. Prison, Dr. C. O. Mohr, and Dr. B. D. Burks, of the Illinois State Natural History Survey. INTRODUCTION Few problems in the Hymenoptera have been so disputed as the gen- eral classification of the sawfly groups. Practically no two authors have agreed either on the subject of major groupings or the basic reasons for their divisions. The present study is an attempt to find fundamental morphological evidence which will separate the sawflies into groups and show the paths of evolution followed in their development. While such a study is almost necessarily incomplete without a lifetime of work, never- theless structures have been found which show definitely some of the relationships and groupings of these insects. It is, I believe, the first time such a study has been based on the comparative morphology of the body and its appendages, rather than on only those characters which may be seen without dissection. It seems worth while to present these findings now in an endeavor to put the classification of the sawflies on a more stable basis. Another grave difficulty confronting the North American sawfly worker has been the almost total lack of illustrated keys to the genera. Little effort has been made in the past to correlate the generic names used in various parts of the world, so that in many cases a considerable re-shuffling has been imperative. The solution of these problems has been linked very closely with the classification of the higher categories. In the unravelling of these problems, palaeontology has given us no clues, since the earliest known hymenopterous fossils belong to specialized groups which occur at the present day. These include such families as the Cephidae, Orussidae, and Siricidae. Fossils showing the nature of the earliest and most primitive Hymenoptera are still unknown. The utiliza- tion of embryological evidence has been impossible because of lack of material except in very few instances; and it is doubtful if a great deal could be learned from this source. This has left open only one avenue of approach, comparative morphology. The problem has therefore resolved itself into piecing together evidence drawn only from the dift'erences and similarities found in the parts of those sawflies existing at the present day. Methods A large number of the drawings were made from pinned specimens. This is especially true of the characters used in generic keys, such as antennae, tarsi, sheaths, etc. The remainder were made from cleared specimens. The specimens were soaked in 25 per cent caustic potash (KOH) for three to twenty- four hours, depending on their darkness, hardness, etc. The male genitalia were preserved in glycerine in small vials ; the saws, wings, and legs were mounted in either balsam or euparal ethyl on slides ; and the larger structures were preserved in 80 per cent alcohol. ;; 8 ILLINOIS BIOLOGICAL MONOGRAPHS Materials The taxonomic portion of this study is based on an examination of about 50,000 sawflies, including 90 per cent of the types of nearctic species. This material has been accumulated through the courtesy of the officers of the following institutions: Academy of Natural Sciences of Philadelphia; Brigham Young University; California Academy of Sci- ences ; Canadian National Museum ; Colorado Agricultural College Cornell University; Deutsches Entomologische Institut; Entomological Branch, Canadian Department of Agriculture; Field Museum of Chicago; Illinois State Natural History Survey ; Iowa State College ; Iowa Wes- leyan College ; Kansas State Agricultural College ; MacDonald College McGill University; Montana State Agricultural College; New York State Museum ; Ohio State University ; Oregon Agricultural College ; Purdue University ; South Dakota Agricultural College ; Texas Agricultural Col- lege; United States National Museum; University of Alberta; University of British Columbia; University of Colorado; University of Idaho; Uni- versity of Illinois; University of Kansas; University of Michigan; Uni- versity of Nebraska; University of Oklahoma; University of Wisconsin; Washington Agricultural College. MORPHOLOGY The first work to be published dealing with the comparative morphol- ogy of sawfiies was Hartig's "Die Familien der Blattwespen und Holzwespen" in 1837. This contains descriptions and many illustrations of the mouthparts, antennae, and other miscellaneous characters. It is, in fact, a veritable classic of sawfly morphology. Since that time several workers have greatly increased our knowledge of sawfly morphology. Cameron in various papers (1882, etc.) gave descriptions and illustrations of many parts in other species. Van Dine (1906) described and illustrated the mouthparts of Pamphilius; Mac- Gillivray (1906) made an exhaustive study of the wings of sawfiies; Snodgrass (1911) illustrated the thoracic structure of many forms; Crampton (1919) described and illustrated almost all types of male genitalia found in the group; Boulonge (1924) made a remarkably fine study of the male genitalia, with special reference to the muscles and their attachment; Bird (1926) made a detailed morphological study of Hoplocampa halcyon (Norton) ; and Taylor (1931) has made a complete histological and internal anatomical study of the head of a few species of Tenthredinidae. Marlatt (1896), Smulyan (1923), Middleton and Rohwer (in several papers) have made some morphological studies, mostly in connection with taxonomic work. Aside from these, references to sawfly morphology are scattered in general texts and papers on insect anatomy and phylogeny. In the following discussion no attempt is made to offer a complete treatise on sawfly morphology. Only those characters have been discussed which, on close study, appeared to offer differences indicating relation- ships of groups, or to possess differences which might be used in such a capacity at a later date. The following parts have been studied and illus- trated in greatest detail: the head, its endoskeleton and appendages; the external anatomy of the thorax and first two abdominal segments; the wing venation and the male genitalia. In addition to these major parts some other characters, such as tibial spurs, have been of considerable help in corroborating the phylogenetic arrangements presented herein. Head and Appendages Head Capsule The head is always hypognathous, having the mouthparts directed downwards in repose. They may be directed forward when in use, but this is caused by a swinging forward of the entire head in a pendulum motion, the hinge being partly at the odontoidea and partly at the dorsal margin. This latter point of articulation is due to the moving of the head and "atlas" together (see p. 23). ; 10 ILLINOIS BIOLOGICAL MONOGRAPHS Many sutures and sclerites found in the heads of primitive insects are obsolescent or entirely absent in the heads of sawflies. See Plate I, figs. 1, 2. Not only this, but no other parts have been added. The epicranial suture (ep) is present in only a few forms (e.g., Pergidae, Megalodon- tidae), and in these it is only a crease ending at a point between the lateral ocelli. The clypealiae and mandibulariae are entirely absent. The clypeus (cl) is not divided transversely into a pre- and postclypeus. It is separated from the front (f) by a furrow instead of a well-marked suture, and separated from the genae (g) by a suture or furrow. The lateral arms of the epicranial sutures are lacking, so that the definition of the vertex and front is practically impossible. The anterior aspect of the head, how- ever, is provided with several furrows, ridges, and areas which bear a fixed relation to the ocelli, eyes, base of antennae, and tentorinae. These have been given a terminology by Marlatt, Rohwer, and MacGillivray and although the names bear no relation to the hypothetical areas of the head, they are of great use in taxonomic description.