By Wattuthanthrige Thelma Thusitha Poojitha Gunawardane B

A STUDY OF THE CLASSIFICATION OF THE CERAPHRONIDAE (HYMENCertARA) By Wattuthanthrige Thelma Thusitha Poojitha Gunawardane B. Sc. (Hons.) Ceylon., F. R. E. S., F. L. S. A Thesis Submitted For The Degree Of Doctor Of Philosophy In The Faculty Of Science Of The University of London. Department Of Zoology And Applied Entomology Imperial College Of Science And Technology, South Kensington, London. S.W. 7. July 1968. 2 ABSTRACT The study of the fatfly Cciraphronideein the British confirmed the desirability of raising this family to the status of a of superfamily. This thesis gives keys to separate the families4Coraphronoideal and the subfamilies and genera of these families. In the subfamily Megaspilidae of the family Megaspilinae, the genus Conostigmus is treated in greater detail and a key to the British species is given. This genus is here divided into two subgenera,Conostigmus and Consulcus (subgenus new). Twenty-eight species are redescribed with figures to facilitate recognition, and twelve new species are described and figured. One new genus Scuto.cervix is erected for a single species previously described in Conostigmus. The genus Megaspilus is redescribed with figures. Among the types studied in the British Museum, seven have been placed as new synonyms. In the course of the study several European species not represented in the British Isles were examined. Several new characters for identifying genera and species have bean discovered; greater attention being given to use of sculpture and comparative measurements than previously. 3 CONTENTS Page Acknowledgements 7 I(a) Introduction 9 (b)Review of literature 17 (c)General morphology and terminology 22 II(a) Material, technique and illustrations 29 (b)Preparation and technique 29 III Biology 30 IV(a) Taxonomy 46 Classification (b) Key to the families 51 (c)Key to the subfamilies of Megaspilidae 53 (d)Key to the genera of Ceraphronidap of the British Tsles 54 (e)Key to the genera of Megaspilinae of the British Isles 56 (f)Key to the genera of Lagynodinae 60 V Genus Conostigmus Dahibom 1958 (a)General description 61 (b)Key to subgenera 62 la Subgenus Consulcus subgen; n 65 b Key to the species of subgenus Consulcus 66 C. (C.) wasmanni (Kieff.). 83 Page C. (Ci)nigriventris Kieff. 87 C. (Co) divisifrons Kieff. 91 C. (C.) levifrons Kieff. 92 C. (C.) testaceipes (Kieff.) 97 C. (C.) planifrons Kieff. 97 C. (C.) lucidus Kieff. 99 C. (C.) myrmecobius Kieff. 104 C. (C.) mullensis (Cameron) 107 C. (C.) multicolor Kieff. 108 C. (C.) tetanoptcrus Kieff. 113 C. (C.) Species I sp. n. 115 C. (C.) versicolcr Kieff. 120 C. (C.) brachypterus (Thomson) 122 C. (C.) leptothorax Kieff. 123 'C. (C.) punctulatus (Cameron) 128 C. (C.) ruficollis Kieff. 131 C. (C. punctatifrons Kieff. 133 C. (C.) scabrivontris Kieff. 138 C. (C.) halteriger Kieff. 141 C. (C.) bipunctatus Kieff. 143 5 Page C. (C.)_ melenopus Kieff. 147 C. (C.) foveatifrons Kieff. 148 C. (C.) hiemalis Kieff. 148 C. (C.) tricolor Kieff. 149 C. (C.) apricans Kieff. 149 C. (C.) subspinosus Kieff. 150 C. (C.) basalis Kieff, 150 2.Subgenus Conostigmus Key to the species of subgenus Conostigmus 151 c. C. (C.) fasiatipennis Kieff. 159 C. (C.) rufescens Kieff. 161 C. (C.) dubiosus Kieff. 165 C. (C.) Species III sp. n. 167 •C. (C.) Species VIII sp.n. 171 C. (C.) Species IX sp. n. 172 C. (C.) Species X sp.n. 174 (-II. (C.) Species VII sp. n. 176 C. (C.) Species XII sp.n. 178 C. (C.) opacus (Thomson) 179 C. (C.) Species. XI sp. n. 181 6 Page C. (C.) alutaceus (Thomson) 185 Co (C.) britta*cus Kieft.- 187 C. (C.) Species II sp. n. 188 C. (C.) marshalli (Kieft.) 191 C. (C.) apterus Kieft. 193 C. (C.) Species IV sp. n. 195 C. (C,) Species V sp.n. 199 C. (C.) Species VI sp. n. 201 VI.Characters separating the genera Conostigmus and Scutocervix 204 VII.Genus Scutocervix gen. n. 206 Scutocervix carpentieri (Kieff.) 206 VIII.Megaspilus hispanicus (Kieff.) 212 IX,Discussion. 217 X, Literature cited 226 7 Acknowledgements This study was carried out in the Department of Zoology and Applied Entomology, Imperial College of Science and Technology, London. I am greatly indebted to Professor O.W. Richards, the former Head of the Department for suggesting and supervising this study, making his collection of this group available to me, and for the encouragement he has given me throughout the course. I am grateful to Professor T.R.E. Southwood, present Head of the Department for the facilities provided. I also extend my thanks to Mr. R.G. Davies, Senior leoturer of the Department, for the interest he has shown in my work and for translating some of the French text. Access to material in the British Museum.(Natural HistorY), was made possible through the courteous permission of the Keeper of Entomology which is gratefully acknowledged. My thanks are also due to Dr. J.F. Perkins, Deputy Keeper, Mr. J. Quinlan, Mr. C. Vardy, Mr. T. Huddleston and Mrs. M. Young of the Hymenoptera Section, and Mr. K. Smith of the Diptera Section, of the museum, for their kind co-operation during the time I spent there studying the material of the group. Among other helpers of my work are included three members of the staff of the Commonwealth Institute of Entomology to whom I extend my very sincere thanks. Mr. G.E.J. Nixon for loaning me his private collection of the group and for the interest he has shown in my work, Mr. R.D. Eady, for his advice and technical remarks, with 8 special reference to sculpture of Hymenoptera; for the translations of several Latin and French papers, for borrowing type species of Conostismus from Professor Lindroth of Lund, Sweden, and for his interest and encouragement. Mr. G.J. Kerrich for so kindly helping me with the Swedish translations. Help received from the following is gratefully acknowledged. Dr. J. Hollebone, Mrs. G. Pike, Mrs. A. Balasubramanium for the translations of French papers, and Mr. C. Vardy for the Russian. I take this opportunity to thank the Director of the Museum National d'Histoire Naturelle Paris and Dr. S. Kellner Pillautt of the Hymenoptera Section of that Museum for the co-operation and kindness extended to me during my short visit to study the types there. My grateful thanks are also due to Prof. E. Tortonese, Director of the Museo Civioo DIStoria Naturale, Genova for his kind oo-operation and to Dr. Delta Guiglia of the Hymenoptera section for the assistance and extreme kindness shown to me during my short visit there to study the type material. I gratefUlly acknowledge the financial aid received from the Association of Commonwealth Universities, United Kingdom, for a Scholarship awarded by them. I also wish to thank the Director Gen3ra.l. and the staff of the Scholarship branch of the British Coundil for the interest they have shown in my progress here. Finally, my grateful thanks to the Director of National Museums, Ceylon, and the Government of Ceylon for granting me study leave for postgraduate studies in the United Kingdom. 9 (a) Introduction This study deals with the classification of the Superfamily Ceraphronoidea (Masner 1956), viz. the Ceraphronidae and Megaspilidae. In the past there has been some controversy about the status of these groups. At various times, they have been given different ranks, such as tribe, super genus, subfamily and family under different names. This is evident in the brief historical review of the literature which will follow later. The majority of authors have referred to a family Ceraphronidae with the subfamilies Ceraphroninae and Megaspilinae within the long established Superfamily Proctotrupoidea. The principle criterion for this classification was based on the very apparent wing- venation, the shape of the pterostigma being regarded as the main distinguishing characteristic between the subfamilies. Ashmead in 1893 states "The group seems to divide naturally into two tribes distinguished as follows: Marginal vein stigmated; antennae, with the same number of joints in both sexes, 11 - jointed Tribe I. Megaspilini. Marginal vein linear never stigmated; antennae, with a less number of joints in the females than in the males; males with 10 - or 11 - jointed antennae; females 9 . or 10 - jointed 000.60101100, Tribe II. Ceraphronini". Due to the importance ascribed to this very clear character the genus Lag nodes F8rster was placed in the subfamily Ceraphroninae. If this character is to be retained for the recognition of the major divisions, one has to ignore other important morphological characters I0 of the metasoma, the absence of the organ of Waterston, and the formula of the tibial spines. In this study, using the above mentioned characters, the genus Lagynodes has been placed in the family Megaspilidae. This supports the conclusions of Masner and Dessert (1966). Consequently, the variation in the wing-venation appear large but not fundamental. Thus one is able to conclude, that it is by a phenomenon of convergence, that the wing of Lagynodes, truly a Megaspilid, has come to resemble that of the Ceraphronidae. This hypothesis is supported in the discussidn at the end of this work. There is a very pronounced tendency continually, to restrict the extent of families and consequently increase their number. This is greatly deplored by many zoologists, and regarded as inevitable, and not necessarily highly undersirable, by others. The reasons for such a change are manifold. The continual discovery of new forms rapidly increases the number of known species and this in itself, merely on the basis of numbers makes it easier to deal with classification if we have a greater number of units of convenient size. The more careful study of anatomical structure frequently leads to the discovery that certain groups are polyphyletic, that is to say, are not of common origin, but represent a convergence or parallelism in the possession of certain closely similar characteristics.

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