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IMAGING SERVICES NORTH Boston Spa, Wetherby West Yorkshire, LS23 7BQ www.bl.uk This PDF was created from the British Library’s microfilm copy of the original thesis. As such the images are greyscale and no colour was captured. Due to the scanning process, an area greater than the page area is recorded and extraneous details can be captured. This is the best available copy TH E BRITISH LIBRARY DOCUMENT SUPPLY CENTRE TITLE k PHYLOGENETIC STUDY OF THE CYNIPOIDEA (HYMENOPTERA). AUTHOR Nigel Donald MacDade FERGUSSON INSTITUTION and DATE The City o£ London Polytechnic C M. A- Attention is drawn to the fact that the copyright of this thesis rests with its author. This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with its author and that' no information derived from it may be published without the author’s prior written consent. THE BRITISH LIBRARY DOCUMENT SUPPLY CENTRE ~n — n - n 20 cm t. United Kkifdom REDUCTION X . K PHYLOGENETIC STUDY OF THE CYNIPOIDEA (HYMENOPTERA), by Nigel Donald MacDade FERGUSSON A thesis submitted In partial fulfilment of the requirements of the Council for National Academic Awards for the degree of Doctor of Philosophy. March 1990 The City of London Polytechnic In collaboration with The Natural History Museum. ABSTRACT A phylogenetic study o£ the Cynlpoldea (Hymenoptera) N.D.M. Fergusson The current classification of the Cynipoldea was subjected to compatibility analysis on the basis of the characters then employed and shown to have a very poor resolution. A comprehensive morphological investigation of 31 exemplar species was undertaken and 234 characters were found, a 450% increase over the established classification. These characters were analysed and the compatibility clique contained 135 apomorphies, 68 of which were synapomorphies. This is an increase of 600% and 300% respectively over the established data, this is a tremendous improvement in the data-base leading to a great improvement in resolution. In addition, the principal morphological character-suites were analysed independently. This technique was used to locate the weaknesses in earlier classifications and, by this method, the distortion caused by the allometric bias to wing-data was recognized. Many extralimital cynipoids were examined and a new tribe was discovered. Other forms of cladistlc analysis. Parsimony and O'Nolan weighting, were undertaken and the J^®®ults considered. After detailed analysis, a phylogeny of the Cynipoidea was reconstructed. The evolutionary biology of the Cynipoidea provided particularly strong support for the phylogenetic reconstruction. Concepts of host defence, host range, gall complexity, gall position, host switching, reproductive cycles, biogeographical distribution, plate tectonics, palaeobotany, palaeoclimatology, the origin of hyper­ parasitism and the adaptive characters associated with xylophagous hosts were all considered. All the available fossil cynipoids were examined and the evolutionary history, holophyly, and relationships of the Cynipoidea with other Hymenoptera were all discussed. Finally the phylogenetic reconstruction was used to provide the first reasoned classification for the super family. Page 2 ACKMOWLEDOBMINTS The following depositories provided Cynlpoldea on loan: American Museum of Natural History, New York; Blosystematlcs Research Institute, Ottawa; Department of scientific and Industrial Research, Auckland; Instltut fOr Pflanzenschutzforschung, Eberswalde; Muse« Royal des I'Afrlque Centrale, Tervuren; Museum fOr Naturkunde der Humboldt-Unlverslttt, Berlin; Museum of Comparative Zoology, Harvard; Museum of Victoria, Melbourne; National Museum of Ireland, Dublin; National Museum of Natural History, Washington; National Museum of Wales, Cardiff; Naturhlstorlsches Museum, Wien; Naturhlstoriska Rlksmuseet Stockholm; Royal Ontario Museum, Ontario; University Museum, Oxford; Zoological Museum, Bergen; Zoological Museum of the University of Helsinki; Zoologlsche Sammlung des Bayerlschen Staates, Munich; Zoologlsk Museum, Copenhagen; Zoological Museum, Lund. I am pleased to record my thanks to the following people In the Natural History Museum. Dr L. Mound (Keeper of Entomology). My colleagues In the Hymenoptera Section, their talent never ceases to Impress. Mr. T. Huddleston for reading the thesis and Miss P. Gilbert for checking the references. Dr P. Eggleton for very many stimulating conversations about our respective Ph.D.'s. I am also grateful to: Dr R. Blackman, Ms. S. Barnes; Dr G. Gibson; Mr. c. Malumphy, Dr A. Menke; Mr. T. Parmlnter; Mr. J. Quinlan; Dr A. Ritchie; Dr J. Spradbery; Dr G. Stonedal A Dr L. Vet. I am particularly keen to thank my supervisors. Dr, G. Underwood and Dr I Gauld, It would be very hard to find better. Dr Underwood kindly provided "state of the art" compatibility programs, and Dr Gauld*s extensive knowledge was a great asset. Most of all, I thank my family for their support. Perhaps when my daughter can read this, she will understand why I could not always come out to play. Page 3 TABLB OF CONTENTS Abstract............................................ 2 Acknowledgements.................................... 3 Table of contents.................................. 4 List of figures.................................... 7 List of tables......................................... 15 1 INTRODUCTION C METHODS...............................17 Introduction..................................... 17 The study g r o u p .................................19 Material...................................... ... Selection of operational t a x a .................. 20 Methods......................................... 21 2 CLADISTIC THEORY.....................................24 Phylogenetic reconstruction .................. 24 Homoplasy.....................................24 Holophyly.....................................25 Cladlstlc analysis...............................25 Parsimony.....................................26 Compatibility.................................27 Choice of analytical methods.................... 28 Computer programs ............................ 29 Compatibility programs .................... 29 Character weighting...........................30 Parsimony program............................. 30 Character scoring ............................ 31 Assignment of evolutionary polarity .......... 31 3 ANALYSIS OF THE ESTABLISHED CLASSIFICATION.......... 34 Establishment of outgroups.......................34 The taxonomic- history of the Cynlpoldea .... 37 Character-states from Weld, 1952.............. 39 Analysis of Weld d a t a ...........................41 Character-states from Quinlan, 1979 ........... 47 Analysis of Quinlan data.........................50 Analysis of combined d a t a .......................56 Page 4 4 SURVEY OF CYNIPOID MORPHOLOGY .................... 59 Head.......................................... ... Antennae.................................. ... Thorax........................................ ... Wings...................................... .. 1<«98........................................137 Oaster........................................ .. Petiole.................................... .. Remaining gastral segments ................ 143 Female genitalia .......................... 145 Female accessory glands ................ 14S Male genitalia............................ .. Karyology.................................... .. Immature stages .............................. I53 The «99......................................153 Larval Instars ............................ 155 Pupal stage................................ .. 5 PHYLOGEMY OF THE CYMIPOIDEA........................176 Compatibility analysis o£ master data matrix. 176 Last characters deleted..................... 179 The clique cladogram ...................... 104 Subset analysis............................ .. Loss characters............................ .. Possible Improvements to tree............... 193 O'Nolan character weighting analysis........... 197 Parsimony analysis............................ .. Comparison o£ compatibility t parsimony trees . 201 The primary division o£ the Cynlpoldea......... 205 Extrallmltal Cynlpoldea ...................... 208 Fossil Cynlpoldea ............................ 219 Relative value o£ the character-suites......... 225 H e a d ........................................226 Antennae....................................227 T h o r a x ......................................227 I«e9S ...................................... 229 Wln9S........................................229 Page 5 Petiole......................................230 O e s t e r ......................................232 Feawle genitalia............................232 What was wrong with the old classification? . 233 Conclusions....................................235 Pinal postulate of cynlpold phylogeny. 235 6 DISCUSSION OP EVOLUTIONARY BIOLOGY................. 236 Introduction....................................238 Phytophagy: the Cynlpldae......................243 Biological trends within the Cynlpldae . 248 Parasitism within trees: the Iballld lineage. 254 Characters associated with xylophagous hosts. 259 Biogeography....................................262 Parasitism outside trees: the flgltld lineage . 270 SuBBsary of cynlpold evolutionary biology . 278 7 CONCLUSIONS ...................................... 281 Relationships with other Apocrlta ............ 281 The holophyly of the Cynipoidea............... 287 A new classification
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  • Hymenoptera, Symphyta) of Japan V

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    Bull. Natl. Mus. Nat. Sci., Ser. A, 46(4), pp. 183–202, November 20, 2020 Taxonomic Notes and New Distribution and Host Plant Records for Sawflies and Woodwasps (Hymenoptera, Symphyta) of Japan V Akihiko Shinohara1 and Hideho Hara2 1National Museum of Nature and Science, 4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan E-mail: [email protected] 2Nishi 4 Kita 3–4–29, Bibai, Hokkaido 072–0033, Japan E-mail: [email protected] (Received 24 August 2020; accepted 23 September 2020) Abstract Aglaostigma kawazoei Togashi, 2007 (Tenthredinidae) is a new synonym of Aglaostigma amoorense (Cameron, 1876). Euura damnacanti (Takeuchi, 1922) (Tenthredinidae) is redescribed and a lectotype is designated. Euura longa (Takeuchi, 1952) is redescribed and divided into two subspecies, E. longa longa (Takeuchi, 1952) from Honshu and E. longa shiretoko Hara and Shinohara, subsp. nov. from Hokkaido. New host records are: Betula platyphylla Sukaczev var. japonica (Miq.) H. Hara (Betulaceae) and Lithocarpus edulis (Makino) Nakai (Fagaceae) for Tremex apicalis Matsumura, 1912 (Siricidae), Quercus crispula Blume (Fagaceae) for Periclista erythrogramma Togashi, 1999 (Tenthredinidae) and Leucaena leucocephala (Lam.) de Wit. (Faba- ceae) for Lataxiphyda nodai (Togashi, 1982) and Kalopanax septemlobus (Thunb.) Koidz. (Aralia- ceae) for Xiphydria ogasawarai Matsumura, 1927 (Xiphydriidae). New distribution records are: Aglaostigma naitoi Togashi, 1972 (Tenthredinidae) from Kyushu and Aglaostigma yasumatsui Togashi, 1970, Euura tibialis (Newman, 1837) and Periclista erythrogramma Togashi, 1999 (Ten- thredinidae) from Hokkaido. New collection records are given for two rare species, Xiphydria kanba Shinohara, Hara & Smith, 2020 (Xiphydriidae) and Xyela kamtshatica Gussakovskij, 1935 (Xyelidae), and a “drumming” behavior of the male is newly recorded for Xiphydria ogasawarai (Xiphydriidae).