FEVISION OF THE HIGHER CATEGORIES OF STIGMODERINI (COLEæTERA : BUPRESTIDAE) JENNIFER ANNE GARDNER B. Sc. (Hons) (Aderaide) Department of ZoologY The University of Adelaide A thesis submitted for the degree of Doctor of PhilosoPhY FEBRUARY 1986 L tn¡o o-, eAP o( ej - 4 -{ BI F s rl T}tE RI],GI.STRY Mr. I-.L. Carrnan Asslstant. ReglsErar- (Sc Lence) Tel 228 5673 ILC;DßA;DPl.7 7l,Lay, l9{Jli )ls. Jennif er A. Gardner, DEPARTMT,NT O}' ZOOLOCY. Dear ]"ls . Gardner, the degree I am oleased to lnform you that you quallfl-ed for the award of of Doctor of Philosophy for your tht.sis entirlecl "Revision of ttre lligher õ;.;fS;i;"-or siig*oà.rini (ôoleoptera ; Bupresttrlae)" on 29 April- I986' Copi¿es of che reports are enclosecl for your lnformaËion. "*"rln"r"r lìfinor corrections are reqttirecl to be ma,le to yotlr Ehesis, therefore would you take up thls lnairer with your supervi-sor as aoon as posslble' In fhe nor$al course of events fhe degree will be conferred at the- annual commemoration ceremony to be helcl fn Aprfl/May 1987 ancl I should be grateful lf you rvould comnlete the enclosed form of appllcatlon for adrnfsslon to a hfgher degree and return it to me as soorì as possible ' I any shoulcl point out, however, that the degree cannot be conferred untll outstanàing tlnion or Library fees have been patd' ltith respect to your application for tìre withho-l ding of ot:rmissj-on for photocopying or ior.t, bof-h the t'acrrlty of Sclence a'cl Lhe B,ard of Research Studles consldereC that your best, rJeferrce against Ëhe posslbí-lity ot plagiarlsnr is Ehe speeciy publlcation of your work' t'¡lth thls in mincl the Boar<l of Research studJ.es has granÈecl pern{ssion for you to wlEìrhokl the consenE Eo loan and phot<)copy iot ott".y""t ln the first lnsÈalrce, with posslbilíty of extendin;1 this permÇ!îãî eor a further ye-ar rrpon rtrceipt by t-he Board of a wriÈten iequerst from you in la¡e 1986 anrl of a reporE on Ehe current situat-ton regarriinfl the possibility of p1a¡4iarlsm' Fi.na]lyr måy L offer you tlly congrattr.lations on the otrtstanclin:'{ reports you receivecl f or -r*our Ehesis. Yours sincert'r1-y, õrs @T llncs , cc Chai.rrnan, DeoarttnenÈ of Zoology Supervisor' Dr. S. llarker To Doug TABLE OF CONTENTS Page i SUMMARY DECLARATION iii iv ACKNOWLEDG]VENTS I I INTRODUCTION IIMA TERIALS AND METHODS I Materials 15 2 KaryologY 15 3 MorphologY and nomenclature 16 4 Scanning electron microscoPY T7 5 Histology 18 6 Taxonomic characters T9 7 Phylogenetic methods 19 I Criteria for suPrasPecific taxa 22 Iil RESULTS t KaryologY 24 2 MorphologY 2.L General- form and colouratíon of adults 26 2.2 Head 27 2.7 Thorax 30 2.4 Abdomen 33 2.5 FemaLe reProductive sYstem 36 2.6 Male reProductive sYstem 40 2.7 AlimentarY sYstem 43 2.8 Nervous and respiratory systems M TABLE OF CONTENTS (cont.) Page IV DISCUSSION 1 KarYotYPe analYsis 46 2 Morphological character analysis 50 7 Phylogenetic analYsis 8I 4 Taxonomic implications of phylogenetic analysis 94 5 Distribution of genera 97 6 Concluding remarks 98 V SYSTEMATTCS I Tribe Stigmoderini 100 2 Tribe Jul-odimorPhini r28 APPENDICES A Museums and private collections from which material was obtained, and abbreviations used I3I of B Species of Stigmoderini dissected and aspects iñlèrnaf anatoñy studied; additional species of Stigmoderini examined externally only L72 C Modified karyological method of Imai, Crozier and 140 Taylor (1977 ) DOtherbuprestidspeciesinwhichtheovipositor and/or fämale reproductive system was studied I42 E Comparison of number of tubul-es/testis in the higher tãiã of Stigmoderini with that in other buprestids L49 FCompari-sonofthearlangementofMalpighiantubules in btigmoderini with thát in other buprestids 151 G Species of Cal-odema: a preliminary revision 154 HspeciesofMetaxymorpha:apreliminaryrevision 179 204 REFERENCES SUMMARY The tribe Stigmoderini (Coleoptera: Buprestidae), comprising about 640 described species, occurs in Australia, New Guinea and South America, suggesting a Gondwanan origin of the tribe. At present' 480 of these are placed in the genus Stigmodera with 3 subgen era: Stigmodera (s. s.), Themognatha, and Castiarina, which represents about half the Austral-ian buprestid fauna. The remaining species of the tribe are placed in Hypostigmodera, Calodema, Metaxymorpha, Conognatha., Dactylozodes and Hyperantha. The inclusi.on of Curis and Julodimorpha in the tribe by Théry Q929) is contentious. In this study, the karyology, internal anatomy, and external- morphology of the higher taxa of Stigmoderini were investigated. Chromosome counts were obtained for 75 species of Stigmodera, and karyotypes of l-5 species are presented. Thirty-three species have a diploid complemenL of 22 chromosomes, two species have l¡ = 20. All s a speci-es studied h.ave an XYO sex-determining mechanism. Mal-e Curis and JuLodimorpha have dipJ.oid complements of 14 and 79 respectively. The most useful- morphological characters are those associated with the femal-e reproductive system. The highly derived ovipositor of Stigmoderini possessing a dorsal val-ve, particle Sac, medial styles and spatulate setae, was compared with that of 84 other buprestid species representing 47 genera in 25 tribes, and was found to be unique, providing strong evidence that the tribe is monophyletic. Curis and Julodimorpha which l-ack this type of ovipositor are excluded from the tribe. f The higher categori-es are delineated by suites of characters and monophyly is established for some, but not al-L taxa. Relationships between the higher taxa of the tribe are investigated' Synapomorphic characters which suggest confl-icting relationships are examined criticalfy and the probabiJ-ity of homoplasy assessed' A cladogram is constructed which best fits the avail-able evidence. The femal-e accessory gland, a structure not recorded previously in any buprestid, is accorded high weight and supports the monophyly of tha - cal_odema - Metaxymorpha. This necessitates removing Themognatha from Stiqmodera and elevating it to generic status' Arguments are also presented to support the elevation of Castiarina to generic status, and to transfer to it the speci-es variegata BJ-ackburnt which at present is placed in the monotypic genus Hypostigmodera' No evidence was found to suggest that the Australian and South American components of the fauna represent different lineages, indeed the data suggest that Conognatha is more closely rel-ated to the Australian fauna than it is to the other south American genera of the tribe. This thesis contributes to the knowledge of Stigmoderini by adding data on aspects previously neglected, such as female genitaliar and by providing an overview of the tribe not attempted by earl-j-er workers. Furthermore, it uses investigative approaches and methods of analysis that may have broader application to the field of coleopteran systematics. t1 DECLARATION This thesis contaj-ns no material acceþted for the award of anY other degree or diploma in any university, and to the best of my knowledge, contalns no material prevÍously published or written by another personr except where due reference is made in the text. ].rt ACKNOWLEDGMENTS Many.individuals and institutions aided my"study. I thank warmly my supervi-sor, Dr Shel-Iey Barker, for his help and enthusiasm. Specimens were l-ent by the following institutions: Australian Museum, Sydney (¡¡r e. Holloway); Australian National Insect Collection' Canberra (Dr J. Lawrence); British Museum (Natural History), London (¡¡r n. o. Pope); Department of Primary Industry, Konedobu, P. N. G. (Dr J. Ismay); Insect Farming and Trading Agency, BuJ-olo, P. N. G. (Mr p. Clark); Museum of Victoria, Melbourne (Dr A. Neboiss); Queensland Museum, Brisbane (Dr G. B. Monteith); and South Austral-ian Museum (Or r. Matthews). SeveraL individual-s sent l-ive material enab]-ing me to expand the Scope of my research: Mrs N. Gawthorne, Toongabbie, N.S.W.; Anthony Hil-ler, Mt Glorious, Qld; Allen Sundholm, Sydney; Stan þlatkins, Summer Hill, N.S.W.; and Geoff. Williams, Lansdowne, N.S.W. Dr James Johnson, University of Idaho; Dr Tomas Moore, Laja, Chile; Prof. Gay1e Nelson, Pomona, California; and George Walters, Los Angeles sent pinned and fluid-preserved material- which was unavail-abl-e in Australia, and of considerable help to my studY. The Directors of the fol-lowing departments gave permission to col-lect in natÍona1 parks and reserves: National Parks and Wildlife Service, S. A.; National Parks Service, Victoria; Department of Fisheries and Wildlife, Vi. A.; and National- Parks and ltJildlife Servicet N. S. W. 1V This work was funded by a Commonwealth Postgraduate Research Award and Austral-ian Museum Trust Postgraduate Grant. Prof. Bill Witli-ams, Zoology Department, University of Adelaide, gave me access to his word processor to produce this thesis. The plates were produced with instruction from Philip Kempster; Figure 24 and overlays of Figures 6 and f0 were prepared with the assistance of Ruth Evans, both of the ZooJ.ogy Department. I am grateful to Alice WelLs, Dr Tom Burton, Prof. Bill Williams and especially Dr Shell-ey Barker for their critical- reading of earlier drafts; and Dr Eric Matthews, Dr Derek Duckhouse and Mike Kokkinn for stimulating discussions. My husband Doug supported me in every possibJ-e way, sharing my moments of elation, and encouraging me j.n times of despondency. He was aLso an indefatigabl-e field assistant. Finally, I acknowÌedge Prof. Frank Beer who first kindled my interest in buprestids, on Christmas eve I97I, in Death ValÌey, CaLifornia. V lt 'lii ì'rililil 'r iii1 .tì ir i,,li. ', ¡.11 i"\. 1 Buprestidae (Col-eoptera) are commonly called jewel beetles because many are brilliantly coloured, with a metall-ic sheen.