Coleoptera : Buprestidae
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
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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 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. The larvae are predominantly wood-borers in trees or shrubs: a few are leaf-miners, ga1l-makers, or exogenous root-feeders living in the soil. Larval- life may be prolonged and emergences after 25 years have been reported (Britton, 19702565). Adults are comparatively short lived, for a single spring oI summel, and feed on fl-owers, foJ-iage, fruit or bark' The family is cosmopolitan and particularly abundant in the tropics. Worldwide there are about L5,000 species in 400 genera (Lawrence, 1982). In AustraLia, there are at least 800 species: the genus Stiqmodera contains more than 480 of these, and according to the most recent checklist of Australian buprestids, Carter (1929), the rest are placed in 47 other genela. The higher classification of the Buprestidae is contentious, with the number of subfamilies worldwide variously reported as: 13 (Cobos, 1980; Nelson, 1981); 12 (Brues, Melander and Carpenter, L954); 7 (Bily , 1974); or 5 (Lawrence , 1982) . Arnett (1960) and Barr Q97I) placed Nearcti-c genera in four subfamilies. The tribe Stigmoderini poses some interesting biological and evol-utionary questions. One of the striking features of the tribe is the prolific speciation and radiation of Stigmodera which appears to paralleI that of Eucalyptus on which many of them feed. There are numerous other examples of speciose coleopteran genera associated with eucalypts e.g. Paroosis ( Chrysomelidae) with some hundreds of species, and Gonipterus and Oxyops (Curculionidae) (aritton I97Oz6]-l-, 619). Ehrlich and Raven (1964) proposed that plant-herbivore interaction was the major factor responsible for generating terrestrial- diversity. In the case of Stigmodera , speciation may have resul-ted from the 2 evoLutionaly pressule on the wood-boring larvae to develop the capacity to detoxify different biochemical defences produced by the different species of host eucalypt, which in turn may have been produced in response to predation by a diversj-ty of phytophagous insects (E. Matthews, pers. comm., l-985). New Q987) estimated that at i-east one quarter, and perhaps one third, of AustraLiats 60,000 terrestrial phytophagous insects depend on Acacia or EucaI tus for some specific resource. Another distinctive feature of the tribe is the great variety of patterns and bright colours displayed by the adults, which would have evolved under a quite different set of selective pressules, possibly associated with the avoidance of predation. Some have general-ized aposematic colouration of red or yellow bands on a bl-ack ground' Other species have been linked with specific mimicry of lycid beetles, bees, asilid flies, and cockroaches, often associated with mimetic behaviour (Nicholson, 1929; S. Barker, pers. comm.' 1980). Not al'l Stigmoderini' however, display warning colouration. Some are cryptic, for examplet those castiarina species which closely resembl-e the pale yel]ow eucalypt blossoms on which they feed and bask. The evol-ution of a great variety of colours and patterns in insects necessarily exposed to avian perception, as are the nectar-feeding Stigmodera I mây have been due to apostasy or frequency-dependent selection. As a strategy, the presentation of a kaleidoscope woul-d enabl-e Stiqmodera species to avoid being the subject of a search image, and take advantage of the hesitation and caution predators may evince towards novel- or unfamiliar items (¡'tatthews and Kitching, 1984)' This seems to me to be a quite likely explanation because it is not uncommon to col-lect a dozen specimens of Stigmodera from one tree, and find that'. they represent half a dozen species. 7 Moore and Brown (1985) found in aLl 14 species of Stigmodera tested, both conspicuous and cryptic, and Julodimorpha bakewelli' a series of bitter substances which they called buprestins. They reported that pure buprestin A produced a long-lasting and intensely bitter sensation when tasted, and a biological test indicated that it rendered sugar solution unpalatabl-e to meat ants. The authors suggested that buprestids are nealer the Mullerian rather than the Batesian end of the mimetic spectrum. 0n the other hand, buprestids are frequently attacked by ants (Gwynne and Rentz, 1987; pers. obs.) and eaten by birds (Sitberglied and Eisner, 1969). I have found scats of emus containing numerous fragments of the elytra of stigmodera tibialis, and a magpie has been observed catching and eating Jul-odimorpha bakeweLl-i (D' Booth and M' Thompson, pers. comm. 1984). In addition, Barker and Inns Q976) recorded predation on Stigmodera tibialis by an asilid fly' Clearly' the efficacy of buprestins in deterring predators is open to question' In the case of avian predation, rather than being an all or none proposition, it is more likely that a sllght seLective advantage is given the conferred when the bird hesitates, or prefers another insect, choice. Yet another possible explanation for the variety of bright colours is mate recognition. Visua] cues play an important roLe as evidenced by the observation of male Jul-odimorpha attempting to copulate with inanimate objects such as beer bottJ-es, which have the colour and surface texture of female elytra (Gywnne and Rentz, 1987). Besides the aesthetic value of the adults which have del-ighted natural-ists for more than a hundred and eighty years, and their intriguing variety of defense strategies, stigmotjerini may play an important biological role as pollinators. PotLen grains are often 4 found adhering to the hairy mouthparts, head and undersurface of the beetles. Despite the interest this group of beetl-es has stimulated, and the avid coll-ecting of specimens from the earLiest days of exploration to the present, the taxonomy of the tribe is poor]y understood. Before evolutionary, biological and ecological questions about the tribe can be addressed, its taxonomy must be cl-arified. Taxonomic historv gur knowl-edge of the AustraLian Stigmoderini can be conveniently divided into four historical periods in a way simil-ar to that used by Holm ( I97gzZ) for the Ethiopian Acmaeoderini-, because the same European entomologists were describing the two buprestid faunas: l. 1805-1860 2. 1860-1910 3. 1910-1960 4. L960-contemporary. Donovan (IBO5) described nine Austral-ian species of Buprestis L.: cancell-ata crenata, grandis, imperialis, Limbatat macuLaria suturaLis variabilis and unduLata. Eschscholtz (1529) erected the genus StÍqmodera into which he transferred three of these: grandj-st variabil-is and macul-aria and two South American Buprestis species: eximia and principalis. other important publications of this period include BoisduvaL (1875), the LaPorte and Gory Q8t7) monograph, and papers by Kirby, Hope, white, Parry, Gehin, Erichson, Boheman and Dejean. The writings of this first period usually consisted of very short descriptions in Latin, and although often accompanied by excel-l-ent figures, aîe quite inadequate for modern systematic studies. 5 The second period was dominated by MacLeay, the first Australian worker, and saunders, Thomson, Blackburn, and Kerremans. It was marked by a flurry of activity naming new species, without considering their I'Catalogus relationships. Saunders published the comprehensive (¿902-1906) Buprestidarumrr in 1871. Subsequently, Kerremans redescribed a great number of old species in his gigantic serj-es of monographs on the Buprestidae of the worfd (Hol-m, 1978). Regarding the other authors of this period, Carter (19]6) l-amented the difficulty of determining the value of their species, with the l-ack of published figures and the- unavailability of the types. He consj-dered the work of Thomson "...so casual-, brief, and unscientj-fic that the greater part is of little valuerr. The third period was dominated by carter, Théry, obenberger and Deuquet. Holm (1978) noted: t'Théry and 0benberger were bitter adversaries, and wasted much or inäir (and theii readers') time on continuous polemics and criticisms of each otherrs work. V'lithholding types and. information from each other, they very often both described the same species as new. As a heritage to their feud, they left the s-yitematics of Buprestidae in-a_deplorable state of ãÀ"or, wnión even after 20 years is still not fully sorted out.rl Since Carterrs I'Check List of the AustraLian Buprestidae" Q929), there have been no new publications dealing with the Australian buprestids as a whole, and only one study (Levey, I978a) of the Australian fauna at the suprageneric leve]. Recent revisions by Barker (1975, 1977, Ig79' 1980, 1983) of Astraeus L. & G. and Stiqmodera (castiarina) L. & G., and Levey (1978b) of Prospheres saunders have concentrated on the generic or subgenerJ-c l-evel' Stigmoderini was first named as a tribe by Kepemans (le97), which (1857). was based on stigmodérides, an earlier grouping of Lacordaire It is placed in a separate subfamily, stigmoderinae (Fowller, l-9I2; 6 Titlyard, 1926; Brues, MeJ-ander and carpenter, 1954; Britton, I97O; Bily, 1974), or in the Buprestinae (Théry, 1929; Cobos, 1980; Lawrence' I9B2; Matthews, 1985; BelÌamyr 1985)' Lacordaire defined Stigmodérides, comprising 6 genera, on the following suite of characters: antennal cavities smaIl, placed near the eyes, broadly cl-osed in front by the clypeus; poriferous fossettes inferior; head prolonged into a nuzz]:el scutel-lum rather large, fJ-at, heart-shaped or rounded; metasternum widely and strongly notched anteriorÌy, sternal- cavity formed al-most entirely by metasternum. Kerremans Qe%) reiterated these characters in his description of Stigmoderini which comprised 9 genera, however, he questioned Lacordai-rers use of the arrangement of antennal pores in forming a cLassification, noting that this character is variabl-e and difficult to determine. Kerremans ( I9O2) l-isted 12 tribes of Buprestidae in genealogical order starting with the most primitive. He placed Stigmoderini 10th, considering it to be advanced' 0n the other hand, Théry Q92g:274) thought Buprestinae (which incLuded Stigmoderini) to be a primitive group. In differentiating Buprestini from Stigmoderini, he found 'rthe characters indicated by Kerremans (the lateral- branches of the mesosternum elongate or very short) impracticablerr. Instead, Théry employed characters of the mouthparts: mouth prolonged into a muzzle, mandibles little bent, Iabrum and mentum longer than wide, and maxiJ-lary l-obes distinctive' I consider that the use of these characters as the sole basis for placement in stigmoderini is unsatisfactory. Mouthparts are not prolonged in the South American genera Dactylozodes, Hyperantha, and some pithiscus, or in some Castiarina. Clearly there is a need for a systematic revision of the tribe, together with a criticaL reassessment of its distinguishing characters. 7 As a base Iine, the 15 genera and subgenera considered in this study are those Listed by Obenberger (1934) and by his contempolaly Théry (1929, 1937). In chronological order they are: Stiqmodera Eschschol-tz, 1829 Conognatha Eschscholtz, 1829 ha Solier, L83t DactvLozodes ChevroLat, !8V7 CaLodema L. & G., 1837 Castiarina L. & G., 1877 Curis L. & G., 1837 Hyperantha Mannerheim, 1877 Metaxymorpha ParrY' 1848 Pithiscus Solier, 1849 JulodimorPha Thomson, 1878 Hvpos tiqmodera Blackburn, 1892 Aqrilozodes ThérY, 1927 Hyper anthel-La Hoscheck & ThérY, 1928 sri deroides Théry, !937 frlhat follows is a brief history of the generj-c or subgeneric status of each of the above taxa and a statement of how they are treated in this study. stiqmodera is at present compri-sed of three subgenera: Stiqmoder a( s. s.), Themognatha and Castiarina (Barker, 1979)' atha was erected as a genus by Sotier (1877), but together with Castiarina Curis, Conognatha, and Calodema was reduced to a division of (1857) Stiqmodera by LaPorte and Gory Q837). Lacordaire elevated curis and calodema to generic status but placed the other divisions together with Metaxymorpha in synonomy with StiEmodera. Matthews (1985) listed Stiqmodera. Th atha and Castiarina as genera. I Blackburn (].ASZ:. 2L5) erected the genus Hypos tiqmodera , based on a I'seems single new species H. variegata which he said to be a perfectJ-y typical Stigmodera in all respects except that the antennae of the male are strongly biflabelLate". Kerremans (1902) incl- uded Hyp ostiomodera as a subgenus of Stigmodera, but Carter Q93I2367) asserted that this species did deserve generic status. Only mounted fragments of one specimen of H. variegata were available untiL this thesis was almost completed, when three J-ive specimens were received: therefore, this taxon was not treated as a higher category during the study, although an assessment of its status is included in the discussion. Théry Ã937) erected the subgenus Stigmodera (sti deroides characterized by the female having a well--developed, al-ways visible' heavily scl-erotized anal tergite, prolonged in the form of a spout or beak. To this subgenus he transferred six species formerly placed in atha. Although the anal tergite is distinctive, I think it is an insufficient reason for considering this group of species separately from Themo tha. Pithiscus was erected as a genus by Solier (1849); synonymized with Stiqmodera by Lacordaire (1857); reinstated as a genus by Kerremans (1SOZ); and synonymized with Conognatha by Obenberger (1974). Hoscheck's (1934) revision of Conognatha identified four species groups. Groups I and II are considered together in this study as Conoqnatha (s. s.), because I perceive little difference between them. Group III corresponds to Pithiscus. Moore (1981) argued that Pithiscus shoul-d not have generic status because there are no substantial differences in genitalia between Conoqnatha ( s. s. ) and Pithiscus; but that it shoul-d be accorded subgeneric status because it differs from Conoqnatha (s. s.) in general morphology, elytral colouration and markings, and distribution. Moore did not provide a nettr subgeneric 9 description, because he considered (in litt. 1983) that the differences taxon noted by So1ier (1849) are adequate. It is treated as a separate in this study. Group IV is not considered in thls thesis because no specimens were obtained. Théry Q927) erected the genus Agrilozodes to accommodate three Perroud and species, Dactyl-ozodes ocularis (Kerremans), D. praecl-ara antennae Hyperantha pygmea Kerremans, primarily on the grounds that the were toothed from the 4th segment, not the 5th as in other Dactylozodes. Cobos Q959) considered Aqrilozodes to be one of five from the subgenera of DactyLozodes which were based on segments toothed 4th, 5th, 6th, or 7th segments. As I consider that this is not a of compelling reason for erecting subgenerar and because representatives only two of these gloups were available, Dactylozodes is treated as a single higher taxon in this study' Finally, Hypera nthelLa was created as a subgenus of Hyperantha by Hoscheck and Théry J928). The placement of Curis and JuLodimorpha in Stigmoderini is controversial. Curis was designated a division of Stigmodera by placed Laporte and Gory Q837). Genus curis was then successively in the tribes: Buprestides vrais by Lacordaire (1857); Anthaxinii by Kerremans :1897); Buprestini by Kerremans o9O2) ; Stigmoderini by Théry (1929); and Buprestini by Qbenberger (1934). The most recent worker to discuss its affinities is cobos (1978) who pJ-aced curis in Melanophilini, cLose to the monotypic chil-ean genus Anthaxioides' BelJ-amyts (1985) catalogue of the higher taxa of Buprestidaé listed Curis in Stigmoderini without explanation' The affinities of the monotypic Julodimorpha have presented a problem from its first description to the present. The species was originally described by white (1859a & b) as stigmodera bakewel-li. Thompson (1878) erected the genus Julodimorpha based on this species, 10 placed and placed it near stigmodera. subsequently, it was in: (J:9O2, l,916)i Acmaeoderini by Kerremans (1891); Polycestini by Kerremans (1934) Stigmoderini by Théry Q92Ð; and Acmaeoderini by Obenberger ' but Holm ( 1979) stated that it has features in common with Julodini' cobos that the whol-e morphology is based on the chalcophorini pattern. (1980) asserted that it constitutes an independent tribe which provisionally would be placed, together with stigmoderj-ni, in the subfamily Buprestinae, while BelÌamy (1985) placed it in Chalcophorinae. There have bee¡ no published descriptions of this tribe, or detailed studies of this species' Thus Hvpos tiomodera Aqril,ozodes and Stigmoderoides are not treated as higher categories in this thesis, but for ease of description and discussion, the remaining J-2 genera and subgenera listed by Obenberger (I9t4) and Théry oozl; 1937) are treated as independent taxa' Stiqmodera ( s. s.), Themognatha, Castiarina, Cal-odemat Guinea: Metaxymorpha, and Julodimorpha are endemic to Australia and New Hyperanthel-la are Co natha Pithiscus, Dactylozodes, Hyperantha, and (]928) and endemic to South America. Curis was reported by Carter al-1 the Théry i.]:92g) to have representatives in both continents, but species l-isted by 0benberger ( I9VOI566) are Australian, and Cobos Qglgz VB) referred to Curis as a southern, non-Ameri-can genus' The close similarity between the Australian and South American (1805) genera was lecognized by many of the earLiest workers' Donovan placed species from both continents in Budrestis' When Eschschol-tz split Buprestis, he placed some of the south American species in a separate genus, conognatha, but LaPorte and Gory Q877) and Fairmaire andGermain(1858)reunitedthemtoformdivisionswithinthesingle genu s Stiqmodera. 1I Backq round to karvoloqical studies Coleopteran cytogenetics was pioneered by Stevens in the first decade of the twentieth century, and two species of Buprestidsr un-named spruce borers, were among the earliest examined (Stevensr 1906). From the early I9AOts to the mid Ì970ts, the work of Stanl-ey G. Smith dominated the literature, and mole recently, Virkki, Yadav, Takenouchi, Suomal-ainen and others have added substantially to our knowledge in this field: the results of all these workers are coLlated in Srnith and Virkki's (1978) monograph which contains a gtobaJ- list of chromosome numbers oî 21160 species of Coleoptera. These include 27 species of buprestid, whose diploid chromosome number ranges from 12 (MeLanophila acuminata, Buprestini) to 26 (Sternocera l-aevigata and S. nitidicollist Julodini). Few cytological- studies of buprestids have been reported since that publication. An ancestral complement for beetl-es of 9 autosomal pairs + sex chromosomes, can be postulated with a high degree of probability, and persists in a very considerable percentage of extant species (Crowson, I9BLz4O2). The highest number recorded in normal bisexual CoJ-eoptera is 2n = 69t and the lowest is 2n = 4 (Virkki, 1984¿ 6t). In most Coleoptera, the male is heterogametÍc and sex-determining mechanisms reported include XY, Xy, XYO, X0, neo-XY and multiple sex chromosomes. XY represents the situation in which the Y approaches the X in si2e. In Xy systems both X and y are }arge but definitely unequal. The Xy^ association in which the y is very small, approaching P the lower limit of visibility is the commonest system in Polyphaga' occurring in practicaÌIy al-I families. In Buprestidae, it has been recorded in 10 of the 21 species reported (Smith and Virkki' 1978t244). Crowson (1981 zh1i) maintained that it is the most primitive condition, and is suspected to have been a feature of the ancestors of T2 the Endopterygota at the beginning of the Permian period' 0n the other hand, virkki (19e+:¿¿) stated that'rit is still an open question whether Xyo is the original sex chromosome system for alL beetl-es'r. In the Xy^ system, the X and y form a characteristic ring bivalent, which P frpatachuterfin stevens (1906) described as a which the X chromosome represents the parachute proper, and the y represents the l-oad' The symbol XyO was adopted by Smith (1950). Opinions about the mechanism of pairing of Xo and yo have oscillated between nucleolar and (Virkki' chiasmate modes and there is now evidence for both associations 1984257). X0, the second most common coleopteran sex chromosome system, represents the situation in which the maLe diploid complement has one Less chromosome than the femaLe. Probably, the y has been translocated onto the X or an autosome thereby reducing the number of chromosomes without loss of important genes (Wnite , !97t:I35) ' The neo-XY sex-determining system is one in which the Y was formerly autosomal and the X is a complex, one arm being homologous with one in the Y, the other comprising the essential part of the original X (Smi.tn I965azL62). It is most easily, but not necessarily derj-ved from an X0 with acrocentric chromosomes. The neo-XY system has been identified in six species of buprestid (smith and Virkki, L978|244). Multiple sex chromosomes have been reported in a number of coleopteran famifies including Dermestidae, Curculionidae, Bruchidae, (smith Tenebrionidae, coccinel-Iidae, cincindelidae and chrysomelidae and Virkki, Ig78). Although the mechanisms by which they arise are various, they are clearly identifiabl-e at meiosis as muLtival-ent associations at metaphase I, and inequality of numbers at metaphase il. vrlhite Q9772 679) suggested that their main adaptive significance is that they lead to fixation of a great deal of heterozygosity in the l7 heterogametic sex, and much genetic recomÞination within its sex, thus alJ-owing for 'sex-]inked' genetic material- in the homogametic a considerable amount of adaptive polymorphism' chromosomes were investigated in this study to determine whether karyological differences are useful in delineating the higher categories. The evolution of genera and subgenera can ultimately be traced back to a speciation event. chromosomal reallangements may result in speciation, or be important in isoLating incipient species' A particutar karyological change at the beginning..of a lineage may'be conserved in the descendants and provide evidence that a taxonomic grouping is natural. Aims of this study The aims of this study were threefold. Firstly, to delimit the tribe stigmoderini. Secondly, to examine and redescribe in full the previously described genera and subgenera of the Australian Stigmoderini; and to devise a key to the higher categories' Third1y, to establish relationships between the higher taxa using a phylogenetic approach. characters of internal anatomy and karyology, which have not been investigated in the tribe prior to this studyr wele employed as well- as a wide range of characters of external- morphology. In addition, new techniques such as scanning electron microscopy were used to cLarify characters such as the alrangement of the antennal sensillat a character frequently given high weight by earlier workers although difficult to see clearly with a light microscope' The study commenced with an examination of Stigmodera (s' s')' Subsequently, Themognatha' Castiarina Calodema, and Metaxymorpha were investigated. Representatives of the South American genera were al-so 14 the study of South examj-ned, but because of the lack of material, Anerican members of the tribe is incomplete' the two It became apparent from preJ'iminary studies of controversial genera, curis and Julodimorpha, that they should be presented in the excl-uded from the tribe, and arguments for this ale thesis. curis probably belongs in Melanophilini where it is placed by 0n the cobos (1978), and consequently it is not considered in detail' other hand, there is no consensus on where Jul-odimorpha should be placed, and therefore it is studied in depth' In the course of the study, most described species of catodema'and were dj'scerned' As Metaxymorpha were examined, and several new species keys to the specific revisions are outside the main aims of this thesis, redescri-ptions are speci-es of these genera, and their descriptions or presented in Appendices G and H respectively' AustraLian The tribe stigmoderini incLudes more than half the of the group buprestid fauna. This thesis contributes to the knowledge byaddingdataonaspectspreviouslyneglected,suchasfemale by genitalia, and by providing an overvi-ew of the tribe not attempted earlier workers. Furthermore, it uses investigative approaches and the taxonomy of the methods of analysis that have broader application to Buprestidae as a whol-e. il MATERIALS AND METHODS T5 I MATERIALS This study is based on adul-ts only: larvae are virtually unknown. The emergence of adults in spring and summer is brief and rather variable, not occurring at all in times of drought. Live specimens were collected from fl-owers by hand-picking beetles as they basked or fed, or by shaking fLowers and foLiage vigorously into a wide-mouthed net. Specimens were placed in individual containers with airholes and a few drops of moisture, and kept in the refrigerator at about 4 oC for up to six weeks. In addition, speci-mens were examined from the museums and private collections l-isted in Appendix A. 264 species of Stigmoderini were examj-ned externally; more than 400 specimens representing L72 species were dÍssected to investigate internal structures; and chromosome studies oî 35 species were undertaken (Appendix B). Voucher specimens of dissected material- will be lodged in the South Australian Museum. 2 KARYOLOGY Adult male specimens were processed as soon aS possibLe after col-lection. A cross-referencing system was used so that each chromosome preparation couLd be matched with the specimen from which it originated. This procedure wiLl remove any uncertainty about the assignation of a karyotype to a species, in the light of any future taxonomic revision. Several techniques were tried, but the most successful was based on that of Imai, Crozier and Taylor 0977) which treats the tissue with a hypotonic coLchicine solution before fixationt and uses Giemsa stain (Appendix C). Photomicrographs were taken on a Zeiss Photomicroscope ModeL III at magnification 400x, using a gleen filter and Agfa-Gevaert Copex Pan Rapid Tri 1l film' 16 Generally I to f karyotypes wele made for each species, and where more than one was made, there was usually good agreement, to the nearest percentage, between the rel,ative total chromosome lengths (TCL) and arms, as measured from the enlarged photographs. chromosomes are (sm), described as acrocentric (a), subacrocentric (sa), submetacentric or metacentric (m) following Petitpierre Q97I) as described in Smith and Virkii (1978), and karyotypes are assigned formul-ae following Smith (1e65b). 3 MORPHOLOGY and NOMENCLATURE Internal anatomy was investigated in freshly killed specimens set i-nto beeswax, and dissected under water, which was then replaced with 70% ethanol to harden and whiten the tissues. South American material was recei.ved freshty preserved by T. Moore as folLows: fixation for two hours in l-:7 :: glacial acetic: absotute ethanol, then storage in B0% ethanol to which a tittle 5% glycerol had been added. GenitaLic preparations were made by transferring the genitalia to l0;¿ cold KO[-l overnight, rinsing in distilled water, dissecting further if necessary, placing in 10% acetic acid for 5 minutes, rinsing again, and storing in 7O% ethanol. Illustrations were made with the aid of a camera lucida attached to a lVild M3 binocular stereoscopic microscope' ' Nomenclature foll-ows Crowson (1981) unless otherwj'se stated' Nomencl-ature of mouthparts follows Richards and Davies Q9n); nomenclature of antennae folLows Chapman QggZ). The problem of rrstriarl termi.noLogy of the elytral surface and in particular the use of as both a generic and descriptive term was discussed by Spilman (I97I). Erwin (og74) coined a new term "i.ntelneursl' for the longitudinal structures or lines between the intervals. Although not 17 widely adopted, I think this term has merit and Ís used in this thesis. Nomenclature of wing venation follows Good (1925) which ís based on Forbest Q922) studies of tracheation, basal sclerites and pupat pads, in preference to the nomenclature of Ponomarenko Q973), Cobos (l9BO) or Crowson (1981): Goodrs nomenclature has been followed by subsequent workers on Buprestidae e.g. Levey (I978b) and Nelson (1981) t which makes for easy comparison. Nomenclature of abdominal musculature follows Kasap and Crowson Í975); only pregenital segments are dÍscussed. As there is no consensus in the literature concerning the terminology and derivations of the components of female genitalia, these are treated more fully as part of the description in chapter III 2-5. 4 SCANNING ELECTRON MICROSCOPY specimens were cl-eaned by soaking overnight in 6o%:4o% :z chl-oroform: absolute ethanol foLlowed by one minute uLtrasound treatment in a Bransonic l-2 unit. Various techniques were used singly, or in combination, to overcome charging problems: dipping specimens in I0% Duron antistatic compound in absol-ute ethanol before mounting; mounting on double-sided conducting tape or generous amounts of silver adhesive; treating lor 24 hours with 0s0O vapour before coating; multiple coatings, both evaporative and sputter; and examining specimens at low voltages, down to 2.5 KV in an ETEC Autoscan scanning electron microscope operating in the secondary electron mode. The most successful method, howeverr was as follows: mounting c1ean, air-dried specimens in a smal-l drop of Aquadhere water-solubIe glue; coating with 15 nm carbon and 20 nn BO%/2O% gold/pal-Iadium alloy using a Denton Vacuum Model 502 high vacuum evaporator, fitted wÍth a rotating-tilting speci-men stage; and examining at 2O KV on the ETEC' adapted to operate in l-ow-vacuum mode (0.1 - 0.2 Torr in the specimen 18 chamber), using a Robinson wide-angle scintillator-photomultiplier backscattered-electron detector. ChargÍng artefacts are not observed with backscattered-eLectron l-ow-vacuum mode, also called the environmental cell mode, a technique developed by Robinson (tZlS, Ig76). Although uncoated specimens can be examined, the image is improved in topographic resolution by coating with a high atomic-number element such as gold (Robinson, 1980). The disadvantage of this method is the lower resolution obtainable at high magnification using a back-scattered detector. Photomicrographs were obtained using Kodak FXP 120 Panatomic-X film. 5 HISTOLOGY The accessory gland was dissected from adul-t females, fixed in l0% formalin, dehydrated in ethanol, cleared in xylene, embedded in oC), Paraplast (mp Sø and sectioned at I Um. The sections ¡¡ele oC, floated onto slides, dried overnight at 47 stained with Mayer's HaematoxyLin{osin, Alcian Bl-ue-Chloratine Fast Red or Polychrome Methylene Blue, and mounted under coverslips with Depex. Slides were examined using a Zeiss Photomicroscope III equipped with an epi-fluorescence condenser system with exciter filter BP 485/20 and Barrj.er fil-ter BP 520-560. Chitin was demonstrated by its birefringence in polarized light, autofluorescence, and failure to break down in 10% KOH overnight. Photomicrographs were taken using Ilford FP4 film. Symbionts were investigated as foLlows. In a laminated airfl-ow cabinet steriLized with UV Ìight for hal-f an hour, the accessory gland was dissected from live specimens. The contents of the lumen were transferred to an agar plate using a fine probe and cultured under microphilic conditions. 19 6 TAXONOMIC CHARACTERS The definition of 'taxonomic characterr adopted in this study is "any attribute of a member of a taxon by which it diffe¡s or may differ from a rnember of a different taxonfl Mayr (19692t2l-). A character in this sense is not an abstraction with different states, as used by the numerical taxonomists; and in the descriptions and analyses in the following chapters, the terms ciraracter and character state are used interchangeably. Schl-ee 0975) demanded that characters used be of principally (not gradually) Oiffering complex structure, but as Brundin Qye:141) observed: "This is the ideal case we are looking for, but rarely meet!" (Saether 19792306). This was found to be especÍally true in the Stigmoderini in which most of the taxonomic characters are simple structures varying only slightly in shape, síze or arrangement. 7 PHYLOGENETIC Ì"ETHODS Wiley (1981 z24O) identified four basic approaches to taxonomy: "traditionalf', phenetic, evolutionary and phylogenetic. In this studyt a phylogenetic (or cladistic) approach based on the principles of Hennig (1966) and elaborated by Arnold (1981), Wiley (1981) and others' was adopted. Phylogenetics is concerned with the discovery of genealogical relatj-onships and with constructing a classification which reflects these rel-ationships. The size of the morphological gap, species richness and other criteria used by evolutionary taxonomists are rejected. The main tenet of this approach is that the designation of a taxon as monophyletic depends on establishing autapomorphies (unique derived characters), and the discovery of genealogical relationships within and between monophyletic taxa depends on establishing a nested pattern of synapomorphies (shared derived characters). It is stressed that the 20 terms apomorphic and plesiomorphic are relative' Since only apomorphic characters are used in the anarysis, the application of this approach of hinges on being able to determine the rel-ative evolutionary polarity characters. stevens (I9go) listed four main groups of criteria used to assign potarity: ontogeny, palaeontology, character state distribution, data on and notions of how evolution tends to proceed' There are no the ontogeny of stigmoderini, and scant information on the fossil record, so these criteria are not discussed further. There are two methods of determining polarity by character state distribution: in-group and out-group analysis. Estabrook (1977) was a proponent of in-group analysis in which a character is considered plesiomorphic if it is widely distributed within a taxon. If the this apomorphy arose early in the evoLution of that taxon, however, assumption, sometimes called the commonality principle, Ís not valid' It is sure to give misleading resul-ts in a very significant proportion at of cases (Arnotd,1981), and consequently, has fal]-en into disrepute' least as a primary indicator of polarity' The method of determÍning polarity most widely accepted and adopted here is out-group analysis in whích a character is considered relatively plesiomorphic if it is distributed over a wide spectrum of related taxa, and apomorphic if it is found only within the taxon under consideration the and not in the out-group. Ideally, comparisons should be made with sister group and with at least one taxon that is cladisticaj-Iy more primitive (Stevensr 1980). This plesupposes that the next higher levels of classification have been resolved, which is not true in Buprestidae, where the higher taxa are in a state of flux' Even when out-group rel-ationships are uncertain, however, a well resolved Maddison' cladogram can sometimes be constructed (Maddison, Donoghue and 19811) . 2I In this study, the out-group used in comparisons is a broad selection of other buprestids at hand, or reported in the literature, for which the appropriate data are avaiÌable. The only serious drawback of this method is if the taxon under study is the cladistically basat group of the higher taxon used as the generalized out-group (Stevens, 1980). When a phylogeny of the Buprestidae as a whole is developed, resolution of sister groups wil-l enable generic relationships within Stigmoderini to be more rigorously analyzed. Where out-group analysis failed to indicate polarityr the l-ess satisfactory fourth criterion, based on ideas about the way evolution proceeds, is sometimes employed in this study. Examples of such notÍons are that evolution is often associated with nr-¡merical- reduction by loss or fusj.on; elaboration or differentiation of structures; i.ncrease in organization or complexity; or ecological or morphological specialization. Such concepts should always be applied with caution. Even after the polarity of characters has been discerned, there are other problems with the application of Hennigian principles. One of these is how to interpret synapomorphies which suggest conflicting relationships i.e. how to distinguish between shared characters which are due to recency of common ancestry from those due to homoplasy (convergence, paralleIism, or reversal). The principle of parsimonyt i.e. choosing the hypothesis that requires the fewest ad hoc statements (Farris, I97V) is the usual way to arbitrate between competing hypotheses. It is adopted Ín this thesis, while bearing in mind that parsimony is only a method of analysis, not an evoLutionary plocess, and should only be applied after consideration has been given to character weighting (Hecht, 1976). V'leighting is a process of estimating the rel-ative rel-iabil-ity of a character by assessÍng the likelihood that the presumed synapomorphic character is in fact homologous, has been rightly 22 identified in the taxa concerned, and has been correctly assigned polarity (Arnold, 198I). If any of those components are doubtful, a low weight must be assigned to the character. In addition, mosaic evolution, in which the derived character is possessed by only some of the descendants of a lineage, is a common occurrence, and complicates the interpretation. All these difficulties are discussed in more detail- as each is encountered in my phylogenetic analysis in Chapter IV 3. 8 CRITERIA FOR SUPRA-SPECIFIC TAXA Natural supraspecific taxa are historical units resulting from speciation events, and are discovered and justified by rrsynapomorphies employed at their correct level of universalityr' (Witey, 1981: 75176). The purpose of placing species in supraspecific taxa is not only to indÍcate relationships, but also to facilÍtate information retrieval. Assignation of a species to a genus is most important because, as part of the scientific name, the affinities of the species are immedÍately indicated. For this reason, I agree with Mayr (Igøgz 279) that "sptitting is particularly deleterious on the generic level|r. Specialist systematists, however, often elevate supraspecific taxa, partly because they "have an inherent tendancy to see their special groups out of perspective in reLation to othersrr, and partly because "they almost invariabLy find themselves wishing to use more categories than the general system allows themrr (Crowson, I97Ozað). In my vÍew, even when substantial morphological differences between groups of species can be discerned, it is more important to indicate affinities and to maintain nomencl-atural stability, than to highlight those 2V differences or to create more tmanageablef sized groups. The only valid reason for splitting existing genera is a cl-ear case of polyphyly or paraphylY. 24 1 KARYOLOGY The material examined had a low mitotÍc index. Often only one or two specimens of a species were availabJ-e, so the following comments are based on limited observations. Neverthel-ess, it is possible to draw some tentative conclusions even from the examination of a single individual (DYer, L979274). chromosome counts were obtained for 75 species of stigmodera: 5 Stiqmodera ( s. s.), 17 Themognatha, and lf castiarina (see Appendix B); and a tentative_ count was obtained for Dactylozodes rouletÍ. 0f theset 34 have a complement of l¡ = 22i two species, S. (T. ) nickerli and species s. (r. ) alternata have 2n = 2O. Karyotypes were made of 17 (Figures 1-3) and the morphology of these is summarÍzed in Table l' All Stigmoderini examined have an XYO sex-determining mechanism (Figures 4A-J). preparations were also made of representatives of two other genera previously placed in Stigmoderini. Cel1s at metaphase I of meiosis of the three species of Curis studied indicate a diploid complement of 14t with a very small X, showing positively heteropynotic at pachytene in Figure 4K. Six cells of Julodimorpha bakewelli examined have 39 chromosomes (Figure 4L), three cells have one or tlvo chromosomes fewer. They appear to lack any mínute chromosome lepresenting a suggests that Julodimorpha has an X0 'py_. The uneven diploid number sex-determining mechanism or muttipLe sex chromosomes. Unfortunatelyt no meiotic ceLls were found which might have confirmed one or other of these al-ternatives. The predominant feature of the JulodimorPha karyotype is that autosome 1 is very large' mole than double the length of succeeding autosomes. Another feature, not apparent from the ceLl in Figure 4L, but observable in other ce]ls, is that many of the smal-l chromosomes are acrocentric. FIGURE 1. KarYotYPes. (A) Stiqmodera ( S. ) porosa (B) I. (r.) tricol-orata (c) S . (r.) viridicincta (D) S . (r.) donovani Arrows indicate NOR, a1t figures at same scale' A lÍ It tt Iß TT tt lr lß II t 7 8 9 10 xy 2 3 4 5 6 5 rrm e B I ft It ll lt ll ¡t ¡rll ilû c (( (( !( ll rD l¡ ti lf )l l( 'I D FIGURE 2. KarYotYPes. (A) Stiqmodera (T.) (B) I. (r. ) heros (c) I . (f.) nickerli (D) g. (f. ) alternata Arrows indicate N0R, all figures at same scale' I 9 10 Xy 4 5 6 7 1 2 3 5rm o B tl ¡I l¡ tr tl I l' c íI lt tt ll ll tt lrtl ìr ls t D tr lI l¡ II lr lt rr¡r ,\ FIGURE ,. KarYotYPes. (A) . ) macul-aria (B) S . (s.) cancel-]ata (c) 9. (S. ) roei (D) !.. (T. ) variabifis (E) S. (T. ) chevrolati (F) s .(r . ) barbiventris (G) q. (I.) mitchelli All figures at same scale. A lf lr f I ¡f l3 ll l, rt l' l¡ u lr r¡ t' B l(U ll Il ll Ir 9 'lO 2 3 4 5 6 7 Xy sPm 3I r¡ r¡ ¡¡ c t( 13 ¡l l¡ ¡s tr I' o D fl llll tl tt ¡lll r¡,r h 0 lt¡ oa aa SSStl¡rr¡ E lf ¡¡ ¡¡ It .¡ o. ., F It !l tt 3t 13 St ¡l ¡l ¡t G ll lr fI t¡It tl l¡ Il 3. tr .' metaphase I FIGURE 4. Meiotic metaphase II (A, B, C); meiotic (D, E, F, Ir J); spermatogoniaL metaphase (G, H' L); and pachytene (K). (LOA X) (A' B ) Stiqmodera regia + Y, l0A + (c) S . sexplagiata (104 + X) (1044 XY (D) S. arqj.l-lacea + p ) (1044 XY (E) 9. oratiosa + p) (F) S . adelaidae (1044 + XY p) (c) 9. f icta (20A + XY) (H) S. decemmaculata (204 +Xy) (r) S . cupreoflava (104A + XVO) (J) Dactvl-ozodes roul-eti (IOAA + XVO) (K) Curis auri-fera (6RR + heteropycnotic X, y not visible) (L) Julodimoroha bakewelli (l8A + X0) XY figures at same scale. Arrowheads indicate p , all -n I C- ¡ o o n )o E ô o (o +.*T t r' o- ('I fl.o t 0 q -J I df TD a ã x T o a t a ? a -I¡-a v a I \ -\ ç I v rn ì rrf '+*ï t 4.v ta CÒ ür+ ID \ r Þ ì o oJ o Þ t t ¡ J T o a ¡ I !\ t--f Ù' 12 a ct t I G' o "lt, TABLE 1 Karyology of 17 sPecies of Stiqmodera metacentric, A = autosomes; superscripts-lo""tocäntrið m, sffir sâr a, represent iuomàtacentrió, and acrocentric respectivg]y; :' M; L smal-L, medium and large size of X chromosome relative to = as a % of totaÌ autosomes; At;% =-ig;gah of aútosome 1 expressed ðniórororå 1ength (TõL); NgR = autosome number on which NgR occurs, where several autosomes are the same lengtli so that the exact order cannot be determined, the number is given as a group' AI% NOR TAXON FORMULA X q. (Stigmodera) cancel-lata 10 Am + Xmyp S t7 qratiosa l0A *XYp S l3 Xsmy L ]8 macularia 10 A * p L l8 7-8 l-o Am * Xsay p Xsmy L L6 l-o Am * p q. (Themognatha) s228 al-ternata 9Am+Xmyp XaY s 15 barbiventris L0 Am * p chevrolati 7Am+7Asm+XsmYp S I7 6-8 donovani 10 A + Xsmyp S 14 10 Am + Xsmyp S T7 8 heros Lo Am + Xsayp L 15 mitcheLli 9 Am + l- Asa + XaYp S 12 S 27 ç9 nickerli 8Am+1Aa *XsmY p regia 9Am+1Aa *XYp M 12 tricolorata 9Am+lAsm+xsayp s T3 7-8 variabilis 8Am+2Asm+XaYp S I7 7-9 viridicincta BAm+2Aa *¡ayp S 15 25 considerable interspecific and intraspecific differences in karyotype size were observed' TCL = 68 Llm to 160 ¡rm' For example, the S. nickl-eri ceLl ill-ustrated (Figure 2C) has chromosomes nearly twice the length and more than double the width of those from another specimen of the same species. The size of chromosomes may be influenced by genetic constitution (incLudÍng intraindividual variation of heterochromatin), phase, nutrition, temperature and prepalative methods. Moreover, synchronous multipl-icative divisions such as gonial mitoses probably always result in a decrease in cell size, mitosis by mitosis, with concomitant increased condensation (Smith and Virkkit l978z2L). V'/ithin the karyotypes there is typically a gradual decrease in size of the autosomes. Abrupt size differences have been reported in other buPrestids: in Sternocera nitidicollis S. laevigata and Julodis whithilli, autosome 1 was 5-6 x the length of the average of the remainder (Smith and Virkki, 1978:25). In Stigmodera, the sex chromosomes appear positively heteropycnotic at zygotene and negatively heteropycnotic at metaphase I of meiosis' In keeping with the interpretation of the sex chromosomes of other beetles, the minute chromosome is presumed to be the y, although this was not confirmed by examining female tissue' Autosomes are predominantly metacentric, but some of the karyotypes analyzed have acrocentrics e.g. s. nickerli (autosome 9), g. viridicincta (autosomes 7 and l0), or submetacentrics e.g. s. tricolorata (autosome 10). changes'in'relative arm length may be due to pericentric inversion, changes in heterochromatin, or simple reciprocal transl-ocation (Imai, crozier and Taylor, 1977). Karyotype variations due to pericentric inversions in congenerj-c species are known in many genera of beettes (Yadav and Pil1ai, 1979) ' 26 0f eight species for which the karyotypes were analyzed in more detail-, six (!. nickerli, S. porosa, S. tricolorata S. goryi, S. alternata, and S. donovani have an autosomal pair with a nucl-eol-ar organizer region (NOR) (Figures I & 2). l'/hen present, it is usually on autosome 8, or possibly one of the adjacent chromosomes which are often so close in J-ength that it is difficult to order them exactly. The position of the nucl-eolar organizer region is either medial as in S. porosa, or subterminal as in S. nickerli and S. alternata. In the latter cases, the distal ends of the arms form satellites. 2 MORPHOLOGY The purpose of this section is to describe the morphology of Stigmoderini, examinÍng diagnostic characters used by previous workers, and presenting new characters especially those of the internal anatomy which has not been studied before. In the Discussion section which follows this chapter, the value of these characters in delimiting the higher categories and elucídating the relationships between them is assessed, and where possible, the polarities of the character states are identified. This provides the basis for the phylogenetic analysis. 2.I GENERAL FORM AND COLOURATION OF ADULT The Þody is elongate or oval and dorsoventrally fl-attened. Body length ranges from small to large (6-55 mm). 0n average, females are larger than mal-es, but usually there is some overlap in size ranges. Stiqmoder a cancellata is an exception in having the female much larger than the mal-e with no overlap in size. Colouration and pattern are very variable within the tribe. Most species have metallic reflections on head, legs, pronotum, undersurface and elytral fasciae, and many 27 Dactylozodes is specj-es are brightly coloured. Pilosity is variable' characterizedbylongdensesetaerparticularlyontheelytraand pronotum. 2.2 I-EAD dense (greater Punctuation is moderate to finer sparse, moderate or In than, equal to, or less than one diameter apart, respectively)' concave and the vertex has most taxa the frons is almost fLat or weakly a plonounced medÍaI a fine medial impressed Line. some castiarina have gEooveonthevertex,orfronsstronglyconcave.Fronshairis variabl-e. The frons is not narrowed at the level- of the antennal of the eyes cavities, which are smaIl, placed 1aterally near the border and/or depression' and clypeus, and each surmounted by a smooth ridge l-ie in a groove between In some species, the first few antennal segments and poorly clypeus and eye, but this character is highly variable defined. (maximum width of both Eyes. Eyes are usually medium to large eyestogethergreaterthanorequaltotheminimuminterocular vertex (Figures distance), oval, subparalleL or converging stightly on have smaller eyesr more 5C-D, 104,C): some Dactvlozodes and Hyperantha widely separated (Figure 108) ' clypeus is longt Mouthoar ts and cl-ypeus. In most species the prolonged into a rostrum weakly to strongty bisinuate, the mouthparts brush-like ' (Figure 5B,D), and the galea round or oval with dense, long, have clYPeus and setae (Figures 6E-G). Dactvlozodes and Hvperantha setae (Figures mouthparts short, and galea oblong, with sparse, short (Figures 5C' 5Dt 5A,6D). Both types occul in Pithiscus and castiarina 6, 6H). (A with FIGURE 5. Comparison of abbreviated mouthparts a c) (E) prolonged mouthparts (a a o); and mandible lacking teeth with strongly toothed mandible (F) ' (A-B) Head, lateraL view with antennae removed (A) Dacty lozodes cupricol-Iis (B) Stiqmodera ( T. ) nickerli (c-D) Head, anterior view (c) Stigmodera (c. ) cupreoflava (D) Stiomodera (C.) subtincta (E-F) Left mandible, ventral- view (E) ÆStigrìtqqqra (T. ) gorYi (F) Hvperantha decorata Scale bars = I mm (A, B); 0.5 mm (C, D, E, F). B # f I E F l-l FIGURE 6. (A - D) Ventral view of mouthparts il]ustrating the attachment of prementum to mentum; (E - H) detail of maxillae, (E - G) nectar-feeding species with dense setae on l_acinia and large galea which exceeds maxillary palps, (H) l-eaf and petal chewer with sparser setae on ]acinia and small- galea which does not exceed maxillary palps. (A) Stismode ra (S.) sanguinosa - attachment anterior without groove (B) Stigmodera ( C.) thomsoni - attachment anterior with groove separating prementum and mentum (C) Stigmodera (T.) nickeri - attachment dorsal, prementum long (D) Dacty l-ozodes con.juncta - attachment dorsal, prementum short, completely concealed; mouthparts abbreviated (E) Stiqmodera ( 9. ) porosa (F) Stiqmodera (C. ) thomsoni (c) Stigmodera (r . ) parvicoLlis (H) Stigmodera ( c . ) cupreofl-ava Abbreviations: g = galea, I = lacinear l-i = Iigula, ]p = fabial palpsr Is = labial suturer l1 = mentum, ftìâ = frìârìdib]e, mp = maxiJ-J-ary palps¡ P9 = palpigert prn = prementum. Scale bars = 0.5 mm (A' C); 0.25 mm (8, D, E, G); 0.1 mm (F, H). A B ; , l-.¿rr t-4 I ', t rl I c D 'j:nè; : -'-- -.- F E , \ ù 1 G H 28 presence The l-abrum is variable in shape, degree of sclerotization' groove' The or absence of translucent border and medÍal longitudinal (Figures 5E-F) mandibles vary in shape and number of incisors ' segme¡ts' the The maxillary palps are comprised of four cylindrical (except in last truncate, rounded, or pointed; setae usually short 2 and 4 longest' 7 Dactvlozodes [Figure 6D]). Segment I always smaLL' short, equal to I or intermediate' pitted The labium is comprised of a heavily scterotized, hairy, prementum; the line of mentum, and a translucent' flexible, retractabl-e attachment of division between the two being the l-abial suture' The deep fissure (Figures the prementum to the mentum is dorsal forming a shallow groove (Figure 6C-D); more anterior so that the junction forms a 69);oralongtheanteriormarginsothatthelabialsutureis the mentum (Figure obl-iterated and the prementum is in the same plane as 6A).Articulatedtotheprementumaretheothercomponentsofthe palp of three labium: a pair of lateral palpigers each bearing a labial The relative segments, and a medial setose ligula rounded apically' genera' The last lengths of the palpal segments are variaþte within digitiform segment of both labial and maxillary palp bears short sensiLla in an apical depression' and strongly' The shape of the mentum varies within the tribe, is transverserL/Wlessthan0'5rinPithiscus'gtigmodera' more elongate, L/W Hvperantha ( s. s.), Hyperanthel-la, and Da lozodes conognatha (s' s')' some more than 0.6, in calodema, Metaxymorpha, and variable in this taxa e.g. T tha and Castiarina are very are constant' character, other genera e.g. cal-odema and Metaxymorpha havlng a pair of The mentum of most Dactvlozodes is distinctive ín l-ateral- notches in the anterior margin' 29 Antennae. Antennae are serrate, l]-segmented, with length less than or approximately equal to the width of the head. The scape is elongate, usually more than 2x length of the pedicel, and constricted at the base: the other untoothed segments are short, cylindrical or obconic, with the pedicel shorter than the meriston. The antennae are toothed from the 7rd, 4th, 5th, 6th or 7th segment' The shape of the toothed segments is variabte (Figures 7A-I), generally broadly rounded, widest behind the apex then constricted towards the base. The terminal segment is usually club-shaped' 4' Antennae of stígmodera (s. s. ) are weakly toothed, especially segment except in s. gratiosa. Those of Hyperantha have very strongly transverse rounded teeth. The other genera are intermediate between these extremes. Dactylozodes have transverse, triangul-ar toothed segments. Toothed segments have peg sensill-a concentrated in a deep subterminal fovea on the ventral surface of the each tooth and also in what I have called a rrsecondary Sensory arearr along the edge of the tooth adjacent to the fovea (Figures 8A-D). segment 1I has a subterminal- fovea on each face in stigmodera (s. s.), Themognatha, The CaLodema. Meta some Castiarina and some Pithiscus. sensilla are referred to as tporest in the literature although no pores are visibl-e even under 8000x magnification. The limited number of that specimens I examined using scanning el-ectron microscopy showed sensill-a are genelally more numerous (up to 200 per segment) in' Themognatha, calodema, Metaxymorpha, and Hyperantha (s. s.) than in sensil-la Stiqmodera (s. s.) or Dactylozodes (about 100 per segment). of the secondary sensoly area a1e either stout pegs with rounded apicest or narrow spine-like structures; those in the lovea are hollow and FIGURE 7. Ventral- view of right antenna, ilJ-ustrating the variation in shape of the segments between representativps of different genera and subgenera. Setae not shown. (A) . ) macul-aria (B) Stigmodera (T. ) variabil-is (c) . ) erubescens (D) Calodema regaLis (E) Metaxymorpha grayi (F) Conoqnatha (ç.. ) pretioÍssima (c) tha (P . ) viridiventri.s (H) Dactvi-ozodes altern ans (r) Hyperantha decorata (J) Hypostigmodera variegata Sca]e bars = 0.5 mm. A B c D E I F G H I J FIGURE 8. Antennal segments, ventral vj'ew' (A) Stiqmodera ( S.) macularia, segment 7 (B) CaLodema regalis, segment 7 (c) Hyperantha decorata, segment 7 (D) Dacty lozodes cuPricollis, segment 6 (E) Calodema regalis, fovea segment 5 (F) Cal-odema regali-s, detail of sensilla in fovea (F)' Scale bars = 0.1- mm (A - D); O-o2 mm (E); 0'005 mm A '"*-Z .\ '\ \ \ q D \ J ì \ \ år p L -¡,t { I t vo conical, with apices acute (Figures 8E-F). In additionr there are usually V or 4 grouped, l-ow-domed sensilla. The antennae of Hypostigmodera variegata are distinct from those of other Stigmoderini in having sensill-a diffuse and dense over most of the surface, not concentrated in foveae and secondary sensory area. The shape of the segments is not atypical in the female, but in the male (Figure 7J), the meriston is strongly toothed, segments 4 to 10 are bipectinate, and segment 11 is tripectinate. 2.' THORAX Pronotum. The pronotum is wider than long (Figures 9A-H). Metaxymorpha and Conognatha (s. s.) are strongly tapered from base to apex (Figures 9E-F): some Calodema and some Castiarina are widest medial-ly with sides convex, diverging from base (Figure 9D). Remaining taxa are sub-parallel on basal half, then incurved towards the apex. ScuJ-pturing is present in Dact l-ozodes some Castiarina particularly lycid mimicking species (S. Barker, pers. comm. ), and some Pithiscus in the form of basal depressions medially and laterally. A pair of smalL basaL notches which may or may not receive projections from the anterior margin of elytra is characteristic of many Themognatha and Castiarina. Conognatha (s. s.) and some Metaxymorpha are entirely smooth and shiny, other taxa show varying sizes and densities of punctuation. The lateral margins are incomplete with sides rounded anteriorly in Conognatha ( s. s.), Hyperanthella, Castiarina and some Dactylozodes; others are moderately or strongly carinate (Figures 104-C). The anterior margi.n is about half the width of posterior in most, with a fine impressed line behind it; weakly bisinuate or straight in most, more strongly bisinuate in Calodema. The posterÍor margin is FIGURE 9. Diversity of pronotal shapes and pronoto-elytral articulations (A - H); and scutellaJ- shapes (I - M). (A) Stigmodera (S. ) macul-aria (B) Stigmodera ( I. ) princeps (c) CaLodema regalis (D) Calodema ribbei (E) Metaxymorpha apicalis (F) Conoqnatha pretioissima (c) Dactylozodes conjuncta (H) Hyperantha sall-ei (r) Stigmodera (S. ) jacquinoti (J) .) mniszechi (K) Stigmodera (C.) andersoni (L) Stigmodera (C.) bremei (M) Calodema regaLis Scate bars = 2.0 mm (A - F); 1.0 mm (G - H); 0.5 mm (I, J, M); o.25 nm (K, L). A B c D E F G H M I J K L FIGURE 10. Characters of the thorax. (A - C) Anterior view of head and prothorax ilJ-ustrating different i-ateral margins. (A) Stigmodera ( S.) macularia, carinate. (B) Hyperantha salLei, globose anteriorly. (c) Stigmodera (!.) gratiosa, explanate. (D - E) Ventral view of mid-thorax ill-ustrating intercoxal process and sternal cavity. (D) Stigmodera (C.) adel-aidae, sternal cavity strongty transverse, intercoxal process with medial- lobe wider than two l-ateral Lobes together, and meso-metasternal suture straight, halfway along the sternaL cavity. (E) Stigmodera (f. ) heros, intercoxal process with medial- Lobe not wider than two l-ateral- l-obes together, and meso-metasternal- suture oblique and basal. The most common arrangement, illustrated in Figure 11 (8, D, F), is intercoxal process with medial l-obe not wider than two l-ateral- Lobes together, and meso-metasternal suture oblique, approximateJ.y halfway along sternal- cavity. scal_e bars = 2 mm (A), I mm (e, c, E)r 0.25 mm (D). A l1 B 1î c 1( D E lntercoxal process meso- meta- sternal suture 7t (s. s. Hyperanthell-a, denticulate in D des. Hyperantha ), many Castiarina' Conoqnatha (s. s.), some Pithiscus, and bisinuate to straight in The pronoto-elytral articulation is weakly (s. and strongly bisinuate most; moderately bisinuate in Conognatha s'); (s. s.)' with mediat robe pronounced in calodema and Hyperantha (except some posterior lateral angles acute, Very acute in some, Calodema in which they are right angled) ' weakly convex Prosternum. The prosternum is variable anteriorly: Dactvlozodes e.g. Stigmodera (s. s.), Castiarina (s. s.); with rounded hump Pithiscus Hvper anthel-1a . and some Hvoerantha e.g.someConognatha(s.g.),Metaxymorphameekigroup;orwithanacute apicalis groupr some anterior projection e.g. calodema, Metaxymorpha (Figures lfA-F)' Conoqnatha (s. s.) and some Hyperantha prosternum trilobed and fits The broad intercoxal process of the is and closely into the sternal cavity formed by the mesosternum retasternum. The mesosternum is divided, and the meso-metasternal along the sternal cavity suture is straight, arched, or oblique, halfway Pithiscus and in most, but croser to the base in some Hyperantha, The metasternum borderÍng the Th (Figures I-OD-E' ltB,D'F)' cavityisraisedinsomeCastiarina,notablyinsomelycidmimicking species (S. Barker, Pers' comm')' but usually large Scutell-um is variable in size within the tribe, in most' l-lvoerantha (s. s.), and Hyperanthella, medium (Figures 9A-H) The shape is smallest in Themo tha and Calodema ' Metaxymorpha' rounded subpentagonal in Stigmodera (s' s')' ( s. HyperanthelIa, and Conoqnatha (s. s. ), Pithiscus Hvperantha s. ), in Castiarina rounded Dactv lozodes; tuLipiform, scutiform or cordiform Themoqnatha triangularrlunaterorcordateinCalodema;andvariablein is (Figures 9I-M). The scutel-lum of some calodema and DactyLozodes (Ar FIGURE 11. Representative prosternal types in lateral C, E) and ventral (8, D, F) views. (A, B) Stiqmodera (S. ) macul-aria , prosternum normal. (c, D) Metaxymorpha hauseri, prosternum enlarged forming a rounded hump anteriorlY. (E, F) Calodema ribbei, prosternum with pronounced conical projection anteriorlY. ScaLebars=2mm. I I I I I A B c D E F t2 strongly transverse. In general, the scutellum is not overLapped by the pronotum, except in some Calodema. Elytra. Stigmoderini basically have 10 interneurs plus a scutellary striole, and 11 major intervals, the last forming the margin of the elytron (Figures f2A-C). In Stiomodera'(s. s.) each interneur is comprised of a single row of large foveoles. The other Stigmoderini have finely seriate-punctate, punctate-striate, or striate interneurs. Fusion or reduction of interneurs occurs to varying degrees. In many species, a translucent spot below the puncture makes it appear larger and deeper than it actually is. The intervals are flatr convexr or alternately costate as in Hyperantha (s. s.), Hyperanthellat Dact lozodes some Castiarina and some atha. Some species have additional serÍal rows of punctures e.g. some Ca]oderna and Metaxymorphat or are scattered with punctures giving a rugose appealance e.9' many T tha. The sides of the elytra are generally parallel on the anterior half, then arcuately narrowed posteriorly. In Metax the sides diverge strongly from the base to the shoulders continuing the line of the pronotum, then narrow apically (Figure 9E). Hyperantha are straight-sided along entire length, Hvperantha (s. s.) strongly tapered, HvperantheLla para11e1. Apices are denticulate in Conognatha (s. s.), some Pithiscus, Hyperantha (s. s.), and some Dactylozodes; bi- or trispinose in ha and Calodema; and variably truncate, rounded, pointed bispÍnose or trispinose in the rest (Figures 1æ-I). The elytral apices are sexually dimorphic in stigmodera sanguinosa (carter, 1916) and Metaxymorpha The lobe at the base of the epipleura may be absent e'g' Hvperantha (s. s.), very weak e.g. Stigmodera (s. s.), moderate or FIGURE 12. Right elytra ill-ustrating different size and density of elytral punctuations (A - C), and variation in elytral apices (D - I). (A) Stismo dera (S.) macuLaria with few, very Ìarge, confluent foveol-es. (B) Stismodera ( T.) barbiventris, a typical- punctate-striate elytron, with punctuations smal-ler and more numerous than (A). (C) Ca]odema regaLis, with very fine, very numerous punctuations, and additional- punctuations between primary interneurs. (D) Stigmodera (S.) sanguinosa male, pointed. (E) Stiqmodera (S.) sanguinosa femal-e, bispinose. (F) Metaxymorpha apical-is, trispinose. (c) Stigmodera (!. ) roei, rounded. (H) Stigmodera (S.) porosa, truncate. (r) Conognatha pretioissima , dentate. Scal-e bars = 2 mm (A - c); I mm (D - I, all- at same scale). OOo c> Þoo o D U NO D o \\ o og O o o c c Oo ñ o L) o D I m - oo 0 o oooo0oo oOOooOOooooo 000 o oooooooooooo oo OooOOOOO o ooo OO oo o o o oooo 000 ooo o oooooo oooooooOOOOç ooooo o o oooOoo oooO ¿ o oo TD oo oooooooooogOoqg o[ooo ooooooooooo oo osoo ooooo I OO o o o oooo o 'Tt o . ' ' ï'..'.3 ..o oo o "'. .ii:'.."i o ".:" " -o ao' oå o I I oo v3 the strong. Metaxymorpha has one elytron overlapping the other towards the apices: in 8v% o1 75 M. glori-osa examined, the left overlapped abdominal right. In Metaxvmorpha . the elytra extend well past the last segment. In other taxa this character is variable between species or (C') less pronounced: it occurs markedly in members of Stigmodera producta mimicry group. The elytra of Dactylozodes have long, dense pitosity: Hyperantha have short, sparse setae' l,ling. Principal venation is constant in the tribe (Figure 13) ' A may An apically pointed wedge ceLl- is absent in alt Stigmoderini; lst join 2d A, Proximally or be incomplete. The degree of sclerotization is very variable, the small-er species are usually less heavily sclerotized. Legs. Tarsi elongate excePt some Dactylozodes and some Castiarina species groups which have compact tarsi . Hj.nd tarsus longer than foret especially in Calodema, Metaxymorpha and some Themognatha where hind is more than 1.5 x length of fore. Tarsomeres 1-4 usually with large setose pulvilli, except in those species with compact tarsi where the pulvilli are reduced. In a few species, the tarsi ale as long or longer than normal, but the pu1villi on tarsomeres l--3 are replaced by one or two spines. Figures 144-D compale mal-es of two species of Castiarina with normal- and reduced tarsal- pads. Tarsal claw usually simple, except in many Themognatha in which the claw has a pronounced basal tooth. 2.4 ABDOI"EN The first visiþle sternite is referred to as S3 although its anterior laterat projections are the remnants of 52' Sternites 3 and 4 are connate, the suture usually entire: tergites 1-4 are also fused' In Hyperantha, Calodema, MetaxvmorD ha. soflìe Pithiscus, and about half of FIGURE \t. Diagram of wing venation of Stigmoderini. There are four anal veins: lst A and 4th A are unbranched, 2d A has three branches indicated by subscriptsr 3d A has two branches. 0ther abbreviations are: Cu = cubitus; M = media; R = radius; Sc = subcosta; r - m = radio-medial cross vein. Sc R 4th A Al A2 3d A2 3d Ar 2d A3 2d FIGURE 14. Comparison of a male tarsus in which the tarsal pads are normal (A a C - stigmodera (C.) subtincta with a male tarsus in which the pads of tarsomeres I - f are reduced to a pair of spines (e a o - Stigmodera (C.) sexplagiata). (C) and (D) show the apex of tarsomere 2. Scale bars = 0.L mm (4, B); 0.05 mm (C); o.o2 mm (D). o / t-, frlr¡ < 9 .tÍ:: i-t \ 34 the Themo tha examined, the mal-e 57 is notched or concave exposing S9t and that of the female truncate or rounded. In Castiarina the difference is slight, the male truncate, the female rounded; both truncate; or both rounded; except Ín the S. (C.) trifasciata and S. (C.) rufipennis species groups in which the female 57 is variably bilobed and the male not (S. Barker, pers. comm. 1985). In StÍgmodera (s. s.) only S. sanguinosa is sexually dimorphic with the male 57 truncate and that of the female bilobed. The female TB is of length 0.8-1.1 x width, with posterior margin rounded or notched medially, and setation variabl-e. TB in Calodema and Me and some Themognatha has a deep lateral groove behind each spiracle. 58 is usuall-y lunate, strongly transverse with medial length less than half width, except in Stigmoderoides gloup of Themognatha in which SB is longer than wide. The female proctiger is usually 1-1.5 x as long as wide and concealed under T8, with posterior margin rounded, notched, or bilobed, and setose. The degree of fusion of the epiproct and lateral paraprocts which together form the proctiger is variable (Figures 154-8). The Stigmoderoides gloup of Themognatha is characterized by a very distinctive exposed proctiger which is a heavily sclerotized, strongly convex, elongate sclerite (length 2-2.5 x width), with upturned posterior margin pointed or notched (FÍgures 15C-D). A few species of Themognatha e. g. S. (T.) maculiventris, S. (!.) nickerli, and !. (T. ) alternata have very long setae on the terminal segments. The male TB is rounded or notched posteriorly, length usually less than, or equaJ- to width, setose on posterior half. TB in Calodema Metaxvmorpha and some Themognatha has a Lateral depression behind each spiracle as in the female. SB is strongly transverse, lunate, medial FIGURE 15. Mal-e and female proctigers. (A) - Female Stigmodera (S.) macularia, dorsaÌ view¡ suture between epiproct and paraprocts pronounced. (B) Female Stigmodera (S.) gratiosa, dorsal view, fuslon of epiproct and paraprocts complete, no suture visibl-e. (c-D) Fema1e Stigmodera (f. ) reichei, dorsal and Lateral- views, showing elongate, strongly convex proctiger. (E) MaIe Stigmodera (S.) macul-aria, dorsal view, apices of paraprocts overlapping base of epiproct, fusion incomplete. (F) MaIe Conognatha etioissima dorsal- view, fusion between paraprocts and epiproct complete, no overlap. Abbreviations: EP = epiÞroct, PP = paraproct. Scal-e bars = l mm (A, C, D)r 0.5 mm (8, E, F). l¡ rlt I ¡¡¡l tt B ¡ | I ¡ r¡ A tt l''' I ¡ ;,t I c D F E ¡ t t¡ tl I I , t¡ ¡ E PP t AI 75 lengthO.2-O.twidthrposteriormarginsetose'roundedorconcave medially. the base The male proctiger has paraprocts distinct and overlapping (s' in which of the epiproct (Figure 15E), except in conognatha s') posterior margÍn is fusion of T9 and T10 is complete (Figure 15F) ' The 59 constant in shape' rounded or notched, with marginal setae. Male is half, and tength equal to or greater than width, with setae on apical posterior margin rounded or truncate' 7 Abdominal musculature. The abdominal musculature of Castiarina SPP" f Stiqmodera ( s. s.) spp., 24 Themognatha sPP'r f0 5 Dactylozodes sPP'¡ calodema sp., I Metaxymorpha s9., 2 Pithiscus sPP. r 16 illustrates the and 2 Hyperantha (s. s.) spp. was studied. Figure arrangement in Themognatha' the Long tergolongitudinaL muscles (tlg) a3e plesent across full widthandfullheightofthesegmentonTl,andT4-16onlyin Calodema, Mele¡Irnorpha. and Stiqmodera (s. s.), Themognathat Castiarina had a few narrowt pithiscus. some of the larger species of Themognatha e.g. of ten medial bands of tlg on Tf, but this varied intraspecifically five had tlg on T3' A single specimen 1.. (I.) mniszechi examined, only of S. (!. ) pubicollis had one narrow band of tlg on T2' Dactylozodes havelongtlgonTlandT2,andshort,fullwidthtlgonT3-T6; ALary muscles Hvoerantha ( s. s. ) have t]g T1-T6, with 12--f6 short' two (a1) arising from the mid lateral antecosta of each tergite comprise same segment, the fan-shaped bands inserting on the heartr one in the by Kasap other in the preceding segment, as recorded in other buprestids (ts) present on and Crowson (1975). A pair of tergosternal muscles is every segment. divergent Long sternomedian longitudinal (smlg) and sternolateral short (sld) muscles are present in all species on s4-s6. Additional FIGURE l-6. Internal- views of abdomen showing the tergal (A) and sternal (B) muscul-ature of maLe Stigmodera (I.) tibialis. On1y segments I - 7 are ill-ustrated. In (A), the heart (h) is shown with the alary muscles on the right and the axrangement of the tergal muscles on the left. Muscles are labelled with the abbreviations: al = a1ary, sld - sternoLateral divergent, sllg = sternolateral longitudinal, smlg = sternomedian longitudinal, tlg - tergolongitudinal, ts = tergosternaL. Sca1e bar = 2 mm. A T1 T2 s T3 Ø T4 pore T5 T6 rls splracle T7 B Ê ts S3 Ë Ø smlg S4 sllg td S5 e S6 S7 v6 in sternolateral longitudinal muscles (srlg) were distinguishable Stiqmodera tha Calodema and Metaxymorpha. rporest present small, paired, translucent slit-like areas or are pores T5 and T6 on T2-T6 in most species of the tribe. usually the in pores are are concealed by the overLapping tergite in front. sternal present in npst speci.es of stigmoderini on s4-s6, but in a few species only on 54, or no Pores visible' 2.5 FEMALE REPRODUCT T\/E SYSTEM The female reproductive system is comprised of an ovipositor' vagina, median and lateral oviducts, ovaries, bursa copulatrixt gland' spermatheca and duct, and in some species, an accessory ovipositors studied in 109 species of Stigmoderini (Appendix B) were 47 compared with those of 84 other buprestid species representing genera, 25 tribes, and 9 of the lJ subfamilies recognized by cobos (1980) (Appendix D). In addition, comparisons were made with descriptions of female genitalia by Tanner Q927) who examined and illustrated 125 genera in 66 famil-ies of coleoptera, and wandolleckrs (fgO¡) study of 73 species of Coleoptera, including 3 buprestids' The ovipositorofstigmoderiniwasfoundtobeunique.Thefemale 28 other reproductive system of stigmoderini was compared with that of of buprestid species representing t7 genera in 7 tribes. The number and ovariol-es per ovary was asceltained for 78 species of stigmoderini (Appendix D)' compared with that oî 28 other buprestid species As stated in Materials and Methods, no single scheme o1 The nomenclature of female genitalia was thought to be satisfactory' (198I), but for terminology adopted here is based on that of crowson by an clarity I have introduced some new terms which are indicated asterisk in the description which folLows. Table 2 presents a TABLE 2. i-son of terminol-o s of the fema le terminalia Scudder Lindroth Tanner Crowson Burmeister Bitsch Bils Mickoleit Gardner Newton+Th. r976 I977 l-96L L957 1927 this thesis L987 1981 I98O, L976 I979 proctiger T9+I0 T10+ll TIO T10 T10 proctiger T9+10 T10 epiproct or T9 T10 EP T9 (part) paraproct T9 (part) T9 Í9 T9 paraproct T9 paraproct 19 (part) r9 T9 (half) divided PP (one half) T9 gonocoxite gonangulum gonangulum T9 (part) val-vifer val-vifer gonocoxite valvifer proximal antero- (part gonocoxite ventral (cranial (appendage of !tr I S9 gonocox. g) (one half) process T9 part) of T9) dorsal vaLve DV lateral- gonocoxite hemi- coxite gonocoxite coxite distaL gonocoxite gonocoxite style- (caudal- 9 sternite ( gonapo- gonocoxite S9 sclerite bearing 2 part) (S9+radix) (se) valve SV physis 59) gonostyle stylus stylus stylus style stylus lamnium style ST stylus stylus of style appendage s10 ventral- s9 vulvar- l-obes membraneux S8 val-ve VV ""1s¡1te(SB) FIGURE 17. Femal-e reproductive system. (A) Dorsal- view of the whole reproductive system of Stigmodera ( T. ) afternata spread to show the oviducts and ovariol-es. The elongate bursa and spermatheca extending beyond it are typical of Stigmoderini. (B) VentraL view of the above ovipositor. (c) Dorsa1 view of Stigmodera (!. ) sanguinosa showÍng only the tril-obed bursa and short, stout spermatheca and duct. (D) Dorsal- view of Conognatha (P. ) viridiventris showing only the bil-obed bursa and spermatheca extending beyond it. Abbreviations: AG = âccêssory gland, B = bursa, DV = dorsa] valve, L0 = lateral oviduct, M0 = median oviductr 0V = ovariole¡ PS = particle sac, SP = spermatheca, SPD = spermathecal- ductr SV = style-bearing valve, VA = vagina, W = vul-Va, VV = ventral valve. Shading represents scl-erotization. Scale bars = I mm (A - C),0.5 mm (D). A V LO MO B PS v SV VU D SP c D I D t I I I I I I I FIGURE 18. Sagittal section through the terminal segments of female Stigmodera (S. ) sanguinosa. Abbreviations: A = ârìus, B = bursa ¡ DV = dorsal vaLve, M0 - medi-an oviduct, p = proctiger, PS - particle sac, R = r€ctuÍì¡ SP = spermatheca, SPD = spermathecal duct, SV = style-bearing valve, 59 = 8th sternite, TB = 8th tergite, VA = v¿gi¡., W = vulva, VV = ventral valve. R T8 A B P9 VU VA SPD MO 8P O.5 m FIGURE 194. Qvipositor of Stigmodera (!. ) sanguinosa, lateral view. Abbreviations: DV = dorsal valve, EP = ePiProct, pp = paraproct, R = rectum, ST = style, SV = style-bearing valve¡ VF = valvifer, VU = vulva, VV = ventral val-ve. Scale bar = 0.5 mm. FIGURE 198. Spermatophore from the right spermatophoral gland rheadl of mal-e Stiqmodera (S.) macularÍa, dorsal vi.ew. The end is on the right. Scale bar = 0.5 mm. A R I EP .=a-_{- PP ..ì.\\\**',Nk \ DV .tl ¡ \ ST \ SV VU VF \a I n\r VV B t at ,Ì ry :-- l:-. - 17 l-isted by Smith comparison of selected terminologies, other synonyms are (Ig6g). The terminalia and reproductive system of adult female Stigmoderini (Figures L7-20) have the following components, whose abbreviations are given in parentheses' the The proctiger (P) (Figures 18-I9A) is formed from the fusion of paired, lateral paraprocts (PP) (derived from the 9th tergite) and the anus (A) on epi ( Ep) (derived from the 10th tergite), and bears the pair lateral its ventral- surface. The 9th sternite is reduced to a of hemisterniùes, o" u"Iuifers (VF) to which the paraprocts are fused anteriorly. The valvifers articulate with the sclerotized base of the ovipositol.TheyareunusuallylonginSofnef¡[email protected]. S. (T. ) maculiventris. The ovipositor of stigmoderini consists of three plates or vaLves: (Figures a dorsal valve, a style-bearing va]ve, and a ventral valve r7-re). The dorsal valve* (DV situated between the style-bearing valve and proctiger, is convex, usually bifidr and scl-erotized baso-laterally,withlongdowncurvedapicalsetae.The stvl VV fused 1atera1lY, valve* SV and membranous ventral- valve* are from forming a genital tube, the vagina (VA), and are probably derived the gonocoxites of the 9th sternite' The thickened style-bearing valve bears on its posterior margin several rows of sclero tÍzed spatulate been setae* and a pair of styles (ST); the Latter are thought to have derived from the appendages of the 9th segment (Crowson, 1981) ' The styles are short, cylindrical appendages, partÍalIy sclerotized, with apical setae; they occur in the medial third of the style-bearing valve' anteri.or to the sPatulate setae' The style-bearing and ventral valves are strengthened by rdorsal rami' (Smith' I97O) sclerotized rods or plates, the and ventral ' and their degree of or 'baculit (Tanner, 1927). The shapes of these v8 sclerotization vary between species groups and higher taxa. The plates are confluent with the scl-erotized base of the valves forming a very rigid structure. The ventral valve is shorter than the style-bearing valve so that the genital opening or vulva VU is ventral rather than terminal as in most other buprestids. Thé ticle sac* PS (Figures 17A, 18) is a chamber which Ís open posteriorly and bounded anteriorly by the membrane connecting the style-bearing and dorsal valves. It may be unilobed or bilobed, and a scLerotized median strut or septum* supporting the dorsal valve and dividing the particle sac longitudinally may be present. The length of the particl-e sac relatÍve to the length of the dorsal valve is variable. The bursa (B (Figures 174, 18) is generally an elongate, unilobed sac with a thick, sometimes convoluted lining which functions as a spermatophoral receptacle. The bursa is bilobed in Conognatha (!.) sagittaria and C. (P.) viridiventris (FÍgure 170), and tril-obed in Stiqmodera (S.) roei., S. (S.) macularia, S. (S.) jacquinotit !.. (9.) porosa and S. (!.) sanguinosa (Figure 17C). In some species e.g. S. (T.) mirandar S. (!.) bonvouloiri and S. (T.) chevrolati, a smalL rectangular basal plate of sclerotized spines is present, and presumably serves to hold the spermatophore in the bursa, or to abrade it. The sperma theca (sp) narrows to a spermathecal duct (SPD) which inserts at the junction of the vagina and bursa. In most species of Stigmoderini the spermatheca and duct are long and curled distally (FÍgures 174,D); in Stigmodera (s. s. ) tney are more robust and shorter than the bursa (Figures 17C, 18). The nedian oviduct (M0). inserted on the ventral- surface of the vagina is usually short, dividing almost immmedÍately into the two lateral oviducts (L0 (Figures 17A, 18). It is attached by two bands 39 of muscle to sternite 8. The length of the lateraL oviducts varies, but may be up to eight times the length of the median oviduct. The ovary i.s comprised of ovarioles (0V closely adpressed at their distal ends. Stiqmodera ( s. s. ) and Castiarina have five ovarioles per ovary, Themognatha and Calodema have sevent Dactyl-ozodes has five or seven, and Met has five, seven or eight. The number of ovariol-es in Conognatha and Hyperanthell-a appears to be five, but this is uncertain because the availabl-e dried material was in a poor state of preservatÍon. The average number of eggs per ovariole is five, but varies according to the time after emergence that the specimen was coll-ected. One specimen of S. (f. ) murravi had an estimated 200 eggs in total-. The eggs of S. macularia, the only species examined using an electron mi-croscope, have two distinct surface textures: a smooth region, J-ess than hal-f the area, and the rest sculptured with ridges arranged in a hexagonal pattern. The accessory gland (AG) is a medial, multilobed, evagination of the ventral surface of the vagina (Figures 178 & 2OA). It is present in Themognatha, Calodema, and one species group of Metaxymorpha, and no other taxa of the tribe. Histologically the gland is comprised of a layer of unspecialized columnar cells, with a thick, lamellate intima of chitin (Figure 2OB). Posteriorly, the gland narrows to form a duct which passes through the thick circular muscl,e surrounding the vagina. The gland may be surrounded by a thin layer of tissue making the outer surface appear smooth, but still revealing the convoLutions beneath. There are al-so thin layers of striated muscle penetrating between the lobes, especially at the junction of the gland and its duct, suggesting the controll-ed release of its contents. The presence of mucopolysaccharides was demonstrated by a positive Alcian Blue stain. FIGURE 20. Accessory gland of Stigmodera (T. ) regia. (A) Whoi-e gland and duct. (B) Detail of longitudinal section through several l-obes of g1and. C = col-umnar hypodermal cel-ls, I = thick intima of chitin, L = lumen containing flocculated material-. A \str¡ateO muscle fibre duct O.5 mm ¡ \ ¿.71...\, I t ) *rì I \t \r t ( I t \ È¿ \ cf ( aa.t .¡ t I ,'{ , f l' I r. '-O.1 m I 1.ì f 1 -"ñ- 40 Baccilli were cultured from the Lumen of Stiqmodera goryi and S. regia, but according to Prof. D. Pinnock (pers. comm. 1984) they did not belong to either of the two principal genera of symbiotic bacteria in insects Bacteroides and Blattabacterium) , and were probably an infection. 2.6 l',tALE REPRODUCTM SYSTEM The male reproductive organs including the aedeagi of 60 species of Stigmoderini were studied, and the aedeagus only was examined in another J9 species (Figures 2I & 22). The testes consist of many tubulest l-10 x length of the abdomen, spirally twisted into an oval or conical shape. Each testis j-s surrounded by a sheath, as is characteristic of buprestids (Lawrence and Newton, 1982). In some specimens, the sheath is bright orange and easily peeled away, but it is often pale and so delicate as to be diffÍcult to discern. The number of tubules per testis is constant in some taxa e.g. 12 in the Dactyl-ozodes and Pithiscus examined, l-0 or J2 in Castiarina; and highly variable in others e.g. 12-94 in Themognatha, 12-50 in Stigmodera (s. s.). Where the number of tubules is high' left and right testes of the same specimen may differ by as many as 20 tubuLes. The testis is connected at its base to the vas deferens. This unites with the accessory gland of that side, before inserting on the ventral surface of the ejaculatory duct at its junction with the large, thick-walled, club-shaped spermatophoral glands. All Stigmoderini have only one paÍr of accessory gj-ands, in addition tå the spermatophoral glands. The length of the thin-walled, tubular accessory gland relative to the length of the abdomen is variable intraspecifically, 2'2O x the J-ength of the abdomen. The paired, club-shaped, thick-wal-led spermatophoral glands are uniform throughout Stigmoderini. 0f the FIGURE 21. Dorsal view of the male reproductive system of Stiomodera (c.) octomaculata with ejacuJ.atory duct extended and only the right testis iltustrated, with sheath removed to expose the spirally coiled tubules. Each accessory gland is 20x longer than the section shown on the right. Scal_e bar = 0.5 mm. spermatoPhoral gland gland -accessory vas deferens stis JaculatorY duct deagus 4t buprestids described by Kasap and Crowson (1975), they most closely much larger in resemble those of Acmaeodera although they are relatively Stigmoderini. The ejaculatory duct which is folded back on itseLf in sitú is lined with chitin and bears a longitudinal band of smaff, posteriorly directed, sclerotized spines. These presumably aid in the movement of the spernratophore along the duct by acting as ratchet' The aedeaqus. Most stigmoderini have sensory hairs on the parameres. The exceptions are Hyperanthella and Dactylozodes spitzi presence or Théry (Cobos , 1959232). Cobos (1980) used the characters Buprestidae absence of sensory hairs on the parameres to separate the into two groups. The shapes of the parameles and median lobe of which the aedeagus are usually is comprised vary within the tribe, but in the smaller taxa similar. In Stiqmodera ( s. s.) (Figures 22A-B) the lobes of the apices parameles are broad and separate for most of their length, with truncate, and basal margin lipped and notched dorso-medialIy' The median lobe is distinctive in having the apical half subparallel' ostium broadening towards the base, and the posterior margin of the medial, 0.5-0.6 length from base. In Castiarina the shape of the (see in Barker aedeagus is very variable between species illustrations (s' Barker' I979r 1980, I9e7), and is uselul in forming species groups pers. comm. 1981). of The aedeagi of Themoqnatha are al-so rather variabl-e, but that the type speci-es, s. (I.) variabilis (Figures 22C-D) is typical of (Figure many. The parameres of Calodema and Metaxymorpha 22E) have sides oblique to rounded shoulders, lobes diverging about 0'7 distance FIGURE 22. Aedeagi: parameres, in dorsal view, (4, C, E, G, I, K); median l-obes in ventral view (8, D, F, H, J, L) and lateral view (M) showing the position of the ostium. (A, B) . ) macularia (c, D) . ) variabilis (E, F) Metaxymorpha (c, H) Conognatha (C. ) pretioissima (I, J) Dactylozodes rouleti (K-M) Hyperantha decorata Scal-ebars=Lmm. A B c D , I , I I I I I I I I I H E F G L M I J K 42 from the base in most, and basal margin lipped and notched dorso-medially. The medÍan lobe is lanceolate, with posterior margin of ostium 0.8 length from the base' (Figure 22G) have The parameres of Conognatha (t. .. ) examined from the base, and lobes rounded apical-ly, diverging at 0.6-0.7 length broader than basal margin with dorso-medial lobe. The median lobe is pointed' in tire preceeding gloups, with apex triangulal or very acutely lateral ridges on the dorsal, surface, and ostium with posterior margin in- 0.8 length from the base. Moore (1981) il-lustrated the variability the shape of aedeagi in 17 species of Pithiscus' (FÍgures The The aedeagi of Dactvlozodes are variable 22]-J)' base palameres examined are elongate, rounded or pointed apically, and narrow' produced into a narlow lobe or rounded, and apophysis long posterior margin The median lobe is elongate, with a small ostium with a medial 0.7 length from the base. Many species of stigmoderini have sclerotized spot or rod anterior to the ostium of the median Lobe' (Figure 22K) are also The parameres of Hyperantha (s. s.) examined at its long and thin, the apophysi-s narrow and strongly curved ventrally the junction. The median lobe is distinctive in that the membrane of the ventral surface is expanded and folded to form a broad flap covering (Figures The small osti.um, which is 0.9 length from the base 22L-M)' spines dorsal surface of the flap is covered with fine dense scLerotized in Hyperantha (s. s.) decorata. Sperma tophores.Spermatophoresfromthespermatophoralglandsof composed of a atl specles examined are of similar form. They are softer transparent or pale amber gelatinous materiaL and an opaque substance;theydissolveinco]-d5%KoH.Theshapeofthe spermatophoral gland' spermatophore conforms closely to the walls of the 4V rheadl Each spermatophore has the same components: a distal tobed and a longer, narrower, curved'taiLr (Figure l9B). Qne of the lobes of the head is a narrow beak-like process terminating in a fine point. This structure resembles that of some Orthoptera as iLlustrated by Tuzet |canal be Q9772286) and labelled ejaculateurl, but it may not homoJ-ogous. The tail sometimes extends into the ejaculatory duct in which case it is Protracted. Spermatophores are found infrequently in the female tract, however presumed spermatophores, often in a state of partial disintegration, have been removed from the bursa or vagina of a number of specimens' Considerable changes to the spermatophore must take place during or after transfer to the femal-e. Recognizable components of the mal-e spermatophore e.g. the beak-like process and tail are absent, and it is possible that the spermatophore evaginates as it is expelled through the ostium. A mal-e specimen was found with a spermatophore in the process rhead' of being extruded from the ostium, apparently first' Moreover' spermatophores taken from the femal-e do not dissolve in 5% KQH' so it seems that sclerotization occurs during transfer' 2.7 ALIMENTARY SYSTEM The alimentary system of 79 specÍes of Stigmoderini studÍed (Appendix B) provided a few characters useful for distinguishing genera. It is described here in detail as it has not been recorded before and is characteristic of the tribe (Figure 23) ' The foregut is usually short, 0.3-0.6 the length of the midgut, with two wide foregut lateral- diverticula which occasionally contain polIen. The midgut is covered with smaIl, evenly distributed evaginations or regenerative crypts which are moderately dense in Castiarina (I-7 dj-ameters apart), or very dense (less than one diameter FIGURE 2r. Alimentary system of Stiqmodera (T.) aLternata uncoj-l-ed to show the Malpighian tuþul-es grouped proximal-ly, typical of Themognatha, Calodema and Metaxymorpha. lateral diverticula cardiac valve -CâêCUllì gut small intestine arge intestine alpighian tubules pyloric valve m ctiger 'l mm. 44 apart). Anterior to the cardiac valve, the junction of the foregut and midgut, ale a pair of caeca. These are very short (less than 0'1 the length of midgut) in Stigmodera (s. s.), and longer (0.1-0.5 the length of midgut) in the rest, and also covered with regenerative crypts' The hindgut is equal to, or greater than, the length of the midgut and coiled anteriorly, so that it overlies the distal end of the midgut. The hindgut is differentiated along its length: anteriorly is a narrow small intestine surrounded by circular muscles; posteriorly is a wider, thin-walled large intestine. The gut terminates in a rectum. The six cryptonephric Malpighian tubules originate at the pyloric valve, the junction of the midgut and hindgut; at about 0.7 along the length of the hindgut they become closely adpressed to it and convoluted. Proximally, they are in two groups of three in Themognatha, Calodema, Metaxymorpha and ungrouped in Stigmodera (s. s.), Castiarina (ei anus on the ventral side of the epiproct. 2.8 NERVOUS AND RESPTRATORY SYSÏEMS The nervous system is invariate j.n all StigmoderinÍ examined, and follows the general structure of Buprestidae as recorded by Kasap and 45 Crowson Q975). The nerve cord consists of the suboesophageal ganglion connected to the brain by circumoesophageal commissures, the prothoracic gangJ-ion, the fused mesothoracic, metathoracic and first abdominal segment ganglion, and four abdominal ganglia of segments 2, 7, 4, and 5-8. The respiratory system is comprised of a pair of lateral trunks and a network of fine tracheae with abundant aÍr-sacs in the abdomen, and is also constant in the trÍbe. Spiracle L is located in the pleural membrane, 2-8 in the tergites. 46 I KARYOTYFE ANALYSIS indicate My karyological studies o1 36 species of stigmoderini genericstabilityofchromosomenumber.Thelimiteddataonother Agrilus studied by buprestids supports this e.g. the four species of Dicerca reported smith ffgt+g, 1957) have 20 or 22; the three species of have20(SmithrI957)randtwospeciesofsternocerahave26(Asana' 2n 39 for male Makino and Niiyama, Ig42). If the tentatÍve count of = reasons' Julodimorpha is confirmed, it will be notabl-e for two more than Firstly, it is doubl-e the ancestral number, and considerably thehighestdíploidnumberrecordedprevj-ouslyforBuprestidae' buprestid reported so far Sternocer¿ 2¡ = 26. Secondly, it is the only sex' This with an uneven number of chromosomes in the heterogametic system' The coul-d represent either an X0 or a multiple sex chromosome The Iatter could aLso contribute to the large number of chromosomes' and their very small many acrocentric chromosomes observed in one cell, that size relative to the one obvious metacentric, suggests strongly the ancestral fission may have been responsible for the doubling of examining meiotic number, rather than polyploidy. Further studies elucidate the cells and al-so using femal-e tissue are required to karyology of the sPecies. The differences in chromosome number of the two Themognathat l'.(].)nicker]iandS.(f.)alternata,withln=20maybedueto are suggested Robertsonian rearrangements. centric fusion or fission but not in the when there is a change in the number of chromosomes, (the f ondamental-e .In total number of major chromosome arms nombre species with 2n 221 view of the lack of acrocentric autosomes in the = 0n the other these mechanisms seem Less likely to be the explanation' the karyotype hand, transLocations have played an important role in evolution of þeetles (Virkki, 1984246) ' 47 autosome I is In Stiqmodera ( f .) nickerli and S. (T.) al-ternata, approximate}ytwicethelengthofautosome2,andrepresentsabout 22-23%TCL(Tab]e1).Intheotherstigmodera,autosomelvariesfrom 1-l.5xlengthofautosome2andrepresentsonly];2-I5%rcL size of (Themognatha), or 13-18% TCL (Stigmodera (s. s)). The large lCL) suggests that it autosome I in s. nickerli and s. alternata Q2% stigmodera by may have evol-ved from a karyotype simil-ar to the other (6-9% autosome transLocation of one of the smaller autosomes TCL) on to in the I (12-18%), followed by a pericentric inversion which resulted more material had been new autosome L reverting to a metacentric. If this avaiLable, C-banding techniques could have been used to test 20 * XYp suggestion. The marked preponderance of species with /¡ = in stigmoderini suggests that 22 chromosomes may be the ancestral condition of the triÞe, with the complements of s. nickerLi and S. alternata representing derived states' stigmodera is The most distinctive difference between species of species the relative size of the X chromosome (see Table f). In some e.g.S.porosaands.heros,theXapproachesthelargestautosomein of length, but in most it is one of the sma]-lest. In the majority Moreover, in species the arms of X are unequal, often markedly so' by both the most of the species, X is heterochromatic as indicated that the differential staining and its diphasic form. It is probable variation in relative size of the X chromosome is due to the duplication ordeletionofconstitutiveheterochromatinwhichcou]-doccurwithout as transLocations may deLeterious effects. structural- alterations such also have been reponsibl-e for some of the variation observed' gloups Variations in the X chromosome do not correspond to species formed on the basis of other characters' 48 Howvaluab]eiskaryotypeanalysisasatoollnsystematics? of coleoptera do crowson (1981:414) asserted that chromosomal features higher levels' not provide very reliable characteristics of taxa at thoughhenotedtwoexceptions:superfamilyCantharoidea-aIlt25 and åuperfamily species studied have an X0 sex-determíning mechanism; a basic 10 AA' not curculionoidea - aLt species studied are derived from species of Aphidae expressed 9AA. Bl-ackman (1980) in his study of 180 data this opinion aLso, although he found that on the whol-e, chlomosome tendency for the corroborate generic concepts since there is a cLear and that differences in chromosome number to be stable at this level; subgeneric groupings' chromosome numbers sometimes agree with recognized phylogenetic Yadav and Pi}lai {.l:97Ð in their study of re].ationshipsofgeneraandsubfamiliesofscarabaeidae,placed they found varies considerable importance on chromosome number which fromn=6-1}¡withl50ofthelg4specieshavingthemodalnumberof n=}0.TheyassertedthattwotribesAdorrhinyptiiniandAdoretini they both have should be considered to be closery related because subfamilies, Geotrupes 2n = 22. HOweVer, when two genera from other close on other grounds were and US , not considered taxonomicatly them as an found to have a unj.form l¡ = 22, Yadav and Pil-lai cited This paper i]lustrates example of parallelism in karyotype evolution. to be just the point that many taxonomists consider chromosome number Low weight if it does not one more morphological character, and one of other reinforce their notions of the taxonomic groupings based on characters. and inversions have More detailed studies involving translocations those of Yates' proved valuabl-e in phylogenetic studies, for example genera of peromyscine Baker and Barne tl 0979). In this study of three outgroups lor rodents, Neotomodon alstoni was chosen as one of the 49 0n the basis of determining relationships within the genus Peromyscus' species of Peromyscust synapomorphic inversions shared with several however,Yatesetal.conc]-udedthata].stoniiscongeneric. Þy some changes in chromosome number and structure are assumed evol-ution of a workers to represent unique historical events in the taxon. As such they may be used to formulate genealogicar hypotheses (Wiley' 198L2727)' Farris in the same vray as other morphological- data phylogenetic (L978) di.scussed the use of inversion data to generate postulate to be trees. He showed that even assuming the uniqueness true, multiple trees coul-d often be drawn since the ancestral inversion couldpersistinaheterozygouspopulationandreemergeinsome using the descendants in the homozygous state. Farris advocated principle of parsimony i.e. seLecting as best the tree which minimized the number of heterozygous ancestors postulated' Acladisticanalysisofkaryotypeswasnotundertakeninthisstudy were not sufficiently because too few species were available, the data possible to decide which detailed, and without outgroup data, it was not state was primitive and which derived' firstly, My exploratory study of the karyology of Stigmoderini, supports the general observation of generic stability in chromosome the exclusion number; secondly, provides one more character supporting suggests that of Curis and Julodimorpha from the tribe; and thirdly, karyologymaybeusefulindelimitingsubgeneraorspecj-esgroups. simirar both have S. nickerli and s. al-ternata which are morphologically more material 20 chromosomes. The use of c-banding techniques when rel-ationships between becomes avail-able could provide insights into the the acrocentrics' and the the complements of 20 and 22, the evolution of evolution of the X chromosome' 50 characters for chromosome data can provide a suite of karyological in some animal the systematist to analyze and has been used successfully which integrates groups. More confidence can be placed in a phylogeny biochemical' data from a wide array of character classes: morphological, biogeographic' behavioural, ecological, physiological, cytological and character classes Moreover, conflicts between analyses using different can highlight areas of special interest' Coleopterancytogeneticsisstillinitsinfancybecauseofthe so size and associated technical difficulties of the chromosomes' uniformity perhaps it is too early to assess its use in systematics' supports the naturafness of of many of the gross chromosomal features Stigmoderiniasagroup,butgiveslessscopeforelucldating relationships within the tribe: however, more detailed karyological information' analyses in the future may provide valuable phylogenetic 2 MORPHOLOGICAL CHARACTER ANALYSIS Idealty,thecharacterschosentode]-imitataxonshouldbe visible, and constant within that taxon, unique to that taxon, clearly natural groupingt well, defined. Furthermore, if the taxon is to be a characters i.e. monophyletic (sensu Hennig lI966D, then only apomorphic were rigidly shouLd be considered. However, if all these criteria there would be such a applied to stigmoderini and its higher categories, paucityofcharacterstouse,thatnoanalysisofthegroupswouldbe not possible. EvoÌution in general proceeds in diverse, and lrequently a mosaic the most parsimonious ways, and the present fauna displays to most distribution of character states. Exceptions can be found large taxa' generalized characters, particularly in the diagnoses of often a character which is distinctive and stable in one taxon and 5] the seemingly useful, may be highly variable in another. Many of character states are part of a continuum rather than discrete. several characters are internal and only discernible in fresh material. What lollows is a discussion of the taxonomic characters used in the tribal, generic and species groups diagnoses. Taxonomic characters are characters of convenience, useful in distinguishing between taxa: not al-l can be used in phylogenetic analysis. Indeed, many must be rejected because they are judged to be plesiomorphic or because no (Lawrence decision can be made on the direction of evolutionary change and H]avac, L979). Those selected for phylogenetic analysis are marked with an asterisk (*). The distribution of these characters is 0 summarized at the end of this section in Table 3. States designated designated Ir lr Ïr are consj.dered to be reLatively primitive. states ' etc. are considered to be rel-atively derived, but the relationship between those states is unknown. States designated 0, I, 2 represent a transformation series with 2 being the relatively most advanced state' characters whose polarity is unknown arei designated A' Br C etc' 0f the characters previously overlooked by other workers, the most useful are those of the femal-e reproductive system' General bodv form (length Character l- Size is arbitrarily classifed as (A) small lessthan20mm);(B)medium(20-]0mm);and(C)large(greaterthan ¡o rm). Most castiarina, DactyLozodes, Hyperanthelta are small; medium; Conoqna tha (s. s.). and Pithiscus are small to Stiqmodera (s.s.),Hyperantha(s.s.),Metaxymorphaaremedium;most considered atha and Cal-odema are medium to large. Théry Q929) (195',) large size a more recent character, an assertion echoed by cobos andHoLmogTg).However,DeJong(1980)notedthatininsects 52 |,increase Ín body size is not known aS a general evolutionary tendencyl|.UpperPermianfossilbeetlesrangedfrom5mmto25mmlong buprestid (crowson, fg75), but it does not necessarily fol]ow that the as to polarity of ancestor was small to medium in sizer so no assumption this character is made here' (all Character 2 tsody shape: (A) dorso-ventrally flattened Stigmoderini) ; (B) cYlindrical Julodimo a). (A) or Character 3 Body shape in Lateral view: wedge-shaped (s' and some sÍnuous (Calodema, Metaxymorpha, Conognatha s') A wedge-shaped Hvoerantha ( s. s.)); (B) suoparalLeL (aII other taxa)' so it is not body would presumably confel an aerodynamic advantage, high-flying surprising to find it in calodema and Metaxymorpha which are while species living 20 m or mole up in the canopy of the rainforest, forms lumber the taxa with more robust, less streamlined, subparallel (A) is a vely over 10w heath and mall-ee. correl-ated with character which may be to smooth, shÍny pronotum, the functional significance of minimize wind resistance in high, fast fliers' Character s of the head (B) Frons hair is Character 4 Frons hair: (A) present; absent' reduced to minute variable in some groups, but always absent or at least and s. (c') iubata setae in castiarina (except s. (c.) rudis carter generic character' Blackburn) and Conoqnatha (s. s.) where it is a useful slightly Character 5 Eyes: (A) large, subparallel or converging Large eyes are on vertex (Stigmoderini); (B) smalI, para}lel' characteristic of the family (Lawrence, 1982), however, most 53 Stigmoderini have eyes larger than those of other buprestids examined' aS diagnostic and the character has been used by most previous workers of the tribe. within the tribe, the eyes of calodema, Metaxymorpha and than conognatha are consistently larger and more strongly converging use in those of the other taxa, however, this character is of timited generic diagnoses because the states are not discrete' Character 6* Mouthparts: (0) not prolonged, galea oblong with sparse, short setae (Hvperantha (s' s'), Hyperanthellat Dact l-ozodes (1) protonged, galea ovoid or many Pithiscus and some Castiarina); (9' globular with }ong dense setae (stigmodera (s. s.), conognatha s')t some castiarina). Themoqna tha. calodema, Metaxymorpha, some Pithiscus, generalized out-group Mouthparts not prolonged is characteristic of the which have and is considered Primitive. Chilean Pithiscus many of short mouthparts, feed exclusivety on pollen (T. Moore' in litt' are IggT). The species of castiarina which have abbreviated mouthparts case' leaf and petal chewers (S. Barker, pers. comm. 1981), and in this or the character may represent the retention of the primitive condition' mouthparts and may Þe a reversal, i.e. secondarily derived. Prolonged and are the clypeus are considered to be an adaptation to nectar-feeding of derived state: a galea with dense 1ong, brush-Ìike setae is typical galea are fl_oricolous beetles. The length and density of setae on the better consíbered as continua, however, because in nectar-feeding Castiarina these characters are variable. Théry Q929) used placement in prolongation of the mouthparts as the soLe criterion for curis in stigmoderini, resulting in his inclusion of Jul-odimorpha and the tribe, whích is not supported by other characters' 54 triangulart character 7 shape of labrum: (A) curvilinear notch, without translucent anterior margin with or without a weak medial- anterior border, medial groove present (cal-odema, Metaxymorpha' with bÍlobed Themoqnatha); (B) trapezoidal, strongly bilobed (súiomodera (s. s.)); (c) translucent border, medial groove pronounced straight' medial rectangular, bilobed, with translucent border almost some groove usually present (conognatha (s' s')' Pithiscust strongly transverse' anterior margin Ca stiarina) ; (D) rectangular, straightorweaklybisinuate,withoutborder,medialgrooveabsent and some (Dact vlozodes HvpeIantha ( s. s.), Hyperanthella, the tribe' it is Castiarina ) . Although the labrum is variabLe in providing a useful diagnostic constant and distinctive in some taxa character. from base' rounded Character I Mandible: (A) elongate, tapered ventral subapical dorsal subapical tooth weak or absent, and a rounded Themoônatha' Calodema' very weak or absent (Stigmodera (s' s')' (B) rounded dorsal Conoqnatha ( s. s.), Metaxymorpha); elongate, subapicaltooth,andalarger,moreacuteventralsubapicaltooth two dorsal subapical and ( Pithiscus); (C) elongate, one or irregular castiarina); (D) squat' one ventral subaPical tooth Da l-ozodes subapicalsr one ventral almost as wide as rong, one or two sharp dorsal (E) cylindrical' apex subapical (Hvperantha (s. s.), Hyperanthelta); (¡ulodimorpha)' The rounded, subapical teeth very weak or absent As expected' there is a mandibLes are a useful- generic character' lack of teeth' and Leaf and correLatj.on between nectar-feeding habit and petalchewinghabitandpresenceofsharpteeth.Strongmandiblesare enable them to emerge from the needed by a1t stigmoderini, however, to pupal stages' wood in which they spend their larval and 55 (A) Character 9 Terminal segment of Labial and maxilJ-ary palps: tapered apically (Dactylozodes, Hyperantha (s. s.), Hyperanthella, and (a11 some Pithisçgq e.g. c. (P.) fisheri ; (B) truncate or rounded other taxa). Character 10 Ligula: (A) rounded apically (atl Stigmoder:'ni); (Jul-odimorpha)' (B) eLongate, triangular, notched or bifid apically Character 11 Prementum attached to mentum: (A) dorsally (s. s. ( tha, Calodema, Me!¡¡Imorpha.t Dact lozodes Hyperantha ) and Hvperanthetla); (B) anteriorly, with grooved labial suture suture Castiarina and some PíthÍscus ; (C) anteriorly, without labial (Stigmodera (s. s.), Conognatha (s. s.), and some Pithiscus)' The generic attachment of the prementum to the mentum is an important character as it is constant in all higher taxa except Eithiscus' In but anterior many of the general-ized outgroup the attachment is dorsal, attachment does occur in some species e.g. schizopus laetus (Schizopodinae) and Jul-odimorpha bakewelli, so it is not possible to assign polarity to these states. All Stigmoderini with anteri-or attachment have prolonged mouthparts, but the convelse is not true' Character ]2* Mentum: (O) not notched; (1) anterior margin notched are lateraIly ( DactvLozodes except undescribed sp.). Lateral notches nor presumed apomorphic si.nce they do not occur in other stigmoderini, were they found in the generalized outgroup' Character If Antennal cavities: (O) smaLl' open anteriorly' (1) situated near the border of the eyes and clypeus (all Stigmoderini); (Julodimorpha cl-osed anteriorly, situated frontally between eyes 56 bakewelli).Sizeandpositionofantenna].cavitiesisacharacter (1857) in given emphasis by earlier workers, starting with Lacordaire ' categorizing the higher taxa. crowson (1981:16) considered antennal the cavities r'low down and lateral between the eyes and the bases of is mandiblesrf to be the primitive state in beetles. If this ässumption correct, all members of the stigmoderini have the primitive state' Character l-4+ Antennal length: (0) approximateJ-y equal to or greater than width of head (most stigmoderini)i (1) noticeably shorter and than width of head (Dact Lozodes Hyperantha (s. s.), Hyperanthella, a few Castiarina e. o. S. (C.) aeraticollis Carter). (1) Character l-5* Antennae: (O) toothed from 4th segment; (2) toothed from 5th segment; (l') toothed from 3rd segment; toothed in from 6th or 7th segment. Antennae are toothed from the 4th segment Stiqmodera ( s. s.), Themognatha, Calodema, Metaxymorphat Castiarina Conognatha (s. s.), and Pithiscus from the 4th or 5th in ( and Hiperanthe]la; and from the 4th, from the 5th in Hyperantha :'-.s. s. ) # 5th, 6th or 7th in Dactylozodes. Male Hypostigmodera variegata have fourth antennae toothed from the meriston. Antenna toothed from the considered segment is widespread in the generalized outgroup and plesiomorphic. Loss of a tooth and its fovea, and shortening of the the antenna as a whole are presumed to be apomorphic: reduction in wíth number of toothed segments is not necessarily concomitant shortening of antennae. Character 16 Toothed antennal segments: (A) triangular, widest apically (Dactylozodes); (B) elongate, length equal to or greater than width, tooth rounded, widest at middle (stigmodera (s. s.), Themognathat 57 Castiarina. Cal-odema, Metaxymorpha, Conognatha (s. s'), and Pithiscus); (C) very strongty transverse, tooth rounded, (Hyperantha (s. s.), Hyperanthella); (D) bipectinate or tripectinate (maLe Hypostigmodera) . The shape of the toothed segments is highly variable in buprestids, so there is no indication of what the polarity of these character states might be. (B) Character 17 Arrangement of antennal sensilla: (A) diffuse; concentrated in a subterminal- fovea and i-n a marginal secondary sensory area (aII Stigmoderini except Hypostigmodera variegata). The arrangement of sensiLl-a, whether diffuse or concentrated in foveae, has been used as a major character in categorj-zing subfamilies by Théry (Ig29), Cobos (1980) and others, however there are several problems with this character. Firstly, the sensilla are difficult to see under a dissecting microscope: none of the earlier workers mentioned the secondary sensory area in Stigmoderini although it is often as large as the fovea. Secondly, many intermediate conditions exist between the (1980) two character states, making that dichotomy in keys e'g' Cobos unsatisfactory. Levey (1978a) also questioned the val-ue of the character, and asserted that regression and perhaps redevelopment of (1901) sensory foveae occur within a phyletic line. Kerremans considered diffuse sensilla to be the primitive condition, while Théry (1927) asserted that pores concentrated in foveae indicate primitiveness. Based on the notion that increased organization is an evol-utionary trend, concentration of sensilla in foveae could be presumed to be an apomorphic character, however, the argument is weak and no polaritY is assigned here. ri ls mo ls m g g lp mp mP g g mp mp 58 (A) dorsal fovea Character 18 Antennal segment 11: with Calodema, Metaxymorphat some Stiqmodera (s. s.), Themognatha, (B) dorsal fovea (all other Castiarina some Pithiscu S ) ; without present in the taxa). A dorsal fovea on segment 1I is rarely generalizedout-group'nevertheless,ithasbeenrecordedinsome (NeLson' 1981), and the Buprestidae e.g. Beerell-us (Polyctesini) polarity of this character is in doubt' Characters of the thorax more than diameter of character l'9 Prosternum: (A) long' length (B) short' Iength less procoxae anterior to them (alt Stigmoderini); (Julodimorpha) This than diameter of procoxae anterior to them ' character is variable within the family' (o) margin character 20 sides of prothorax: with lateral castiarina' some incomplete (conognatha (s. s.), HyperanthelLa, Dact l-ozodes ;(1)withlateralmargincarinatetoexplanate(allother taxa).Théry:]92Ðregarded|'prothoraxmoreor].essbordered||asa that incomplete primitive character. However, Lawrence (1992) stated lateraledgesarecharacteristicofthefamily,sothisisconsidered heretobethemoreprimitivestate.Thedivisionissomewhat on basal I/4 only arbitrary as a continuum can be seen from a cari'na pronounced carina' to e. g. Conoqnatha amoena, through a weak carinat strong}yexplanatemalgnse.g.Stigmode¡agrätiosa,sothischaracter has low weight. pronotum (0) not character 21* Postero-Lateral angles of (conognatha (s' s')); (2) pronounced (most taxa); (I) prominent, acute prolonged, very acute (Hyperantha (s' s')' Hyperanthella)' 59 (A) almost straight Character 22 Pronoto -elytraI articulation: ( Stiqmodera ( s. s.), Metaxymorphat Pithiscus (Conognatha (s' s')); (c) strongly Hyperan thella); (B) bisinuate and Castiarina bisinuate Calodema Hypera ntha ( s. s. )). Themoqnatha with prominent medial are variabLe. The strongly convex articulation most striking features of lobe in front of the scutellum, is one of the Calodema,andisusefulfordistinguishingthatgenus.The pronoto-elytralarticulationinthesubfamilyBuprestinaeisnearly straightorfeeblybisinuate,withmedianlobeofthebaseofthe (Cobos, I98O), however' all three pronotum generally not differentiated statesoccurÍnthegeneralizedoutgroupsonopolarityisassigned. and shape of the HoIm (1978) considered that the sculpturing pronotumprovidessomeofthebestgenericcharactersinthetribe Acmaeoderini.InStigmoderini,however,mostcharactersofthe between higher categories pronotum are of limited use in distinguishing þecausetheyarerepresentedbyacontÍnuumofstatesandarethus difficult to quantifY' (0) enlarged Character 2f* Anterior prosternum: not des. (Stiqmodele (s. s-)' Castiarina Themoonatha Dactvl-ozo (s. (I) with low hump (some Hvoeranthe IIa. and some Hyperantha s.)); (2) acute conical process Conoqnatha. some Metaxymorpha); with (s' s')t and some Calodema some Metaxymorphar some Hvperantha Conognatha' concurred Conoqnatha). Hoscheck Q934), in his revision of demarcation between these with lrlaterhouse 1.f9I2) that there is no sharp he divided Conoqnatha character states in that taxon' Nevertheless intofourgroupsonjustthischaracterr'whilstconcedingthatthe 60 species are difficult to groups are contrived and that some intermedlate place least in I think that this structure is a useful one' at groups of Metaxymo rpha characterizing Cal-odema and the two species those groups' The because it is easily seen and constant within presenceofaprosternalprojectionisconsideredtobeaderived characteralthoughitsfunctionisunknown.Itaccentuatesthewedge shapeofthebody,possiblyenhancingitsaerodynamicproperties'o1 actingasastabilizingstructureinhighfliers.T.Moore(inlitt. 1987)suggestedthatConognatha(s.s.)usetheprojectiontoopenthe presence of a prosternal rotting fruits upon which they feed. If the projectionlsconsideredtobesynapomorphic'thenthosegloups possessingitvizCalodema,Metaxymorpha'Conognatha(s's')and thamaybemorecloselyrelatedtoeachotherthantheyareto prosternal projections the other taxa of the tribe. 0n the other hand, (Julodini) occulinotherquiteunrelatedbuprestidSlê.9. Sternocera hasapronouncedmeso.metasternalprojection.Moreover,itisnota complexstructurebutsimplyanenlargementoftheexoskeleton. plocess invoLve characters which are expressions of an altometric minimalgeneticchangetoproducesubstantialmorphological in such characters differences. Paratlelism probably occurs frequently phylogenetÍc analyses which should therefore be given low weight in (Hecht and Edwards, 1976) ' (A) mesosternum only' Char actet 24 Sternal cavity formed by : mesosternum divided which is entire; (B) mesosternum and metasternum, (l-893) the first (Stigmoderini); (C) metasternum onJ-y. Kerremans was an important character for to consider the form of the sternal cavity as stigmoderini in the group delineatÍng higher categories. He classj-fied 6T rn most ,sternal cavity formed almost entirely by metasternumr. along the cavity so stigmoderini, the meso-metasternal suture is halfway In some that the mesosternum and metasternum contribute equally' suture is basalr so Hvperantha (s. s.), Pithiscus and tha the it is that the cavity is formed primarily by the mesosternum, although his analysis of the always divided, never entire. Holm Q97Ð in sut'ure at buprestid tribe Jul-odini, asserted that meso-metasternal posteriorendofcavityisthepJ-esiomorphicstate,whj-Iesuture dubiously interrupted is apomorphic, and suture well behind cavity, suture would have apomorphic. I concur with Hol-m that primitively the been entire. than or equal character 25* sternal cavity: (o) length greater (1) much l-ess than width to width (all- taxa except castiarina); length the suture is Castiarina . The condition in Castiarina in which are short, is the widely interrupted and the mesosternal branches relativelY derived state' Character26scute]-IumsizewasmeasuredaSaveragediameter The scutetlum is (n) very relative to maximum width of the elytron' (B) sma1l' 0'1-0'2 smatl, up to 0.I width of elytron (Themognatha); Calodema, and width of elytron (Stigmodera (s' s'), Castiarina (Conognatha (s. s.) and Pithiscus ; (c) larger, o.2-o.' width of elytron than 0'l width of elytron Dact Lozodes ; (D) very large, greater The unusuaÌIy (Metax vmorDh a. Hvper antha ( s. s.), and Hyperanthella)' large scuteLlum is useful for characterizing Metaxymorphat shape of scutellum is Hvperantha ( s. s.) and Hyperanthel-ta, however, the too variable to be of use in most taxa'. 62 Characters of the elytra Fossilevidenceofbuprestide}ytrafromtheLowerMesozoicinthe was reported by Ipswich coaL Measures, Denmark HilLr QUeensland elytron of a new Etheridge and olliff (1890). They described the specj.esMesostj-gmoderatypicaaS||coveledwithVelylegularrowsof base to apex' the large shallow pits or punctures which extend from their interstices moderately broad, and closely covered throughout .'r and observed entire length with very sma]l el-evations or tubercles" to that of that the fossil elytron appeared to be most cLosely allied gratiosa and S' roei' except for Stiqmodera (s. s.) sanguinosa, also !. the elytron (P1' V' the interstitial elevations. Their illustration of Dunstan Q927) described fig. L) showed 15 complete serial rows. ttcylindlaceous formrr a another elytron of the same species as having a ' trdi-sc rows of ilmoderately hightt convexity, and a with about seventeen pitsrr. I question the placement of Mesostigmodera tvpica in Buprestidae' sinceitisbasedonsotittleevidence.Thedescriptionand illustration of the pits and smaIl tubercl-es seem to correspond more closelytotheelytraofextantArchostemataOmmathanto does represent a Stiqmodera ( s. s.). However, if Mesostigmodera typicalearlybuprestid,thenthefossi]-evidencesuggeststhat early in the evolution interneurs comprising regular rows of pitS arose of buprestids. 10 rows was probably crowson (1981) considered that punctures in theprimitiveconditioninbeetles.Furtherfusionand]'ossof several transformation interneurs is apparent in many stigmoderini and seriescanbeproposed.Interneursgandl0maybefusedmediallye.g. e'g' Meta s. Castiarina) skusei, fused on anterior half species,orinterneurgwithananteriorgapandfusedwithl0a}ong 6V antha ( s. s.), interneurs rest of length in many Dactylozodes' In Hvper half to one third length of the 7 and I are very short, terminating at ale elytron. In H. (Hyperanthella) sa]Iei, only eight interneuls evident. margin undifferentiated Char acter 27x Elytra: (O) convex' l-ateral edges roLl-ed over (most stigmoderini)i (I) flatter, costate, with Hyperantha (s' s')t forming a pronounced rim (Dactyl-ozodes' eranthel-la and some Castiarina). (O) completely separate Character 28 Interneurs: 10, (1) 9 + 10 partly fused (Stigmodera (s. s.), some Themognatha); ]0, with (mostStigmoderini)i(2)9,with9+10comp}etetyfused(some (1t) 10, with 7 and I short' fused Dactv lozodes . some castiarina); (2') I (some Hvperanthella)' It is posteriorlY H erantha (s. s.)); has arisen by the ross of unclear whether the situation in Hyperanthella In either case it is 7 + 8, or by the fusion of 7 with 8' 9 with 10' of states exists both inside the most derived state. since a continuum of limited use in determining and outside the tribe, this character is relationships,eventhoughthepolaritycanbedeterminedwithsome certaintY. (most Stigmoderini); Character 29* E]ytra: (0) punctate-striate (})withlarge,deep,lessnumerousfoveo]-es(Stigmodera(s.s.)).The (s. s.) in which the Iarge, often coalescent foveoles of Stigmodera sclerotized differ markedly fLoor of the foveole is thick and heavily fromthemembranous-f].ooredwindowtypeoffoveolesfoundinthe ArchostemataandDerodontidae(.consideredtobeoneofthemost primitivePolyphaganfami]-iesbyLawrenceandHlavaclllgTg)).The 64 to distinctive interneurs o1 Stigmodera (s. s.) are therefore considered progressive decrease in size and be derived. other stigmoderini show a primitive extant increase in number of pits compared with fossit and smooth elytron e'g' types of elytra. This trend towards a completely Calodema regalis is also considered to be derived' (B) Character l0 EIytraI api-ces: (A) subserrate; denticulate; (G) pointed' (c) bispinose; (D) trispinose; (E) rounded; (F) truncate; is not possible Given the variety of elytral apices within the family it the elytron of to assign a polarity to any given type, atthough I'obtusely (Dunstan, 1927256)' similar Mesostiqmodera had apex pointed" given the to that of Stiqmodera (s. s.) macularia. Indeed, intragenericdiversityofsometaxa,thevalueofthischaracterin margins occur generic diagnoses is questionable. However, dentículate (s. some Pithiscust only in the South American genera (Conognatha s'), and the elytral apices of Hvp erantha ( s. s. ), and Dactylozodes), The Hvper anthellaaredenticu]-atebetweentwoterminalspines. for distinguishing character, apices bispinose or trispinoser is useful þetween species groups of Metaxymorpha' (most Stigmoderini)i Character 3l-* Elytra: (O) not overl-apping (Metaxymorpha)' The latter (1) with one elytron overlapping the other unique within the character is considered to be apomorphic since it is Buprestidae' in tribe. The only other record of this character in Strandiol-a paradoxa'wasbyObenberger(192oz162)ascitedbyHoscheck (Ie3Iz773). Character32Elytra:(A)greaterthanlengthofabdomen (s. s.), some (Meta xvmorpha some Castiarina Hvperantha 65 abdomen' Conognatha (s. s.) and some pithiscus); (B) equal to length of or not exceeding it appreciably (all other taxa) ' Fossil beetles had elytra longer than abdomen (Crowson, 1975; Lawrence and Newton, 1982): however, it is likely that in some species with extended elytra e'g' c Castlarina ) producta, elongation of elytra is associated with the evoLution of a particular mimicking facies. No polarity is assigned here because of the difficulty of distinguishing between primitive states which have been retained and those secondarily derived. (B) Character 37 EIytra: (A) not setose (most Stigmoderini)i setose (Dact vlozodes. Hvoerantha (s. s.) and Hvperanthella). E1ytra with long, dense setae j-s one of the most striking characters of Dactvlozodes. The setae of Hvperantha ( s. s.) and HvPeranthel-la are j-s shorter and sParser. Conoqnatha ( Pithiscus) vul-nerata Perty an excepti-on to the above generalization, having elytra with long dense setae. Setose elytra occur sporadically throughout the family as well as in the primitive genus Omma (Ommatídae): the polarity of this character is uncertain. Character 74* Wing: (O) with wedge cell present (Julodimorpha); pointed wedge (1) wedge cell absent (a11 Stigmoderini). An apically cell is characteristic of Buprestidae, the generalized out-groupt like (Lawrence , Ig82)., and is present Ín primitive coleopteran families Cupedidae (Crowson, 198t). The absence of a wedge ceLl l-t Lot unique ( Astraeusini) toStigmoderini,however,becausethewingofAstraeus1and@ À (Buprestini) also lacks one. 66 Characters of the leqs posterior margin Character f5 Hind coxa: (A) dilated medially, posterior margin oblique (all Stigmoderini); (e) parallel-sided' The other genera of of hind coxa is more strongly angled Ín Calodema than in the tribe. and mid; Character 76 Hind tarsi: (O) slightly longer than fore (I) very elongate (1.5 x J.ength of fore). Buprestids typically have tarsi long, about half the length of the tibiae (cobos' 1980)' some Conognatha (s' s') Calodema and Metaxymorpha, some Themognatha and out-group, and have hind tarsi noticeabLy longer than the generalized this state is considered relatively apomorphic. It is not a character hind tarsi of high weight, however, because the relative length of the is a continuum in the tribe' (most Character f7 Tarsomeres: (O) I to 4 with large pulvilli pulvilli (some Stigmoderini)i (1) I to 4 compact, globular, with reduced two spines only, without Dact lozodes ; (I') I to 3 bearing one or (1857:56) pulvilli (mal-es of some Castiarina Lacordaire character, but distinguished Dactylozodes from Zemina solely on this character may be they were synonymized by saunders (1871:60). This more useful at the species group level-' tooth' Character fB* Tarsal claw: (O) simple; (1) with basaL by nTarSaI claw hooks l-obed and toothed at baseil was a character used In 74 Théry Ã92g) to separate Themognatha in a key to Stigmoderini' only 46 species of Themognatha examined for this character, however, (62%)hadaplonouncedtooth,]0hadaweaktooth,andintheremainder 67 be more useful the tooth was very weak or absent, so this character may at the sPecies group Level' Characters of the abdomen sexually Character j9* sternite 7: (0) not or scarcely (s' Castiarina dimorphic, male 59 not exposed (Stigmodera s')' Hyperantha (s. s.), Dactylozodes, Conognatha (s' s'), most Pithiscus Hyperan thetla);(1)sexuallydimorphic,maleSTnotchedexposing59 57 of male (Calodema, Metaxvmorpha . about hal-f Themognatha). the apex of 59' The Conoqnatha pretioissima is concave, just exposing notched a1so, but 59 is male of Hvperan tha ( s. s.) decorata ís weakly scarcelyexposed.BothofthesestatesoccuTinthegeneralized confidence' out-group, so it is not possible to assign polarity with a general evol-utionary however, it is argued that if differentiation is trend, sexual dimorphism is the derived state' suture: character 40 Female proctiger T9+J-0, oblique transverse (1) (0) entire (most stigmodera (s. s.), Pithiscus); partial Castiarina (conognatha (s. s.)); (2) not discernible, fusion complete The suture calodema, Metaxymorpha, Hyperantha (s. s.), Hvperanthella)' also in the outgroup' is variable in Themo atha and Dactylozodes, and most primitlve subfamily The 9th and 10th tergites ale separate in the fusion and subsequent of buprestidsrschizopodinae, and therefore complete The character is obliteration of the suture is considered apomorphic' givenlowweight,however,becauseitisacontinuuminthetribe. Theverydistinctive,heavilysclerotized,visibleproctigerof be derived: perhaps stigmoderoides group of Themognatha is presumed to A few species' those the scoop-Like structure assists in oviposition. group have very long setae on of Stiomodera (Themognatha) maculiventris 68 their terminal- segments. Long pilosity is uncommon in Buprestidae and is considered an apomorphic character' (most Character 4L* Male proctiger T9+10: (O) fusion incomplete (s. Stigmoderini); (1) fusion complete, no overlap (Conognatha s'))' character 42* Tergolongitudinal muscles (!rg): (0) T1-T6 (Stigmodera (s. Dact lozodes and Hvperan tha (s.s. ) ) ; (l) 1I, T4-T6 s'), Castiarina cal-odema, Metaxymorpha, conognatha (s. s.), Pithiscus, and (1975) Themognatha). AII the buprestids studied by Kasap and Crowson had tlg T1-T6, which is considered here to be the primitive state' The loss of tlg muscLes on T2 and Tl is associated with the fusion of the first four abdominal tergites. The state in Hyperanthella is unknown' Character 47* t:Ig T2t (0) fu]l height (Dactylozodes); (1) reduced (Hyperantha (s. s.)). This character is not available for Hvperanthel_la and not applicable to the other taxa. Character 44* 54-56 slIg: (0) absent (CastÍarina ( Conoqnatha ( s. s. ), Pithiscus DactyIozodes, and Hvperantha s. s.)); (1) present (stigmodera (s. s.), Themognatha, Metaxymorpha, and muscles on S7 in Cal-odema ) : HVperanthell-a is unknown. The absence of and 4' most buprestids is associated with the connation of segments f (]975), two Among the buprestoids studied by Kasap and crowson only s119, and species estÍs aurulenta L. and Jul-odis onopordi F' have outside Buprestoidea, sIÌg are recorded in only three species' Kasap (the being and crowson noted that sllg and sl-d rarely coexist exception Bupres tis) and suggested that s1d are possibly modified sIIg' 69 If Kasap and crowsonts suggestion is correct, the transformation would be as foflows: modification I smlg + sIlg smlg + s1d sIlg In this case, (smlg + sld) is the derived state. However, since alternative are uncommon in buprestids and retated families, an interpretation is that sl]g represent a differentiation of smlg' The transformation would then be as follows: differentiatíon ll + sld smlo+sllq+sId possibility In this case, (smlg + sld) is the primitive state. A third is that sl]g differentÍated from sld' differentiation I I smlg + sld smlo+sIIq+s1d (smlg + srd), the whatever the more primitive state, (smrg + srlg) or probably presence of three distinct muscle bands (smIg + slIg + sld) presence of sI]g represents the relatively derived condition' Thus the (s. Themognathat represents a synapomorphy uniting stigmodera 9.), Me and Calodema. (B) present Character 45 Tergal and sternal pores: (A) absent; The 12-T6, 54-56 (most Stigmoderini)i (C) present on other segments' occurlence of pores i-s variable in the Buprestidae' Kasap and Crowson 70 (:Ìg75) suggest that they may function as heat receptors' Buprestids Diadoxus ervthrurus (l'/hite) and D. scaLaris L. & G' attack fire damaged Cal]-itris(HadlingtonandGardner,Ig5g),andlhavefoundevidencethat discovered Stiqmodera porosa oviposits in burnt wood. Evans Q966) pits that the buprestid MelanophiLa acuminata DeQeer possesses sensory detect containing 70 to over L00 spine or hair-like sense organs which infra-red radiation and enable the beetle to ]ocate fire-damaged trees while they are still smouldering (Komarek, 1969)' Another buprestid' (Britton, 197O2565; Merimna atrata L. & G., is attracted to campfires pers. obs.). Crowson, in the newsletter Bupreltis (1985) suggested that the poles are associated with glandular function, probably glands defensive rather than pheromonal, but I have found no evidence of adjacent to the pores in Stigmoderini' Characters of the female repro ductive system general The ovipositor of stigmoderini is unique. A search of the literature as well as papers dealing mole specifically with buprestids dorsal (see Appendix D) reveaLed no mentj-on of a structure such as a (1975)' valve in any other beetl-e. Figures 2 and 3 in cobos illustrating the ovipositors of Microcastalia Ditriaena incerta Cobos fromMexicoandM.(D.)sexpinosa(|Vaterh.)fromCentralAmerícasuggest obtain that they may have dorsal valves, but I have been unabl-e to of material to confirm this. According to Cobos Q975) the ovipositor same This does Microcastalia (M. ) lobithorax Thomson is of the type' from that have a deeply divided dorsaL valve, but the ovipositor differs of Stigmoderini in that it lacks the conventional styles and possesses the instead a pair of long, nalrowr setose appendages dorso-l-ateral to style-bearingvalveandconnectedtoitonlyattheirbases. 7T The particle sac is often packed with charcoal, sand, or fragments of plants. Cobos ¡975), in discussing the ovipositor of Ditriaenat earth or noted that ovipositors of that kind are frequently packed with of sand which was indicative of oviposition in. the ground at the base the larval- host plant. Dodd (1912, L9I6), after observing sand in the served sac of Stiqmodera regia Blackburn considered "that the particles as ballast enabLing the beetle, when its body had become light from the loss of eggsr to fly steadily in the violent gusts of the hilly country female, which it frequentsrr. An alternative explanation is that the some of the when excavating a depression in which to oviposit, stores particles in the sac. Anteriorly directed setae on the ventral surface of the dorsal valve help to retain the contents of the chamber' After some of the eggs are 1aid, they could be covered over by the ejection of the particles, perhaps by hydrostatic pressure of the haemocoel-ic fluid' since there are no muscles attached to the particle sac itsel-f' I suggest that the short ovipositor of stigmoderini is a highly plant derived form, adapted to scraping a depression in the soil or host in which to oviposit. The presence of heavily scl-erotÍzed spatulate setae and particle sac packed with sand, charcoal or plant fragments supports this idea. This is in contrast to the typÍcal long, flat and membranous buprestid ovipositor suitable for probing ovipositing studies under bark and in crevices. Future ecological and behavioural of ovipositÍon may illuminate the function of the particle sac' Indivídua11y, the features of the ovipositor of stigmoderini are rruritiformrr less not unique in buprestids. short ovipositors of l-ength (Volkovitsch' than, or equal to, width occur sporadically in Acmaeodera 1979)' re78), Acmaeoderella (Volkovitsch, 1977; Volkovitsch and Bily, AgriLus and Sphenoptera (Volkovitsch, pers. comm.), Notomorpha and (Cobos' Xirosce]-is (Cobos, 1955); Bordonia (Cobos, 1980), Microcastalia 72 Ig78) and Ethon, Paracephala and Melobasis (pers' obs')' Spatulate setae have been illustrated in Microcastatia ( Ditriaena and Bordonia (cobos, 1980), and other coleoptera e.g. Neoclosterus 918 and Doracercus barbatus where they occur on sternites 4 and I respectively (wandolleck,1905, plate 28). I have already noted the presence of a dorsal vaLve in Microcastalia ([.) globithorax. Also, I have found sand particl-es retained in the ovipositor of Paracephala sp', however, in this case, the chamber is formed by a ventrally curved ventral valve with long mediaJ.Iy dj.rected, incurved hairs. The ovipositor of- Ethon also 1S similiar to that of Paracephala. Pockets containing humus have been recorded in the female terminalÍa of Elateridae (Calder, 1979275) ' Although the form of the ovipositor and the presence of dorsal valve, particle sac and spatulate setae are not unique in themselves, their occurrence together represents an important suite of synapomorphic characters that unites the genera of Stigmoderini and distinguishes it from all other buprestid tribes, providing a strong argument that Stigmoderini is monophyletic. Smaller differences in the components of the ovipositor provide useful generic, subgeneric or species group characters. The question arises of how a structure such as the dorsal- valve coul-d have evolved. V'/hen I dissected Schizopus laetus, I was struck by the presence of a membranous foLd between the proctiger and genital tube, which, by a process of elongation and sclerotization, possibly couLd be transformed into a dorsal- valve. This fold is also present to a much lesser extent in Dystaxia murrayi and Glyptoscelimorpha marmorata, two otherSchizopodinae,as well as other buprestids e'g' Catoxantha opulenta as illustrated by Wandolleck (1905). Two al-ternative pathways can be postulated for the evolution of the buprestid ovipositor. Either the ovipositor of the typical buprestid FIGURE 24. Schema of one possible path of evol-ution of the buprestid ovipositor from a Dystaxia like ancestor (a) - 'bY elongation and increasing sclerotization of the membranous fold (arrowed) to form a dorsal valve, (a) - (b) - (c); or elongation and decreasj-ng sclerotization of style-bearing and ventraL val-ves, with the loss of the membranous fold, (a) - (d) - (e). (a) Dystaxia murrayi, (b) Schizopus laetus, (c) Stigmodera spp. r (d) Cisseis sp., (e) Buprestis aurulenta. Membranous areas are indicated by shadÍng. Abbreviations: EP = epiproct, pp = parêproct, R = rectum, VU = vulva. (c) +VU (b) R Development of dorsol volve çVU lncreosing sclerotizotion (o) +VU Decreosing sclerotizotion (d) PP Elongotion of volves (e) PP VU 73 represents the primitive state with the Dystaxia type an early transformationleadingtotheStigmoderineform; or the Dystaxia ovipositor resembles the ancestral form of the buprestid ovipositor' as ill-ustrated in Figure 24, with the typical buprestid ovipositor representing a reversal to the primitive coleopteran type. In either case, the ovipositor of stigmoderini is highly derived. From a basis of this framework, the variations observed within stigmoderini can be assigned polarity. The plesiomorphic states are: valves and proctiger short, weakly sclerotized; dorsal valve weakly valve, notched, setae short; particle sac short, not exceeding dorsal and unilobed; septum absent; and spatul-ate setae few. The corresponding apomorphic states are: val-ves and proctiger elongate, increasingly sclerotized; dorsal valve deeply bifidr setae long; particle sac proglessively longer, increasing the volume of the chamber, spatulate and bilobed; septum present and increasingly sclerotized; setae numerous. Character 46* Ovipositor: (O) long, flat, membranous, with 2 valves, vulva terminal (most buprestids); (1) shortr lightly (It) sclerotj-zed, with J val-ves, vulva ventral (a11 Stigmoderini); long, subcylindrical, heavily sclerotized, with 2 valves, vulva terminal (Julodimorpha) . Character 47* Dorsa] vaLve: (0)'scarcely divided (Hyperantha (s. s.), Hyperanthella, most Dactylozodes); (1) deeply divided (the remaining taxa). Character 48 Particle sac tength: (O) short, not exceeding va1ves (Stiomodera ( s. s.), Conognatha (s. s.), Pithiscus, 74 valves Hyperantha (s. s.), Hyperanthelfa); (1) long, exceeding anteriorly by about one third their length (calodema and Metaxymorpha); length' (2) very long, exceeding valves by more than one third their where it may It is variable in Castiarina , Dactylozodes and Themognatha makes it a be up to twice the length of dorsai valve. Its variability character of Low weight. (0) absent character 49* sclerotized septum in particle sac: (1) present (the (Hyper antha ( s. s.), Hyperanthell-a, Dactylozodes); character' remaining taxa). This is an important generic or round, Character 50* Bursa copulatrix: (0) unilobed, elongate (Themognatha' Castiarina spermatheca + duct longer than bursa Metaxymorpha, Conognatha (s' s'), most Pithiscust longer than bursa (some Hyperanthelta); (1) bilobed, spermatheca + duct PithÍscus); (lt) trilobed, spermatheca + duct shorter than bursa of the Stiqmodera ( s. s.)). A unilobed bursa is characteristic generalizedout-group,andthusispresumedtobeplesiomorphic. A species of long spermatheca and duct curLed distally is found in and Buprestis' Melobasis Ethon, Ast¡eeus, Merimna, Cisseis bursa found in curis' spermatheca more robust and shorter than the is and Acmaeodera' The Julodimorpha, chrysobothris, Dj-adoxus, Nascioides (s. are treated trilobed bursa and short spermatheca in stigmodera !.) as a single character as these are correlated' (s' character 51 0varioles/ovary: (A) 5 (Stigmodera s'), (B) 7 (Themognathat Dactvl-ozodes (except undescribed sÞ.), Castiarina); M. meeki has 5' A CaIodema). Meta a grayi has 7 or 8, (s. and tentative count of 4 or 5 was made in conognatha s') 75 number of ovarioles per ovary in eranthelLa. Appendix D, l-ists the 21 genera in 15 tribes' Within 28 other buprestid species representing 4 in Astraeus and the Buprestidae, the lowest numþer recorded is ova¡ioLes is stable in many highest 22 in Buprestis' The number of genela'althoughdifferentcountshavebeenobtainedfordifferent Iozodes as weII as for species of Curis, Metaxvmorpha, and Dact Julodimorphabakewe].li.Thereisnoindicationofwhatwasthe ancestral number. (0) absent (most Stigmoderini)i Charac ,er 52x Accessory gland: apicalis group). (1) present (Themognatha, Ca]-odema, and Metaxvmorpha accessory glands in female Heberdey (I9r1) summarized reports of he placed the coleoptera. Based on the position of the opening, that emerge on the accessory glands in three groups: those 8th and 9th sternite; those that intersegmentaL membrane between the forminvaginationsíntheareaofthevulva;andthosewhichopeninto been reported in a number of the vagina. Paired vaginal pouches have (198') who assumed that these organs chrysomelídae by Mann and crowson symbiotic bacteria' although the ale concerned with the transmission of ol even looked fort authors gave no evidence of having found' in a Limited sample symbionts. Although the failure to find symbionts more likely that the accessory is not conclusive evidence, it seems gland in Stigmoderini has another function' previously in severar other col_reteriar glands have been recorded (crowsont 1981) Elateridae (Becker' families of beetles: chrysomelidae ' coccj'nellidae and possibly Cantharidae 1956; Williams , L945), carabidae, (Wi1liams,1945),butneverbeforeinBuprestidae.Inmostofthese reports,pairedglandsinsertonthemedianoviductordorsalsurfaceof the vagina. 76 the Martjoa Q964) and Mateu and Martjoa (1964) noted that in glands, the buprestid Sterapsis speciosa which Lacks colleterial secreting oviducts and ovarioles have taken over the function of glycogen, varÍous protei-ns, calcium oxalate crystals, ald small amounts glandular tissue must be of mucoPolYsaccharides. In St the while in mesodermal in origin since it is associated with ovarioles, Stign,oderini, it is ectodermal' Thesecretionisprobablyassociatedwithsomeaspectof oviposition e.g. an eggshell which hardens on contact with air to to protect against water Loss or attack by bacteria or fungi; a toxj-n so that they discourage predators; an adhesive to make the eggs sticky the eggs to a acquire a protective covering of particles, or to cement substrate or to each other; a trophic substance; or a tropho-stimuLant to encourage newly emerged larva to eat the eggshell' that other functj.ons of the secretion could be postulated, e'g' it from the is a digestive enzyme which helps to release the spermatozoa spermatoza from the spermatophore, or involved in the translocation of spermatophoretothespermathecaasdescribedinLyttanutalli (Meloidae) by Gerber, Church and Rempef (1971). However, this would not explain the absence of the gland in the other taxa which have structure' simil-ar spermatophores. As the accessoly gland is a complex buprestids' its absent in most coleoptera, and not reported in other presence is an apomorphic character of high weight' Characters of the male reproductive system (O) (10-12); (t) hign Character 57* Number of tubules/testis: low (2o-5o); Q) verY high (so+¡' Castiarina Dactyl-ozodes and Pithiscus (t. have a high number have a l-ow number of tubules, most stigmodera t') (20-50)(excepts.gratiosawith12),andmostThemognathaexaminedto 77 (except (T') date have a high to very high numbet (2o-94) s' varrelli with 12). crowson (1981:l-07) suggested that a single tubule per testis, as occUIS in three of the four suborders of Coleoptera may be the ancestral type, but he notes that the number and form of tubules differs greatly in the Polyphagä. 0f the nine other buprestid genera studied (Appendix E), eight have 12 or fewer tubules per testis' Although the sample is very small, I suggest that in Buprestidae the primitive condition is a l-ow number of tubules per testis' Furthermore, I postulate that 12 tubules per testis is the ancestral one member number in stlgmoderini since that numbel occuls in at l-east In Castiarina of each of the higher taxa for which data is available' but at most species have the reduced number of l0 tubules per testis, (s. and least I species retain the primitive number' In Stiqmodera s.) tubules per Th tha most species have undergone an increase in testis, but at least one species of each taxon retains the primitive number of 12. Julodimorpha with an avelage of about 200 tubules pel testis of represents the most advanced state in buprestids. A large number and a very short tubules may be associated with high sperm production aLso be adult mafe life. A large sperm production per individual may correlated with the observation of a marked sexuaL imbalance in populations of Jul-odimorpha, males often outnumbering females 50 to one or more (S. Barker, pers. comm' 1982; pers' obs')' character 54 Male accessory g)-ands: (0) one pair (all stigmoderini); (1) two pairs (Julodimorpha)' Julodimoroha is the onL v mal-e in buprestid reported so far with two accessory glands in the addition to the spermatophoral gland, so this character is considered 78 apomorphic.Therelativelengthofaccessoryglandlistoovariable stigmoderini. to be of use in discriminating between genera of (A) and cl-ub-shaped' character 55 Spermatophoral- glands: large (B) The buprestids curved ventrally (aII Stigmoderini); otherwise' of forms' described by Kasap and crowson (1975) have a variety membranous flap (most Char acter 56* Median Lobe: (0) without covering ostium Stigmoderini)i (I) with large ventral membranous flap (Hyperantha (s. s.), Hyperanthellat one species of Castiarina [s.Barker,pers.comm.1986],andundescribedDactylozodessp')' the tribe' It The shape of the aedeagus is highly variabl-e within perhaps in the smaller therefore has 1imited diagnostic value except and in taxa where some features are constant and distinctive, such as castiarina' determining species groups within the larger taxa Characters of the aI svstem crop; (I) with Character 57* Foregut: (0) without diverticula or (Julodimorpha)' lateral- diverticula (a11 Stigmoderini)i (1') with crop are uncommon This character is unknown for Hyperanthell-a' Diverticula beetles in Coleoptera and recorded only in fl-oricolous TJånnba""ti" are known only in (Crowson ISBI:166). l;itnin the Buprestidae they Acmaeodera Sp.and@sp.(KasapandCrowson,I975),where,itis The enlargement of the suggested, they serve as a food reservoir' recorded in Julodis posterior end of the foregut to form a crop has been Since diverticula and crops and Acmaeodera (Kasap and Crowson, L975)' and are án elaboration are usually absent in the generalized out-group of the foregut, their presence is presumed to be apomorphic' 79 (O) (more than 0'1 length of Character 58* Midgut caeca: long (Less than 0'1 length of midgut) (most stigmoderini)i (1) reduced the onty South American midgut) (Stigmodera (s' s'))' Dactvlozodes is genusforwhichthischaracterwasavailable.Thecharacteris variableinthegeneralizedoutgroup.0fthespeciesstudiedbyKasap have short caeca (fess than 0'1 and crowson (1975), Agrilus and Trachys and Nascioides have tength of midgut); Acmaeodera Julodis, Blgestis longercaecaC]/6 coI/Tlenglhofmidgut).Itispossiblethatthe midgutcaecaaleassociatedwithmaintena¡ceofsymbioticbacteria,at ].eastinwood-boring]arvae'aSleportedinsomeChrysomelidaebyMann length in the nectar-feeding and Crowson (1981). Reduction of caecal presumed to be an aduLts, which would not require symbionts' is apomorPhic trend. and caeca: (A) dense' Character 59 Regenerative crypts of midgut (B) unevenly distributed' spalser evenly distributed (a]] Stigmoderini)i posteriorlyorarrangedinrings.ThischaracterisvariableÍn (19811.166) observed that buprestids (Kasap and crowson, Ig7Ð. crowson activelyfeedingadultstendtohavelonger'moreconspicuous regenerative crYPts. length than midgut character 60 Hindgut: (A) equal or greater in (allstigmoderini);(B)shorterthanmidgut.Inalfspeciesof (1975), except perhaps Buprestis, buprestid studied by Kasap and crowson gut' the midgut formed the longest part of the (O) proximally Character 6l-* Malpighian tubules: ungrouped (C') pa tlidiventris (Stigmodera (s. s.), most Castiarina except S' Dactvlozo¿es); (1) grouped L. & G. and S. (C.) erubescens Blackburn' and BO tubules proximalty (Themognatha, Calodema, Metaxymorpha)' Malpighian but this needs appeared to be ungrouPed in C' Pithiscus viridiventris (s' s') to be confirmed: the character was not available for Hyperantha (1975) different or Hyper anthella. Kasap and crowson reported four From theír arrangements of the Malpighian tubutes in Buprestidae' findings together with additional species examined in this study (Appendix F), it is apparent that the arrangement of Malpighian tubuLes is stable at generic level, but varies within tribes. Regular spacing is primitive in beetles (crowson 1981:82), and independent attachment is characteristic of Buprestoidea (Lawrence, 1982). It is therefore considered that Malpighian tubules grouped proximally is a synapomorphy of Themo tha , Cal-odema and Metaxymorpha. character 62* scl-erotized rods anterior to junction of rectum and hindgut: (O) absent (most Stigmoderini); (1) present Dact Iozodes I have [except D. rousseliJ, Hyperantha (s' s'), and Hyperanthella)' laetus and found anterior rods in only two other buprestids: Schizoous a separate Glyptoscelimoroha marmorata bothSchizopodinae(considered a species of family by some workers), although they were absent in third by Kasap and that subfamilYt Dystaxia murrayi. Nor were they reported presumed to be Crowson Q975). In this study, anterior rods are in the apomorphic because their absence is the prevaÌent condition generalized outgroup. Crowson (198] zL77) suggested that the sclerotizations of the hindgut provide purchase for muscles to expel hard faeces, which would be an adaptation to an arid environment' (1) fused Character 6f Nervous system, ganglion 5: (0) separate; (Stigmoderini). Of the species examined by Kasap and Crowson (1975) stigmoderini is and myself, only Buprestis has ganglion 5 separate, so o- P. an character /É IDAC .t= tDo= D0) sÐsgHustsùìàÈùsÈssHÐu\uNÈGEßo\ -toH FoooHoHoooooooH Stiomodera õ r< ootsPoqoTHHtsH c+ ctO F Ðo{, -È,.)J(l)oO HoqqHoooooclooo|- Themoonatha F-0) oHoHo?tsoHFH}J orHr0) N HH FØO OOcf(I)o r-J ¡ Hcro oHooqHoo qqoq Castiarina c+A ooo,-o o oo H,-¡-o v)(Cto HHPts c+A P.o aoo -r, O- H(fHOHOOOONOOOOH Cal-odema |-.HP.c)= oPoPo'otsotsHHP rD o- c+ }.' 1õ fi)an= uto not unique in this character. The evolutionary trend is towards fusion of the ganglia. 3 PHYLOGENET IC ANALYSIS This section presents hypotheses relating to the monophyly of the tribe and each of its higher categories based on suggested apomorphic characters described in the preceding section and summarized in Table 7. Relationships between taxa supported by synapomorphies are proposed, and synapomolphies which suggest conflicting relationships are di.scussed, leading to the establishment of a tentative cladogram' I of the tribe The monophyly of the tribe stigmoderini is supported by a suite of apomorphic characters assocÍated with the ovipositor: the short' sclerotized form with dorsal valve, particle sac, spatulate setae, and medial styles directed posteriorly. This combination of characters provides a strong argument that the grouping is natural because it is present in every species of Stigmoderini studied here and has been found in no other buprestid examined by me or reported in the literature' Nor have I found any report of such an ovipositor in any other famiJ-y of beetles. The development of a dorsal valve and particle sact presumably as an outgrowth and elaboration of the membrane connecting the proctiger to the genital tube is clearly apomorphic. The presence of a pair of foregut diverticula, and lack of a wedge cel] in the wing are other synapomolphies of stigmoderini. Individually, they are not unique in the family, but they may be unique when considered together: of the two buprestid genela in which I have noted the absence of a wedge celL Ín the wing, Neocuris has a single 82 1argecIop'[email protected] the mouthparts' apomorphic character of the tribe, the prolongation of in the although used as a diagnostic character, is given low weight phylogenetic analysis because it is not constant, and is not unique to since it occurs in genera which on other evidence do not belong Stigmoderini. curis is property excluded from stigmoderini in which it is pJ-aced where is by Théry Q929), and probably belongs in Melanophitini it placed by Cobos (1978). The ovipositor of Curis is of the typicaL buprestid form lacking a dorsal valve, particle sac and spatulate setae. It j-s, however, exceedingly long, equal to the length of the evolutionary abdomen, which represents an apomorphy in a different direction from that of stigmoderini. The styles are medial and directed latera1lY. per other autapomorphies of curis are reduced number of tubules testis (six) and a male karyotype of 2n = 14 (6AA + Xy)' Prolonged Presence mouthparts are also derived and convergent with stigmoderini' in of six, eleven or thirteen ovarioles per ovary is another character polarity. The which curis differs from stigmoderini; it is of unknown midgut caeca midgut is short, only about half the length of the hindgut, tubules are are long, about I/7 Iength of the midgut, and the Malpighian and constricted ungrouped proximally and distal]y. The bursa is short at the point of insertion forming a short neck. The spermatheca and are duct are very short, much shorter than the bursa. Paratergites fu1] present, and abdominal musculature is smtg + sld s4-S6; tlg TI-T6 the height and width. A lateral prolongation of the abdomen conceals pointed wedge metathoraic epimera. The wing has a closed, apically ce11. 8V Julodimorpha should also be removed from the tribe, because it lacks the stigmoderini type of ovipositor. Julodimorpha is distinguished from stigmoderini by a number of characters which warrant ptacing this taxon in a tribe of its own. These characters incfude the following (those marked with an asterisk are considered autapomorphic): cylindrical body shape; mandibles cylindrical, rounded apically; Iigu]a elongate, bifid or notched apically; antennal cavities round, placed frontally between the eyes; frons narrowed at the l-evel of antennaL insertions; *eyes sexually dimorphic, those of the male being larger and 2 + 3 + 4 more protruberant; antennal scape very ]ong, equal to segments equal or together and 0 .25 total length (cf Stigmoderini with scape greater than the following two together, but less than 0.25 total length); antennal teeth triangular, with sensilla in a terminal, shallow depression, and in a large marginal secondary Sensory area whÍch wraps around both faces of the tooth, including segment Il; prosternum shortt length less than diameter or procoxae anterior to them; intercoxal process unilobed or scarcely trilobed, process not nearLy fitling the sternal cavity; meso-metasternal suture entire and raised; wing venation with closed, apicalJ-y pointed wedge celI; *tergal muscles tmlg + tpd + tug on T2-T6 (the only buprestid recorded to date with both tergoparamedian divergent tpd and tergolateral longitudinal tllg muscles); *sternal muscLes sld full height (cf in Stigmoderini and all buprestids studied by Kasap and crowson (1975), sld are short); *ovipositor long, cylindrical, with a very heavily sclerotized, medial, erect disc protruding beyond the posterior margin, and bearÍng small, oblique styles, two val-ves only present, and vulva terminal; 9 to 12 *two ovarioles per ovary; female tergite 9 without setae; pairs of (cf accessory glands in the male in addition to spermatophoral glands by Kasap one additional pair ín stigmoderini and aJ.I buprestids studied 8lr per about 200' and Crowson [t975]); *very high number of tubules testis' buprestids; which greatly exceeds any number previously recorded for parameres very Iong, narlow, straight-sided and flat; median Lobe similar, with sides curled inwards apicalIy, ostium well towards the apex; midgut caeca very long and thin (0.4 length of midgut); regeneratlve crypts decreasing in size towards the pyloric valve; midgut)' Malpighian tubules ungrouped; foregut long (0.7 length of taxa: Thus Stigmoderini is comprised of the fol-lowing higher Metaxymorphat Stiqmodera (s. s.), Themognatha, Castiarina Cal-odema, ( and Conoqnatha ( s. s. ), Pithiscus Dact lozodès Hvpe rantha s. s.), Hvperanthel-Ìa. Having established with a reasonable degree of the structure confidence, that the tribe is monophyletic on the basis of generic and of the ovipositor, phylogenetic analysis of the subgeneric, supra-generic groupings can proceed' l-v of the qenera and subqenera presumed The monophyly of stigmodera (s. s. ) is supported by three the autapomorphies: the trilobed bursa and short, stout spermatheca; short midgut caeca; elytra with large foveoLes often coalescing, resulting in the Loss of intervals. A reduced number of antennal another sensÍLla and reduction or absence of secondary sensory area is possibleapomorphythoughnotnecessari}yunique.Theshapeofthe the parameres, the shape of the median lobe and position of its ostium, (s' s') form of the Labrum etc. are other characters in which Stigmodera dilfers from other taxa in the tribe, but the polarities of these are not known. supporting the I was unable to identify any autapomorphic character 76 species of monophylY of Themoqnatha. Nevertheless, most of the beLonging to this taxon Themo tha I examined are easily recognized as 85 to suggest that by their overall facies, and I found no evidence taxon' Some characters' Themoqnatha should be lumped with any other (50+) of tubul-es e.g. tarsal claw with basal hook, and very high number pertestisrarearguablyderived,butarenotpossessedbyall species.Manyofitsdistinguishingchatacters,e.g.sma]-Iscutellum' areofunknownpolarityatpresent,butmaybeshowninfuturestudies there is evidence of to be aPomorPhic. Within Themoqnatha however, (!.) reichi group characterized by some natural species groups e.g. s. femaLe proctiger; the heavity sclero tized, strongly convex slipper-like by long marginå] and the S. (T.) maculiventris group characterized pilosityonthefemaLeproctigeranddorsalvalve'andelongate valvifers. by the autapomorphy of The monophyly of castiarina is supported branches reduced' sternal cavity strongly transverse with mesosternal wider than the two lateral lobes and medial Lobe of intercoxal process together.0fitsotherdistinguishingcharacters,smallsizeisnot lack of frons hair' or unique, and most are of unknown polarity e'g' presumedplesiomorphice.g.]-ownumberoftubulespertestis.Some charactersaleapomorphicbutdonotoccurina].lspeciesandarenot proJ-onged (secondary reduction)' unique in the tribe e.g. mouthparts not the number of toothed antennal mal-e tarsal pads reduced, reduction in segments or in the length of the antennae' ThemonophylyofCalodemaisnotestablishedclearly.Mostofits projection' anterior distinguishing characters e.g. acute prosternal marginofpronotumprojecting,smallscutellum,andpronoto-elytral articutationstronglybisinuateetc.arenotuniquetothegenus they are unique and individuaJ_Iy. As a suite of characters, however, presentareadilyrecognizab]-efacies.Theposteriormarginofthe than in other generat hind coxa is noticeably more oblique in calodema 86 of high but as part of a continuum is not considered a character weight.0nthebasisoftheshapeofthepronotumandother distinguishing characters, three species groups are suggested: C.regalisL.&G.,C.ribbeivandePoll,andC.p]-ebiusJordan. The monophyly of Metaxymorpha is supported by a single represents a simple autapomorphy, overlapping elytra. Although this constant in the transformation of doubtful adaptive signifj-cance, it is Principle, the genus and unique in the tribe. According to the Darwin habits, the more less any part of an organism is concerned with special characters are important it becomes-for classification, since adaptive morelikelytobeconvergent.Overlappingelytraaretherefore o1 the taxon is accorded high weight. Another distinctive character theverylargescutellum.Twospeciesgroups'M.meekiParryand M.apica]-is(vandePolI)aresuggestedandaredistinguishedbyasuite of characters, although the placement of the nominate species with each M. grayi Parry is prob}ematical, sharj-ng some characters groups the plesence group. The most striking difference between the is orabsenceoftheaccessoryglandwhichisdiscussedlater. by the complete The monophyly of Conognatha (s. s.) is supported fusionofthemaleparaproctandepiproct.Thisgroupisalso surface of the characterized by transverse scl-erotization on the ventral and style-bearing valve. The prosternal expansion is not unique differs in degree within the taxon' pithiscus is a diverse group of species and no autapomorphy was One found to support the monophyly of this group as a whole' in the character, bilobed bursa, is apomorphic and not found elsewhere taxon. The tribe, however, it is not shared by al] species in this found no synapomorphy maÌe proctiger is relatively plesiomorphic and r to support its incl-usion in Conognatha' 87 characters, of which Dac tvlozodes is characterized by a suÍte of only autapomorphy I anterior margin of mentum notched laterally is the found.Pronotumstronglysculpturedwithposteriorlateralandmedial toothed antennal depressions and explanate rim, reduced number of 9 and ]0 fused segments, compact tarsomeres, and elytraÌ interneurs characters' but completely from the humerus, are all presumed apomorphic The longt dense occur in only some species and to varying degrees' pilosity of the dorsal surface, characteristic of this genus is of unknown PolaritY. by Hyperantha (s. s.) and Hvperanthella are characterized distinctivebodyshapes,butacharacterasvariabtewithinthefamily Hyperantha (s. s.), as body shape cannot be assigned a pol-arity. In 7 and I is an the reduction in length of the elytral interneurs possessed by some apomorphic character as is the prosternal projection the relatively species. The elytra of some Hyperanthella represent HVPEr antheLla mostapomorphicconditionwithonlyBinterneurs'andsome possess an unsclerotized septum in the ovipositor' some of the genela Although monophyly has not been established for been found' The and subgenera, neither has evidence to the contrary monophylyt but apparent absence of an autapomorphy does not disprove .may As each of simply reflect the workerrs inability to identify one' they are these groups can be recognized by a suite of characters, which examj-nes treated as discrete units in the following discussion their synapomorphies and the reLationships they suggest' Supra- qeneric qroups Thefirstdichotomywithinthetribeappearstobethe Dactvlozodes lineage versus the rest - HVp erantha ( s. s.) - Hyperanthella (s' s') (Figure 25). The monophyly of the Dactyl-ozodes - Hvperantha - gnaÈha- S ti gmodera-Themo Dactvlozodes Castlarlna-Calodema- gwoerantha (s. s.) Metaxl¡mo rpha-P ithi s cu s- (s. s.) sclerot. rods mouthParts anterior to prolonged hindgut,/recÈum muscles antennae !þ short orl T2, T3 lost dorsal valve reduced I divicled toothed deeplY anten. seg. elytra with scleroÈÍzed ri¡rs septum ctivicling lateral particle sac wing lacks wedge cell foregut divertlcula present ovipositor with dorsal valve, Particle sac¡ spatulate setae, a¡¡d ¡nedial styles FIGURE25.Ctadogramshowingthetwoprimalylineagesof Stigmoderini.InFigures25-29,So]-idlectangles indicate the acquisition of a re]-atively apomolphÍc state; incompleteshadingindicatesthatsomeSpeciesretainol haverevertedtotherelativelyplesiomorphicstate;and stripesindicatethatspecieshaveanintermediateState of a multistate character' B8 the presence Hyperan thella lineage is supported by four synapomorphies: of six sclerotized rods anterior to the junction of the hindgut and rectum; the lateral margins of the elytra rolled over to form a distinct rim; shortened antennae; and the reduced number of toothed antennal segments, five in Hyperantha and five or fewer in most Dactyl-ozodes' Regarding the antennal characters: within Castiarina. number of toothed segments, but 1.. (C.) adelaidae group has a reduced obconic antennae of normal length; S. (C.) trilasciata group has three group has both segments but shortened antennae; and S. (9.) flindersi (S' Barker' reduced number of toothed segments and complessed antennae reduction in pers. comm. 1985). These three conditions suggest that have the number of toothed segments and shortening of the antennae arisen independently in Castiarina and that the similarity to the Hyperanthella antennae of the Dactvlozodes - Hyperantha (s. s.) - lineage is due to homoP1asY. In addition to the synapomorphies listed above, Dactylozodes - of Hyperantha (s. s.) - Hyperanthella share several- other characters than or uncertain polarity: elongate parameres (length/width greater equal to 7.5); labial and maxiLlary palps with terminal segments the pointed; and setose elytra. Finally, Cobos Qg>g272), noting parameres without sensoly hairs in D. spitzi, suggested that this constituted, up to a point, a link between Dactylozodes and Hvoeranthella. In The monophyly of Dactylozodes has already been postulated' is the support of the monophyJ.y of Hyperantha (s. s.) - Hyperanthel-la structure of the median Lobe of the aedeagus which has a large a simple membranous ventral flap. Although this character is (198I)' innovat.lon, i,t can be argued, following Arnold that it represents a special case for according high weight because the 89 efficiencyofgenitaliadependsnotsomuchonanyparticular morpho}ogy,butonintraspecificrecognition.Onceanewgenitalic featureariseslsâY'aSaresultofpJ-eiotrophiceffects,andthe oppositesexaccomodatesthechange,thenitislikelytopersistin descentlineagesbecauseofalackofselectionagainstit. also shared by other characters of unknown polarity which are incl-ude the shape of the toothed HVp erantha ( s. s.) and Hyperanthella transverse; mandibles antennal segments, rounded, and very strongly squat,almostaswideaslongwithtwoorthreesharpsubapicalteeth; proctiger notched medial}y; ]-atera} angles of the pronotum acute; female the base' The tlg muscLe and parameres strongly recurved ventrally at (compared with Dactylozodes in which on 12 short in Hyperantha (s. s.) so it may be an it is long) was not avail-able for Hyperanthelta, synapomorphy of autapomorphy of Hyperantha (s' s') or a Hvperantha (s. s.) and HvPeranthella' Themognatha Castiarina The monophyly of the Stigmodera (s' s') - - (s. s.) - Pithiscus lineage is - calodema - Metaxymorpha - conognatha with a deeply divided supported by four synapomolphies: ovipositor longitudinally dividing dorsal, valve; presence of a sclerotized septum theparticlesac;Iossoftlgmusclesfromtergites2andT;and mouthpartsprolongedintoarostrum.Thisgroupingisalsosupported eyes' by characters of unknown polarity e.g. large conflicting with this hypothesis are the following observations' onespecies,Dactylozodesalternans,hasadeeplydivideddorsalvalve' butlacksaseptum.Thedivisionofthedorsalvalveisasimple this species' some derivation which may have arisen independently in CastiarinaandmanyPithiscushaveshortmouthparts.Thiscouldbe state, or a Îeversal' as explained as the retention of the plesiomorphic mode of the ancestor those species groups changed from a nectar-feeding 90 ofthatlineage,toleafandpetal-chewingmodeoffeeding.Ifdueto of those species a reversal, this character would be an autapomorphy possessingit.Ithinkthattheargumentssupportingthetwomain evidence against them' lineages expressed above are stronger than the Withinthissecondmentionedlineage,themonophylyofthe Metaxvmorpha group is Stiqmodera ( s. s.) - Themognatha - Calgdema ' presence of sIIg muscles in supported by the synapomorphy of the character' increased sternites 4-6 (Figute 26). A second apomorphic mqny, but not all number of tubu}es per testis, is found in character needs to be Stiqmodera (s. s.) and Themognatha' This Metaxymorpha are confirmed, however, when data for calodema and available. chatacter, Against this proposed grouping is the apomorphic increasednumberoflobesofthebursa,whichsuggestsalinkbetween have a bilobed bursa' pithiscus and Stigmodera (s. s.). Some Pithiscus bursa (although that of and most Stiqmodera ( s. s. ) have a trilobed s.cancelrataisscarcelyso)rhoweverrtheshapesofthebilobedand bursa Looks as if it tril-obed bursae are quite different. The bilobed of an elongate uniLobed may have been derived by longitudinal division form;whilethetrilobedbursalooksasifitmayhavebeenderivedby Therefore' these l-ateral outgrowth of a very short unilobed form' in the two taxa' derived states may well have deveroped independently Theanteriorattachmentoftheprementumtothementuminwhich (s' s')' and some Pithiscus Stiqmodera (s. s.) resembles Conognatha cannot be assigned PolaritY' an unresolved polychotomy The suggested cladogram at this point has (Figure26).Nosynapomorphieswerefoundtosupportacloser relatior¡shipbetweenanytwoofthesegloupsalthoughtheclose (s' s') and similarlty in shape between the maLe genitalia of Conognatha South Àme¡ica ? Austlalia & Nee, Guinea Pithiscus high # tubules,/ testis bursa male sternal bilobed proctiger cavity fusion transv. , sIlg courplete mesost. Present sho!t s4-s5 mouthpartE prolonged !þ muscles on T2' T3 lost dorsal valve deeply divi.ded sclerotl,zed septum dividing particle sac FIGURE 26. Postul-ated relationships within stigmodera- T hemogna tha-c astiarina-c alodema-Metaxvmolpha-Pithiscus- Conoqnatha (s. s.) lineage. 9T Pithiscussuggestthatthosemaybemorecloselyrelatedtoeach with other. Neither did I identify any synapomorphy to link castiarina group' the Stiqmodera ( s. s. ) - Themognatha - Calodema - Metaxymorpha assertion that the This means that no evidence was found to support the group' Australian component of stigmoderini is a monophyletic A close relationship between Themognatha, Ca]-odema, and Metax ha(Figure27)hasnotbeensuggestedpreviously,butthe presence monophyly of this group is supported by three synapomorphies: proximally; and sternite of accessory gland; Malpighian tubul-es grouped This 7 sexuaLly dlmorphic, notched in the male exposing sternite 9' do not Last character must be given low weight as many Themognatha with male possess it, moreover, there are a few other stigmoderini sternite 7 excavated. The grouped Malpighian tubules may not be an ì¡/as unavailable for some autapomorphy of this J-ineage, as this character found of the south American taxa and two exceptional Castiarina were (s. (c.) pallidiventris and s. (c.) erubescens). Another derived Conognatha (s' s')' character, elongate hind tarsj-, al-so occurs in some seven ovarioles per ovary is a character shared by Themognatha - character is Calodema Metax however the polarity of this uncertain. stigmodera (s. s.) with three autapomorphies is the rpha proposed sister gloup of the Themognatha - Calodema - Metaxymo lineage. The several- arguments can be marshalled against this hypothesís' presence of the prosternal projection suggests that calodema and South Amerícan genera Metaxymorpha may be more closely related to the Parry (1848) erected Conoqnatha (s. s.) and Hyperantha (s. s.)' When the opinion that Metaxymorpha as a subgenus of stigmodera, he expressed M.grayiwasaconnectinglinkbetweenStigmoderaandConognatha' the resembling the former in some aspects and the latter in having Themognatha Calodena nidgut 57 sexuallY caeca dfmorphic short bursa I accessory trilobecl gland present Malpigh. tubuleE elytra grouped proxirnallY foveolate hieh S tubules/testis g,!þ muscles present stigmodera- FIGURE 27. Postulated reLationships within the Themoqnatha lineage 92 Greek for rbetween prostelnum prol-onged. The name is derived from the genus near Conognatha' formr. Van de PoIt (1886) also placed this of each discordant frlhen conflicts like this occul, the credibility Synapomorphymustbeassessed.Thepresenceofanaccessoryglandisa complex structure' character of high weight because it is an innovative, prosternal- not recorded to date in any other buprestid; while the projectionisofÌowweight,becauseasasimplecharacterduetoan allometric process, it is more difficult to discount homoplasy' of Anterior margin of elytra strongly bisinuate is a synapomorphy these taxa' Calodema and Conoqnatha ( s. s.), but Ís not unique to Conoqnatha ( s. s.): Similarly the characters shared by Metaxymorpha and elytra strongly tapered pronotum, large scutel-Ium, and sides of are therefore diverging from the shoulder, are of doubtful polarity and accorded 1ow weight- of one of the main problems with the hypothesis of monophyly absence of the accessory Themo tha CaLodema - Metaxymorpha is the interpreted in gtand in the Metaxymorpha meeki group. This could be related to the several, ways : 1 - the meeki group is not so closely accessory gland has other Meta xvmorPha species (Figure 284); 2 - the an autapomorphy of been secondarily lost in meeki gloup and represents thatspeciesgroup(Figurezs})i]-theaccessolyglandevolved independentlyinThemognatha,Ca]-odemaandMetaxymorphagrayi- M. apicaLis groups (Figure 2æ)' IfthemeekigroupofMetaxymorphaweretoberemovedfromthe because it lacked the Themognatha - cal-odema - Etaxvmorpha lineage problems of requi-red synapomorphy for incLusion (Figure 28A), more analysis would be created than soLved. Firstly, the overLapping elytra, the similarity of general facies with other Metaxvmorpha' and all the other shared characters woul-d have to be explained as M. g- l,t. elytra elytra o/L o/L a9 I'lt A 9,/u M. M. M. M. M. M. T Cal. apic. gr-"vi r.ãrt T. cal. apic. grayi ¡neeki a9 lost ag ag ag (as) elytra ÀNC. 3 p o/L elytra o/L Ar¡c. 2 P(as¡ ag B A¡TC. 1 p(as) Mt g,/u Mt g,/u c FIGURE 28. Three alternative schemes of argumentation explaining character distribution in the Themognatha- Calodema-Metaxymorpha lineage. Apomorphic characters are: Mt g/u = Mal-pighian tubules grouped proximally' ungrouped distally; â9 = accessory gland present; and elytra o/l = elytra overlapping. Based on notation of Saether (I98t), p(ag) represents an underlying potential to develop an accessory gland in a hypothetical ancestor (ANC.) represented by a circle. 97 grayi seems to occuPY a Position homoplasY. Furthermoret Metax and M. meeki grouPs, sharing some intermediate between the M . apicalis unjustifiable to split Metaxymorpha characters with each, and making it l' into different genera' Thus I reject alternative invoking secondary reversal The second scheme of argumentation inside parallelism (Figure 288) and the third scheme representing 1987) termed 'underlying (Figure 28C) are forms of what saether Q979, when the evolutionary synapomorphy" Inside parallelism occurs potentialtodeve]-opaparticu}archaracterisrealizedonfyinsome' group. saether's concept not a]1, of the descendants in a monophyletic ofinsideparallelismechosideasputforwardmuchearlierbyBock in evol-ution' parallelism (}gez) and Throckmorton (1965) who argued that andreversalmaybetheruleratherthantheexception'andaccordingly relationships' and exploited should be taken into account in assessing in the construction of cladograms' Ithinkthatinsideparallelismaloneshou]-dnotbeusedtoassert monophy}yforacollectivetaxon,asitj-smoredifficulttofalsify thansynapomorphybecauseoftheadhocnatureofitsexplanations. by other criteria however' the concept once the monophyly is established ofunderlyingsynapomorphycanbeausefultoo}toexamineandexplain apparentconf]-icts.Inmostcases,asintheabsenceofanaccessory glandinM.meekigloup,itisnotpossiþIetodistinguishbetween secondaryreversalandinsideparallelism,onlytoargueastowhichis morelike}y.Agreaterdegreeofconfidencecanbeplacedincomplex featureswhenjudgingcharacterstobehomo}ogousratherthan reason of the complexity of the accessory homoplasous (Bock, ]981). By group' I think that g1and, and its absence in only one species explanation for the alternative 2 (Figure 288).is the most parsimonj-ous Met. cas' S T cal. D. H. HIA P. co' Pro- proc. sternal I bilob Proc. elytra intern. male o/r proct. fused antennae toothecl 50+ from tub. prosternal 6Eh/7r}]' Process prostern tarsal claw stronglY toothe¿l sculPt. elYtra s7 sexually ctirnorphic mentun median lobe foveolate flaP notched ventral mg caêca accessory glancl IaterallY short present bursa Malpighian tubules trilobeal groupecl ProximallY sternal cavity 20+ tubules/testis transvèrse mesosternu¡n muscles Present s4-s6 very short El¡g mouthParts Prolonged antennae toothecl fron 5th segment' sclerotizecl sePtum shorÈ dividing Particle sac valve rods dorsal sclerotizecl deeplY divideil anterior to hg,/reclurn from 12' 13 9þ rnuscles ]ost foregut cliverticula presenÈ wing lacks vrealge cell ovipositor with dorsal valver parlicLe sac, sPatulate setae FIGURE29.C}adogramSummarizingtheavailableevidenceof phylogenetic relationship within Stigmoderini' 94 observed character state distribution, and consider M. meeki group best left in Metax ha. hlithin the Themo atha - Calodema - Metaxymorpha lineage no apomorphic charactels were found that were unique to Themognatha - Calodema or Themognatha - Metaxymorpha. Cal-odema and Metaxymorpha both have prosternal projections which may be due to cl-ose relationship or parallelism. These two genera are almost indistinguishable in many characters of unknown polarity e.g. shape of the mentum, Labrum and mandible; shape of 8th sternj-tes and tergites, aedeagus and median lobe, and I suggest that they are most closely related to each other. They are the only Australian genera confined to the rainforests of north-east Australia and Papua New Guinea: the other taxa are usually found in dry sclerophyll forests, mallee and heathlands. The higher the taxon, the more difficult it is to find apomorphic characters that are both unique to, and possessed by, all members of the taxon. Often the only evidence that a higher category is a natural group is derived from overlapping suites of characters, and it may be that not one species possesses al-I the characters which together delimit the higher taxon. Figure 29 summarizes the available phylogenetic evidence of relationship withÍn Stigmoderini. 4 TAXONOMIC IMH- ICATIONS OF PHYLOGENETIC ANALYSIS lVithin StigmoderÍni, the main implication of this phylogenetic analysis is that if Themognatha Ís more closely related to Cal-odema and Metaxymorpha than it is to stigmodera (s. s. ) and castiarÍna, then it should be removed from stiqmodera and elevated to generic status. There is no evidence to suggest that it should be lumped with any other 95 made in response to clear cases genus. Changes of status are rightly of polYPhYIY. Secondly,thereisnojustificationforretainingCastj-arinaasa contrary, the avaiLable evidence subgenus of Stigmodera' 0n the suggeststhatstigmoderaismorecloseJ-yrelatedtotheThemognatha- than to Castiarina' Each taxon is Calodema - Metax lineage as a number of characters of distinguis hed by autapomorphies as well discontinuity between these two unknown polarity, and the morphological taxaisatleastasgreatasþetweenanytwotaxaofgenericrankinthe tribe.IthereforeconsiderthatCastiarinashou].dbeelevatedto generic status also' in I think that Hypostigmodera varieqata is ProPerlY Placed foregut diverticula, lack of wedge Stigmoderini on the basis of having cellinthewingrandanovipositorwithdorsalvalve'particlesacand spatulatesetae;butthatitbe}ongsinCastiarina.Itshareswith cavity strongly transverse with Castiarina the apomorphy of sternal mesosterna}branchesreduced,andmediallobeofintercoxalprocess widerthanthetwolaterallobestogether.Inaddition,itsharesthe not hairy; prementum following characters: smatl size; frons concave, suture; scutellum not round or attached to mentum with grooved labiar per testis; Malpighian tubures ovar; 5 ovariotes per ovary; 10 tubul-es ungroupedproximatly;lackofsllgmuscles;andgeneralfacies.The differs from Castiarina is in its only character in which Hvpostigmodera distinctiveantennae,whichhavesensilladiffuseovermostofthe surface,notconcentratedinfoveaeandsecondarysensoryaleas,and which,inthema}e,arealsoofunusualstructure,withmeriston and segment 1l strongly toothed, segments 4-Ì0 bipectinate' differs also from a}l tripectinate. In this character, Hypostigmodera 96 other Stigmoderini. Therefore, I tecommend that this monotypÍc genus be synonymized with Castiarina. while the M. meeki and M. apicalis gloups of Metaxymorpha are distinct, and the inclusion of the M. meeki group in the Themognatha - problems, the intermediate Calodema - Me tax ha lineage presents some form of M. grayi and the synapomorphy of overJ-apping elytra algue against splitting Metaxymorpha into two or three genera' Similarly' I think that elevation of the rank of the three species groups of calodema woul-d be unjustified. Regarding Hyperantha ( s. s.) and HyperanthetLa, there are a number of morphological- differences which suggest divergent lineages including the prosternal Projection in some Hvperantha ( s. s.), and the reduction of elytral interneurs in Hyperanthell-a. However, I have not been abl-e to examine aIl of the species in these groups and it may be that those I have examined were by chance from opposite ends of the spectrum of variability. 0n the other hand, the presence of a large the ventral membranous flap on the median Lobe of the aedeagus supports hypothesis that these two taxa form a monophyletic group. I agree with Ehrlich and Murphy Q983) that obligatory categories shoul-d be conserved whenever possible, especially the generic namet because by indicating affinity it conveys maximum information about those species' So, I think that while these taxa are rightly separated at the subgeneric l-evel, there is insufficient evidence at this time to warrant changing their congeneric status. Similarly, the limited number of representatives of Pithiscus and Conognatha (s. s.) I examined have a number of morphological differences, and further studies, especially those of internal anatomyt genera' may indicate that they are sufficiently distinct to be fotlowing Solier (1849) and Kerremans (1902), and against waterhouse 97 Jl9lz)andHoscheck(1974).0ntheotherhand,theirverysimilar Thus' at present' in the genitalia suggests a close relationship' interestsofnomenclaturalstabi}ity,andintheabsenceofphylogenetic evidencetothecontrary,Iproposethatthediversetaxashouldremain subgenera of Conognatha' 5 DISTRIBUTI ON OF GENERA NoevidencewasfoundtosuggestthattheAustralianandSouth Americanfaunasrepresentdifferentlineages.Onecharactercommonto most,butnota]'lsouthAmericanStigmoderini,isthattheelytral apicesaredenticu]-ate,butthischaracterisofunknownpolarityand couldeasilyhavearisenseveraltimes,asitalsooccursinother plesent which support a cLoser buprestid tribes. some characters are relationshipofConognathatotheAustralianfaunathantotheother is correct' it would mean that South American taxa' If this hypothesis split off from the rest of the Dactvlozodes - Hyperantha lineage Stigmoderini].inebeforethebreakupofGondwanaland.Secondly,it wouldmeanthateitherthecontinentalsplitoccutredafterthesecond lineagegaverisetoConognathawhichcoincidentally'wasrestrictedto and the other taxa to the area the area which became South America, breakup of Gondwanaland resurted in which became AustralÍa; or that the theformationoftwopopulationsfromtheoriginalancestralone,the SouthAmericanpoputationgivingrisetoConognatha,andtheAustra]-ian Calodema Cas tiarina population giving rise to Stigmodera' seems to me to be the most and Metax vmo This last aLternative plausibleexplanationfortheobserveddistributionofgenera. Therealeanumberofexamplesofparallelevolutioninthe Australia-NewGuineaandSouthAmericancomponentsofthefauna.In species evolved with reduced tarsal both Castiarina and Dactvlozodes, 98 pads'compressedantennae,andreducednumberoftoothedantennal with prolonged segments. Both Castiarina and Pithiscus have species (eithel as a mouthparts and species with mouthparts abbreviated plesiomorphiccharacterretainedrolanapomorphysecondarily (s. show derived). Cal-odema. Meta and Conognatha s.) projection, pronotum convergence in shape of the body: the prosternal beyond the abdomen and elytra smooth and lustrous, and elytra extending presumablyallconferaerodynamicadvantagetohigh,fastf].iers. 6 cONcL UDING REMARKS Apaucityoflgood'characerswasoneofthemajorproblemsofthis they were anatysis. Many characters had to be discarded because within taxa invariable in the tribe e.g. wing venation; too variable e.g. number e.g. scutel]-um shape; judged not time rewarding to measure e'g' degree of sensilla in the antennal fovea; distributed haphazardly not discrete e'g' of fusion of female proctiger; or part of a continuum, that they relative eye size. some characters are so closely correlated €.9. dorsal valvet are considered col-Iectively as a single characterr particlesacandspatulatesetae.Anothermajorproblemwashowto the apomorphic deal with taxa in which not all the species possess rrone true character al-ternative. ltliley (1981:82) observed: Synapomolphyisenoughtodefineauniquegenealogicalrelationship. may or may not be The problem is that the synapomorphies we hypothesize we have true synapomorphies. Thus, the more shared derived characters tocorroborateagroup'spurportednaturalstatus,thebetterour hypothesis.rl Somegroups}endthemselvestophylogeneticanalysismorethan orders' and others. coleoptera is probably one of the more difficult 99 be achieved for it is doubtful if a thorough phylogenetic analysis can characters and their many beetl-e taxa. The initial selection of analysis (Doyen and poJ,arity has overwhelming influence on the cladistic of assessing polarity and Tschinket , 1982). Given the difficutties with appropriate homoplasy, and the scarcity of apomorphic characters distributions,Iconsideracompletephylogeneticanalysisof Stigmoderinitobeprematureatthisstage.Aschemeispresented' As more species are which I think þest fits the avaitable evidence' newly discovered and examined, and as fresh information (in the form of be required' reinterpreted characters) is integrated, a reappraisal will and its resolution may be improved' study, basing while I have adopted a phylogenetic approach in this relationshipsonsynapomorphyasmuchaspossible'inthefaceof groups, my classification inconclusive phylogenetic reso}ution of some relationship such necessarily rests also on less rel-iable indicators of the value of the as overal-I morphological similarity. Nevertheless, only directed phylogenetic method of argumentation is that it is not organisms, but also towards discovering the relationships between (ltliley' This aspect towards explaining character evolution 198L:I47)' of phylogenetics is one of the most challenging' I00 1. Tribe STIGMODERINI Lacordaire, 1857 L' & G" I9V7zI Buprestidae (part) Div. 2, subdiv. I solier 1873:267; Lacordaire 1857t52 Buprestides vrais (part) Group II Stigmodérides Heyne & stigmoderini Kerremans l}gSzLor'LI4, 190I:171, L9O2zJ'98; 19342677; Cobos Taschenberg 1907:J-40; Théry 1929227I; obenberger l980z29 and stigmoderinae Fowl-er 19122151; Tillyard I926t2I7; Brues, MeJ-ander Carpenter 19542555; Britton L97O:565; BIIy 1974274 Stigmoderae Carter 1916:80; Théry I929227L Eschschol-tz IB29 Type genus (by absotute tautonomy): stigmodera , stic characters: Body SmaLl to large, dorsoventrally flattened' HeadEyesgenerally}arge,subpara]-]-e].orconvergingslightlyon vertex; vertex with medial tongitudinal impressed line; clypeus and most, or clypeus mouthparts prolonged, galea with long dense setae in sparse setae and mouthparts abbreviated, galea with short' some Castiarina); Da l-ozodes Hvper antha. some Pithiscus, and prementum mandibres acute apically; ligura broadry rounded apicarly; prominent, or short and concealed, attachment to mentum variable; clypeus and eyes; antennal cavities open anteriorly, situated between antennalscapelengthgreaterthanorequaltopedicel+meriston together, but l-ess than 0.25 total length; toothed antennal segments with sensilla concentrated in a subterminal fovea and in a marginal in which sensil-la secondary Sensory area (except in Castiarina variegata are diffuse on both faces) ' 101_ than the Thorax Prosternum in front of procoxae longer of meso-metasterna]. suture of procoxae; mesosternum divided, position the cavity in variable; intercoxal process trilobed, almost filling most; wing without wedge cel-I' AbdomenMuscurature:tfgpresentonTl,T4-T6randaLsol2rTv T1-T8; short smlg + sld present on 1n Dactvl-ozodes and Hyperantha; ts 54-56,stlgalsoon54-56inStigmodera,Themognatha,Calodema,and pores present at least on f2-f4 and 54' Meta xvmorpha Ite rgal and sternal on 5th and 6th segments concealed' and in most on 12-16 and s4-s6, poles ovipositor with Female re ive system Short, sclerotized valve with marginal three varves: a dorsar vaLve, a style-bearing a ventraL valve; a particle spatulate setae and short medial styles, and varves; ovary with 5 or 7 sac present between dorsal and style-bearing ovarioles closely adpressed at distal ends' 10-94+ tubul-es' spirally MaIe reproduct ive system Testis with twisted;oneventralpairofaccessoryglands,2-2oxthelengthof thick-walled spermatophoral abdomen, in addition to a pair of targe, glandsreachcontainingaspermatophoreofcharacteristicform' than midgut, with a pair Alimentarv svstem Foregut much shorter (up to 0'2 length of of 1ateral foregut diverticula; midgut caeca short regenerative crypts dense and midgut) in most, longer in Dactyl-ozodes; even;hindgutaslongorlongerthanmidgut,anddifferentiatedinto a large intestine (0'7 small intestine (0.3 length of hindgut)' and grouped or ungrouped proximally' tength of hindgut); Malpighian tubules ungroupeddistally,becomingadpressedtohindgutabout0.Tlengthfrom pyloric va1ve. + 4 + 5-8' Nervous svstem I + II-III-I + 2 + 7 10 AA + Xy (most), or 9 AA + Chromosomes MaIe karyotype l¡ = O *to (t. nickerli and T. alternata)' l:O2 with the structure of the Remarks: The.suite of characters associated that the tribe is ovipositor is unique to stigmoderini and evidence monophyletic.Wingvenationandpresenceofforegutdiverticulaare not unique' The tribe is characteristlc of the tribe and constant, but Stiqmodera comprisedofmorethan640describedspeciesinSgenera. Castiarina L' & G' L837 Eschscholtz, !829, Themognatha Solier ' L8t3' ' ' Parry' 1848 are found in Calodema L. & G.' 1837, and Metaxymorpha Eschscholtz, 1829, AustraLia and PaPua New Guinea' Conoqnatha Mannerheim, 1837 are south DactvLozodes chevrolat, 1877 t and Hyperantha American. Key to the qenera of Stiqmoderini t Sternalcavitywidthmuchgreaterthanlength,meso-metasternal suture never basal " 'Castiarina equal to length' 1r Sternal cavity width less than or meso-metasternaL suture medial to basal " " "2 3 z(tt) Mouthparts not prolonged into rostrum" 5 2t Mouthparts prolonged into rostrum' 3Q)Antennaewithsetaeaslongorlongerthansegments,toothed towards apex" Dactvlozodes segments with tooth triangular, widest ' segments, toothed segments 3t Antennae with setae much shorter than with tooth rounded, widest towards middle ""4 Hvperantha 4(7t ) Antennae toothed from 5th' Conoqnatha pithiscus) (part) 4t Antennae toothed from 4th' LO7 5(2,)Bodywedge-shapedorsinuousinlateralview;prosternum 6 enlarged view; prosternum not enlarged I 5t Body subparalleL in l-ateral ..Conognatha (s' s') 6(5) Elytral aPices denticulate trispinose' 7 6t Etytral apices bidentate or Met a 7(6') Elytra over1aPPing.. ' ...Calodema 7t Elytrq not overlaPPing Stiomodera (s. s.) 8(5' ) EJ-ytra foveolate 9 8r E1ytra not foveolate' elytron; Iabrum g(g,) scutelLum very small, Iess than 0.1 width of curvilineartriangular,withoutborder;tarsa].clawwithbasal toothinaþout75%olspecles;elytralapicesneverdenticu]-ate; Themoonatha Australian. . . . of elytron; Iabrum rectangular' 9t Scutellum larger' O'L-O'2 width withmedia]Þorder;tarsalclawsimple'neverwithbasaltooth; elytral apices denticulate in most; South American' Conoqnatha (Plthiscus) (Part) Genus Stiqmodera Eschscholtz, 1829 I877t297; L' & G' I877:1-I0; stiqmodera Eschscholtz 1829:9; Solier ' Saunders Lacordaire 1857 257; laitmaire & Germain 1858272o-775; & Harold 1869: 1397-1404; IB68:23-5v, 1870:6+' 1871 266-76; Gemminger L9o2z2o7-2L4; Masters rg7LzL29-.r41; Kerremans r}gz:r43, L897zrr4, 104 Carter Froggatt t19O7l tI64; Heyne & Taschenberg 1907214]-147; 1926:217 1916:78-178, :f92g ¡ 289-3oo, 19312777-768; Tillyard ; Obenberge r 1974 :678-75O; McMilLan L9522I45-7' Polvchroma Dejean 1836289 lnomen nudum] TVpe species(designatedbySo]-ier1877):S.macularia(Donovan),].805 ostic characters BodyLengthl5-TOmm,subpara}Ielinlatera]-view. prolonged, Head Frons hairy; eyes large, subparallel; mouthparts galea, with long dense setae; Iabrum trapezoida}, anterior margin bilobedwithbisinuatetrans].ucentborderandwidemediallongitudinal subapical tooth and groove; mandibles elongate, with one rounded dorsal segment of labial and one Very weak ventral subapical tooth; terminal anteri-orly' maxillary palps truncate; prementum attached to mentum is dorsaf); l-abial suture absent (except S. gratiosa in which attachment ]aterally; antennae mentum strongly transverse, L/I¡'l=o.5, not notched eJ-ongate, tooth toothed from 4th segment; toothed antennal segments rounded,widestatmidd]-e;antenna}segmentllwithdorsalfovea. Thorax sides of prothorax carinate, or strongly explanate (S. gratiosa); posterior lateral angles of pronotum not prolonged; not enlarged; pronoto-elytral articulation straight; anteri'or prosternum process sternal cavity as long as wide, medial l-obe of intercoxal 0']-0'2 width of rounded, equal in width to Lateral Lobes; scutell-um elytron, scutj-form. with Elvtra convex, lateral margins undifferentiated; interneurs length (except large, deep, cl-ose foveae , 9 * 10 separate fOr mOst of va¡iable but !. gratiosa in which they are fused anteriorly); apices never denticulate; not overlapping; not setose' 105 Legs Posterior margin of hind coxa not strongly obJ-ique; hind tarsus only slightly longer than fore or mid; tarsomeres fl-attened; tarsal claw simPle. Abdomen 57 not sexually dimorphic (except S' sanguinosa in which pronounced mal-e truncate, female bilobed); female proctiger with transverse suture (except S. gratiosa); male proctiger T9+l-0 fusion smlg + incomplete; tergal muscles tlg : TI, T4-16, long; sternal muscles sllg+s1d:54-56. sac not Female reproduct ive svstem Dorsal valve divided; particle with exceeding val-ves, bilobed, with sclerotized septumi bursa trilobed, spermatheca shorter than bursa (except S. gratiosa); 5 ovariol-es/ovary (except !. gratiosa with 7); accessory gland absent' median lobe of Mal-e reoroductive svstem I2-5O tubules/testis; parallel-sided on aedeagus without ventral membranous flapt nal¡ow and anterior half, broadening posteriorly, ostium 0.5-0.6 length from base' apex triangular. Alimentarv system Midgut caeca very short (less than 0'1 length (except of midgut); Malpighian tubules ungrouped proximally s. gratiosa); sclerotized rods anterior to junction of rectum and hindgut absent. specj-es: s. cancel-lata (Don.), 1805; !. gratiosa chevr., L847; 9. jacquinoti (Boisd.), }875; S. macularia (Don.), 1805; 9. porosa Carter, 1916; S. roei Saunders, 1868; S. sanguinosä Hopel 1846. elytra' Remarks: The monophyly of stigmodera is supported by foveolate trilobed bursa and short midgut caeca: it is the sister group of transferred rudis Themognatha - Cal-odema - Metaxvmorpha. Barker Q979) Carter to Stiqmodera ( s. s.), but subsequently reinstated it in 106 Castiarina (Barker' 1980). g. goryj-, formerly placed in Stiqmodera (s.s.)istransferredtoThemognathaonthegroundsthatit with has none of the features unique to stigmodera and it shares an accessoly gland' Themoqnatha aII characters including the presence of shares with The affinities of S. gratiosa are problematical. It characters stiqmodera foveolate elytra, and with Themognatha a number of with including: prementum attached dorsally; bursa unil-obed, elongate, Malpighian spermatheca longer than bursa; 7 ovarioles per ovaly; and tubules grouped proximal-Iy. I propose to l-eave gratiosa in Stiqmodera are because most of the characters shared with Themognatha diagnostic symplesiomorphic or of unknown polarity; foveolate elytra are in of stigmodera; and it lacks an accessory gland required for inclusion from the Themogna tha. I suggest that s. gratiosa diverged first Stigmodera lineage, retaining a number of primitive characters' Distribution: S. macuLaria, S. iacquinoti and S' porosa are eastern Australian species, s. cancellata, s. gratiosa, and s. roei are western Australian species, S. sanguinosa occurs on both sides of the continent' Genus tha So1ier, 1833 T atha Solier 18732 29I 1837 z76i Lacordaire 1857 Stlgmodera Mannerhej-n "59; Heyne & Gemminger & Harold I869:1797i Kerremans 19o2z2j4-206; Taschenberg I9O7:J-41; Carter 1916:86192-96, 19242 577, 1973 :66-68' 19342678 1929 2289-292, I93L:337 ; obenberger L928262-68, Stigmodera sri deroides) Théry 1937224 Temoqnatha (sic) So1ier 18732 26v 107 G. ]:877zl-]--22; Kerremans I89vzII4 Stiqmodera (remoqnatha) ( sic) L. & , Sticmodera (Temnognatha) (sic) Imhoff 1856247 Tvpe soecies(byoriginaldesignation):T'variabilis(Donovan)r1805 tic characters Bodv Length Ì5-60 mm, subparallel- in lateral view' Headfronshairy;eyeslarge,subparallelorconvergingslightlyon vertex;mouthpartsprolongedrgaleawithlong'densesetae;Iabrum With medial curvilinear trlangular, anterior margin scarcely notched, groove present; mandibles setae, without border, medial longitudinal or absent; terminal segment elongate, with dorsal and ventral tooth weak prementum attached to of labial and maxillary palps truncate or rounded; suture; mentum variable in shape mentum dorsally, with plonounced labial (L/W=0.5-0.9)'notnotchedlateralty;antennaetoothedfrom4th tooth rounded, widest at segment; toothed antennal segments elongate, fovea towards apex' the middle; antennal segment 11 with smalL dorsal posterior Thorax sides of prothorax carinate or explanate; lateralanglesofpronotumnotprolonged;pronoto-elytralarticulation almoststraight,oranteriormarginofelytronwithsmall]-atera]. cavity as long as projection; anterior prosternum not enlarged; sternal cavity, or basal; scutellum wide, meso-metasternal suture hatfway along variabre. ress than or equar to 0.r width of elytron, shape Elytraconvex,lateralmarginsnotstronglydifferentiated; apices variabre' interneurs punctate-striate , 9 * 10 fusion variabte; setose' but never denticulate; not overl-apping; not LegsPosteriormarginofhindcoxanotstronglyobtique;hind tarsusslightlyormarkedlylongerthanforeormid;tarsomeres flattened;tarsalctawwithbasa]-toothinT5%ofspecies. 108 the species, s9 Abdomen Mar-e s7 widely notched in about half T' flavicollis and exposed in most (except a few species groups e'g' variable; male I. saundersi); female proctiger with transverse suture T4-16 long; proctiger T9+]0 fusion incomplete; tergal muscles tlg: TI, sternal muscl-es sml-g + smlg + sl-d: 54-56' particle sac Female ive svstem Dorsal valve divided; unilobed elongate' long or short, bilobed with scLerotized septum; bursa accessory gland spermatheca + duct longer than bursa; 7 ovariotes/ovary; present. median Lobe Male reproductive svstem 12-94+ tubules/testis; 0'8 length from without ventral- membranous flap, sides curved, ostium base, apex variable. (O'1-O'2 of midgut); Alimentarv system Midgut caeca short length rods anterior to Ma}pighian tubu]-es grouped proximal]-y; sc].erotized junction of rectum and hindgut absent' (carter, 1929) lists 76 species: The most comprehensive checklist specj-es. Carter lJfg7u places 12 oî these in synonymy' obenberger (]g:'L)listsonly12species.Afewspecieshavebeendescribedsince 1964 and then: l. gemmelli Deuquet , L947, T. macqueeni Deuquet, six species from L. coronata Peterson, 1982' Barker 0979) transferred and Castiarina to tha. A revision of the group is needed, over 100 (S' Barker' would probably bring the total number of species to pers. comm.). Metaxymorpha on Remarks: The taxon is closely related to Calodema and thebasisoftheSynapomorphies:presenceofaccessoryglandand Malpighiantubulesgroupedproximally.Todate,theonlypublishedkey 109 of species groups are to species is carter (1916:92-96). A number are described below' evident, three of them which I examined in detail Distribution: Mainland Austratia and Tasmania' atha reichi group Gehin, 1855; soecies: T. bonvoul_oiri saunders, Ig6g; T. chevroLati 1927., T. murrayi Gem. & T. congener Saunders , 1869; T. miranda Carter, Har., 1869; T. reichei L' &G', 1877 antennae longer than Di tic characters: Size large; ligu1a bifid; pointed behind; basal the width of the head; scutellum round or oval, ridge between s2 & tooth on tarsat claw pronounced; plonounced internal 53;maleSTverystronglynotched;femaleSSelongate;maleTgelongate' visible, heavily tapered, pointed or bifid apically; female T9 always apically; sclerotized, Very elongate, concave and narrowly rounded anteriorly; sclerotizatÍon on ventral surface of ventral valve narrow plate near base (in dorsar valve ronger than wide; bursa with spiny (average 55) some) ; 40-75 tubules/testis ' as Stiomodera Remarks: This group was given subgeneric status (sti des) by ThérY Qglt:24) ' Themoqnatha maculi ventris group I862i Soecies: T. alter¡a'þ Lumh., 1889;. T. maculiventris Macleay, T. nickerli Obenberger' L922 110 medium; antennae noticeabry longer than the D tic characters: size pale lateral margin; width of the head; pronotum tapers rapidly, without tarsi elongate; intercoxal process with pronounced lateraL depressions; with metal]ic tarsal claw simple or weakly toothed; elytra yellow margin of elytron with blue/black fascia and red ñrargin/fascia; anterior lateralprojection,butnotasstrongasinT.donovanigroup;abdomen pa}e;elytralapicestruncateorwidelyrounded;maleSTtruncate; proctiger and ovipositor femal-e s8, T8, proctiger and valvifers tong; T' fringed with long setae (especially in T' nickerli and ventral valve; particle sac macuLiventr 1S ) ; dorsal val-ve extends beyond 20 shorter than valves; 2l-22 tubules/testis; chromosots5 !¡ = available)' ( T. alternata and T. nickerl-i T. maculiventris not proctiger and Remarks: The long termi_nalia, and rong setae of the ovipositorareautapomorphiesofthisspeciesgroup.g.alternataand subgenus Themognatha (Kerremans, I9O2; 1.. macul-iventris were placed in with s' nickerli obenberger, 1922), in subgenus castj-arina together tha (Carter, !924, 1929, I97D, and transferred back to subgenus (Barker , 1979). tha donovani gloup 1874; soecies: T. donovani L. & G., 1877; T. tricoLorata waterhouse, T. viridicincta V'laterhouse, 1874 but not as long as in Dia tic characters: size medium; antenna long, pronotum T. macuLiventris group; eyes elongate, converging apically; anterior margin metal-lic with yellow oI red margin; tarsaL claw simple; of elytra with strong lateral projections fitting into basal notches on tll the pronotum; elytra yellow with metallic green or violet fascia or sutural margin; elytral apices bispinose, sutural spine longer; abdomen pale with metal-lic blue or green markings; s9 short; 23-25 tubules/testis; chromosomes 2n = 22. Genus Castiarina L. & G., 1877 Stigmodera castiarina) L. & G., 183724,22-47 ; Imhoff L856247; Lacordaj-re 1857:60; Gemminger & Harold 1869:1397; Kerremans I9O2z2O4 1206-214 Froggatt [1907] :I64i Heyne & Taschenberg l9O7 :l4I; Carter J.916 :96-110, 1929 :292-3OO, 19712349-367 ; obenberger 19282 69-72, 728-775, 1977268-76, IO4-II2, 19342 678-75O; Barker 1979 zI-2t, 1980:1-7, 1987:179-169 Polychroma De jean 1836:89 [nomen nudum] Hypostigmodera Blackburn, 1892 Tvpe species: C. pertyi L. & G., 1837 (Barker, 1979) Diaqnostic characters Body Length less than 20 mm (except c. deyrolLei Thomson, C. magnificollis Barker, C. al-ternozona Thomson, and C. rol-lei Kerremans tS. Barker, peIS. comm.l); subparallel in Lateral view (except 9. producta group, attenuated behind). Head Frons without hair (except C. rudis Carter, 9. iubata Blackburn and C. erubscens Blackburn), with medial longitudinal sul-cus in most (except in C. adeLaidae Hope, other bee mimics, C. flava Saunders, and C. obscura.saunders); eyes large, subparallel or converging slightly on vertex; mouthparts prolonged or abbreviated; trz labrumrectangular,anteriormarginweaklybisinuateorstraight; mandibleselongatewithoneortwodorsa].subapicalteethandone ventralsubapicattooth;terminatsegmentof]-abialandmaxillarypalps with deeply grooved variable; prementum attached to mentum anteriorly ÌateralJ-y; antennae ]abial suture; mentum variable in shape, not notched (except mal-e c' variegata from the frd) toothed from 4th or 5th segment ' segments with tooth rounded' widest compressed in some; toothed antennal (except C. variegata segments at midd}e, almost as wide as long male segment 11 with dorsaL 4-10 bipectinate, 11 tripectinate); antennal & G' fovea in few species only e'g' C' ctuenta L' (except c' bremeÍ Thorax sides of prothorax with incomplete edge l-ateral angles of pronotum Hope and c. insularis Blackburn); posterior notpro}onged;pronoto-elytralarticulationalmoststraightor sternal cavity strongly bisinuate; anterior prosternum not enlarged; transverse,mesosterna]-branchesshort,mediallobeofintercoxal lobes together' and with process truncate, wider than the two Lateral in many; scutellum lateral longitudinal grooves; metasternum raised or scutiform, never oval o.I-0.2 width of elytron, cordiform, tulipiform or round. in most; interneurs Elvtra convex, Iateral margins undifferentiated punctate-striateorcostate,g+]-0fusedonanteriorhalfinmost,or completelyfused(e.g.c.rufipesMacLeay,c.pulchripesBlackburn' g.sexualisCarter,C.ro]-]-eiKerremans,C.viridiventrisMacLeay,and denticulate but C. victoriensis Blackburn); apices variable' never L' & G'); not overl-apping; sometimes subserrate (e.g. c. austral-asiae (c. producta group); not equal to length of abdomen, or exceeding it setose. hind Legs Posterior margin of hind coxa not strongly oblique; ftattened or tarsus only slightly tonger than fore or mid; tarsomeres TT7 Distat ha].f of fore-tibia compact and globular; tarsal claw simple. pl-anipes Barker and c' ratipes frattened in severar rycid mimics e.g. c. Carter. 57 not deeply Abdomen 57 not sexually dimorphic, at least male notchedexposingsg;femaleTg+]-0withfusioncomplete,suturenot discernible,proctigersmallerthanvalvesinmany;ma].eT9+]-0fusion incomplete;tergalmusclestlg:T1'T4-T6Iong;sternalmusclessmlg+ sld onIY : 54-56. particle sac Female re ve svstem Dorsal valve divided; longorshort,bilobed,withsclerotizedseptum;bulsaunilobed' than bursa; 5 elongate or round, spermatheca + duct longer ovarioles/ovary; accessory gland absent' median lobe Male reproductive system ]]O/12 tubules/testis; lobe and parameres without ventral- membranous flap, shape of median variable between sPecies groups' (0.1-0.2 length midgut); A1imen tarv svstem Midgut caeca short (except C' pallidiventris Malpighian tubules ungrouped proximally L.&G.andC.erubPscensBlackburn);sclerotizedrodsanteriorto junction of rectum and hindgut absent' Species: This Ís a large and diverse genus, currently undergoing many of which revision by s. Barker. It contains more than 374 species exhibitstrikingmimicryoflycids,cantharj-ds,orotherinsects (Blackburn), 1892, initially (s. Barker, pers. comm.). c. variegata placedintherronotypicgenusHypostigmoderaonthebasisofits distinctivemaleantennae,properlybe}ongsinCastiarina. branches Remarks: sternal cavity strongty transverse, mesosternal than the two short, medial lobe of intercoxal process truncate wider 114 lateral lobes together are characters unique to Castiarina, and presumed groups apomorphic, supporting the monophyly of the genus. The species are characterized by genitalic structure as well as a suite of characters of external morphology such as body shape, size, modified tarsal pads and antennae (S. Barker, pers' comm')' Distribution: MainLand Australia and TasmanÍa, New Guinea, Thursday Island. Genus Calodema L. & G., I8V7 Stiqmodera (Calodema) L. & G., 1877z7O-72 1868222' Calodema Lacordaire 1857:60; H. Deyrolle 1864278; Saunders 187Iz65; Gemminger & Harold 1869:1405; Masters 1871:J-28; Kerremans I8922L42; I897;II4; u9O2:2O2-2O7; Heyne & Taschenberg 1907zI47; Froggatt t19o7l:I43; Carter 19292288; obenberger 19342 750-751 & L837 Tvpe species ( by original designation): C. regalis L' G', Dia characters Body Length: 24-5o rrn, notum strongly convex, body wedge-shaped or sinuous in Lateral view. Head Frons with medial longitudinal- groove, hair variable; eyes large, converging slightly on vertex; mouthparts and clypeus prolonged; without Iabrum curvilinear triangular, apex lounded or weakJ'y notched, border, medial longitudinal groove present; mandibles elongate with weak dorsal subapical tooth, and no ventral subapical tooth; terminal mentum dorsally' segments of labial palps rounded; prementum attached to u5 ]abialsuturepronounced;mentumelongateL/|tl=0.7-o.S,notnotched laterally;antennaetoothedfrom4thsegment;toothedantennalsegments the middle; antennal segment }onger than wide, tooth rounded, widest at 11 with dorsal fovea. angles of Thorax sides of prothorax explanate; posterior lateral strongly bisinuate' pronotum not prolonged; pronoto-elytral articulation posterj-ormarginofpronotumwithaprominentmedianlobeinfrontof scute]']-um;anteriorprosternumwithVerypronounced,acuteconical suture halfway process; sternal cavity as long as widet meso-metasternal process rounded, equal in width along cavity, medial lobe of intercoxal elongate, rounded to lateral lobes; scutetlum 0. I-o.2 width of elytron, in front pointed behind, lunate or subcordiform' Elytraconvex,lateralmarginundifferentiated;interneursfinely (e'g' C' regalis) or punctate with additional punctuations on intervals stronglypunctate-striate(e.g.C.ribbei),9*l0fusedonanterior half;apicesbispinoseortrispinose'neverdenticulate;not setose' overlapping; exceeding length of abdomen; not oblique; hind tarsi Egs Posterior margin of hind coxa strongly markedlylongerthanforeormid;tarsomeresflattened;tarsalclaw simple. AbdomenMaleSTbroad}ynotched,exposingSg;fema]-eproctiger mare proctiger T9+10 T9+10 fusion complete, suture not discernibl-e; fusionincomplete;tergalmusclestlg:TIrT4-16long;sternalmuscles smlg+s]lg+sld:54-56' deeply divided; particle Female reProd uctive system Dorsal valve Sacexceedingvalvesbyonethirdtheirtength,bi}obed,with + duct longer sclerotized septum; bursa unilobed elongate, spermatheca gland present' than bursa; 7 ovarioLes/ovary; accessory TABLE 4 groups Distinguishing characters of Cal-odema species plebius group C. regalis group C. ribbei group C' sinuous supra- weakl-y sculptured frons flat with antennal- ridges converging on vertex transverse more elongate transverse pronotum L/I^,l < 0.6 L/W > o.7 L/W < 0.6 diverge from base converge from diverge stronglY sides of with sides curved pronotum base where from base widest angular Projections less than equal to elytral slightly more pronotal width than pronotal pronotal concave flat, flat, posterior t rounded pronotal truncate rounded median l-obe subcordiform scuteLlum elongate t lunate, triangular overlapPed bY lobe overlapPed bY lobe abutting it overlapPing lobe or bisPinose male bisPinose male tridentate elytral maJ-e femal-e tridentate apices female bisPinose femaLe tridentate or not interneurs not striate strongly striate striate 116 not available; Male reproductive svstem Number of tubuLes/testis curved, ostium 0'8 median Lobe without ventral, membranous flap, sides length from base, apex drawn into a point' of Alimentarv system Midgut caeca slightly more than 0't length rods anterior midgut; Matpighian tubules grouped proximatly; sclerotized to junction of rectum and hindgut absent' G. c. ribbei soecies: c. prebeius Jordan, rg95; c. regaris L. & r 1877; speci.es van der PoIl, 1885; C. wallacei Deyrolle, L864; and four with a key to all described for the first time in Appendix G, together the species. Remarks: This genus is closely related tO Metaxymorpha and group tha. Three species groups can be discerned: C' regalis containing containing c. regalis and c. wallacei; c. riþbei group plebeius C.ribbeiandonenewspecies;9'plebeiusgroupcontainingC . characters' and three new species. Table 4 lists their distinguishing Distribution:North-easternAustraliaandNewGuinea. Genus Metaxymorpha Parry, 1848 1857t59 Stigmodera (Metaxymorpha) Parry 1848:82; Lacordaire MetaxvmorphaGemminger&Harotd1869z1404;Saundersl87l:65;Masters 187}:128;KerremansI892zL42,J:897I.J.I4;I9o2:2o3;Heyne& 19742 75I-752 Taschenberg 1907 zl47; carter L9292288-289; obenberget Semnopharus van de PolI 1886 2279; Kerremans I892zM 117 gravi Parry, 1848 Tvpe soecies ( by original designation): M' Di tic characters BodvLength:22.SSnn,notumconvex,bodywedge-shapedinlatera] view. HeadFronshairy;eyeslargerconvergingslightlyonvertex; triangular' apex mouthparts and clypeus prolonged; labrum curvilinear longitudinal groove rounded or weakly notched, without border, medial and no present; mandibles elongate, with one dorsal subapical tooth, palps ventra] subapical tooth; terminaÌ segment of l-abial and maxiLlary with pronounced broadly rounded; prementum attached to mentum dorsally laterally; fabial suture; mentum elongate LlW = 0.7-0.8' not notched segments longer than antennae toothed from 4th segment; toothed antennal 11 with dorsal wide, tooth rounded, widest at middle; antennal segment fovea. of Thorax sides of prothorax carinate; posterior lateral angles or weakly pronotum not prolonged; pronoto-elytral articul-ation straight bisinuate; anterior prosternum with low hump or plonounced, acute meso-metasternal coni.cal process; sternal cavity longer than wide, process rounded' suture halfway along cavity, medial Lobe of intercoxal O'3 width of elytron' width equal to l-ateral lobes; scutellum more than pentagonal or cordiform' 'E]-ytraconvex,lateralmarginundifferentiated,divergingfromand scutellum overlap base, continuing line of pronotum, base of elytra punctate-striate' pronotum and are raised above it; interneurs finely g+Lofusedanteriorly;apicesbidentatewithmedialtoothrounded'or overlapping trispinose, Sexes dimorphic in M. grayi; with one elytron sternite 7 by the other; markedly longer than abdomen, extending beyond a length greater than 57; not setose' t]8 Legs posterior margin of hind coxa not strongly obrique; hind flattened; tarsal- tarsus markedly ]onger than fore or mid; tarsomeres claw simPle. proctiger T9+r0 Abdomen Male 57 notched, exposing sg; femar-e proctiger T9+L0 fusion fusion complete, suture not discernible; male incomplete;tergalmusc]-estlg:T1'T4-T6Iong;sternafmusc]-essmlg+ sllg+sld:54-56. divided; particle Female reProductj-ve svstem Dorsal valve deeply bilobe.d with sclerotized sac exceeding valves by one third their length, + duct longer than bursa; septum; bursa unilobed elongate, spermatheca (one specimen of grayi had 7 and number of ovarioles/ovary variable [' accessory gland g, one specimen of M. meeki or near M. meeki had 5); present in M. grayi and M' gI¡icalis group' not available; Male reproduct ive svstem Number of tubules/testis sides curved, ostium 0'8 median l-obe without ventraL membranous flap, length from base, apex acute or obtusely angled' to 0.1 J-ength of midgut; A system Midgut caeca equal Malpighiantubulesgroupedproximally;sclerotizedrodsanteriorto junction of rectum and hindgut absent' Théry, 1922; Species: M. apicalis (van de Po11), 1886; M. apicerubra 1894; M' grayi Parry' M. dohertyi Théry, 1922; M. gloriosa Blackburn, 1922; 1"1 . sternalis 1848; M. hauseri Théry , L926; M' meeki Théry ' for the first time in Appendix Hoscheck , l97L; and two species described H, together with a key to all the species' related to Calodema and Remarks: This is a monophyletic genus, closely be discerned: M. meeki group Themo atha. Two main species groups can gloup containing containing M. meeki, and M. hauseri; and M. apicalis TABLE 5. DistinguishingcharactersofMetaxymorphaspeciesgroups M. meeki group M. apicalis group groove vertex very fine, shallow wider, deeper impressed line edges slightlY raised antennal segs longer (seg.8L> v'/) wider(seg.8L femal-e stern. 7 rounded truncate emarginate male sternite 7 weakly emarginate more stronglY (mediaL LlW O.7) male sternite I broader, (L/W = 0.4) narrower = present femal-e acc. 91. absent 0.7) longer 1¡7¡r¡ = 0.9) shorter 1¡7¡1 = style-bearing margin straight vaLve posterior margin curved posterior 0.8) dorsal valve lonoer (L/V'J = 1.0+) shorter 1¡7Y'¡ = posierior margin oblique posterior margj-n straight ventral valve narrow PosteriorlY broaden PosteriorlY sclerotization 119 and two new M. apicalis, M. gloriosa, M' a cerubra M. sternalis species.Table5].iststheirdistinguishingfeatures.Typespecies M.grayiParry'ls4SresemblesM.meekigloupinfivecharacters,and therefore M. apicalis group in 17 cha¡acters. This species is because I was ungrouped. I cannot judge the affinities of M' dohertyi unab]-etoobtainaspecimen,andthepresenceorabsenceofthe description, it appears accessory gland is unknown. From the original to share some characters with each group' are from Distribution: M. grayi, M. glorj-osa and M' hauseri are from Papua New Guinea' north-eastern Australia, the remaining species Genus Conoqnatha Eschschol-tz, L829 1837237; conognatha Eschscholtz L82929; Solier 1837"294; Mannerhein 187Iz62; Kerremans Gemminger & Harold I869:L795; Saunders I87Oz3+, I8922L37,1893:1}4,I9o22U99-2oo;Heyne&Taschenberg1907:140; Hoscheck 1934295-289' PI' waterhous e I9:12z57; obenberger 19342752; r-17. Stiqmodera (Conognatha)1.&G.,1837t45o;Lacordaire1857z58-59; Fai.rmaire & Germain 1858272O-774 Polydora Gistel I8V4zI0 [nomen nudum] Pithiscus Solier 1849:. 49I amoena (Kirby) IB18 Tvpe species (designated by Sotier I8V72295): !' ' L912:58). (not c. haemorrhoidal-is oliver as stated by waterhouse r20 tic characters or Bodv Length 9-70 mm (Hoscheck, 1974:100) ' wedge-shaped subparallel- in lateral view' Head Frons hair variabte; eyes large, subparaÌIel; mouthparts margin scarcely prolonged or abbreviated; labrum rectangular, anterior groove bisinuate, many with translucent border, medial longitudinal with teeth present (except c. subdilatata and c. insignis); mandibl-es palps usually variabl-e; terminal segment of l-abial and maxillary truncate or rounded; prementum attachment variable; mentum shape 4th segment; variabl-e, not notched laterally; antennae toothed from but always widest toothed antennal segments weakly or strongly toothed, at the middle; antennal- segment 1l with or without dorsal fovea' Thorax sides of prothorax variable, incomplete with weak lateral rimonbasa}quarteron}y,tostronglyexplanate;posteriorlateral angleofpronotumprominentornot;pronoto-elytralarticulation pronounced' variabLe; anterior prosternum variable with low or very enlargement in acute conicaL process in conognatha (s. s.), without g.. (Pithiscus); sternaL cavity as long as wider meso-metasternal suture medial lobe of halfway along cavity or basal (e.g. c. (P.) kLugi), intercoxal process rounded; scutel-lum 0.1-0., width of elytron, shape variabLe. (except El-ytra convex, lateral margin not strongly differentiated in costate species e.g. c. compta and c. chiLiensis); interneurs 9 + 10 costate, or punctate-striate with punctuations fine and shallow' denticulate; fused anteriorly and deepl-y grooved; apices variable' many not overl-aPPing; not setose' LegsPosteriormarginofhindcoxanotstronglyoþIique;hind tarsimarkedlyorslightlylongerthanforeormid;tarsomeres flattened; tarsal claw simPle. T2T AbdomenMaLeSTstronglynotchedinsome'exposingsg;femaLe proctigernalroulmedially,exceedinglengthofvalves,Tg+l0fusion (Conognatha (s' s')) or variable; male proctiger T9+I0 fusion complete TI, T4-f6 long; sternal incomplete (Pithiscus); tergal muscl-es tlg: muscLes smlg + s1d onJ-Y: 54-56' deeply divided to base; Female reprod uctive svstem Dorsal valve sc].erotized septum; particle sac not exceeding valves, bilobed with style-bearingandventra]'valveswiderthanlong,ventra}va].vemuch shorter;styIe-bearingvalveswithlighttransversesc]erotization ventratly;bursaelongate,unilobedorbilobed,spermatheca+duct longerthanbursa;5ovarioles/ovary;accessoryglandabsent' (based on one speci'es MaIe reProduct ive svstem L2 tubules/testis flap, short (0'5-0'7 only); median lobe without memþranous ventraL lengthofparameres),sidescurved,ostium0.Slengthfrombase'apex acuteornarrowedtoathinpoint,withlateralridgesondorsal lobe' surface, ostium large + O'25 length of median Malpighian tubules Alimentarv svstem Midgut caeca not avair-abre; notavailabte;sc].erotj-zedrodsanteriortojunctionofrectumand hindgut absent. listed 87 species' species: Hoscheck's Q9V4) monograph on conognatha Sincethen,4newspeciesand2subspecieshavebeendescribed: Cobos, 1959; C' reichardti Cobos' C. obe i OLave , 1978; C. leechi 1967;C.iriscayenensisCobos,Ig6g;C.amoenaanaphitritesCobos' undescribed sp. (T. Moore' 1969; and C. penai Moore' 1981; and one in litt. 1985). (1974) concurred that Remarks: Waterhouse (1912) and Hoscheck that the character used Conoqnatha and Pithiscus are synonymsr arguing the subgenera of Conoqnatha TABLE 6 Characters distinguishing Conoqnatha ( s. s.) Pithiscus not stronglY downturned body stronglY downturned bodY wedge- anteriorl-Y, bodY sub- anterloilY, l-ateral view shaped in l-ateral view parallel in hairy (excePt excell-ens frons without hair verY abbreviated or Prolonged mouthParts prolonged, cIYPeus Iong transverset almost as long as wide more stronglY Labrum length Less/= 0.7 width more robust, strong ventral mandibles elongate, weak ventral subapical subapical attachment variabl'e prementum attached anteriorlY 0.7-0'8 transverse, L/W = 0'4-0'6 mentum longer, L/W = toothed I'l head, toothed L less than W head, antennae L equal rounded segments weaklY rounded segments stronglY often with strong medial pronotal weak dePressions in basal angles and lateral basal sculpture depressions prominent posterior acute, Prominent not lateral angles of pronotun almost straight pronoto-e1YtraI bisinuate articulation not enlarged anterior enlarged with low prosternum hump to acute Process I/3 small to intermediate scute]Ium size intermediate I/5 - elYtron width of elYtron LlLO - 1/5 width punctate-striate or costate interneurs punctuations fj-ne and shallow absent suture entire female Proctiger suture Partial or fusion incomPlete, male proctiger fusioncomPlete,L=W elongate,L=l.5xW unilobed or bilobed bursa uniLobed 0.5 L parameres L of median lobe 0.6-0.7 L Parameres L22 the prosternum' was not by Kerremans ( I9O2) to distinguish them, form of or less valid since there was evely gradation between the states "more tthumping an angular projectionrr' convexrr and of the prosternum into only difference between However, the form of the prosternum is not the study two the two taxa, although it is the most obvious. In this ( to Hoscheckrs subgenera are lecognized: Conoqnatha s. s.) equivalent Hoscheckrs Group III' Groups I and II, and Pithiscus equival-ent to Table 6' Pithiscus They are distinguíshed by the characters listed in including is a divelse group: a thorough examination of all species, their internal anatomy, is needed before it can be properly characterized. Dístribution: Central and South America' Genus Dactylozodes Chevrolat, Ie37 & Harold Dactvlozodes chevrolat I8V7z79; Lacordaire 1857:56; Gemminger 1897 : 19O2z214-216; 1869 zI7g4-I795; Kerremans l-892 ;I75-I77' II5, Heyne&Taschenberg1907zI47;Obenberger1974z769-776 Lasionota Dejean 1876:94 [nomen nudum] ZeminaL.&G.,1839;LacordaireIS5T:55;Fairmaire&Germain1858z774; Kerremans I897zII5 Zemima (sic) Saunders 1871:60-62 AqriLozodes ThérY 1927zITI a type species' but Tvpe species: Chevro]ra:- (1877) O:-O not designate Chevrolat, 1877 and erected the genus on two sPeciest D . alternans r23 D. tetrazonus Chevrolat, 1877 (synonym of D. gadrifasciata Mannerheim, r8t7). Di stic characters Body sma].I' nallow, hirsute, subparaltel in lateral view. Head Frons with long, dense hair; eyes smaller, parallelt more (except widely separated than most other taxa of the tribe D' robustus); mouthparts not proJ-onged, clypeus very short; labrum strongly transverse, anterior margi-n straight or weakly bisinuate, without border or medial longitudinal groove; mandibl-es with one or two dorsal- subapicat teeth and one ventral subapical, long dense setae; terminal palps segments of labial and maxillary palps tapered apically, maxillary with very long setae; prementum very short, attached to mentum dorsally' margin and not exceeding it; mentum strongly transverse, anterior or 7th notched laterally in some; antennae toothed from 4th, 5th' 6th segment; antennae short, with long setae, tooth on antennal segments triangular, widest apically, strongly transverse; antennal segment 1I without dorsal fovea. Thorax sides of prothorax with incomplete edge, carinate or explanate; posterior l-ateral angles of pronotum not prolonged; pronoto-elytrat articulation almost straight; anterior prosternum not lobe enlarged; sternal cavity in most, markedly longer than wide, medial of intercoxal process rounded, not reaching meso-metasternaÌ suture; scutelh^rn up to 0., |^l elytron, subpentagonal or triangular. Elvtra flatter, with edges rol-Ied over forming a pronounced rim; interneurs costate in most, or punctate-striate with large punctuations' 9 + l-0 incompletely fused for most of length, scutellary striole reduced or absent; apical third of elytra finety denticulate; not overlapping; with long dense setae. 124 Legs posterior margin of hind coxa not strongry obrique; hind globular tarsi slightly longer than fore or mid; tarsomeres compact and tarsus; tarsal cl-aw in most, with small tarsal pads especially on fore simple. proctiger T9+L0 Abdomen s7 scarcely sexually dimorphic; femal-e fusion with or without transverse suture; male proctiger T9+1,0 incomplete;tergalmusclestlg:Il-12long'T7-T6short;sternal muscles sml-g + sLd onIY : 54-56' scarcely divided (except Female reproductive svstem DorsaL -valve q. alternans ;particlesacslongorshort,unilobedorweaklybilobed + duct without scl-erotized septum; bursa unilobed elongate, spermatheca (except species with Ìonger than bursa; 5 ovarioles/ovary undescribed 7)i accessory gland absent' (I species only); Mal-e reproduc tive system 12 tubules/testis curved or pointed' median l-obe without ventral membranous flap, sides ostium0.Tlengthfrombaserapexpointed,bilobedorrounded' A].imentarysystemMidgutcaecao.2-o.5lengthofmidgut; rods anterior to Malpighian tubules ungrouped proximally; sclercjtized junction of rectum and hindgut present in most' Dactylozodes in five Remarks: Cobos if-]95g) placed the species of toothed from the 4th' subgenera based primarily on the antennae being 5th, 6th, or 7th segment, a character also variable in castiarina' I9t5 are monotYPic' Neodactylozodes cobos , 1958 and stenozodes Théry, majority of species in parazodes cobos , rgSg has 5 species, with the Théry, 1927 wittt Dactvlozodes (s. s.). Cobos asserted that Aqrilozodes Sspecies,shouldbereducedtosubgenericstatus.Arevisionof 1985) and contai'ns Dactylozodes is in preparation by cobos (in litt., about 55 species (T. Moote, in litt, 1985)' L25 Distribution: South America' Genus HVperan tha Mannerheim, 1837 Gemminger & Harol-d Hvperantha Mannerheim IBfT:99; Lacordaire 1857l.54; 186921393-4; Saunders I87o..7, 18, 7I1 187]:60; Kerremans Heyne & I8922I33-L35, 1893:Ì15, 1897:47-47 (key), 1902:216'217; (revision); Taschenberg I9O7zI47; Hoscheck & Théry I928t125-154 Obenberget 1934 277 6-777 poecil-onota solier 18772298 (not Eschschottz, 1829) paecilonota L. & G., 1838:1-5; Gory 184I zIgI-I9V (not Eschscholtz, 1829) Hiper antha (sic) Gistel, 1874:10 [nomen nudum] (K1ug) Type species (designated by solÍer, l8t3): H. interrogationis t 1825 Diaqnostic characters BodvLength!5-25nn,sidesstraight'convergingtowardsapexor parallel, body subparal-lel or wedge-shaped in lateral view' HeadFronshairy;eyessmal]-erandmorewidetyseparatedthan clypeus very most stigmoderini, paralleJ-; mouthparts not prolonged, straight or short; labrum thick, strongly transverse, anterior margin groove; weakly bisinuate, without border or medial longitudinal 2 sharp mandibles squat with 2 or I dorsal subapical teeth, and I or maxiLlary paJ-ps ventraL subapical teeth; terminaL segment of labial and concealed; tapered apicalÌy; prementum attached to mentum dorsally and antennae toothed from mentun stronglY transverset not notched laterally; 126 transverse, tooth 5th segment; toothed antennal segments very strongly without dorsal fossa' rounded, widest at the middte; antennal segment 11 Thorax sides of prothorax carj-nate, or with incomplete edge; posterior lateral angles of pronotum Very acute, pro}onged in a hook curvedoverthehumeralanglesoftheelytra,especiallyin almost straight Hvperantha (s. s.); pronoto-elytral articulation (Hyperanthella) or strongly bisinuate (Hyperantha (s' s')); anterior as 1ong or prosternum with angular projection in some; sternal cavity scutelLum longer than wide, medial l-obe of intercoxal process rounded; wider than O.f W elytron, cordiform or subpentagonal' (in Elytra flat, interneurs punctate-striate and costate costa forming a Hvperanthel-la costae diminish basally), the outermost apices round pronounced rim, 9 + 10 often fused, and/or 7 + 8 short; with serrated with denticul-ate margin (Hyperantha (s. s.)), or bispinose very sparse interval (Hyperanthella); not overl-apping; most with short' setae. LegsPosteriormarginofhindcoxanotstronglyoblique;hind tarsi slightly longer than fore; all tarsomeres complessed l-aterally' but with large pu1villi; tarsal claw simple' female' Abdomen s7 straight or weakly notched in both mal-e and suture not but s9 not exposed; female proctiger T9+10 fusion complete, proctiger T9+l-0 discernible, posterior margin strongly notched; male fusionincomplete;tergalmusclest}gT11ong,T2-f6short;sternal muscles smlg + sld onlY: 54-56' Female reProdu ctivesvstemDorsalvalvenotdividedorscarcely sclerotized divided; particle sac shorter than valves, unilobed, without longer than bursa; 5 septum; bursa unilobed elongate, spermatheca + duct ovarioles /ovary; accessory gland absent' I27 not available; Mal-e reproductive system Number of tubules/testis conceaLing small median lobe with inflated ventral membranous flap drawn into a short thin ostir¡m 0.9 length from base, sides curved, apex point. tubules not Alimen tarv svstem Midgut caeca and Malpighian junction of rectum and hindgut avail-abl-e; sclerotized rods anterior to present, bifurcate in some' . hoschecki Species:Obenberget(Ggl,Ðlisted}gspecies,sincethenH H . hoschecki Théry, I9V4 has been added as welL as 4 new subspeci'es: Cobos L963, ornat is Cobos , L959, H. thervi oseudotestacea , cobos I96t H. decorata mauri cobos, 1963, and H. decOrata sanguinolenta ' (T. Moore, in litt., 1985)' the heading Poecilonota Remarks So1ier 0837) described a taxon under LaPorte and Esch. which he þased on Buprestis interrogationis Klug. (sic) Esch' listing Gory 0877) also described a taxon under Paecilonota to other species of what is now Hyperantha in addition described interroqationis. Mannerheim 1G817) recognized that the taxon replacement name Hyperanthat was a ne¡r genus for which he proposed the synonymizingitwithPoecilonotaasusedbySolierandL.&G. Hyperantha, attributed Hoscheck and Théry e9zÐ, in their revision of had not authorship to Lacordai-re, 1857 on the grounds that Mannerheim providedagenericdescription.Inmyopinion,however,Mannerheimhad Code of given a cl-ear indication of the genus, and under International (b) this satisfies the ZootogicaL NomencLature (1985) Article 12 O), correctly attributed to requirements of availability, so authorship is him. Furthermore, since sol-ier erected the genus now designated is the type species. H antha on the species interrogationis, this TABLE 7 Comparison of the speci-es groups of Hvpe rantha H, interrogationis group H' testacea group (27-25 nn) size small (20 mm) Iarge anteriorlY pronotun not tapered tapered more stronglY curved sides of Pronotum almost straightt not carinate anteriorlY, carinate for most of length bisinuate pronoto-elYtral straight strongJ-Y articulation never entirelY black scutel-1um* entirelY bLack elytra parallel for most narrowed from baset of length, costate strongly costate towards aPex onlY rounded, denticulate elytral aPices* truncate , denticulate between 2 strong spines * from Hoscheck and Théry (j928) L28 Hvoerantha Therefore, Hoscheck and Théry were in ellor' when splitting containing into two subgenera, to have designated the subgenus subgenus' The interroqationis as Hyp eranthella not the nominate avoid confusion' I problem needs to be addressed, but in this thesis, to ( Hvperanthell-a as they are have used the names H antha s. s.) and currently understood. The two groups were distinguished by Hoscheck and the coLour of and Théry Q928) on the basis of the elytral apices this study' the scutel-Ium. In the I species of Hyperantha examined in are listed in several additional differences wele observed and these Table 7 together with those already noted' Distribution: South America' 2 Tribe IJL0DIMORPHIN I Kerremans, I9Oz (Figures 70 -32) Julodimorphites (Polycestini Group 1) Kerremans L9O2zJ:6' 19062768; (Acmaeoderini subtribe I) Obenberger 1926235 Julodimorphae (Stigmoderini subtribe) Théry 19292 274 Julodimorphini Matthews 1985224; Be}lamy 1985:408 (White) Tvpe species (by monotypy): !.' bakewelli ' 1859 D nostic characters BodY Large, cYlindrica]' HeadEyeslarge,subpara}l.el,thoseofmalelarger'morerounded very short, mouthparts and more prominent than those of female; clypeus a rostrum, galea with long, dense setae (Figures prolonged into '04-B); mo pg c lp t¡ FIGURE 10. JuLodimorpha bakeweLl-i, characters of the head. (A) Dorsal- view of head showing small rounded antennal cavities placed frontally between the eyes, very short clypeus, and elongate mouthparts with large, robust mandibles. (B) Ventral view of mouthparts showing short, strongly transverse mentum and very long, membranous prementum and palpigers. (c) Detail of Labium showing long, narrotv ligula, and basal- segments of l-abia1 palp. (D-E) Dorsal- and ventral views of antennal segment 57 showing sensil-Ia distributed along the Lateral- margi-n of both faces. (F) Apex of antennal- segment I showing detail of the fovea. Abbreviations: c = cfypeus, li = figula, Ip = labial palp, m = mentum, ma = mandiblesr pg = palpiger, pm = pr€mentum. Scale bars = 0.5 mm (4, B); 0.1- mm (C, D, E); 0.02 mm (F). B J ¡ t tl I I Df !¡ I I \ I I !, \ a E O FIGURE ,1. Julo dimoroha bakewell-i, sternum and male reproductive system. (A) Ventral view of thorax showing the very short prothorax; divided mesosternum with meso-metasternal- suture entire; and unil-obed intercoxal- process just overlapping the sternal cavitY. (B) Dorsal- view of the male reproductive system. Only the right testis and accessory gland 1, and left accessory gland 2 are il-lustrated, and for clarity' only 9 of the approxlmately 200 tubules are drawn. (C) Ventral view of median lobe. B accessory gland spermatophoral gland accossory gland 2 A vas deferens testis lntercoxal process sternal cavitY meso- meta- sternal suture atory duct a 2 mm. FIGURE 12. JulodimorPha bakewelli, wing and ovipositor. (A) Wing venation. Arrow indicates wedge cell formed by the apical coalescence of veins 2d{t and ,dAl. In some specimens, vein ZdA, is vestigial. (B-D) Dorsal, ventral- and lateral views of the ovipositor. Shading.i¡dicates sclerotization. Abbreviations: SV = ;:]|lnu"tu"r \F = valvifer, VU = vulva, VV = ventral valve. Scale bars = 5 mm (A), 2 mm (B - D). A R Sc M cu-a 1st A A1 4thA A2 A.+3d4.' A2 B c D VU V -vy F VF I29 mandibles cylindrical, apices rounded subapicaJ- teeth very weak or absent; Iigula narrow, notched apically (Figure lOC); prementum very 1ong, attached anteriorl-y to mentum, labial suture absent, palpigers very 1ong; antennal cavities round, situated cl-ose together, between the eyes and above their l-ower margins; antennal scape very Iong, equal in length to segments 2 + 3 + 4 together, and 0.25 total length of antenna; sensill-a in shallow, terminal fossa and in secondary sensory area along edge and extending to both faces of toothed segment (Figures 70D-F). Thorax prosternum in front of procoxae shorter than diameter of procoxae (Figure 7IA)i mesosternum divided, meso-metasternal suture entire, at base of cavity, raised; intercoxal process unilobed or scarcely trilobed with nedial longitudinal depression, not filling sternal cavity; wing pointed apically, with wedge cell (Figure 32A)' Abdomen Musculature: tmtg + tpd + tllg T2-f6, tmlg fu1] height 12, 2/3 height ß-16; ts T1-T8; smlg + sÌl-g + sl-d s4-s6' smlg and sld fuI1 height, sllg 1/4 - I/2 height; tergal and sternal pores variablet often absent, esPeciallY sternal. Female reproductive svstem Long, very heavilY sclerotized ovipositor without dorsal valve, particle sac or spatulate setae, styles lateral (Figures V2ÙD) ; 9'I2 ovarioles/ovary. Male reoroduc tive svstem Testis with about 200 tubul-es/testis; two pairs of accessory glands, a ventral pair 2-3 x length of abdoment and a dorsal- pair 0.1 length of abdomen in addition to a pair of larget thick-walled spermatophoral glands (Figure rlB). Alimenta s em Foregut long, 0.7 length of midgut with crop; midgut caeca more than 0.3 length of midgut; regenerative crypts denset decreasing in size towards pyloric valve; hindgut I.5-2 x length of midgut, differentiated into small intestine (0.2 length of hindgut), and Lto large intestine (0.8 length of hindgut); Malpighian tubules ungrouped proximally and distatly, inserted about 0.6 length of hindgut. Nervous svstem I + II-III-1 + 2 + 3 + 4 + 5-8 Chromosomes Karyotype ,*7¿ 2¡ = 39 Distribution: Western Australia, South Australia, Victoria. Remarks: This tribe is monotYPic. L7l APPENDIX A was obtained' and Museums and private coll-ections from which material- abbreviations used AH Mr Anthony Hil1er, Mt Glorious, QId (tqr Holloway) AM Austral-ian Museum, Sydney e' (Dr J. Lawrence) ANIC Australian National Insect coll-ectioñ, c.s.I.R.0. AS Mr Al-Ien Sundholm, SYdneY London (lqr n' D' Pope) BM British Museum (Natural History), Papua New Guinea (Dr J. Ismay) DPI Dept primary Industry, Konedobu, Medicine of the GN prof. Gayle Nelson, college of osteopathic Pacific, Pomona, CaLifornia, U'S'A' U'S'A' GÌ'll Mr George Walters, Los Angeles, cl'ti Mr Geoff. Williams, Lansdowne, N'S'W' P.N.G (Mr P. B. Clark) IFTA Insect Farming and Trading Agency, Bu1o1o, JJ Dr James Johnson, Univ. Idaho, U'S'A' (Dr Neþoiss) MV Museum of Victoria, Melbourne A' (Or e' Monteith) QM Queensland Museum, Brisbane (Dr E' Matthews) SAM South AustraLian Museum, Adelaide N'S'W' Sv'l Mr Stan Watkins, Summer Hil-l' Chile T'N4R Dr Tomas Moore Rodriguez, Laja, I72 APPENDIX B Species of Stigmoderini dissected and aspects of internal- anatomy (A) studied (+): (M, F) male and femal-e reproductive systems; alimentary (ov) /É and nervous systems, abdominal musculature; (C) chromosomes; ovarioLes/ovary; ftlt) /É tubuLes/testis; (-) data not obtained' Species u F  c OV t/t Stiqmodera (s. s. ) cancell-ata (Donovan), l-805 + + + + 5 20 qratiosa Chevrol-at, 1874 + + + + 7 12 ìacquinoti Boisduval , l8t5 + + + 72-V4 + + 5 24-28 macularia (Donovan) ' 1805 + + porosa Carter, 1916 + + + + 5 26-34 roei Saunders, 1868 + + + + 5 70 sanquinosa Hope, t840 + + + 5 70 (Themognatha) +St¿gIqEeIa + + + 7 22 alternata Lumh. ' 1889 + barbiventris Carter, 1916 + + + + 7 85-9I bonvoufoiri Saunders, 1868 + + + + 7 43-5I brucki Thomson, 1878 + chal-codera Thomsonr 1878 + + + 5l-56 chevrolati Gehin , 1855 + + + + 7 54-75 congener Saunders, 1869 + conspicilJ.ata White' 1847 + donovani L. & G., 1877 + + + + 7 24-25 flavicollis Saunders, 1869 + + 7 fl-av inata Gem. & Har., 1869 + + + 7 28-70 gorvi L. & G., IB77 + + + + 7 72 r77 A OV t/t Species T T ! Stiqmodera (Themognatha) (cont. ) 83-94 heros Gehinr 1855 + + + + 7 maculiventris MacLeaY, 1862 + + 58 miranda Carter, 1929 + + + 7 mitcheLli HoPe, 1846 + + + + 7 19-30 + 7 3t-48 mnj-szechi Saunders, l-868 + + + 45-56 murrayi Gem. & Har., 1869 + + + 7 nickbrli 0benberger, 1922 + + + + 7 20-22 7 obscur ioennis Mannerheim , 1877 + parvicollis Saunders, 1869 + + + + 7 38 pictipes Bl-ackburn, 1894 + + 7 46 princeps Blackburn' L89I + pubicollis lVaterhouse, I874 + + + + 7 26 regia Bl-ackburn, 1892 + + + + 7 24-36 reichei L. & G., 1877 + + 7 sangu inea Saunders, 1869 + saundersi. Waterhouse, 1876 + secularis Thomson, J.B57 + + 5 tibialis V'laterhouse, I874 + + + 7 61-80 tricol-orata Waterhouser L874 + + + + 7 22-24 variabilis (Donovan), 1805 + + + + 7 28-40 viridicincta Waterhouse, 1894 + + + + 7 24-25 vitticollis MacLeaY, I86t + varrell-i L. & G. , I8t7 + + + 7 T2 sp. near saundersi + 134 Species M I A c OV t/t Stigmodera Castiarina adel-aidae HoPe, 1846 + + + + 5 l0 arqil-Lacea Carter, 1916 + + + + 5 10 bicol-or L.& G. , 1877 + + 5 biquttata MacLeaY, 1867 + + 5 chinnocki Barker, 1983 + + + 5 l0 clancul-a Obenberger, 1922 + + + 5 T2 + + 5 crenata (Donovan) ' 1805 cupreoflava Saunders, 1869 + + + 10 cupricoll-is Saunders,1868 + + 5 decemmaculata ( Kirby), l-818 + + + + 5 l0 erubescens Blackburn , 1901- + + 12 erythromelas (Boisd.), 1875 + + 5 fLavooicta (aoiso. ) , 1875 + + + + 5 10 grata Saunders, 1869 + + + + 5 L2 iospil-ota L.& G. , 1837 + + 10 + 10 kirbyi (Guerin) ' 18f0 + media Hope, 1847 + + + 5 l_0 mustelamaior Thomson , 1857 + + + 5 10 ocell-igera GorY' 1841 + 5 octomaculata Saunders, 1868 + + + 5 12 palLidipennis Blackburn' L890 + + + 5 pallidiventris L.& G., 1877 + + + 5 10 parallela lVhite , 1859 + + + 5 12 parallelipennis Obenberger, 1974 + + + 5 10 oiliventr.is Saunders , 1868 + + 10 roLlei Kerremansr 1908 + 5 t75 Species T F A ! OV tlt St Castiarina) (cont. ) rufipennis (Kirby), 1818 + + + + 5 10 sexplagiata GorY,. 1841 + + + + 5 simulata L.& G., l8V7 + + + + 5 10 subnotata Carter, 1933 + + + + 5 subtincta Carter, 1977 + + + + 5 10 terraereoinae Blackburn , 1893 + + l0 thomsoni Saunders, 1868 + + + 5 L2 triramosa Thomson, 1879 + + + + 5 10 vegeta Hope, 1847 + + + 5 10 vittata Saunders, 1868 + + + 5 12 xanthopilosa Hope, 1847 + + + 5 IO/12 Hypostigmodera variegata Blackburn, 1892 + + + 510 Calodema bifascia sp. nov. + + 7 plebeius Jordan, 1895 + + regalis L. & G., 1837 + + + 7 ribbei van de PolJ.r 1885 + + rubrimaroinatum sp. nov. + violifascia sP. nov. + 7 wallacei Deyrolle, 1864 + rv6 Species M F 4 c OV t/t Meta (van + apicalis de Pol1) ' L886 apicerubra Th êry, 1922 + cordata sp. nov. + hauseri Théry, 1926 + + qloriosa Blackburn, 1894 + + grayi Parry, 1848 + + + 7/8 meeki Théry, 1922 + + + E niqrisutura sp. nov. + sternalis Hoscheck, I93I + Conoonatha ( s. s.) q? amoena (Kirby), 18L8 + + pretioissima Chevrolat , 1878 + + Conognatha (Pithiscus) azurea PhiliPPi, 1859 + chal-ybeofasciata Ker-Ger, 1906 + compta Perty, i-8r0 + errata Fairmaire, L867 + + + excell-ens (Klug) , 1825 + fisheri Hoscheck, 1974 + germaini Théry, 1906 + klugi Gory, 1841 + laticollis PhiliPPi' l-864 + lebasi Mannerheim, I8V7 + saqittaria Fairmaire , 1856 + + 12 rt7 Species M F A c OV tlt Conognatha Pithiscus) (cont. ) sellovi (Klug), 1825 + qQ souverbi Germ. & Fair. r L855 + + subdilatata Saunders, L869 + viridiventris Solier, 1851 + + + Dactvlozodes aÌternans Chevrolat' 1878 + + conjuncta Chevrolatr l-878 + + + 5 12 cupricollis L. & G., ]838 + + + 5 12 + dispar Kerremans ' I9O2 + minor Solier, 1851 + + + 5 okea Gory, 1841 + + tr t2 rouLeti Faj-rm. & Germ. ' 1858 + + rousseLi Solier, 1851 + + 5 undescribed + + + 7 T2 Hvoerantha cardinalis (Donovan), L825 + decorata GorY, 1841 + + q? inte tionis (Klug), 1825 + + 'Lanqsdorffi (Klug), 1825 + sallei Rojas, 1855 + + st ticol-1is Desm., 1843 + testacea (Fabricus), 1808 + + + 118 The fo]-lowing species were examined external]y only: Stiqmodera ( tna) affinis Saunders, 1868; bifasciata Saunders, 1872; 1868; carpentariae Blackburn, 1892; duboul-ayi saunders, excisicollis Macleay, L862; flavocincta L. & G., 1877; fortnumi Hope, L842; gemmelli Deuquet, 1947; gigas Carter, 1916; grandis (Donovan), I8O5; haematica Hope, 1846; imperiaLis carter, 1916; jansoni saunders, Ì868; LaevicoLl-is saunders, Ì868; lessoni L. & G., 1877; limbata (Donovan), J-805; lobicollis Saunders' 1868; macfarl-ani V'jaterhouse, I88l; macqueeni Deuquet, 1964; marqinaLis carter , 1929; martini Saunders , 1869; menalcas Thomson, 1879; oleata BLackburn, 1894; pascoei Saunders' 1868; rectipennis Bl-ackburn, 1891; sanguinea Saunders, 1869; sanquineocincta Saunders, J-868; Stigmodera (Themognatha) sanguinipennis L' & G.' 1868; 1877; sanquiniventris Saunders, 1868; sexmaculata Saunderst similis Saunders, 1868; spencei L. & G., l8V7; suturalis (Donovan), 1805; thoracica saunders, 1868; unicincta saunders, L872; viridicauda Carter, l92O; westwoodi Saundersr 1868; wimmerae BLackburn, 1890; 7 sPP. G't Stiqmodera Castiarina aeraticol-lis Carter, I93O; andersoni L. & IB37; aurantica Carter, I97I; austral-asiae L. & G. , I8V7; bella (Hope) Saunderst 1871; Saunders , I87I; bilasciata ' 187L; binotata bremei (Hope) , 1845; brutell-a Thomson, 1879; carinata MacLeay' 1867; campestris Blackburn, 1897; carminea saunders' 1868; castelnaudi Saunders, 1869; cl-ancula 0benberger, 1922; coLorata Hope, 1847; cruenta L. & G., 1837; cupida Kerremans, 1898; cyanipes saunders, 1868; danesi Obenberger, 1973; delectabilis Hope, 1847; discofl-ava Carter, I97O; distincta Saunders' 1868; 139 Stiqmodera (Castiarina) distinguenda Saunders, 1869; doddi Carter, I9I3; elderi Blackþurn , !8|21 erythroptera (Boidsuval) ' 1815; flava saunders, 1869; fl-avopurpurea carter, t90B; genitilis Kerremanst 1900; qibbicollis Saunders, 1868; hatelyi Barker, 1980; hilaris Hope, 1846; hypocrita Barker, l98V; ignea Blackburn, 1892; ignota saunders , 1869; imitator carter, I93O; ;þcunda saundersr 1868; kerremansi Blackburn, 1890; l-uteipennis GorYr J-84I; malleeana Carter, I97I; marqinata Barker , 1983; nigriceps Barker, 1979; G'' obscura saunders, 1869; ocelLigera Gory, 184Ì; octospilota L. & 1837; ovata Barker, 1979; perLonga Carter, I97I; phaeorhaea (Kirby), 1818; praetermissa carter, I92I; producta saunders' 1868; pulchripes Blackburn, 1897; punctatosulcata saunders, 1869; quadriguttata MacLeay, 1863; recta saunders, 1869; robusta MacLeay 1863; rutila Deuquet 1947; Saunders , 1869; rufipes , ' scalaris (Boisduval),1835; sexuai.is Carter, 1929; skusei BJ-ackburn' IB92; unduLata (Donovan), 1805; victoriensis Blackburn, 1890; viridiventris MacLeay, 1867; ?near crockerae Barker, 1979 or subtestacea Barker, L987; + 20 sp. nov' Calodema viridiventris sP. nov. 1870; Conoqnatha ( Pithiscus) chiliensis Guerin, L87O; insionis Perty, Perty' l-8f0 ma cl-eavi (Donovan) , IB25 IHOLOTYPE]; vulnerata Dactvlozodes okea GorY, 1841 Hvperantha menetriesi Mannerheim, 1877 140 APPENDIX C Modified karyological method of Imai, Crozier and Taylor Q977) I Dissect out testes in coLd hypotonic (1% sodium citrate sol-ution) the and remove sheath and tracheae, taking care not to damage del-icate distal ends of the tubuLes where most of the cell- divisions take Place 2. Tease tubules apart and transfer to fresh hypotonic at room temperature, for 50 minutes (0.005% w/v 3 Transfer the testes to colchicine-hypotonic solution colchicÍne in 1% sodium citrate solution) for 15 minutes (cold, freshly 4 Transfer the testes to three changes of Fixative I 70 mixed 7:1 absol-ute ethanol : glacial acetic acid) for a total of minutes wiped' 5 Remove the distaL ends of the tubules and place on a freshly pre-cleaned slide slide at about 6 with one end resting on a paper tissue, incline the l-5o and apply fourteen drops of Fixative 2 (freshly mixed 2:4zI absolute ethanol : glacial acetic acid : distilled water) so that theyflowoverthematerialanddrainofftheend Fixative 2 and 7 with the slide horizontal, apply two more drops of using macerate the material as completely and quickly as possible glass two fine entomological pins bent at right angJ-es and set into rods for handles on the I with the same implements smear the cel-l suspension evenly sl-ide 141 9. Immediately, before the suspension dries, add 2 drops of glacial acetic acid to the centre, allow them to move outwards (about 20 seconds), then drain off by tilting the slide latera1ly (with long edge bottorn-most) on a paper tissue. Allow the slide to dry compJ.etelY, usuallY overnight. 10. Stain the preparation for 15 minutes with freshly mixed and filtered Giemsa stock solution diluted l:L0 with Sorensents pH 6.8 buffer. 11. Rinse the slide with distil-ted water, flick off the excess, and leave to drY. L2. Mount under a coverslip with DePex. 142 APPENDIX D Qther buprestid species in which the ovipositor and/or femal-e reploductive system was studied and compared with that of Stigmoderini (+), indicating the author, and the number of ovarioJ-es per ovary where availabLe. Author abbreviations ale as foLlows: B - BlIy 1974, !977, L980, 1982, 1983, 1984; C - Cobos 1955, !973, 1978, 1980; G - Gardner (tnis thesis); H - Holm 1979; K+C - Kasap and Crowson 1975; M - Martoja 1927; 1964; MR . Moore 1985; R - Robertson 196I; T - Tannet V - Votkovitsch Ig77, 1978, 1982; V+B - Volkovitsch and Bil-y 1979; 1945' V,l - Wandoll-eck I9O5; Wi - Williams Taxon ovr- bursa /É ovar. source SubfamiJ.y ACMAEODERINAE ACMAEODERINI Acmaeodera anatolica Vol-k. & Bily + V+B Acmaeodera chivensis Vol-k. + v78 Acmaeodera connexa Lec. + MR Acmaeodera hol-zschuhi Volk. & Bily + V+B Acmaeodera inqui-rienda VoIk. + w7 v78 Acmaeodera koenigi Ganglb. + Acmaeodera st Í Volk. + v78 v78 Acmaeodera subscalaris Reitt. + Acmaeodera sp. + I K+C Acmaeoderella alePidota Volk. + v77 Acmaeoderella candens Volk. + vTl Acmaeoderella cinerea VoLk. + v82 Acmaeodere]Ia coeLestina Volk. + v77 Acmaeoderella insueta Volk. + v77 Acmaeoderella pseudovirgulata Volk. + V+B r43 Taxon ovip. bursa lÉ ovar. source ACMAEODERINI (cont.) AcmaeoderelLa therYana (Ab. ) + v77 Acmaeoderell-a zarudniana VoIk. + v77 Atacamita biimpressa (PhiI' & Phil.) + MR Atacamita chiliensis L. & G. MR Xantheremia jelineki BilY + B,87 Xantheremia volkovitshi BiIv + B8' ACMAEODEROIDIN] Acmaeoderoides rossi (Cazier) + c55 NOTHOMORPHINI Nothomorpha verrucosa L. & G. + c55 Subfamily AGRILINAE AGRILINI Agril-us diaguita Moore + MR Agrilus sulcipennis Solier + MR Agrilus thoracicus L. & G. + MR Agrilus sP. A + 5 K+C Agrilus sp. B + + 5 G CORAEBINI Cisseis sp. + + 5 G Ethon sp. + + 5 G Meliboeus vi-olaceus Ki-esw. 5 K+C Paracephala sp. + + G 144 source Taxon ovip. bursa /l ovar. Subfamil-y BUPRESTINAE ANTHAXIINI Anthaxia al-adin Obenberger + 877 Anthaxia aspera BilY + 877 Anthaxia brodskYi BilY + 882 Anthaxia manca (L.) + B,82 Anthaxia morio F. 6 874 Anthaxia roxana BilY + BgV Anthaxia stateira BilY + B8' BUPRESTINI Buprestis auruLenta L. + + 22 G; K+C Microcastal-ia qlobithorax (Thoms.) + G Microcastal-ia incerta Cobos + c78 Microcastalia sphericollis C. & G. + c78 Nascio vetusta Boisduval + + 10 G Nascioides enysi (Sharp) + + 6 K+C 6 G Neocuris sp. A + + G Neocuris sP. B + + Philandia araucana Kerremans + MR Philandia valdiviana (Phi]. A Phil) + MR Pterobothris corrosus Fair. & Germ. + MR DICERCINI Dicerca divaricata SaY T2R r45 source Taxon ovip. bursa ll ovar. I.ELANOPHILINI Curis caloptera (Boisduval) + + 6G Curis ol-ivacea Carter + + TI/I7 G Curis splendens MacIeaY + + 6G + + 6G + + 6G + + l0 l4ri Melobasis sexpLaqiata C. & G. + + 6G Melobasis sP. A + + 6G Melobasis sP. B + + 6G Subf amily CHALCOPHORINAE CATOXANTHINI Catoxantha opulenta GorY + vll Steraps is speciosa KIug 5 M CHALCOPHORINI Capnod is tenebrionis L. + v'l EPISTOMENTINI Diadoxus erythrurus (V'lhite) + + G Diadoxus scalaris C. & G. + + 12 G PSILOTERINI 880 La S persica BilY + 146 Taxon ovip. bursa /l ovar. source Subfamily CHRYS0BOTHRINAE ACTENOD]NI Merimna atrata Hope + + T2G CHRYSOBOTHRINI Chrysobothris debilis LeC. + T Chrysobothris sP. + + 10 G Subfamily JULODINAE JTJLODIN] Aaata finchi (Waterh. ) + H Amblysterna natalensis (Fahraeus) + H Julodel-Ia qlobithorax Steven + H Julodis caffa Castel-nau + H Julodis onopordi (Fabricius) + H Neo.julodls hirta (L.) + H Sternocera orissa Buquet. + H Subfamily P0LYCESTINAE ASTRAEUS]NI Astraeus crassus V. de PoLl + + 4G POLYCESTINI Strigoptera obsoLeta Chevr. + I''t r47 Taxon ov].p. bursa /É ovar. source POLYCTESINI Chrvsoohana hol-zschuhi BilY + 884 c55 Chry soohana pl-acida LeC. + Xenopsis laevis Saund. + c80 TYLAUCHENINI Ancylotela oculata Waterh. + c77 c77 TV lauchenia crassicoLlis L. & G. + TYNDARINI Bordonia venezolana Cobos + c80 Tyndaris planata L. & G. + c55 XIROSCELINI Xyroscelis crocata L. & G. + c55 Subfamily SCHIZOPODINAE DYSTAXINI Dystaxia murrayi LeC. + G Glvotoscelimorpha marmorata Horn + + G SCHIZOPODINT SchizoPus laetus LeC. + G 148 oviÞ. bursa /É ovar. source Subfamily TRACHYINAE TRACHYINI Trachys pygmaeus F. + + 5 K+C Incertae sedis .].JLODIMORPH]NI Julodimoroha bakewelli (t'lhite) + + 7-T2 G (average of I ovaries = l0 ovarioles) I49 APPENDIX E Comparison of the number of tubules per testis in the higher taxa of stigmoderini with that of other buprestids as recorded by Kasap and Crowson (og75) or Gardner (this thesis). ll tubules Source STIGMODERINI Stismodera (7 sPP.) 12-50 G Th atha (2v spp.) 12-94 G Castiarina Q7 spp.) 10-12 G Conoqnatha (1 sP. ) 12 G Dactvl-ozodes (1 sp. ) 12 G Subf amily ACMAE0DERINAE ACMAEODERINI Acmaeodera sP. T2 K+C Subfamily AGRILINAE AGRILINI Agrilus spp. 4 K+C CORAEBINI Cisseis sp. A 5 G Cisseis sP. B 5 G MeLiboeus violaceus Kiesw. 4 K+C Subfamily BUPRESTINAE BUPRESTINI Nascioides enysi (Sharp) 4 K+C 150 /t tubules Source MELANOPHILINI Curis caloptera (goisduval) 6 G Curis corusca V'laterhouse 6 G Curis obscura Carter 6 G Curis olivacea Carter I G Curis valqoensi-s Carter 6 G Subf amilY CHRYSOB0THRINAE ACTENODINI Merimna atrata HoPe t2 G SubfamilY TRACHYINAE TRACHYINI Trachvs pygmaeus F. I K+C Incertae sedis .T.JLODIMORPHINI Julodimorpha bakewelli (White) L56 - 22L G (average 12 testes = 199) 151 APPENDIX F Comparison of the auangement of Malpighian tubules in Stigmoderini Crowson compared with that in other buprestids as recorded by Kasap and points e975) or Gardner (this thesis): (u/u) evenl-y spaced at both of insertion; (g/u) grouped proximally; (u/g) grouped distally; ß/g) grouped at both ends- Source STIGMODERINI Stiqmodera ( s. s. ) (7 spP. ) u/u* G tha (24 spp.) g/u G ,+ Castiarina (15 spp. ) u/u G CaLodema (1 sp. ) g/u G Meta (1 sp. ) glu G Conognatha (1 sp. ) u/u G DactyLozodes (5 spP. ) u/u G Subfamily ACMAEODERINAE ACMAEODERINI Acmaeodera sp. s/g K+C Subfamily AGRILINAE AGRILINI Agrilus sPP. g/u K+C Mel-iboeus viol-aceus g/u K+C .tt except S . qratiosa + except S. (c.) pallidiventris and S. (C.) erubescens L52 Source CORAEBTNI Cisseis (2 spp. ) g/u G Subfamily BUPRESTINAE BUPRESTINI Buprestis aurul-enta L. u/u K+C Nascio vetusta Boisduval u/u G Nascioides envsi SharP u/u K+C Neocuris sP. A, C, D glu G Ì"ELANOPHILINI Curis caloptera (Boisduval) u/u G Curis corusca lVaterhouse u/u G Curis obscura Carter u/u G Curis ol-ivacea Carter ulu G Curis splendens MacIeaY u/u G Curis yalgoensis Carter u/u G Curis sp. u/u G Melobasis (l spp.) g/u G Subfamily CHALCOPHORINAE EPISTOMENTINI Diadoxus erYthrurus V'lhite u/u G D. scal-aris L. & G. u/u G Subfamily CHRYSOBOTHRINAE ACTENODINI Merimna atrata HoPe g/u G L57 Source CHRYSOBOTHRINI Chrvsobothris sp. g/u G Subfamily JJLODINAE TJLODINI Julodis sp. u/g K+C Subfamily P0LYCESTINÆ ASTRAEUSINI Astraeus crassus van de PoLl glu G Subfamily SCHIZOPODINAE DYSTAXINI Glypt osceLimorpha marmorata Horn ulu G Subfamily TRACHINAE TRACHYINI Trachys pygmaeus F. glg K+C Incertae sedis JULODIMORPHINI Jul-odimoroha bakewelli (White) ulu G L54 APPEND]X G Species of Cal-odema: a preliminary revision During the course of my work, specimens of the four species of which wal]-acei Calodema is at present comprised: rega].is L. & G. , 1837; DeyrolÌe, 1864; ribbei van de PoIl, 1885; and plebius Jordan, 1895 were As examined, and four probable new species were discerned. specific revisions are outside the main aims of this study, the species descriptions and redescriptions and key to species presented in this appendix are preliminary on1y, and form the basis of a revision of calodema currently being undertaken by the author. Type specimens of al-I described species must be Located and examined, or Neotypes designated, and al-I species illustrated, before the revision is complete. Additional specimens of new species are being sought, and at present, Types have not been selected' The generic name Cal-odema is derived from two Greek words: kal-lost beauty and demas body, both of which are nèuter. under the (b) International Code of ZoologicaL NomencLature (1985), Article ,l- r "A j-n participle species-group name, if it is or ends a Latin adjective or in the nominative singular, or is l-atinized, must aglee in gender with and Articl-e the generic name with which it is at any time combined...rr, 32 (d) (ii) states that I'an adjectival species-group name that is published with an incorrect Latin termination, is to be correctedrr' The fo]lowing are two such "justified emendationsn of specific epithets: regal-is adj. from L. regal-is, -þ, -er royal should be regale; pl-ebeius adj. from L. plebeius, -3, -uffir coflìrìon shouLd be plebeium. The corrected'spellings are used in this appendix, but in 155 the main text, the currently accepted spellings have been used to avoid confusion. Key to the sPecies of Calodema I Sides of pronotum converge from base where widest; median fobe of pronotum concave, truncate; scutell-um elongate triangular, rounded in front, pointed behind, overlapping median Lobe of pronotum; elytral api-ces of female bispinose " " " '2 fr Sides of pronotum diverge from base, maximum width one third to one half Iength from base; median lobe of pronotum flat, rounded; scutelLum cordiform or lunate, abutting or overlapped by median Lobe of pronotum; elytral apices of female trispinose.... --..t 2(I) Body strongly tapered posteriorly; elytra yellow with narrow green borders on anterior, sutural and apical third of lateral margins ..regale L. & G. 2t Body subparalleL for most of length; elytra basal two fifths yell-ow, with narrow metall-ic anterÍor and sutural margins, apical three fifths dark brown with blue-green and violet reflections, with a pair of oblique yelJ-ow fasciae two thirds length from base, not reaching suture or lateral margins...... wallacei Deyrolle 156 ,(lt) Pronotum wider than elytra, sides diverging strongly from base, with angular Lateral projections; scutell-um Lunate or semicircuJ-ar; elytral api-ces of maLe bispinose; frons with sinous supra-antennal- ridges converging on vertex" ' " " "4 7t Pronotum same width as elytra, sides curved; scutel-lum subcordiform; elytral apices of male trispinose; frons sculpture irregular """'5 4(7) Lateial quarters of pronotum red, enclosing a green spot; elytra metaL1ic with single broad yeIlow fascia occupying medial third or more; apices of femaLe with sutural tooth obtuse and anterior to medial and lateral teeth, which are al-most l-eveL; shoulder rugose "'ribbei van de PolI 4t Lateral quarters of pronotum yellow, not enclosing a green spot; elytra metaLLic with two yellow fasciaer one broad, occupying the second quarter, the other only half as wide, in apical thirdt discontinous at suture; apices of female with sutural- tooth acute, just anterior to medial and cl-ose to it, lateral spine most anterior; shoul-der smooth. ....viridiventris sp. nov. 5(31) Epipleuron strongly emarginated at the leve1 of hind coxa; elytra not striate, with punctuations fine and shaLlow, and additional l-ines of punctuations, red with the following black markings: narrow border on anterior margin and suture, a transvetse, bilobed, v-shaped marking in basal third, a J-arge transverse post-medial fascia, and a pair of small medial spots in apical quarter; length greater than 35 mm...... p1-ebeium Jordan L57 5t Epipleuron not emarginated; elytra punctate-strj-ate, coJ-ouratJ-on and markings not as above; length less than 35 nrn. . " " '6 6(5') AþdominaL sternites 4-7 of femal-e predominantly yellow; elytra bl-ue-black with two narrow orange or yelIow fascj-ae, discontinuous at the suture, at half and two thirds length from base ....bifascia sp. nov. 6t Abdominal sternites 4-7 of femaLe green; elytral colouration and marking not as above ...... 7 7(6,) Elytra orange-tan to reddish-brown, with the foll-owing markings black with violet reflections: wide anterior border with bisinuate posterior margin, diminishing rapidly at shoulder; broad band on third quarter, the anterior margin almost straight, the posterior margin a shallow V, and apical fifth viol-et with blue-gIeen reflections medial-Iy...... vio1ifascia sp. nov. 7t Elytra anterior half yellow with narrow anterior border black; posterior half blue-black with bright red margin on apical quarter; the bl-ack haLf extends anteriorly with a pair of oblique bands, confJ-uent at the suture, then diverging to reach the . . rubrimaroinatum sp. nov. 158 Calodema regal-e L. & G., 1837 Stiqmodera ( CaLodema) regalis L. & G. I877t7L, Pf' 16' fig' 88 calodema regal-is saunders 1868222, PI. 4, iig. 7, 187l:63; Gemminger & Harold 1869zI4O5; Kerremans 18922142, I9O2z2O3; Froggatt 13' lIgoT):J-65, Pl. f8, fig. 7; carter 19292288, PI' 3I, fig' calodema kirbyi Hope 18402!77, Pl. 1; obenberget 1974275I regal-e: not located kirbyi: g (Hope Museum, oxford #1016) (not seen by author) var.blairi:d(BM)label-ledilwaLl-acei-varBl-airei";Ekelkei, N.Guinea,iv.!9o3,Pratt.(photographonlyseen). Colour: (live) Head and antennae bril-l-iant green with gold refl-ections. Pronotum brilliant gleen with coppery-gold refLections and pair of large, irregular, lateral red spots. scuteLlum briLliant green. Elytra yellow (often tan or brown after death) with narrow dark green anterior border diminishing laterally, narrow sutural border which posterior diminishes posteriorly, and very nallow lateral border on third only. Undersurface and legs brilliant green with copper or gold reflections. Proepisternum with pair of large, i'rregular, Ìateral red spots corlesponding to those above. Abdominal sternites 7-7 with smaller lateral red or orange spots usually increasing in size posteriorly, but in some, absent from sternites 7, 4 and 5. Presence of one or two pai-rs of spots on coxa 3 variable' frons Shape and sculpture: Head punctuation fine, event dense; moderateLy hairy. Pronotum J-arge, L/W -- O.7, broadest at base; punctuation fine, moderately dense, stronger and denser Ìaterally, 159 smooth along anterior malgin; sides al-most paralIel for haLf length' explanate anteriorly; anterior margin bisinuate, projecting strongly medially and at lateral- angJ-es; posterior margin bisinuate with concave, bi-pronged medial lobe. ScutelLum elongate triangular, rounded anteriorly, pointed behind, f]at, smooth, width 0'17 width of elytron' Elytra slightty wider than thorax; elytral interneurs long and joined basally, with fine, shalIow, dense punctuations; scutell-ary striole weak, often obscured by dark mediaL margin; shorter secondary interneurs inserted between primaries I & 2, 3 & 4, 5 & 6, 7 & 8, with additional punctuation on shouLders; intervals flat; sides subparalJ-e], then tapering strongly to pre-apical area; apex bispinose, both spines smaÌ], interval between spines oblique, the Lateral spine anterior to the mediaL. Prosternum smooth mediaJ-Iy, hairy and fineLy and densely punctuated on the sides; proepisternum with coarser punctuations, dense medially, very spalse almost smooth laterally, with deep smooth fossa in posterior angle. Mesosternum and metasternum smooth medially, hairy with fine moderately dense punctuation 1atera1ly. Abdomen smooth and shiny, punctuations very fine; male sternite 7 deeply concave. (16) Size: Males (Aust.): L 36.I-.42.8 mm, lll I4.8-I7'7 mm (PNG) : L 46.2 nm, W 17 -7 mm (1) FemaLes (Aust.): L V7.9-44.6 mm, W 15.7-18'8 mm (71) (PNG) ; L 46.8-49.3 mm, W 18.0-18.2 nn Ø) Type (from original description): L 40 ffi, W l-6 mm Distribution: Australia - Queensl-and and N.s.lv.; Papua New Guinea. General- remarks: The three female specimens examined from Papua New Guinea differ from the Austral-ian female specimens examined as follows: 160 longer; meta]lic green borders of elytra, both sutural- and postero-latera1, wider; lateral pronotal punctuation less coarse; 57 notched medially; and pronotal and sternal spots Ìarger. The type rrwal-lacei specimen in BM labelLed var Bl-airei" is regale, but the pronotal sPots are Yellow, not red. C. regaLe is distinguished from C. wallacei Deyrolle, to which it is most cJ.osely related, by the following: body slightly broader and more strongly tapered apicaIly, elytral- and sternal colouration, scutell-um more pointed behind, l-ateral apical spine of the elytron anterior to medial, posterior margin of 7th sternite of mal-e deeply concave. Specimens examinedz 4 6 *Z g Cooktown, QId (¡4V); 3 ó Cooktown, Qld' 19.v. I9I5, C. 01ive (t'¿v); 5 6 &.3 I Endeavour River, Qld (t'lv); t ó northern Queensland (MV); 1 ó Cape York' QId (t'lv); 1 ç Kuranda, N'Q' (sAM); (t"tv); 1 g Queensl-and (MV); 5 6 & 7 9 (MV); I d AustraLia, French 1g Coomra River, Qld, 9.xii.I968, S. V'J. V'lhite (SAM);2 q Queensland (sAM); I g N. Queensland (sAM); 2 6 (sAM); I ó ?cairns? (aM); I I Tannymorelr 3O.iv.I934, C. Hughes; 19 Killarney' IV.i.L9VO' E. J. D', E. Sutton (aM); 1 g Eumundi, N. C. Line (aM); I I Queensland (aM); I I Mt Tambourj_ne, Igzt, V'l. H. Davidson (QM); I I Cairns, N. Q., C. Smith (av); NG' (aM); J_ ?g Tooloom, N.S.W., i. 1926, H. Hacker 1 I 5000rAkuna I2-I3.iv.1960, J. M. Barrett (0M);2 6 &2 g (AM); 1 o Rigo C P' PNG' iv.I979 (IFTA); 1 d Queensland, Illidge (ANIC); I d cooktown, Qld (nrurc¡; I óAcacia PLateau, N.S.W., ii.1948, H. Davidson (ANIC); t I N.S.W. (ANIC); 1g Cooktown, 19.v.J-91-5, C. Olive (ANIC); J- I Mt Tambourine, QId, Davidson (ANIC); I ó (ANIC); 1 dW.A. (AM); J- ó& I I Cooktown, Qld (AM); I ó Kyogle' N.S.W., xii.!976, H.C. Hayes; I d & t I QueensLand (AM); I d'?Beltinjer River, N.S.W. (AM); I dN' QId, 161 (AM); F. Bradshaw,(AM); I I Endeavour R.' QId (nM); 1 I Richmond River I d& I q (AM); 1 ç Morobe Province, BuloJ-o Area, Papua N. Guinea, Ív.1980, P. Clark (DPI);1 9 [live] Mt Glorious, Qld' 9.L1.I983, A. Hiller, on Euodia micrococca F. MuelL. (SAM-JG iÉ8r02114). Cal-odema wal-Lacei Deyrolle , 1864 Cal-odema waLlacei Deyrolle 1864278, Pf . 2, fig. 9; Gemminger & Harol-d 1869:1405; Saunders L87I:65; Kerremans 18922142, I9O2|2O7; Blackburn l896zv5; Carter 1929:288; obenberget 1934275I. Calodema .johannae van Voltenhoven 1865:61-r Pl. l-, fig. Ir2i Gemminger & Harold 186921405. Calodema walLacei var. meeki Théry 1922:259- Types: not located. Col-our: Head and antennae briLliant green. Pronotum brill-iant green with coppery-gold reflecti.ons, and two large spots occupying the lateral- thirds, just reaching the anterior margin l-aterally but not touching the posteri-or margin, yelIow, or red (Deyrolle, 1864) or chocoLate brown (van Vollenhoven, 1865). Scutel-l-um bril-liant green. Elytra basal two fifths yellow, apicaJ. margin and suture bordered with the same colour as the posterior three fifths, dark brown with brilliant metallic blue-green and violet reflections, anterior border diminishing laterally, sutural border broadest just behind the scutellum; two obJ-ique yelIow fascia two thirds from base, not reaching the suture or lateral margins. Prosternum britliant green with violet reflections, with a pair of large, yellow spots occupying most of the proepisterna, 162 and corresponding to those above. Abdominal sternites yellow with metall-ic green posterior borders, sternite 7 metallic green medially' sternite 4 green on anterior border. Abdominal colouration may be variabl-e as Deyrolle (1864) and van Vollenhoven (1865) described the abdomen as green with lateral yellow spots, increasing in size towards the ]ast segment. Legs green. Shape and scul-pture: Head punctuation fj-ner even, dense, with smal-l- smooth spot on middle of frons; frons with short, dense hair. Pronotum Iarge, L/W = 0.8, broadest at base; punctuation very fine, moderately dense, slightly coarser: laterally, smooth along anterior margin; sides subparallel- for one third length, explanate anteriorly; anterior margin bisinuate, projecting medial-ly and at lateral angles; posterior margin bisinuate, with concave, bipronged medial lobe- ScuteLlum elongate' rounded anteriorly, narrowed behind, flatr smooth' width 0.14 width of elytron. Elytra slightly wider than thorax; striate-punctate, with fine, shallow, moderateLy dense punctuations; scutel-lary striole weak; secondary interneurs between 3 & 4, 5 & 6, 7 & 8; intervals flatr very wide between interneurs 3 & 4, 5 & 6 and scattered with additíonal- punctuations; sides subparallel- for two thirds their lengthr then curved to pre-apical area; apices truncate, bispinose, lateral spine almost level with medial-. Prosternum smooth medially, finely and very densely punctuated on the sides, almost rugose, and hairy; proepisternum with fine, sparse to moderately dense punctuation with a deep smooth fossa in posterior angle. Mesosternum and metasternum smooth mediallyr with moderateLy fine and dense punctuations laterally. Abdomen smooth and shiny medially, punctuations very fine, coarser on Lateral projection of sternite 3; male sternite 7 broadly and shalLowly notched. 167 (1) Size: Male (Aust.): L 34.8 mm, W I7'5 mm (PNG): L 40.9 mm, f'l L6 nm (fr photograph of specimen in BM) Types (from original descriptions): waLlacei L 46 mm, Itl 17 mm ohannae L 41 mm, W 16 mm Distribution: Australia - Queensland; Papua New Guinea' General remarks: The Australian male specimen examined is considerably given in sma]Ler than that from Papua New Guinea in BM or the dimensions the original descriptions. C. wal-laiei i-s grouped with C' regale lobe L. & G. on the basis of: shape of pronotum, with posterior median truncate, bipronged, not overl-apping scutellum; elongate scutellum; bispinose elytra in both sexes; and elytra not striate. It differs strongly from c. regal-e in the following: body slightly nalrower, less less tapered apicaÌ]y; elytral and sternal colouration; scutell'um pointed behind; lateral apical spine of elytron almost level with medial; maLe sternite 7 not so deeply concave' French collection (MV)' Specimen examined: I dQueensland, 5.xi.1908, C. Cal-odema ribbei van de Pollr 1885 Calodema ribbei van de Pol] 1885 z7I-32, PI. 3 fig' 5; Kerremans 1892:142, I9O2:2O3; Obenberger 1934 :75I' Types: (not seen by author), according to original descriptiont I 9 in Leyden Museum, 1 q in Collection of van de PolL' 164 Co1our: Head brilLiant metal-Lic blue-green-violet, antennae bronze. pronotum briLtiant blue-viol-et or metal-lic green mediaIly, red on tateral- quarters, the dividing line oblique from base, and not reaching anterior margin; a small blue-green spot within each red marking. Scutellum metalLic green. Elytra typically with basal and apical thirds brilliant blue-violet, middle third yetlow [in g 82O7OIK, basa] and apica] sixths are metallic green, the middle two thirds yeIlowl. Prosternum brilliant blue-greenr proepisternum gleen mediallyt red on lateraL two thirds, enclosing a small blue-green spotr colresponding to pronotum. Mesosternum and metasternum blue-gleen or violet' green, Abdominal- sternites 3r 4 and 7 of female yellow, bordered bl-ue or sternites 5 and 6 entirely blue-violet or green; sternites of male predominantly ye]Iow, sternite f with broad W-shaped green anterior border, sterni-tes 4-7 wj-th narrow green anterior and posterj-or borders and green antero-lateral spots. Legs green. The metall-ic colour in specimens varies from golden-green to blue-green to brilliant violet. Shape and scul-pture: Head finely and densely punctuated except for smooth antero-medial clypeus; vertex with deepr nalrow medial groove; frons with two sinuous supra-antennal ridges converging towards the vertex, frons depressed between the ridges. Pronotum L/W = 0'5-0'55 in dorsal- view; punctuation Very fine, moderately denset event giving a smooth appearance; two weak humps in middle of disc on either side of longitudinal impressed Iine, with a pair of shallow fossae anterior to them in some specimens; anterior margin bisinuate, with strong media] projection; posterior margin strongly bisinuate, median l-obe elongate' rounded; sides strongly angled out from base to form prominent lateral ang1e, then narrowed in a straight line to apex; maximum width about half way from base; sides with narrow rim anteriorly. Scutellum 165 Lunate, smooth, width 0.2 width of elytron, overlapped anteriorJ-y by median ]oþe of pronotum. Elytra much narrower than thorax; interneurs punctate-striate, punctuations smaLl, close, evenr with an additiOnal- row of punctuations between 7 & I and on the shoulders; intervaì-s convex, scattered with fine points; sides subparallel for two thirds length then curved to pre-apical area; apices bispinose in mal-e, the l-aterat spine more acute and slightly anterior to medial, tridentate in female, the sutural tooth obtuse and anterior to the medial and lateral which are acute and al-most Ievel. Prosternum punctuation very fine and sparse nedially, Very hairy, sides of conicai p"o."tt with coarser and with very dense punctuation; proepisternum punctuation fine and dense smooth l-ateral and posterior border, fossa in posterior angle' Mesosternr.m and metasternum finely punctuated, sparse medialIy, dense 1aterally, ridged obliquely from sternal cavity, continuing the line of anterior conical process. Abdomen smooth and shiny, scattered with very fine points; male sternite 7 broadly and shallowly notched' (1) Size: MaIe z L 75.8 mm, W (at thorax) l4'8 nm (3) FemaLe z 139.2-45.6 frrrì, w (at thorax) 17.0-18.1 mm Type (from original description) L 40 mm, I'l (at humerus) L5 mm' Distribution: PaPua New Guinea. General remarks: Differs from c. regale gloup in the shape of the pronotum, transverse, lunate scutel-L¡m overlapped by median lobe of pronotum, strongly striate elytra, and elytral apices sexually dimorphic' t66 (IFTA); Specimens examined: I d Yakopai Aseki M. P., PNG, viii.1981 I I Garaina, M. P. PNG, Vi.I98O, S I M (IFTA); 1 I Dimaga, New Guinea' iv.I974, H. ohlmus (ANIC); 1 I Papua New Guinea, central District, Brown River forest, 25 miles N Port Moresbyr iii.I97I, Shinji Nagai (op1 tfv44I7); I (abdomen missing) Mt Lamington District, Northern (AM) Division, Papua , 22.i?.1929, C. T. McNamara ' Cal-odema plebeium Jordanr 1895 Calodema Pl-ebeius Jordan 1895:22O CaJodema plebeja Kerremans I9O2z2O7; Carter 1929 1228 Calodema Plebeia Obenberger I9V4z75l Hon. f'/alter Rothschild according Type: Ç (not 1ocated), in collection of to originaL descriPtion. colour: Head dark metallic blue-green, antennae black with bronze reflections. Pronotum red with the following markings: broad black band with bronze reflections behind anterior margin, dilated medially into a triangular projection extending approximately half length of pronotum; posterior margin with narrow border and median lobe black; sides wÍth narrow metall-ic bl-ue border on anteri-or haLf' Scutellum black with violet reflections. Elytra red with the following black markings: border on anterior margin diminishing laterally; very narrow sutural- border; a transvelse, bilobed, V-shaped marking in basal third' of variabte size, constricted at suture; a large transverse post-medial fascia, wider at the suture, narrowing towards the sides, with margins sinuous; in the apical quarter, a smaller medial spot of variable shape, 167 diamond or inverted tvr or reduced to a pair of dots; apices bl-ack. Thoraic sternites brilliant blue-green-violet; proepisternum with small marginal red spots. Abdominal sternites of femal-e with small- red lateral spots on sternj-tes 4-6, and large orange-red patches on stçrnite 7; maLe sternites 4-7 red with blue posteriór margins, sternite 9 testaceous. Legs metallic blue-green. Shap e and sculpture: Head punctuation dense, moderately fine with smooth medial triangle on anterior margin of clypeus; frons with weak sinuous supra-antennal ridges, depressed between ridgesr with }ong, moderatel-y dense hair. Pronotum L/W = 0.6, broadest 0.4 J-ength from base; punctuation fine and dense, medial longitudinal groove pronounced; a pair of light i.mpressions medial-Iy towards the front and/or lateralJ-y at the middl-e in some; sides curved out from base, only very s]ightly in Some, then rounded or straight to the apex, explanate anteriorly; anterior margin bisinuate with medial projection; posterior margin strongly bisinuate with rounded median lobe. Scutellum transverset 0'2 L/W = 0.7r. rounded cordiform, anterior margin concave, smooth, width width of elytron. Elytra just equal in width to thorax; punctate-striate, punctuations smal-l- to very smaII, shallow, dense; interneurs I & 21 7 & 4, 5 & 6 joined at base, with secondary interneurs j-ntervals between them, and additional punctuations on the shoul-ders; flat; sides subparallel for two thirds of their length, widest at shouLders, then curved to pre-apical area; apex tridentate in both sexes, the sutural and lateral teeth acute, intermediate tooth obtuset teeth approximately ]evel; epipleuron strongly emarginated at Lhe l-eveL of the hind coxa. Prosternum smooth and shiny medially, hairy and rugose lateralÌy; proepisternum with moderately coarse, VeIy dense punctuations except laterally towards the base and on excavated 168 posterior margin. Mesosternum and metasternum smooth medially, sides hairs' with very dense, moderately coarse punctuation and dense fine concave' Abdomen very finety punctuated; mal-e sternite 7 deeply (ll) Size: MaIes z L 75.2-78.7 mm, V'/ 14'5-15'9 mm (10) Females: L t6.8-41.7 mm, V'l L5.4-I7 '8 mrn (sic) mm' Type (from originaÌ description), female: L 40 mm, W 27 Distribution: Austral-i-a - North Queensl-and' the GeneraL remarks: 9. plebeium differs from other Calodema in and is epipleuron strongly emarginated, co]-ouration and markings, groupedwithC.bifasciasp.nov.'C.violifasciasp.nov.and group on the C. rubrimaroinatum sp. nov. It differs from C. regale basis of: pronotum with sides diverging from base, with rounded posterior median lobe; scuteLLum transverse; and elytral apices tridentate. It differs from c. ribbei group in: pronotum same width as elytra, sides curved not with angular projections; male elytral apices tridentate; and elytra not strongly striate' (MV); V'l' Dormer' Specimens examj-ned: I 6 Ayr, N. 01d I 6 Cairns, c' F' P' Dodd (rqv); I é (Mv); r I N. Qld, Hacker (t'lv); t 6 a 2 I Kuranda' (SAM); N. Qld, I2.ii.1950' (SAM); 1 d& 1 9 (SAM); 1 9 Qld 1 óEdmonton, (aM); A. G. E. Sutton (aM); 2 ó Babinda, N' Qld' 2'ü'l-95I' E' Sutton I6 &.I g (aM); 1g N. Qld (QM); t I Edmonton, N. QLd' 22.Li'1950' A' G' F' P' E. Sutton (aM); I ó Kuranda, i.1948 (ANIC); I d Kuranda' N' Qfd' (AM); (AM)' Dodd (ANIC); I d Kuranda, ii.I9O7' F' P' Dodd 1 ó I69 Calodema bifascia sP. nov. Types: Holotype ó Lufa, New Guinea, xj-i.1979, D. A. S. F. BuloLo (ANIC) (æ #83O201E); Allotype g Lufa, New Guinea, xii.1979; D. A. S. F. Buloto (ANIC) (Jc tl87O201G); Paratypes l- ó Saurere Waria UAL M. P.' vi.I979 (IFTA) (æ #82O701I); 1 d Papua N. Guinea, owen stanley Range, Karamuki, 2,000 m, 24.ix.I973, R. Straatman (DPI) (¡e ll83IO27H); 1ç Menbi, S. M. P. PNG, iv.l98o (IFTA) (¿e #eZÚ01L); I g Papua, Kandep, 24OO n, xii.1961, f''l. ll1. Brandt (ANIC) (Jc tf83O2O1D); 1g New Guinea, Henganof, viii.1976, H. ohlmus (ANIC) (JG ç8aO2O1F); 1g New Guinea, sirunki, viii.1974, H.0hl-mus (ANIC) (¿e #A7ftzofi); 1g Toowoomba, s. Q., E. Sutton ?sneth (QM) (û #87O201A); 1 ç Dengalagu, Gembogl, iv.1974 (DPÍ ll7442)) (¡e +e¡toz7lc)i 1g Papua N. GuJ.nea, Chimu Province, Mt v'/ilhelm' 8'ooo', v.L978, P' clark {oPÍ t134427, tl.64o) (¡e #87ro27a); 19 Papua N. Guinea, chimu Province, Mt wilhelm, 8r000t, Vi.1978, P. clark oPI #744261 31679) (¡e #ÚIo27F); 19 Papua N. Gui.nea, owen stanley Range, Karamuki , zr2oot , 24.ix.I977, R. Straatman (oPI tf744r9) (¡e /¡s¡toz7D) . Colour: Head and antennae metaLlic blue-gleen. Pronotum olange oI yel1ow, with narrow anterior and posterior borders wider medialJ-y, black with green or purpte refl-ections, and a variabLe bLackish marking mediaIly, in some specimens occupying the medial third and extending from posterior to anterior margin, in others reduced to an irregular medial spot, with or without a pair of small-, lateral spots a short distance behind the anterior margi-n. scuteLlum black. Elytra shiny blue-black with two orange or yellow fasciae, discontinuous at suture; the anterior fascia one third from base, of varying width, usually (but 170 not necessarily) reaching the lateral margin, and in some following the sides a short distance before curling back towards the suture behind the anterior margin; the posterior fascia two thirds tength from the base, transverse, sinuate, not quite reaching the lateral- margin' Thorax and legs brilliant blue-green-viol-et, orange on lateral edge of proepisternum. AbdominaL sternite 3 same colour as thorax, but yellow postero-lateralJ-y; sternites 4-6 yeJ-Iow, green medialÌy and on posterÍ-or margin, the amount of green diminishing towards the apext and femal-es tending to have more green than mal-es; sternite 7 entirel-y yellow in both sexes. Shape and scuLpture: Head finely and densely punctuated, frons with somewhat irregular surface, very hairy. Pronotum L/w - 0.5; punctuation fine, moderately dense, sides explanate and centre of disc rather humped; anterior margin bisinuate; posterior margin bisinuate' median lobe rounded or pointed; sides strongly curved out from base then narrowed to apex, maximum width halfway along length; spalse short hairs on anterior pronotum. Scutellum subcordiform, width 0'1 width of elytron, overLapped anteriorly by medial l-obe of pronotum. Elytra same width as thorax; interneurs punctate-striate, punctuations very close, more conspicuous on fasciae; shallowly rugose on shoul-ders and apicaÌ]y; intervals convex; sides curved outwards from base, indented very slightly at the first fascia, subparalleL for half length, then curved to pre-apical area; apices tridentate, the sutural and lateral teeth small, pointed or obtuser VeIy weak or absent in some, the medial tooth more posterior, sharply pointed or obtuse. Prostelnum with fine, dense punctuation medially, rugose on the sides; anterior conical process acute, but less pronounced than in other Calodema e.g' C' regale or C. ribbei; proepisternum densely punctuated medially, with fine sparse T7I points laterally, and posteriorly; posterior angle almost squaret without fossa. Mesosternum and metasternum with fine, very dense 7; punctuation. Abdomen with scattered fine points, denser on sterniEe Entire male sternite 7 broadly and very shallowly excavated. undersurface covered with moderatel-y dense, erect hairs. (3) Size: Male t L 24.4-28.9 mm, W 9.1-10'5 mm Femalez L Vl.I-74.6 mm, W 11.4-I4'I mm (]0)' Distribution: Papua New Guinea. [The locality label on the Sutton specimen rToowoomba, s. Q.t is viewed as unreliablel. pronotum General- remarks: Placed in c. plebeium group on the basis of: posterior same width as elytra, sides curved, diverging from base, median lobe rounded; scutellum transverse, subcordiform; and elytral apices of both sexes tridentate. c. bifascia differs from C. viol-ifascia sp. nov. and C. rubrimarginatum sp' nov' t which it most closely resembles, in the colour and pattern of elytra, and abdominal sternites of femaLe not entirely green' two, and L' Etymo loqy: the specific epithet is derived from L' bi- fascia f. band, and refers to the two narrow orange el'ytral fasciae, which distinguish thís sPecies. Specimens examined: TYPes onIY. L72 Calodema rubri.marqinatum sp. nov. Type:1gWatutM.P.ilG,xi'1980(IFTA)GGtl92O70IJ)' colour: Head brilÌiant metallic green with goJ-d reflections, antennae blue-green. Pronotum yeIlow, posterior margin black, median lobe golden-green; anterior margin with goLden-green border confLuent with a half large dark green almost black marking with medial Lobe extending length of pronotum, and l-ateral bands parallel to anterior margin reaching sides of pronotum, middle of the l-obe shaltowly depressed and golden-green. ScuteLlum green. Elytra anterior half yellow with red nalrow anterior border black; posterior half blue-black with bright with apical margin on apical quarter; the black hal-f extends anteriorly a pair of oblique bands, confluent at the suture then diverging to reach gold the humeral call-us. Legs and undersurface brilliant greenr with and blue reflections, anterior conical process blue-violet; proepisternun with broad yellow border enclosing a small green spot' hairy, Shape and s UTC: Head finely and densely punctuated; frons shal]owly depressed. Pronotum L/!tl = 0.6; punctuation fine, moderateJ-y dense; sides explanate anteriorly; large circul-ar depression strongly antero-medialJ-y; anterj-or margin bj-sinuate; posterior margin bisinuate, median lobe rounded; sides strongly curved from base then length narrowed in a straight line to apex; maximun width about a third width of from base. ScutelLum subcordiform, smooth, concave, width 0'1 elytron. Elytra approximately same width as thorax; interneurs punctate-striate, with small-, dense punctuations, additional punctuations on shoul-ders; intervals convex; sides curved out from base' subparallel for half length, then curved to truncate pre-apical area; 177 apices of female tridentate, teeth smaLl, obtuse, the J-ateral- pair most anterior. Prosternum conj-caL process with finer sparse punctuation medially, rugose and hairy Iaterally; punctuation on proepisternum fine, moderatel-y dense, smooth laterally and posteriorly, with depression in Iateral- angJ-e. Mesosternum and metasternum with punctuation fine and spalse medially, denser and hairy laterally. Abdomen with finet moderatel_y dense punctuations and fine hairs laterally. Size: Female: L 32 nm, W I2-5 mm. Distribution: Papua New Guinea. General remarks: Placed in the C. plebeium group on the basis of: pronotum same width as elytra, sides curved, diverging from base, posterior median lobe rounded; and scutetLum transverse. Most closely resembles C. bifascia sp. nov. from which Ít is distinguished by elytral colouration and pattern, and abdominal sternites of female entirely green; and C. violifascia sp. npv. from which it is distinguished by elytral and pronotal colouration and markings. The erection of this new specÍes is tentative, pending the acquisition of more material. Etymology: specific epithet derived from L. adj. ruber, -bm, -brum red (combinÍng form tub"i-), and L. adj. marginatus, -3, -um bordered, and refers to the distinctive red border on elytral"apices. specimens examined: Type on1y. I74 CaLodema violifascia sP. nov. (IFTA) (JG #820701M) 1 g Telefomin w. s. P. PNG, iii.L979 coLour: Head metallic green with gold reflections, antennae blue-green. Pronotum orange-brown with the following dark metal-lic medialJ'y; an green markings: Very narrow border on anterior margin wider of the irregular marking antero-medially, extending for half the length pointed behind; pronotum, longer than wide, notched in front and sharpty border smaller spots behind the anterior margin; and broad pósterior to diminishing laterally. Scutel-Lum blue-green. Elytra orange-tan reddish-brown, with the foltowing markings al-most black with violet refl-ections: anterior border, width about four times J-ength of then abruptly scutell-um, with posterior margin bisinuate to interneur 5 on quarter, convergi.ng on anterior margin at shoulder; broad band third the anterior margin almost straight, the posterior margin a shallow V; and apicat firth viol-et with brilliant blue-green reflectj-ons mediatly. Legs and undersurface briLliant green, legs and conicaL process with violet reflections; proepisternum with broad reddish Lateral border enclosing a smalL green spot; and narrow yellow markings along lateral edge of abdominal sternites f-6' punctuated; vertex shape and sculpture: Head finely and very densely to with a deep furrow which divides into two shallow grooves extending antennal cavities; frons with short recurved hairs, depressed between the grooves. Pronotum L/W = O.6i punctuation Very fine, moderatel-y and fairly evenly dense; fine, tongitudinal impressed line; lateral and anterior margins with pronounced rims; a pair of shall-ow depressions on mediaL disc either side of black marking; anterior margin bisinuate; L75 sides posterior margin strongly bisinuate, median lobe narrowly rounded; about a third angled out from base then narrowed to apex; maximum width width 0'1 width from base. scuteLLum rounded cordiform, smooth, flat, ofelytron.Elytraaboutsamewidthasthorax;interneurs punctate-striate, punctuations dense, only visible on light coloured areas, with additionat punctuations on the shoulders; intervals convex; to sides curved out flom base, subparallel for half length then curved truncate pre-apical area; apices ol female tridentate, the sutural and almost medial obtuse, the l-ateral acute, sutural and lateral.teeth leveI, the medial- posterior to them. Prosternum with fine' spalse punctuation media}]y, sides of conical process rugose, hairy; proepisternum with very dense punctuation medially, less dense l-ateral J-aterally, smooth posteriorly; with shaLlow depression at punctured angle. Mesosternum and metasternum hairy, very densely laterally, fj-ner and l-ess dense mediaIly. Abdomen with fine dense punctuation. Size: Female: L 34 mm, V'l 1,.5 nrn (1)' Distribution: PaPua New Guinea' of: General remarks: PLaced in the C. plebeium group on the basis baset pronotum same width as elytra, sides curved, diverging from posterior median lobe rounded; and scutellum transverse, subcordiform' Most closely resembles c. bifascia sp. nov. from which it is distinguished by elytral colouration and pattern, and abdominal sternites of female entirely green; and C. rubrimarginatum sp' nov' from which it is distinguished by elytral and pronotal coLouration and 176 markings. The erection of this new species is tentative, pending the acquisition of more material. Etymology: specific epithet derived from L. viola f. violet, transf' vioLet-coJ-oured, (combining form -Ylo]i-), and L. fascia f. band, and refers to the violet reflections on medial and apical elytral- fasciae. Specimens examined: TYPe on1Y. Calodema viridiventris sp. nov. Type: g Papua New Guinea, Central District, Vesilogor 800t, 7.iü.I972, oPr #t44r8) (JG /É811027P). Colour: Head and antennae briLl-iant green. Pronotum brilLiant green with purpLe reflections medialJ-y, yellow on l-ateral quarters, the dividing line weakly bisinuate, paralIeI to midline starting just in front of base, and returning to the sides behind the apex, Ieaving a narrow border of green along posterior margin and wider border along anterior margin. ScuteLlum dark metallic green with purple reflections. Elytra dark bLue-green wÍth purple refLections and two full-width yellow fasciae, one broad, occupying the second quarter of the elytra, the other only half as wide, in the apical third, the left and right halves not quite meeting at the suture. Entire undersurface and legs bril-liant green except for lateral two fifths of proepisterna which are yeJ-low encLosing a small green spot. 177 Shap e and scu]-pture: Head finely and densely punctuated except for smooth antero-medial clypeus; vettex with deepr nalrota medial- groove; frons with two sinuous supra-antennal ridges converging towards vertex, frons depressed between ridges. Pronotum L/Yl = 0.56 in dorsal view; punctuation Very fine, moderately denser even, giving a smooth appearance; two weak humps in middle of disc, either side of Iongitudinal impressed l-ine, with a pair of shallow fossae anterior to them; anterior margin with strong medial projection; posterior margin strongly bisinuate, median l-obe elongate, rounded; sides strongly angled out from base to form prominent lateral angles, then narrowed to a straight line to apex; maximum width half length from base; sides with narrow rim anteriorly. Scutell-um semicircul-ar, smoothr concaver width 0.15 width of elytron, overlapped anteriorly by median Lobe of pronotum. Elytra narrotver than thorax; interneurs punctuate-striatet shoulders smooth; intervaLs convex, scattered with fine points; sides subparallel for two thirds of length then curved to pre-apical area; apices of fema1e trispinose, the sutural spine just anterior to medial- and cLose to it, the lateral spine most anterior, the interval between medial and lateral strongly oblique. Prosternum punctuation very fÍne and sparse medially, sides of coni-cal process very hairy with coarsert very dense punctuation; proepisternum densely punctuated medialJ-y, sides less dense and posterior border smooth. Mesosternum and metasternum smooth medially with fine, very dense punctuation and setae lateral-l-y; ridged obliquely from sternal cavity, continuing line of anterior conical process. Abdomen with fine, moderately dense punctuation; sternite 7 with dense setae. Size: Femal-e z L 44 ffi, W (at thorax) l8 rlil, W (at humerus) 15.5 mm' 178 Distribution: PaPua New Guinea' pronotum General remarks: Placed in C. ribbei gloup on the basis of: wider than elytra, sides diverging strongly from base to form pronounced lateraL angles, and etytra strongly striate. It is distinguished from C. ribbei by the following: two fasciae on elytra; lateral spot on pronotum ye]Iow, not enclosing a small- spot, and not reaching base of pronotum; abdominal sternites of female entirely green; elytral apices trispinose, the sutural spine acute, just anterior to medial and close to it, l-ateral spine further removed and most anterior; and humeral cal-lose smooth, not rugose. The erection of this new species is tentative, pending the acquisition of more material. gleen (combining Etvmolo OV: specific epithet from L. adj. viridis, -e, form viridi-), and L. venter -tris m. betJ-y, and refers to the distinguishing green abdominal- sternÍtes of this species. specimen examined: TYPe onlY. 179 APPENDIX H spe cies of Metaxymorpha: a preliminary revision During the course of my work, I examined specimens of seven of the eight described species of which Metaxymorpha is at present comprj-sed: gravi Parry, 1848; apicalis (van de Pol-l), 1886; gloriosa Blackburn, I894; apicerubra Théry t 1922; meeki Théry, 1922; hauseri Théry, 1926; and sternatis Hoscheck, irgtJ-. No specimens of M. dohertvi Théry were obtained. In addition, two probable new species *""" ài..erned. The species descriptions and redescriptions and key to species presented in thÍs appendix are preliminary only, and form the basis of a revision of Metaxymo rpha currentl- y being undertaken by the author. 0n1y some type specimens of described species have been seen, the rest must be located and examined, and all species illustrated, before the revision is complete. Additional specimens of new species are being sought, and at present, Types for these have not been sel-ected. Key to the species of Meta (M. dohertyi is not included, because no specimens were obtained) 1 Pronotum very fineJ-y textured, dark purple-blue; anterior prosternal process acute, projecting beyond anterior inargin; elytra bl-ue-bl-ack with sanguineous Lateral margins, weakJ-y punctate-striate, with additional punctuations on intervals; apices of mai-e bidentate with medial- tooth rounded, female trispinose; scuteLLum rounded pentagonal, medial length greater than width. .grayi ParrY 180 1r pronotum finely textured, dark blue-green' purple or black; anterior prosternal process bLunt, not exceeding anterior margin; elytra weakJ-y punctate-striate, with additional punctuations on intervals; apices of both sexes narrow, bidentate with medial tooth rounded; scuteLlum rounded cordiform, medial length less than width 2 (except I lt Pronotum smooth, briLliant green, with copper reflections gloriosa); prosternal process acute (l-ess so in apicaLis), projecting beyónd anterior margin; elytra punctate-striate' striations strong, without additional punctuation except on shoulders; apices of both sexes broadly rounded, trispinose; scutellum rounded pentagonal, medial length greater than width. 7 2(L') Epipleuron strongly emarginated at I'evel of hind coxat and deeply grooved to emargination; interneurs with very small, irregular punctuation, wi-thout transl-ucent spot belowr more than their diameter apart; elytra uniform orange or reddish-brown with bLue-black markings: anterior border nalrow, half length of scutellum medially; and marking on apical quarter, with anterior margin dilated at the suture, sides straight converging to pre-apical- area, then bordering apices; undersurface brilliant blue-green with viölet refleitions.. ' 2t Epipleuron Less strongly emarginate at l-evel of hind coxa' anteri-or gloove not nearl-y reaching emargination; interneurs with larger,.even punctuation, with a transl-ucent spot bel-ow each, less than their diameter apart; elytra basaL two thirds yelIow merging 181 into red apically, with three markings, bl-ack with blue-gleen reflections: anterior border very wide medially, trianguJ'ar, enclosing scutellum; a Large irreguJ-ar transverse fascia half-way along length; and an elongate marking extending to the apices; the three markings joined by narrow sutural borders; undersurface bril-tiant green with bluer gold or copper reflections"'meeki Thery f (1") Apices strongl-y divergent " " "4 3t Apices not strongly divergent. " " "6 4(7) pronotum brilliant green with violet reflections; sides of elytra rounded on shoulders; elytra orange with reddish l-ateral margin apicalJ-y, with very narrow dark anterior border one eighth length of scutellum, and a dark, blue-green apical marking beginning at the suture three quarters length from the base, maximum width about hal-f width of elytron, anterior margin convexr constricted in front of the apices then fl-ared to form a narrow apical border; sternite of female entirely gleen...... g1oriosa Blackburn 4t Pronotum with brilliant copper and gold reflections; sides of elytra strongly angled at the shoul-ders; elytral markings not as above; abdominal sternites 6 & 7 of femal-e fulvous " " "5 5(4') Anterior prosternal process nalrow, terminating abruPtlY, flat medially, exceeding anterior margin; elytra with large, transverse, cordiform btack marking with concave anterior margin at the junction of red and yeIlow, and two thirds width of elytra at widest, and not nearly reaching the apex. ..cordata sp' nov' I82 5t Anterior prosternal process broader, rounded apically and mediaIIy, scarcely exceeding anterior margin; elytra with transverse black marking with violet, bfue and green refl-ections occupying the apical firth, and reaching the sides and apex' anterior margin of marking sinous ..apical-is (van de Poll) process; 6(3t ) Prosternum inflated medial-Iy behind the conical elytral apices with medial spine noticeabl-y l-onger than sutural; elytra basal two thirds ful-vous, apicaJ- third red, with the following blue-black markings: anterj-or border hal-f the length of scutell-um medially; a small marking on the suture, at a quarter length from base, nallow anteriorly, at widest one fifth that of elytra together, posterior margin bisinuate; occupying the third quartert a large, transverse, bilobed marking across the suture but not tMr reaching l-ateral margins, anterior margin a flattened shapet posterior margin short, almost straight but notched at the suture; an apical marking with sinuous anterior margin, extended anteriorly at the suture sternalis Hoscheck 6t Prosternum fLat medial-ly, behind conical process; elytral apices with medial and sutural spines al-most ]evel; elytral malkings not as above ""'"'7 7(6t) Anterior border of elytra nalroy¡, I/6th width of scutell-um mediatly; scutellum rounded behind; elytra basal two thirds yellow, apical third, with or without small- black mediaL spot on the suture at the junction of red and yellow...... apicerubra Théry r83 7t Anterior border of elytra wider, I/2 width of scutellum medialj-y; scutellum more pointed behind; elytra basal half yel}ow, apicaJ- hal-f red, with elongate black sutural marking more than five times as long as wide, extending from the posterior margin of the scutel-Lum along the suture for 5/6ths the length of elytra, with irregular sinous lateral margins; sutural and medial spines bJ-ack-tipped.. niqrisutura sp. nov. Metax a grayi Parry, 1848 st a (Metaxymorpha) grayi Parry 1848:82-87 PI. 11' fig. 7 Metaxvmorpha grayi Gemminger & Harold 1869:I4O4; Saunders 187Iz65; Kerremans I892:I42, L9O2:2O3; Carter 19292289; Obenberger 1974:752 Met nata Théry 1922226I-262. Types: grayi g (BM' photograph only seen by author) rubromarginata I Moreton Bay (ANIC - on permanent Loan from Macleay Museum, SYdneY). Colour: (live) Head, pronotum, scutel-Ium and undersurface bril-liant violet blue. Antennae bl-ack with bronze reflections. E1ytra bl-ue-black with broad, sanguineous lateral borders, border irregular on inside edge, widest ãpically and narrowing towards anterÍor margin. Legs dark blue with violet or coppery reflections. Sha and sculpture: Head with fine, dense, even punctuations; vertex with shallow medial longitudinal gloove; hairs fine, short' spalse. pronotr¡m L/W = 0.4r appears smooth and shiny, punctuations very fine, r84 sparse to moderately dense, except for smooth medial- line; impressed line behind anterior margin; a small, deep, basal fossa on either sidet hal-fway from the Lateral margin to the scutel-lum; sides narrowing rapidly. Scutellum large, rounded pentagonal, width 0.4 width of etytron. Elytra punctate-striate, punctuations shal1ow, small-er laterally and apically; interval-s fLat and scattered with fine points; widest just behind the base, tapering strongly and evenly to apex; apices divergent, in the male bispinose, the lateral spine more acute and anterior, in the female trispinose, spines equal, the lateral pair directed sJ-ightly outwards and anterior to the sutural and medial pair which are Level. Prosternum with a strong, conj.cal projection exceeding anterior margin; punctuation smooth mediaIly, rugose laterally; proepisternum finely punctuated, dense mediallyt sparsel on the sides. Mesosternum, metasternum and abdominal sternj-tes smooth, scattered with fine points mediaJ-ly, with fine, moderately dense punctuation 1aterally. Male sternite 7 truncate, sternite 9 exposed. Size: Mal-e : L 2O.6 mm, Vl 8.7 nrn (1) Female z L29.8-32.0 mm, W 12-7'12.8 mm (2) Distribution: Australia - N. S. l./. and Queensland' General remarks: This species occupJ-es a position intermediate between the M. meef;i and M. 4icalis species groups. It shares with M. meeki gloup the characters: pronotum dark metallic blue or green; impressed line behÍnd anterÍor margin of pronotum weak; elytral apices of male bidentate; male sternite 7 not emarginate; and elytra shallowly punctate-striate, intervals punctuated. It shares with M . apicaLis group to which it is cl-oser, the characters: anterior prosternal process 185 acute, exceeding anterior malgin; antennal segments more broadly toothed; shape of pronotum, and sides not strongly carinate; scutel'lum rounded pentagonal, longer than wide; antero-l-ateraL margin of elytron bisinuate in l-ateral view; eJ-ytral apices of femal-e trispinose; female sternite 7 truncate; male sternite I narrow; dorsal val-ve short, L/W=O.E,posteriormarginstraight;style-bearingvalveshort' of ventral valve L/W = 0.6, posterior margin straight; sclerotization broadens posteriorly; and femaLe accessory gland present. The elytra are more strongly tapered than in M. apicalis group, but less strongly than in either M. meeki or M. hauseri.. It is easily distinguished from the other species of the genus by its colouration and pattern' Soecimens examined: I g Queensland, C. French's Coll-ection, 5'xi'I908 (MV) (.X #821I22G)i I9 (live) t'lt elorious, near Brj-sbane, Queensland' (SAM on Euodia micrococca F. Muell, I2.ii.I983r A. Hiller - JG tl87}2t5l); I ó Upper Tweed River, N' S' l"' (MV) (JG #821122F)i I? (terminalia missing), Tambourine Mt, i.L9I2, H. J. C. (ANIC); and Type of rubromarqinata (ANIC). (van l-886 ha apicalis de PolI) ' Semnopharus apicalis van de PolI 1886:240 Metaxymor pha apicalis Kerremans 1892'137, ¡gozizoz; Obenberger I934t75I' Type: not l-ocated. Colour: Head golden green with coppery vertex. Antennae green' Pronotum brii-liant coppery green with pink and gold refl-ections' 186 Scutell-um dark green with violet reflections. Elytra basal three fifths yellow, reddish behind; with violet anterior border one quarter length of scutellum medially, diminishing laterally; and a transverse bl-ack marking with violet, blue and green reflections occupying the apical fifth, anterior margin of marking si-nuous. Undersurface and legs brill-iant blue-green with gold reflections; female sternites 3-5 green, 5 with fulvous Ìateral spot, sternites 6-7 fulvous; mal-e sternites (from originaÌ description) fuLvous except for sternite f green with J-arge fuLvous spot at each side' Shape and scuLpture: Head with fine, denser even punctuation; vertex with medial longitudinaL impressed line pronounced; hairs very shortt Sparse. Pronotum trapezoid, L/lr/ = 0.5i smooth and shiny, scattered with very fine points; with well-defined impressed line behind anterior margin, and anterior tateral angles acute; þasal- depressions weak; sides just aLmost straight. Scutellum large, rounded pentagonal, widest behind anterj-or margin; width 0.4 width of elytron. Elytra striate' punctuations very small; intervals convex; interneurs 6-8 not nearly reachj-ng shoulders which are scattered with additional punctuation; sides divergent from base and strongly angled at the shouLders where widest, subparaLLel for half length, then curved to pre-apical area; apices of both sexes strongly divergent with three equal spi-nes, the l-ateraL anterior to the medial and sutural which are almost level. prosternum with conical process broader, rounded apicatJ-y and medially' scarcely exceeding anterj-or margin, smooth mediallyr lugose laterally; proepisternum finely and moderatel-y densely punctuated. Mesosternum and metasternum smooth medially, with fine, very dense punctuation Iatera11y. Abdomen smooth medially with fine points laterally; male 187 sternite 7 strongly emarginate (according to original description); not hairy Size: Female: L 28.5 mm, W 11.9 mm (1) Type, maLe (according to original description): L 30 mm, W 11 mm. Distribution: Aru Islands. General remarks: Grouped with M. gforiosa Blackburn, M . apicerubra Théry, M. sternalis Hoscheck, M. cordata sp' nov', ahd M. niqrisutura sp. nov. on the basis of shared characters listed in Table 5. It is distinguished from those species by the pronounced humeraL angles; anterior prosternal process broader, more rounded apically and medialJ-y, scarcely exceeding anterior margin; and elytral colouration and Pattern. Another specimen, ó Papua New Guinea, Owen Stanley Range, Karamuki, (æ 2r2OO n, 24.ix.L97V, R. Straatman (DPI t174427, tf3I64I) ll87IO27L) ]L26.5 mm, W 12.4 rml, is close to M. apicalis, especially in the pronounced humeral angles, but differs from the above description in having elytra entirely ful-vous except for narrow, black anterior and apical borders; scutellum pointed, not rounded behind; anterior prosternal process apex narlowr terminating abruptly, flat medialty; and sternites 3-7 green, 7 with a medial transverse spot. More specimens need to be obtained in order to determine whether this specimen lies within the range of variaþility of M. apicalis, or should be given specific status. (JG specimens examined: I g Aru Islands, H. Elgner (SAM) #821122A); I ó Aru Island (e¡,l - photograph only seen). 188 Metaxvmoroha qloriosa Bl-ackburn, 1894 Meta gloriosa Bl_ackburn 18942207-208; Kerremans l9o2z213; Carter 19292288; Obenberger I9V42752. Holotype: ó labelled: rC. Frenchts ColI. 5.11.08', 'HgLgTYPE T-7977 Metaxymorpha gloriosa BIkb.r + L handwritten and 1 printed 'Type' l-abeL (MV). Colour: Head, antennae, pronotum, and scutel-lum brilliant green with violet reflections. Elytra orange with reddish lateral margin apically; with very nalrow dark anterior border one eighth length of scutellum; and a dark, blue-green apical marking beginning at the suture, three quarters length from the base, maximum width about hal-f width of elytron, anterior margin convex, constricted in front of the apices then flared to form narrow apical border. Entire undersurface and legs of both sexes þrilliant blue-green, conical process violet. Shape and scufpture: Head punctuation finer very dense, less so medially, vertex with medial impressed line; frons hairs short' Pronotum trapezoid, LlW = 0.5; smooth and shiny, scattered with very fine and moderately dense points; with wel-I-defined impressed line behind anterior margin; basal- depressions small- or absent; sides weakly curved, almost straight. Scutellum large, rounded pentagonalt width 0.4 width of elytron. Elytra punctate-striate, punctuations sma1l, regular, dense, smaller lateralJ-y and apically; interneurs 6-8 short, not reaching shoulders which are scattered with additional punctuations; intervals convex; sides diverging from base, widest at shouJ-derst continui-ng the line of the pronotum, but not strongly angled' 189 subparallel for about half length, then curved to pre-apical area; apices strongLy divergent, with three equal spines, the lateral anterior to the medial- and suturaL which are a.l-most level-. Prosternum with conical process nalrow, terminating abruptly, flat medially, projecting beyond anterjor margin, smooth medially, rugose lateraIly; proepisternum finely and densely punctuated. Mesosternum and metasternum smooth medially, very densely punctuated J-aterally; thorax hairy. Abdomen smooth medially, with fine punctuations laterally; male sternite 7 concave, sternite 9 exPosed. Size MaIe z L 22.9-7I.7 nn, W 9.5-17.0 mm (65) Femal-e z L 2I.9-3O.7 nn, lrl 9.1-1r.7 nn (2I) Type, male; 70.4 mm, W I2.7 mm (cf from original description: L L3I tines = 27.5 fiint W 5 L/2 Lines = l-l-.6 mm). Distribution: Australia - Northern Queensland. General remarks: Pl-aced in M. apical-is group on the basis of shared characters listed in Table 5, atthough the brilliant green pronotum lacks the coppery reflections typical of the group. It is distinguished from other specì-es of the gloup by elytral marking, humeral shoul-ders rounded not angled, and prosternal process narrower and more pronounced. Specimens examined: I dKuranda, Queensland, F. P. Dodd (SAM); I d Queensland (SAM) ; 2 6 & I g Endeavour RÍver, QueensLand (SAM); I I Kuranda, QueensJ-and, i.1905, F. P. Dodd (SAM); I I Northern Queensland (SAM); 1g (SAM);21Kuranda, Q., i.L953, Geo. Brooks, (MV)r 2 6 & 19 P. c. French CoII., 11.xii.1918 (uV); 5 6 & 7 I (MV); 10 d &2 I 190 (MV); Northern QueensLand (MV); 6 6 & 1 9 Queensl-and V 6 & 4 I Endeavour River, Queensland (MV):1 dCooktown,0. (NV); t d& I I cooktown, 19.v.L915, C. Ol-ive (tqv); I d cooktown, Hacker (MV); r d c. York, O. (t4V); I d N. Queensland (OM); 2 6 (aM); L d Edmonton, N'Q', 12.ii.l-950, A. J. E. Sutton (aV); 1 dEndeavour R., Queensland (QM); I d (aM); ó Woodstock, Queensl-and (0M) i I 6 Cooktown, 19.v.191-5, C. Olive I N. Q., E. Sutton (aM); 19 Cooktown, Q., 28.xii.I948' C. Vallis, E. Sutton (aM); I ç N. QId (QM); 1 I Cairns, E' All-en (aM); I dNorthern (ANIC); Queensl-and (ANIC); I ó Cooktown, W. J. Tillyard I ú & I I (ANIC)i Queensl-and (ANIC); I g Kuranda, i.1950, G. B. 2 6 Queensl-and (nM); 1 dKuranda, i.I947 (nM); I dNorthern Queensland (nM); 1 d cooktown (nM); I óEndeavour R., Queensland (AM); I ó (AM) + Type. Metaxymorpha meeki ThérY, 1922 Meta ximoroha meeki (sic) ThérY 1922226I'262 Metaxymorpha meeki 0benberger J-9742752. Type: not located CoLour: Head, antennae and pronotum dark green with iridescent bluet purple, copper or gold reflections. ScuteLlum dark green or black. Elytra basal two thirds yellow merging into'red apically, with three markings, black with bLue-green refl-ections: the basal one triangular' enclosing scutellum, narrowing to a thin anterior border at the shoulders; a large irregular, transverse fascia haLf-way along lengtht not reaching lateral margins; and an elongate marking extending to the apices; the three markings joined by narrow sutural borders. 191 undersurface and legs brilliant gleen with blue, gold oI copper refLections. Shape and sculpture: Head punctuation fine, moderately dense, anterior clypeus smooth; frons hairy. Pronotum ¡7¡'¡ = 0.5f punctuation fine' moderately dense, very dense on lateral margins; sides curved to apext strongly carinate; lateral basal depressions vely shallow or absent' Scutell-um large, rounded cordiform, width 0.3 width of the elytron' Elytra punctate-striate, punctuations shallow, smaller lateralJ-y and apically, additionat punctuations between interneurs 3 & 4, 5 & 6, 7 & 8, especialJ-y on the shoulders; intervals flat; sides angled outwards from base, widest just behind the base, subparallel' for half J-ength, then narrowed 1n a straight line to pre-apical area; apices divergent, bidentate in both sexes, the medial tooth obtuse, angular or rounded, the l-ateral spine small, anterior to medial. Undersurface with fine punctuation, spalse mediallyr very dense ]aterally; hair dense, unevenly distributed; anterior prosternal process blunt, broadly rounded, not extending beyond anterior margin, and rugose laterally' MaIe sternite 7 sha1J-owly notched, sternite 9 exposed. SÍze: Males z L 29.I-tO.7, W 12.0-12.6 mm (2) Femal-es t L 29.5-32.5, W I2.1-1r.8 mm (5) Type (from original. description): L 73 mm, V'l l-1'5 mm' Distribution: PaPua New Guinea. General remarks: M. meeki differs from M. gravi and M. apicalis group in a number of characters (listed in TabLe 5), and is most closely rel-ated to M. hauseri Théry. The specimens examined varied in the size 192 and shape of the elytral markings. In specimen ($ l1zO7OIF), the anteri-or marking is a diamond shape enclosing the scutelLum and scarcely reaching the anterior border; the mediaÌ marking is much smalLer than in other specimens, each half resembì-ing a normal cuIve with the base along the suture, flanked by a small spot on each side; and the apical marking is smaller, parallel-sided for most of length, constricted at the apex' to form a nallolv border which extends to the lateraL spine' In addition, the pronotum is coppely, the sutural tooth of the elytron rounded, and the femqle proctiger mole strongly notched on the posterior border. In specimen QG llt'zO7OID), the anterior triangular elytral marking is larger than in the preceding specimen, but smalLer than the others examined; from the anterj-or marking a narrow sutural border extends a third of the length then expands into an elongate marking which almost reaches the apices, the marking is about haLf as wide as the elytron, constricted hal-f-way along its length, then parallel-sided; the apices are ted, except for a faint black tip on the sutural tooth' preceeding Specimen Qg +AÐZO2-A) has markings similar to that of the specimen, but the basal- one is reduced to a narrow border on anterior margin and around scutellum; and the posterior margin is narrower, l-ess than half the width of elytron, and not connected to the basal by any sutural- border. V,/ithout more specimens it is not possible to ascertain the range of variability of elytral markings in this species, and these specimens are best considered as conspecific with M. meeki' (SAM) (¡C Specimen s examined: I g Hudewa, New Guinea, Rev. L Wagner (æ lt82ll221);1g V,latut R., vlatut M. P., PNG' viii.1981 (IFTA) Papua #82O7O]:D); 1g l/rlatut M. P., PNG, v.1980 (IFTA) QG tt82O701F); 19 N. Guinea, Madang Distr., Karamuki 2r2oo m, in carcase of dead marsupial, i.L974, Petrus Banga (DPI) (JG #8r1027K)i I ó New Guinea, l9v Menyama, x.I97I, M. Mackel-Iar (ANIC) (æ ll87O2OZA); I g ltlatut M' P'' PNG, v.1980 (IFTA) (æ tl82o701F); 19 Watut River, watut M. P.' PNG' viii.1981 (IFTA) (æ tf82o70lD). Metaxymorpha apicerubra ThérY, 1922 Meta apicerubra Thery 19221259-26O; Obenberger I934:75I. Type: not located. Col-our: Head and antennae brilliant green, vertex with coppery-gold reflections. Pronotum and scutellum briLliant green with coppery-red and gold reflections. Elytra basal two thirds yellow, apical third red; with narrow bLack anterior border one sixth J-ength of scutellum and with or without small black medial spot on the suture at junction of red and yeJ-low. Undersurface and legs brilliant green with copper, gold and víol-et refl-ections; mal-e abdomen entirely green except for small, l-ateral fulvous spots on sternites 6 and 7' Shape and scuLpture: Head with fine, dense, even punctuation; vertex with strong medial longitudinal impressed Line; hairs short' Pronotum trapezoid, L/w = 0.5; punctuation fine, moderately dense; with well-defined impressed Line behind anterior margin; basal depressions smal-l; sides almost straight. scuteLlum rounded-pentagonal, posterior margin rounded; width 0.4 width of elytron. Elytra punctate-striate; punctuations smal], even, with translucent spots below; shoulders scattered with additional punctuations; intervals convex; sides of elytra diverge from base, contS-nuing the line of the pronotum; angled at 194 the shoul-ders where they are wi-dest, subparallel- for half length' then curved strongly to pre-apical area; apices not divergent, with three equat spines, the Lateral spine anteri-or to the mediaL and sutural which are al-most level. Prosternum with conical process rounded apically, fl-at medialÌy, exceeding the anteri-or margi-n, smooth and shiny medially, rugose and hairy laterally: proepisternum punctuation fine and moderately dense. Mesosternum and metasternum smooth medially, with fine, dense punctuation laterally. Abdomen smooth medially with fine points laterally; posterior margin of male sternite 7 strongly and broadJ-y emarginated, sterniie 9 exposed. Size: Mal-es z L 26.5-70.2 mm, Ìrl 10.6-II.7 mm (2) Type (from original description) male: L 32 nm, ftl 12 mm' Distribution: Papua New Guj-nea. General remarks: Placed in M. apicalis group on the basis of characters listed in Table 5, although elytra are more strongly narrowed than other species of the group. It is distinguished from M. apicalis (van de Poll) by having apices not strongly divergent, red, lacking broad apical marking; from M. cordata sp. nov. by the elytral marking a smaLl sutural spot or absent altogether, and abdominal sternites of male green; and from M. nigrisutura sp. nov. by eJ-ytra] markings and scutellum more rounded behind. Another specimen examined, JG /É8302028, resembled M. apicerubra, but differed in havÍng elytral apices with narrow black border. Without additional specimens, it is not possible to determine whether it is conspecÍfic with M. apicerubra, or if a new species shoul-d be erected to accommodate it. t95 vesilogo, 800 m, Specimens examined: I d Papua N. Guinea, central Dist., 7.iii.1972, R. Straatman @PI #71643, t134422) (JG /É8',IO7I); and I d seen by Ekeikal, New Guinea, iv.1903, Prat (BM - photograph only author).Specimenwithblack-borderedapices:dNewGuinea'Wabag' ií.L975, Mark Feist (ANIC) (¡c + 83O2O2B) lL 26.5 mm' lrl t0'6 mml' Metax a dohertvi Théry, L922 1974275I' Metaxvmorpha dohertvi Théry 19222262-263; Obenberget from the original No specimens wele obtained for examination. Judging description,M.dohertyishareswithM.meekigroup:scute].lum strong cordiform; elytral apices wíth smal], external, acute tooth and punctuated; and obtuse tooth; sutural angle rounded; elytral intervals prosternal projection flattened, J-ess convex; and with M . aoicalis group:pronotumbri]-]iantgreenwithcopperref]ections,with strongly wel-l-defined impression behind anterior margin, and sides not carinate. ha hauseri ThérY, 1926 Obenberger MetaxvmorPha hauseri Théry 19262167-168; Carter 19292289; 19742752. Type: not located. 196 co]our: Head, pronotum, scutel-Ium, undersurface and legs brill-iant blue-green with violet refl-ections. Antennae bronze' Elytra orange or reddish-brown with blue-black anterior border half length of marking on scuteLLum nedially, diminishing J-aterally; and blue-black apical quarter, with anterior margin dilated at the suture, sides straight, converging to pre-apical area, then diverging to reach the lateral margin just in front of the apical spines' w shortt Shape and scuJ-pture: Head finely and densely punctuated; ith sparse hairs. Pronotum L/W -- 0.6; punctuation moderately fine, dense present or and even; sides curved to apex, carinate; basaL depressions absent. scutell-um J-arge, rounded cordiform, width 0.4 width of etytron. Elytra interneurs punctate, weakly striate, with additional punctuations between interneurs J & 4, 5 & 6, 7 & 8, especially on the shoulders; sides angled out from base, widest just behind the base' then tapered to apex; apices bidentate with medial tooth rounded, lateraL tooth small and sharp; epipleuron deeply grooved and very strongly emarginate at the level of hind coxa. Thoracic sternites moderately finely and densely punctuated, less dense medially and very dense on proepisternum; anterior conical process blunt, broadly rounded, not exceeding anterior margin. Abdomen finely punctuated, density increasi-ng apically; maLe sternite 7 shallowly notched, sternite 9 exposed. undersurface with short, moderately dense hair. (14) Size: Males ; L 25.2'VO.3 mm, lt'/ 10.3-12.5 mm Females z L 26.9-VO.9 mm, ltl 11.2-13'1 mm (5) Type (from original description): L 50 mm, W 12 nrn' I97 Distribution: Austral-ia - North Queensland' General remarks: Grouped with M. meeki Théry on the basis of shared characters Listed in Tabte 5, but differs from it in col-ouration and pattern; in the epipleuron deeply grooved and very strongly emarginate at ]eveI of hind coxa; and interneurs scarcely striate. (MV); (0M); I d & I Specime ns examined: 2 ç Kuranda, F. P. Dodd I I I (ANIC); Kuranda, i'1947 Kuranda, ii.I95I, G. 8., (.:. B. Brooks Beq.) t ç (AM);2 ó(MV); 1 óQueensLand (MV); I dKuranda, N.Q.,1l.xii'1908' P' (¡¡V); C. French (¡,tV); 1 d Kuranda, N.Q., i'I95V, J' G' Brooks I ó (SAM),7 ô Kuranda, Q., ii.I952, Geo. Brooks (VV); I óCairns, Q., Dodd (aM); d (aM); I d Hambledon, N. Q., 2.ii.1950, A. Johnson, E. Sutton I Kuranda, N. Queensl-and, F. P. Dodd (ANIC); 1 dKuranda, iii'I907, F'P' Dodd (AM); and 19 Kuranda, G. Brooks (gl'4 ' photograph only seen)' Metaxvmoroha sternaLis Hoscheck, I9VI Metaxvmoroha sternalis Hoscheck I97I:73I Metaxymorpha aternalis (sic) Obenberget 19742752. Types: not seen bY author. green colour: Head, antennae, pronotum and scutellum briLliant metall-ic with coppery-red and gold reflections. Elytra basal two thirds fulvous, apical third red; with the following blue-bl-ack markings: anterior border half the length of scutellum medially, narrowing rapidly quarter towards l-ateral margin; a small marking on the suture, at a 198 length from base, narrow anteriorly, at widest one fifth that of elytra quarter, a together, posterior margin bisinuate; occupying the third large, transverse, bilobed marking across the suture but not reaching tMr lateral margins, anterior margin a ftattened shape, posterior margin short, almost straight but notched at the suture; an apicaJ- marking with sinuous anterior margin, extended anteriorly at the suture. undersurface and legs brilliant green' prosternum with blue reflections, proepisternum with coppery refJectj-ons' punctuation, and deep Shape and sculpture: Head with fine and dense medial longitudinal groove with edges raised; hairs short and dense' fine Pronotum trapezoid, LlW = O.4i smooth and shiny, scattered with points; with well-defined impressed line behind anterior margin; basal depressions weak or absent. scutellum large, rounded-pentagonal, 0'4 widest just behind anterior margi.n; posterior margin rounded; width width of elytron. Elytra punctate-striate; punctuations smal]' event dense, smaller laterally and apically; interneurs 6 and 7 not reaching the shoulders which are scattered with additional- punctuations; intervals convex; sides divergent from base and angled at the shoulders pre-apical where widest, subparatl-el for half length then curved to area; apices not divergent, with three subequal spines, the lateral with spine most anterior, the medial spine most posterior' Prosternum anterior conical plocess rounded apically, exceeding anterior margin, with a definite medial hump posterior to it behind a short, ttansverse gEoove,sidesrugose;proepisternumfinelyandverydensely punctuated. Mesosternum and metasternum with fine punctuatÍon, sparse medially, dense taterally. Abdomen scattered with very finet moderately dense points; underside hairy on thorax and third abdominal- sternite. 199 Size: Female: L 25.7 M, W 10.9 mm (I) Type (from original description) z L 26 ffi, W 11 rnrn' Distribution: PaPua New Guinea' General remarks: Placed in M. apicalis group on the basis of shared characters listed in Table 5. It is distinguished from other species of that group by the distinctive elytral markings; medial- elytral spine anterior most posterior; and medial hump on the prosternum, behind the conical process. AnotherspecimenrgSambioM.P.PNG'iii.1981'(IFTA) to (JG #82O701C), L 22.5 mm, W 9.4 mm, was examined which is close from it in the u. sternaLis Hoscheck in shape and sculpture, but differs following ways: posterior margin of scutelLum more pointed; anterior sides border of elytra one quarter the width of the scutellum medially; large' of elytra J-ess angled at the shoulders; anterior marking rounded; fMr transverse marking with anterior margin almost straight, not shapedt posterior margin narrowel and joined to apical marking by a broad sutural band; and sutural spine level with medial, not anterior to it' speci-men is More specimens need to be obtained to determine if this be accorded within the range of variability of .M. sternal-is or should specific status. Wagner (SAM) (¡g specimen examined: I g Warco Finsch Haven, Rev. L. #870r20A). 200 MetaxvmorPha cordata sP. nov' (¡C Holotvoe: 1 d Maprik, E. S. P. PNG (IFTA) # 8207014) ALlotype: I I Papua N. Guinea, Central Dist', Vesilogo' 800 m' 7.iii.1972, R. Straatman (DPI # 3L64t, tlt4422) (¡c + 87IO27J)' green with colour: Head, antennae, pronotum and scutelLum britLiant yelIow, apical third coppely-gol-d reflections. Elytra basal two thirds red; with narrow bl-ack anterior border one fifth length of scutellum; margin at and large, transverse, cordiform marking with concave anterior at the junction of red and yellow, and two thirds width of elytra legs bril-liant widest, and not nearLy reaching apex. undersurface and and 4 green, green with gold and vj-ol-et reflections; male sternites f sternite 5-7 orange-red, femaLe sternites 7-5 green, 5 with olange spot, green, and 6 orange with broad anterior and narlow posterior margins sternite 7 orange-Yel}ow. punctuation; vertex Shape and sculpture: Head with fine, dense' even with medial tongitudinal impressed line; hairs short' Pronotum with trapezoid , L/w = 0.5; punctuation fine, moderately dense; depressions weLl-defined impressed Line behind anterior margin; basal- posterior small; sides al,most straight. scutellum rounded pentagonal, punctate-striate; margin rounded; width 0.1 width of elytron. Elytra 6 to punctuations smaLl, even, with translucent spots below; interneurs I not reaching the shoulders which are scattered with additional punctuations; intervals convex; sides of elytra divergent from base, shoulders continuing the l-ine of the pronotum; strongly angled at the curved to where they are widest, subparallel for hal-f length, theri pre-apical area; apices strongly divergent with three equal, 20l- black-tipped spines, the l-ateral spine anterior to the medial and sutural which are almost level. Prosternum with conj-cal process narrou/, terminating abruptly, fl-at medially, exceeding the anterior margin, rugose and hairy lateraIly: proepisternum punctuation fine and moderatel-y dense. Mesosternum and metasternum smooth mediallyr with fine, dense punctuation laterally. Abdomen smooth medially with fine points J-aterally; posterior margin of male sternite 7 concave, sternite 9 exposed. Size: MaIe z L 29.0 mm, W I2.0 mm (t) Female: L 27.9 mm, !f ll.5 mm (1). Distribution: PaPua New Guinea. General remarks: Placed in M. apicalis group on the basis of characters listed in Table 5. It most closely resembles M. apicerubra Théry and M. apicalis (van de Po11), and is distinguished from M. apicerubra by the J-arge, transverse, cordiform black mark, two thirds the width of the elytron at the widest; elytral apices strongly divergent; and male sternites 5-7 orange; and from M. apicalis by anterÍor prosternal process narrow, terminating abruptlyr flat medially' exceeding anterior malgin, and by elytral markings. The erection of this new species is tentative, pending the acquistion of more material. Etymology: from L. adj. cordatus -3-Um, heart-shaped, and refers to its distinctive elytral marking. Specimens examined: Types only. 202 Metax niqrisutura sp. nov. (IFTA) (JG Type: t ç Namiwe via Menyamya, M. P. v.1980 #8207018). Colour: Head green with brilliant coppery-pink and gold reflections especially on frons; antennae green with blue refLections. Pronotum coppery-pink and metal-lic green. scutellum green with coppery reflections but less brilliant than pronotum. E1ytra basaL half bright yel1ow, apical haJ-f bright red, the two colours merging in an irregular transverse l-ine; with the following duIl black markings: anterior border half the Ìength of the scutellum medially, narrowing rapidly towards the sides; elongate sutural marking more than five times as long as wide, extending from the posterior margin of the scutellum, along the suture for 5/6ths the length of elytra, with irreguJ-ar sinous Lateral margins; sutural and medial spines black-tipped. undersurface and legs green with copper and gold reflections, sides of prosternal process with violet reflections. Shape and scul-pture: Head hairy, with fine, moderately dense punctuation; vertex with fine medial longitudinal impressed Line, frons with a pair of weak supra-antennal ridges which converge where j-mpressed line ends. Pronotum trapezoíd, L/W -- O.5; with smooth medial line, scattered with fine punctuations, denser anteriorly and laterally; wel-I-defined impressed line behind anterior margin; l-ateral angles acute; no basaL depressions; sides gently curved, aJ-most straight. Scutetlum large, rounded-cordiform; width 0.4 width of elytron. EJ-ytra punctate-striate, punctuations sma1l, even, dense, with transl-ucent spot under each; intervals weakly convex; interneurs 6 to I not reaching shoulders which are scattered with additional punctuations; sides 207 divergent from base continuing the l-ine of the pronotum, angled but not strongly at the shoulders, then curved strongly to pre-apical area; apices scarcely divergent, with three equal spines, the lateraL anterior to the medial- and sutural which are level. Prosternum with conical process rounded apicaJ-ly, flat medially, exceeding anterior margin. Undersurface, smooth or scattered with fine points mediallyr finely and densely punctuated laterallY. Size: Femal-e: L 29.2 mm, W 11.3 nm. Distribution: Placed in M. apical-is group on the basis of shared characters l-isted in TabLe 5. Closely resembl-es M. apicerubra Théry in elytra more strongly narrowed than other species of the group, and abdominal sternites green; but differs from it in the broader anterior margin, scutel-lum more pointed behind, Ìong prominent sutural mark, and red of apices extending hatf-way along elytra. The erection of this new species is tentative, pending the acquisition of more materiaL' Etymology: from L. adj. ni.qer -gra -grum black (combining form nigri-), L. sutura -ae f. suture, and refers to the distinctive elytral mark. Specimen examined: Type on1y. 204 Arnett, R. H. (1960). rrThe beetles of the United States (a manual for identification)". (The Catholic Univ' Press, V'lashington' D. c.). 11J2 PP. Arnold, E. N. (1981). Estimating phylogenies at low taxonomic Levels. Z. zool. Syst . Evol-ut.-forsch. 19, l-t5. Asana, J. J.r Makino, S., and Niiyama, H' Og+Z)' A chromosomal- survey of some lndian insects. IV: On the sex chromosomes of some species of beetles (Col-eoptera). Cytofoqia E, r87-2O5. Barker, S. Q97Ð. Revision of the genus Astraeus LaPorte & Gory (Coleoptera: Buþrestidae). Trans. R. Soc. S. Aust. ee(3), 105-141. Barker, S. 0977). Astraeus (Col-eoptera: Buprestidae): A description of three new species and new loca.l-ity records' Trans. R. Soc. S. Aust. 101(1) ' l-l-l-4. Barker, S. Q979). New species and a catalogue of Stiqmodera (Castiarina) (Coleoptera: Buprestidae)' Trans. R. Soc. S. Aust. lol(1) , r-27. Barker, S. (1980). New species and new synonyms of Stiqmodera Castiarina (CoLeoptera: BuPrestidae). Trans. R. Soc. S. Aust. 104(1) , r_7. Barker, S. Q987). New synonyms and new species of Stigmodera Castiarina (Coleoptera: BuPrestidae). Trans. R. Soc. S. Aust. ro7(7), 179-169. Barker, S. and Inns, R. Qe76). Predation on Stigmodera tibialis bY a fIY. lrtest. Aust. Nat. 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A check l-ist of the Australian Buprestidae. Aust. ZooI. 5, 265-704. Carter, H. J. (I93D. Notes on the genus Stiqmodera (Family Buprestidae) together with descriptions of new species of and a retabul-ation of the subgenus Castj-arina . Aust. ZooI. 6, 337-367. Chapman, R. F. (L982). rrThe insects, structure and functionr'. lrd ed. (Hodder and Stoughton, London). 819 pp. Chevrolat, A. (1877). rrRevue entomologiquefr. vol. 5. (G. Silbermann, Strasburg). Cobos, A. (I95V). Revision de los Jul-odis Esch., de Ia Peninsula Ibérica (Coleoptera, Buprestidae). 8u11. Inst. Sci. nat. Belq. 29, 1-16. 208 cobos, A. o95Ð. Estudio sobre los Ptosimites de ch. Kerremans (Coleoptera, BuPrestidae). BuLl. Inst. Sci . nat. Belo. 3I, I-24. cobos, A. Q959). Novena nota sobre Bupréstidos neotropicales, rectificaciones y descripciones diversas (coleoptera, Buprestidae). Archos Inst. Acl-im., ALmeri-a 8, 29-4t. (Coleoptera, Cobos, A. Q977) . Dos nuevas tribus de bupréstidos Buprestidae). Eos 9, 87-104. cobos, A. 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Q964). ButterfLies and plants: a study in coevoLution. Evolution, Lancaster. Pa 18,586-608. Erwin, T. L. (1974). Studies of the subtribe Tachyina (Col-eoptera: Carabidae: Bembidiini), Part 2: A revision of the New Worl-d - Australian ge nus Pericompsus LeConte. Smithson. Contr. Zool-. 162, l-96. 2ro Eschschortz, J. F. {ogzg). "Zoolgischer At1as, enthaltend Abbildungen und Beschreibungen neuel Theirarten, wahrend des FLottcapi-tains van Kotzebue zweiter Reise r,¡m die we1t, auf der Russisch-Kaiserlichen (G. Kriegsschlupp Predpriaetie in den Jahren 1827-1826. Reimer, Ber1in). Erstes Heft' f7 PP' Estabrook, G. F. Q977). Does common equal primitive? Syst. Bot. ?, 36-42. Etheridge, R. and Olliff, A. S. (1890). The Mesozoic and TertiarY insects of New South Wales. Mem. I. Surv. N.S.l¡'l. Pal-aeontofogy, No. 7. forest Evans, W. G. (1966). Perception of infra-red radiation from fires by Melanophil-a acuminata De Geer (Buprestidae, Coleoptera). Ecology 47, 1061-1065. Fairmai-re, L. and Germain, P. (1858). Révision des Coléoptères du ChÍIe, FamilY BuPrestidae. Ann]s Soc. ent. Fr. 6, 709-742. Farri-s, J. S. (re73). A probability model for inferring evolutionary trees. Syst. ZooI. 22, 25È256. Farris, J. S. (1978). Inferring phylogenetic trees from chromosome inversion data. Svst. ZooL. z, 275-284. Forbes, W. T. M. Qe22). The wing venation of the Coleoptera. Ann. ent. Soc. Am. Þ, 328-752. (1887-1913), Fowler, W. IÂl. Q9I2). "British Coleopterarr. 6 vols' and Paussidae. "Coleoptera, general introducti.on and Cicindelidae Fauna of British Indiar'. (London). (original not seen)' Froggatt, w. w. tI9O7l. rrAustral-ian insects". (lllilliam Brooks & Co., Sydney). zLt I'CataJ-ogus Gemminger, M. and Harold, B de. (1869). coleopterorum' 5 hucusque descriptorum, synonymicus et systematicusr'. vol' Buprestidae, Trixagidae, Monommidae, Euchemidae, Elateridaet Cebrionidae. (f. Deyrolle fils, Paris) ' Gerber, G.H., Church, N.S. and Rempel, J'G' Q97I) ' The anatomy' histology, and physiology of the reproductive systems of Lvtta nuttalti Say (Coleoptera: Meloidae). I. The internaJ. genitalia. Can. J. 7ool-. 49, 523-533. Gistel-, J. N. F. X. {j'Ú4). rrDie Insecten-Doubletten aus der (X' Sammlung des Hrn. Grafen Rudolph van Jenj-son Wa]worth, b' Kammerer und obertforst-meister zu Regensburg), wel-che sowohl- in Kauf al-s Tausche abgegeben werden I. Kaferrr. (Munchen). 36 pp. (original not seen). Ann. ent. Good, H. G (1925). Wing venation of the Buprestidae' Soc. Am I8, 25j.-276. Gory, H. (1841). t'Histoire natureLle et iconographie des insects Paris) coléoPteresrr. voI. 4 Suite aux buprestides. (P. Dumenil, ' male Gwynne, D. T. and Rentz, D. C. F. (Gg83). Beetles on the bottle: buprestids mistake stubbies for femal-es (coleoptera). J. Aust. ent. Soc 22, 79-80. (white) Hadlington, P. and Gardner, M. J. Q959). Diadoxus ervthrurus (Coleoptera - Buprestidae) attack on fire-damaged Call-itris spp. Proc. Linn. Soc. N.S.W. 84, 325-7VL. Heberdey, R.F. Q9VD. Zur Entwicklungsgeschicte, vergleichende Anatomie und Physiologie der weibLichen GeschlechtsausfÜhrwege der Insekten . z. Morph. öXol. Tiere 4, 416-586. Hecht, M. K. Q976). Phylogenetic inference and methodoLogy as appl-ied to the vertebrate record. Evol. Biol. 9r 335-36t. 2L2 Hecht, M. K. and Edwards, J. L. 0976). The determination of parallel or monophyletic reLationships: the proteid salamanders - a test case. Amer. Nat. 1l-0' 653-677. (univ. Illinois Pr. Hennig, v,/. (1966). "Phylogenetic systematicsr'. ' Urbana). (reprinted 1979). 263 pp- Heyne, A. and von Taschenberg, E. C. trl. (1907)' "Die exotischen Kafer in Wort und BiLd't. (Heyne, Leipzig). Holm, E. (1978). Monograph of the genus Acmaeodera Eschscholtz (coleoptera: Buprestidae) of Africa south of the sahara. Entomoloqy Mem. DeP. aqric. tech. Serv. Repub. S. Afr. No. 47. HoIm, E. 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