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Revision of the dung beetle genus Temnoplectron Westwood (Coleoptera: Scarabaeidae: Scarabaeini) PDF

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Preview Revision of the dung beetle genus Temnoplectron Westwood (Coleoptera: Scarabaeidae: Scarabaeini)

REVISION OF THE DUNG BEETLE GENUS TEMNOPLECTRONWESTWOOD (COLEOPTERA: SCARABAEIDAE: SCARABAEINI) C.A.M. REID AND R.I. STOREY Reid, C.A.M.& Storey,R.I.2000 1231: Revisionofthedungbeetlegenus Temnoplectron Westwood (Coleoptera: Scarabaeidae: Scarabaeini). Memoirs ofthe QueenslandMuseum 46(1): 253-297. Brisbane. ISSN 0079-8835. Temnoplectron Westwood is revised and five new species described, four from North Queensland: cooki, finnigani, lewisense, monteithi, one from New Guinea: wareo. Temnoplectron reyi Paulian is removed from synonymy with T. politulum Macleay, TemnoplectronlaevigatumMatthewsisplacedinsynonymywith T. boucomontiPaulian, T heurrti Paulian and T. howdeni Paulian are synonymised with T atropolitum Gillet, and T. majorPaulianisrecognisedinAustraliaforthefirsttime.Allknownspeciesareredescribed. Akeyisprovidedforthe19speciesofTemnoplectronandnewdistributionrecordsarenoted. Acladisticanalysisofthegenusispresented,theresultsofwhichsuggestatleasttwoorigins for flightlessness in the genus. The biogeography of Temnoplectron is discussed with referencetoisolationofrainforestblocksduringperiodsofmaximumaridity.OColeoptera, Scarabaeidae, Temnoplectron, Australia. New Guinea. CAM. Reid. Co-operative Research Centre for Tropical Rainforest Ecology and Management, JamesCook University, Smithfield4878, Australia(currentaddress: Centre forBiodiversityandConservationResearch, AustralianMuseum. 6 CollegeStreet, Sydney 2000):R.I. Storey, QueenslandDepartmentofPrimaryIndustries, POBox 1054, Mareeba 4880. Australia:'received24 October2000. ' ThescarabaeinefaunaofAustraliaisoneofthe More than 7,500 Australian specimens of best known speciose beetle groups on this Temnoplectron have been examined, collected continent, thanks to taxonomic revisions ofall from all ofthe major rainforest blocks in north the genera by Matthews (1972, 1974, 1976). Queensland.Thisrecentcollectinghasdiscovered These works were partly based on surveys of fournew flightless montane species and several pastoral countrypriortotheintroductionofexotic cryptic-species complexes, which are described species (Bornemissza, 1976) and collections by here. Nineteenspeciesarenowrecognised inthe Matthews himself. The rainforests of north genus, 16 in Australia and 5 in New Guinea. A Queensland were relatively under-explored for key tothe species ispresented. scarabaeines until systematic collecting by Ross Storey, Geoff Monteith (Queensland Museum) The species ofTemnoplectron are confined to andothers,from 976. Somematerial from these the tropics and subtropics, as far south as the collections has b1een described (Storey, 1977, Brisbane area. The feeding and nidal behaviour 1984, 1986, 1991; Matthews& Stebnicka, 1986; of Temnoplectron laevigatum Matthews was Storey & Weir, 1990; Storey & Monteith, 2000; described by Matthews (1974), and the biology Reid, 2000), but many new species remain ofT involucre Matthews has been examined in undescribed, especially in the largergenera. detail bv Asjnes Rortais,James Cook University, TownsvilM1999, unpubl. PhD). To this can be Temnoplectron Westwood is a genus of added published studies of altitudinal range Scarabaeini with 16 described species prior to (Monteith, 1985 perching(Howden,Howden& ), this revision; 10 in Australia (Matthews, 1974) Storev, 1991), seasonality in two species (Hill, and 6 inNewGuinea(Paulian, 1985). Thegenus 1993), edge effects (Hill, 1995), habitat fidelity, is well-defined and probably monophyletic. die! activity and diet (Hill, 1996). These studies althoughitispossiblethatTemnoplectronispara- show that, for the commoner species in the phyletic with respect to Monoplistes Lansberge southern part of the Wet Tropics, collecting and Diorygopyx Matthews (Matthews, 1974). Temnoplectron isbestwithinrainforest,between The lasttwoappearto besister-taxa. Thecurrent December and April, at night, using dung baits. revision is concerned with attaching names to Temnoplectron species are alsoattracted to liver, species,preparatorytoforthcomingphylogenetic mushroom and banana baits (Hill, 1996). Other analysis of the Australian Scarabaeini, which species occur in open woodland or dry forest may result in changes to the generic concepts. (Matthews, 1974). 254 MEMOIRS OF THE QUEENSLAND MUSEUM MATERIALS AND METHODS sclerotised lump adjacent to the flagellum; the ringsclerite,asclerotisedringatorbeyondthetip MORPHOLOGY. The morphology of Tenmo- ofthe flagellum; the median sclerite, a deeply plectron was studied in detail to obtain as many folded irregular shape in the middle of the endophallus.Otherareasofsclerotisationmaybe characters as possible for phylogenetic analysis. Most terms for external characters should be present, but when observed through the inverted obvious or selfexplanatory. The microsculpture wallarepoorlydefinedandgenerallyonlylightly is described as seen under 50x magnification. sclerotised. The flagellum, being almost in- Several ratios are used in the descriptions as variable, is not illustrated, the othersclerites are convenient short-hand descriptors ofattributes, illustrated foralmostall species. but it should be noted that many specimens are Female genitalia were removed by tearingthe asymmetricandthereforetheratioscanvaryfora softened integument along one side of the singlespecimen.Theeyewidthisthewidthofthe abdominal tergum and around themarginsofthe dorsal part ofthe eye at its widest point, which abdominal apex, freeing the genitalia plus gut maybebasalormedial,and iscompared withthe fromtheabdominal walls.Thisunitwassoftened shortest dorsal distancebetween theeyestogive in dilute KOH, then water, and cleaned with the interocular ratio. The hypomeral stria is the removal of most of the gut, tracheae and ridgeandgroovefromthebaseofthehypomeron, glandulartissue.Thespennalhccawasexamined parallel to the lateral margin (Fig. 4); it is in glycerol. The female genitalic system of comparedwiththelengthofthehypomerontothe Temnoplectron is unusual amongst Scarabaeinae posterioredge ofthe femoral excavation, at that inhavingthespermathecal ductopeningdirectly point, to give the hypomeral ratio. The subtle to the external surface ofthe animal, not viathe greenish colour of the elytra is best seen by vagina,whichisaseparateventrallysituatedduct comparison with the pronotum (always black), (Fig. 23). There isoftensecondarysclerotisation understronglighting.Striae8-1 donotreachthe around the entrance of the spermathecal duct, basal edge of the elytra but are always ab- which may form atransverseorquadratebar,the breviated by shortdistances whichare useful for spermathecal sclerite. The characteristic sclerite diagnosing species. These distances are most between the female genitalia and the anus of conveniently measured against the length ofthe Coptodactyla (Reid, in press) is absent, and the mesepimeron, where it touches the epipleuron. hemisternites are small and insignificant short The outer margin ofthe foretibia hasthree large quadrangular struts, internally placed on either major teeth, and an indeterminate number of side ofthe anus. The spermatheca, in common minorteeth (Fig. 7). The length ofthe male hind with many Scarabaeinae, has a transparent tibial spine (Fig. 14) is often diagnostic (in fresh semi-circularwindowatthemiddle,ontheinside specimens) and is best compared with the width margin ofthe point of inflexion (Fig. 24). The ofthe tibia at the base ofthe spine, which gives spermatheca showed little variation and is the tibial spine ratio. therefore not illustrated for all species, but the Male genitalia were preparedby immersion in spermathecal sclerite, ifpresent, is illustrated. KOH dilute for several hours then rinsing in Descriptions are based on freshly emerged water. The endophallus was removed by cutting specimens, if available. Older specimens may the membrane between the parameres and basal havescratcheddorsal surfaces,eroded foretibial piece and separating these. In all species it is a teeth (compare Figs 7G & H), tibial spurs, and simple tube without lateral lobes. The endo- hind tibial spines (compare Figs 15A & B), and phalluswaspulledaparttoexposethesclerites.A the frontoclypeus may beblunted. nomenclature of male endophallic sclerites in Coprini was provided by Genier (1996), which MATERIAL. Abbreviations for repositories are was modified for Coptodactyla Burmeister as follows: Australian Museum, Sydney (AMS); (Reid. 2000) and this system is appropriate for Australian National Insect Collection, Canberra Temnoplectron species. In the latter, there arc 4 (ANIC); Canadian Museum for Nature, Ottawa endophallic sclerites (Fig. 20, flagellum omit- (CMN); Deutsches Entomologische Institut, ted), in a single ejaculatory sac, which form the Berlin(DEB); Hope DepartmentofEntomology, sperm pumpwheneverted: theflagellum, almost Oxford (HDO); James Cook University, uniformly shaped, with broad, trilobed and Townsville and Cairns campuses (JCU); Museo ridged base and single whip-like apex; the basal Civici, Genoa (MCG); Museum Zoologicum sclerite, an irregular folded plate or almost solid Bogoriense, Bogor, Indonesia (MZB); Natural ' I"NOF7Y:uv Oh)DUNGBEETLES HMiusmto:r1y1inM,usPeruamg.uePar(iNsM(FMiN;HNQ)u:eeNnasilfoannadJ t(oMaptatrthietwioSn,in1g97s6p;ecRieetsd,in2t0h0e0)r.otAusnadurmo-uggmhugpu-iwdce Department of Primary Indusiiv, Matceba first relied on male secondary sexual characters (DPIM): Queensland Museum. Brisbane lOMj. on fiesh majormalespecimens,arguingthatthese South Australian Museum. Adelaide (SAM); mporlaiil in mate-recognition and therefore University ol Queensland Insect Collection, spccies-ciisciimmation by the organisms Wfc Brisbane (UQX Western Australian Museum. believe litis is likely because in the rotnndum Perth(WAM) gpecies-grrjuptheprimaly malesexual organ,die Abbreviations lor collcclor's names (with itedeagUS, shows little variation, whereas in the more than two entries) are as follows: CB, C. otherspeciesofTbmnoplecfwHthere is littlem no Burwcll;GB,G, Bometniissza; EB f B Britton; en em oi secondary sexual charactersbut JGB, J.G. Brooks; JDB. J.D. Brown; IC, I.F B. large and constant differences in acdeagal Common; DC, 1)1 Coofc; DIC, I) J. & I. Cook: morphology [foi example in the poiin JPDK.P..FTP.eDrorvaefnj;GEHK.,GAE..DH.oEldlwoawradvs;;HJAII.JL. FIIecShanA; psopiencti,esit-cboemcpalemxe.oFbigv.ioIu'sKitHhait. Fchraormacttheirssstoafrtitnhge Howden; R1I. R. IfuppatZ; HJ, H. Janctzki; EM, surface sculpture and elytra! striae are too F.G. Matthews; DM, U.K. McAlpir.c, SM. S variablein\\\cmiiinjiiinspecies-grouptohi. Misko; (iM, GB Monteith; GSM, GJ3. & S.R. diagnoslically. although these may be valuable Momcith: IViBM, M.S. & B.J, Moulds, S.IP, s. & characters in the other species-groups, On the .1. Peek.i. k.( .A M. Retd; 1R. L Real;DR. D.C.F. other hand, there are excellent male leg char- RSetnotr/e.y;;LRRT,,L.RRWoberTtasy;lo.rIS, G.1.T;SeyGmlourt,hoRmSp,soR.nI;. iaccstuelrtsswhoifcohutarestduidaygnoosftitchefoiwieaucmhlSupmecisepse.ciIchs-e ML). MS Upton: AWU, A. Wal&rd-Huggii grouparesomenewsynonymyandtherecognition AMVVII, A. & M. Walford-Ilugiiins, IW, .1,1 new species liu ilu- Australian fauna. WasselF I VV I A Weir; DY,D.K. Yeates: PZ, F Wehave used the family name Laporrc inplace Zborovvski. Bislelnau lor the author of Temnbple< .. Abbreviations for geographic features: Bell, 'dinni isanhonorifictitle, lot rraoyois Beget) C. Cape; Gk, t reek: I., island; Ml. I cHiis ' ompar de Caumonl Laportc. self-styled Mount/Mountain, NP, National Park Pi, I fotnptf de Castelnau (Miisgrave, l°o2). Ra. Range: R, River; Rd. Road; SI'. State forest; Tbld, Tableland. Teninoplection Wcstwood. 1841 DlyEdFeIsNigInTaIteOdNSsu<c>hFbSyPFaCtIaxIosnoAmisspteicnksiginsoursaunacle- IikYIM-.iySpPyEC(MIaEtSt.heIwesn,iii1n^i7>4k)'.cw>n>;,tiindinnWestWBCd,hv ofthe full range of sanation ol the organism DISTRIBUTION. Australia, from near Brisbane concerned. This morphological study, based on north and west io Cape York and the ktmberley more than 7,800 specimens, has allowed a Ranges, and New Guinea. reasonable understanding of intra- and mter- population variation within speeies. liven the DIAGNOSIS The genii', was comprehensively rarest Australian speeies is represented by SO described by Matthews (1974). Amongst Aus- specimens. tralianCanthonini it is identifiedby thefollowing me ofthe new taxonomic decisions giver attributes: elytron without pscudepiplcuron h are the result of discovery of hitherto outside stria 7i each claw with sharp basal loolh; unknownmorphotypes,oi ofcryptic specieswith mid andhind tibial spursnot fused to tibial apex; distinctive male genitalia. Oiese species are pj gidjum wiiiiout basal groove oi depression fairly obvious. However. Tcnino/>k\1roii has a A few minor additions 01 alterations \^;<\ be .ulailvdifficultspecies-complex which was made to the features listed by Matthews (1974): only partly resolved in the last revision hypomeral stria present (minute in some (Matthews, 1974): the roiimJiun spccics-gioup specimens oi / ,7m Blackburn); elytron In this group, surface sculpture varies, perhaps with stria H present or absent, lore tibia ofmale elinally, so that fresh specimens from range nol internally lobed at apex, with 3 large outer extremesmayhavequitedillereiii punctalionand teeth and many small subsidiary teeth; mid tibia micro.seulpturc. Thisissueis furtherconfusedby with two articulated spurs; hind tibia with piti tmhe high freqiicicv ufold abraded :.po„rncnv as longation [spine) present w absent: a species oi (. - - i spurpresentorabsent; firsthindtarsalsegmentas MEMOIRS OF THE QUEENSLAND MUSEUM 256 long as or shorter than second; parameres Clypealmargincurvedbetweengenalanglesandmedian symmetrical or asymmetric, apices not strongly teeth,convexnearmedianteeth(Fig. 2B)(lessobvious inwornspecimens); $ pygidiumwithstraightorevenly deflexed ventrally: endophallus with four curved basal margin (Fig. 16A); apex of 3 hind tibia selerites; female with entryto spermathecal duct without articulated spur(Fig. 14B)(ifhypomeral ratio externally exposed on ridge (often sclerotised) 0.6. 1-4 minor teeth between major teeth of fore betweenvaginaandanus:spermathecalductlong tibia) .' 5 and tightly coiled. 4(3). Frontoclypeus surface entirely finely punctate, or slightly rugoseatanteriormargins: venterof$ and 2 The male can be distinguished by: apical fore mid femora evenly curved, not lobed or expanded al tibial spurbroader,ovateandbladelike(compare apex(Fig. 11A);parameresmoreelongate,almostsym- Fig.9partsE&F);pygidium longer;lastventrite metrical(Fig.17A-B)(NQ) . . bomermsszaiMatthews medially foreshortened. Secondary sexual mod- Frontoclypeus more strongly punctate and rugose ifications may be presenton themale pronotum, towardstheedges; 6" mid femurwithstrongpreapieal mid and hind femoraand hind tibia. Mostmales pvernctarpailcallolbyeex(Fpiagn.deIdI;E)p,avreanmteerreosfshSortmeird,lfefetmtuhricskleirghatnldy haveextendedhindtibialspinesincontrasttothe lesspointedthanright(Fig. 17F)(NQ) /WereLaporte u1n5sBpi&neCd).feFmoaulressp(eccioemspasrheowFisg.ex1u3alpadritmsoEr&phFisomr 5(3).mOaujtoerrmtaeertghin(Foifg.for7eB-tiDb)i;awiintnher1-m2amrigni.onr. ftoereethtbiebitaweneont in elytral surface sculpture. emarginate; 3 mid femur relatively flat, broadest in apicalhalf,withlargepreapiealventral lobeand 8 mid Thelarvalandpupalmorphologyisundescribed. femur slight!) ventrally expanded in apical half(Fig. KEYTO SPECIWEESSOTFWTOEOMDNOPLECTRON nIaIrBr);ow3edhaitndapfeexm(uFrigb.ro1a1dHe)s;t3inhaipnidcatlibhiaallf,spviennetrraaltliyo 1.5-2,spinegraduallytaperedinprofile(Fig. I4B-D); 3 pronotum not antero-medially depressed (parameres Supplementarycharacterstatesforahalfcoupletaregiven asymmetric, left blunt, right narrower with sharp flat iT.nbcroaockkie,tsT..NIoewtiesetnh.asteboatnhdsTt.atmeosnloeficlloiuip.leTth1e2caoruepplreetssenmtaiyn apicallob,e)(S,C&NQ,NT,WA,.NGb)oucomoniiPauYian appearcumbersomebutappeartowork forthethousands Outermarginofforetibiawith2-4minorteeth between ofspecimenswehaveseen,includingdwarf,defonnedand majorteeth(Fig.7E-FI); innermarginforetibiausually teneralspecimens. distinctlyemarginateinbasalhalf; 3 and 9 midfemora not expanded in apical half, broadest at middle (Fig. I. Lrasotwtowfolavregnetrpiutnecst(u5raensd(a6n)nsuelpaarrpaittesd);bhyedaedeapngdrporoovneowtiutmh (1F1iCg-.DI)I;F,3J)h;ianpdicfaelmuspribnreoa3dehsitndintmibiidadlloenograinndbatshailckh,aolrf impunctate.orextremelyfinely punctured(fronto-clypeal short and flat; 3 pronotum often antero-medially margin not produced between genal angle and median depressed 6 atenedth;trimaancgruolpart,erloeusss;th6anpoasptiecrailorwitdibtihalosfpitinbeias,howritt,hofluatt 6(5).Outerfaceofhindlibiaevenlycontractedtobase(Fig. 8sp-uIr:Omhimn;dNtGa)rsi long, 0.35-0.5 lengthahtirnodpotliibilau;mlQeilnlgetth Sl1le2hnBg)t;ehliyftnrnraeolrmmsbatarrisgaeina(Fboibfgr.feov7riFea.t-tiFeb)di;ahebymyapro0gm.ie5n-ra1at.l5eraXabtoiulote0n0.g.4t3-h-00..7o4;f Suturebetweenlasttwoventritesnotorweaklygrooved, mesepimeron; pronotum and elytra dull, slrongly and without row of punctures: head, and usually also evenlymicroreticulate(exceptpronotaldiscshiningand pronotum,distinctlypunctured 2 notmicroreticulateinsomeCapeYorkPeninsulaspeci- 2(1). Larger, length 8-13mm; basal segment oflabial palp mens); major 3 pronotum deeply antero-medially muchwiderand 1.3-2x longerthan2ndsegment(eyes depressed with lateral tubercles; 3 mid femur without large, interocularratio4-7; lateralmarginsofpronotum swollenexternal face; 3 hind femurbroadest at middle, complete; macropterous; 8th elytral stria present; 3 evenlytaperedtoapex(also V)(Fig. I IJ);apicalspineof posteriortibial spineelongate,aslongasorlongerthan 3 hindtibiamassiveandblunt,almostasthickasbroad tibial width: hind tarsi short, <0.3x length hind tibia; (Fig. 14F-G); parameres strongly asymmetric, left base of metaslernal process without triangularly strongly curved, blunt, right with large fiatapical lobe expanded[margins)(raftmdumspecies-complex). . . 3 (Fig."l7G-H)(NQ&NG) majorPaulian Smaller,length3.5-7.5mm;basalsegmentoflabial palpi Outerfaceofhindtibiaabruptlycontractedatbase(Fig. aswideasand1-1.5x longerthan2ndsegment(margins 12C); inner margin of fore tibia emarginate 0.2-0.25 ofefebflryo,nptrocoldyupceeudsbneostiderumgeodseilayntpeuentch;tu6r"edmiadnfdemnoutr,nootr 7lGen-gItIh):frhoympboamseer(aelmarrgaitniaoti0o.n6-r0a.r9e;l"y8atbhsenetlyitnra9l)(sFtirgi.a modified;A hindtibiawitharticulatedspur) 7 abbreviated by 1-3x length mesepimeron; disc of 3(2). Clypeal margin almost straight between genal angles pronotum shining, not orshallowly microreticulate, in andmedianteeth,notexpanded(Fig.2A);basalmargin contrast to dull elytra; major 3 pronotum shallowly of6 pygidium medially swollen(Fig. 16B):apexof3 depressed without lateral tubercles; tj mid femur hind libia with small articulated spur (Fig. I4A) broadestatmiddle,outerfaceswollen(lesssoin minor (hypomeralratio0.3-0.6; 1-2minorteethbetweenmajor :' I; ' hind femur broadest before middle, with apex teethofforetibia; 3 pronotum notmedially depressed; ventrallylobed(slightlysoin V)(Fig. IIF);apicalspine 8th stria almost reaching base ofelytra, abbreviated at oft? hindlibiashort,equaltoapicaltibialwidth,andflat base by s length ofmesepimeron; 3 mid and hind inprofile(Fig. 1411-1);parameresasymmetric,leftthick femora broadest inapical half; 3 hindtibial spineratio and blunt,rightwithshort flat lobeatapex(Fig. 17I-.I) 1.5-2,spinethickandblunt). 4 (NT,NQ) rotundumWestwood , ' REVISION OF TEMNOPLECTRONDUNG BEETLES 257 7(2).B2asxallqeunagrttherraoefseelpyitrmaervovinl,hu1susallrliyae,re8atchhaibnbgraepviicaatledhablyf girneteenr)spaces equal to puncture diameters; elyt1r4a ofelytra(eyeslarge,interocularratio3.5-4.5;pronolum 13(12). Eyes smaller, interocular ratio 11-12 (Fig. 3P-Q); shining, disc not orshallowly microreticulate, strongly body less elongate, and more convex (Fig. IB), punctured and lateral margin complete; elytra dark lengttuwidth ratio 1.25-1.45; remnant of 8th stria bronze-green;macropterousoralmostso,wingsextend shorter,presentasshortgrooveinelytralsecondquarter beyondabdominal apex; c? hindtibial spineratio0.75- orreduced to punctures(Fig. 411); elytra black (length 1.25;length4-6.5mm) 8 5.5-7.5mm) cookisp.nov. Basal quarterofelytrawithatmost9striae.8thabbrev- Eyes larger, interocular ratio 8-9 (Fig. 3F); body more iated by >3x lengthmesepimeron, usuallyconfinedto elongate, less convex (Fig. 1C), length:width ratio second quarter ofelytra (T diverxicolle. T.fmmgemi, 1.40-1.65; 8th stria longer, at least throughout second some T. cooki),orreducedtoa fewelongatepunctures, quarter (Fig. 41^); elytra dark bronze-green (length orabsent 9 6-6.5mm) diversicollcBlackburn 8(7).Rightparamerewithoutpreapicaldorsalnotchinlateral I4(12).Stria9deeply impressed, likestria 10,delineatedby- view ("Fig. 19B) (length 5-6.5mm; head strongly but fineridge,withatmost3-5foveolatepuncturesinapical sparsely punctured,dullandstronglymicroreticulate: i fifth; lore tibia with shorter, broader major teeth, elytraentirelydullandmicroreticulate, 9 withintervals separated by 3-5 convex or sharp minor teeth (Fig. 5-10 or 6-10 shining, not obviously niicrosculptured lOF-FIl;baseofupperepipleuralmarginnotdepressed; exceptatextremebase;striae 1-7withobviousfoveolate apices ofboth parameres concave, laterally depressed psuhnacrtpuremsinionrapitceaelthhablfetowfeeelyntram;ajfoorre ttiebieathw)it(hN3Q-)7 (tFoicgu.rv18eEd)t;ipap(Fiicga.l1s0p1ur1)of9 foretibiateevweinsleynsaettsepn.unaotve.d aeneopiceimiMatthews Stria 9 shallow, much shallower than stria 10 and not Right paramere with preapical dorsal notch (Fig. I9C) delineatedbyaridge,withscatteredfoveolatepunctures (length4-5.5mm(5-5.5mmonCarbineTbld);punctures throughout;foretibiawithlonger,narrowermajorteeth, and microsctilpture ofhead usually uneven in density separated by 2-3 feebly convexorfiatminorteeth(Fig. andsize,withpart(atleastpatchanteriortoeye)orallof 10B-E);baseofupperepipleuralmargindepressed;apex headshining;basalthirdtohalfof6 elytrashiningand ofleftparamerebluntly rounded,apexofrightparamere without microsculpture, apex microreticulate, 2 with triangularly produced (Fig. 18C-D); apical spur of 9 elytraentirelyshiningoronlymicroreticulateonapical l'oretibiaangulateoninnermargin(Fig. 10C-E) .... halfofintervals 1-4;striae 1-7not.indistinctly,orrarely monteithisp.nov. mdiisntoirnc,tluys,uaplulnyctbaltuent,inteaeptihcabletthwierde;nfmoarjeotribtieaetwhi)th(N2Q-)5 15(11ey).eSstrilaar1gesrt,ronignltyerdoeceupleanredraitnioapi3c-a4l.5qua(rFtiegr.of3eAl-yBtr)a;; subvolitansMatthews frontoclypeus slightly concavebesidemedian teeth; 6 9(7). Basal border of pygidium with middle strongly hindtibiaevenlycurved,tibialspineratio0.6-0.75(Fig. produced (Pig. 16D) (elytra dark bronze-green; lateral I3A-B); parameres symmetrical, with apico-venlral pronotal border complete; brachypterous, wing scale tooth(Fig. I9I-JHNG) 16 halfelytra!length;8thstriapresentasimpressed lineon Stria I not deepened at elytral apex; eyes smaller, secondquarterofelytra; interocularratio7-8.5; S hind interocular ratio 5.2-6.5 (Fig. 3C); frontoclypeus not tibial spine ratio 0.65-0.75, spine blunt; parameres concavebesidemedianteeth; 6 hindtibiaslightlybent symmetrical, each with preapical fringe oflong hairs) 0.25 from apex,tibial spineratio0.75-1.25 (Fig. 13C); (NQ) prmiganisp.nov. parameresnot symmetrical, without ventral tooth (Fig. Basal border ofpygidium not medially produced, but 19E) (middle of pronotal disc finely and sparsely straight orevenly curved (fig. 16C);without theabove punctured;apexofelytranotproduced;NQ) combinationofcharacters 10 disruptionMatthews 10(9). Lateral margin of pronotum partially or entirely 16(15).Elytrainprofileevenlycurvedtoapexofsuture(Fig. pefrfoacceesdst(rFiiagnsg4uAla-rBl,yEe,xHpa)n(adnetde)riorcornersofmetastern1a1l 4anAt)e:riofrrotonteoycelsy;peeuysessstmraolnlgelry, mpounrcetuerveednl,yinnacrlruodwiendg L(atNeOr)al margin ofpronotum complete(Fig.4C-D, F-G17) danitsecrmioorrley,stirnotnegrloycualnadrdreatnisoel3y.8p-u4n.c5tu(Friegd. 3A); pronotal utTO'd/iOTiLansberge 11(10).Lateralmarginofpronotummissingmedially,atleast Elytra in profile produced and slightlyraisedatapex of presentinapicalandbasalfifths,usually onlyeffacedin suture (Fig. 4B) (intervals 1-3 with subapical middlequarter(Fig.4E, H);brachypterousorwingless depression); frontoclypeus finely and sparsely punc- ((Fmiegs.o5-Dm)e;teaysetsersmnaalll,miantregriocnulawritrahtiosm8a-l1l2(tFriig.an3gPu-lQa)r tured, anterior to eyes impunclate or almost so; eyes excision;NQ) 12 (lFarigge.r,3aBb)r;uptplryoncoutravleddainstcerimoorlrye, ifnitneerloyculaanrdrastipoar3s-e3l.y5 Lateral margin completely effaced, or only present in punctured wttreosp.nov, corners (Fig. 4A-B); macropterous; eyes larger, inter- 17(10). Eyes larger, interocular ratio 3.5-4.5 (Fig. 3.1-K); ocularratio3-6,5(Fig.3A-B)(length4.5-5.5rom). . 15 macropterous;hindbodyelongate,sidesevenlytapering 12(11).Sizelarger,length5.5-7.5mm; 6 hindtibiawithlong from prominent elytral humeri to almosttruncate apex apicalspine,equaltowidthoftibia(Fig. 15L-G)(linear (Fig. IF) (length 6-7.5mm; black; 9 outer elytral segment of8th stria usually partially present in second intervals shining, contrasting with dull microreticulate quarterofelytra) 13 inner intervals; 6 hind tibial spine sharp, tibial spine Sizesmaller, length 3.5-4.5mm; A hindtibia with short ratio0.85-1.25) 18 apical spine, less than halfapical width oftibia (Fig. Eyessmaller,interocularratio7-12;hindwingsvestigial; 13N-0) (8th stria absent or reduced to 2-3 punctures; hindbodybroadandrounded,withoutprominentelytral pronotum strongly punctured, sides ofdisc with some humeri 19 258 MEMOIRS OF THE QUEENSLAND MUSEUM 18(17).Leftparamereevenlyattenuatedtotriangularlip(Fig. TYPE. Notseen (in MCG). The species is recognisable 19G); basalthirdofstriae5-6very lightly impressedto from the redescription of the type material given by almostinvisible, lessimpressedthanonsecondthirdof Paulian. elytra,vvholeofstria7similar. . . . politulumMacleay Leftparameresharplyproducedinprofile,withflattened MATERIAL.(5)PAPUANEWGUINEA:5.34miEPort tip (Fig- 19H); basal two-thirds of striae 5-6 evenly Moresby,KokodaTrail,2200',dunstraps, 14-18.vii.1974, impressed, stria 7 similar or more stronglyw/impressed S. Peck(CMN,DPIM) Paulian 19(17). Length 3.5-5mm: elytra distinctly bronzed or DESCRIPTION (male). Colour. Body black, greenish; meso-metasternal border with small median appendagesreddish-brown. transversetubercle; 8 hindtibiawithshortapicalspine, Length: 4.5-5.5mm. tibial spine ratio <0.5; 9 elytra shining and evenly shallowly microsculptured; one or both parameres Head(Fig.3A). Stronglybutsparselypunctured, concave,roundedorpointedinprofile 20 anterior,genaeandmiddleoffrontoclypeusmore Length 5.5-7.5mm; dorsum pure black; meso- finely punctured and microreticulate than base; metasternal border with flat triangular excision (Fig. eyeslarge,evenlynarrowedanteriorly,interocular 61B-)1;.25S(hFiingd.ti1b5iFa-CwTi)t;hlVonogutapeircaellystprailne,inttiebrivalalsspisnheinriantgi,o ratio 3.8-4.5; anterior margin frontoclypeus shallowly microsculptured, contrasting with dull inner strongly upraised and curved anteriorly from intervals;bothparamereswithbluntortruncateapicesin rounded genal angles, but slightly excavate profile (Fig. 19D-E) (disc metaslernum strongly beforenarrowapicalteeth;firstsegmentoflabial spturnacptluikree)d, mesosternum almost impunccloaolkei;spw.innogv.s palpi 1.2-1.5x length second segment. 20(19), Stria 9 similar to stria 10, deeply impressed, Thorax (Figs 4A. 8A, 13A). Pronotum: shining, delineated by a fine ridge, with at most 3-5 foveolate not microreticulate except extreme lateral punctures inapical fifth; 2-5 sharpminorteethpresent margins; pronotal disc strongly and closely betweenmajorteethofanteriorborderofforetibia;base punctured, not medially depressed; lateral ofupperepipleuralmarginnotdepressed;apicalspurof V foretibiaevenlyattenuatedtocurvedtip 21 margin pronotum entirely effaced, or almost so, Stria 9 shallow, much shallower than stria 10 and not at least 75% absent; hypomeral stria weakly delineatedbyaridge,withscatteredfoveolatepunctures expressed, hypomeral ratio 0.25-0.3; elytra throughout;foretibiawithlonger,narrowermajorteeth, entirely microreticulate, but shallowly and separatedby2-3feeblyconvexminorteeth(Fig. IOB-F); irregularly in basal third; elytra strongly arched obVafslfeeofrotefptuaiprbipaaemraenregepuilbpalltueneutrolanylrminoanurengrdiemndad,regapiprneesx(sFoeifgd.;riaIgpOhitCc-aplEa)rs(apamupreero\ef ianpipcraolfihlael,f,hwiightehs0t-a2tpmuindcdtluer;ess;trsitari1a8dereepdeunceeddtino triangularly produced; both parameres with short a few elongate punctures in second elytral subapicalrowofsetae) monteitlusp.nov. quarter, stria 9 abbreviated by 1.5-2x length 21(20). Elytra dark bronze-green, only slightly contrasting mesepimeron;baseofepipleuronnotconstricted; with pronotum, basal thirdofintervals 1-3 shiningand macropterous; meso-metasternal border slightly cuosnuvaelrlgyenwtitwhitohutsidmeicmrarogsicnu;lpitnutrereo;culhayrproamtieor7a-l8,setyreisa raised, with narrow transverse median tubercle; broadestatbaseofdorsalportion(Fig.3L);& hindtibial median lobe of metasternum shining, without spur 1.5x length firsttarsal segment(Fig. 13M);head, microsculptureexceptatextremeapex,finelybut pronotum and metastemal median lobe usually finely closely punctured, with margins triangularly arnodundspeadrsaelpyex,punrcigthutredp;aralemfetrpearwaimtehreansgiunluaattee,bwaistahl expanded in apical comers; outer margin fore projection, neither with ventral setae (Fig. 18B) tibia with 2-4 convex minorteeth betweenthree involucreMatthews acutemajorteeth, innermarginslightlyconcave; Elytra brassy-green, contrasting strongly with black mid and hind femora elongate-ovate; hind tibia pronotum, strongly microreticulate throughout; with short apical spine, tibial spine ratio 0.75, hypomeral stria almost parallel to side margin; with prominent articulated spur; hind tarsi long, dionrtsearlocpuolratriornat(iFoig8.-310M,);eySeshinbdrotaidbieasltspnuerarasmliodndglaesoorf 0.4X length hind tibia, segment 1 ventrally shorter than first tarsal segment (Fig. 13N); head, lobed,2,3and4elongaterectangular,decreasing pronotumandmetastemalmedianlobeusuallystrongly in length, segment 5 almost equal length and closely punctured; apices of both parameres segments 3+4. concave, laterally depressed, with short \entral row of smallsetae(Fig.l8E). . lewisensesp.nov. Abdomen (Figs 19J, 21E). Suture between last two ventrites simple, not deeply grooved and punctured; basal margin of pygidium evenly Temnoplectron aeneolum Lansberge curved, without prominence in middle; (Figs3A, 4A, 8A, 13A, 19J, 21E, 23L, 24D. 25) parameres almost symmetrical, short and broad Temnoplectron aeneolum Lansberge, 1885: 375; Paulian. with triangular tooth on venter ofapices, which 1934: 285; Paulian. 19*5: 224. are reflexed and overlapping, without fringe of REV(Sil >N ( >F TEWOPLECTBON DUNG RUM S TYPi;. Ilolorspc: Paluma Dam Rd, Ml Spec, 30.iii.IWiS, FA (ANIC). Lxamined. I MATERIAL. (479)NORTH QUEENSLAND: Kfthdaj Ck Falls (AN1C. BMNHl; Bluewaler Ka(()\1/: Boulder Ck, lulls (MM): KJot '!V. Caidwcll(ANIC);ChamuUJu W Ck (QM); 40km Ingham (ANIC); 22km Nl (sic, MW'I Kenned) (ANIC); k.irrama Ra. (JCU, QM): klcllbcrgRdtumofl(IJM):I..aminsHill(QM);Mal;ianRd 2km otrilVav (QM); Malaan SF (QM); MlUaa Mfflaa I alls(QMi. Mifather("lanes (QM);MtFislieriOM);Ml Graham,( ardwell Ka. (QM); Mt Halifax(K'lJ. QM); Mi Hugh Nelson (QM); 7km S Mi ftajroomool (QM); Ml MaCallSter(QM MtSpec(ANIC.JCU);2miWMlSpec I; (l)IMM);ImmersionNP(QM);Paluma ANFC,.ll to: Imi WNWI IHirk.imliiWiiMiPpaarlaurmvpae:(.\AfN\IICCi);;6k12m1m) E PPaalluummaa((AANNIICC));; Paluma Dam Rd (including 27 paratypes; ami.. QM); Ravenrfioe SF(ANIC); 11km& lSkm SSw Raveush* lulls Falls SF iDPIM): 5km NE Ravenshoc (DIMM), iskm ssw Ravcnslme(DPIM); SmokoCk r.ic'fi>:Fully i nils (AMS. QM); Fulls R Dam (QM); Tulh R Ciii (QM): l.f'kin N lulls R Xing (QM); Upper Buuld PI:'G'!'.,i'iii'/,i)iniiil'oaput\tiivin,^ <I w ipi ipc.eaavb.;oCd,)iiivoeurt.liiiin:e(>ll<i (WQoMn)g:abWealll6anktmanSFa^lJlhserRtdon((QDMP)1;M)W;inWdospgaPobseil,(A?TknIl S Bnlofvl.c;kl!u.ir;n;i;IPFaualtirannp.aiNiotliiIm"Csiiltet; I , (eivwmse sp MluatOll I \NIC);YueeahincCk (QM) DESCRIPTION (malei. Cnlow: Black, clylta seiae; endophallus with roughly C-shaped basal dark greenish appendages darkreddish-brossn. sclerite and appendage, short and uanoss ring Length. 5.0-6.5mm. msceldeiraitnescwliertihtoe.ut appendage, strongly lobed rHeelaatdiv(e1lyig.sp3aDr)s.elSytiopnugnlcytuarnedd,alsmtorsotngelvyeiimlicrIot-il Female (Figs 23L, 24D). Spermathecal sclerite reticulate; Irontoelypeus not rugoscly punctured divided into two feebly selerotised smooth towards anterior margin, which is evenly ridges, separated by base ofspermatheeal duct; shallowly curved between genal angles mil spermatheca falcate, gradually narrowed to median teeth; eyes large, broadest at middle. pointed apex. interoeular ratio 4.5-5; first segment of labial palpi 1225 x length ofsecond. REMARKS. Contrary to Paulian's observations Thorax (Figs 5A. SG. 13D-EF Pronotum: ( 1985: 224), this is a typical member ofTemno- strongly and closely punctured, disc shining. pleciron,sharingmanycharacterstateswithseveral shallowly or not microreticulate, sides strongly Australian congeners and T, Wareo SJV, nov. mieroreticulaie, disc not anteriorly depressed, DISTRIBUTION AND BIOLOGY (Fig. 25). lateral margins pronotum complete; hypomeral ratio 0.2-0.4: elytra entirely microreticulatc. Tenmoplt'ttranueiienhimwasdescribed from Kly usually strongly and evenly so, rarely shallow*t RbiyveDr'A(lLbaenrstbiesrgien1e8i8t5he),rfrNoomvemmatbeerri-alDecoclelmecbteerd in basal half; intervals moderately strongly 1875 or May-July 1876, within 580km of the punctured; apical half stria 1 punctate hui lujt depressed; striae 1-7 with sparse fcneolatc amomuotshaiocftohfeFslwyaRmipvefro(reGsotodaned19s7a7\a).nnIahhiswaroeoadi-s pbausnacltutrheirsdoonfaepliyctarla whiatlhf r1a0resltryiateo.b8alsheeolflacelevdtima land. Italsooccurs inasimilarrangeofhabitatat apical half, rarely apical 0.7; stria 8 abbreviated 3-700maltitudenearPortMorseby<Paulian 1985). by 1-2 X mesepnneronlength;baseofepiplcuron notconstricted:macroptcrous;meso-metasiernal Temnoplectron aeneopiceum Matthews margin with almost Hat triangularmediantubercle: (Fins 50; 5A. XG-H. 13D-K, I9B. 21C, 23C, rnetasternum strongly punctured throughout. 24F, 26) shining except anterior of median lobe micro- TwrWnennN.tt1i9e9m3mtiuntmuuUxum Matthews. ic)74 15?; Cassis& rtertiiacnuglualtaer,lyanetexrpiaonrdceodrnmearrsgionfs;lobfoeressittihbianarrow 260 MEMOIRS OF THE QUEENSLAND MUSEUM margin with three acute major teeth separated by 3-7 sharp or convex minor teeth (varies within individuals), inner margin almost straight; mid femur elongate- ovate; hind tibia evenly curved. outer face abruptly or gradually contracted towards base, almost parallel-sided forapicalhalf; hind tibial spine sharply pointed, tibial spine ratio 0.75-1.5. with apical spur as long as first tarsal segment; hind tarsi long, c.0.3 length hind tibia, segment I ventrally lobed, 2, 3 and 4 elongate rectangular, decreasing in length, segment 5 almost equal length segments 3+4. Abdomen (Figs OB, 21C). I Ventrites 1-5 with basal row of small sensory pits: last two ven- trites with shallowly impressed impunetate boundary; basal margin of pygiclium evenly curved or rarely with median swelling; parameres wilhout setal fringe, roughly sinusoidal with deep ventral c\cavation towards base, but asymmetric, left with obliquely truncate apex, apex dorsally minutely toothed and ventrally produced; right paramere not dorsally notched, gradually constricted to blunt FIG.2. Temnoplectrunspecies,ratundumspeciesgroup,dorsal view of inwardly folded apex; endo- head; A, bornem/sssai Matthews: B. boueomonri Fatilian (holutype phallus:basalscleritepear-shaped lueVigalumMatthews):C,baucomonti(wornspecimen,Dimbulah);D, withangularlateral lobeandsmall boucomonii (PNG): E. baucomoYrtl (Cairns): F, lucre (Laporte) adjacent sclerite: flagellum long, (Windsor Tbld); G. taeve (Paluina): II, major Paulian; I, rotundum lobesnotequidistant; ringsclerite Westwood. All tosamescale. withthick-vvailed narrow ringand blauttersapllliyt bfylamreeddilaonbe:clemfetdian selcrile triangular REMARKS. Adetailed description isgiven here because this species was not clearly distin- Female (Figs 811, I3F, 23C. 24F). As above, guished from T. siibmliians when originally except: inner halfofelytra (intervals 1-4. or 51 described (Matthews, 1c>74). Two paratypes of microreticulate and dull (intervals 1-4, or -5), 7^ aeneopiceurrt (from Mount Lewis) belong to contrasting with shining, non-microreticulate 7^ sitbvolitans. Outer half(intervals 5- or 6-10); fore tibial spur DISTRIBUTION AND BIOLOGY (Fig 26), slightlyflattened,attenuatedtocurvedapex;hind Present all year, but most active or abundant tibia with short apical lobe, less than halfapical during the wet season, from December to April ttirbainalsvweirdsteh;wegaenkiltyalias:clesrpoetrimsaetdh,ecbarloasdclerriitdege,a o(Hnilvl,eg1e9t9a3ti).oTnhiatssnpiegchitesancdomimsoanttlryacpteerdchtoeslligohwt nnnslucent around orifice; spermatheca small (Hovvden etaL, 1991 ), Widespread andcommon andC-shaped,withslightlyswollenbaseandthin in rainforest from Bluewater Range north to (he curved apex. southern end oftheAtherlonTableland, where it REVISION OF TEMNOPLECTRONDUNG BEETLES 261 FIG. 3. Tenmoplectronspecies,dorsal view ofhead; A,aeneolum Lansberge; B,wareo sp. nov.; C, disruptum Matthews; D, aeneopiceum Matthews; E, subvolitans Matthews; F, diversicolle Blackburn; O, atropolitum Gillet;H,henrniPaulian;I,howdeniPaulian;i,politulumMacleay;K,reyiPaulian;L,involucreMatthews;M, lewisensesp.nov.;N,monteithisp. nov.; O.fvmigmiisp. nov.; P,cookisp.nov. (MtSpurgeon); Q,cooki(Mt Haig). All tosamescale. 262 MEMOIRS OF THE QUEENSLAND MUSEUM FIG.4.Temnoplectronspecies,lateralviewofthorax;A,aeneolumLansberge;B,wareosp.nov.;C,boucomonti Pauiian;D,laeve(Laporte);E,diversicolleBlackburn;F,atropolitumGillet;G,finniganisp.nov.; H,cookisp. nov.; epipleuron stippled.Notto scale. overlaps slightly with its sister-species, T. DESCRIPTION (male). Colour. Black, subvolitans. appendagesreddish-brown. Length. 8-10mm. Body relatively elongate (Fig. Temnoplectron atropolitum Gillet ID). (Figs ID, 3G-I, 4F, 6A, 9A-D, 12D, 13I-K, 17K-M, 20A, 23B, 24C,25) Head (Fig. 3G-I). Lemon-shaped, anterior margin offrontoclypeus evenly curved between TemnoplectronatropolitumGillet, 1927: 252; Pauiian, 1934: genalanglesandmedianteeth,exceptslightnick 285;Pauiian, 1985:224. at base ofclypeus; frontoclypeus impunctate or TemnoplectronheurniPauiian, 1985:225;syn.nov. apparently so (sometimes minute punctures TemnoplectronhowdeniPauiian, 1985: 227;syn.nov. visible at x 50), densely and evenly finely TYPES.LectotypeofT.atropolitum(inDEB),designated microreticulate, shining but duller than pro- by Pauiian (1985) from Dormanpadbivak, not seen, but notum; eyes large, interocularratio 3.7-5.3; first contemporarytopotypicmaterialexamined.HolotypeofT. maxillary palp segment 1.2-1.5 x length of heurninotexamined(inDEB).HolotypeofT.howdeninot second segment. seen (in CMN), but part ofthe same series (topotypic) Thorax (Figs 4F, 6A, 9A, 9C-D, 12D, 13I-K). examined. Pronotumshining,shallowlyorobscurelymicro- MATERIAL. (336, onlythose examined in detail listed) reticulate, minutely and sparsely punctured; INDONESIA:4, Dormanpadbivak, 1410m,x.1920,W.C. pronotal disc anteriorly slightly depressed or vanHeurn(ANIC,MZB); 1,FreeportConcession,Timika, evenlyconvex;lateralmarginofpronotumentire 4.76145°S 136.86369°W[sic], inner mature mangrove, to completely obliterated from base tojunction dungpitfall, 15m, 16.iii.1997,Ubaidillah(MZB); 1,ditto, withfemoralhollow,oftenwithdorsaltriangular except 4°39'43"S 136°53'50"E, peat swamp, thickenedareaatextremelateraledge(abradedin 13-16.iii.1997 (MZB); 4, ditto, except 4°17'23"S oldspecimens?);hypomeralratio0.25-0.4;elytra 138°59'98"W[sic], open heath forest, 600m, shining, shallowly or obscurely microreticulate, pSIurlyi-am1n4a.trioyi,if.oA1r.e9s9tS7,a(piMamZnB(t)rMa;Zp,B1),1;3J0a20y,amdw,iitj6t-ao2y6ea.x,Xc.Keep1l9ti9l51a,,5E0W.0uCrmhiog(leMilZke,Bb)Au;.r ainbtseernvtalsorfifneeelbylypuinmcptrueresds;edeloyntraellysttrrailaedi1s-c6, PAPUA NEW GUINEA: 3, Western Highlands, Mt shallowly impressed in apical third or with Hagen,oakforestdungtraps,6000'[1800m],5-S.vii.1974, scattered deep foveolate punctures, striae 5-6 S. Peck(CMN, DPIM). sometimes almost entirely punctate; stria 7

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