Perameles sobbei sp. nov.(Marsupialia, Peramelidae), a Pleistocene bandicoot from the Darling Downs, south-eastern Queensland more

rW.-IMM.KS SlWtinSI' MA iVIARSUPIAI.LA, PI RAMI I.IDAli. API.EISTIM l-.NI BANDICOOT FROM THE DARLINCJ DOWNS, SOUTH-EASTERN QUEENSLAND GILBERT J. PRICE Price uj. 2002 531: PmuNle* .rrtfirVtsp. wv- (MBfupMit, Pcnmcttdwli a PMhoown handiL-uui iKini tlx: Darling Downs, sautn-caHefn OuL'cnsland. \fi-muir* <if lilt- QutvndauJ 4H. I l"; I"" linear,.-. ISS\ 'lir-l SS.»> i'tni/atrlcx .•coiibri sft. ik>v. is described from Pleistocene fluviatik sniimcnls from King Crwk cmtheeAsliirn rXirhnu.Dnwus. PiTumelmxtiblm lulls w illlintlxv ii/Ciariefftf modem Ptrameia -specks, bui ils niokir nmrphnlosy inttie»les M elnva- affinity »ilh Ihu Laily I'Ikktl-iil- spttifv, I', htnwniis. Both species tetani lhc plesioninrphic slates of possessing slTiigh) erislKl obliqfliax with irlnv-ly aprmminatcil tngtiriid eusjmlv aild ihe syriupuMorpliic stale of the reduction of ihe hvpocoiHiiid on M. However, the ciwi sire diftVrWM and slighi molar miimhnliig)' diilcrvncvs between iIk iwo ate considered sufficient to warrant spec ific separation. Ptttnnhts ft>t>fvi sp nov, is tbc thmi IokmI /Vjuh.v/, * specie* described and the first fnim Pleistocene deposits. ^1 Bawihnvl. Ptnmelvi. PIi-m.u • >u . rtarlinglimtm. Kh\f> Ovc* (iHhi-ii -J- Prirc, AlMMnMd Vnnrrxily \>( lieluohgy, Sihtntf of S'tlUtt\lt RtUNiW? Si i'cNct'.t. GPO flflt JJ'.S-*'. nrtibiini: Quetnxtaatl 4t)t)l. Ailttralla femuii: fiiyy/i hp ■■hnk nl.cfnr.ithi.iru: feit'fiYti ?i ,lfi)i\-!l 2fH>?. Pleistocene fossils have been known from the eastern Darling Downs for over M) years, with •in- In si fossilsColleLied lv. s.r I'ln. ■ riL.i - \l hy ]S42 (Owen. 1877}. Recent fauna I lists, such as that of MoItkit & Kure (I W). indicate a high degree of laxonnmic diversity and wide distribution of Pleistocene vertebrates within I he Darling Downs, Such lists an; domiiuicd h> iik'm;ii';iiim;:I species, notably mammals, in part- ietikr member s (it- the Diprolodonlidae. Vom- b«iidi*e. Thylacolejomdae and Macro- pndidae, plus laiye reptiles fnun the Croeodylidac and Varuiiidac. Molnardt rstir/ (I W) suggested thai past collecting mi the Daring Downs has been biased towards the recovery of larger specimens, thereby leading to an underestimation of the smaller vertebrate component. Mohai & Kur/'s I IW7) view is supported by recent systematic cvlhxting. from un eslensive Hnvialiledeposit kieatedSkin Wof niftonalong llK hanks of King Creek I Queensland Museum site L7'>6>. Alonji with nrprvsentulives. of the ali>reineiiiioni_d Uailinu. Downs meLiat'aunal famil ies, other vertebrate species recovered from this site include members of the Tachyglossida?, OrriiiluiThynchidae. Dasyuridac. Peramclidac and Muridac, plus Agamidac. Scincidue ami tluptdue Among ihe pcramelids is a previously undesct itied species otPeramdtt. To date, only two tonsil species of Peramei*3 have been described, tmlh frculi PIilkciic deposits: P. attinghamensis from die Bluff Downs Local Fauna (Archer. 1976i, and /'. beiwnsix \xv>m the Bow, Bi{t Sink (Muirhcad el aL, IW7) and Chinchilla Local Faunas(Macknesset ill,. 2000), Perumele\ stthtwi sp. nov.. is the lirst extinct species ot the genus desenbetl from Pletstoeene deposits. I }.•"!.i. i-. ■■■<•. ■ . I.iln:.' fiillnvH-, I i.lIilV. < '.•!'••[ where ihe adult unreduced check tooth formu ll of :i.i; ■ -i.p.aK -■ V '' ..lid M --I ll >.i|h .Ippct .Mul lower dentil ions Tooth morphology nomenclature tblloivvs Precdman 4lW7p. Stmt distingiiishinu features of Peiiiawln J'ollow those described by Smith (1^72) Higher systcmaties follow Apltn & Archer < 19H7). Types uedeposited in iheQtieeiisland Museum I.QMF). SYSTEMATIC PALAEONTOLOGY Suptu iihort MARSUPIAUACavwr, ISI7 CoHart AUSTRALIDELPKlASzalay. 1982 Order PERAMELEMORPKIA Kirseh. \Wti Family Pl;.RA\1 ELIDAE Cray, l»25 Ptriiindrs GeoiTn.iy. 1S04 TYPI;. SPirituS. /VjwJiriCTJiiiMrtatJuoltluj, 1WM. OTHER SPECIES. Ptnimeia rV^iXwt'rV Quo)' and (iarirtml, IK24.1' tfmn/i i iray, miS,/?er™iVwurS|M>.vr. 1897; P, otfowtKimcnw Arclwr. \17&. P bmnwiM MuirliL-ad, l)a»>«Tl & Arcner. I»7. IM MEMOIRS OF THE QUEENSLAND MUSEUM Fl(i I ./V™.nr1^ji.wiAhi'i'sp.Tniv.,liLil<rt>i>.-Qyi'-JJHTJ(,riglil dtnlaxy: A, Li. buccal hcwlC D. lingual s lew: E F, occlusal view'. Scale tare; 5rmn, Peraaiele* snhheL sp. mjv, {Figs 1-1) ETYMOLOGY. For Ian H. Sobbe of Clifton for hi; ra-iliikiluuiMi.' na>a>iviii: lauri.ii<'l'irK.f;jriiilg Downs. MATFJIIAL. ElOUlTYPt:: yHHJtiTJUKiu ]J,pwtion of the horizontal rarnas of a riajhl d? nt»v «-iih P-M, and M, rwftATVPES: QMF43S79 (Fig. 2A. B), isolated right M-; QMFilHKHFin. 2C.D1 imlalL-d Idl Mj. All typ« an) from QML.7%. KingC reek. E Darling Downs. DLAtiNOSIS. Hypoconulid reduced to absent do Mi.j; Mi long, Mj narrow, Mj tiduisid, markedly reduce*! in comparison to similar-siied species: uigonid cuspids equidistant: anterior cinguLLd cm M; and Mj. pttunintfli, rouudfd in Ms anteroltngual corner. DESCRIPTION. Dummy. Deepest below MJ: mandibular symphysis emending posteriorly 10 below Pi: mental foramen is anterovemtid lo anterior alveolus, of the Pi; P: in missing, though length of its alveoli sugge^is prvrri"lu>^, iai^r^iasL- in size from P, to P,. Mj is [he Khortesl molar tnoill, followed in sending order by Mn, M :md M ■ ifj'jf^f P.-. Laterally compressed: sub-rec1angular in occlusal outline, wi1h slight lateral constriction one-lhird IV™ anterior margin; anlerior, central and fKMlcrior cuvpids oecupy midline of crown; central cuspid tallest and posterior cuspid shortest: cenlral cuspid base in midline of lootk (hough ape* is curved lingually; wear Ibcet present on posterior ape* of teni-rsl cuspid: abrasion from upper premolars has obscured any unique structures nil posterior cuspid. Right Pi. Sub-rectsnguLjj in occlusal outline: ulterior, central and posterior cuspids on crown: central cuspid tallest and ameriOT CUSpidshOfiesi: anterior cuspid in lingual corner of tooth, its buccal surface sloping more genlly towards crown base than does lite lingual surface; central cuspid slightly lingual to midline, not curved linguallj like corresponding Cuspid of Pi; small blade-like crest running posteriorly alonu, midline from apex, of centra) cuspid 1o posterior extreme of posterior cuspid; buri/outa! shcll'-likc structure an Lingual side of crest of posterior cuspLd extending anteriorly to base of main cuspid; small liuriiontal shelf, positioned posterior cuspid pnsteroling'uaL shallow basin on clone to banc of main cuspid; slopes steeply to base of crown on buccal side of crest; faint ridge runs antero-poslcriorly along longitudinal extent of the buccal sloped surface Of posterior cuspid. Rigftt M>, Anlerior One-third Iriangular, remainder rectangular in occlusal outline; taJonid ntfir"k&JLy wider lltfm uigonid; rc*lac<joid lalftt than proloconid. both taller than paiaconid; relative heights of hypoeonid and entneoniel indelerminuble due to wear: paraconid forms anterior margin of tooth, positioned slightly I■ i:lI lo ii.ill in-. : ..n il [iM.tii in:;u.i In paraconid:protoconid occupies- buccal portion of trigonid. slighLly antcrobuceal to melaconid; e-nlocOnid lies directly posterior to melaconid; hypoeonid posteroboccal to proloconid; t:eiKm>iiid and hypoeonid lie on ume transverse plane; hypoconulid reduced to slight swelling at posterior base oL'cnloconid; faint posthypoeristld Connects hypoconulid to hypoeonid; form of cristid obliqua has been obliterated by wear; no anterior or posterior ciugulid present. Right Mj. (Based on QMF4W7% Fig. 2/\. B); Anterior one-third triangular, remainder illuh^iiLu in occlusal outline; lalnnid markedly wider than trigcinrd: proloconid is tallest cuspid on crown, followed in descending order by niL-LiLiiiiid, eiitoeonid, hypoeonid, and pantconid: panuconid LS just posterior to anterior margin, positioned slightly lingual to midline; melaconid is postero lingua I to paTaconid: melaconid and proloconid arc in same transverse plane; hypoeonid is diredly posierim to protoeomd: entoconid is postcrolmgual to DARLING DOWNS PLEISTOCENE BANDICOOT 193 mctmjoimJ: cntoconid ii lin^ua.1 slightly anterior to hypoconid; no hypoconulid is present: ii taint pnsthypocriMW runs po*lerolinBually to posterior base of cntoconid: cristid ohliqua descends anterolingually from apex of hypoconid, curving slightly anteriorly to terrnirtaie at pcstetohueeal base of protoconid; anterior cingulld is low on the base of erow-n, rounded and hulhouR in its lingual corner, tapering buecally lo icntiiiuiic at anlcrohuceal lia-vc ofprotoconid. Right jW<. Sub-rectangular in occlusal outline; irigoiiid and talonid arc of approximately equal width: lurinuuid is talk-si cusp followed by protoconid then paraconid; comparison of heights of talonid cuspids to ihoseofihe iri^unid not possible due lo wear and broken posterior lingual turner of talonid; paraconid sli.nlnly 11-.: 101 in .ulterior iv.ire.x p-.-.>11-1■ r-i^i.t in- lingual portion of tooth; metaconid directly posterior to paraconid; mctaconid and protoconid are in same trans\ei\e. plane: cnloconid directly posterior to mctaconid; hypoconid directly posterior to proioconid: cntoconid and hypoconid are in ihe ww. nans-verse plane, hypoconulid reduced to a slight swelling at posterior base of cntoconid: no posthypocristid present, cristid obliqua runs anterol ingual ly from hypoconid curving slightly anteriorly to terminate lingual to midline of touth at posterior base of mctaconid; anterior cingultd Jst"tsitioncd low 00 base of crown, rounded at Us lingual corner, tapering bltccally lo terminate at anterobuecal base ofprotoconid. Left AtV (Based on OMF438W, Fi$- ZC. D). Sub-rectangular in occlusal outline; tngonid wider lhan lalonid; relative heights of cuspids indeierminahlc owing to wear, but Irigomd cuspids arc clearly taller than those oflalonid: paracnri id slightly posterior lo<in1crior margin of tooth, posilioned lingual to midline; metaconid posterolingual to paraconid: metaconid and proioconid are on same transverse plane: entoeonid and hypoconid arc closely iippriniiiti.ned, positioned lingual to midline of tooth; lalonid has been worn such that hypoconid imd ntioconid appear to be fused lo form one main cuspid, however a slight constriction on apes suggests, demarcation between very small cntoconid and hypoconid cuspids, liriy.aal eornei 11 fanterior eingul id is one half the way up anterior face of paraconid: anterior cingulid rounded and bulbous in lingual corner, descending buccal!)1 to terminate al anlembuccal base of proioconid. A IR i l is c FIG 2. Penimttes wihM sp nov„ pnnnvpes, A. n, RH7 QMF'4J»7S), occlusal view: C, D, LMj (JMVJVUHCI, Hiccliisal view. Statu bars I mm. with its lowest point just lingual to its buccal coiner. REMARKS. This species is placed in Perameha based on iK anterior cingulid heing significantly lower than the apex of the paraconid. the result of iln- low position of ihe amerior eingdiJ or me crown (Smith. I TO), P. gfffagAamavti is fcnown only from an isolated right M:, which is larger than that of all other extant Perametes. Whi Ic P. lobbei is known only I'mm lower ilentition, it is smaller than extant species such as I' ativiita and P gunnii. hence is extremely unlikely to be referable to P. aHingfiameitiis, P xtMwi is clearly distinguished from P. hongamviik: P. eivmiww. and P hmvnsis by its much larger size. P. sobbeiis most similar in size to /' mnula and P. gUMtt, but differs from those species by pos- sessing 1hc lolloping eooibinoiiiin of features: (I) smaller P- and Pi. (2) longerMu(3) narrower M.i, (4) rounded lingual comer of the anterior cingulid on M_m. (51 hypoconulid reduced Oil M, and absent from M, -.. and (o) closer approx- imation of enlOeOrtHi and hypoconid on Mj. AFFINITIES. Pt'rtimele.i .itibbei shares a combination of plesiomorphic ami upornntphk- I9h itfnMoiRS of the Queensland museum TABLI: 1. Mi;;isiircmLHU ol' type spcjtiTnuni Of F&Oattiet stibhri feint QML796. AO maOliremtOta alt maximum <l wanees in mm. Length is flie nnterior-poMwior di*Hine«. Cent, width is the widih hcchws the central l-uspiJ dI" ;i pn.-tiKtlur. rt.ni. «iditi is me liikgual-biiceal disianLe accross iltc Biganid, Post, width is ittc lineaal-buccill distance JKCIOSS the latmtid Fiirail - puratinnil, mdmI incULiimd. pmrtixl pnihM.i<Kiid. l-iiIihI rtlloCGrtid. llvpoJ - hypoeonid. Twih 1 i ill CW. niljlh ■Mr ^ih Mii-ri wi^Jh mrlijl llr.'lj,l - |> .'. ,1 Krjml Iil|^l. HA PiA ItfA Sva Vlfi 1 111 ■\ ■, KA Mf* RH i IJ9S 1,14 . K< EM, V 2JG 1 : 1.17 I.7B (Hi til JJJ II l.ll> ll^l-i^um 1 I.Mh 3.41 h.A : I - \M IjH LJ3 imjrph(>ki<iK^il characters with muM modern Pereimtlet species. Muirhcad ct aL. (1997) vujiee-.lal lli.il I' tvmpnini-iil* is the fliOSt pIcHionioTphic of the let en t species by reason <>l its possession of ill* following coniblnKtioTi of characters: \ 1 ) equidistant paratonids, inclaconids and nnTtoconidK, (2] a more widely wparfllLv) rttntcLHtid sod mdseonid in compar- ison to other recent species, and (J Mn incomplete jimerioreinc.nlLd on M|. The plcsiomurphic char- acters of equidistant liigutiid Cuspid* flsiJ ■ l-I alivcly widely separated paraconid-mctacomd JllsOOCeUr ifl P. ft(iiiMTt.™-{\1ililhe;id el ill . I'M 7 I and. in P. svbbet In common with all recent Peramefvx eac-eptr? ertmisna, P- whfvi has a relatively slraijjhl crlstld obliqua, a condition regarded as pLcsiomorpliic by Muirhcad d al. (1997), wht> mm-d ili.: :nii:i|ii>tnorp1iic trail was a more .vrwo crMid obliqua as fvirl <>r" u narrower Lalmnid. Like J? mpmiama, a narrower tilonid is also prewnt in P. sobbti, but alternatively may have arisen by the closer approximation of ihe dii<xoitid and. hypoconid. Pn-umelrs xoNn'i and P. bttwetaa are the only species of ihe genus with the synapwiHiipliie reduced hypoconultd cm M i- In all other known species {with th<r possible e?tceplion of iillinfihtiniL'niii in which the M< is unknown) the ll<. |1ml I • 11.111.1 |% | ■ I ,.......11. 1 .11'. L I i i Ml i I! l ■■>, .1 i extension of the crown, as well us providing supfton for (he sueeceding. tomli. The Mi allrihuted to P. wyfrhvi puHsessc! u clowly jppfoaimaiwd cnl«oiiid J4»d hyp«cpiiid1 a trait regarded by Mutrihcud A P i Inn il^?Ms rtppfseritin^the plcsNimLWiihie eondilinn. This is unlike all recent Pcramvtrn species, whicli exhibit the anomorphie condition in whKh the eslocotrid is. conical in shape and is clearly distinct from (he hypoconid. Comparison with IheCOlMlilii....../•■ !'i\\,-i;sis mi! (Hy^ible iIul'Io thepoof swlt of ptreservaliortofilK Mj talonid in the known material. P. nrJiitfFUj'.v is marc plcisomorphic Ihan P. wlrhei in the possession of a pre-endxrislid on the M;. However, on the basts of ihe synaponftorph.be condiiion of the reduction oflhc hypocOnultd un Mi and its absence frOftl Mj, Peraim'les mttrfvi is here awarded as the sister tiLiion to the early Pliocene P. h/wenm. ki other respects, notably in sharing the characters of equidistant trigonid cuspids and wider pariiConid-mciiKxniiil, IilhIi sfecic^ are more plesinnio-rphie than P. nasuier, P. gunnii and P. crvntitriifi. The relationship of the P. trthhfl-P. bcwensii clade to P. bougaiaville remains unclear, huJ followint; Muirhcad eta!. (1997}, the P. sobhvi-P. AowwisiitcladeidBy represem asdster clade to the P. nmuta - P. gunnii - P. etvmiiAna clack Barth»lornai t1977) nuKd sliahtlv wnyller loo-th dimensions in the Peramelea nasuia population from the Pleistocene fissure-fill deposits of Ciore. stnith-eastcrn Queensland. 1nnn those in modem Queensland pnpalatjnns, hut did nci i^msider the differences syfikient lo warrant speeillc separation. While the possibility of a siinihn limiphivliji;!! ichition may also exist between the Pliocene P. bowensis and Pleislocene P. sablM, the much greater si-'e dillcrcnce and slight morpliologiciiI differences between the two. is here considered sufficient to wiin*anl spocifie sepurjiion ACKNOWLF.DOFVirNT!> I thank ^ietut 1 locknull. Bernie CoukeL Gre^j; Wcbh and Brendan Brooke for guidance in ptcpar;ii iim nl litis, manuscript- Joanne Wilkinson, Debbie Dolby.Or-iiham Jordan, Philip Colqhoun.. Ricky Bell. Brad Martin and Trevor Sutton are thanked for assistance in fossil preparation. Desmond and JcnniJcr Price are daki iMiixmNsn i isitH i ni nwnifoOT th linked for continued sup purl ;ind citcoufajjement. I especially Ihank lan Snhbe few his continued iissislunire inlo (he research on Pleistocene iaunas of the Darling Downs. This tesearelt has. been funded by lite Queeivshmd University of Technology and the Queensland Museum. LITERATURE! CITED AFI.IN. k P. ii AROII'R, M. I9K7 Recent advance* in marsa n. i systematics with a new syncretic e I as si tl cat ion, pp. xy-Iadui in M. Arctici (ed. lc Possums and Opossums; Studies in Evolution. Surrey. Beany & Sons and the Royal Zoological ShjciuIv ui New South Walev, Sydney. ARCHER, M. l^'fc. BlulV Downs Ideal laura, Pp. 383-197, In Archer,. M. A Wade, M., Result* of the Ray EL. Leniley expeditions. Pari 1. Tltc Allingham Formation and a new Pliocene vertebrate fauna from northern Queensland. Memoirs of the Ouecnsland Museum 17: 37<MU7. EiARTIIOI.OMAI, A. N77. Thcih™ilvcrtchralL-iaiin.i from Pleistocene depnsitsal Cement Mills. Gore, southeastern Queensland. Mcinuirs pi llie Oucenslamd Museum IX: 41-51 I't VIIR.d I..C. f" D. 1X17 ].crxiii.ncanimaldi.<irihu? d'aprcs son uiuymiviiion. pour servir de nase •* ITiisluirc nulmvlk des animaxiK ut d'lnliudiiLikin i I'linnlomx; enmparcc. Volume I. (Dctervillc Pa/is) «4up. FREEDMAN. L-1967. Skull and tooth variation to dm t;eniis /YrvjmrfVv Tart I: Anatomical Ic-.itur^v Records of the Australian Museum 27: 147-UhS. GEOFFROY. E. 1804. Mctmoirc sur un nouvcau genre de itlarluOl leres. a bourse. iMKmnc PertimcUw. Annalcs de la Musce Natioanl d' Histinrc Nature-He de Puns -J U*fA. GRAY. J.E. 1825. Outline of an attempt at the dupo-olion -al lit.' Mammalia into tnhes and Hi nil I '.uii i I.' ..I :h:' miierii ipptwnllj appenuinuij! to eaeh tribe. Annals of Philosophy- (new series) Ut: .117.>M. {iRAY, I.V.. llOH. On a new speeiizs cif Piftimrlw ftweedinp of the Zoological SocieP, ofLondon 1HJ»: 1. KIR St'I I. J. A. W l'Kik. I'loduuiiusul the eoniparilnc serology nl' Marsupial ia. Nature, I Liolnn 2 I 7' 41 Ma LI ( kt I I. W.P. IMUJ. An imUiKenelje assuMmcnt of denial homologies in therion mammals. Pp. IS2-204 in Szalay, F.S.. Novacck. M.J. £ Mcktrma. M.t . ledsl Mammal phylojcny. Volume I Springer-VcrlnK. New York. MACKNESS, B.S.. WROE. S.. MUIRHEAD. J. WILKINSON. C". t. & WILKINSON. D. M. 200(i. Fird fo«il bandicoot from the Pliocene < liiivhilla Local l-.iunu. Australian Mammalogy 22: LtJ-IJb. MOLNARuR E.&KURZ.C. IW7 The distrrtHilwn of Pleisioceite ventbraies on not eaaaara Darling M. v. i" . l-.a-.-d on the Queensland Museum eolleelKins. Ptoceedi ngs of the L innean Society of New SourJl Wales 117: 107-134. Ml IRIU AD. I., DAWSON, I..H ARCHI R.M l'W7 frnwirffs bowensa. a new specks of /Vnaamfa (Peramclcniurphia, MarMipialia) Ihim PliiKcne taunns of Bow arid Wellington Cnves. New South Wales. Proctodin.LLs of the Linnean Society of New Souui Walts 117: 163-173. MURIIPAD. J & FiLAN, S. L. IW5 YamSa bmdh flttdi, a piesioniorphie bandied** (Marsupialia. PeianiekiiiiirpJiial I'nicn OligiH-Mioeene deposits, ■ •\ Rhxt< eish, ntirth-.n-siem i.j nv lslund lou nnl of Paleontology 69: 127-Ii4. QIJOY, J.R.t iAIMARD. I P. IX2J. ?wloiiie-P Sfs in Voyage auiotirdu Monde. Freyeincl. L. C. de <exL), Pillet Aiuc. lifi|3Tinu.-ur-Lihraire: Pans. OWI'iN', R. 1K77. Rettcarehes on the fossil remains of the eninet mammals of Australia. witJi a ikuIlc-oI lite extinct uur^ipiaK of Lnglarxl. ). Prxlchcn: I nr.don SMITH, M. J. 1972. Small fossil vertebrates Iron. Vietivia i a\c, Naraeixirte, South AuslTalia, II. Peramelidae, Thylncindae and Dasyuridae < Mai supialta). Transactions of die Royal Society ofSnudi Australia "6: 125-L>7 SPENCER* W B I rB7. Oeseription of two new species otlna Nupi.ih rnuu ...t-Ii a 'mi-.i ;i i.i Pi.'i.......lii;.". of the Rnj-al Society of Victoriu n s 0: 5-11, SEA LAY. F. IW2. A new appraisal of marsupial Iihyloacriy and eliLssiliealum. I"fi. (i2 I W(l in M. An;hcr (ed,k Cnmivonou* marsupials. Stttiey Beany A Sons and Royal /uoHsgieal Society of New South Wales, Sydney.