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Nimbadon, a New Genus and Three New Species Oftertiary Zygomaturines (Marsupialia: Diprotodontidae) from Northern Australia, with a Reassessment of Neohelos

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Nimbadon, a New Genus and Three New Species Oftertiary Zygomaturines (Marsupialia: Diprotodontidae) from Northern Australia, with a Reassessment of Neohelos NIMBADON, A NEW GENUS AND THREE NEW SPECIES OFTERTIARY ZYGOMATURINES (MARSUPIALIA: DIPROTODONTIDAE) FROM NORTHERN AUSTRALIA, WITH A REASSESSMENT OF NEOHELOS S. J. HAND, M. ARCHER, H. GODTHELP, T.H. RICH AND N.S. PLEDGE Hand, S.J.,Archer, M., Godthelp,H., Rich, T.H. & Pledge,N. S. 199306 30: Nimbadon,a new genusand three new speciesof Tertiary zygomaturines(Marsupialia: Diprotodontidae) from northern Australia, with a reassessmentof Neohelos. Memoirs of the Queensland Museum33(l): 193-210.Brisbane. ISSN 0079-8835. Three new speciesof Oligo-Miocene zygomaturinediprotodontids ire describedfiom northernAusiralia. All are small,plesiomorphic and appearto comprisea distinctiveclade of zygomaturines, naqred here Nimbadon The clade is partly defined on the basis of the posteiiorlyi4clined P' parastyleand bladeextending from the parametaconeto the lingual half of the P' crown and then to the anterolingualcingulum. Two of the speciesare known from Oligo-Miocenelocal faunas of RiversleighStation, northwestern Queensland. The third is from the middle Miocene Bullock CreekLocal Faunaof Camfield Station,northwestern Northern Territory. Description of additional fossil material referableto Neohelos tirarensis Stirton, 1967,a middle Miocene zygomaturinefrom the KutjamarpuLocal Faunaof South Australia, enablesthe genus to be distinguished fiom Nimbadon. The chronostratigraphic significanceofthe new zygomaturinesis consideredand found to approximatelycorrespond to currentunderstanding of the relativeages of the depositsfrorn which the specimenswere obtained. I Diprotodontidae, Zygomaturinae, Nimbadon, Neohelos, Oligo-Miocene, Riversleigh, Bullock Creek Incal Fauna, Henk's Hollow Incal Fauna, Fig Tree I'ocaL Fauna. SuzanneHand, M. Archer & H. Godthelp, School of BiologicaL Science, University of New South WaLes,PO Box l, Kensington,New South Wales 2033, Australia; T.H. Rich, Pakteontology, Museum of Victoria, Russell St,Melbourne, Victoria 3000, Australin; N.S. PLedge,PaLaeontology, South Australian Museum, Norlh Terrace, Adelaide, SouthAustra- Iia 5000,Australia;24 November,1992. Specimens representing a very small zygoma- (Rich et al., 1982, l99l; Archer & Flannery, turine were collected between 1984 and 1986 by 1983; Archer & Hand, 1984; Woodburne et al., Archer, Godthelp and Hand from the Henk's Hol- 1985; Munay, l99Oa; Murray & Megirian, 19,90; low Site, in an as yet unnamed freshwater lime- Archer et al., 1989,1991). Recent reappraisal of stone on the Gag Plateau of Riversleigh Station the age of central Australian Tertiary mammalian (Archer, Hand & Godthelp, 1986; Archer & Flan- faunas, based on the study of foraminifera (Lind- nery,1987; Archeret al., 1989, 1991). Refened say, 1987), suggeststhat some of these faunas material includes maxillary and dentary frag- (e.g. the Ditlimanka, Pinpa and Ericmas l.ocal ments and isolated teeth. A second,. larger Faunas) are appreciably older than once thought Riversleigh species, represented by a maxilla, (e.9. Woodbume et al., 1985; but see Tedfbrd, was collected in 1986 by Pledge from Fig Tree 1966), possibly late Oligocene. By comparison, Site Locality adjacent to Godthelp Hill, the Bullock Creek Local Fauna is estimated to be Riversleigh Siation. ?middle Miocene in age, i.e. immediately post- A small Bullock Creek zygomaturine is known Wp-ajiri- (Munay, 1990a; Munay & Megirian, (Archer et al., l99l) of from a palare collecred i;"1981, by Rich and 1990). On-going ^studies the colleagu'es,from Horseshoc West Locality (WV the Riversleigh- fossil-faunas suggest -th.at I l3) i; the Camfield Beds of Camfield Stuiion, Henk's Hollow Lxrcal Fauna is ?middle Miocene northwestern Northern Territory (Rich et al., in age and the Fig Tree Local Fauna somewhere lgg1, lggl). between late Oligocene and early Miocene in age. Estimates of the age of the Henk's Hollow, Fig The new zygomaturines described here are Tree and Bullcrck Creek Local Faunas are based most similar to forms currently referred to the mainly on stage-of-evolution comparisons of par- genus Neohelos. Their generic distinction, how- ticular diprotodontian marsupials with those of ever, has been clarified by discovery at the type other northern and central Australian faunas lcrcality of new material referable to NeoheLos 194 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE L Measurements of Neohelos tirarensis P3 ing unchallengedconvention, is called the hypo- (AMF87625) and M'J (AMF87626) from rhe teaf cone.Cheektooth homology is that proposedby t-ocality,Lake Ngapakaldi,South Australia.In mm. Archer (1978). Higher level systematicnomen- Abbreviations: ant, anterior; post, posterior; paramet, claturefollows Aplin ( parametacone;protoc, protocone; parac, paracone. & Archer 1987). Length 15.8 SYSTEMATICS width 13.9 c SuperorderMARSUPIALIA Illiger, l8l I width(paramet to Drotoc tiDS) 5.5 Order DIPROTODONTIA Owen, 1866 heisht(oaramet) t1.4 FamilyDIPROTODONTIDAE Gill, 1872 height(parastvh) 6.5 SubfamilyZYGOMATURINAE Stirton, length 18.8 Woodburne& Plane,1967 M] width(ant ) t].2 width(oost) 15.8 NeohelosStirton, 1967 height(parac.1 7.8 (Fig. l, Table l) tirarensis,the type andonly namedspecies ofthe Typespecies. Neohelos tirarensis Stirton, 1967 genus.The original description(Stirton, 1967) of this Miocenezygomaturine from the Kutjamarpu RevrssnGsNeRrc DrncNosrs Local Fauna of central Australia was basedon The following combinationof featuresappears five isolated teeth collectedin 1962at the teaf to distinguishspecies of Neohelosfrom all other Locality, Lake Ngapakaldi, Etadunna Slation, zygomaturines:P' with large,erect parastyle that South Australia.The holotype,a brokenP', pre- is well isolatedfrom the parametacone;parastyle serves posterior only the portion of the tooth. is conicalwithout tip developedas.a blade; an- Thoughless complete than the otherteeth, it was terolingualbasin well defined in Pr by distinct selectedas the holotype hcause of the impor- basalcingulum extending between parastylar cor tanceof premolar morphologyin diprotodontoid nerof toothto anterolingualbase of protocone;P' (Stinon, systematics 1967;Stirton, Woodbume parametaconewith an undivided tip; parameta- & Plane,1967). cone tip not developedinto anterobucallyori- In 1982 two isolatedteeth referableto N. rl- ented blade; line between protocone, rarensiswere collectedfrom the type locality by parametaconeand widest buccalpoint on crown Archer,Hand, T. Flannery,G. Hic^kie,J. Caseand is approximatelyrectilinear and dividescrown P. Bridge: a well-preservedRPr (AMF87625) either in half or leavesanterior division shorter and RM' (AMF87626). Theseadd considerablv than the posteriordivision; Pi with^well-devel- to knowledgeof this species.Additional materi- opedhypocone or hypoconeshelf; M'--'with well als referableIo Neohelosfrom the Bullock Creek developcdparastyle and metastyle. Local Fauna(Plane & Gatehouse,1968; Plane, 1971; Munay & Megirian, 1992) and various Nimbadon n.sen. Oligo-MioceneRiversleigh local faunas(Archer (Figs2-5. Table 2) et al., 1989,l99l) arepresently under study. lnstitutionalabbreviations used here are as fol- Type species. Ninbudon lavarackorunr t't.sp. lows: QMF, fossilcollection of the Queenslanc Additionol species. Nintbadon vyhitelawi n.sp., Nlrn- Museum; AMF, fossil collection of the Austra- badon scotton'orunt n.sD. lianMuseum; SAMP, palaeontological collection of the South Australian Museum; NMVP, pa- GnNrnrcDrAGNosrs laeontologicalcollection of the Museumof Vic- Speciesof Nitnbadondiff'er tiom those of all toria; AR, researchcollection of the School of other zygomaturines(with special referenceto Biological Sciences,University of New South Neohelos,the taxon to which they are otherwise Wales. Cusp nomenclature follows Archer most similar) in the following combinationof (1984) and Rich, Archer & Tedford (1978).The fcatures:small size (although they arelarger than poslerolingualcusp of theupper molars, conven- RaemeotherianRich, Archer & Tedford, 1978); tionally calledthe hypocone(e.g. Stirton, Wood- in havinga molar gradientthat does not apprecia- burne & Plane, 1961), is now called the bly increaseposteriorly (in contrastto all other metaconule following Tedford & Wqodburne zygomaturinesexcept Raemeotheriam); posteri- (1987)but theposterolingual cusp of Pj, follow- orly inclinedparastyle on P' (ratherthan conical Nts,WDIPROTODONTIDS FROM NORTHERN AUSTRALIA l9-5 I:IG.l.Neoltelostirurett.si.;,teaf[-mality,LrkeNgapakaldi,$outhAu.^ttaliaAMI;87625, I{I'}]:A-A',treltrsitl stereopair;B, buccal vicw; C, lingual view. AMF87626, I{M-': D-D', crcclusalstcrcopitir Brtrindiclrtcs l()rrtrtt andcrcct as in othcrzygomaturines exccpt posst- z-ygonralurincscxccpt Neohclo.v und Alkwu turlt- bly Plaisiodott centrali.sWoodburne, 1967); an eriutnMurray, 1990b);an_antcrobuccal blade on antcrolingualbladc on thc parastylcof P' that thc paramctaeortcof P' that sun)lout)lsan surmountsa posterolinguallyoricnted thegotic obliquelyoricntcd anterolingual thcgotic facette lacctte(in contrastto no apical bladc such as (in contrastto eitherno bladcor a poorlydcvcl- may[?]charactcrisc Neohelos): completcly undi- opcd antcriorlyorientcd bladc such as may[?l vidcdparametaconc on F (in contrastto all other charactcriseNcolrclos); vcry snrallto absenthy- 196 MEMOIRSOF THE QUEENSLANDMUSEUM TABLE 2. Measurementsof Nimbadon lavarackorum n.sp.,N. whitelawin.sp. and N. scollorrorum n.sp. from Riversleighand Camfield Sta- tions, northern Australia. In mm. Brackets around figures indicate estimates. Abbrevra- tions: ant, anterior; post, poste- rior; *Gag Site specimens. UPPER DENTITION N. Species
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