A new genusand speciesof dasyuromorphianfrom the Miocene of Riversleigh,northern Australia STEPHENWROE

WROE, S.,2001:12:20.A new genusand speciesof dasyuromorphianfiorn thc Mioccncof Riverslcigh,nofihem Australia.Menoirs of the AustrulianAssociution of Pulueontologists25. 53-59.ISSN 0810-8889

Jocttlttsiuntntttizoni, new genusand spccies,is the leastderived dasyuromoryhian yet described from the Oligoccnc-Miocenedeposits of Riversleighin northwesternQueensland. Within ,J.muizoni is plesiomorphicfbr all dcntalfeatures examined excepting metacristid orientation.Oblique alignmcnt of this cristidwith respectto the long axisof the dentaryin./. tttuizoni is an apomorphywithin Dasyuromorphiasharcd with some representativesof both Thylacinidaeand . Conscqucntly. placement of this newtaxon at the family levelis untenable.Analysis of the materialdraws attentionto the absenceof well supportcd synapomorphiesfor the dasyuromorphianorder. Considcred together with temporalas well as geographicposition, the prcscnccof a featurewhich is possiblysynapomorphic with some Thylacinidaeor Dasyuridae,is trcatcdas a reasonablebasis for the placementof J. ntuittni within Dasyuromorphia.

S. Wroe,(s.v:roe(cl,un,sw.edu.au) School o./Biologit'al Sciences,Universitl, of'S.v"dne1,,Nev.'South Wales,2006 and S<:hoolo.f Biologic'al Sc'ience,Universitl, o.f Nev South Wales,Nev'South Wales, 2052.Received I June2000

Keywords:Joculu.s iuttt muizoni, Dasyuromorphia, Mioccne. Riversleigh

DASYUROMORPHIANS havc been Recentlydiscovered material from earlyTertiary suggestedto be structurally ancestralto the depositsof Murgon, southcasternQueensland remainderof the Australianmarsupial radiation (Godthclpet al. 1999)hasadded further complexity (Bcnsley1903, Ride 1964.Szalay1994). Howcvcq to our understandingof dasyuromorphian the f-actoroften flagged to suppofta basalposition evolution.Australia's oldest marsupicamivore, for the clade,that is, the retentionof essentially Djurthia mtu'gonensis,may bc refcrableto either primitivemarsupial rnorphology, hinders attempts Didelphidae(sensu Marshall et a\.,1990)or to demonstratea spccialrelationship between Australidelphiaon the basisof dcntalevidence. dasyuromorphiansand any specificSouth If the latter is ultimatcly supportedby the Amcricanclade. The discoveryand interpretation unambiguousassociation of D. mttrgonensi.vwith of new Australianfossil material has added new australidelphiantarsal material, then it becomes dimensionsto debateon the topic of dasyuro- more difficult to define Australidelphia, morphiancvolution (Wroe 1998.| 999a).Archer Dasyuromorphiaor Didelphidaeon the basisof (1976b,1982), Wroe (l 996,1991a, I 997b, 1999b, in dcntal morphologyalone. A conservative prcss),Wroe e/ al. (2000) and Wroe & Nasscr approachis requiredin the taxonornicplacernent (2001)have denronstrated that cvcn generalised ofgeneralisedAustralian fossil taxa, such as that dasyurids,commonly thought to approximatethe describedbelow, particularly in the absenceof plesiomorphicaustralidelphian statc for many basicranialmaterial. features,arc highly derived in their cranial morphology.Moreover, Wroe ( 1999b)argues that SYSTEMATIC PALAEONTOLOGY althoughthe family Dasyuridaecan not bc defined by thepresence ofdcntal synapomorphies.shared OrderDASYUROMORPHIA Gill" I 872 derivedbasicranial features demonstrate a special relationshipbetween the middle Miocene Joculusiumgcn. nov. 'modern dasyurid,Barinl:a wungala. and all Dasyuridae'(seir"su Wroe 1999b),with 'modern T1'1tt':pL'cit'.s..loctrltt.:itrnr nrtri:ttni sp. nov. Dasyuridae'united by a furthcr suite of synapomorphicbasicranial features. Diagno.sis.Joculusiunt muizoni differs from all 54 AAPMemoir 25 (2001)

known dasyuridsand thylacinids in the following combinationoffeatures: Prequal to or exceeding P, dorsoventrally; Mr paraconid large, circumscribedby a well definedanterior cingulid; cristid obliqua terminatesjust buccal to the carnassialnotch of the metacristid on Mr-o; trigonid shorter,anteroposteriorly, than the talonid on M,_,;metacristid almost equal to the paracristid in length on Mr-o;postprotocristid-metacristid and hypocristid strongly torsioned away from transverseorientation to dentary on M,_o; posteriorface of the trigonid on M,_ois anteriorly inclined;entoconid large on M,_.and distinct on Mo; hypoconulidnotch preseniin the anterior cingulid of Mr-., but not deeply recessed. Joculusium muizoni can be distinguishedfrom known bandicoots(peramelomorphians) by the abovecombination of charactersand possession ofa well definedposterior cingulid, a morebuccally positioned hypoconulid and a less lingually positionedcristid obliqua. 'a Etymologt. From the Latin joculus, meaning little joke', in referenceto the type locality (Gag Site).

Joculusium muizoni sp.nov. (Fig. l) Fig. I . Joculusiummuizoni gen. et sp. nov., QMF36442, holotype,left dentarycontaining the posteriorroot of Diagnosis.As for the genus. P1,P2 (exceptingthe occlusal tip), and P3, Mt-+. Stereopairin occlusalview. Scale bar indicates 2 mm. Etymologt. After Frenchpalaeontologist Christian de Muizon, in recognitionof his greatcontribution The crown of P,, the anterior root, and the to marsupialpalaeontology. lingual portion of the anteriorroot alveolusare broken away. Only the posterior root remains.A Holotype. Qli{F36442, a left dentary containing small diastemaseparates P, and Pr. theposterior root ofP,, P, (exceptingthe occlusal In P, a portionof the crown is missinganterior tip), andP.,, Mr_4. to the posterior root. A clearly defined cristid ascendsthe posterior face of crown, from a small Locality and age of material. Gag Site, centralcuspule on the heelto wherethe crown is RiversleighWorld Heritagefossil property,Lawn brokenaway. Another cristid continuesfrom the Hill National Park,northwestem Queensland. The posterocentralcuspule along the buccal crown fossil material recovered from Gag Site is margin to a point between the anterior and interpretedto be middle Miocene in age on the posteriorroots. From the distancebetween the basis of the site's stratigraphicrelationships to anterior and posterior roots it is inferred that P2 is other Riversleigh depositsand its contained the longestpremolar anteroposteriorly. (Dwornamor) local fauna (Archer et al. 1995, P. is wider than P, transverselyand gently Creaser1997). recurvedin lateralview. A cingulidcircumscribes the heel and is bisectedby anothercristid which Description. In buccalview the dentaryis roughly ascendsthe posteriorface of the principal cusp uniform in depth posteroanteriorly from Mo to from the small, posterocentralcuspule. The beneathanterior root of M, , but tapersfrom M, to principal cusp and M, protoconid are of nearly beneaththe posterior root of P,, and is broken equalheight. It is infened that P, probably exceeds away from 2 mm posteriorofM, and I mm anterior P, in height, but damageto P, precludescertainty. ofP root. In M, the paracristid is the main vertical shearingcrest. Principal cusps in order of AAP Memoir25 (2001) decreasingheight are: protoconid,paraconid, Meristicgradient,s /rom M, ,. The orientationsof metaconid,cntoconid, hypoconulid and the paracristid,metacristid and hypocristid are hypoconid.Although taller than the metaconid, increasinglytransverse with respectto the long the paraconidis the smallerof the two with the axis of dentary, the angle fon.nedbetween the protoconidthe largestcusp of thc M, trigonid. paracristidand metacristid is increasinglyacute; The metaconid is largc and positioned protoconidheight increases M, . (M, protoconid posterolinguallywith respectto the protoconid. tip rnissing):metuconid height increases liom M, A right angleis formed betweenthe paracristid- to M,, then decrcascsfbr M.,;the metaconidis preprotocristidand the postprotocristid- largcr than thc paraconidon all molars but n.retacristid.An acuteinternal angle is formcd dccrcascsin sizcfcrr M,_..,: talonid width increases betweenthe cristid obliqua and hypocristid. Both from M,_,,but dccreasesfrom M._.,. the metacristidand hypocristidarc oricntcdat about forty-five degreesto the long axis of the CHARACTER STATEPOLARITIES dentary.The cristid obliqua ten.ninates anteriorly Method. Most f-eaturesconsidered in the present just buccalto the 'camassialnotch' separating study have been examinedby Archer (1976a, the metacristidand postprotocristid.In occlusal 1916b), Reig et al. ( 1987),Marshall et al. (1990), vicw thc surfacearea ofthe talonidis muchgreater Wroe( 1996, 1991 a, 1991b.I 999b).Wroe e r al. (2000) thanthat ofthe trigonid.The posterior face ofthe and Godthelpet al. (1999).Further details trigonid is inclined.A well deflnedanterior regardingtheir distributionamong dasyuro- cingulidruns anteriorly from the buccalbase of morphiansand relevantoutgroup taxa (i.e. theprotoconid to beneaththe carnassial notch of Peradectidae,Didelphidae, Djarthia murgonensis theparacristid, whcrc a smallnotch accommodates (Marsupialiaince rtae sedi^r),Microbiotheriidae, thc hccl of P.. The posteriorcingulid is well- Peramelemorphia)is contained therein. Systematic developed,running from the hypoconulidto the nomenclature for Dasyuror.norphia fo II ows Wroe baseofthe hypoconid.A distinctbuccal cingulid ( 1996)with highcrlcvcl n.rarsupial systcmatics after is almostcontinuous with the anteriorand Marshallet al. (l 990). Dentalnomenclature posteriorcingulids. The hypoconulidis not followsFlowcr ( I 867)and Luckett ( I 993)regarding deeplyrecessed into antcriorcingulid of M.. the molar-premolarboundary, such that thc adult M. dificrsfrom M in thelollowing wayi: the (unreduced)postcanine cheektooth formula of metacristidis ahnostas long as the paracristid; marsupialsis Pl -3 andMl -4. thetrigonid is longerthan the talonid; the trigonid and talonid are aboutequal in width; in occlusal Morphologl,o/ P, A P. that equalsor exceedsP. view the internalangles fbrmed by the paracristid in heightis plesiomorphicfbr Marsupialia(Archer and metacristid,and the cristid obliqua and 1976a, Muirhead & Wroe I 998,Wroe 1996,1991 a, hypocristid,are more acute;the entoconidlarger; 1997b,1999b).Marked hyperlrophy of P., as in and a distinct hypoconulidnotch beneaththe some Thylacinidae (r.e. Thl,luc:inus)and, anteriorface of the paraconidaccommodates the Borhyaenidae,as well as reductionor lossas in M, hypoconulid. some Dasyuridac(c.g. Dasyurus)are derived. M, morphologyis sirnilarto that of M. except Joculusittmmttizoni is plesiomorphicfor this as follows: the trigonid is wider but more feature. compressedanteroposteriorly, with the angle betweenthc paracristidand metacristidmore Morphologl;ol theM ,paraconid. The paraconid acute:on the transverseaxis the talonid width is is well developedon M, in unspecialiscd lcssin both absoluteterms and relativeto that of representativesof outgroupsto Dasyuromorphia thc trigonid;the anteriorend of thecristid obliqua (ArcherI976a, Wroe 1996,1991a, I 997b),and many terminatesin a more buccal position and the dasyuridand thylacinid taxa. This cuspis smallto entoconidis smaller. abscntin some dasyuromorphians,especially Thc tip of the M. protoconidis brokenoff. M.' among carnivorousDasyuridae (e.g. Da.svtrru.s diff'ersfiom M. asfollows: both thetalonid width spp.).The largeM, paraconidof .1.muizoni andthe hypoconidare reduced; the paraconidis represents a plesiomorphy within larger,the entoconidand metaconid are smaller: Dasyuromorphia. unteriorly.the cristid obliqua tcrminatcs at a point Iingualto thecarnassial notch of thc mctacristid; Morphologyof themetaconicl. Metaconids which and the angle formed between paracristidand are well developedon M, as well as M._,,are metacristidis moreacute. treatedas plesiomorphic for Dasyuromorphiaby AAP Memoir25 (200I )

Archcr (1916)ancl Wroe ( 1996,I 991a,1991b). Calurontl,s)and all Microbiotheriidaehavc Bothunifbrm reduction of themetaconids on M, talonidsthat are clearly longer than the trigonids , (c.g. in rrost thylacinids),and differential on all lowermolars. In manymarsupials trigonid reduction,with reductionof thc M, metaconid andtalonid lengths are almost equal, including in greutlycxceeding lhul ol'M.., te.g. in specicsol' some Didclphidae(e.g. Didelpftis),some Da s I urrt r, Sa rc' o p hi Ius), are derived. J oc tr I trsi tt m Dasyuromorphia(e.g. Ankotarin ja tirarensis)and rrtuizoniexhibits thc plesiomorphicstate. Perameler.norphia(e. g., Pe r or,- c' te s), Dj nrI h i ct mtrr go ne n s i s and .lo c' uItt,s i u m mtt i z o n i. At-nong Orientationof the metacristid. This cristid these taxa, slight variation in subjective (protocristidof Marslrallet a|.1990) is oricnted determinationof the trigonidlalonid boundary 'trigonid transversclywith respectto the long axis of the might result in their receivingeither 'talonid dentary in most rcpresentativcs of longerthan talonid' or shorterthan dirsyuromorphianoutgroup cladcs. Anktturinia trigonid' status.Bccause of variation among tiraren.si.sand Wukamutha losselli (both outgroup taxa and disagreementbetween Dasyuromorphiaincertae sedis),and among authorities,Godthelp et ul. ( 1999) were non- dasyurids,r.nost sminthopsines. Previous authors committalregarding the assignmentol polarity havetreated this character state as plcsiomorphic for this f'eatureamong Marsupialia. However, the 'sub-cqual' for Marsupialiaand Dasyuromorphia(Archer prcsenceoltrigonids and talonids of 1976a,1976b;Van Dyck el al. 1994:Godthelp et lcngthin the leastdcrived Australian al. 1999).Thc derived statefor this f'eature(i.e. (Diarthia murgonensis)as well asothcr gcnerally obliqucorientation of thc metacristid)is prcsent unspecialiscdfossil taxa (e.g.Ankotarinja in somePeradectidac and Didelphidac(Archer tirarensis,Keeunu v,oodburnei),supports thc 1976a).at leastone taxon treated as plesiomorphic view that this reprcsentsthe plesiomorphic fbr mostdental characters within Dasyuromorphia conditionfor Dasyuromorphia.Consequently, J. (i.e.Ke eu no v'o o d bw'ne i), thylacinidswhich retain ntuizoniis treatedas plesiomorphicwithin the a mctaconid (e.g. Muribac'inusgadivuli, orderfor this f-eature. Batllcinusturnhulli), and all Phascogalinacand .In thc presentstudy it is accepted Anterior point o./'termination o./'the cri.stid that transverseorientation of thc metacristidis cthliqua. The distributionof this featurc is plesiomorphicfor Dasyuromorphia.However, consideredby Archer( 1976a,197 6b), Marshall et pervasivehomoplasy must be acknowlcdged a/.( 1990),Muirhead & Filan( 1995)and Godthelp regardlessof thc polaritydecision taken. Thus, et al. (1999).Termination of this cristidbencath within Dasyuron.rorphia,the derivcd state (oblique the carnassialnotch of themetacristid is probably alignrnent)must have evolvcdindependently plesiomorphicfor metatheriansand thc presence within Thylacinidaeand Dasyuridae,unless the of both buccal (c.g. Djarthia rnurgonensis, presenceof transverselyaligncd metacristids in .Jo c' u I us iunt mu izo n i, most Di delphimorphia and mostSminthopsinae is trcatedas an apomorphic Dasyuromorphia)and lingual placcment (e.g. nrost revcrsalto a primitive state.Evcn among Peramclemorphia)are derivcd(Godthelp et a/. srninthopsincsoblique orientation of this cristid 1999).Termination ofthe cristidobliqua just buccal is prescnt in Sninthopsi.sleucopus requiring to the carnassialnotch. evidentin Joculusiunt acccptanceol at least three indepcndent nuizoni, rcpresentsa likely dasyuromorphian derivationsof this featurewithin the order. plesiomorphy. Jot:ulusiummuizoni is apomorphicfbr this characteramong dasyuromorphians,which Positictnof'the h-vpoconttlid.Placement of thc rcpresentsa possiblesynapomorphy with any of hypoconulidslightly posterobuccalto the threetaxa within theorder. entoconidis plcsiomorphicfor Dasyurornorphia andDidclphimorphia. Presencc of a hypoconulid Reltttivelength.s of trigonid.sand talonids. The dircctly posterior to the entoconid is a trigonid may be anteroposteriorlylonger, equal peramelemorphiansynapomorphy (Muirhead & to. or shorterthan the talonid. The distributionof Filan l 995). Joculttsiuntmuizt.nri shows the this featr.rreamong polyprotodontmarsupials is plesiorlorphiccondition. considcredby Reiget al. (1987),Marshall et a/. ( 1990),Marshall & Muizon( 1995),Springer et a/. Developmentof'the anterior cingulid. A distinct (1996)and Godthelpet ul. (1999).Some anterior cingulid is presentin peradectids, Peradectidac(e.g. A lphadon), Didelphidae (e.g. microbiotheriids,peramelemorphians, Dj arthio AAPMemoir25 (2001) )/ nrurgonensi"i,and most Ameridclphiaand Hypcrlrophyof the Ma preparacrista,although Dasyuromorphia.This featureis oftcn poorly certainlyderived within Marsupialia and common developcdor absentin largecarnivorous taxa (e.g. l()most dasyuronrorphians. isnot present in somc Borhvuena, ThylaT'irt,.tr.In a number of generaliseddasyurid taxa (e.g.lVeophascogule). omnivoroustaxa the anterior cingulid is markedly However,it is presentin somepossible sister taxa hypertrophied(e.g. Didelphis. Wukumatha to Dasyuronrorphia(i.e. Djarthitt murgonen.sis, ta.sse I I i, andPeramelcmorphia). Both hypeftrophy Peramclemorphia).Regarding the relative size s of and hypotrophyof this feature,rclative to that thc trigonid and talonid,there is disagrccrnent cviderrtin peradcctids.are apomorphicfbr both over whicl.rconstitutes the plcsiornorphic Metatheriaand Dasyuromorphia..Ior:ulusiunt nrarsupialcondition (Godthelp et al. 1999). muizoniis plesiornorphicfor thisf'eature. Within a phylogenetic framcwork the elucidationof higher level relationshipsfbr J. Morphologt;ofthe M,talonid. A well dcveloped muizctniis problematicat both the familial and M, talonid with thc hypoconid,entoconid. and ordinallevels. In ten.nsof overallsimilarity, this hypoconrrlid cach pt'esenl.is cotntttottttr speciescould bc placedwithin Dasyuridae.but Peradectidae,most Didelphidae, some only on thc basisof symplesiomorphy.Whilst Peramclcrnorphia.Djurthio mLtrgonensis,some somedasyurids retain a dcntitionthat may bc Dasyuridaeand Thylacinidae,and Joctrlttsitrnt plesiomorphicfor a higher clade within ntuizotti.Reduction of oneor moreof thcsccusps Dasyuromorphia,ifnot Dasyurornorphiaitself, all is evidentin microbiotheriids.son.re didelphids Dasyuridaecan be dcfined by a number of andperamelemorphians, and most dasyurids and basicranialsynapomorphies, as demonstratcd by thylacinids.Presence of a wcll developedM. Wroe( 1996,1997a, 1999b) and Wroc c1a1. (2000). talonid has conrmonly been treated as a The singlc derived dasyurouorphianfeature dasyuromorphianplcsior.norphy (Archer 1976a, identifiedtnJ. ntuizoni,oblique orientation of thc 1976b).Reduction has clcarly occurred metacristid,is sharcdwith sometlrylacinids and independentlywithin severalmarsupial lineages dasyurids.Clonsequcntly, in the absence ofcranial and at leasttwicc within Dasyuromorphia(i.e. rrraterial,it is irlpossibleto positionJ. muizoni Dasyuridacand Thylacinidae)..Jot'ttlusitutt within the ordcr. Even at the ordinal level ntuizoniis plesiornorphicamong dasyuro- circurnspectionis required.Loss of Isremains the morphiansfor thisfeaturc. only wcll sr,rpporteddental synapor.norphyfor Dasyuromorphia,a fcature not yet known in .,/. DISCUSSION ntuizoni.Howevcr, the presenceof a derived Many researchers in the field of' f-eaturethat may indicatespecial relationship with dasyuromorphianphylogeny have treatedthe either sorneThylacinidae or Dasyuridae,in ordcr as indeflnablcon the basis of dental conjunctionwith stratigraphicand geographic rrorphology(Ride 1964,Archer 1916b,Szalay position.arc together treated as a rcasonablebasis 1994),with thepossible exception of a reduction for thc placement of .J. mttizorziwithin in incisorfonrrula to 4/3.Marshall et al. (1990)put Dasyuromorphia. fbrward two additional dental t-eaturesas synapomorphicfor the clade, that is, M, ACKNOWT-EDGEMENTS preparacristaelongated and talonidsreduced I am indebtedto M. Archerand R. Cifelli for relativeto thc trigonids.Nonc of theseputative theirconstructive criticism and comment on drafts synapomorphiesare uncquivocal within of this manuscript.Funding was providcdto S. Marsupialia.Four upper and thrce lower incisors Wroe through grants from the following are found in some borhyaenoid and institutions:Univcrsity of Sydney(U2000 peramelemorphiantaxa, as well as all PostdoctoralResearch Fellowship), French Dasyuromorphia.While therecan bc little doubt Ministry of ForeignAtfairs, Linnean Society of that lossof I5occurred independcntly in eacho1' Ncw SouthWales, Australian Geographic Society, thescthree clades, this is notnecessarily truc with Instituteof Wildlif'eResearch, and thc University rcspectto lossof I,. The fburth lower incisoris ofNew SouthWales. Supporl has also been given absentin all Australian taxa and thcrefore by theAustralian Research Clouncil (to M. Archer); representsa possiblesynapomorphy fbr the National E,stateGrants Schcme(Queensland) Australian rnarsupialradiation. Polarity (grantsto M. Archer and A. Bartholomai); determinationsfor the additional f-eatures Deparlmentof Environment,Sports and Territories; proposcdby Marshallet al. (1990')are debatable. QueenslandNational Parks and Wildlife Service; 58 AAP Memoir 25 (200 I )

Commonr.vealthWorld I {eritageUnit (Canberra); MARSHALL, L.G. & MUIZON, C.de.,1995. ICI AustraliaPty Ltd; QueenslandMuseum; Put:ude lph.t:.s anrllira,r (Marsupialia, Mammalia) fronr AustralianMuscum; Ccntury Zinc Pty Ltd: Mt the early Paleoceneof Bolivia.Part II. Thc Skull. lsa Mines Pty Ltd; SurreyBeatty and SonsPty Mlntoires du Mushtm Nationul d'Histoire Nuluralle Ltd; RiversleighSociety Inc., RoyalZoological 165,21-90. Socictyof Ncw SouthWalcs and many privatc MUIRHE,AD,J. & FILAN, S., 1995. Yurulu suppofters.Skilled preparation of mostRiversleigh burch/i e I d i (Peramelemorphia)frorrr Oligo-Miocene materialwas carried out bv AnnaGillesoie. dcpositsof Rivcrslcigh,nofthwestern Queerrsland. JournuI ol Puleontuktgt,59, 127-134. REFERENCES MUIRHEAD. J. & WROE, S., 1998. A ncw gcnus ARCHER.M.. 1976a.The dasyuridclentition and its anclspecies, Budjtinus turnbulli (Thylacinidae: relationshipto that of didelphids.thylacinids, Marsupialia),from the late Oligoccnc of Rivclslcigh, borhyacnids(Marsr-rpicarnivora) and pcramclids nofthcrnAustralia, and arr investigation of thylacinid (Pcrarrclina: Marsupialia).Austruliun Journul of phylogcny..hturnal oJ'Vertehrtfte Paleontologv 18, Zrtologt:.Strpltlamenlurt. Serias 39, 1-34. 612-626. ARCHEI{, M., 1976b. Miocenemarsupicarnivores MUIZON,C.de., 1994. A newcamivorousmarsupial (Marsupialia)l'rom ccntral South Australia, fiom the PalaeoceneofBolivia andthe problerlof Ankttarin.ju tirurensis gcn. ct. sp. nov.. Keeunu marsupialmonophyly. Nature 370, 208-21 I . yrtxtdlntrneigcn. ct. sp.nov., and thcir significancc RIDE. W.D.L.. 1964. A reviewof Australianfossil in tenrs ofearly marsupialradiations. Trunsuctiotrs .../orunulofthe Ror:alSot:ietv ol Western ol the Rowl Societ.t,ol SoutltAustroliu 100,53-13. Atrstrulia17.97-129. ARCHER.M.. HAND, S.& C;ODTHELP,H., I995. SPRINGER,M.S., KIRSC]H, .I.A.W, & C]ASE,J.A. Tcrtiary cnvironrrcntaland biotic changc in 1997. Thc chroniclco1'marsupial evolution. 129- Australia.77-99 in Vrba. 8.S.. Denton,G.ll., l6l in Givnish.T.J. & Sytsrna,K.J. (cds), Molecular Partridge,T.C. & Burkle.L.H. (eds),Paleoc'limate Evoltrtionand Atlaptive Rudiation.Cambridgc and EvolLrtion,vrith Emphasisot't Huntan Origins. UniversityPress, Cambridge. YaleUniversity Press, New Haven. SZALAY, F.S., 1994. Evolutionaryhi,ttort' of the BL,NSLt,Y,ts.A., 1901. On the evolutionof the Mar.supialia and un analv.si.sof osteological AustralianMarsupialia with rcmarkson thc c'huructers.Cambridge University Press, New York. rclationshipsof marsupialsin general.Trunscripts 2155n'"r' ol the LinneunSocietv of Lontktn (Zoologyl 9,83- VAN DYCK. S.V, WOINARSKI,J.C.Z, & PRESS, 2n. A.J., 1994. The Kakadu ,Sntinthopsi.t FLOWER,W.H., 1867. On thc dcvclopmcntand blzrfi(Marsupialia: Dasyuridae), a new speciesfiorn successionof teethin the Marsupialia.Philo,sophical the stony woodlandsof thc Northcrn Tcrritory. Transuction,so/ the Royul Sociel) o/ l.ondon 157, Memoirsol the Queenslant{Musettm 37,3ll-323. 63t-641. WROE, S., I 996. Mttribucintrsgarlr_r,a/l (Thylacinidae, GILL, T., 1872. Arrangementof the familiesof Marsupialia),a veryplesiomorphic thylacinid fiom nramnrafswith analyticaltables. Smithsoniun the Miocene of Riversleigh,northwestern Mi,sce/l uneousCo I lecti on 2, I -98. Quccnsland,and thc problen.rofparaphyly for the GODHELP,H.. WROE, S. & ARCHER,M.. I999.A Dasyuridae.Joa rnal of-Puleontologv70, I 032- I 044. new marsupialfrorr the early E,oceneTingamarra WROE, S., 1997a. A reexaminationof proposed local faunaof Murgon,southeastern Queensland: morphology-basedsynapomorphies for the farnilies thc prototypicalAustralian marsupial'/.kturnul of of Dasyuromorphia(Marsupialia): Part I, Muntmulian Evolution 6, 289-313. Dasyuridae..lottrnal of MuntnruliunEvohtlion 1, LUCKETI W. P., 1993.An ontogcncticasscssmcnt t9-52. of dentalhomologies in therianmammals. 182-204 WROE. S., 1997b.Mat,igriphus orbus, a ncw gcnus in Szalay,F..Z., Novacek, M..1. & McKenna,M.C. andspecies of dasyurorlorphian(Marsupialia) fiom (eds),M ammu I Ph.t;krye nt, ; M esozoit: Dil/ere n tiution, the M iocene of Riversleigh,northwestern Multitttberculute,s,Monotremes, Eurly Theriansand, Queensland.Menutirs oJ'the QtreenslundMtrseum M urs upi a ls.Springer-Verlag. New York. 11, 421)-138. MARSFIALL,L. G,,CASE, J.A. & WOODBURNE, WROE.S., 1998.A new genusand spccics of 'bonc- M.O.. 1990. Phylogeneticrelationships of the cracking'daysurid (Marsupialia) from theMiocene familiesof marsupials.433-502irrGeno'"vays, H.H. of Riversleigh,northwestem Queenslancl. A Icheri nga (erl.), Current Mamntahtgl,,I/o1. 2. Plenum Press, 22.271-284. Ncw York. AAPMemoir25 (2001) 59

WROE. S.. 1999a. Killer kangaroosand other WROE,S., EBACH, M.. AHYONC, S.. MUIZON, C. murderousmarsupials. Scientific Amerit'un, Ma1,, dc & MUIRHEAD, J., 2000.Cladistic analysis of 68-71. dasyuromorphian(Marsupialia) phylogeny using WROE, S., 1999b.The gcologicallyoldest dasyurid, cranialand dcntal features. Journal of Mantmulogt from Miocenedeposits of Riverslcigh,northwestcm 81,1008-1024. -521 Qucensland.Pu lueontoktgt' 42, 50 | . WROE, S. & MUSSER,A., 200l. The skull of WROE,.S.. irr prcss. Maxintut:intts mttirheadue, gen. er Nimbacinusdicksoni (Thylacinidae: Marsupialia). sp. nov. (Thylacinidac:Marsupialia), fiorrr the Australian.-/.),rnalof ZooIog1-49. |-29. M iocencof Riverslcigh,northwestcrn Queensland. AtrstruLianJournuI of Zoolog,,-.