Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 O:10.1080/03115510601123601 DOI: (online) Ó 1752-0754 (print)/ISSN 0311-5518 ISSN ae lc kbakus.d.u,Vrert aaotlg aoaoy colo ilgcl at and Earth Biological, of Maradidae, 1.6.2005. revised words: School 17.1.2005, Key received Australia; Laboratory, 2052, Sydney, Wales, Palaeontology South New Vertebrate of University known Sciences, [[email protected]], Environmental the Black Of requires Karen features. Vombatomorphia within apomorphic family crania. Maradidae new and and of dentitions a position plesiomorphic upper to phylogenetic of both the assigned discovery of is of clarification it Further combination dentary, diprotodontids). unique single a a only vombatomorphians, on from based known Although Maradidae, Queensland. northwestern Property, arcanum Marada Queensland. northwestern Riversleigh, B BLACK KAREN Riversleigh, of Queensland Oligocene northwestern late the vombatomorphian from of marsupial family new a Maradidae: otwsenQenln.I xrse a Property, expresses It Heritage Queensland. northwestern World Site, Hiatus Riversleigh Oligocene right late single the a from from dentary has known is morphology herein, scribed dental difficult. relationships on proven solely phylogenetic morphology, based postcranial of and resolution cranial diverse supported by strongly more is infraorder mono- far the of Although phyly a radiation. of pre-Oligocene remnants that repre- the noted families sent (1987) vombatomorphian Archer known & Aplin denti- autapomorphic tions. highly exhibit Miocene, all early and than younger deposits known from families not are two Ilariidae) extinct, and (Wynyardiidae are Of families five Diprotodontidae. these, and Palorchestidae, Vombatidae, Thylacoleonidae, nyardiidae, the Wy- Ilariidae, within Vombatomorphia: recognized infraorder are FAMILIES SIX LACK 07Ascaino utaainPalaeontologists Australasian of Association 2007 . ac 07 aaia:anwfml fvmaoopinmruilfo h aeOioeeof Oligocene late the from marsupial vombatomorphian of family new a Maradidae: 2007. March K., , e.e p o.i ecie rmtelt lgcn itsSt,RvrlihWrdHeritage World Riversleigh Site, Hiatus Oligocene late the from described is nov. sp. et gen. Marada Marada aaaarcanum Marada sms iia opiiiewnadisaddpoootis(aocetd and (palorchestids diprotodontoids and wynyardiids primitive to similar most is e.nv,de- nov., gen. lhrna31 Alcheringa obtmrha obtfre,Mruila Riversleigh. Marsupialia, Vombatiformes, Vombatomorphia, , 73.IS 0311-5518. ISSN 17-32. , osAln&Ace (1987). Archer fol- & (1967). Aplin nomenclature lows Stirton systematic follows Higher-level terminology Mandib- (1867). ular Flower follows Premolar homology (1993). Luckett follows homology follows Mu- nomenclature Rich Cusp Queensland (QMF). the seum of collection logical unl,anwfml,Mrdde a been has Maradidae, established. family, Conse- new a family. quently, vombatomorphian any de- to assigned known and be cannot primitive and features rived of combination unique aiyMRDDEnwfamily new MARADIDAE Family 1987 Archer, Aplin & VOMBATOMORPHIA Wood- Infraorder 1984 VOMBATIFORMES burne, Suborder 1866 Owen, DIPROTODONTIA Order 1811 Illiger, MARSUPIALIA Superorder palaeontology Systematic aeili eoie ntepalaeonto- the in deposited is Material tal. et 17)adAce 18) Molar (1984). Archer and (1978) Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 eeoe upt iga iglmo M on cingulum well- lingual a row cuspate and developed a obliqua; molar ramus); cristid horizontal short well-developed of length teeth; (50% length cheek lower to process; crowned lateral post-alveolar post-alveolar weak less extensive and and less shelf row; behind origin; molar point the of a at point and inflated posteriorly its more originates or ramus at ascending crest-like ramus non ascending symphysis; mandibu- lar extensive com- posteriorly less mediolaterally pressed; gracile, and Dentary elongate narrow, features: of following combination the in families vombatomorphian Diagnosis. ikno,18,56-59 1982, Wilkinson, char- proceeding the anaylsis.) in acter detailed are groups, respective which the the of using members made primitive are below. comparisons listed that are (Note groups differences specific Further more from diagnosis. in familial listed previously the are taxa batomorphian Remarks. is Gender masculine. attachment. shal- muscle medio- the for fossae narrow, and low the dentary to compressed reference laterally word in Riversleigh) flat for of language Aboriginal Marada Etymology. are taxa additional until known. family the for that diagnosis. Generic nov. sp. et om (except forms arcanum Marada species. only and Type Marada 8KRNBLACK KAREN 18 h ln ftehrzna ramus). horizontal the 60 of than plane less the is 1C) ascending the (Fig. the ramus of where border anterior ramus the of angle ascending inclined iorly 94 50 1974, Marada 8 e.nov. gen. ;and itntosfo l te vom- other all from Distinctions aaia iesfo l other all from differs Maradidae iesfo l te vombati- other all from differs aedawakefieldi Warendja Wakaleo h eei igoi is diagnosis generic The steWay telocal (the Waanyi the is 8 nhvn poster- a having in ) lmn Plane, & Clemens 8 rltv to (relative oe& Hope gen. 1 . ntemlr;abte-endparaconid M better-defined on paracristid a and molars; the on nP on cingulum posterior well-developed a lacking otro upda h emnso the of terminus the P on at postprotocristid cuspid posterior aigmr pn -hpdtransverse molars. U-shaped lower the on open, valleys more having iga rsso h rtcndo P on protoconid the of crests lingual n yyrid nhvn nM an having in palorchestids, wynyardiids and diprotodontids, cingulum. from posterior differs posthypo- and posterolingual entoconid, cristid, the a between having pocket except in vombatomorphians wynyardiids, other all from igal oiindpoooi nM transverse more on a protoconid a lophids); positioned the lingually than across wear on (rather of cuspids pattern buccal molars the longitudinal lower along a pattern molars; wear lophodont less ing: Marada M (i.e. molars posterior reduced agrP larger cept rmwnadisi htteP the that in wynyardiids from lrisadwnadisi akn the M Tedford on lacking 1987) 1987a, Woodburne & in (Pledge cuspid wynyardiids neomorph and ilariids n scnieal hre hnM than shorter considerably is and esta 1. than less rsso P on crests 5 rmiaid n akn rcsi,bulbous tricuspid, P a lacking in: ilariids from ium n ipe iupdP bicuspid simple, diastema elongate a an and pa- having and in lorchestids diprotodontids except morphians oohd;abte-endprcndo M convex on paraconid ‘pro- better-defined anteriorly a and tolophid’; ‘hypolophid’ an concave than teeth; rather smaller hav- in ing: diprotodontoids) (i.e. palorchestids rsi,penorsi,adpreprotocristid M (in and preentocristid, cristid, posthypo- postentocristid, postmetacristid, 3 ) eswl-eeoe preprotoconid well-developed less 1); n eldvlpdatro cingulum anterior well-developed a and Marada Marada Marada Rich 3 n akn h otro,bca,and buccal, posterior, the lacking and ; 2-4 Raemeotherium ). iesfo irtdnisi hav- in diprotodontids from differs 3 tal. et eaiet M to relative 3 iesfo irtdnisand diprotodontids from differs iesfo l te vombato- other all from differs eue otro cingulum posterior reduced a ; iesfo aocetd in: palorchestids from differs 98;adawell-developed a and 1978); 3 ;aproportionately a ); 1 . (except Marada 1 3 n distinct a and ; . 1 ALCHERINGA . Marada Marada Marada 3 sntbladed not is Raemeother- iesfrom differs 4 1 /M /M 1 Marada ;andin 1 4 length length differs differs differs 1 (ex- 3 1 . ; Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 egh R,mlrrwlnt;S,smhsslnt.Hthn ersnsboe ra ndnay Bar ramus dentary. horizontal on HRL, areas broken length; represents diastema Hatching DL, length. ramus; symphysis SL, ascending length; of row border molar MRL, anterior length; of angle AAR, of Abbreviations: dentary right of diagram schematic ALCHERINGA i.1 Fig. . aaaarcanum Marada e.e p o.Hltp QMF42738. Holotype nov. sp. et gen. .arcanum M. E OBTMRHA AIY19 FAMILY VOMBATOMORPHIAN NEW hwn esrmnsue nTbe4adtruhu h text. the throughout and 4 Table in used measurements showing A uclve frgtdentary; right of view buccal , B iga view; lingual , ¼ 0mm. 50 C , Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 troaro M478 Bar QMF42738. of stereopair ae ewe h roots. the tooth between respective taken below ramus horizontal of depth oend nlcig neogt,bae P bladed elongate, an lacking: in coleonids ntelwrmlr;adi aigless having in and molars; molars. complex lower the BLACK KAREN on 20 W7480847.2 6.3 26.1 8.4 7.3 10.0 25.9 8.0 10.8 26.3 7.5 10.9 27.4 7.4 7.3 arcanum. Marada 1 11.4 Table 27.4 5.2* 8.7 DR PW AW L 2 Fig. 1-4) arcanum Marada premolar. ep one ypyi;ahgl inclined I highly a caniniform a fossa; symphysis; rounded masseteric deep, deep well-developed, a . aaaarcanum Marada esrmns(m fdniinof dentition of (mm) Measurements . P 3 1 n nhvn igelower single a having in and ; Mxmmwdho P of width *Maximum Marada M p o.(is12 Tables 1-2, (Figs nov. sp. 1 e.e p o.occlusal nov. sp. et gen. ¼ M iesfo thyla- from differs 0mm. 10 2 M 3 3 DR, . M 3 4 ; P Holotype. 46m.Tedatm srltvl long relatively is diastema The mm. 14.6 etr Fg 1). (Fig. Dentary Description known de- single specimen. the and unknown by exhibited primitive the features the rived and of species to combination the strange alludes of position Latin, taxonomic in tery Arcanum etymology. Specific additional known. until are family taxa the for that is species diagnosis. Species age. is Oligocene faunas, late of contained be to and interpreted stratigraphy basis the on of which, 1997) (Creaser deposit A age. and Distribution 1997). Creaser (see northwes- Queensland tern Property, Fossil Heritage World locality. Type coro- angular process. and the condyle, missing articular process, and noid damaged is dentary poieM opposite I au s2. mblwteatro base horizontal anterior the M the below mm of of border. 27.9 ventral is depth ramus the maximum is as The is straight, ramus and horizontal flat the of surface (medial) yatkolai ?diprotodontid delicate relatively the primitive of characteristically dentaries exception horizontal the the vombatomorphian with narrow unlike robust a is more It with ramus. slender gracile very is and dentary The process. and angular condyle, articular missing process, and coronoid preserved the poorly is dentary the M 1 3 ,C ,M 1-4 .I 0 ,P 1-4 1 1 h oiotlrmsthickens ramus horizontal The . rw rkn h otro raof area posterior The broken. crown .I 3 ,M Rich 1 M478 ih etr with dentary right QMF42738, rw rkn etlformula Dental broken. crown 3 1-4 ihamxmmwdhof width maximum a with h otro rao the of area posterior The . tal. et itsSt,Riversleigh Site, Hiatus ih etr ihP with dentary Right h igoi o the for diagnosis The itsSt saSystem a is Site Hiatus 98 h lingual The 1978. ALCHERINGA Raemeotherium enn mys- meaning , 3 , Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 1 astrcfrmnAsn Present deep Elongate, Absent Short ramus horizontal to relative length Diastema 12. foramen Masseteric symphysis 11. of extent Posterior 10. HRCE LSOOPI APOMORPHIC PLESIOMORPHIC CHARACTER ALCHERINGA h isea mle uretforamen of nutrient surface smaller dorsal A the diastema. to the ventral mm 3 and below P directly of root point anterior a the termi- at symphysis posteriorly shallow. nates The dipping to diastema, posteriorly. parallel the and slightly of angle surface an narrow dorsal 12 at the sub- lies is symphysis The The 1B). fossa 11 Fig. sym- unfused long, lingual short mm (50.3 rough, very physis 10 a to steeply drops away diastema The the of diprotodontoids. surface diastemal 9 lingual of dorsal reminiscent distinct a ridge with mm) 8 (38.7 7 6 3 Table Ngapakaldia 5 Raemeotherium Propalorchestes 4 Namilamadeta Muramura 3 Warendja Kuterintja Wakaleo 2 Madakoala Trichosurus 1 Marada CHARACTER state e is1. matro otebs fP of base the to anterior mm 11.6 lies men .P 2. .Peec facitdolqaWl eeoe eko absent or Weak developed Well M gradient molar increasing obliqua Posteriorly cristid 7. a of Presence 6. .Conhih o-rwe High-crowned Low-crowned ( Low ramus ascending of border anterior of Angle 9. height Crown 8. .Dge fblpoot betFlybilophodont Fully Absent absent or Weak developed Well M on protostylid a of paracristid Development and 5. paraconid of Development 4. bilophodonty of Degree 3. .Iciainageo I of angle Inclination 1. eaieylrecnllk etlfora- mental canal-like large relatively A ¼ 3 opooyBcsi Multi-cusped/bladed Bicuspid morphology ;ms pmrhcstate apomorphic most 1; hrce tt oaiisfrslc obtmrhas lsoopi state Plesiomorphic vombatomorphians. select for polarities state Character . al 2 Table 102212121 111 1?2201120 1?0 102211121 111 111111110 110 111111110,111010*11*11*020110210021 1?? 010?12010 010 010000001 1?? 011000000 000 101210010 011 120 hrce ttsue ncaatraayi fVombatomorphia. of analysis character in used states Character . 3 . 1 ¼ ;msigdata missing 2; E OBTMRHA AIY21 FAMILY VOMBATOMORPHIAN NEW 1 4 /M 1 3 ¼ ;caatrplrsdusing polarised character ?; eri vdn ln h ot o mainly row tooth the along evident is of line wear longitudinal Vomba- A other 4). (Table any tomorphian the than of ramus) length the horizontal to relative (50% shorter row relatively molar a has specimen Proportio- this dentary. with nately, the of compared length overall row the molar short relatively iswti h nrnet h mental M the foramina ven- to the mm tral nutrient 15.5 to approximately positioned small are entrance Three the foramen. within lies otro oa rdeti vdn (M evident is decreasing gradient slightly molar A mm. posterior 51.8 is row tooth length rsn Absent 5 Present ( High 5 h egho h lgtycre cheek curved slightly the of length The 1 30% ¼ 5 .) tiigfauei the is feature striking A 0.9). 30 70 8 8 o ( Low ) )High( 1-2 alveoli. .ursinus V. 1 30% ¼ ;apomorphic 0; 5 70 30 ¼ 8 8 ) ) *. 4 /M 1 Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 isealnt maue rmP from (measured length diastema 4 Table novaculacephalus Propalorchestes tedfordi Ngapakaldia yatkolai Raemeotherium Namilamadeta pinpensis Muramura ngama Kuterintja aae oldfieldi Wakaleo wakefieldi Warendja arcanum Marada egh(esrdfo nio lelst otavoa rcs) R,mlrrwlnt (measured length row molar MRL, process); post-alveolar P to of alveolus anterior from incisor from (measured length ntebca up ihtehpcndof hypoconid the with M cusps buccal the on CHARACTERS DENTARY BLACK KAREN 22 rsn ntevnrlbre fthe M poster- of of the below border mm mm ior 27.1 border 12.0 and approximately to ventral is posterior the fossa the foramen of masseteric masseteric on face out small present lateral A fading the dentary. border, onto anterior anteriorly distinct with shallow and the no flat is fossa of masseteric determined. The be position cannot speci- foramen the mandibular the Consequently, added of for men. preparation paper rice during pterygoid in strength the covered of was face. the area fossa lateral fragile the of the on Medially, area preserved mas- fossa the the seteric by encompassed only ramus ascending with incomplete wear. most on poietepseirbre fM of border at posterior the originates opposite point ramus a ascending The ramus. ol,isatro odrpstoe tan 55 at positioned of border angle anterior its iorly, the dentary. the of been on have low flatness would and the weak it suggests but ramus preserved, vertical not is sn itntps-lelrpoeso the on process post-alveolar distinct lateral no There less vombatomorphians. is other and all in posterior than further is which 1 h otro rao h etr is dentary the of area posterior The h sedn au sicie poster- inclined is ramus ascending The n h rtcndo M of protoconid the and esrmnsadpootoso h etr sdi hrce nlss brvain:DL, Abbreviations: analysis. character in used dentary the of proportions and Measurements . 8 p 404. 2406 .21.0 0.22 0.67 62.4 41.8 14.0 sp. eaiet h horizontal the to relative 4 3 h astrceminence masseteric The . opseiro M of posterior to 2 3 hwn the showing (mm) 036. 1. .703 1.1 1.0 1.0 0.36 0.16 0.24 0.57 0.64 112.2 0.71 82.0 64.0 52.7 51.0 40.3 13.0 36.3 12.0 683. 7805 .908–0.9 – 0.8 0.9 0.29 0.38 0.56 0.51 57.8 101.0 32.5 51.8 16.8 38.7 lelst I to alveolus DL . 695. .001 n M (no 0 0.11 0.70 52.7 36.9 6.0 95–––1.0 0.9 – – – – – 69.5 – – 44.2 – 4 ae tbs fcrown). of base at taken , MRL (mm) 4 , 1 lelso oslsrae;HL oiotlramus horizontal HRL, surface); dorsal on alveolus m)MLHLD/R M DL/HRL MRL/HRL (mm) HRL etto Fg ,Tbe1). I Table is 2, formula (Fig. Dentition shelf. post-alveolar the below posteriorly ftehrzna au.I sdeetbelow deepest is It M ramus. horizontal border ventral the the the of to of parallel linear runs shallow, flaking that a fossa is the fossa digastric by The bone. slightly obscured area this is However, expansive. laterally very non-carinate. and of ascending thick point the is of its ramus ‘crest’ determine anterior to The rise difficult origin. gradual is the a it rise such of that is to margin It seen shelf. buccal post-alveolar be the can along gradually ramus view, ascending occlusal In the dentary. the of face lingual n a aels M lost have premolars may lower two and retain thylaco- which leonids, some (except vombatomorphians acoae nmli itiue nthe on I The distributed surface. lateral and is ventral was it Enamel that the lanceolate. but suggests preserved, cross-section is incisor elliptical the of mm 16.2 ee fteM the of level o t neoiga ae,sub-central face), anterolingual its worn very (on large, A two-rooted. and bicuspid h diastema. the 28 of angle an 3-4 h trgi os svr a n not and flat very is fossa pterygoid The h rmlri ipe ml,narrow, small, simple, is premolar The n xed neiryt eo the below to anteriorly extends and 1 8 rmtehrzna ln of plane horizontal the from C 1 0 aoi n ae out fades and talonid P 3 M 4 .Ol h distal the Only ). 1-4 ALCHERINGA iealother all like 4 /M h dental The 1 1 rjcsat projects LENGTH 4 ) Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 -hpdtases aly.TeM The wide, valleys. very transverse by U-shaped characterized and phodont, main the slightly of is base enamel posterior cuspid. the The at the crown. is wrinkled to the sloping lingual of steeply cingulum are base and tooth the the buccal buccal, of faces anterior, and opposite A The large point absent. cuspids. the a between small at valley crown base transverse the the into fading poster- of cuspid, small weak anterolingually the from extends A cuspid cingulum crest. olingual posterior steeply posterior weak, small a sloping by metaconid) a the (possibly is to and premolar connected the dominates protoconid otrlnult h otro updof margin, cuspid tooth posterior anterior the P to the posterolingual slight at a in the swelling terminates from and apex anterolingually protoconid extends but para- cristid) well-developed (or preprotocristid A worn is considerably rounded. protoconid and the of large dipro- less base and The though wynyardiids todontoids). protolophid, in than a developed as tentatively to is a (which referred by ridge joined transverse been short have protoconid the may the In of metaconid protostylid. and apices the no state, is unworn There of tooth. midline horizontal the the to have buccal to just appears been apex the that such lingually ALCHERINGA ooi n yooi xii h most the exhibit hypoconid wear. and pro- meta- The the toconid hypoconid. by and the followed entoconid, to cusp state, conid, tallest that unworn due the its appears been in cuspid have It may each protoconid wear. the of determine considerable to height difficult large is relative a It in talonid. hypoconid and the buccal trigonid and entoconid the lingual buccal on least large and (the protoconid cuspids) metaconid major the lingual of small worn a of consist- ing tooth four-cusped sub-rectangular, 3 hsseln a ersn weak a represent may swelling This . h oasaelwcond bunolo- low-crowned, are molars The h rtcndo M of protoconid The 1 spositioned is E OBTMRHA AIY23 FAMILY VOMBATOMORPHIAN NEW 1 sa is u ttelnulbre ftetransverse the of border valley. lingual cingu- extends the lingual at the lum postmetacristid meet the to weak of posterolingually base A the the to crown. of steeply faces ante- slope The metaconid posterolingual apex. and protoconid rolingual the to lingual h uclfc fteprcndslopes crown. paraconid the of the base the of to face steeply buccal paraconid. The the from buccally runs anterior cingulum short, weak, similarly A paraconid. lih lhuhntt h xetse in seen extent River- the B, to Marada not System (Dirk’s although Fauna, specimens sleigh) 1979, Local Archer, Towers preen- some & the in on Rich evident enamel is the tocristid of thickening and cusp diprotodontids, a similar in A expressed koalas). variably in is postmeta- does the it of (as swelling does cristid a it of as originate because metastylid not doubtful the with is homology cusp phascolarctids well-developed lingual The this a cingulum. of by lingual lingually cuspate off is It the talonid. closed posterior separates and trigonid and anterior U-shaped, and shallow, h rw.Tehplpi slne than longer is protolophid. hypolophid the The of base crown. the the to steeply slopes the base entoconid a the lingual of of The be apices hypoconid. to and the entoconid unlikely linking unworn crest is its continuous hypolophid In the cusps. state, these between longitudinal the the valley as traverses to that referred hypolophid) (tentatively a the in lophid at resulting weak hypoconid, swelling the of similar base lingual a meets entoconid cuspate. state, been have unworn may its obliqua cristid In the buccally. trans- valley the and verse off protoconid sealing the effectively of hypoconid, apices the connects preentocristid. weak the to poster- iorly and postmetacristid the to anteriorly h rnvrevle fM of valley transverse The mm 2.0 lies metaconid the of apex The ekseln ttebca aeo the of base buccal the at swelling weak A obliqua cristid worn well-developed, A h iga iglmconnects cingulum lingual The . Namilamadeta 1 swide, is Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 oaso yyrid uhas lower such deta the wynyardiids in of found also molars is feature This cingu- lum. posterior the and entoconid, posthypocristid the the of base posterior the the formed tooth. between is the pocket runs of postero-lingual resultant border A that posterior the cingulum of length a the posterior with in continuous weak bulge are posthypocristid postentocristid or weak and A swelling point. this slight at enamel a is there hplpi’aesmlri length. in similar are ‘hypolophid’ h rsi biu swl developed well M is in than so obliqua less lingually. although effectively valley cristid transverse preentocristid, The the off weak sealing and postmetacris- weak tid the crescen- short connecting a ridge reduced, is tic cingulum is lingual cuspid the lingual and The worn. more iea nM in lophid- not as is and like bases occurs their be- down hypoconid further and connection entoconid The the tween cingulum. of buccal border lingual the the and of obliqua base cristid buccal crenulated the the small between formed A is enamel. pocket the of swelling hni M buccally trigonid. in more than the positioned on is protoconid cusps The the of arrangement ieta nM in than ridge- more like is cingulum buccal The cuspate. mrsino ekpooohd The protolophid. M in weak than that larger the is a metaconid gives so of tooth the connected impression across are wear transverse protoconid and 4KRNBLACK KAREN 24 on n h nooi suwr.In unworn. the is entoconid by slightly the followed only and is cuspids worn, most metaconid the the the hypoconid; is of protoconid worn the the than longer hypolophid; being developed, protolophid protolo- noticeably better the the are with is hypolophid trigonid; and talonid the phid is than the cingulum narrower lingual and the reduced, following: the for h M The however, absent; is cingulum buccal A h eswr M worn less The . 1 2 u h ae ftemetaconid the of bases the but , ssmlrt M to similar is 1 h pooohd and ‘protolophid’ The . 1 hr tcntttsa constitutes it where 3 ssmlrt M to similar is 1 1 n ti non- is it and , xetfrthe for except 1 n t apex its and , Namilama- 2 except Tyaoende.Seisof Species (Thylacoleonidae). and lorchestes eetcasfiain eg pi Archer Marshall & Aplin 1987, (e.g. classifications Recent analysis Character hntepoooi,adteetcndis entoconid hypoconid). the the than anterior and anterior more protoconid, more the is metaconid than the buccal (i.e. opposing ante-cuspid transversely more its positioned than riorly is wherein cuspid phascolarctids lingual and the taxa ilariids semi-lophodont as the such other in more of is feature reminiscent This posteriorly molar. the of curve half buccal hypolophid and protolophid entoconid, the Consequently, and and respectively. metaconid the protoconid than pos- teriorly more the slightly positioned deeper; are the hypoconid is between posthy- pocristid and cingulum, pocket posterior entoconid, posterolingual the protolophid; the the the worn; than only shorter are is cuspids, hypolophid hypoconid major and the protoconid of less of cristid and result a wear, the as prominent margin; more is obliqua tooth posterior linear olwn:M following: n nooi r eaieytl n well and peaks). sharp tall metaconid no (i.e. relatively the rounded are of entoconid apices and the view, lingual imrhcfrterrsetv families: wakefieldi Warendja respective ngama their ple- Kuterintja considered for are siomorphic material, dentary able Suborder the a of as outgroup Vombatiformes. used distant was more Phalangeridae) todontia, plesio- vulpecula Trichosurus the using Woodburne phascolarctid determined morphic were of ties polari- group character sister Hence, the Vombatomorphia. are phascolarctids that h M The h olwn aa hc aecompar- have which taxa, following The aae oldfieldi Wakaleo 4 ury 96(Palorchestidae); 1986 Murray, ssmlrt M to similar is tal et 4 mle iharudd less rounded, a with smaller tal. et 97,a noutgroup. an as 1987c, . lde 97 (Ilariidae); 1987b Pledge, lmn ln,1974 Plane, & Clemens (Vombatidae); Kr,19)(Dipro- 1792) (Kerr, 99 acknowledged 1989) aaol devisi Madakoala ALCHERINGA 3 xetfrthe for except Muramura Propa- Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 Muramura hrceso h etto antbe cannot the dentition of Three the for polarized taxon. of are this dentitions characters lower for as unknown analysis, of the Species from considered. also Koobor 1990, is 1997) Murray Black see diprotodontid: tionable tedfordi but group, kaldia the of most member the primitive possibly is and analysis. Diprotodontidae to cladistic yatkolai be a meotherium to by yet their determined have because relationships considered phylogenetic both are diidae) .P 2. Marada lorchestes .Iciainageo I of angle Inclination 1. 2 Tables in respectively. listed 3, are and polarities char- and state significantly states acter Character a analysis. in be biased resulting here, cannot employed phylogeny delineate char- to vombatomorphian analyses postcranial previous in and missing used acters of cranial level many high data: the to due performed not was analysis parsimony un- puter-generated are for postcranium known Only and cranium, the 3. dentition, as denti- analysis, upper Table the lower in in used and are dentary tion 6 the and of characters 4, polarize 2, to used characters been have specimens 1800) non juvenile Consequently, molars. unworn, the on wear extreme and 1987a Pledge, sdb Marshall by used nlnto nl fI of angle inclination aocetd oss ipebcsi P bicuspid simple a possess palorchestids is ihiciainage( angle inclination thylacoleo- high and A ilariids of nids. exclusion the to clade vombatid/diprotodontoid wynyardiid/ osdrdpiiieadi on in surus found is and primitive considered ALCHERINGA , 3 Madakoala , rhr&Wd,17,aeexcluded are 1976, Wade, & Archer morphology. , Kuterintja and , Namilamadeta Marada aedawakefieldi Warendja tro,16 a unques- (an 1967 Stirton, Warendja and , obtsursinus Vombatus Namilamadeta osqety com- a Consequently, . , stnaieyassigned tentatively is tal. et 1 Ngapakaldia irtdnisand Diprotodontids ( 1 5 synapomorphy A . , Wakaleo . 30 Raemeotherium 8 speetin present is ) 18)fra for (1989) (Wynyar- low A . E OBTMRHA AIY25 FAMILY VOMBATOMORPHIAN NEW , Ngapa- Tricho- 30 u to due Propa- (Shaw 8 Rae- )is 3 , rmlrmrhlg a hne little P changed plesiomorphic the has from morphology This postprotocristid. premolar a by of joined apices are the which small (metaconid), a and cuspid protoconid posterior central large a with ut-upd lddP bladed multi-cusped, with polarized been rmlr fbt ugop r highly are outgroups both specialized. The of condition. premolars bicuspid plesiomorphic the h otro hr ftebae The blade. the of third P bicuspid the along posterior lingually above curving the positioned then apex root, anterior an from cally aueo the of nature Raemeotherium ietedpoootis a ipeP simple a has diprotodontoids, the like dasyuromorphians. and phians eutn nabae pernet h P crest the longitudinal to appearance a Kuterintja bladed a by in linked resulting medial are These cusps protoconid, hypoconid. posterior longitudinal and anterior the metaconid, along an lie midline: cusps three but xii eiblpootmolars.arcanum semi-bilophodont exhibit Namilamadeta Muramura, molars. bilophodont ngama plesiomorphic Vombatomorphia. considered for is bilophodonty of Absence molars). unworn (in Vombatidae and Diprotodontidae, Palorchestidae, Wynyar- diidae, extent), Macropodidae, lesser a (to including Phalangeroidea lineages diproto- dontian several in independently developed bilophodonty. of Degree 3. condition. primitive the ered i mtcnd,adawell-developed a P A and cuspid. posterolingual (metaconid), cus- pid posterior well-developed a protoconid, int otrbca upadamuch a are and cusp cusp. cusps lingual posterobuccal addi- weaker a 4 in to crest premolar: tion longitudinal a along 6-cusped situated a has lddP bladed ,and 3 osse eiblpootmolars semi-bilophodont possesses h ld xedn posterobuc- extending blade the , 3 a rcsi P tricuspid a has of rcouu vulpecula Trichosurus .oldfieldi W. Warendja u oteeteeyworn extremely the to Due . aaaarcanum Marada .ursinus, V. aae oldfieldi Wakaleo rcouu vulpecula Trichosurus aaol devisi Madakoala 3 . P 3 3 aaol devisi Madakoala iohdnyhas Bilophodonty hsfauehas feature this , ,and 3 fperamelamor- of ontpossess not do sukonfor unknown is 3 ihalarge a with hc shows which Muramura sconsid- is Warendja Marada a a has a a has , K. 3 3 , . , , Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 .Dvlpeto h aaoi n para- and ( paraconid the cristid of Development 4. phodonty .Dvlpeto rtsyi nM on protostylid a of Development 5. omn ekpooohdadhypo- and protolophid weak lophid. respectively, a hypoconid the forming and linking entoconid and protoconid, the crests and metaconid the transverse of apices weak with oas Selenodonty/bunodonty molars. Propalorchestes osbyiaid Tdod&Woodburne & (Tedford ilariids possibly (Wood- burne miralinids petaurids, pilkipildrids, (Woodburne docheirids pseu- 1997), (Cooke macropodoids morphic betprcndadwa rabsent or weak and paracristid paraconid devel- absent well paracristid oped) and para- (i.e. present trigonid conid triangular ‘un- is: a coding of designated Character presence taxon. is this for character known’ trigonid. this the on cuspid Hence, main is the as for uncertain, case is the however, ridge this a paracristid; of homology occupies a the to that position ridge similar (the anterior cusp an trigonid this From extends the 1956). Woods on of metaconid cusp apical anterior rtsyi nM on protostylid 6KRNBLACK KAREN 26 ec hi rsnei osdrdplesio- considered is including morphic. presence taxa their hence marsupial peramelemorphians; devel- most and dasyuromorphians well in are oped paracristid and paraconid and Propalorchestes paraconid. Raemeotherium, weak Kuterintja, a in culminating state. primitive Namilamadeta the sess h M The ursinus antb oaie for polarized be character cannot This paraconids. and paracristids Ngapakaldia ¼ 1 tal et ;prcitdpeetbtlow but present paracristid 0; hw h pmrhccondition. apomorphic the shows Raemeotherium of ¼ ¼ Madakoala ¼ ;bilophodonty 1; aae oldfieldi Wakaleo rpoorsi)o M on preprotocristid) 97) hsoacisand phascolarctids 1987a), . 2. oss itntparacristid distinct a possess oss ul bilophodont fully possess oss eko absent or weak possess 1 speeti plesio- in present is and , Ngapakaldia Trichosurus ¼ tal et Muramura Warendja 2. a simple a has ¼ 1987b), . ;semilo- 0; Marada, 1 The . ,and ; 1 pos- ¼ and .A V. 1; , 18)itrrtdtepeec faproto- a M of on presence stylid Marshall the and interpreted (1989) (1978) Archer 1987). antb oaie for polarized ursinus be in absent cannot is obliqua Ngapakaldia cristid A protoconid. in ium present is obliqua cristid ted direc- anterolingually moderate A ‘midlink’. rhr18) h bec faprotostylid a of in absence The the 1987). Archer in present idi is 1997), protostylid (Cooke phalangerids plesiomorphic small ridge a a to but reduced is stylid rhr19) In 1997). Archer 1997, al. Archer, et & of Black plesiomorphic species as such the forms derived in weak genus protostylid from the ranges phascolarctids, Alter- group. in the natively, of in members absent derived or more reduced is the but plesiomorphic macropodoids is that in protostylid suggested a of (1997) formation Cooke dontia. biu swl eeoe ihnmost within Kuterintja developed vulpecula, In groups. well obliqua. diprotodontian cristid is a obliqua of Presence 6. (1997). Archer follow- & protostylid Myers ing a lacking as interpreted derived. as interpreted sibly pos- (except diprotodontoids and coleonids, uclt ro h ogtdnlmdieof midline longitudinal M the the on or just to protolophid the buccal the on meets position and lingual origi- hypolophid more directed, a from anteriorly nating more is obliqua In buccally. valley Muramura transverse the off effectively sealing postprotocristid, hypoconid the meet the to apex from anterolingually tends Marada and u emntsa h aeo the of base the at terminates but Raemeotherium 1 Madakoala 97 and 1967, ec h eiaino h term the of derivation the hence , a oeaeydvlpdcristid developed moderately a has rcouu dicksoni Trichosurus obtd,wnadis thyla- wynyardiids, vombatids, , and 1 speimrhcfrDiproto- for plesiomorphic as and Propalorchestes Wakaleo .vulpecula T. Phascolarctos ovr ag nmore in large very to and , [Rich ueitangama Kuterintja Litokoala hscharacter This . ALCHERINGA Marada tioucsre- Strigocuscus Madakoala tal. et Warendja Fanr & (Flannery h proto- the , Raemeother- h cristid the , Nimiokoala Bak& (Black cristid A 98)is 1978]) tex- it , Stirton tal. et ; , V. T. is Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 euigsm ftesrs lcdo the M on 1977). placed (Beecher stress symphysis the effectively mandibular of increases some joint, molars reducing towards temporomandibular posteriorly posterior the force the masticatory the the in of increase An the size diet. in the increase of an abrasiveness with is associated It possibly diprotodontoids. the is particular in vombatomorphian taxa, gradient derived several molar in found increasing gradient. molar posteriorly increasing Posteriorly 7. obliqua odto iha M an with condition and ietdmidlink directed irtdnin n r consequently Both Vombatomor- phia. are within among plesiomorphic common coded and are diprotodontians molars selenodont height. Crown 8. primitive. oldfieldi surus aea M an have etdt postprotocristid to well- nected con- obliqua coded: cristid are anterolingual developed states Character obliqua. ooe nM An one. to ALCHERINGA cdd1,weesM whereas 1), (coded nlnto fteacnigrmsfor of ramus angle ascending the the the listed of of 116) border p. inclination anterior (1982, inson the ramus of ascending Angle 9. nature. vestigial their of consequence a as et (P teeth In high-crowned 2). are diprotodontoids, (coded ilariids and and 1) vombatoids 0). (coded (coded crowned molars Marada low-crowned possess M therium one. 1 aisaelse nTbe4. Table in listed are ratios L , Marada aiaaea Muramura Namilamadeta, , Madakoala ec hscaatri oe as coded is character this hence ; ¼ 3 Ngapakaldia n yyrid r moderately are wynyardiids and n M and 2. 4 /M 4 Trichosurus 1 scmltl otin lost completely is ai rae hno equal or than greater ratio L .oldfieldi W. xii h plesiomorphic the exhibit ¼ 1 r oeaeycrowned moderately are ) Fg C.Hp Wilk- & Hope 1C). (Fig. o-rwe bunodont/ Low-crowned , ;Wa rasn cristid absent or Weak 1; Kuterintja ,and 4 2-3 /M r oe crowned lower are 1 ai esthan less ratio L and h functional the , ¼ Propalorchestes ;anteriorly 0; , Madakoala , Warendja E OBTMRHA AIY27 FAMILY VOMBATOMORPHIAN NEW Wakaleo Raemeo- Tricho- A 4 / , oiotlrmslength ramus horizontal proportionately length Diastema 12. for plesiomorphic foramen foramen be masseteric to Vombatomorphia determined koala xedn oblwteatro otof root anterior the below M to extending oblwtepseirro fM of root posterior the below in to evident is sis nl slwin low upright is more ( angle a ramus and ascending plesiomorphic, ered ( inclined 70 angles than have less diprotodontians most and perame- lids, Dasyurids, taxa. marsupial various aeysotrin shorter nately otro oto P the of below root to posterior posteriorly further extends but mura Marada in absent is men n ag in large and foramen. Masseteric 11. Vom- for batomorphia. apomorphic deemed is symphysis for meotherium in Kuterintja elongate posteriorly more vulpecula ndja Marada (70 xed otrol oteatro odrof border P anterior the to posteriorly extends 80 ce ttsaedslydb pce of species by displayed are Muramura states acter 0 otro xet(lnain fsymphy- sis. of (elongation) extent Posterior 10. 3 1 8 h otpseiryetniesymphy- extensive posteriorly most The . 8 Fg C.Tesmhssi proportio- is symphysis The 1C). (Fig. ,and ), Raemeotherium ), h adblrsmhssin symphysis mandibular The , bec famseei oae is foramen masseteric a of Absence . Ngapakaldia (56-59 Propalorchestes , ¼ (55 5 (65 , Wakaleo Propalorchestes 2. (65-70 aiaaea uaua Rae- Muramura, Namilamadeta, , 70 8 8 8 8 ), .I surgtin upright is It ). osqety posteriorly a Consequently, . ), 8 Warendja sedn au sconsid- is ramus ascending ) Raemeotherium Warendja Namilamadeta aae oldfieldi Wakaleo Propalorchestes 8 Trichosurus Trichosurus ¼ 3 ). , h ypyi becomes symphysis The . (70 , ;salo moderate or small 0; ¼ Kuterintja Namilamadeta 70 8 Dasyuromorphians, . and , ,paclrtd (70- phascolarctids ), ;lremasseteric large 1; and , 8 sdrvd The derived. is ) astrcfora- masseteric A ti unknown is It . (70 ti ml in small is It . 8 and .Bt char- Both ). Ngapakaldia Ngapakaldia and , (62 Madakoala extending , (65 Kuterintja 1 Wakaleo 4 8 long A . (50 , ), 8 Marada ,and ), 0 of 30% Mada- Mura- Ware- 8 ), T. , , , , Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 8KRNBLACK KAREN 28 18)ntd h ihyautapomorphic highly the Archer & noted, Aplin as (1987) However, 1989). burne I between diastema a diprotodon- lack tians most and peramelemorphians rhr18,Tdod&Wobre1987, & Woodburne Marshall & Aplin Tedford 1986, 1987, Archer Murray (e.g. phascolarc- tomorphians of vomba- exclusion the uniting tympanic to synapomorphy tomorphians the a as the to cavity recognize contribution studies squamosal and phylogenetic cranial recent most both particular, In morphology. by postcranial supported strongly Vombatomorphia. addition families infraorder seven the the now within With are there years. deposits Maradidae, 30 of Riversleigh past material the the fossiliferous over from of tonnes processed de- the Maradidae new spite the family Riv- of occurrence Site, first 2001 vombatomorphian the Hiatus is in Oligocene It ersleigh. recovered late dentary the from single a from arcanum Marada Discussion P and ol mi egh u otepresence the to due length) P of in mm 6 (only h rsneo C of presence the surus for polarized Kuterintja be in cannot length ramus lamadeta Nami- horizontal Raemeotherium, Wakaleo, the Warendja, of 30% ramus is Ngapakaldia horizontal length in the diastema of length The 30% than 4. greater Table in are study listed current the in horizontal taxa for and length ramus derived. diastema deemed of is Measurements ramus horizontal the length the of to respect elonga- with diastema therefore the of and tion plesiomorphic, be regarded is to diastema functional a of absence h oohl fVmaoopi is Vombatomorphia of monophyly The 2 . 1-2 u otepeec fvsiilI vestigial of presence the to due n emdasn for absent deemed and , ,and iseai betin absent is diastema A . tal et Marada h iseai esthan less is diastema The . 99 pigr&Wood- & Springer 1989, . Muramura e.e p o.i known is nov. sp. et gen. 1 , n P and Propalorchestes hscharacter This . Madakoala 1-2 1 ec,the Hence, . n P and Wakaleo Tricho- 3 and , u to due and 2 titdmniua oae;ad4, and foramen; uncon- pterygoid 3, mandibular ramus; long horizontal stricted the 2, flaring of widely flange crest; 1, include: the submasseteric Woodburne of These & synapomorphies dentary. (Tedford recognized one 1987), only 1998), aoopi aeas enquestioned. been also Murray Vom- have with thylacoleonids batomorphia of affinities macropodoid. The primitive a or diprotodontid Similarly, meotherium Vombatomorphia. Phascolarctomorphia the and both to assigned hlgn eg rhr18,Aln& Aplin 1987, Woodburne 1984, Marshall & Tedford Archer 1987, Archer (e.g. vombatomorphian the of phylogeny Of analyses to data. previous postcranial due all and cranial difficult of proven absence has morphia to respect group. and the with of plesiomorphic position rest the distant highly a are recognized occupy they but that than Vombatomorphia lions & marsupial phalangerids within the included Aplin to (1987) Archer closer However, be vombatomorphians. to nids is h ngai genus enigmatic palorches- The with tids. sug- specifically (1990) affinities gested Murray and diprotodontoids, nw o h yespecies those type to the similar bassiana presumed for were a limbs known the on material limb wherein based with dentitions of tentative, association is dae obtmrhaso h ai fits between Archer Namilamadeta affinities of greater & recognized other Aplin basis (1987) all morphology. the basicranial to on group vombatomorphians sister plesiomorphic concluded (1987) aytx antb sinda the alone. both at of dentitions lamadeta placement assigned on the example, based For be level cannot familial taxa analyses. families Many phylogenetic in use vombatomorphian limited provide of dentitions lcmn of Placement tal. et tal. et and (Tedford ol eete plesiomorphic a either be could Muramura 99 usn19,Murray 1992, Munson 1989, 18)rgre thylacoleo- regarded (1987) and Marada Wynyardia tal et Muramura ihnWynyardii- within ihnVombato- within Koobor 97.Aplin 1977). . ALCHERINGA ob the be to Wynyardia ihthe with a been has Nami- Rae- Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 atclr h osrcino h M the of in construction the the and of particular, preentocristid, development and the postmetacristid in molars, pocket lower posterolingual respect the the with other, cuspids The each major to 3). the the of (Table molars, position five each semi-lophodont with sharing low-crowned states Diprotodontidae, derived and Palorchesti- dae, Wynyardiidae, primitive the to of members similar most morphologically aey oeo hs etrsaepreserved Unfortu- are the features on these condyles. of none nately, mandibular dorsally and curved elongate transversely narrow, biu;adteasneo rtsyi on protostylid cristid a M reduced of absence a the molars; and obliqua; lower well-developed the in a metas- tylid and of postmetacristid, absence lophodont premetacristid, more the of its presence molars; the on: size; based large Phascolarctomorphia) (rather than Vombatomorphia the within placed aaoi,adprcitdo M on paracristid weak a and and size, large paraconid, diprotodontids its include the palorchestids with shared exclusively Features molars. lophodont towards ALCHERINGA a ig,adalremna foramen. mental large a and diaste- ridge, prominent mal a with diastema elongate tiigysmlrto similar strikingly nl ficiaino I low of a inclination include of palorchestids angle and diproto- wynyardiids, dontids, with shared Derived features palorchestids. and diprotodontids u otepeec fapostmetacristid, a preen- of and Diprotodontoidea presence postentocristid, posthypocristid, the of to due member primitive h ipepeimrhcbcsi P of bicuspid plesiomorphic premolar simple bicuspid the the Marada ‘neomorph’ and a of the cuspid, absence on the valleys molars, lower transverse U-shaped a more open, of the lack cingulum, the anterior are in well-developed taxa these differences of main dentitions the The Riversleigh. from 1 . ihnVombatomorphia, Within aaaarcanum Marada Marada Marada h atfauei otsmlrto similar most is feature last The . spelddfo en a being from precluded is dentary. Namilamadeta s oee,tentatively however, is, 1 n tendency a and Marada E OBTMRHA AIY29 FAMILY VOMBATOMORPHIAN NEW material 1 1 ,an are , 3 of is h eret hc hs etrsare features these which to degree fossae). The digastric and pterygoid, shallow the masseteric, pronounced (i.e. attachment of muscle for absence areas and dentary linear no length. or row little molar in with increase diastema by the achieved was lengthening this but diprotodontoids larger, the becoming to in path similar a along egho h oiotlrms twould It the the ramus. of that horizontal 50% appear the only of occupies length row which molar short length, the is vombatomorphians Marada diprotodontoids. lophodont lophs fully transverse ilariids, the the in seen across in wear even found the to opposed as that phascolarctids and of wynyardiids, reminiscent is otro rgno h sedn ramus. except ascending phians the more a of Marada and origin; origin of point posterior its at ramus ascend- ing inflated or non-crested extensive a posteriorly symphysis; less a gracile, dentary; a narrow including: features, primitive ique few the of Dipro- superfamily todontoidea. one the represent of synapomorphies may for- curve and pa- lophs ward and crescentic diprotodontids the lorchestids, In a the molars. in lies in lophid lower oriented resulting hypoconid or entoconid, curved posteriorly the the to to and posterior posterior wynyardiids metaconid, lies protoconid and the the ilariids more wherein the are of lophids and The reminiscent molars. lophids molars lower weak the the on lower of construction the primitive on tocristid oas npriua,tepoooisof protoconids the the of particular, M cuspids most in buccal wear molars, the of on pattern pronounced oriented itudinally small a ramus. ascending and the talonid of inclination the of than angle wider is that 2-3 nte nqefauei h flattened the is feature unique Another tiigfaueo h etr in dentary the of feature striking A naddition, In arcanum Marada n h yooiso M of hypoconids the and iesfo l te vombatomor- other all from differs n n hti nqeamong unique is that one and Wakaleo Marada Marada osse eea un- several possesses nhvn trigonid a having in , xiisalong- a exhibits etr evolved dentary 1-2 hswear This . Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 trbtbet euldmrhs snot is dimorphism The sexual known. to attributable 0KRNBLACK KAREN 30 eeoii o obtmrhaadcon- and Paleo- Vombatomorphia a difficult. for suggested origin (1987) differentiation gene Archer & familial the Aplin of of assessment refined time million any 26 fossil makes and Australian years 55 an between record of mammal Oligocene. lack late the the However, by distinct was vombatomorphian seven (Creaser families the age of in Each Oligocene 1997). late sedi- is and oldest ments stratigraphically Riversleigh’s Maradidae. family establishment new a the of and, other warrants known any consequently, to presently this different vombatomorphian makes molars) strikingly lower lingual dentary the the on of cuspid cingulum distinct with the a row and of length; tooth presence ramus cheek horizontal to the respect of length short dentition, known. are specimens for complete more niche masticatory ecological arcanum unique mandibular and the a action of suggest of construction symphysis attachment, lack the muscle the the the and for case, row, wear, areas any tooth pronounced tooth In cheek of lip. shortened mobile pattern suggestive large longitudinal be may a to foramen jaw) of lower innervation mental anterior increased the large suggests (which The muscle complexes. masseter/pterygoid and the of strengths temporalis relative the assess to difficult ftouiu etrsfor features unique of two possession the of combination with features the primitive Vombato- these to However, respect morphia. with define plesiomorphic to diagnosis in used generic features the they Most defined. as currently are family, vombatomorphian known articular Unfortu- in preserved and not species. are process condyle the coronoid of the nately, female gracile a itsSt sitrrtdt efrom be to interpreted is Site Hiatus lower and dentary the of basis the On htwl eanamseyuntil mystery a remain will that Marada Marada osntfi noany into fit not does etr a represent may dentary Marada Marada Marada Marada ,soitis (the are euto optto ihdiprotodon- with macropodoids. competition and toids of as possibly result Miocene, a the beyond failed survive that to evolution vombatomorphian Miocene, in middle the Marada fossil than a younger have not record do which Like wynyardiids, Vombatomorphia. the of a diversity supports higher certainly Riversleigh at deposits tdnso h nvriyo e South New of University postgraduate Wales. the and volun- of staff of students with hundreds together many teers field from the & come in Sue has assistance and Vital Lavarack, Scott-Orr. Jim Don & Martin Riversleigh Sue Beavis, Margaret Dickson, the Clark, includ- Elaine Wales; supporters Ltd; ing private South and the Pty Inc.; Society New Wales; Zinc South of Century New Society of Linnean Zoolo- Society Royal the Geo- gical Museum; Museum; Australian Australian Queensland the the the Society; Ltd; graphic Pty ICI the Unit; Australia Service; Queensland Heritage Wildlife World the Commonwealth and Parks Territories; National Environment, and of the Sports Com- the Department Bartholomai); Wales; A. monwealth South New and of Archer University M. (grants to (Queensland) Scheme National Grants the Estate Archer); Grant M. to Research (grants Australian Scheme the from has Riversleigh manu- come at research this for comments. support of useful Vital many draft provided a and Pledge, script read Neville critically and who Sue Murray, Archer, Peter Mike their Hand, to compara- Thanks Anna for to material. access thank tive Godthelp and support to and Henk advice like and would Gillespie author The Acknowledgements in h rsneadol nw occur- known of only and rence radia- presence the pre-Oligocene The diverse tion. represent a families of remnants known that cluded a ersn neryexperiment early an represent may Marada noeo h oldest the of one in ALCHERINGA Downloaded By: [University of New South Wales] At: 04:45 25 May 2007 C B C A B A B A A References ALCHERINGA A C F LANNERY LACK LACK EECHER OOKE REASER LEMENS RCHER RCHER RCHER PLIN PLIN psus tde nEvolution In in Studies Queensland. 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Society Sydney, Zoological Wales, Royal South the and Ltd Pty Evolution classi- syncretic In new fication. a with systematics marsupial n oe oa vlto ihntesubfamily. the 41 Museum within Queensland the evolution of Memoirs molar lower and 314. topogra- of significance phy. potential the Riversleigh: Australasia In In evolution. marsupial in significance ..B T.J. uem41 late Museum early north- Queensland. Riversleigh, to western Fauna, middle Local Encore late Miocene the from (Marsupialia) . 97 airna ntm fteearly the of anatomy Basicranial 1987. K., , ,K.&A 209-228. , ,K.&A Palorchestidae of species new A 1997. K., , .. 97 w e abrn kangaroos balbarine new Two 1997. B.N., , ,M.&W radiation. marsupial Australian The 1984. M., , their and (phascolarctids) Koalas 1978. M., , etbaeZoegah n vlto in Evolution and Zoogeography Vertebrate rcouu dicksoni Trichosurus . 97 lgcn-icn eiet of sediments Oligocene-Miocene 1997. P., , .. 97 ucinadfso tthe at fusion and Function 1977. 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Sydney, Royal Wales, the South New and Ltd Pty Sons h aspai n hlgn ftePhalanger- the of iformes. phylogeny within a and characters Marsupialia basicranial the some of distribution icn gm oa an fSuhAustralia. South medial of In Fauna the Local from Ngama Miocene marsupial vombatoid primitive I ogas re,London. Green, Longmans, II. aoda rmtemda icn fSouth of Miocene medial the Australia. from batoidea) snideri fSuhAsrla In Evolution in Australia. Studies South Formationof Etadunna Miocene middle the from dea) Vombatoi- (Marsupialia: ?wynyardiid of species and .. A T.H., , A & T.H.V. , . 80 eea olg rsseai natural systematic or Zoology General 1800. G., , . 1866. R., , RCHER ..&A & T.J. , osm n psus tde nEvolution in Studies Opossums: and Possums .. 1987a. N.S., , .. 97.Anwgnsadseisof species and genus new A 1987b. N.S., , .. 97 h irtdnia from Diprotodontidae The 1967. R.A., , ,P.F.,W ..&W & M.S. , e irtdna Mruila Vom- (Marsupialia, diprotodontan new a , . udues .Kase,Lno,249-552. 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Australian fauna South South and Desert, formation Tirari Tertiary the new A 1967. eainhp,eouinadboegah.In course short biogeography. a for and Notes Mammals: evolution relationships, fGooia cec,Suisi elg 8, Geology in Studies Science, 48-71. Geological of fNwSuhWls yny 293-317. Society Sydney, Zoological Wales, Royal South the New and of Ltd Pty Sons & oacia)fo ot utai,adanew a and Australia, of South species from (Marsupialia:phas- koalas colarctidae) Miocene of species two eor fteQenln uem13 Museum Queensland the of Memoirs tde nEvolution in Studies LEDGE RCHER .. 96 h kl of skull The 1956. J.T., , .. T R.A., , .. A R.H., , ..&W & R.H. , d,Uiest fTneseDepartment Tennessee of University ed., , LEDGE .. 1987c. N.S., , d,Sre ety&Sn t t n the and Ltd Pty Sons & Beatty Surrey ed., , .. T M.O., , T M.O., , P M.O., , marsupials: of Families 1984. M.O., , Perikoala ,N.S.,R EDFORD RCHER ,M.A EDFORD EDFORD ,M.A LEDGE ICH OODBURNE ,M.,B .In ..&W & R.H. , osm n psus Stu- Opossums: and Possums Madakoala ,T.,R RCHER ..&A & N.S. , RCHER osm n Opossums: and Possums .. A R.H., , A & R.H. , ARTHOLOMAI 427-462. , d,Sre ety& Beatty Surrey ed., , ICH .. 97 The 1987. M.O., , hlclocarnifex Thylacoleo ALCHERINGA d,Sre Beatty Surrey ed., , ,P.&W e eu and genus new a , M ora of Journal BMR OODBURNE 53-57. , ,T.W.B eod fthe of Records ,M.A RCHER RCHER . P A., , RCHER 125-140. , ELLS Possums ,M.& M.O., , RCHER R.T., , ,M., ROAD- ,M., LANE ,M. , , .