Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 O:10.1080/03115510701305082 DOI: (online) Ó 1752-0754 (print)/ISSN 0311-5518 ISSN icn fRvrlih otwsenQenln n mlctosfor implications and Queensland northwestern Riversleigh, of Miocene e words: Key 17.12.2005. revised 17.6.2005; received Australia; 2052 Wales South L GODTHELP HENK AND LOUYS JULIEN for implications and Queensland Miocene northwestern the Riversleigh, from of fossils koala of Descriptions en oedsic in and distinct more metacrista of posterolingual being basis the the features: on two species these (1987) distinguished Springer fragments. and molar de- molars upper lower two two was of basis Australia the on scribed South Formation, Kanunka, Etadunna Lake RV-8453, Locality molar. upper single on kanunkaensis a described of basis was Local the Australia, Ngapa- Kutjamarpu South Lake kaldia, the Formation, Wipajiri from Fauna, 1967, ton, KUTJAMARPENSIS LITOKOALA 1 02Asrla[[email protected]]. Australia 2052 pce of species Litokoala 31, Alcheringa arn pcfi itnto,telte en h uirsynonym. junior the being latter the distinction, specific warrant between variation morphological nqecmiaino ohpeimrhcadaoopi etlfaue,mkn t hlgntcposition phylogenetic its making features, Litokoala dental phascolarctid apomorphic Miocene and early plesiomorphic both of within combination unique a Phascolarctidae) colo ilgcl at n niomna cec,Uiest fNwSuhWls yny e ot Wales South New Sydney, Wales, South New of University Science, Environmental and Earth Biological, of School OUYS 07Ascaino utaainPalaeontologists Australasian of Association 2007 . B J., , Litokoala .kanunkaensis L. Litokoala rmJm aoslSt,Sse sas ecie.O h ai fti etto,i ssgetdta the that suggested is it dentition, this of basis the On described. also is C System Site, Carousel Jims from srvsdo h ai fnwmtra rmteeryt ideMoeedpst fRvrlih new A Riversleigh. of deposits Miocene middle to early the from material new of basis the on revised is LACK Litokoala 910 SN0311-5518. ISSN 99-110. pigr 97 rmteUCR the from 1987, Springer, . A K., , icl oitrrt nsm set fdna morphology, dental of aspects some In interpret. to difficult sdsrbdfo h ytmBOtst oa Fauna. Local Outasite B System the from described is hsoacia,Moee ieseg,Australia. Riversleigh, Miocene, Phascolarctidae, , 1 AE BLACK KAREN , RCHER iikaagreystanesi Nimiokoala . H M., , ossigaless a possessing .kutjamarpensis L. .kutjamarpensis L. 1 AND ..&G & S.J. , Litokoala Litokoala 2 1 aut fSine nvriyo e ot ae,Sde,New Sydney, Wales, South New of University Science, of Faculty IHE ARCHER MICHAEL , Stir- ODTHELP tro,16 and 1967 Stirton, ; lc rhr 97 h etto faprilsulof skull partial a of dentition The 1997. Archer, & Black . ue 07 ecitoso ol osl rmthe from fossils koala of Descriptions 2007. June, H., , olw npriua,teM the Henks particular, from In recovered Hollow. fragment dentary a to assi- it and material gned new to this Archer Riversleigh, described & early (1997) of Black the Queensland. deposits northwestern from Miocene described middle been of species have to referable specimens with kutjamarpensis compared L. margin tooth buccal convex ecie n sindto assigned and described 19)asge tcnpcfi status. conspecific it Archer assigned simi- & (1997) of Black of disparity, material geographical level and holotype high the kanunkaensis a with such larity shared tary ic httm,svrlmr complete more several time, that Since nM An (Marsupialia, 3 .kanunkaensis L. Litokoala iool garyjohnstoni Litokoala .kanunkaensis L. rmHnsHlo a also was Hollow Henks from htdsieischronological its despite that .garyjohnstoni L. 1,2 UAN .HAND J. SUZANNE , . Mruila Phascolarctidae). (Marsupialia, pigr 97de not does 1987 Springer, .kanunkaensis L. scnegn nthe on convergent is ntebssof basis the on p o.possesses nov. sp. 2 fti den- this of Litokoala 1 L. . Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 omrrpeetda M an of represented holotype the former and kutjamarpensis molar L. this between kanunkaensis. for represented recovered L. molar specimen upper complete this first the such, assign- North reasonable As a Kanunka ment. was partial this from Fauna, the Local and recovered molar simi- molar this the between and larity species Riversleigh, koala from of recovered number small the Hollow, osqec,tecnpcfiyo hs two discussed. these is of species conspecifity the consequence, ie h soito fti oa with molar this of the association the Given sltdM isolated emd ewe h M the between made to comparisons be allowed has skull tooth this complete of the row of (Louys Description study prep.). is separate in a description of cranial subject the the here; described is as described C System referred also to Site, was and Carousel Riversleigh, deposits, Jims from ered ytmBdpst,Rvrlih a referred to was Riversleigh, deposits, B System pensis atrrpeetda M an represented latter ne nteOutasite the in differ- ences morphological significant show skull with Comparisons kutjamarpensis level. L. species to defined oa opooyflosAce (1978) Archer follows (1987). morphology Archer Molar & Archer Aplin and follows menclature (1997) al. Archer et & fol- Black nomenclature (UCR). lows systematic Riverside level at Species California the Uni- of the of and versity (QMF) collection Museum palaeontological Queensland the as in herein described distinction. is garyjohnstoni Litokoala specific it warrant Consequently, to sufficient 0 UINLOUYS JULIEN 100 Litokoala etmxlawt P with maxilla left A ata kl ihP with skull partial A aeilrfre ohri sdeposited is herein to referred Material Litokoala .kanunkaensis L. n h M the and 19) ihrlvlsseai no- systematic Higher-level (1999). 4 rmteOtst oa Fauna, Local Outasite the from .kanunkaensis L. yBak(92 u a not was but (1992) Black by Lus20)adi here is and 2004) (Louys 3 n h partial the and oee,acomparison a However, a o rcia sthe as practical not was of .kanunkaensis L. etr rmHenks from dentary 1 p nov. sp. . Litokoala 3 1 -M of 3 3 -M hra the whereas , 1–4 t dentition Its . .kutjamar- L. 1–3 ta.ALCHERINGA al. et a recov- was Litokoala maxilla, n an and .Asa tal et . xedn rmteae ftemetaconid the of M apex on crest the posterobuccal from a extending ‘having in colarctids differentiated Diagnosis. nov. ietdcetfo h rtcndapex protoconid M the from of crest directed eldvlpdpseoiga upon cusp a P having posterolingual following: well-developed the in phascolarctids M opi upl tteanterolingual M of the metaconule the at of base cuspule morphic Litokoala Owen, PHASCOLARCTIDAE 1839 Family 1987 Archer, & Aplin PHASCOLARCTOMORPHIA Infraorder 1984 Woodburne, VOMBATIFORMES Suborder 1866 Owen, DIPROTODONTIA Order palaeontology Systematic diinlseis iool kanunkaensis 1987; Springer, Litokoala species. Additional Stirton (in 1967 ton, kutjamarpensis Litokoala species. Type 1967) 19;M fAce 17)i P of dP3 Woodburne is 1999), Archer (1978) follows Luckett ture Archer nomencla- Biostratigraphic (1993)]. of and Luckett older M1 the ‘hypocone’) [1993; to homologous convention, the is that understanding metaconule the ter- with Woodburne neometaconule minology, & and Tedford metaconule (1987; by revisions with livle 86weei svariably is it where present). 1816 of species Blainville, of exception (1997). 3 n aigawl-eeoe neo- well-developed a having and ; 4 ;awl-eeoe posterolingually well-developed a ); 2–4 4 .I lodffr rmalother all from differs also It ’. atog hsceti eue on reduced is crest this (although tro,16 i Stirton (in 1967 Stirton, lc rhr(97 .218) p. (1997, Archer & Black Litokoala iool garyjohnstoni Litokoala tal et tal. et tal et 19)adCreaser and (1993) . rmohrphas- other from 18,19,1997, 1994, (1989, 1967). . Phascolarctos 1 wt the (with tal et Stir- sp. de . Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 oi rltv ohpcnd nM on ento- hypoconid) displaced to anteriorly (relative ‘an conid having in upt aatl nM on well-developed a parastyle having: cuspate in 1997 Archer, of & species for except tids ALCHERINGA n yooi fM of metaconid the hypoconid on and ribs lower internal the and molars; on preentocristid is bifurcate ante- an riorly cristid preentocristid; the postmetastylid with continuous the metastylid wherein ante- a tooth; fold squarer that of much margin rolingual a consequently, and, features, eto h otrtcitdadcristid and M postprotocristid on the obliqua of ment o etedt n’(lc rhr1997, Archer 219). & – 218 (Black pp. end’ to end meet not isexcept tids eaoueo M on metaconule 97 .218). p. 1997, tldadamr iga rtcndo M on protoconid lingual more more proto- a and and a developed stylid obliqua larger molars; cristid a the lower postprotocrisitd; of the junction in lingual more entos- ridge a an tylid paraconule; neometaconule; linear less larger ‘being crescentic, a namely: selenodont; by more having (1997), crowned; listed higher Archer features smaller; & those in Black 1957 Stirton, Woodburne h neirbs ftenoeaoueof neometaconule the of M base anterior at cuspule the neomorphic ‘a having to addition otro agno M on margin posterior togydvlpdetsyi’(lc & (Black 219). p. 1997, entostylid’ Archer developed strongly essrnl eeoe,mr bucally more M on developed, paraconid strongly positioned less a upt aaoueadnoeaoueon neometaconule M and paraconule cuspate rtcnd nM to on (relative protoconid) metaconid displaced anteriorly akn h otrbca upo P of cusp posterobuccal the lacking hn a poe oblos P bulbous) to opposed (as chant 1–3 1 natrlnulpoooeceto M on crest protocone anterolingual an ; Litokoala Litokoala Litokoala ral eue eaoeand metacone reduced greatly a ; Litokoala Phascolarctos iesfo te phascolarc- other from differs iesfo te phascolarc- other from differs tal. et 4 iesfrom differs hri hs rsisdo cristids these wherein 4 4 n aallarrange- parallel a and ; 97and 1987 , n oerounded more a and ; iesfrom differs 4 1 nadto othese to addition In . Bak&Archer & (Black ’ 1 well-developed a ; Nimiokoala nhvn tren- a having in 1 n more a and ; Madakoala Nimiokoala Perikoala 3 n in and Black 4 3 ;an ,in 1 1 , , ; troaro ih P right of stereopair mm. 5 Holotype. 1) Table 1, kanunkaensis Litokoala Cst,Sse ,Riversleigh. C, System site, JC etadrgtdniinadpalate. and dentition right and left 1 Fig. ot utai.Ti eoi slate is deposit (Woodburne This age Kanunka, in Lake Oligocene Australia. of Etadunna side South Fauna, west Local Formation, North Kanunka Locality. Type . iool kujamarpensis Litokoala ICN OLS101 KOALAS MIOCENE AP29,argtM right a SAMP32397, 3 C oaiyRV-8453 Locality UCR n M and pigr 97(Fig. 1987 Springer, 1–4 A eerdmtra from material referred , M532 showing QMF51382, , cl a indicates bar Scale . tal. et B 2 . Occlusal , 1993). Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 al 1 Table AW M532LP QMF51382 ie C295 ih M right a UCR21945, Site: material. Referred LOUYS JULIEN 102 ie M289 ih M right QMF20809, JC9 protocone; Site: of half buccal and paracone lgcn nae(Woodburne late age is in Formation, Oligocene Etadunna Fauna, Local age. and Distribution nitc pe hetohdniin The dentition. with cheektooth complete palate upper partial intact almost a orbito- an and one arch, intact zygomatic relatively region, a temporal an with neuro- partial is cranium a Preserved basicranium, intact worn. almost molars lightly four all but with adult erupted young a of 1) Fig. General. Description Miocene middle as 1997). (Creaser interpreted north- are deposits, which C Riversleigh, System are from Queensland, western faunas local and JC9 JC Gotham, Gag, Hollow, Henks The M532 ata kl ihlf and left with skull P right partial Riversleigh: a Site, JC QMF51382, from herein material described Additional (1997)]. Archer & Black eaoeo ih M right a of metacone o afo P poster- of with half fragment ior dentary right 13079, eaoeo etM left a of metacone oa an:QF00,rgtP right QMF30500, Hollow Fauna: Henks Local Riversleigh: From (1987)]. a ie M351 ih M right QMF30501, Site: Gag ie M353 M QMF30503, Site: ¼ neirwdh PW width; anterior 3 esrmns(m fuprtehof teeth upper of (mm) Measurements . ,M ata kl QF51382, (QMF skull partial A 1–4 3 ,M . LM LM LM LM 1–2 3 auk ot Local North Kanunka M352 ih M right QMF30502, ; 4 3 2 1 x 1 h auk North Kanunka The rfre ySpringer by [referred rgetcontaining fragment ¼ otro it;* width; posterior 3 .941 .0RM RM RM 3.10 RM RP 4.37 5.01 5.03 4.13 – 5.03 5.38 4.69 5.08 5.37 3.16* 5.46 5.85 5.29 C299 a UCR21979, ; 4 WP WPW AW L PW AW L C290 a UCR21980, ; 3 rfre by [referred tal et 1 ta.ALCHERINGA al. et ;Gotham .; 3 1993). . ;QMF ¼ 3 ; aiu width. maximum etto Fg ,3A). 1, (Figs Dentition broken is fossa. glenoid cranium the of the forward away of portion dorsal n neouclyadtels extending last the posterobuccally, and one anterobuccally tooth, posterior one the the of descending, joins margin crests which posteriorly, four poster- one has The and cusp cusp. posterior opposite olingual arm the situated to short is anterior cusp pos- one The terolingual sinu- posterolingually. cusp, other extending the ously the anterolingually, of extending base at bifurcates the crest lingual buccally, The lingually. then and extending anterobuccally margin crests posteriorly, tooth well-developed the anterior three the of with situated behind is apex cusp mm 1.5 posterior the The cusp. from crest medial buccal A buccally cusp. three posterolingual extends the the the and in between cusps abruptly formed ending the trench apex, from cusp anterolingually crest medial well-developed slightly ante- A the extends margin. to tooth long- posteriorly rior mm cusp short 2.6 medial situated the by of is apex anterior cusp The the crests. posterior itudinal to and connected apex. cusp is margin. its cusp from lingually Medial tooth and poster- buccally anterior anteriorly, iorly, extend the crests pos- Prominent mm to 1.5 situated teriorly is cusp The cusp. anteriormost lie posterolingual additional which crest an of longitudinal with crescentic three slightly anteriorly, a cusps, along major than four posteriorly with wider chant, .kutjamarpensis L. 3 4 3 2 1 brvain:L Abbreviations: . .840 3.43 4.86 5.24 5.49 4.02 – 5.35 5.53 4.68 5.15 5.25 3.04* 5.55 6.15 5.03 h RP The 3 ¼ stren- is length; Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 neiry iigrs oasalcsueat LP cuspule The small base. a its to rise giving anteriorly, n rmtemda cusp. descend- medial crest the lingual from the ing to base and broader cuspule, a lingual developed better a with ALCHERINGA ue onn h otrlnulparacrista, posterolingual cus- the posterior joining the cingulum. of pule, apex anterior the from buccally postero- the extends paraconule joining the stopping Posteriorly, the of anteriorly, buccally, extending short extending arm arm other short bifurcates, a cuspule, The from anterior with the posterior. of anterobuccally apex the the extends than anterior paraconule the higher with bicuspate cuspule the is of base It lingual paracone. anterior the at paraconule situated The is tooth. the of and corner poster- obuccal the defined to postmeta- posterobuccally weakly extends The crista very posterobuccally. arm extending postparacrista, other the the meet to bifur- extending crest, anterobuccally main linear, its arm, is one with cates, premetacrista The transverse the valley. closing with and up premetacrista postparacrista, meet the to the posterobuccally the continuing extending margin; arm tooth buccal other the and down anterobuccally fading extending arm one postpara- bifurcating, with posterobuccally, The extends B. crista swelling cusp slight stylar a in representing margin tooth buccal the at ending ( linearly, extends A buccal preparacrista cusp The the stylar The at as margin ending developed. anterolin- tooth crest, an poorly directed as gually E extends corner and and parastylar well-developed, developed, B moderately is cusps D and A C cusp are cusps Stylar metacone slightly paracone deeper. and the with paracone open sur- and Buccal shallow the corner. of parastylar faces the the posterolingual the of proto- of and base the metacone base of and base cone posterior posterior the and paracone, anterior valley, the transverse the in shallow crenulations with surface posterior, the towards slightly h RM The 1 ssbudae tapering subquadrate, is 3 ssmlrt h RP the to similar is ¼ parastyle). 3 but , xed uclyt h ogtdnlvalley. longitudinal the other to the anterobuccally buccally postprotocrista, extends continuing the meeting arm bifurcates and one the anterobuccally, premetaconule with and extends The protocone crista the cuspule. of poster- metaconule the side on ridge the olingual by lingually and postprotocris- ta, and crista premetaconule by the buccally bound is valley the This and a metaconule. protocone forming the between thereby valley metaconule, closed anterolingual the the of on base cuspule a ridge meets poster- poster- it The anterior; iorly the the protocone. towards to abruptly the stops of crest face the olingual the from of extending base crenulations well-developed posterolingual and with is protocone, the crest paraconule at well-developed situated the very A between protocone. valley in paraconule the protocone the the from ante- crenulation the of a extends joining apex the ridge of from A robuccally metaconule end protocone. the the buccal and between valley the pre- transverse the at meeting and metaconule sharply, turning val- ley, transverse the towards posterobuccally extends It crescentic. more The is cingulum. postprotocrista anterior crescentic the becoming with long, continuous anterobuccally, a extending of crest, pre- consists The protocrista buccally. and gently protocone slope the metaco- metaconule of surfaces and Buccal metacone nule. the the longitudinal of between the valley arm into transverse linear fades neometaconule the the Posteriorly, in valley. ending extending arms of anterobuccally, parallel base with anterolingual metacone, the the at situated neometaco- is bifurcate nule anteriorly cuspate, protocrista. anterolin- A anterobuccal the extends joining crest gually connects The cuspule preprotocrista. it posterior the until to cuspule anteriorly perpendicularly anterior extends and The crest anterior cuspules. the from crest. posterior arise cuspule crests posterior Lingual postero- the extending from ridges lingually short three with ICN OLS103 KOALAS MIOCENE Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 n aaoueaerdcdrltv oM to relative reduced are paraconule protocone, and metacone, metacone/meta- paracone, the The crista. by is formed posi- margin posterior crescentic, metacone The the posteriorly. tioned and anteriorly posi- the tioned paraconule with and metaconule, paracone, and protocone, metacone the in oebcal,bcmn otnoswith directed cingulum. continuous posterior then the becoming postmetaconule, buccally, the more of the tip from posterobuccally extending crescen- ridge long tic a is crista postmetaconule The xed otebs fteprotocone. the of spur base directed the lingually to valley. short extends a apex, metaconule transverse the its of and the end anterior the into Between paraco- extends the nule posteriorly ex- preprotocrista; joining the other of short falling the but anteriorly anterobuccally tending preprotocrista, extending the ante- arm meeting bifurcating one single-cusped, riorly; para- is The crenulation. conule paraconule not the does protocone join ante- the from the ridge shorter; robuccal are arms neometaconule ipa:bte eeoe rnltoson crenulations developed better display: otrlnulypstoe hni M more in and than reduced positioned posterolingually greatly is metacone The hr neoigal ietdsu;teM the spur; a directed being anterolingually third the short arms, three into splits anteriorly neometaconule the bifurcate; premetacrista not does the paracrista; no is with anterolingual and developed D posterolingual distinct less less cusp is crenulations; protocone stylar the B; base of posterolingual the and on of ridge the juncture A reduced; the cusps at parastyle swelling stylar small the a reduced, parastylar as exists relatively the is metacone; is corner the paracone the than of deeper basin buccal shallow the on ntetinua alyformed valley triangular cusps. major three the the between in crenulations from found indistinguishable is neometaco- The nule margin. on cuspule posterobuccal small the a to reduced is metaconule 0 UINLOUYS JULIEN 104 M h RM The LM 2 n M and 1–4 r iia oRM to similar are 4 stinua u oareduction a to due triangular is 3 r iia oM to similar are 1–4 1 xetthey except ta.ALCHERINGA al. et xetfor: except 3 The . 3 3 . eaieymr endprsya onron M corner deeper parastylar defined a more a relatively protocone, metaconule, and of protocone between valley face anterior the een hr xssasrkn similarity kaensis striking and a of molars upper exists the between there herein, the oa erbetween wear in differ- differences molar the interspecific account the into than Taking ences. along rather differences row tooth represent to appear h aatlrcre nM on ante- corner far parastylar with as continuous the is extend preprotocrista the not riorly; does paraconule M546 etM left QMF51406, aaoueo M on paraconule otwsenQueensland. northwestern Property, Fossil Heritage World Riversleigh locality. Type ail ihP with maxilla Material. 2) Table garyjohnstoni Litokoala of of bounds synonym the within sis fall variation. or intraspecific row are, tooth the along taxa differences by these accounted therefore, between 1992). differing variation (Black koala of Features modern range the in the observed within this is However, difference metaconule. and protocone pensis jamarpensis distinguish to (1997) Archer sis of toothrow molar complete ouei etrdvlpdo h M the on para- developed of better basin is is conule buccal metaconule the developed; the better whereas developed, less 1 h igotcfaue sdb lc & Black by used features diagnostic the , h neirydrce r fthe of arm directed anteriorly The . Remarks s ecfrh osdrdtejunior the considered henceforth, is, .kanunkaensis L. .ktaapni.Ltkaakanun- Litokoala kutjamarpensis. L. nlcigtede alybtenthe between valley deep the lacking in osdffrfrom differ does ooye M545 etpartial left QMF51405, Holotype, ae ntedsrpino the of description the on Based . iool kutjamarpensis. Litokoala from uaie otepHill, Godthelp Outasite, 3 h rtcnl nM on protoconule the ; 4 iool kanunkaensis Litokoala . 3 ,M .kutjamarpensis L. iool kanunkaen- Litokoala pcmndescribed specimen p o.(is2 3B, 2, (Figs nov. sp. iool kutjamar- Litokoala 1–3 .kanunkaensis L. iool kut- Litokoala .kanunkaen- L. 2 Paratype, . oeworn more ; 4 . and 3 is Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 aaoe cprcnl;p—rtcn.Saebar mm. Scale 5 indicates pr—protocone. pc—paraconule; paracone; pa— ne—neometaconule; mt—metaconule; metacone; ( M545 clslseepi flf ata ail ihP with maxilla partial left of stereopair occlusal QMF51405, etto of dentition of flf M left of ALCHERINGA i.3 Fig. 2 Fig. ¼ aatl) ,C n .Abeitos me— Abbreviations: D. and C, B, parastyle), iool garyjohnstoni Litokoala iedaig fdniin described. dentitions of drawings Line . . iool garyjohnstoni Litokoala 4 cl a niae mm. 5 indicates bar Scale . iool kutjamarpensis Litokoala hwn tlrcssA cusps stylar showing ooyeadprtp,fo uaie ytmB Riversleigh. B, System Outasite, from paratype, and holotype , ; B etdentition left , A right , 3 ,M .garyjohnstoni L. 2 M Table QMF51406 n neoiga rtcn rs on crest protocone anterolingual M and cusp stylar C parastyle, pyramidal developed kanunkaensis from L. garyjohnstoni differs Litokoala diagnosis. Specific is Miocene middle faunas, to age. early of contained be to and interpreted basis the stratigraphy on of which, 1997), (Creaser deposit age. and Distribution M545P QMF51405 1 1–3 rsetcprcnl n bicusped and paraconule crescentic a ; . B–B esrmns(m fuprtehof teeth upper of (mm) Measurements . ICN OLS105 KOALAS MIOCENE 0 aaye M546 clslstereopair occlusal QMF51406, Paratype, , brvain sprTbe1. Table per as Abbreviations . M M M 3 4 3 2 1 .kutjamarpensis L. nhvn:abetter- a having: in uaiei ytmB System a is Outasite .242 3.44 4.56 5.45 4.20 5.87 5.29 – 5.86 4.02 5.48 5.05 3.47* 5.27 6.15 4.95 LAWPW A–A 0 Holotype, , and Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 neoiga aeo h eaoe(pro- M metacone in the reduced gressively of base anterolingual emtcnl nM on neometaconule Dentition present. is canal interorbital large A 2). (Fig. ihyceuae otro oeyo M on and moiety reduced posterior relatively crenulated a highly view; occlusal square base in protocone anterolingual the postprotocris- ta; the the and with crista junction premetaconule its between pocket lingual otrol,pseoigal n lingually and posterolingually medially, posteriorly, anteriorly, positioned cusps main iglma h otrlnulbs fthe of M base on posterolingual protocone a lacking the in at and cingulum outline; in circular is which rmta of that from oueadteatro n fteneome- para- the M of the on end of taconule anterior end buccal the and posterior the conule the to connecting margin ridges transverse 0 UINLOUYS JULIEN 106 for ueuli egtin height (whereas are in cusps cusp the posterior subequal tallest and medial together; the anterior, the is closer cusp and and are medial anterior the anterior cusps of apices the posterior the a cusps; between medial cingulum; link anterolingual reduced reduced mely rhi rsre n rgntsopst M opposite originates and zygomatic preserved the is of arch portion jugal A preserved. General. Description water. more to access have to capacity philan- engineer the Australians and helping Ltd is who Jaycar thropist of CEO Johnston, Etymology. trenchant) essentially still premolar. (albeit bulbous- more slightly looking crests. a is better-developed result associated overall a The and and cusp crown lingual posterolingual the the of along side midway cuspule distinct h P The .kutjamarpensis L. h rnhn P trenchant The . 3 uho h ail spoorly is maxilla the of Much of h pcfi aehnusGary honours name specific The .kanunkaensis L. iool garyjohnstoni Litokoala 1–3 1–4 ag upl tthe at cuspule large a ; . nhvn:a extre- an having: in ) .kanunkaensis L. 1–3 2–4 well-developed ; ,wihfrsa forms which ), 3 ossso five of consists (P 3 sunknown is ta.ALCHERINGA al. et differs ;a ); 4 2 , ntetoh h P The tooth. the on pxt h neircnuu fM cusp of cingulum posterior anterior the the to apex connects crest posteriorly directed prominent A medial short crest. the a by longitudinal of connected are apices cusps posterior The and point of cusp. apex a medial the at to the anterior margin slightly but lingual is opposite the cuspule on lingual anterior The situated subsequently into parts. ‘trench’ posterior cuspule, and lingual the lingual dividing the anterolingual crest directed The from posterobuccally crown. a the meets crest of at base anteriorly the curves apex. and crest this anterobuccal anterior from The anterobuccal extend the crests Strong to anterolingual posterior margin. cusp the medial the tooth to the of of close apex apex lies the The from cusp. anterobuc- medial an crest meets directed it cally where base crown the the to apex of cusp anterior anterobuc- the A from extends crown. cally the crest of buccal base prominent and the towards apex anterior fade buccal, is the the and from extend Anterior, cusp from crests lingual anterior margin. mm 1 most anterior approximately and situated tallest margin. tooth anterior The the to width in tapers a upocpigteatrbca corner anterobuccal the pyrami- occupying large, cusp the a dal with is parastyle comparison The in metacone. are reduced paracone slightly valleys the is of The surface buccal longitudinal The anterolingually. shallow. and slightly transverse slopes mar- is buccal gin margin paracone The buccal convex. strongly metacone buccally The is elongate. tooth The orientation. rolingual per ob eptec ewe this between cusp. posterior trench the deep and cusp a There posterolingual be crown. cusp the to of posterolingual base appears a the the into of fades by and apex cuspule the extends crest from lingual posterior A the crest. is longitudinal It to corner. the forming connected base the its of well with corner tip tooth posterolingual its the with of situated forward is cusp developed h neireg fM of edge anterior The 3 swdrpseiryand posteriorly wider is 1 a nante- an has 1 well- A . Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 ietdam iial ag,crescentic, buccally large, similarly posterolingually a A short arm. a with directed and bifurcates, arm directed the paraconule then the posteriorly of from apex, base extends A crest the directed paracrista. at terminates posterolingual paraconule and anterobuccally the from apex An extends paracone. crest directed the anterolingual of the point at base crescentic situated this large is A at paraconule E. swelling of cusp slight stylar corner represents A posterobuccal tooth. the the at minating ter- crescentic posterobuccally, The extends premetacrista. postmetacrista the junc- with its tion before C postparacrista cusp the stylar of intersects apex posterobuccally – the anterior linear from ridge short an directed A – A 3B). create cusps (Fig. D to ‘cingular’ of as alignment so posterior of points cristae and bifurcation the C, the over B, C taken cusps have though cusp D as appears Stylar It pyramidal apex. well-developed valley. a of consists transverse itself the end buccal the of arm, off buccal closing posterobuccal the effectively of at margin a premetacrista apex the and the meets which C, to cusp connects stylar that antero- short arm a poster- buccal into extends bifurcating the postparacrista obuccally, D. linear cusp between A stylar The and margin. but occurs premetacrista buccal bifurcate apex connection the paracone to similar lingual the slightly a opposite at B cusp point stylar meets which the arm, posterobuccally directed ante- meets a and an which crista, postparastylar into arm, bifurcates directed tooth robuccally it buccal where the margin towards anterobuccally parastyle. worn extremely an of base the (as in cingulum 1967) marpensis the Stirton buccal between by the described connection and buccal paraconule the that ftetoh trsmlsbti slightly is in but found pensis resembles cusp that It than larger tooth. the of ALCHERINGA hr,cecni rprcit extends preparacrista crescentic short, A and .kanunkaensis L. ooyei nfc h posterior the fact in is holotype ti o evident now is It . .kutjamar- L. .kutja- L. agn ag upl sstae tthe M in at metaconule situated the the of is base cuspule lingual anterolingual in large the A to crista buccal margin. well premetaconule valley transverse the the meets postprotocrista bifurcate paracone and crescentic, the A opposite apex. paraconule point longitudinal a the at the protocone between in terminates valley and protostyle preproto- crista the from well-developed posterobuccally extends A its posterolingual the at of metacrista. base terminates to- the and with junction mostly posterior the extends wards posterior crest A valley. neometaconule runs transverse metacone that the the of through ridge base anterior transverse the A to contin- parallel the is crista. with spur uous premetaconule anterobuccal short the second from directed spur short buccally a neo- meets A crest The anterolingual anteriorly. metacone. bifurcates ante- the metaconule the of at base lies rolingual neometaconule bicusped neiryt h neircnuu;the cingulum; anterior the connects paraconule to the anteriorly reduced; is meta- crista posterolingual the premetacrista; postparacrista and the are join valley and continuous transverse parallel more the occupy transverse that the ridges are metacone, reduced; the on relatively extent, lesser paracone a the to on and, cusps stylar poster- the and more olingually; margin slope margin buccal buccal convex paracone metacone less the is both margin and buccal parastyle; metacone pyramidal the the lack they features: M n sporsieyrdcdfo M from reduced progressively is and o agno h tooth. the of poster- margin the ior with The continuous becomes view. it occlusal until directed posteriorly in is the crista square postmetaconule of is proto- base base anterolingual the cone the result, a into As crown. apex protocone from extends the strong crest A cuspule. directed the anterolingually of base the postproto- and crista, crista, junc- premetaconule the the between of tion formed is pocket lingual 1 oee,te ie ntefollowing the in differ they However, . h M The ICN OLS107 KOALAS MIOCENE 2–3 r iia nms epcsto respects most in similar are 2–4 .A 1 Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 fawl-eeoe otrlnulcs on cusp P posterolingual well-developed a of o.in nov. ftemtcnl nM on metaconule base the anterolingual the of at cusp neomorphic ouei oerdcdi M in meta- reduced the more and is rounded; conule is more metaconule and the reduced at of base cuspule anterolingual the is the the preprotocrista; and at the protocone to ridge the is joined posterior of base short protostyle anterobuccal a the the to reduced to crista; connects premetaconule variably neometaconule nlso of Inclusion Discussion n aayeae otpoal,fo a from probably, individual. most single are, paratype holotype and the morphology, general and wear, Remarks. weakly is paracrista the developed. anterolingual crown; an the crenulated; is and of paracone apex the base of metaconule surface the buccal into the fades from and extends lingual median crest prominent a pattern; from transverse crenulation indistinct the is neometaconule the the valley; towards posterior the margin from are fine radiate ridges a poster- of cusps respectively; network and margins, these posterobuccal tooth olingual severely of the at the apices are situated that: the metaconule in reduced; and differs it metacone However, moiety. 0 UINLOUYS JULIEN 108 anoetcmuiiso northwestern of (Archer Oligo-Miocene Queensland communities the within biodi- rainforest of present levels This high versity prep.). formally commonly in be the Archer to reflects & yet (Black others described with Area, Heri- World tage Riversleigh the from described garyjohnstoni iaino etlfaue htaeboth are that features dental of bination 3 h M The iool garyjohnstoni Litokoala n h rsneo well-developed a of presence the and Litokoala 4 ntebsso ie ereof degree size, of basis the On ssmlrt M to similar is stetidseiso koala of species third the is iool garyjohnstoni Litokoala sbsdo h presence the on based is tal. et 91 1999). 1991, osse com- a possesses 3 3 1–3 nisanterior its in . . ta.ALCHERINGA al. et Litokoala sp. fprilsul fboth of skulls partial of of analysis analysis conver- However, phylogenetic of dentitions. a phascolarctid result on the based gence as interpreted similarities authors these latter the 1997; Archer, eerha ieseg a oefo the from come has Riversleigh for at Anna support research Vital thank herein. fossil described to the material of preparation like skilled for would Gillespie authors The Acknowledgements e,tececni aaoue n the in and M found the also paraconule, are neometaconule, crescentic bicusped the cor- parastylar ner, cuspate developed strongly ulbs ftemtcnl (M metaconule anterolin- the of the base on gual cuspule neomorphic and nlsspromdb lc Archer pola- character & therein, determined subsequent rities Black the and by (1997) within performed analysis apomorphic Litokoala and plesiomorphic tlrcs .I smr eie nits in parastyle derived well-developed more a is (M of It C. possession cusp stylar tRiversleigh. at B in species of garyjohnstoni L. into between affinity close genera. these a indicates prep.) (in Litokoala oesLclFua ugssthat suggests koala Faunas kutjamarpensis collected Local L. Dirks and recently Towers Sputum Camel the middle from Preliminary of material the deposits. to C analysis restricted System been Miocene has sleigh pce nisrltvl oebulbous, more relatively its P cuspate in other to species relative plesiomorphic ytmBLclFuaa Riversleigh. at Fauna Local Previously, B System garyjohnstoni 1 eea fteefaue,icuigthe including features, these of Several h usmto of subsumption The ,prcnl,bcse neometaconule, bicusped paraconule, ), .kutjamarpensis L. 1 of ntebsso h character the of basis the On . 3 yLus(04 n Louys and (2004) Louys by iikaagreystanesi Nimiokoala n h rae eeomn of development greater the and Litokoala skonfo uaie a Outasite, from known is Litokoala ean h oa number total the retains sas rsn nSystem in present also is aeilfo River- from material n h icvr of discovery the and ttwo. at .garyjohnstoni L. .kanunkaensis L. Nimiokoala 1–3 Litokoala Litokoala ). lc & Black tal et and is . Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 A A References this on comments to thoughtful manuscript. like their Turnbull would William for We and Pledge Wales. Neville thank South New University of the of staff students as from postgraduate well come and as volunteers has Scott-Orr. of field hundreds Don the many & in assistance Sue & Vital and Sue Lavarack Dickson, Clark, Jim Martin Elaine Beavis, Johnstone, Margaret supporters River- Gary private the and including Ltd; Inc.; Society Pty South sleigh Zinc New South of Century New Society Wales; Linnean of the Society Wales; Zoological the Museum; Royal Australian the the Museum; land Ltd; Queens- the Pty Society; Geographic Australia Australian ICI Unit; World Heritage Commonwealth and Terri- the Parks Service; and National Wildlife Queensland Sports the Environment, tories; Depart- of Commonwealth ment the Wales; New of South University Archer the M. Bartholomai); A. to and Grants (grants Estate Queensland National Scheme, the Archer); M. (grants to Scheme Grant Research Australian ALCHERINGA A A A RCHER RCHER RCHER RCHER PLIN rpy orlto n niomna change. environmental 25 biostrati- Zoologist and Australian of correlation overview graphy, preliminary northwestern Queensland: Riversleigh, of mammals Fossil 1989. asadifrne bu aaoaias In mam- Vegetation palaeohabitats. Australia’s Australian about of the of inferences history History and the mals in Patterns Riversleigh d,CmrdeUiest rs,Cmrde 80- Cambridge, Press, 103. University Cambridge ed., n oa olgclSceyNwSuhWales, South New xv-lxxii. Society Sydney, Zoological Royal and Evolution classi- syncretic In new fication. a with systematics marsupial rtto ftehmlg fmruilcheek- marsupial of reinter- homology a teeth. the and of marsupials pretation in boundary premolar 31-35. ,K.&A . H M., , G – M., molar , the of nature The 1978. M., , . H M., , eor fteQenln uem18 Museum Queensland the of Memoirs ,M.A ed yny 6 pp. 264 Sydney, Reed, , ODTHELP RCHER AND AND osm n psus tde in Studies Opossums: and Possums RCHER ..&G & S.J. , ..&G & S.J. , . H H., , . 97 eetavne in advances Recent 1987. M., d,Sre ety&Sons & Beatty Surrey ed., , 29-65. , AND ..&M & S.J. , ODTHELP ODTHELP ..H R.S. , . 1994. H., , 1991. H., , EGIRIAN ,D., ILL , , C O A S O B A L L B B B L PRINGER OUYS LACK OUYS UCKETT LAINVILLE LACK LACK REASER RCHER RCHER WEN WEN &W chroM’97 h ieseg ol eiaefsi property, fossil In Heritage of Australia. sediments World Queensland, Cainozoic Riversleigh the the of Correlation 1997. 1816 B G . M J., K us tde nevolution in studies In sums: bearing evolution. their phascolarctid on and Phascolarctidae) (Marsupialia: M ety&Sn n oa olgclSceyNew 319-325. Society Sydney, Zoological Wales, Royal South and Sons & Beatty I ogas re,London. Green, Longmans, II. 7 London of Marsupialia. Mruila hsoacia)fo Riversleigh, la of from revision a with Queensland, northwestern Phascolarctidae) (Marsupialia, eMnplir otele,Fac,131-152. France, Montpellier, Montpellier, de d. E eds., acia)adterfntoa interpretation. functional their and larctidae) Phasco- (Marsupialia: skulls koala Oligo-Miocene of eor fteQenln uem41 Museum Queensland the of topography. Memoirs of significance potential the Riversleigh: e cecsprl Socie la par sciences des syste distribution eeo ieseg,nrhetr Queensland. northwestern Riversleigh, of Mio- cene early the from Marsupialia) (Phascolarctidae, turnbullae: eclts oorms al uhrasand eutherians early marsupials monotremes, multitu- differentiation, berculates, Mesozoic In Phylogeny: mammals. therian mal in homologies dental aoy21 malogy mammals. Australian of diversity iyo e ot ae,Sde,6 pp. 63 Sydney, Wales, Univer- South Science, New Biological of of sity School thesis, ours Riversleigh from phascolarctids ne,Uiest fNwSuhWls yny 5pp. 95 Sydney, Wales, South New Sci- of University Environmental ences, and Earth Biological, of School on characters based phylogeny cranial vombatiform of examination an RAMMAL . . 89 ulnso lsicto fthe 1866. of R., , classification a of Outlines 1839. R., , EAR ILLESPIE ,J.,2004. ,J.,A ,K.&A C ,K.&A 1992. K., , eor fteQenln uem41 Museum Queensland the of Memoirs K . A M., , H M., , . 97 lgcn-icn eiet of sediments Oligocene-Miocene 1997. P., , .. 93 notgntcassmn of assessment ontogenetic An 1993. W.P., , 113-124. , ICN OLS109 KOALAS MIOCENE .. 97 oe oasof molars Lower 1987. M.S., , ROE USSER ENNA . K B., , ´ ...d,11.Porm ’n nouvelle d’une Prodrome 1816. de, H.M.D. , oePaiu e atsE Hautes des Pratique cole PLIN . 99 h vltoayhsoyand history evolutionary The 1999. S., , . G A., , ,J.-PA . C J., , 1-45. , ,F.S.S d. pigr e ok 182-204. York, New Springer, eds., , . M A., , AND h otpeimrhcdfiiiekoala definitive plesiomorphic most the RCHER rna ecitoso rhi olsand koalas archaic of descriptions Cranial ,K.,A RENA 5-19. , RCHER RIKMAN ytmtc n aaoilg ffossil of palaeobiology and Systematics nteAaoyo Vertebrates of Anatomy the On rceig fteZooia Society Zoological the of Proceedings .. G S.J., , OOKE ODTHELP GUILAR ´ . B R., , aiu ure du matique . 1997. M., , nulse oor thesis, honours Unpublished . . nprep. in M., , YERS RCHER ZALAY . M A., , . C B., , . P T., , ODTHELP ,S.L ASSAROVA . npe.Adescription A prep. in M., , ,M.J.N . G H., , ,M.A ´ te ACKNESS ´ REASER ce uCongre du Actes EGENDRE hlmtqed Paris de philomatique LEDGE Nimiokoala ` nulse hon- Unpublished . n animal. gne ,H.&C osm n opos- and Possums OTT RCHER . B M., , rsaol lucy- Priscakoala utainMam- Australian . C P., , OVACEK . M B., , . W N., , . H M., , ´ ue Institut tudes &J.M 303-314. , d,Surrey ed., , REASER 209-228. , ROSBY LACK Litokoala e.nov. gen. UIRHEAD Litokoa- M.C. & AND ANG Bulletin ICHAUX ` Bio- s Mam- Vol. , K., , K., , ,P., ,S., ,Y. , Downloaded By: [University of New South Wales] At: 23:19 23 May 2007 W S T 1 UINLOUYS S JULIEN 110 TIRTON TIRTON EDFORD OODBURNE vlaino h oooyo oa up nthe in an cusps and molar In of Australia Diprotodontia. homology South the of of evaluation strata marsupials Miocene vombatiform of from family new a Ilariidae, nEvolution in ia cecs tde nGooy8 Geology in Studies Sciences, gical ed., n oa olgclSceyNwSuhWales, South New 401-418. Society Sydney, Zoological Royal and h ot utainMsu 13 Museum Australian South the Australia. South of Fauna Palankarinna ot utainMsu 15 from Museum Australia. Australian fauna South South and Desert, formation Tirari Tertiary the new A 1967. as oe o hr Course Short a In for Notes biogeography. mals: and evolution tionships, .. T R.A., , Pliocene the from koala new A 1957. R.A., , nvriyo ense eateto Geolo- of Department Tennessee of University ..&W & R.H. , .. 94 aiiso aspas rela- marsupials: of Families 1984. M.O., , ,M.A EDFORD RCHER osm n psus Studies Opossums: and Possums OODBURNE ..&W & R.H. , d,Sre ety&Sons & Beatty Surrey ed., , 427-462. , .. 97 The 1987. M.O., , ,T.W.B 71-81. , OODBURNE 48-71. , eod fthe of Records ta.ALCHERINGA al. et ROADHEAD eod of Records M.O., , Mam- , W W OODBURNE OODBURNE P S W fteEaun omto lt lgcn)of Oligocene) 13 ogy (late Formation Australia. South Etadunna the of magnetostratigraphy and biostratigraphy mammal osadRylZooia oit e South New Society 293-317. Sydney, Zoological Wales, Royal and Sons & w pce fMoeekaa (Marsupialia: koalas new a of and Miocene species Australia, South of from Phascolarctidae) species two tde nEvolution in Studies PRINGER LEDGE OODBURNE 483-515. , .. 1987. N.S., , .. M M.O., , T M.O., , .. P M.S., , Perikoala ..&S & J.M. , ora fVrert Paleontol- Vertebrate of Journal EDFORD ,M.A AC .In LEDGE Madakoala F PRINGER ADDEN RCHER osm n Opossums: and Possums .. A R.H., , .. P N.S., , .. C B.J., , .. 93 Land 1993. K.B., , d,Sre Beatty Surrey ed., , e eu and genus new a , RCHER OWER ASE ,M.& J.D., , J.A., ,