palaeontologia electronica · 2019-09-10 · travouillon et al.:riversleigh basal macropodoid 2...

Palaeontologia Electronica palaeo-electronica.org

Revision of basal macropodids from the Riversleigh World Heritage Area with descriptions of new material ofGanguroo bilamina Cooke, 1997 and a new species

K.J. Travouillon, B.N. Cooke, M. Archer, and S.J. Hand

ABSTRACT

The relationship of basal macropodids (Marsupialia: Macropodoidea) from theOligo-Miocene of Australia have been unclear. Here, we describe a new species fromthe Bitesantennary Site within the Riversleigh’s World Heritage Area (WHA), Ganguroobites n. sp., new cranial and dental material of G. bilamina, and reassess material pre-viously described as Bulungamaya delicata and ‘Nowidgee matrix’. We performed ametric analysis of dental measurements on species of Thylogale which we then used,in combination with morphological features, to determine species boundaries in thefossils. We also performed a phylogenetic analysis to clarify the relationships of basalmacropodid species within Macropodoidea. Our results support the distinction of G. bil-amina, G. bites and B. delicata, but ‘Nowidgee matrix’ appears to be a synonym of B.delicata. The results of our phylogenetic analysis are inconclusive, but dental and cra-nial features suggest a close affinity between G. bilamina and macropodids. Finally, werevise the current understanding of basal macropodid diversity in Oligocene and Mio-cene sites at Riversleigh WHA.

K.J. Travouillon. School of Earth Sciences, University of Queensland, St Lucia, Queensland 4072, Australia and School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia. [email protected]. Cooke. Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101, Australia. [email protected]. Archer. School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia. [email protected]. Hand. School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia. [email protected]

Keywords: Macropodoidea; Miocene; Riversleigh; Kutjamarpu; new species; marsupials

PE Article Number: 17.1.20ACopyright: Palaeontological Association April 2014Submission: 17 May 2013. Acceptance: 11 March 2014

Travouillon, K.J., Cooke, B.N., Archer, M., and Hand, S.J. 2014. Revision of basal macropodids from the Riversleigh World Heritage Area with descriptions of new material of Ganguroo bilamina Cooke, 1997 and a new species. Palaeontologia Electronica Vol. 17, Issue 1;20A; 34p; palaeo-electronica.org/content/2014/711-riversleigh-basal-macropodoids

http://www.zoobank.org/3529571D-961B-4D09-9656-DA287F8A6E3C

TRAVOUILLON ET AL.: RIVERSLEIGH BASAL MACROPODOID

INTRODUCTION

Archer (1979) described the first macropodoidfrom the Riversleigh World Heritage Area (WHA),Wabularoo naughtoni, on the basis of a single den-tary found at D Site. It differed from potoroines inhaving bilophodont molars, and from macropodidsin having an enlarged masseteric canal, but shareswith both a tall and wide plagiaulacoid p3. Flanneryet al. (1983) identified an additional left p3 of W.naughtoni from the Riversleigh’s G Site anderected a new potoroid (their potoroines) subfam-ily, ‘Bulungamayinae’, to accomodate W. naughtoniand a new species, Bulungamaya delicata. The lat-ter was described from damaged, heavily worndentaries found at D Site and G Site. It is smallerthan W. naughtoni, has less well-developedlophids (although heavily obscured by wear) and aposthypocristid at least on m3 (their m4; othermolars were too worn to tell). Case (1984) andWoodburne (1984) suggested B. delicata and W.naughtoni represent macropodids (their macropod-ines) rather than potoroids, but Flannery et al.(1984) provided further evidence for their originalassignation based on an additional dentary of W.naughtoni from D Site.

Cooke (1997a) described two new macropo-doid species from Riversleigh WHA, ‘Nowidgeematrix’ and Ganguroo bilamina. ‘Nowidgee matrix’is clearly bunolophodont (sensu Flannery et al.,1984), with the buccally-directed cristid from theentoconid forming the hypolophid. The cristid obli-qua is also well-developed in this species andascends to the apex of the protoconid. In contrast,G. bilamina is completely bilophodont, the hypolo-phid crest being formed entirely by the much ele-vated posthypocristid, the posterior cingulid beinglost and no trace remains of a buccal cristid associ-ated with the entoconid apex. Cooke (1997a) alsodiscussed differences between Bulungamaya deli-cata, ‘N. matrix’ and G. bilamina. Additional mate-rial referable to B. delicata was reported from anumber of Riversleigh local faunas (LFs) in FaunalZones B and C (Cooke, 1997b; Archer et al.,2006). Some of these specimens have been usedin phylogenetic analyses (Kear and Cooke, 2001;Kear et al., 2001a; Kear et al., 2007; Kear andPledge, 2008). However, none of these specimenshave been described. We also question theirassignment to B. delicata. Upper molars with aclearly bunolophodont morphology have been con-fidently assigned to N. matrix (Cooke, 1997a) butupper molars of B. delicata and G. bilamina haveyet to be recognised. Considerable numbers ofmaxillae and even partial skulls of small macropo-

dids of a size range appropriate for either B. deli-cata or G. bilamina have now been recovered fromRiversleigh. Unfortunately none of these involvesunambiguously associated upper and lower denti-tions. Similarities in morphology of associatedspecimens (skulls and dentaries) from a smallmacropodid species from Faunal Zone C, Gan-guroo sp. 2 (Cooke, 1997b; Archer et al., 2006),suggest that all maxillae and partial skulls recov-ered to date from Faunal Zone B are likely tobelong to G. bilamina. These specimens aredescribed in the current paper. Previous phyloge-netic analyses have also used postcranial charac-ters for G. bilamina based on specimens from themiddle Miocene AL90 Site (Kear et al., 2001a).However, these postcranials were found in associ-ation with unnamed species Ganguroo sp. 2, not G.bilamina. In view of previous confusion and thedescription here of additional materials thatincrease understanding about basal macropodidtaxa, we have conducted a new phylogenetic anal-ysis for B. delicata, N. matrix, G. bilamina and G.bites n. sp. from Bitesantennary LF. Finally, wereassess the diversity of taxa currently known fromthe Riversleigh WHA.

MATERIALS AND METHODS

Materials and Dental Measurements

Specimens of Bulungamaya delicata, ‘Nowid-gee matrix’, Ganguroo bilamina and G. bites n. sp.used in this study were collected from fossil sites inthe Riversleigh World Heritage Area, northwesternQueensland (18°59'-19°08'S, 138°34'-138°43'E).Specimens with the prefix QM F are from theQueensland Museum fossil collection, Brisbane,Australia. One specimen of G. bilamina in the fossilcollections of the University of California at Berke-ley (UCMP) was recovered from the Wipajiri For-mation at the Leaf Locality, Lake Ngapakaldi,South Australia. Other specimens from the Eta-dunna Formation of South Australia in the collec-tions of the South Australian Museum (SAM P)used in metric analyses represent Ngamarooarcheri (Kear and Pledge, 2008) and Purtia mosa-icus (Case, 1984). Specimens of the modern Red-legged Pademelon (Thylogale stigmatica) andRed-necked Pademelon (Thylogale thetis) wereexamined from the Queensland Museum mammalcollection (QM J or JM).

All specimens were measured using digitalvernier callipers. Maximum anteroposterior lengthand buccolingual width measurements of decidu-ous and adult premolars were made at the base of

2

PALAEO-ELECTRONICA.ORG

the crown. Width measurements for molars weremeasured across the anterior and posterior lophs/lophids. All specimens examined and their mea-surements are provided in Appendices 1, 2, 3, 4, 5,6 and 7.

Metric Analysis

All metric analyses were performed using thecomputer software PAST Version 1.51 (Hammer etal., 2001). To assess the amount of variation inupper and lower premolars and molars in modernmacropodoids, basic univariate statistics were cal-culated for Thylogale stigmatica and Thylogale the-tis. These two taxa were chosen because they arecongeneric, have very similar dental morphologyand are sympatric. Intraspecific dental variation inthese species may help to illuminate speciesboundaries within the fossil samples. In order toassess the potential for failure to recognise distinctspecies in the fossil samples, basic univariate stat-ics were calculated for combined measurements ofThylogale stigmatica + Thylogale thetis. Bivariateplots of tooth measurements (length versus ante-rior or posterior width), for males and females ineach taxon were also generated. Principle Compo-nent Analyses (PCAs) were performed using thesame measurements, log transformed, to assesswhich dental measurements appeared to distin-guish these two taxa. A Multivariate Analysis ofVariance (MANOVA) was performed to testwhether the means of the log transformed dentalmeasurements were statistically different betweensexes of each species and between species. ACanonical Variates Analysis (CVA) was also usedto produce a scatter plot of specimens along thetwo canonical axes, producing maximal and sec-ond to maximal separation between all groups(multigroup discriminant analysis), which is thenable to reclassify specimens. This method is usedhere as a way to test whether fossil specimenswould be able to correctly reclassify with the cor-rect taxon using log transformed dental measure-ments.

Basic univariate statistics were also calcu-lated for G. bilamina (G. bites sample size too smallto include) and B. delicata + N. matrix. Bivariateplots, PCAs, MANOVAs, and CVAs were repeatedfor all fossil taxa listed above.

Terminology

Cheek-tooth homology follows Flower (1869)and Luckett (1993). Dental terminology followsArcher (1984), Tedford and Woodburne (1987),Szalay (1969) and Cooke (1997a, 1997b). System-

atic nomenclature follows Aplin and Archer (1987)and Kear and Cooke (2001) for higher-level marsu-pial relationships. Cranial terminology followsWible (2003) and Murray (1989).

Biostratigraphic nomenclature follows Archeret al. (1989, 1997), Creaser (1997), Arena (2004)and Travouillon et al. (2006, 2011).

Phylogenetic Analyses

We assessed the phylogenetic relationshipsof basal macropodid species to other macropo-doids using the matrix of Kear and Pledge (2008).While the matrix of Prideaux and Warburton (2010)is more recent, they acknowledged that their princi-ple purpose was to investigate the relationship ofbilophodont macropodids, and therefore not suit-able to analyse more basal forms. They mentionedthat their matrix would require an extensiveincrease in characters and additional outgroupsbefore it could be used for that purpose. Hencewhy we used the matrix of Kear and Pledge (2008),because it was explicitly generated for assessingthe relationship of basal macropodiforms. Theirmatrix contained 108 qualitative morphologicalcharacters including cranial, dental and postcranialfeatures. We removed skull and upper dentitionscores for Wabularoo naughtoni because materialscored previously was unpublished and may repre-sent a different species. Only the holotype of thatspecies, QM F9177 (Archer, 1979) which is a den-tary containing p3 m1-3, was used to score charac-ters. We scored the new specimens of Ganguroobilamina, which included cranial and upper denti-tion characters, and removed postcranial scoresfor G. bilamina because the postcranials describedby Kear et al. (2001a) belong to a different speciesof Ganguroo found in Riversleigh’s Faunal Zone C(see Ganguroo sp. 2 in Archer et al., 2006). Sev-eral skulls and dentaries from the AL90 Site thatwere associated with postcranials will be describedshortly (Cooke et al., in review). The new Gan-guroo species from the Bitesantennary Sitedescribed in this paper was also scored and addedto the matrix. Finally, we deleted ‘Nowidgee matrix’from the matrix (see discussion) and rescoredBulungamaya delicata based only on the materialassigned to this species in the current paper.

A parsimony analysis was performed follow-ing Worthy et al. (2006) and Beck et al. (2008) withPAUP* 4.0b10 (Swofford, 2002) using a two-stagesearch strategy. An initial search comprising 1,000heuristic replicates, saving 10 trees per replicate,was followed by a second heuristic search withinthe saved trees. This method allows optimising the

3


heuristic search and minimising the number ofmost parsimonious trees. All most parsimonioustress produced were summarised as a strict con-sensus tree. Bootstrap values for each node werecalculated using 1000 bootstrap replicates of 10random addition sequence replicates and decayindices were also calculated using TreeRot.v3(Sorenson and Franzosa, 2007).

SYSTEMATIC PALAEONTOLOGY

Family MACROPODIDAE Gray, 1821Genus GANGUROO Cooke, 1997a

Type Species.- Ganguroo bilamina Cooke, 1997aAdditional Species.- G. bites n. sp. Revised Generic Diagnosis. Species of Gan-guroo differ from all hypsiprimnodontids and potor-oines by having bilophodont molars and m4 is notgreatly reduced in size relative to other molars.They differ from all macropodines and sthenurinesby having a combination of: a buccally expandedmasseteric canal; a sharp ventrally convex dentarymargin below m1-2; i1 with enamel confined to thebuccal surface and extensive on that surface; ven-tral and dorsal flanges present on i1; p3 and P3elongate with fine transcristids and a bulbous base;a small precingulid present anterobuccal to theparacristid; lower molars and dp3 bilophodont. Itdiffers from balbarids in lacking a posterior cingulidon any lower molars and m1 lacks a protostylid(Kear and Cooke, 2001).Remarks. A third species of Ganguroo, referred toin Cooke, 1997b (and consequently in Archer et al.,2006; Travouillon et al., 2006, 2009, 2011) as Gan-guroo sp. 2, has yet to be described. This speciesseems to be common but restricted to FaunalZones C and D Sites at Riversleigh.

Ganguroo bilamina, Cooke, 1997aFigures 1, 2, 3, 4

Revised Species Diagnosis. Ganguroo bila-mina is characterised by the following features: flatfrontal region; broad parietal-alisphenoid contact;alisphenoid contribution to ventral floor of second-ary foramen ovale; carotid foramen anterior to pos-terior margin of basisphenoid; upper caninespresent; P3 elongate with a bulbous base and 7cusps and associated transcristae (in more wornspecimens, the 7th and sometimes even the 6th

cusps/transcristae are worn off); upper molars anddP3 bilophodont; stylar cusp C present on all uppermolars but decreases in size posteriorly; protolophin upper molars formed mainly by the preprotoc-rista and always joins to preparacrista, but alsoforms a small forelink anteriorly; neometaconule

and associated crest (postlink) present on dP3 andall upper molars, decreasing in size posteriorly(presence may be affected by wear); and smallalveolus for i2 present directly posterior to i1.

Ganguroo bilamina differs from G. bites in:having 7 cuspids and associated transcristids (con-tra Cooke, 1997a) on p3 (instead of 8 in G. bites); ashorter and wider p3; smaller and narrower lowermolars that are rectangular in occlusal outline;paraconid absent; and a buccal crest from theentoconid in some juveniles (contra Cooke,1997a).Holotype. QM F19915, left dentary with i1, p3, m1-4. Referred Material. From Riversleigh WHA, Cad-bury’s Kingdom Site: QM F56318, left P3; CamelSputum Site: QM F19870, left dentary with m1-4,QM F19868, left dentary with i1, p3, m2-3, QMF19966, left dentary with p3, m1-4, QM F30400,left dentary with m2-4, QM F56996, left dentarywith dp3, p3, m1-3, m4 in crypt, QM F19607, leftdentary with p3, m1-3, QM F20248, right dentarywith dp2-3, m1, QM F20254, right dentary with m4,QM F23486, right dentary with worn m2-4, QMF20246, right dentary with p3, QM F20019, leftdentary with dp2, m1, QM F56994, left dentary withp3, QM F19903, left dentary with i1, dp2-3, p3, m1-3, QM F56254, right dentary with i1, dp2-3, p3, m1-3, QM F56259, right dentary with p3, m1-4, QMF24744, right dentary with dp2-3, m1-3, QMF56995, right maxilla with dP3, P3, M1, QMF19696, skull with right and left P3, M1-3, QMF19958, left maxilla with M1-3, QM F20705, leftmaxilla with P3, M1-2, broken M3, QM F19900, leftmaxilla with M1-4, QM F19695, left maxilla with P3,M1-2, QM F20253, right maxilla with P3, M1, bro-ken M2, QM F19861, left maxilla with dP2-3, M1-3,QM F19976, left maxilla with dP2-3, M1, QMF19957, left maxilla with broken M1, M2-4, QMF19857, right maxilla with dP2-3, M1-3, QMF19975, left maxilla with M1-3, QM F19959, leftmaxilla with M2-3, QM F20017, right maxilla withP3, M1-4, QM F56990, left maxilla with broken M2-3, M4, QM F30436, left maxilla with M1-3, QMF30343, left maxilla with dP2-3, M1-2, QM F36230,right maxilla with M1-3; Inabeyance Site: QMF20006, left dentary with dp2, p3, m1-3; Judith’sHorizontalis Site: QM F36351, right dentary with i1,p3, m1-4; Mike’s Menagerie Site: QM F19988, rightdentary with m3-4, QM F19844, left dentary withm1-2, QM F19668, right maxilla with M1-2, QMF19692, right maxilla with M1-4; Mike’s PotatoPatch Site: QM F234604, left maxilla with dP2, M1;Neville’s Garden Site: QM F24190, right dentary

4


5

FIGURE 1. Ganguroo bilamina skull (QM F56265) from Wayne’s Wok Site, in dorsal (1), right lateral (2) and ventral(3) views. (4) Line drawing of the upper right cheek teeth of QM F56265. Abbreviations: ac, anterior cingulum; as,alisphenoid; bo, basioccipital; bs, basisphenoid; cf, carotid foramen; eam, external auditory meatus; ec, ectotym-panic; fl, forelink; fm, foramen magnum; fr, frontal; gf, glenoid fossa; iof, infraorbital foramen; I1, I3, upper incisorone and three; inf, incisive fenestra; ip, interparietal; jf, jugular foramen; ju, jugal; lac, lacrimal; lacf, lacrimal fora-men; M1-M4, upper molar one to molar four; ma, mastoid; map, mastoid process; mcl, metaconule; me, metacone;ml, metaloph; mp, maxillopalatine fenestra; mx, maxilla; na, nasal; ne, neometaconule; oc, occipital condyle; os,orbitosphenoid; P3, upper third premolar; pa, parietal; pac, paracone; pal, palatine; pc, precingulum; pe, petrosal;pl, postlink; pmx, premaxilla; pomc, postmetacrista; pomclc, postmetaconulecrista; popc, postparacrista; poprc,postprotocrista; pr, protocone; prl, protoloph; prmc, premetacrista; prpc, preparacrista; ps, presphenoid; pt, ptery-goid; sfo, secondary foramen ovale; smf, suprameatal foramen; sof, sphenorbital fissure; sq, squamosal; St C-D,stylar cusp C and D. Scale bar equals 10 mm.


6

FIGURE 2. Juvenile maxilla of Ganguroo bilamina (QM F57020) in occlusal view (1), juvenile maxilla of Bulungamayadelicata/’Nowidgee matrix’ (QM F57024) in occlusal view (2), line drawing of upper cheek teeth of Ganguroo bilamina,QM F57020, (left), and Bulungamaya delicata/’Nowidgee matrix’, QM F57024 (right), (3) and juvenile dentary of Gan-guroo bilamina (QM F57022) in buccal (4), occlusal (5), lingual (6) views and line drawing of lower cheek teeth (7).Abbreviations: ac, anterior cingulum; co, cristid obliqua; dP2-3, deciduous upper premolar two to three; dp2-3,deciduous lower premolar two to three; ecd, buccal crest from entoconid; end, entoconid; fl, forelink; hl, hypolophid;hyd, hypoconid; i1, lower first incisor; m1-m3, lower molar one to molar three; M1-M2, upper molar one to molar two;mas, mandibular symphysis; mcl, metaconule; me, metacone; med, metaconid; mf, mental foramen; ml, metaloph;ne, neometaconule; p3, lower third premolar; pac, paracone; pacd, paracristid; pad, paraconid; pc, precingulum;pcd, precingulid; phc, posthypocristid; pl, postlink; pomc, postmetacrista; pomclc, postmetaconulecrista; popc,postparacrista; poprc, postprotocrista; pr, protocone; prenc, preentocristid; prd, protoconid; prl, protoloph; prld, pro-tolophid; prmc, premetacrista; prmcd, premetacristid; prpc, preparacrista; prprc, preprotocrista; St C, stylar cusp C.Scale bar equals 10 mm.


7

FIGURE 3. Lingual view of dentaries of: 1, Bulungamaya delicata (holotype, CPC 22187) from G Site; 2, ‘Nowidgeematrix’ (QM F19991) from White Hunter Site; 3, ‘Nowidgee matrix’ (QM F19937) from Wayne’s Wok Site; and 4, Gan-guroo bilamina (QM F57021) from Wayne’s Wok Site (reversed for convenience of comparison). Scale bar equals 10mm.


8

FIGURE 4. Occlusal view of dentaries of and associated line drawings: 1 and 5, Bulungamaya delicata (holotype,CPC 22187) from G Site; 2 and 6, ‘Nowidgee matrix’ (QM F19991) from White Hunter Site; 3 and 7, ‘Nowidgeematrix’ (QM F19937) from Wayne’s Work Site; and 4 and 8, Ganguroo bilamina (QM F57021) from Wayne’s WorkSite (reversed for convenience of comparison). Abbreviations: co, cristid obliqua; ecd, buccal crest from entoconid;end, entoconid; hl, hypolophid; hyd, hypoconid; mcl, metaconule; med, metaconid; pacd, paracristid; pad, paraco-nid; pcd, precingulid; phc, posthypocristid; pomcd, postmetacristid; prenc, preentocristid; prd, protoconid; prld, pro-tolophid; prmcd, premetacristid. Scale bar equals 10 mm.


with broken m2, m3, QM F19987, left maxilla withM1-3, QM F23202, left maxilla with M1-4; RossScott-Orr (RSO) Site: QM F20039, right dentarywith m2-3, QM F20269, right maxilla with M1, bro-ken M2; Upper Site: QM F30396, left dentary withi1, p3, m1-3, QM F19642, left dentary with p3, m1,QM F30397, right dentary with m1-4, QM F20293,right dentary with m2-4, QM F56999, right dentarywith dp2-3, p3, m1-2, QM F20271, right dentarywith m4, QM F19643, left dentary with damagedm1-4, QM F20295, right dentary with broken m1,m2-3, QM F19947, right dentary with dp2-3, p3,m1, QM F56991, left dentary with dp2-3, p3, m1-2,QM F20272, left dentary with m3, m4 in crypt, QMF19640, left dentary with dp2-3, p3, m1, QMF19646, right dentary with m3, QM F19688, leftdentary with i1, dp2-3, m1-3, QM F56993, left max-illa with P3, M1-4, QM F19801, posterior half of askull, QM F19673, left maxilla with dP3, M1-3, M4in crypt, QM F56992, right maxilla with P3, M1-3,QM F19985, right maxilla with M1, QM F19885,right maxilla with P3, M1-3, QM F19687, right max-illa with dP2, M1-4, QM F19651, right maxilla withP3, M1, QM F19689, right maxilla with M1-4, QMF19629, right maxilla with dP2-3, M1, QM F19674,left maxilla with P3, M1-4; Wayne’s Wok Site: QMF19814, right dentary with p3, m1-2, QM F30399,right dentary with p3, m1-4, QM F19591, left den-tary with m3, m4 in crypt, QM F19810, left dentarywith m1-4, QM F19835, left dentary with dp2-3, p3,m1, QM F57022, left dentary with i1, dp2-3, p3,m1-3, QM F19836, left dentary with i1, QMF57021, right dentary with i1, p3, m1-4, QMF56998, left dentary with i1, m1-4, QM F19834,right dentary with p3, m1-2, QM F19827, left den-tary with broken m1, very worn m2-4, QM F20082,left dentary with m4, QM F19597, left dentary withp3, m1-4, QM F24480, right dentary with m1-2, QMF56997, left dentary with p3, m1-2, QM F20108,left dentary with dp3, p3, m1, QM F20109, rightdentary with p3, QM F19602, left dentary with m1-3, QM F50519, left dentary with i1, p3, m1-4, andassociated right dentary with p3, m1-4, QMF24050, left dentary with i1, p3, m1-3, QM F30721,left dentary with p3, m1-2, broken m3, QM F20081,right dentary with i1, p3, m1-4, QM F36364, rightdentary with i1, dp2-3, m1-2, QM F24477, left den-tary with m4, QM F30398, right dentary with p3,m1-4, QM F56257, left dentary with p3, m1-4, QMF24482, left dentary with m2-3, QM F31405, leftdentary with p3, m1-4, QM F56260, left dentarywith m1-4, QM F36363, right dentary with m3-4,QM F36470, left dentary with broken p3, m1-4, QM

F56264, right dentary with p3, m1-3, QM F52814,Skull with left M3-4, right P3, M1-4, QM F57020,associated left and right maxilla with dP2-3, M1-2,QM F20261, left maxilla with dP2-3, P3, M1-3, QMF19589, left maxilla with M1-4, QM F19617, rightmaxilla with P3, M1-2, QM F19590, left maxilla withP3, M1-4, QM F24219, right maxilla with dP3, P3,M1-3, QM F20163, left maxilla with M1-3, QMF56256, right maxilla with dP3, M1-3, QM F56258,right maxilla with P3, M1-4, QM F56265, Skull withleft I1, and left and right I3, P3, M1-4, QM F24475,right maxilla with M1-2, QM F24476, right maxillawith dP3, M1-2, QM F50518, right maxilla with M3-4, QM F56261, right maxilla with M1-2, QMF56255, skull with left and right P3, M1-4, QMF24216, left maxilla with dP2-3, M1-2, QM F56263,right maxilla with P3, M1-2; Kutjamarpu LocalFauna, Leaf Locality, Wipajiri Formation: UCMP88221, right dentary with i1, p3, m1-4.Age and Distribution. The holotype is fromWayne’s Wok Site, D-Site Plateau, RiversleighWorld Heritage Area, northwestern Queensland,Australia (18°15'35"S, 138°06' 41"E). Paratypesand referred specimens are from Riversleigh’sCadbury's Kingdom Site, Camel Sputum Site,Inabeyance Site, Judith's Horizontalis Site, Mike'sMenagerie Site, Mike's Potato Patch Site, Neville'sGarden Site, RSO Site, Upper Site, Wayne's WokSite, and the Leaf Locality in the Wipajiri Forma-tion, Lake Ngapakaldi, South Australia. All the Riv-ersleigh site LFs are interpreted to be part ofRiversleigh’s Faunal Zone B as distinguished byArcher et al. (1989, 1997), Arena (2004) andTravouillon et al. (2006, 2011), except Cadbury'sKingdom LF which is thought to be part of FaunalZone C, but see discussion below. They are inter-preted to be early Miocene in age on the basis ofbiocorrelation of their contained faunas (Travouil-lon et al., 2006, 2011). The Leaf Locality (Kutja-marpu Local Fauna) is also interpreted to be earlyto middle Miocene in age (Megirian et al., 2010;Metzger and Retallack, 2010).Remarks. The original description of Ganguroo bil-amina was based on dentaries and lower dentitiononly. Additional specimens are described below,including a near complete skull, the completeupper dentition and a juvenile upper and lowerdentition. Isolated postcranial elements from AL90Site referred to G. bilamina by Kear et al. (2001a)have since been found associated with a skull anddentaries referable to Ganguroo sp. 2 rather thanG. bilamina.

9


Description

Nasal. Anteriorly, the nasals are level with theanterior part of the premaxilla (Figure 1). The nasalsutures are almost parallel anteriorly but widenposteriorly in the premolar region. The widest partof the nasals occurs above the most anterior partof the lacrimal. The dorsal surface of the rostrum isventrally curves. The nasal-premaxillary suture isshort, half the length of the nasal-maxillary suture.The nares are wider than tall.Premaxilla. The premaxilla is taller than it is long(Figure 1). The most posterior part of the postero-dorsal process of the premaxilla is a thin flangethat posteriorly contacts both the nasal and themaxilla. From this point, the maxillary-premaxillarysuture is almost vertical, and ends ventrally at thecanine alveolus, which is bordered anteriorly by thepremaxilla and posteriorly by the maxilla. Thecanine is not preserved but the alveolus is consid-erably larger than the incisive alveoli. Anterior tothe canine alveolus, three incisive alveoli are pres-ent (with I1 preserved on the left side, and I3onboth sides). The alveolus for I3 is ventrallydirected, while the alveoli for I1 and I2 are antero-ventrally directed. The alveolus for I1 is almostthree times the size of the alveoli for I2 and I3. Twothin incisive fenestrae are present in the ventralplate of the premaxilla. The most anterior part ofthe incisive fenestrae is level with the posterior endof the alveolus for I2. The posterior end of the inci-sive fenestrae is bordered by the maxilla. Maxilla and Palatine. The maxillopalatine fenes-trae are broad, boarded anteriorly by the maxillaand by the palatine laterally at the level of the mid-point of M2, and posteriorly to that point (Figure 1).The maxillofrontal suture is almost horizontal,crossing the facial region just superior to the levelof the superior lacrimal foramen. The infraorbitalforamen is vertically elliptical, anteriorly directedand located well anterior to the orbital marginabove the anterior end of P3. There is little devel-opment of the masseteric process of the maxilla,which barely reaches as far ventrally as the alveo-lar margin which is a normal condition for potoro-ines (Kear and Cooke, 2001). The suborbital shelfof the maxilla is relatively flat dorsally, broaderanteriorly, tapering posteriorly and terminating justposterior to the anterior end of the sphenorbital fis-sure. Small foramina for the passage of the supe-rior alveoli nerves and vessels open within narrowsulci on the lateral margin of the suborbital shelf.There is a deep infraorbital fossa within the anteriorlateral corner of the suborbital shelf and a smaller,more shallow fossa medial to this and superior to

the maxillary foramen. The circular opening of thisforamen is overhung dorsally by a shelf posterior tothe smaller of the two infraorbital fossae and themargin of the shelf continues as a ridge extendinghorizontally across a dorsal wing of the maxilla,through the maxilla-palatine suture and extendingonto the palatine dorsal and posterior to the sphe-nopalatine foramen located posterior to the maxil-lary foramen. The sphenopalatine foramen is onlyjust larger than the maxillary foramen and oval inshape. The palatine forms the ventral mesial wallof the orbit. Lacrimal. The lacrimal extends anteriorly onto thefacial region and dorsally beyond the maxillofrontalsuture (Figure 1). There is a large inferior lacrimalforamen on the facial region, a superior foramenopening posteriorly on the anterior orbital marginjust dorsal to a small prominence. The broad orbitalwing of the lacrimal forms the anterior mesial wallof the orbit.Frontal, Parietal and Interparietal. Anteriorly, theoverlapping joint between the frontal and the nasalcrosses transversely from the anterior of the maxil-lofrontal suture to the frontal/frontal suture (Figure1). The frontal region is broad in the interorbitalregion, contrasting markedly with the constrictionof this region seen in balbarines, and is remarkablyflat. Strong supraorbital crests on the lateral mar-gins of the flat dorsal portion of the frontals con-tinue posteriorly along the lateral, dorsal margin ofthe parietal as sagittal crests, converging on themidline on the posterior dorsal surface of the neu-rocranium. Here they form a single sagittal crestthat continues posteriorly through the overlappinginterparietal sutures. The interparietal and supraoc-cipital sutures form a tall crest that ends at thejunction of the parietals and the supraoccipital. Therostrum is very broad anterior to the orbital regionbut does not exhibit the marked inflation seen inthis region in balbarines (Cooke, 2000; Kear et al.,2007). The parietals are laterally flared at their pos-terior ends and curve gently dorsally, indicating awell-developed nuchal crest. There is a clear,broad contact between the alisphenoid and parietalon the lateral wall of the neurocranium. While shortparietal-alisphenoid contact is known to occuramong Riversleigh macropodoids, e.g., in Hyp-siprymnodon bartholomaii (Flannery and Archer,1987a) and Bettongia moyesii (Flannery andArcher, 1987b), the condition seen here is muchmore extensive and far more resembles that seenin macropodoids rather than that in potoroines andhypsiprymnodontids (Kear and Cooke, 2001).

10


Zygomatic arch. The jugal ends anteriorly at thelevel of the anterior margin of the lacrimal (Figure1). Its lateral surface has a crest arising anterior tothe masseteric process and running posteriorlybelow the orbit to the posterior end of the jugal.The crest marks the anterior and dorsal limit ofattachment of the superficial layer of the masseter.The orbital part of the jugal is overhung anteriorlyby the ventral margin of the orbit. The zygomaticarch makes a smooth transition to the facial region,there being no antorbital sulcus, absence of thisfeature being the condition in potoroines. The mostanterior part of the jugal-squamosal suture is V-shaped, with the jugal extending dorsally as well asdorsoventrally. The posterior end of the jugal ven-trally overlaps the zygomatic process of the squa-mosal and forms the anterior margin of the glenoidfossa. The zygomatic process of the squamosal iswide transversely at its posterior end, surmountedby a broad, shallow sulcus. The process is rela-tively shallow and has a flat lateral surface. Theglenoid fossa is broad and flat and mergessmoothly with the ventral surface of the zygomaticprocess, which projects anteriorly for a consider-able distance. There is a well-developed pneuma-tised postglenoid process posterior to the glenoidfossa. Neurocranium. The parietals form most of the roofand walls of the neurocranium, having a broad lat-eral contact with the squamosals (concave inshape) which form the posteroventral lateral walls(Figure 1). The anteroventral component of theneurocranial wall is contributed by the temporalwing of the alisphenoid. The anteriorly facing fora-men rotundum is visible at the anteroventral mar-gin of the temporal wing of the alisphenoid, lateralto the sphenorbital fissure (foramen rotundum isbetter preserved in QM F19801). Retained suturelines between parietals and the supraoccipital indi-cate that the later contributed significantly to theposterior roof of the neurocranium.Basicranium.The basioccipital is roughly hexago-nal in outline with a relatively broad anterior contactwith the basisphenoid (Figure 1). Paired foraminalie within fossae overhung by the ventral margin ofeach of the occipital condyles. Very short canalsleading from these foramina open posteriorly withinthe foramen magnum. The most anterior of theseforamina opening on the ventral surface of thebasioccipital are presumed to represent the hypo-glossal foramina, the more posterior foraminabeing the condylar foramina. The jugular foraminalie lateral to the condylar foramina, posterior to thealisphenoid tympanic wing and mesial to the ante-

rior ends of the paracondylar process of the exoc-cipital. The foramina for the greater petrosal nervesare mesial to the alisphenoid tympanic wing, pos-teromesial to the secondary foramina ovale andclosely posterolateral to the carotid foramina. Thealisphenoid tympanic wings are relatively flat andpoorly inflated and terminate posteriorly at thebase of the short, laterally compressed paracondy-lar process of the exoccipital. The secondary fora-men ovale opens on the anteromesial side of thealisphenoid tympanic wing and is floored by a thin,very narrow process of the alisphenoid tympanicwing that separates it from the foramen for thegreater petrosal nerve which lies posteroventral tothe secondary foramen ovale. Contribution of aventral floor to the secondary foramen ovale by aprocess of the alisphenoid was regarded by Murray(1991) to be typical of macropodoids, contrastingwith the incomplete ventral floor provided for thesecondary foramen ovale by the process of the ali-sphenoid which is the usual condition in potoro-ines. The carotid foramen is anterior to theposterior margin of the basisphenoid in QMF56265 and QM F19801, but more posteriorlylocated and lateral to the basisphenoid-basioccipi-tal suture in QMF 52814. Murray (1991) suggestedthat an anterior position for the carotid foramenwas a potoroine synapomorphy. The transverseforamen of the basisphenoid in QM F56265 andQM F19801 is single with a shallow sulcus, pre-sumably for the Eustachian tube where it crossesthe ventral border. In QMF 52814, the transverseforamen is incompletely divided by dorsal and ven-tral spurs, and there is no sulcus ventral to it.

The pterygoid fossae are oval in outline withventrolaterally sloping mesial walls provided by thepterygoid and an anterior wall provided by the ali-sphenoid. The internal surfaces of the pterygoidfossa are relatively smooth.

The posterior vertical face of the occipitaldoes not exhibit obvious muscle attachment scarsfor the neck musculature. The foramen magnum isteardrop-shaped. The width of the posterior sur-face of the skull is extended by the vertically elon-gate mastoids.Auditory Region. The ectotympanic is roughlysquare in ventral outline and concave in its centralregion (Figure 1). The external auditory meatus isnot enclosed dorsally by the ectotympanic but isclosed by a thin plate of the squamosal. The smallstylo-mastoid foramen is located immediately ante-rior to the ectotympanic/mastoid suture. The mas-toid process is relatively long, more massive than

11


the paracondylar process of the exoccipital andinclined ventrolaterally.

The articular eminence of the glenoid fossa isroughly square. The postglenoid foramen is mesialto the low triangular postglenoid process and ispartially divided by a spur of the ectotympanic fromthe smaller and more mesial residual Gasserianfissure. A large opening into the epitympanic sinusposterior to the postglenoid process represents thezygomatic epitympanic sinus of the squamosalwhich is enclosed entirely by the squamosal.

The lateral wall of the petrosal is visiblethrough the suprameatal foramen, on the posteriorlateral wall of the squamosal, dorsal to the exit ofthe auditory meatus. The dorsal opening of thetemporal canal lies within this opening, anterior tothe visible portion of the petrosal. The cerebellarface of the petrosal (visible in QM F19801) moreclosely resembles that of Macropus than Potorousor Hadronomas in so far as these are illustrated byMurray (1991). The parafloccular fossa is deep,undercutting the edges of the orifice. The endolym-phatic duct opens on the cerebellar face of thepetrosal, as it does in M. robustus (Murray, 1989).Upper Dentition. The description which follows isbased primarily on the upper dentition preserved inthe skull (Figure 1), with variation evident in otherspecimens noted where appropriate.

The chisel-like I1 is the largest upper incisor. Ithas an oval occlusal outline but in buccal view it isstrongly anteriorly curved. Its alveolus is antero-ventrally orientated. The I2 is absent in QM F56265but its anteroventrally-orientated alveolus is small,circular in shape and more ventrally situated thanthe alveolus of I1. The I3 is oval in occlusal outlinewith a central bladed crown. It would have beenlarger than I2 but it is smaller than I1. While itsalveolus faces ventrally, the crown of I3 is antero-ventrally orientated.

In occlusal view both cheek teeth rows areslightly buccally convex. The long axes of the pre-molars are aligned with the molar row. In lateralview the posterior molars are more ventrally situ-ated than the anterior molars and the occlusal mar-gins of the premolars project beyond those of theadjacent molars. Molars are low-crowned, bilopho-dont and larger in the middle of the row than at itsends.

dP2 is preserved in QM F19857, F19861,F19976 and in both tooth rows of F57020 (Figure2). The tooth is short, blade-like and slightly longerthan dP3. The occlusal margin is subhorizontal andshorter than the crown base. It begins posterior tothe anterior margin of the crown base and extends

to the level of the posterior margin of the tooth. Thelingual face of the crown is steeper than the buccalface. Three minor cuspules anterior to a longercusp are present on the occlusal crest and all fourcuspules support transcristae. Some lingual por-tions of these transcristae are obscured by wear insome specimens. A strong posterior crest runsfrom the apex of the posterior cuspule and sweepslingually as it approaches the crown base. In QMF57020, an additional posterior, lingual transcritaruns parallel to and just anterior to this posteriorcrista.

dP3 is preserved in QM F56995, F19857,F19861, F19976, F57020 and F20261 (Figure 2).The tooth has a straight buccal margin and a con-vex lingual margin which may be constricted at thelevel of the interloph. The occlusal surface is domi-nated by a longitudinal ridge continuing the line ofthe occlusal blade of dP2 and formed by the promi-nent, laterally compressed parastyle, similarlycompressed paracone and metacone and theirassociated pre- and postcristae. The postparac-rista and premetacrista form a continuous centroc-rista that is interrupted by a narrow interloph cleft inQM F57020. This tooth is bilophodont, with apoorly developed protoloph and metaloph. Aneometaconule and associated crest (postlink) arepresent at the midpoint of the metaloph.

P3 is a long and relatively low blade-like toothwith an occlusal edge that is horizontal over mostof its length but has a taller buccally-orientatedposterior ‘heel’ (Figure 1). In occlusal view thecrown base has a sub-elliptical outline but isslightly constricted just anterior to the posterior‘heel’. There is a very well-developed lingual ridgebuttressing the posterior end of the occlusal blade.A shelf-like lingual cingulum extends from the ante-rior margin to the posterior lingual ridge, and theexpanded buccal crown base forms a relativelyrounded buccal cingulum. There are six cuspuleson the occlusal margin anterior to the posterior‘heel’, each having associated transcristae thatextend to the crown base. The number of cuspulesvaries in other specimens from five to as many asseven (e.g., in QM F20253). The variation in thenumber of cuspules and transcristae is caused bydental wear. A crest extends anteriorly from theapex of the most anterior cuspule to the crownbase. A better developed but shorter crest is asso-ciated with the posterior surface of the posterior‘heel’.

M1 is almost square in occlusal outline, theprotoloph and metaloph being subequal in length(Figure 1). Lingual cusps are more massive and

12


have more rounded apices than those of the buc-cal side. The buccal surfaces of the crown slopemore steeply than the lingual surfaces. The shortpreparacrista slopes very slightly lingually from theapex of the paracone to the anterior margin of thecrown. The protoloph is formed by a lingually slop-ing crest from the paracone which meets an anter-olingually-inclined preprotocrista at a low point onthe approximate midline of the loph. The preprotoc-rista continues anteriorly as a ‘forelink’, although itsprominence has been reduced by wear. An anteriorcingulum extends across the buccal anterior mar-gin from the preparacrista to the ‘forelink’, and aprecingulum extends from there to the lingual mar-gin. The postparacrista slopes directly posteriorlyfrom the paracone apex to the interloph valleywhere it is linked to the curved premetacristathereby forming a continuous centrocrista. A shortridge on the posterobuccal flank of the paraconeconnects to a distinct stylar cusp C which occupiesthe anterior half of the buccal end of the interlophvalley. Stylar cusp D is not developed but a stylarcrest on the anterior buccal face of the metaconelinks the premetacrista to the base of StC. Stylarcusp C is reduced in size in some specimens, andthe crest representing StD similarly reduced. Thepostprotocrista slopes posterobuccally to the inter-loph valley and extends almost to the occlusal mar-gin of the metaloph, meeting it just buccal to themetaconule. There is a distinct prominence, theneometaconule, along the midline of the crest ofthe metaloph. A broad, rounded neometaconulecrista (the ‘postlink’) extends from thisneometaconule onto the posterior face of themetaloph. The postmetacrista extends from theapex of the metacone to the crown base where itmeets the postmetaconulecrista which sweepsacross the posterior face of the loph.

M2 is similar in shape to M1 except in the fol-lowing features (Figure 1): it is larger than M1 andwider anteriorly than posteriorly; stylar cusp C isnot as large; the postparacrista and premetacristaare flexed lingually in the interloph valley; and sty-lar crest D is not as well developed.

M3 is similar in shape to M2 except in the fol-lowing features (Figure 1): it is narrower posteri-orly; stylar cusp C is reduced to a stylar crest(absent in some specimens); the postparacristaand premetacrista sometimes fail to meet in theinterloph region; and the neometaconule and itscristae are not as well developed.

M4 is similar in shape to M3 except in the fol-lowing features (Figure 1): it is considerably nar-rower posteriorly; its metaloph is narrower than its

protoloph; and both stylar crests are absent fromthe interloph valley.Lower Dentition. The lower dentition is describedby Cooke (1997a). However, one of the specimensused in that description as dp2 and dp3 (QMF23777) have been referred here to G. bites. Fur-ther, QM F19385 is a worn and damaged specimenof G. bilamina. Additional juveniles of G. bilaminahave since been recovered (e.g., QM F57022, QMF56996, F20248, F20019, F19903, F56254,F24744, F20006, F56999, F19947, etc.). The mor-phology of these specimens is consistent with thedescriptions of the taxa provided by Cooke (1997a)except as follow (Figures 2, 3, 4): a buccal crestfrom the entoconid is present on dp3 and all lowermolars (decreasing in development posteriorly) insome juvenile (e.g., QM F57022, F56996, F20019,F56991) but absent in others (e.g., QM F19640,F19947, F20108). This buccal crest is absent in alladult specimens, possibly due to wear. Unworn p3reveal a total of seven cuspids and associatedtranscritids, instead of six described by Cooke(1997a). Wear tends to occur posteriorly, wearingdown the seventh cuspid and transcristid first, thenthe sixth and so on.

Ganguroo bites, n. sp.Figure 5

http://www.zoobank.org/C7E35201-45C6-4319-99F7-852743890585

Species Diagnosis. Ganguroo bites differs fromG. bilamina in the following features: it has eightcuspids and associated transcristids on p3 (insteadof 7 in G. bilamina); a longer but much narrower p3;larger and much wider molars that are almostsquare in occlusal outline; a paraconid present asa small cusp in m2-3; and no buccal crest from theentoconid.Species etymology. Named after the Riversleighfossil locality, Bitesantennary Site. Holotype. QM F23776, left dentary with p3, m1-3. Paratypes. QM F23775, right dentary with m1-2,m3 in crypt; QM F23777, juvenile left dentary withdp2-3, m1. Age and Distribution. The holotype and para-types are from Bitesantennary Site, D-Site Plateau,Riversleigh WHA, northwestern Queensland. Bite-santennary LF is interpreted to be part of River-sleigh’s Faunal Zone B as distinguished by Archeret al. (1989, 1997), Arena (2004) and Travouillon etal. (2006, 2011). It is interpreted to be early Mio-cene on the basis of biocorrelation of its containedfaunas (Travouillon et al., 2006, 2011).Remarks. QM F23777 was originally listed as aparatype of Ganguroo bilamina. In light of addi-

13


tional material found in the Bitesantennary Site,this specimen and others from the same siteexhibit morphological features that clearly distin-guish it from that species.

Description

The holotype is a left dentary with p3 to m3and alveoli for m4 (Figure 5). The dentary is incom-plete anterior to p3. The ventral margin of the hori-zontal ramus is convex and deepest below theroots of m2. No distinct digastric prominence ispresent. Two mental foramina are present on thebuccal margin of the dentary, one larger foramenjust anterior to p3, and one smaller foramen belowthe anterior root of m2.The ascending ramus is notpreserved in the holotype but it is almost com-pletely preserved in QM F23775 and partly pre-served in QM F23777. The angle formed by theocclusal margin and the ascending ramus is about120 degrees. The masseteric foramen is large anddeep, extending at least below the anterior region

of m3. Posteriorly, the masseteric foramen is con-fluent as a large oval opening with the mandibularforamen in the holotype, but is confluent as a smalloval opening with the mandibular foramen in QMF23775 and QM F23777. This difference may belinked to age; QM F23775 and QM F23777 arejuveniles while the holotype is an adult. The con-dyle, present in QM F23777, is broad and ellipticalin dorsal view.

The dp2 and dp3 are preserved in QMF23777 (Figure 5). The dp2 crown is plagiaulacoidin form and slightly longer but narrower than m1.The occlusal margin is shorter than the crownbase. The buccal face of the crown is less steepthan the lingual face. Four cuspids are presentanterior to a much larger posteriorly positionedcuspid. Transcristids are associated with the fouranterior cuspids only.

The dp3 is a molariform tooth, shorter butwider than dp2 (Figure 5). It is almost rectangularin shape with the buccal margin concave and the

FIGURE 5. Ganguroo bites n. sp., holotype QM F23776, from the Bitesantennary Site, left dentary in lingual (1), buc-cal (2) and occlusal (3) views; Paratype QM F23775, right dentary in buccal (4), lingual (5) and occlusal (6) views;and Paratype QM F23777, left dentary in buccal (7), occlusal (8) and lingual (9) views. Abbreviations: ang, angularprocess; dp2-3, deciduous lower premolar two to three; i1, lower first incisor; m1-m3, lower molar one to molarthree; maf, masseteric fossa; mafo, masseteric foramen; manf, mandibular foramen; mas, mandibular symphysis;mf, mental foramen; p3, lower third premolar; psmf, posterior shelf of the masseteric fossa. Scale bar equals 10mm.

14


lingual margin convex. The trigonid is laterallycompressed. The protoconid is the tallest cusp onthe crown, followed by paraconid, entoconid andhypoconid. No distinct metaconid is present. Theparacristid is anteroposteriorly orientated, while themetacristid is posterolingually orientated. The ento-conid is situated posterolingual to the protocone.No distinct preentocristid is present. The entoconidis joined to the hypoconid laterally by the hypolo-phid. The cristid obliqua extends from the hypoco-nid anterolingually and ends in the interlophidvalley, posterobuccal to the protoconid.

The p3 is represented in the holotype (QMF23776) and the paratype (QM F23777; Figure 5).In the latter specimen, p3 is erupted but the poste-rior alveoli for dp2 and dp3 are still visible, and thep3 blade is not aligned with the molar row. Thisarrangement indicates that the specimen may rep-resent a young adult (the m4 alveoli indicate that itwas fully erupted) in which p3 had not yet comeinto alignment with the molars (dp2 and dp3 arealigned with the molars in QM F23777). The p3 is along, plagiaulacoid-like tooth, about one and a halftimes the length of m1, but narrower than the ante-rior width of m1. The p3 width is greater anteriorlythan posteriorly. Eight cuspids are present alongthe occlusal edge of the tooth. The seven smalleranterior cuspids have associated transcristids butthe large posterior cuspid does not.

The m1 is subrectangular in occlusal outline,but wider posteriorly than anteriorly (Figure 5). Thehypoconid is the tallest cuspid, followed by theentoconid, metaconid and protoconid in decreasingheight. The m1 is bilophodont. The protoconid isconnected to the metaconid lingually by a nearlystraight protolophid. A short premetacristid extendsanteriorly to the anterior margin of the tooth. Theparacristid extends from the protoconid anterolin-gually to join the premetacristid anteriorly therebyenclosing a broad anterior cingulum. No distinctparaconid is present. A small precingulid is presenton the anterobuccal flank of the paracristid. Theentoconid is directly posterior to the metaconid andis joined to the hypoconid by a concave hypolo-phid. A preentocristid extends anteriorly from theentoconid and terminates in the interlophid valleydirectly posterior to metaconid. The cristid obliquaruns from the hypoconid anterolingually and endsin the interlophid valley posterolingual to the proto-conid. No posterior cingulid is present.

The m2 is similar in morphology to m1 but dif-fers as follows: it is larger, wider and longer inocclusal outline; the paraconid is present as a dis-tinct cusp on the paracristid, just anterolingual to

the protoconid; the protoconid is the second tallestcuspid (after the hypoconid), followed by the ento-conid, metaconid and paraconid; the anterior cin-gulid and precingulid are wider; the premetacristidis less well-developed; the preentocristid is absent.

The m3 is similar in morphology to m2 but dif-fers as follows: it is larger, wider and longer inocclusal outline; the trigonid is wider than the tal-oned; the paraconid is taller.

Genus BULUNGAMAYA Flannery et al., 1983Bulungamaya delicata Flannery et al., 1983

Figures 2, 3, 4

v. 1997 Nowidgee matrix Cooke, 1997a, p. 282-288, figs. 1-2.

Revised Diagnosis. Bulungamaya delicata ischaracterised by the following features: a buccallyexpanded masseteric canal; a sharp ventrally con-vex dentary margin below m1-2; i1 has its enamelconfined to the buccal surface where it is exten-sive; ventral and dorsal flanges are present on i1;p3 and P3 are elongate with seven cups and cus-pids (contra Flannery et al., 1983) with associatedfine transcritids and transcristae (although coarserthan in species of Ganguroo) and a bulbous base;a small alveolus for i2 is present directly posteriorto i1; lower molars and dp3 are bunolophodont withwell-developed posthypocristids, no connectionbetween the entoconid and hypoconid, protoconidconnected to cristid obliqua and no posterior cin-gulid present; m1 lacks a protostylid; m4 is notgreatly reduced in size relative to the other molarsin contrast to the condition in potoroines and hyp-siprymnodontids (Kear and Cooke 2001); dP3 isbilophodont but the upper molars are bunolopho-dont, having an incomplete the protoloph (no con-nection between protocone and paracone), but acomplete metaloph (the metacone connects to themetaconule); and a large stylar cusp C on all uppermolars.

Bulungamaya delicata differs from species ofGanguroo, Wabularoo and all macropodines,sthenurines and balbarids in having bunolophodontmolars instead of bilophodont (Cooke, 1997a;Archer, 1979; Kear and Cooke, 2001; Prideaux,2004). It differs from hypsiprymnodontids in havingelongate premolars, but differs from all potoroinesin having shorter and more bulbous premolars(Kear and Cooke, 2001).Holotype. CPC22187, left dentary containing par-tial i1, p3, m1-4. Referred Material. Camel Sputum Site: QMF30390, right dentary with i1, p3, m1-4, QMF19961, left dentary with m1-4, QM F20255, rightdentary with m3-4, QM F22761, left dentary with

15


m1-2, QM F30392, right dentary with dp3, p3, m1-2, QM F30394, right dentary with p3, m1-4, QMF30395, right maxilla with P3, M1-4, QM F19962,right dentary with m4, QM F19984, left dentary withm1, QM F19974, right dentary with m1-2, QMF30723, right dentary with m1-3, QM F57023, leftmaxilla with P3, M1-4, QM F19965, right maxillawith P3, M1-2, half of M3, QM F19954, left maxillawith M2-4, QM F19953, left maxilla with dP3, M1-2,QM F23491, right maxilla with M1-3, QM F20285,left maxilla with dP3, M1-2, QM F19675, right max-illa with P3, M1-2, QM F20310, right maxilla withM2-4; D -Site: QM F10650, right dentary containingthe root of i1; Dirk’s Towers Site: QM F24651, leftdentary with p3, m1-4, QM F30718, right dentarywith p3, m1-4, QM F29702, right maxilla with P3,M1-3; Judy's Jumping Joint Site: QM F57024, leftmaxilla with dP2-3, M1-2; Keith’s Chocky BlockSite: QM F57012, right m1; Neville’s Garden Site:QM F20011, right maxilla with M1-2; Upper Site:QM F30393, right dentary with dp3, m1, QMF19986, left maxilla with M1-2, QM F19619, rightmaxilla with P3, M1; Wayne’s Wok Site: QMF19937, left dentary with p3, m1-4, QM F20069,left dentary with dp2-3, p3, m1-2, QM F20080, rightdentary with dp2-3, p3, m1-3, m4 in crypt, QMF30391, left dentary with m1-2, m3 in crypt, QMF56988, right dentary with i1, p3, m1-4, QMF19579, left dentary with p3, m1-3, QM F24184,left dentary with m2-3, QM F19586, left dentarywith dp2, p3, m1-3, m4 in crypt, QM F24191, leftdentary with dp2-3, broken p3, m1, QM F20165,right dentary with m1-2, m4 in crypt, QM F56253,left dentary with dp2-3, m1-3, QM F19616, leftmaxilla with M1-3, QM F19939, left maxilla withM1-4, QM F56989, left maxilla with P3, M1-2, QMF19918, right maxilla with P3, M1-4, QM F30831,left maxilla with M1-4, QM F24478, right maxillawith M2-4; White Hunter Site: QM F19991, left den-tary with m3-4.Age and Distribution. The holotype is from GSite, D Site Plateau, Riversleigh WHA, northwest-ern Queensland, Australia. The age of G Site isinterpreted to be equivalent to the RiversleighLocal Fauna (=”D -Site LF”), Riversleigh FaunalZone A, late Oligocene (Archer et al., 1989, 1997;Arena, 2004; Travouillon et al., 2006, 2011). WhiteHunter LF is also interpreted to be a part of River-sleigh’s Faunal Zone A. Camel Sputum LF, Dirk'sTowers LF, Judy's Jumping Joint LF, Upper LF andWayne's Wok LF are part of Riversleigh’s FaunalZone B, early Miocene in age. Keith's Chocky

Block LF is part of Riversleigh’s Faunal Zone C,middle Miocene in age.Remarks. Dental descriptions of Bulungamayadelicata are provided by Flannery et al. (1983) andCooke (1997a). Arguments for subsuming Nowid-gee matrix into B. delicata can be found in the dis-cussion below. Description of the previouslyunknown upper deciduous premolars follows.

Description

The dP2 and dP3 are preserved in QMF57024 (Figure 2). The dp2 is plagiaulacoid, aboutas long as M1 but only two thirds the widths. Theocclusal margin is relatively horizontal and aboutone third shorter than the length of the crown base.The lingual face of the crown is steeper than thebuccal face. Five cusps are present on the occlusalcrest and have transcristae associated with them,although the posterior transcristae are much lessobvious than anteriorly.

The dP3 is a molariform tooth (Figure 2), bilo-phodont and shorter than dP2 and wider (but notas wide as M1). The tooth has a straight buccalmargin and a convex lingual margin. The paraconeis the tallest cusp on the occlusal surface followedby the metacone, metastyle, parastyle,neometaconule, metaconule and protocone, indecreasing order. A tall preparacrista joins theparastyle at the most anterobuccal corner of thetooth. The postparacrista descends from theparacone posterolingually and meets thepremetacrista, forming a complete centrocrista atthe buccal end of the interloph valley. Thepremetacrista joins the metacone posterobuccally.The postmetacrista departs from the metaconeposteriorly and joins to a small cusp, possibly themetastyle, before curving posterolingually to meetthe postmetaconulecrista at the most posterioredge of the tooth. The postmetaconulecrista con-tinues anterolingually and connects to themetaconule. The metaloph joins the metaconule tothe metacone transversely, through a smallneometaconule, located about halfway betweenthe metaconule and the metacone. Similarly, theprotoloph joins the paracone to the protocone, butthis crest is shallow lingually. The postprotocristadeparts the protocone posterobuccally and ends inthe interloph valley, at the anterior flank of themetaloph, anterobuccal to the metaconule. Noanterior cingulum is present; instead, the lingualflank of the preparacrista is wide and steep.

16


RESULTS

Coefficient of Variation Analysis

Coefficients of variation (CVs) for Thylogalestigmatica (Appendix 8) range between 3.54 and12.5, generally falling within the expected range(4–10) for a single mixed-sex population (Simpsonet al., 1960). The second lower molar was the leastvariable tooth in overall size (CVs 3.54–4.83) whilep3 was the most highly variable tooth (CVs 7.6–12.5). P3 was the only other tooth to have valuesover 10 (CV = 11.35).

Thylogale thetis (Appendix 9) was slightly lessvariable with CVs ranging between 4.98 and 11.16.The third upper molar was the least variable toothin overall size (CVs 5.28-5.95), but m1 was themost variable tooth (CVs 9.06-11.16), followed byp3 and P3.

When Thylogale stigmatica and T. thetis arecombined (Appendix 10), the range increases tovalues between 5.06 and 16.33. The second lowermolar (CVs 6.01–6.52) and third upper molar (CVs5.86–6.19) remained the least variable teeth, andp3 (CVs 11.75–16.33) remained the most variabletooth followed by P3 (CVs 9.71–13.25) and m1(CVs 7.09–9.09).

CVs ranged between 3.28 and 7.99 for Gan-guroo bilamina (Appendix 11) and between 2.06and 8.86 for the combined sample of Bulungamayadelicata and ‘Nowidgee matrix’ (Appendix 12). Inboth cases, they followed the trends seen in spe-cies of Thylogale, with p3 being the most variabletooth followed by P3 and m1, but in the combinedsample of Bulungamaya delicata/’Nowidgeematrix’, m4 and M4 were more variable than m1.

Bivariate Plots

Bivariate plots of dental variables for speciesof Thylogale (Appendices 13, 14, 15, 16) show astrong separation for p3 and P3, except for onespecimen overlapping in p3 length versus posteriorwidth. Length of P3/p3 seems to correlate withgender, with one sex having slightly longer premo-lars than the other, but while males generally havelonger premolars in T. thetis, they are generallyshorter than those of females in T. stigmatica.

In contrast, upper and lower molars do notshow a clear separation and mostly overlap in theirrange (Appendices 13, 14, 15, 16). There is no dis-tinct separation of males and females either,except in Thylogale thetis m4s which are distinc-tively larger in males. There is no significant differ-ence between using anterior or posterior widthversus length. In upper molars, females of each

species greatly overlap. Males overlap much lessand tend to occupy opposite spectra (short andwide versus long and narrow). This is also reflectedin the overall size of the upper teeth where T. thetishas smaller premolars but larger molars than T.stigmatica. This pattern does not seem to be pres-ent in lower molars.

Bivariate plots of upper dentition of Ganguroobilamina and ‘Nowidgee matrix’ are shown in Fig-ure 6. These two species are almost completelyseparated by P3 measurements except for onespecimen of N. matrix from Dirk’s Towers (QMF29702) which falls well within the range of G. bila-mina. Upper molar measurements overlap greatlybetween the two species.

A similar pattern is found in the lower dentition(Figure 7) where very little overlap exists in p3measurements of Ganguroo bilamina and Nowid-gee matrix but greatly overlap in m1-4. A specimenof G. bilamina from the Leaf Locality (KutjamarpuLocal Fauna) falls within the range of Riversleighspecimens for all dental measurements. The holo-type of Bulungamaya delicata falls within the rangeof N. matrix for all comparable teeth. Ganguroobites does not fall within the range of any otherspecies for p3 and m2. It overlaps with both G. bila-mina and N. matrix in its m1 measurements. Purtiamosaicus overlaps with N. matrix in its p3 mea-surements only, while Ngamaroo archeri overlapswith P. mosaicus and N. matrix only in its m1dimensions.

Principle Components Analysis

In the PCA of the upper dentition (Appendix17.1), Component 1 accounts for 39.32% of thevariance while Component 2 accounts for 27.98%of the variance. Other components accounted forless than 8% of the variance (e.g., Component 3 =8%, Component 4 = 4.83%, etc.). P3 length andwidth were the most useful measurements for sep-aration, followed by M4 length, M2 to M4 anteriorwidth and M1 posterior width.

In the PCA of the lower dentition (Appendix17.2), Component 1 accounts for 44.02% of thevariance while Component 2 accounts for 30.84%of the variance (Component 3 = 7.93%, Compo-nent 4 = 4.71%, etc.). The p3 length and widthwere the most useful measurements followed bym1 and m3 lengths.

The PCA of the upper dental variables of Gan-guroo bilamina and ‘Nowidgee matrix’ (Figure 8)shows significant overlap. Component 1 accountsfor 27.36% of the variance while Component 2accounts for 14.97% of the variance (Component 3

17


18

FIGURE 6. Bivariate plots of upper dentition of Ganguroo bilamina and ‘Nowidgee matrix’. L – anteroposterior length,AW = anterior width, W = maximum width, P = premolar, M = molar.


19

FIGURE 7. Bivariate plots of lower dentition of Ganguroo bilamina (+UCMP 88221 from Kutjamarpu LF), Ganguroobites n. sp., Bulungamaya delicata, ‘Nowidgee matrix’, Purtia mosaicus and Ngamaroo archeri. L – anteroposteriorlength, AW = anterior width, PW = posterior width, W = maximum width, p = premolar, m = molar.


20

FIGURE 8. Principle Component Analysis with convex hulls of log transformed upper (1) and lower (2) dentition mea-surements of Ganguroo bilamina (+UCMP 88221 from Kutjamarpu LF), Ganguroo bites n. sp., Bulungamaya deli-cata, ‘Nowidgee matrix’, Purtia mosaicus and Ngamaroo archeri.


= 11.17%, Component 4 = 8.65%, etc.). P3 lengthand width were the most useful measurements fol-lowed by M1 anterior and posterior width. The PCAof the lower dental variables (Figure 8) also showssignificant overlap between G. bilamina and N.matrix. A specimen of G. bilamina (UCMP 8822)from the Leaf Locality falls within the range of Riv-ersleigh specimens. The holotype of Bulungamayadelicata falls within the range of N. matrix. Two outof three specimens of G. bites fall within the rangeG. bilamina and N. matrix, but the third is well out-side their range. Purtia mosaicus and Ngamarooarcheri are well separated. Component 1 accountsfor 45.23% of the variance while Component 2accounts for 14.62% of the variance (Component 3=8.91%, Component 4 = 7.07%, etc.). The p3length and width were the most useful measure-ments followed by m2 and m3 lengths and poste-rior widths.

Multivariate Analysis Of Variance (MANOVA)

When the means of log transformed dentalmeasurements were compared using MANOVA,we found significance between comparing meansof each sex of each taxon (Appendix 18.1-2) andbetween means of each taxon (Appendix 18.3-4).When sexes are separated, the MANOVA found nosignificant differences between the means of eachsex of each taxon, for both upper (Appendix 18.1)and lower dentition (Appendix 18.2), with P-valuesranging from 0.34 to 0.87. The MANOVA wasunable to calculate p-value for the comparisonbetween males and females of Thylogale thetis, asa result of a smaller sample size. In contrast, whensexes were combined as a single taxon, theMANOVA found significant differences between

taxa for both upper (Appendix 18.3) and lower den-tition (Appendix 18.4), with P-values below 0.05.This suggests that when used on fossil specimens,the MANOVA should be able to differentiatebetween fossil taxa, but will not be able to differen-tiate between sexes of the same taxon.

The results of the MANOVA for the fossilspecimens are shown in Table 1. The results of theMANOVA found significant differences (P-value =1.48E-03) between Ganguroo bilamina and‘Nowidgee matrix’ for the mean upper dentitionmeasurements (Table 1.1). For the mean lowerdentition measurements, the results indicate thatthere are significant differences between all spe-cies examined (G. bilamina + UCMP 88221 fromKutjamarpu LF), G. bites, ‘N. matrix’ (+B. delicata)and N. archeri, except between G. bites and N.archeri, for which the MANOVA couldn’t calculate aP-value, due to sample size (only 3 specimens forG. bites, and 4 for N. archeri), and between G. bitesand ‘N. matrix’ were the P-value was above 0.05(P-value=0.053).

Canonical Variates Analysis (CVA)

The results of the CVAs for the log trans-formed dental measurements of Thylogale taxa areshown in Appendix 19. Whether sexes were sepa-rated or not, the CVAs separated each group quitedistinctively, with only a slight overlap betweenmales and females T. stigmatica in the upper denti-tion. The CVA reclassifications are shown inAppendix 20. When sexes were separated, onlyfour specimens incorrectly misclassified as theincorrect sex, but never as a different species(Appendix 20.1). When sexes were combined, notmisclassification occurred (Appendix 20.2).

TABLE 1. Results of the Multivariate Analysis Of Variance (MANOVA) analysis showing the P-values for the meancomparison of upper (1) and lower (2) log transformed dental measurements of Ganguroo bilamina (+UCMP 88221from Kutjamarpu LF), G. bites n. sp., ‘Nowidgee matrix’ (+Bulungamaya delicata) and Ngamaroo archeri.1

2

G. bilamina ‘N. matrix’

G. bilamina 0 1.48E-03

‘N. matrix’ 1.48E-03 0

G. bilamina G. bites N. archeri ‘N. matrix'

G. bilamina 0 5.71E-05 9.10E-26 2.58E-07

G. bites 5.71E-05 0 Fail 0.053

N. archeri 9.10E-26 Fail 0 2.66E-08

‘N. matrix' 2.58E-07 0.053 2.66E-08 0

21


22

FIGURE 9. Canonical Variates Analysis with convex hulls of upper (1) and lower (2) log transformed dental measure-ments of Ganguroo bilamina (+UCMP 88221 from Kutjamarpu LF), Ganguroo bites n. sp., ‘Nowidgee matrix’ (+Bulun-gamaya delicata) and Ngamaroo archeri.


The results of the CVAs for the log trans-formed dental measurements of the fossil taxa areshown in Figure 9. In the upper dentition CVA (Fig-ure 9.1), there is a lot of overlap between Gan-guroo bilamina and ‘Nowidgee matrix’. This is alsoseen in the lower dentition CVA (Figure 9.2), withG. bites also overlapping with ‘N. matrix’. Ngama-roo archeri is, however, very well separated. CVAreclassifications are shown in Table 2. Some of theGanguroo bilamina are misclassified by the CVAas ‘Nowidgee matrix’ and vice versa. No Ngama-roo archeri is misclassified, while one specimen ofG. bites is misclassified as ‘Nowidgee matrix’.Specimen UCMP_88221 from Kutjamarpu LF isclassified by the CVA as G. bilamina, and the holo-type of Bulungamaya delicata, CPC_22187, isclassified as ‘Nowidgee matrix’.

Phylogenetic Analysis

Seventy-eight most parsimonious trees (treelength = 265; consistency index excluding uninfor-mative characters = 0.4651; retention index =0.7320) were recovered from the analysis of themorphological matrix (see Appendix 21 for matrix).The strict consensus tree of these is given in Fig-ure 10, with bootstrap (above branches) andBremer (below branches) support values.

The strict consensus is highly unresolved,with few branches supported by high bootstrap orBremer values. Macropodoidea is divided in twomajor clades, one comprising macropodines,sthenurines, Riversleigh fossil taxa and potoroines(bootstrap = 67%) and the other comprising bal-barids and hypsiprymnodontids (bootstrap =<50%). The relationship between Riversleigh fossiltaxa (Bulungamaya delicata, Ganguroo bilamina,G. bites and Wabularoo naughtoni) and potoroines(Bettongia penicillata, B. moyesi, Potorous tridacty-lus, Purtia mosaicus, Ngamaroo archeri and Wak-iewakie lawsoni) is unresolved, with monophyly ofPotoroinae not recovered while P. mosaicus, N.archeri and W. lawsoni form a clade distinct fromthat of other potoroines (bootstrap =<50%). Wan-buroo hilarus is recovered as the sister taxon(bootstrap = 59%, Bremer = +2) to all macropod-ines and sthenurines. The macropodine andsthenurine clades are recovered with moderatesupport (bootstrap = 61%, Bremer = +2) and reso-lution within each clade. The relationship of River-sleigh fossil taxa (Bulungamaya delicata,Ganguroo bilamina, G. bites and Wabularoonaughtoni) within Macropodoidea remains uncer-tain.

DISCUSSION

Coefficients of variation of dental dimensionsof a single, mixed-sex population, usually fallsbetween 0 and 10 (Simpson et al., 1960).Theresults of our metric analyses demonstrate thateven though p3 and P3 are the most variable teethin terms of length and width (CV values >10) mea-sures, they remain the most useful teeth to distin-guish species of Thylogale in thebivariate plots,Principle Component Analysis (PCA) and Canoni-cal Variates Analysis (CVA). These results are con-sistent with the metric analyses of species ofMacropus in Bartholomai (1971). However, Bar-tholomai (1971) urges that all cheekteeth shouldbe considered together, taking into account boththe variation in size and morphological characters.

Dental measurements for Ganguroo bilaminaare within the range expected from a single spe-cies (CV values <10) for both the lower and thenewly described upper dentition (Simpson et al.,1960). Molar measurements significantly overlapwith other fossil species (e.g., Bulungamaya deli-cata/’Nowidgee matrix’ and Ganguroo bites),although anterior molars are generally smaller insize and upper and lower third premolar measure-ments overlap very little with Bulungamaya deli-cata/’Nowidgee matrix’ and not at all with G. bites.Morphologically, G. bilamina differs from the bunol-ophodont B. delicata/N. matrix in being completelybilophodont (sensu Cooke, 1997a) and differs fromG. bites in having seven cuspids instead of eight onp3, no paraconid present on any molars and a buc-cal crest from the entoconid, which is evident injuveniles but worn away in adults. Both morphol-ogy and size variation strongly support G. bilaminabeing treated as a species distinct from B. delicata/’N. matrix’ and G. bites.

While Bulungamaya delicata (Flannery et al.,1983) and ‘Nowidgee matrix’ (Cooke, 1997a) weredescribed as separate species in part because thetype specimens of B. delicata are heavily worn,making it difficult to compare their morphology withrelatively unworn specimens. Here we argue thatthey represent the same species. First, metricanalyses do not support the view that there are twospecies involved. All dental measurements of B.delicata fall within the range of ‘N. matrix’ (seebivariate plots, Figures 7). In both the PCA andCVA, B. delicata wfell within the range of ‘N. matrix’and was reclassified by the CVA as ‘N. matrix’ (Fig-ures 8, 9, Table 2). Combined univariate statisticsfor the two species had coefficient of variation val-ues within the range of a single species (CV = <10,Appendices 4, 5) and the MANOVA (Table 1) sug-

23


TABLE 2. Results of the Canonical Variates Analysis (CVA) reclassification using log transformed dental measure-ments of fossil specimens, for the upper (1) and lower (2) dentition. Group 1 = Ganguroo bilamina; group 2 = ‘Nowid-gee matrix’; group 3 = Ngamaroo archeri; group 4 = G. bites. Incorrect reclassifications are highlighted in grey. 12

Upper dentition Lower dentition

Specimen Given group CVA Classification Specimen Given group CVA Classification

QMF56318 1 1 QMF19870 1 1

QMF56995 1 1 QMF19868 1 1

QMF19696 1 2 QMF19966 1 1

QMF19696 1 1 QMF30400 1 1

QMF19958 1 1 QMF56996 1 1

QMF20705 1 1 QMF19607 1 1

QMF19900 1 1 QMF20248 1 1

QMF19695 1 1 QMF20254 1 1

QMF20253 1 1 QMF23486 1 2

QMF19861 1 1 QMF20246 1 1

QMF19976 1 1 QMF20019 1 1

QMF19957 1 1 QMF56995 1 1

QMF19857 1 1 QMF19903 1 1

QMF19975 1 1 QMF56254 1 1

QMF19959 1 1 QMF56259 1 2

QMF20017 1 1 QMF24744 1 1

QMF56990 1 1 QMF20006 1 1

QMF30436 1 1 QMF36351 1 1

QMF30343 1 1 QMF19988 1 1

QMF36230 1 1 QMF19844 1 1

QMF19668 1 1 QMF24190 1 2

QMF19692 1 1 QMF20039 1 1

QMF234604 1 1 QMF30396 1 1

QMF19987 1 1 QMF19642 1 1

QMF23202 1 1 QMF30397 1 1

QMF20269 1 2 QMF20293 1 1

QMF56993 1 1 QMF56999 1 1

QMF19801 1 1 QMF20271 1 1

QMF19673 1 1 QMF19643 1 2

QMF56992 1 1 QMF20295 1 1

QMF19985 1 2 QMF19947 1 1

QMF19885 1 1 QMF56991 1 1

QMF19687 1 1 QMF20272 1 1

QMF19651 1 1 QMF19640 1 1

24


TABLE 2 (continued).

QMF19689 1 1 QMF19646 1 1

QMF19629 1 1 QMF19688 1 1

QMF19674 1 1 QMF19814 1 1

QMF52814 1 2 QMF30399 1 1

QMF52814 1 1 QMF19591 1 1

QMF57020 1 1 QMF19810 1 1

QMF57020 1 1 QMF19835 1 1

QMF20261 1 1 QMF57022 1 1

QMF19589 1 1 QMF19836 1 1

QMF19617 1 1 QMF57021 1 1

QMF19590 1 1 QMF56998 1 1

QMF24219 1 2 QMF19834 1 1

QMF20163 1 1 QMF19827 1 2

QMF56256 1 2 QMF20082 1 1

QMF56258 1 1 QMF19597 1 2

QMF56265 1 1 QMF24480 1 1

QMF56265 1 1 QMF56997 1 1

QMF24475 1 2 QMF20108 1 1

QMF24476 1 1 QMF20109 1 1

QMF50518 1 1 QMF19915 1 1

QMF56261 1 1 QMF19602 1 1

QMF56255 1 2 QMF50519 1 1

QMF56255 1 1 QMF50519 1 1

QMF24216 1 1 QMF24050 1 1

QMF56263 1 1 QMF30721 1 1

QMF57023 2 2 QMF20081 1 1

QMF19965 2 1 QMF36364 1 1

QMF19954 2 2 QMF24477 1 1

QMF19953 2 2 QMF30398 1 1

QMF23491 2 2 QMF56257 1 1

QMF20285 2 2 QMF24482 1 2

QMF19675 2 2 QMF31405 1 1

QMF20310 2 1 QMF56260 1 1

QMF29702 2 1 QMF36363 1 2

QMF57024 2 2 QMF36470 1 1

QMF20011 2 2 QMF56264 1 1

QMF19986 2 1 UCMP_88221 1 1

QMF19619 2 2 CPC_22187 2 2

QMF19616 2 2 QMF30390 2 2

QMF19939 2 2 QMF22761 2 2

QMF56989 2 2 QMF19961 2 2

25


TABLE 2 (continued).

QMF19918 2 2 QMF20255 2 2

QMF30831 2 1 QMF30392 2 2

QMF24478 2 2 QMF30394 2 2

QMF19962 2 1

QMF19984 2 2

QMF19974 2 1

QMF30723 2 2

QMF24651 2 2

QMF30718 2 2

QMF57012 2 2

QMF30393 2 2

QMF20080 2 2

QMF20069 2 2

QMF19937 2 2

QMF30391 2 2

QMF56988 2 2

QMF19579 2 2

QMF24184 2 2

QMF19586 2 2

QMF24191 2 2

QMF20165 2 1

QMF56253 2 2

QMF19991 2 2

SAMP23626 3 3

SAMP27817 3 3

SAMP27818 3 3

SAMP31834 3 3

QMF23776 4 4

QMF23775 4 2

QMF23777 4 4

gested that the mean of combine specimens B.delicata and ‘N. matrix’ were significantly differentfrom other taxa. Morphologically, while it is very dif-ficult to compare the worn type specimens of B.delicata to other specimens, several features areshared between B. delicata and ‘N. matrix’, sug-gesting synonymy of these species (Figures 3, 4).These features are: the occlusal surface of i1reaches the level of occlusal plane of the molars(Figure 3; it is below the molar occlusal plane in G.bilamina); p3 transcristids are relatively coarse(Figures 3, 4; transcristids are much finer in spe-

cies of Ganguroo); cristid obliqua connects to pro-toconid (visible on m3 of the holotype of B.delicata); buccal crest from the entoconid does notconnect to the hypoconid (also visible on m3 in theholotype of B. delicata); and the talonid of m4 issignificantly smaller than the trigonid (contra thecondition in species of Ganguroo). The lower molarproportions of type specimens of B. delicata fromRiversleigh’s D and G Sites are almost identical tothe only known specimen of ‘N. matrix’ from WhiteHunter Site. All three of these sites are interpretedto be late Oligocene in age (Archer et al., 1989,

26


27

FIGURE 10. Phylogenetic relationships of Bulungamaya delicata, Ganguroo bilamina, Ganguroo bites and Wabularoonaughtoni (highlighted in bold), based on 108 character craniodental matrix. Fossil taxa are indicated by †. Strict con-sensus of 78 most parsimonious trees (tree length = 265; consistency index excluding uninformative characters =0.4651; retention index = 0.732) from maximum parsimony analysis of the matrix. Numbers above branches representbootstrap values (1000 replicates); numbers below branches represent decay indices.


1997; Arena, 2004; Travouillon et al., 2006, 2011).Considering the overall similarity in size and mor-phology, we propose that ‘Nowidgee matrix’ beregarded as a junior synonym of Bulungamaya del-icata.

The phylogenetic analysis failed to resolve therelationship of basal macropodids to other macrop-odoids. Ganguroo bilamina, G. bites, Bulungamayadelicata and Wabularoo naughtoni were recoveredas a clade containing potoroines and macropod-ines+sthenurines, but the relationship was notresolved. Kear et al. (2007) and Kear and Pledge(2008) recovered ‘bulungamayines’ as a paraphy-letic group, more closely related to macropodinesthan potoroines, but with no support (bootstrap val-ues = <50%). They recommended ‘Bulungamay-inae’ to be treated as incertae sedis. However,there are some issues with the character coding for‘bulungamayines’ in those studies. First, charac-ters scored for Wabularoo naughtoni include cra-nial and upper dentition scores based onundescribed material. Second, characters scoredfor G. bilamina include postcranial scores belong-ing to another species from AL90 Site, Ganguroosp. 2, which is also undescribed. Third, charactersscored for B. delicata include cranial and upperdentition scores also based on undescribed speci-mens but which in this case have been determinedhere to represent G. bilamina.

In contrast, Prideaux and Warburton (2010),who only included in their analysis Ganguroo bila-mina and ‘Nowidgee matrix’, recovered ‘N. matrix’as the sister group to all macropodids (lagostro-phines, sthenurines and macropodines), and G. bil-amina as the sister group to sthenurines andmacropodines with stronger support values (boot-strap = >50%). It is noteworthy that Prideaux andWarburton (2010) completely rescored G. bilaminaand ‘N. matrix’ based on type material assigned toboth species by Cooke (1997a), but they didinclude postcranials assigned incorrectly to G. bila-mina by Kear et al. (2001a).

New material of Ganguroo bilamina describedin this paper may also shed some light on its rela-tionship to other macropodoids. The flat frontalregion observed in G. bilamina is also present inskulls of Ganguroo sp. 2. This feature may be asynapomorphy for species of Ganguroo althoughthis region of the skull is not preserved in speci-mens of G. bites. Some features of the skull of G.bilamina, however, are almost certainly plesiomor-phic for macropodoids. For example, absence offlexion of the skull in the basicranial and rostralregions, smooth transition of the zygoma to the

cheek region and lack of development of the mas-seteric process are also features of hypsiprymno-dontids, potoroines and balbarids (Kear andCooke, 2001). However, G. bilamina sharesderived features with macropodines including abroad parietal-alisphenoid contact and an alisphe-noid contribution to the ventral floor of the second-ary foramen ovale which suggest a closerrelationship with macropodines (Kear and Cooke,2001).

Upper molars of G. bilamina are also morederived than Bulungamaya delicata in protolophmorphology, being bilophodont rather than bunolo-phodont. The M1 of G. bilamina has a ‘forelink’derived from the preprotocrista, which contributesa lingual component to the protoloph crest beforeturning anteriorly. In contrast, in B. delicata the pre-protocrista makes no contribution to the protolophcrest. The M1 of G. bilamina, however, retains aremnant of stylar cusp D in the form of a stylarcrest which is absent in B. delicata.

The neometaconule of Ganguroo bilaminaoccurs also in Bulungamaya delicata. It is alsocommon in relatively derived balbarines (Cooke,2000) and relatively derived macropodids such assthenurines (Prideaux, 2004). Tedford and Wood-burne (1987) note that the neometaconule is aneomorphic structure in quadritubercular uppermolars of diprotodontian marsupials. In agreement,it is therefore more parsimonious to presume thatthe neometaconule of macropodoids is a plesiom-orphic feature rather than independently developedin the different macropodoid lineages in which itoccurs. Absence of this feature in some balbarinesand basal macropodids would accordingly be theresult of secondary loss rather than a plesiomor-phic macropodoid condition.

Lower molars of Ganguroo bilamina areclearly more derived than those of species ofBulungamaya delicata in terms of hypolophid for-mation. In G. bilamina, the entoconid is linked tothe hypoconid by the posthypocristid, which hasbecome transversely orientated and elevated, inaddition to a second crest that departs bucally fromthe entoconid and is sometimes present in juve-niles (contra Cooke, 1997a). This situation con-trasts with the condition in balbarines where both abuccal crest from the entoconid and the linguallydisplaced posthypocristid contribute to the hypolo-phid (Cooke, 1997c). While both balbarines andmacropodids have achieved bilophodonty in lowermolars, the process of doing so appears to be theresult of parallel evolution. In contrast, loph forma-tion in upper molars has apparently proceeded in

28


similar ways in both groups. The preprotocristacontributes the lingual component of the protolophand continues anteriorly to form the forelink(Cooke, 2000).

Cooke (1997b) provided the first speciesoccurrence list of Riversleigh macropodoids, whichwas later updated in Archer et al. (2006) andTravouillon et al. (2011). In the course of compilingdata for the present paper, we reviewed all avail-able macropodoid specimens from Riversleigh(Table 3) other than hypsiprymnodontids and bal-barids, which will be the subject of a separatereview. A number of taxa listed in Cooke (1997b),Archer et al. (2006) and Travouillon et al. (2011)have been left unchanged, removed from or modi-fied in our species lists here for the following rea-sons.

Gumardee pascuali is probably a large potor-oine, as described by Flannery et al. (1983) on thebasis of a single worn maxilla from D Site whichcontained an elongated P3, typical of potoroines.No further material has been reported until now.Cooke (1997b) reported the presence of an unde-scribed species, ‘Nowidgee’ sp. 2, based on twodentaries from the White Hunter Site. New materialfrom White Hunter Site, including an almost com-plete skull and several maxillae and dentaries, per-fectly matches G. pascuali. The two dentariesreferred by Cooke (1997b) to ‘Nowidgee’ sp. 2 alsoappear to be referrable to G. pascuali. Added sup-port for this conclusion comes from the fact that thedentaries perfectly occlude with the maxillae, cor-responding precisely in size and morphology(bunolophodont dentition). Hence we conclude that‘Nowidgee’ sp. 2 is synonymous with G. pascuali(Table 3) and should no longer used in the litera-ture, at least until it is formally described.

Wabularoo naughtoni was first described byArcher (1979) on the basis of a dentary from D Siteand a p3 from G Site. Flannery et al. (1984)reported an additional specimen from D Site.Cooke (1997b) reported the occurrence of this spe-cies from undescribed specimens in White HunterSite as well as six younger Faunal Zone B sites.After re-examining this material, we do not agreewith Cooke (1997b) that these specimens belongto W. naughtoni, as none possesses the distinctiveenlarged p3 characterising this species, apart fromthe specimen from White Hunter Site. These spe-cies occurrences should not be used at least untilthe material is formally described.

Wakiewakie lawsoni was described by Wood-burne (1984) on the basis of a dentary and two P3sfrom the Kutjamarpu Local Fauna from Lake Nga-

pakaldi in northern South Australia. This specieswas also reported from Riversleigh on the basis ofa single Faunal Zone B dentary from Upper Site(Godthelp et al., 1989).

Bulungamaya delicata, first described byFlannery et al. (1983) on the basis of specimensfrom D Site and G Site, was also reported byCooke (1997b) from a number of Faunal Zone A, Band C specimens. After reviewing these specimensin the context of the present review, we determinedthat some of these additional specimens appear torepresent other taxa (e.g., Ganguroo bilamina andG. sp. 2). While it is clear that B. delicata is repre-sented by specimens from Faunal Zones A and B,the only Faunal Zone C deposit occurs in theKeith’s Chocky Block Site (KCB), which is a verticalfissure fill. As noted by several authors (e.g.,Creaser, 1997; Archer et al., 1997; Travouillon etal., 2006, 2011), the KCB Site may have been col-lecting fossils for a longer period of time than othersites and potentially taxa from both Faunal ZonesB and C.

Cooke (1997a) described Ganguroo bilaminabased on dentaries from Faunal Zone B sites.Here, we describe additional material from thisspecies and reassign specimens from the Bitesan-tennary Site originally assigned to G. bilamina to anew species, G. bites. As noted above, all speci-mens of G. bilamina occur in Faunal Zone B,except for Cadbury’s Kingdom (CK) Site, whichwas previously thought to be a Faunal Zone C site(Travouillon et al., 2006, 2011). However, assign-ment of the CK faunal assemblage to Faunal ZoneC had low support (<50%, Travouillon et al., 2011).Eight of the nine species present in this site arepresent in at least Zones B and C (and in somecases all Faunal Zones). Nimiokoala greystanesi isonly present in Faunal Zone B and Cadbury’s King-dom, as is Priscakoala lucyturnbullae (Black et al.,2012). Additionally, while reviewing the whole col-lection, Balbaroo fangaroo was identified fromCadbury’s Kingdom, which otherwise only occursin Faunal Zone B, while a more derived speciesoccurs in Faunal Zone C (Balbaroo sp. 4 of Cooke,1997b). Similarly, the relatively derived species ofGanguroo, Ganguroo sp. 2, occurs in FaunalZones C and D, but not B. There is, on balance,more support for referring the CK assemblage toFaunal Zone B than C, suggesting that G. bilaminais restricted to Faunal Zone B (Table 3). The pres-ence of G. bilamina in the Kutjamarpu Local Faunastrengthens biocorrelations with Riversleigh’s Fau-nal Zone B. A description of Ganguroo sp. 2 is inprogress.

29


TO

TAL

The suggestion that Ganguroo bilamina andBulungamaya delicata are present in the Price IsRight LF (Archer et al., 2006) is not supported herebecause specimens used to identify these two spe-cies are larger and morphologically distinct. Allspecimens from the Price Is Right Site examinedduring the present study appear to represent Bal-baroo fangaroo, Ganawamaya acris and a newspecies related to Gumardee pascuali.

Bettongia moyesi was described on the basisof an associated skull and dentary from the TwoTrees Site on the Gag Plateau at Riversleigh as theearliest record of a modern potoroine genus (Flan-nery and Archer, 1987b). Since then, another den-tary has been recovered from the Henk’s Hollow

Site, confirming its presence in Faunal Zone C(Archer et al., 2006). Two specimens from the Mel-ody’s Maze Site, also on the Gag Plateau, mayrepresent a new potoroine (Table 3).

Wanburoo hilarus was described as a ‘bulun-gamayine’ on the basis of several dentaries andupper teeth from Faunal Zone C sites and EncoreSite (Cooke, 1999; gen. Wan. sp. 1 in Cooke,1997b). However, Prideaux and Warburton (2004)suggest that it may be in fact a sthenurine. In ourreview of Riversleigh material, we have identifiednew specimens referable to this species, includinga near complete skull. Dental specimens from theEncore Site (gen. Wan. sp. 2 in Cooke 1997b),however, are synonymous with Rhizosthenurus

TABLE 3. Site occurrence (serriated) of pre-Pliocene Riversleigh basal macropodid species, excluding hypsiprymno-

dontids and balbarids.

SpeciesFaunal Zone A F

ZA

?

Faunal Zone B

FZ

B/C

?

Faunal Zone C

FZ

D

G S

ite

D S

ite

Wh

ite

Hu

nte

r S

ite

Ju

dy'

s Ju

mp

ing

Jo

int

Sit

e

Dir

k's

To

wer

Sit

e

Bit

esA

nte

nn

ary

Sit

e

Cad

bu

ry's

Kin

gd

om

Sit

e

Cam

el S

pu

tum

Sit

e

Inab

eyan

ce S

ite

Ju

dit

h H

ori

zon

talis

Sit

e

Mik

e's

Men

ager

ie S

ite

Mik

e's

Po

tato

Pat

ch S

ite

Nev

ille

's G

ard

en S

ite

RS

O S

ite

Up

pe

r S

ite

Way

ne

's W

ok

Sit

e

KC

B S

ite

AL

90 S

ite

Cle

ft O

f A

ges

Sit

e

Do

me

Sit

e

Ga

g S

ite

Hen

k's

Ho

llow

Sit

e

Tw

o T

rees

Sit

e

Jim

's C

aro

use

l Sit

e

Jim

's J

aw S

ite

Las

t M

inu

te S

ite

Mai

n S

ite

NJC

08 S

ite

Ric

k S

ausa

ge

Sit

e

Me

lod

y's

Maz

e S

ite

Ara

chn

id R

idg

e

An

ge

la's

Sin

kho

le S

ite

Go

lden

Ste

ph

Sit

e

Jaw

Ju

nct

ion

Sit

e

Sku

ll S

ite

En

core

Sit

e

Rhizosthenurus flanneryi

1 1

Wanburoo hilarus

1 1 1 1 1 1 1 1 1 1 1 1 13

Bettongia moyesi

1 1 2

Ganguroo sp. 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 18

G. bilamina 1 1 1 1 1 1 1 1 1 1 10

G. bites 1 1

Bulungamaya delicata

1 1 1 1 1 1 1 1 1 1 10

Wakiewakie lawsoni

1 1

Wabularoo naughtoni

1 1 1 3

Gumardee pascuali

1 1 2

Total 2 3 3 1 1 1 1 2 1 1 1 1 2 1 3 2 3 2 2 2 2 3 1 2 1 2 2 2 2 1 1 1 1 1 1 2

30


flanneryi (Kirkham, personal commun., 2013),which was first described as Wanburoo sp., thenrenamed in a following publication, based on post-cranial material from the same site (Kear et al.,2001b; Kear, 2002). Cooke (1997b) also notes anundescribed species, Wanburoo sp. 2, from theEncore, Last Minute and Henk’s Hollow Sites.However, specimens of Wanburoo sp. 2 from theEncore Site are synonymous with R. flanneryi andfrom the Last Minute and Henk’s Hollow Sites withW. hilarus. A complete description of the skull anddentition of R. flanneryi is currently in progress(Kirkham et al., in prep.).

CONCLUSION

Based on morphological assessment andmetric analyses, ‘Nowidgee matrix’ is synonymisedwith Bulungamaya delicata. This bunolophodontspecies is now known from upper and lower denti-tions, from several of Riversleigh’s Faunal Zone Aand B sites as well as Keith’s Chocky Block Site(arguably Faunal Zone C but as a vertical fissurefill it may have also accumulated fossils from Fau-nal Zone B). We have described additional materialfor Ganguroo bilamina, including the previouslyunknown skull and upper dentition, and describeda new species, G. bites, from the BitesantennarySite. These two species are currently restricted toFaunal Zone B. Our phylogenetic analysis wasunable to resolve the interrelationships of basalmacropodids. However, cranial and dental mor-phology of G. bilamina suggest closer affinities withmacropodines than potoroines. Finally, we haverevised understanding about some of the othermacropodoid taxa in pre-Pliocene Riversleighsites.

ACKNOWLEDGMENTS

Support for research at Riversleigh has comefrom the Australian Research Council (LP0989969,LP100200486 & DP130100197 grants to M. Archerand S.J. Hand at the University of New SouthWales); XSTRATA Community Partnership Pro-gram (North Queensland); a National GeographicSociety grant to M. Archer; the University of NewSouth Wales; P. Creaser and the CREATE Fund,the Queensland National Parks and Wildlife Ser-vice; Environment Australia; the QueenslandMuseum; the Riversleigh Society Inc.; Outback atIsa; Mount Isa City Council; and private supportersincluding K. & M. Pettit, E. Clark, M. Beavis and M.Dickson. Assistance in the field has come fromhundreds of volunteers as well as staff and post-

graduate students of the University of New SouthWales. For access to and loans of specimens, wethank H. Janetzki, K. Spring, A. Rozefelds and S.Hocknull of the Queensland Museum, S. Inglebyand A. Divljan of the Australian Museum and C.Stevenson from the Western Australian Museum.We thank R. Day for providing funding to the Uni-versity of Queensland to create a postdoctoralposition for K. Travouillon. We thank the UNSWPalaeosciences Lab and the UQ Palaeo Hub fortheir support and we thank anonymous reviewersfor helpful comments.

REFERENCES

Aplin, K.P. and Archer, M. 1987. Recent advances inmarsupial systematics with a new syncretic classifi-cation, p. xv-lxxii. In Archer, M. (ed.), Possums andOpossums, Studies in Evolution. Surrey Beatty andSons, Sydney, Australia.

Archer, M. 1979. Wabularoo naughtoni gen. et sp. nov.,an enigmatic kangaroo (Marsupialia) from the middleTertiary Carl Creek Limestone of northwesternQueensland. Results of the Ray E. Lemley Expedi-tions, part 4. Memoirs of the Queensland Museum,19:299-307.

Archer, M. 1984. The Australian mammal radiation, p.585-625. In Archer, M. and Clayton, G. (eds.), Verte-brate Zoogeography and Evolution in Australasia.Hesperian Press, Western Australia, Carlisle.

Archer, M., Godthelp, H., Hand, S.J., and Megirian, D.1989. Fossil mammals of Riversleigh, northwesternQueensland: preliminary overview of biostratigraphy,correlation and environmental change. AustralianZoolologist, 25:29-65.

Archer, M., Hand, S.J., Godthelp, H., and Creaser, P.1997. Correlation of the Cainozoic sediments of theRiversleigh World Heritage fossil property,Queensland, Australia, p. 131-152. In Aguilar, J.-P.,Legendre, S., and Michaux, J. (eds.), Actes duCongrès BiochroM’97. École Pratique des HautesÉtudes, Institut de Montpellier, France.

Archer, M., Arena, D.A., Bassarova, M., Beck, R.M.D.,Black, K., Boles, W.E., Brewer, P., Cooke, B.N.,Creaser, P., Crosby, K., Gillespie, A., Godthelp, H.,Hand, S.J., Kear, B.P., Louys, J., Morrell, A.,Muirhead, J., Roberts, K.K., Scanlon, J.D., Travouil-lon, K.J., and Wroe, S. 2006. Current status of spe-cies- level representation of faunas from selectedfossil localities in the Riversleigh World HeritageArea, northwestern Queensland. Alcheringa SpecialIssue, 1:1-17.

Arena, D.A. 2004. The geological history and develop-ment of the terrain at the Riversleigh World HeritageArea during the middle Tertiary. Unpublished PhDThesis, University of New South Wales, Sydney, Aus-tralia.

31


Bartholomai, A. 1971. Morphology and variation of thecheek teeth in Macropus giganteus Shaw and Macro-pus agilis (Gould). Memoirs of the QueenslandMuseum, 16:1-18.

Beck, R.M.D., Godthelp, H., Weisbecker, V., Archer, M.,and Hand, S.J. 2008. Australia's oldest marsupialfossils and their biogeographical implications. PloSONE, 3(3):e1858. doi:10.1371/journal.pone.0001858

Black, K.H., Archer, M., and Hand, S.J. 2012. New Ter-tiary koala (Marsupialia,

Phascolarctidae) from Riversleigh, Australia, with a revi-sion of phascolarctid phylogenetics, paleoecology,and paleobiodiversity. Journal of Vertebrate Paleon-tology, 32:125-138.

Case, J.A. 1984. A new genus of Potoroinae (Marsu-pialia: Macropodidae) from the Miocene NgapakaldiLocal Fauna, South Australia, and a definition of thePotoroinae. Journal of Paleontology, 58:1074-1086.

Cooke, B.N. 1997a. New Miocene bulungamayine kan-garoos (Marsupialia: Potoroidae) from the River-sleigh, northwestern Queensland. Memoirs of theQueensland Museum, 41:281-294.

Cooke, B.N. 1997b. Biostratigraphic implications of fossilkangaroos at Riversleigh, northwestern Queensland.Memoirs of the Queensland Museum, 41:295-302.

Cooke, B.N. 1997c. Two new balbarine kangaroos andlower molar evolution within the subfamily. Memoirsof the Queensland Museum, 41:269-280.

Cooke, B.N. 1999. Wanburoo hilarus gen. et sp. nov., alophodont bulungamayine kangaroo (Marsupialia:Macropodoidea: Bulungamayinae) from the Miocenedeposits of Riversleigh, northwestern Queensland.Records of the Western Australian Museum, Supple-ment 57:239-253.

Cooke, B.N. 2000. Cranial remains of a new species ofbalbarine kangaroo (Marsupialia: Macropodoidea)from the Oligo-Miocene freshwater limestone depos-its of Riversleigh World Heritage Area, northern Aus-tralia. Journal of Paleontology, 74:317-326.

Cooke, B.N., Travouillon, K.J., Archer, M., and Hand,S.J. in review. Ganguroo robustiter sp. nov. (Macrop-odoidea, Marsupialia), a middle to early late Miocenemacropodid from Riversleigh World Heritage Area,Australia. Acta Palaeontologica Polonica.

Creaser, P. 1997. Oligocene–Miocene Sediments of Riv-ersleigh: the potential significance of topography.Memoirs of the Queensland Museum, 41:303-314.

Flannery, T.F. and Archer, M. 1987a. Hypsiprymnodonbartholomaii (Potoroidae: Marsupialia), a new spe-cies from the Miocene Dwornamor Local Fauna anda reassessment of the phylogenetic position of H.moschatus, p. 749-758. In Archer, M. (ed.), Possumsand Opossums: Studies in Evolution. Surrey Beattyand Sons, Sydney.

Flannery, T.F. and Archer, M. 1987b. Bettongia moyesi, anew and plesiomorphic kangaroo (Marsupialia: Poto-roidae) from Miocene sediments of northwesternQueensland, p. 759-767. In Archer, M. (ed.), Pos-sums and Opossums: Studies in Evolution. SurreyBeatty and Sons, Sydney.

Flannery, T.F., Archer, M., and Plane, M. 1983. MiddleMiocene kangaroos (Macropodoidea: Marsupialia)from three localities in northern Australia with adescription of two new subfamilies. Bureau of MineralResources Journal of Australian Geology and Geo-physics, 7:287-302.

Flannery, T.F., Archer, M., and Plane, M. 1984. Phyloge-netic relationships and a reconsideration of higherlevel systematics within the Potoroidae (Marsupialia).Journal of Paleontology, 58:1087-1097.

Flower, W.H. 1869. Remarks on the homologies andnotations of the teeth of the Mammalia. Journal ofAnatomy and Physiology, 3:262-278.

Godthelp, H., Archer, M., Hand, S.J., and Plane, M.D.1989. New potoroine from Tertiary Kangaroo WellLocal Fauna, N.T. and description of upper dentitionof potoroine Wakiewakie lawsoni from Upper SiteLocal Fauna, Riversleigh. Conference on Austral-asian Vertebrate Evolution, Palaeontology and Sys-tematics (CAVEPS) Abstracts:6.

Hammer, Ø., Harper, D.A.T., and Ryan, P.D. 2001. PAST:Palaeontological statistics software package for edu-cation and data analysis. Palaeontologia Electronica,4.1.4:9pp., 178KB; http://palaeo-electronica.org/2001_1/past/issue1_01.htm

Kear, B.P. 2002. Phylogenetic implications of macropo-did (Marsupialia: Macropodoidea) postcranialremains from Miocene deposits of Riversleigh, North-western Queensland. Alcheringa, 26:299-318.

Kear, B.P. and Cooke, B.N. 2001. A review of macropo-doid systematics with the inclusion of a new family.Memoirs of the Association of Australasian Palaeon-tologists, 25:83-101.

Kear, B.P. and Pledge, N. S. 2008. A new fossil kanga-roo from the Oligocene-Miocene Etadunna Forma-tion of Ngama Quarry, Lake Palankarinna, SouthAustralia. Australian Journal of Zoology, 55:331-339.

Kear, B.P., Archer, M., and Flannery, T.F. 2001a. Postcra-nial morphology of Ganguroo bilamina Cooke, 1997(Marsupialia: Macropodidae) from the middle Mio-cene of Riversleigh, northwestern Queensland.Memoirs of the Association of Australasian Palaeon-tologists, 25:123-138.

Kear, B.P., Archer, M., and Flannery, T.F. 2001b. Bulun-gamayine (Marsupialia: Macropodoidea) postcranialelements from the late Miocene of Riversleigh north-western Queensland. Memoirs of the Association ofAustralasian Palaeontologists, 25:103-122.

32


Kear, B.P., Cooke, B.N., Archer, M., and Flannery, T.F.2007. Implications of a new species of the Oligo-Mio-cene kangaroo (Marsupialia: Macropodoidea) Nam-baroo, from the Riversleigh World Heritage Area,Queensland, Australia. Journal of Paleontology,81:1147-1167.

Luckett, W.P. 1993. An ontogenetic assessment of dentalhomologies in therian mammals, p. 182-204. In Sza-lay, F.S., Novacek, M.J., and McKenna, M.C. (eds.),Mammal Phylogeny: Mesozoic Differentiation, Multi-tuberculates, Monotremes, Early Eutherians andMarsupials. Springer-Verlag, New York.

Megirian, D., Prideaux, G.J., Murray, P.F., and Smit, N.2010. An Australian land mammal age biochronologi-cal scheme. Paleobiology, 36:658-671.

Metzger, C.A. and Retallack, G.J. 2010. Middle Mioceneclimate change in the Australian outback. AustralianJournal of Earth Sciences, 57:871-885.

Murray, P.F. 1989. The cranium of Hadronomas puckridgiWoodburne, 1967 (Macropodoidea: Macropodidae) aprimitive macropodid kangaroo from the late MioceneAlcoota Fauna of the Northern Territory. The Beagle,6:115-132.

Murray, P.F. 1991. The sthenurine affinity of the LateMiocene kangaroo, Hadronomas puckridgi Wood-burne (Marsupialia, Macropodidae). Alcheringa,15:255-283.

Prideaux, G.J. 2004. Systematics and evolution of thesthenurine kangaroos. University of California Publi-cations in Geological Sciences, 146:i-xvii, 1-623.

Prideaux, G.J. and Warburton, N. M. 2010. An osteology-based appraisal of the phylogeny and evolution ofkangaroos and wallabies (Macropodidae: Marsu-pialia). Zoological Journal of the Linnean Society,159:954-987.

Simpson, G., Roe, A., and Lewontin, R. 1960. Quantita-tive Zoology. Harcourt Brace, New York.

Sorenson, M.D. and Franzosa, E.A. 2007. Treerot ver.3.0. Department of Biology, Boston University, Bos-ton, MA, 02215, USA.

Swofford, D.L. 2002. PAUP*. Phylogenetic AnalysisUsing Parsimony (*and Other Methods). Version 4(updated to 10 beta). Sinauer Associates, Sunder-land, Massachussets.

Szalay, F.S. 1969. Mixodectidae, Microsyopidae, and theInsectivore-Primate transition. Bulletin of the Ameri-can Museum of Natural History, 140:193-330.

Tedford, R.H. and Woodburne, M.O. 1987. The Ilariidae,a new family of vombatiforme marsupials from Mio-cene strata of South Australia and an evolution of thehomology of molar cusps in the Diprotodontia, p.401-418. In Archer, M. (ed.), Possums and Opos-sums: Studies in Evolution. Surrey Beatty and Sons,Sydney.

Travouillon, K.J., Archer, M., Hand, S.J., and Godthelp,H. 2006. Multivariate analyses of Cenozoic mamma-lian faunas from Riversleigh, north-westernQueensland. Alcheringa, Special Issue 1:323-349.

Travouillon, K.J., Legendre, S., Archer, M., and Hand,S.J. 2009. Palaeoecological analyses of River-sleigh’s Oligo-Miocene Sites: Implications for Oligo-Miocene climate change in Australia. Palaeogeogra-phy, Palaeoclimatology, Palaeoecology, 276:24-37.

Travouillon, K.J., Escarguel, G., Legendre, S., Archer,M., and Hand, S.J. 2011. The use of MSR (MinimumSample Richness) for sample assemblage compari-sons. Paleobiology, 37:696-709.

Wible, J.R. 2003. On the cranial osteology of the short-tailed opposum Monodelphis brevicaudata (Didelphi-dae, Marsupialia). Annals of Carnegie Museum,72:137-202.

Woodburne, M.O. 1984. Wakiewakie lawsoni, a newgenus and species of Potoroinae (Marsupialia: Mac-ropodidae) of medial Miocene age, South Australia.Journal of Paleontology, 58:1062-1073.

Worthy, T.H., Tennyson, A.J.D., Archer, M., Musser,A.M., and Hand, S.J. 2006. Miocene mammalreveals a Mesozoic ghost lineage on insular NewZealand, southwest Pacific. Proceedings of theNational Academy of Sciences of the United Statesof America, 103:19419-19423.

33


34

APPENDIXES

For appendix material, see palaeo-electronica.org/content/2014/711-riversleigh-basal-macropodoidsAPPENDIX 1. Measurements of the lower dentition of type and referred material of Ganguroo bilamina in mms. L –anteroposterior length, AW = anterior width, PW = posterior width, dp = deciduous premolar, p = premolar, m = molar.APPPENDIX 2. Measurements of the upper dentition of type and referred material of Ganguroo bilamina in mms. L –anteroposterior length, AW = anterior width, PW = posterior width, dP = deciduous premolar, P = premolar, M = molar.APPENDIX 3. Measurements of the lower dentition of type and referred material of Ganguroo bites in mms. L – antero-posterior length, AW = anterior width, PW = posterior width, dp = deciduous premolar, p = premolar, m = molar.APPENDIX 4. Measurements of the lower dentition of type and referred material of Bulungamaya delicata/’Nowidgeematrix’ in mms. L – anteroposterior length, AW = anterior width, PW = posterior width, dp = deciduous premolar, p =premolar, m = molar.APPENDIX 5. Measurements of the upper dentition of type and referred material of Bulungamaya delicata/’Nowidgeematrix’ in mms. L – anteroposterior length, AW = anterior width, PW = posterior width, dP = deciduous premolar, P =premolar, M = molar.APPENDIX 6. Measurements of the lower dentition of additional specimens used in analyses, in mms. L – anteropos-terior length, AW = anterior width, PW = posterior width, p = premolar, m = molar.APPENDIX 7. Measurements of the uppers dentition of additional specimens used in analyses, in mms. L – anteropos-terior length, AW = anterior width, PW = posterior width, P = premolar, M = molar.APPENDIX 8. Univariate statistics of specimens of Thylogale stigmatica. N = number of specimen, Min = minimummeasurement, Max = maximum measurement, L = anteroposterior length, AW = anterior width, PW = posterior width,p = lower premolar, P = upper premolar, m = lower molar, M = upper molar.APPENDIX 9. Univariate statistics of specimens of Thylogale thetis. N = number of specimen, Min = minimum mea-surement, Max = maximum measurement, L = anteroposterior length, AW = anterior width, PW = posterior width, p =lower premolar, P = upper premolar, m = lower molar, M = upper molar.APPENDIX 10. Univariate statistics of combined specimens of Thylogale stigmatica and T. thetis. N = number of spec-imen, Min = minimum measurement, Max = maximum measurement, L = anteroposterior length, AW = anterior width,PW = posterior width, p = lower premolar, P = upper premolar, m = lower molar, M = upper molar.APPENDIX 11. Univariate statistics of type and referred material of Ganguroo bilamina. N = number of specimen, Min= minimum measurement, Max = maximum measurement, L = anteroposterior length, AW = anterior width, PW = pos-terior width, p = lower premolar, P = upper premolar, m = lower molar, M = upper molar.APPENDIX 12. Univariate statistics of type and referred material of Bulungamaya delicata/’Nowidgee matrix’. N =number of specimen, Min = minimum measurement, Max = maximum measurement, L = anteroposterior length, AW =anterior width, PW = posterior width, p = lower premolar, P = upper premolar, m = lower molar, M = upper molar.APPENDIX 13. Bivariate plots with convex hulls of lower dentition of Thylogale stigmatica and T. thetis, using anteriorwidth. L – anteroposterior length, AW = anterior width, p = premolar, m = molar.APPENDIX 14. Bivariate plots with convex hulls of lower dentition of Thylogale stigmatica and T. thetis, using posteriorwidth. L – anteroposterior length, PW = posterior width, p = premolar, m = molar.APPENDIX 15. Bivariate plots with convex hulls of upper dentition of Thylogale stigmatica and T. thetis, using anteriorwidth. L – anteroposterior length, AW = anterior width, P = premolar, M = molar.APPENDIX 16. Bivariate plots with convex hulls of upper dentition of Thylogale stigmatica and T. thetis, using posteriorwidth. L – anteroposterior length, PW = posterior width, P = premolar, M = molar.APPENDIX 17. Principle Component Analysis with convex hulls of upper (1) and lower (2) dentition of Thylogale stig-matica and T. thetis.APPENDIX 18. Results of the Multivariate Analysis Of Variance (MANOVA) analysis showing the P-values for themean comparison of upper (1 and 3) and lower (2 and 4) log transformed dental measurements of Thylogale stigmat-ica and T. thetis, with separation of sexes (1 and 2) and without separation (3 and 4).APPENDIX 19. Canonical Variates Analysis with convex hulls of upper (1 and 3) and lower (2 and 4) log transformeddental measurements of Thylogale stigmatica and T. thetis, with separation of sexes (1 and 2) and without separation(3 and 4).APPENDIX 20. Results of the Canonical Variates Analysis (CVA) reclassification using log transformed dental mea-surements of Thylogale stigmatica and T. thetis, with separation of sexes (1) and without separation (2).1, group 1 = T.stigmatica male; group 2 = T. stigmatica female; group 3 = T. thetis female; group 4 = T. thetis male; 2, group 1 = T.stigmatica; group 2 = T. thetis. Incorrect reclassifications are highlighted in grey.Results of the Canonical Variates Analysis (CVA) reclassification using log transformed dental measurements of Thylo-gale stigmatica and T. thetis, with separation of sexes (1) and without separation (2).1, group 1 = T. stigmatica male;group 2 = T. stigmatica female; group 3 = T. thetis female; group 4 = T. thetis male; 2, group 1 = T. stigmatica; group 2= T. thetis. Incorrect reclassifications are highlighted in grey.APPENDIX 21. Nexus-formatted character matrix for the 29 taxa included in our phylogenetic analyses. Outgroup taxawas Trichosurus vulpecula. ? = missing data; - = inapplicable.

palaeontologia electronica · 2019-09-10 · travouillon et al.:riversleigh basal macropodoid 2...

Documents