Memoirs of Museum Victoria 74: 189–207 (2016) Published 2016 ISSN 1447-2546 (Print) 1447-2554 (On-line) http://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/ New material of Gumardee pascuali Flannery et al., 1983 (Marsupialia: Macropodiformes) and two new species from the Riversleigh World Heritage Area, Queensland, Australia KENNY J. TRAVOUILLON 1,2,*, KAYLENE BUTLER 2, MICHAEL ARCHER 3, AND SUZANNE J. HAND3 1 Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia (Kenny.Travouillon@museum. wa.gov.au) 2 School of Earth Sciences, University of Queensland, St Lucia, Queensland 4072, Australia ([email protected]; [email protected]) 3 PANGEA Research Centre, School of Biological, Earth and Environmental Sciences, UNSW, New South Wales 2052, Australia ([email protected]; [email protected]) * To whom correspondence should be addressed. E-mail: [email protected] Abstract Travouillon, K.J., Butler, K., Archer, M., and Hand, S.J. 2016. New material of Gumardee pascuali Flannery et al., 1983 (Marsupialia: Macropodiformes) and two new species from the Riversleigh World Heritage Area, Queensland, Australia. Memoirs of Museum Victoria 74: 189–207. New specimens of the late Oligocene macropodiform, Gumardee pascuali, are described, as well as two new late Oligocene to early Miocene species, G. springae sp. nov. and G. richi sp. nov. Species of Gumardee exhibit a unique combination of features, including very long upper and lower third premolars, partially lophodont molars and, in dorsoventral plane, concave lower molar row and convex upper molar row. We combined two morphological matrices to assess the phylogenetic relationships of these species. Our analysis recovered species of Gumardee as a well-supported monophyletic group, nested within Potoroinae. Gumardee richi and G. pascuali appear to be more derived than G. springae in having a more strongly developed posthypocristid that is almost hypolophid-like. Keywords Macropodiform; systematics; late Oligocene; early Miocene; new species. Introduction M4 (their M5) and an elongate P3, aligned with the molar row and with buccal and lingual cingula and 11 buccal and lingual Flannery et al. (1983) described Gumardee pascuali from a transcristae. Its relationship with potoroines was further single maxilla containing worn P3-M4. The holotype was supported by Flannery et al. (1984), who discussed the found at D-Site, in the Riversleigh World Heritage Area monophyly of this group in which they also included other (WHA), north-western Queensland, Australia. Current Oligo-Miocene taxa such as Purtia, Wakiewakie, Bulungamaya understanding of biostratigraphy in the Riversleigh WHA and Wabularoo (Archer, 1979; Case, 1984; Flannery et al., 1983; suggests that D-Site is part of Faunal Zone A, which is Woodburne, 1984). However, monophyly of potoroines, and interpreted to be late Oligocene in age (Archer et al., 1989; including these extinct taxa, have not been found in more recent Archer et al., 1997; Creaser, 1997; Travouillon et al., 2006, morphological phylogenies (Kear et al., 2007; Kear and Pledge, 2011). Three other Riversleigh Faunal Zones, B, C and D, have 2008; Prideaux and Warburton, 2010; Travouillon et al., 2014). been interpreted to be early, middle and early late Miocene in Here we describe new material of Gumardee pascuali, as age, respectively (Archer et al., 1989; Archer et al., 1997; well as two new species of the same genus, from the Riversleigh Creaser, 1997; Travouillon et al., 2006, 2011). WHA. We re-assess the phylogenetic relationships of all three Additional specimens of Gumardee were reported from Gumardee taxa, using a new combined morphological matrix. the Namba Formation of Lake Pinpa, South Australia (Flannery and Rich, 1986). However, due to the fragmentary Materials and methods nature of the fossils recovered (mostly isolated teeth), they were referred to Gumardee sp., and await re-assessment when The new specimens of Gumardee pascuali and of the two new additional material is known. species were collected from Riversleigh WHA, north-western Flannery et al. (1983) suggested that G. pascuali was a Queensland (18°59'–19°08'S, 138°34'–138°43'E). All potoroine because it had ‘bunodont’ molars, a greatly reduced specimens are registered in the Queensland Museum fossil 190 K.J. Travouillon, K. Butler, M. Archer and S.J. Hand collection, Brisbane, Australia (prefix QM F). All dental supplementary information for details), and added additional measurements were taken with digital vernier callipers. characters to the matrix (see characters 115–119 in supplementary Terminology follows Luckett (1993) for cheek tooth information). One such character appears to be particularly homology, Archer (1984) and Cooke (1997a, b) for dental important for distinguishing basal taxa from other morphology, Meredith et al. (2009) and Prideaux and Warburton macropodiforms: ‘molar morphology’ (character 23), which we (2010) for higher systematic nomenclature (using redefined to be applicable to the whole range observed. Macropodiformes sensu Meredith et al., 2009; and using ‘Bunodont’ taxa (e.g. Burramys and Hypsiprymnodon spp.) subfamilies Potoroinae, Macropodinae, Lagostrophinae and have poorly defined lateral crests (no distinct lophs or lophids). Sthenurinae as part of Macropodidae, sensu Prideaux and ‘Bunolophodont’ taxa (e.g. Trichosurus, Ekaltadeta, Warburton, 2010), and Travouillon et al. (2006, 2011, 2013) for Aepyprymnus spp.) have defined lateral crests, but these crests biostratigraphic nomenclature (see also Archer et al., 1989, not functioning as lophs or lophids (no physical connection 1997; Creaser, 1997; Arena, 2004). between the crest and the opposing cusp). ‘Partially lophodont’ In a previous attempt to analyse the phylogenetic relationship taxa (e.g. Bulungamaya, Purtia, Ngamaroo and Gumardee of basal macropodiforms, using the matrix of Kear and Pledge spp.) have functional anterior lophids (well-developed lophid (2008), the tree recovered was highly unresolved (Travouillon et connecting protoconid to metaconid) and posterior lophs (well- al., 2014). We suggested that combining the matrices of Kear developed loph connecting metacone to metaconule), but and Pledge (2008), and of Prideaux and Warburton (2010), incomplete (non-functional) posterior lophids (entoconid and might provide better resolution of macropodiform relationships. hypoconid are not connected by a well-developed lophid) and Here, we combined those two matrices, using the matrix of anterior lophs (protocone and paracone are not connected by a Prideaux and Warburton (2010) as the base, because it included well-developed loph). ‘Bilophodont’ taxa (e.g. Gangaroo, more taxa and contained all macropodid subfamilies Wabularoo, and all balbarids, sthenurines and macropodines) (Lagostrophinae was not represented in Kear and Pledge, 2008). have fully functional anterior and posterior lophs/lophids. Our We added four new out-group species, including the Miocene combined matrix has a total of 119 characters. bandicoot Galadi speciosus (Travouillon et al., 2010), the extant The parsimony analysis was performed using 1,000 heuristic Common Brushtail Possum Trichosurus vulpecula, the extant replicates, saving ten trees per replicates, and the analysis Common Ringtail Possum Pseudocheirus peregrinus, and the repeated using the saved trees. The analysis resulted in multiple Miocene Pygmy Possum Burramys brutyi (Brammall and parsimonious trees, which were summarised as a strict Archer, 1997). We also added several other fossil and extant consensus tree. Bootstrap values were calculated using 1000 macropodiform taxa as follows: replicates of ten random addition sequence replicates. The -Hypsiprymnodontidae: Hypsiprymnodon bartholomaii analysis was done in the software PAUP* 4.0b10 (Swofford, Flannery and Archer, 1987a, H. philcreaseri Bates et al., 2014, 2002). Decay indices were calculated in the software TreeRot. Ekaltadeta ima Archer and Flannery, 1985, Jackmahoneya v3 (Sorenson and Franzosa, 2007). toxoniensis Ride, 1993 and Propleopus oscillans De Vis, 1888. -Balbaridae: Nambaroo saltavus and N. tarrinyeri Flannery Systematic palaeontology and Rich, 1986, N. couperi Cooke, 1997b, N. gillespieae Kear et al., 2007, Wururoo dayamayi Cooke, 1997b, Balbaroo Macropodidae Gray, 1821 gregoriensis and B. camfieldensis Flannery et al., 1983, B. fangaroo Cooke, 2000, Ganawamaya acris, G. ornata and G. Potoroinae Trouessart, 1898 aediculis Cooke, 1992. -Basal macropodiforms: Bulungamaya delicata , Purtia Gumardee Flannery et al., 1983 mosaicus Case, 1984, Wakiewakie lawsoni Woodburne, 1984, Ganguroo bites Travouillon et al., 2014, Wabularoo naughtoni Type species. Gumardee pascuali Flannery et al., 1983 Additional species. Gumardee richi sp. nov. and G. springae Archer, 1979, Gumardee pascuali Flannery et al., 1983, G. richi sp. nov. sp. nov. and G. springae sp. nov (this work). -Potoroinae: Bettongia moyesi (Flannery and Archer, Revised generic diagnosis. Potoroine with elongated and narrow 1987b), and the extant Rufous Bettong Aepyprymnus rufescens. sectorial P3/p3 aligned with molar row, buccal and lingual -Lagostrophinae: Tjukuru wellsi (Prideaux and Tedford, cingulae on the medial portion of P3, concave lower molar row 2012). and convex upper molar row in dorsoventral plane, buccinator From the Kear and Pledge (2008) matrix, we added to our sulcus on the dentary, reduced m4/M4 (smaller than m3/M3), matrix only cranial and dental characters