New South Wales, 2052, Australia Version of record first published: 08 Jan 2013.

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1 This article was downloaded by: [ ] On: 06 February 2013, At: 03:54 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Journal of Vertebrate Paleontology Publication details, including instructions for authors and subscription information: The genus Galadi: three new bandicoots (Marsupialia, Peramelemorphia) from Riversleigh s Miocene deposits, northwestern Queensland, Australia K. J. Travouillon a b, Y. Gurovich b, M. Archer b, S. J. Hand b & J. Muirhead b a School of Earth Sciences, University of Queensland, St Lucia, Queensland, 4072, Australia b School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales, 2052, Australia Version of record first published: 08 Jan To cite this article: K. J. Travouillon, Y. Gurovich, M. Archer, S. J. Hand & J. Muirhead (2013): The genus Galadi: three new bandicoots (Marsupialia, Peramelemorphia) from Riversleigh s Miocene deposits, northwestern Queensland, Australia, Journal of Vertebrate Paleontology, 33:1, To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 Journal of Vertebrate Paleontology 33(1): , January by the Society of Vertebrate Paleontology ARTICLE THE GENUS GALADI: THREE NEW BANDICOOTS (MARSUPIALIA, PERAMELEMORPHIA) FROM RIVERSLEIGH S MIOCENE DEPOSITS, NORTHWESTERN QUEENSLAND, AUSTRALIA K. J. TRAVOUILLON, *,1,2 Y. GUROVICH, 2 M. ARCHER, 2 S. J. HAND, 2 and J. MUIRHEAD 2 1 School of Earth Sciences, University of Queensland, St Lucia, Queensland 4072, Australia, kennytravouillon@hotmail.com; 2 School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia ABSTRACT We describe three new bandicoot species of the genus Galadi from the Miocene of Riversleigh World Heritage Area in northern Australia. The first species, which is represented by a complete dentary and several isolated upper molars, is restricted to Riversleigh s Faunal Zone B. It is the largest bandicoot as yet known from Riversleigh. The second species is represented by 19 specimens, including a partial skull and several maxillae and dentaries, from Riversleigh s Faunal Zone C. Several features distinguishing this species from the similarly sized type species G. speciosus are of interest, notably the presence of larger maxillopalatine fenestrae and additional maxillary fenestrae, incomplete centrocrista on all upper molars, a more complete posterior cingulum on upper molars, and higher degree of dental wear, which together suggest a more omnivorous diet. The third species is represented by a single maxillary, which exhibits a quite different combination of dental features compared with other Galadi species. All Galadi species appear to be restricted to Riversleigh s Faunal Zones B and C, which are interpreted to be early and middle Miocene in age, respectively, with rainforest habitats persisting throughout. SUPPLEMENTAL DATA Supplemental materials are available for this article for free at INTRODUCTION Peramelemorphia (bandicoots and bilbies) is an extant order of marsupials from Australia, New Guinea, and surrounding islands. It consists of the two extant families of Peramelidae: Thylacomyidae and the recently extinct Chaeropodidae (which contains Chaeropus ecaudatus, the pig-footed bandicoot). Eight extant Australian species are referred to four genera (Perameles, Isoodon, Macrotis, andechymipera), with at least 13 New Guinean species (in the genera Echymipera, Peroryctes, Microperoryctes, andrhynchomeles; see also Aplin et al., 2010). Currently, nine fossil species of peramelemorphians are known from the late Oligocene to the Pleistocene of Australia; six of these are attributed to modern families and two are referred to the extinct family Yaralidae (Muirhead, 2000). Galadi is the second genus of fossil bandicoots recognized from the Oligo-Miocene of Riversleigh World Heritage Area (WHA), northwestern Queensland, Australia. It is currently known only by the type species Galadi speciosus, represented by an exceptionally well-preserved skull and a few dentary remains (Travouillon et al., 2010). Galadi speciosus exhibits a mixture of plesiomorphic and apomorphic dental and cranial characters, with the relationship between Galadi speciosus and Yarala burchfieldi (the first peramelemorphian described from the same Oligo-Miocene deposits of Riversleigh by Muirhead 2000) being unresolved (Travouillon et al., 2010). However, Galadi speciosus appears to lie outside crown group Peramelemorphia but closer to the crown group than Yarala burchfieldi, and for this reason, was not referred to the family Yaralidae (Travoullion et al., 2010). A total of three species of fossil bandicoots have been described from the Oligo-Miocene of Australia: Yarala burchfieldi and Galadi speciosus from the Riversleigh WHA; and Yarala kida Schwartz, 2006, from the late Oligocene Kangaroo Well Local Fauna. At least six additional bandicoot species are yet to be * Corresponding author. described from Riversleigh Oligo-Miocene sites (Muirhead, 2000; Archer et al., 2006) and eight from the Etadunna and Wipajiri formations of South Australia (Woodburne et al., 1993). Here we describe three new bandicoot species from the Miocene of Riversleigh that appear to be referable to Galadi. The three species are represented by numerous specimens, including partial cranial remains and isolated maxillae and dentaries. We assess the phylogenetic relationships of these three taxa within Peramelemorphia. We also discuss their paleoecology and document the temporal distribution of species of the genus Galadi. MATERIALS AND METHODS All specimens were collected from the Riversleigh WHA, northwestern Queensland, between 1983 and 2010, and recovered by acid processing of limestone blocks. Biostratigraphic nomenclature follows Arena (2004) and Travouillon et al. (2006). Institutional Abbreviations AR, Specimens temporarily held in collections at the University of New South Wales, Sydney; QMF, Queensland Museum fossil collection, Brisbane. Anatomical Abbreviations Dental terminology follows Archer (1976), Muirhead and Filan (1995), Voss and Jansa (2003), and Travouillon et al. (2010) and with molar loci homology following Luckett (1993). Cranial anatomy follows Muirhead (1994, 2000), Voss and Jansa (2003), Wible (2003), Beck et al. (2008a), and Travouillon et al. (2010). Peramelemorphian systematics follows Groves and Flannery (1990), Muirhead (1994, 2000), Muirhead and Filan (1995), and Travouillon et al. (2010). Higher-level marsupial systematics follows Aplin and Archer (1987). Phylogenetic Analysis We analyzed the matrix from Travouillon et al. (2010) and made a number of changes. The original matrix contained 51 qualitative morphological characters (28 dental and 23 cranial). 153

3 154 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 We made the following changes and additions, resulting in a total of 74 characters: 1. Character 15, morphology of the centrocrista, was split into two characters, to distinguish between the morphology of the centrocrista of M1 and M2 from that of M3. 2. We added 14 characters from Voss and Jansa (2009; characters 52, 55, 57, 59, 82, 85, 89, 93, 96, 97, 114, 116, 120, and 125). 3. We added four characters from Aplin et al. (2010; characters 3, 6, and 8 from page 22 and character 11 from page 17) 4. We added four new dental characters (shape of I2 4, shape of upper canine, lower molar crown height, and presence of metaconule). All new characters were left unordered, leaving the original 23 ordered characters in all analyses (see online Supplementary Data 1 and 2 for a complete list of characters) We included the 21 peramelemorphian species surveyed by Travouillon et al. (2010) and 12 additional taxa, of which four are extant (Peroryctes broadbenti, Microperoryctes papuensis, M. ornata, Echymipera davidi) and eight are fossil taxa from the Pleistocene (Perameles sobbei; Price 2002), Pliocene (Ischnodon australis, Stirton, 1955; Numbigilga ernielundeliusi, Beck, Archer, Godthelp, Mackness, Hand, and Muirhead, 2008a; cf. Peroryctes tedfordi and cf. Peroryctes. sp., Turnbull et al., 2003), and Miocene (the three new taxa described here). Following Travouillon et al. (2010), we used the following taxa as outgroups: the early Eocene australidelphian Djarthia murgonensis Godthelp, Wroe, and Archer, 1999; the early Miocene dasyurid Barinya wangala Wroe, 1999; and the middle Miocene dasyuromorphian Mutpuracinus archibaldi Murray and Megirian, Parsimony Analysis Parsimony analysis of the matrix (see online Supplementary Data) was carried out using PAUP 4.0b10 (Swofford, 2002), following Travouillon et al. (2010), using an initial search comprising 1000 heuristic replicates, saving 10 trees per replicate, followed by a second heuristic search within the trees saved from the first stage. Multiple most parsimonious trees were summarized using strict consensus. Bootstrap values were calculated using 100 bootstrap replicates; each bootstrap replicate itself comprised 10 random addition sequence replicates. We performed two analyses, one containing all taxa, and a second in which we removed all taxa that were represented by maxilla or dentary only. If one taxon is known only from a maxilla and another is known only from a dentary, a phylogenetic analysis can never reconstruct them in a clade (even if they are in fact closely related) because they have no characters scored in common, so there is nothing that can link them. The taxa we removed from the second analysis were Galadi adversus, Perameles sobbei, Ischnodon australis, Numbigilga ernielundeliusi, cf. Peroryctes tedfordi,andcf. Peroryctes sp. SYSTEMATIC PALEONTOLOGY Order PERAMELEMORPHIA (Kirsch, 1968) Aplin and Archer, 1987 Superfamily incertae sedis Family incertae sedis Genus GALADI Travouillon, Gurovich, Beck, and Muirhead, 2010 Type Species Galadi speciosus Travouillon, Gurovich, Beck, and Muirhead, Remarks A nearly complete skull, with associated femur and pelvis (QM F54568), of Galadi speciosus is now known from Inabeyance Site, Faunal Zone B, Riversleigh WHA, northwestern Queensland. Revised Generic Diagnosis Species of Galadi differ from all other bandicoots in the following combination of features: short and robust snout; upper molar crown margins rounded, short and wide, lower molar crown anteroposteriorly reduced; anterior cingula on all upper molars; large entoconid, blade-like with low anteriorly directed preentocristid; small metaconular hypocone; parastylar region of M1 less blade-like than in modern peramelemorphians; and large size. Upper and lower premolars are large and wide. Anterior cusps of lower premolars are absent or minute. Five incisors without separating diastemata, nasals extend posteriorly past the anterior region of the orbit; short infraorbital canal; no lachrymal orbital rim; moderate nasolabial/antorbital fossa; large orbitosphenoid; alisphenoid and parietal contact; unspecialized sphenorbital fissure and foramen rotundum; one pair of incisive foramina and one pair of maxillopalatine fenestrae; large primary foramen ovale bordered by alisphenoid and petrosal; no secondary foramen ovale; small, uninflated alisphenoid tympanic wing; small knob-like rostral tympanic process of the petrosal; uninflated epitympanic recess; and very weakly defined squamosal epitympanic sinus. GALADI GRANDIS, sp. nov. (Figs. 1 4) Specific Diagnosis Galadi grandis differs from the type species G. speciosus in the following combination of features: having a complete centrocrista on all upper molars that does not breech the ectoloph; stylar cusp E present on M1 and M2; smaller metaconular hypocone on upper molars making the overall shape more triangular; laterally compressed lower canine; m1 paraconid is more buccally positioned, curving the anterior part of the tooth between the paraconid and metaconid in occlusal view; and the dentary is longer and less robust at the anterior end. Size large ( kg compared with 900 g). Specific Etymology Grandis, Latin word meaning grand, in reference to its large size (being the largest bandicoot so far known from the Riversleigh WHA). Holotype QM F23394, left dentary with c to m4 (Figs. 1 3). Paratypes QM F23395, isolated LM1; QM F23396, isolated RM2; and QM F23397, isolated LM3 (Fig. 4). Referred Material Camel Sputum Site: QM F54575, isolated right M2; QM F54576, isolated right m2; QM F54577, isolated right m3; QM F54578, isolated left m3; QM F54579, isolated left m1. Creaser s Rampart Site: QM F30788, left dentary fragment with unerupted p3 and loose m3. Mike s Menagerie Site: QM F56066, isolated left m2; QM F56065, isolated left m3; QM F56067, isolated right m3. Ross Scott-Orr (RSO) Site: QM F56062, isolated right M2. Wayne s Wok Site: QM F54574, isolated left M2; QM F56063, isolated right M2. Type Locality The holotype and paratypes are from Camel Sputum Site, Godthelp Hill, Riversleigh WHA, northwestern Queensland, Australia. Age and Stratigraphy These sites are part of Riversleigh WHA, northwestern Queensland, and contain terrestrial carbonates that appear to represent vadose cave deposits (Arena, 2004). They are part of Faunal Zone B, estimated to be approximately early Miocene in age (Travouillon et al., 2006, 2011). Description Dentary Dorsal surface of the ramus remains at constant height (Fig. 1), with depth of ramus greatest below m3. The symphysis extends posteriorly to below the posterior root of p2. Two mental foramina are present: the anterior foramen is below p1, the posterior foramen below m1. The largest diastema occurs between c and p1 and a smaller one between p1 and p2. The coronoid process extends from the dentary at approximately 95 degrees.

4 TRAVOUILLON ET AL. FOSSIL BANDICOOTS FROM AUSTRALIA 155 FIGURE 1. Galadi grandis, sp. nov., mandible (holotype QM F23394). A, buccal view of left mandible; B, lingual view of left mandible. Abbreviations: ang, angular process; c, lower canine; coc, coronoid crest; con, mandibular condyle; m1 m4, lower molar 1 to lower molar 4; maf, masseteric fossa; manf, mandibular foramen; mas, mandibular symphysis; mf, mental foramen; p1 p3, lower premolar 1 to premolar 3. Scale bar equals 2 cm. Lower Dentition The canine is laterally compressed (Fig. 1). It is broken at its tip but would have been taller than the molars. It extends from the dentary at an oblique angle and is not strongly curved. The posterior roots of the premolars are thicker than the anterior roots (Fig. 2). Only remnants of an anterior cusp are visible on p1 to p3. The anterior margin of the main cusp is rounded on p1 to p3. The main cusps of p2 and p3 are anteriorly positioned above the gap between the roots, and lingually so that the buccal surface bulges in occlusal view. The posterior cusp increases in size from p1 to p3 (Fig. 2). The posterior crest curves from the lingual main cusp to terminate at the middle of the posterior edge of the crown. No enamel ridge is present around the posterior margin of p1 but is present on p2 and enlarged on p3 on both lingual and buccal surfaces. A crest runs around this ridge on p3. The crown and central cusp increase in size from p1 to p3. On m1, the protoconid and metaconid are closer together than either is to the paraconid (Fig. 3). The paraconid is much lower on the crown, lower than the entoconid. No anterior cingulid is present. The metaconid is posterolingually positioned with respect to the protoconid. The paraconid lies anterobuccal to the metaconid, curving the anterior part of the tooth between the paraconid and metaconid in occlusal view. The talonid is wider than the trigonid. The entoconid, oval in shape in occlusal view, is posterolingually positioned relative to the metaconid and joined to the base of the metaconid by a low preentocristid. The hypoconulid is not raised but extends posteriorly; it is not joined to the entoconid but connects to the hypoconid by a deflected posthypocristid. The hypoconid lies directly buccal to the entoconid. The cristid obliqua is slightly concave, terminating in the buccal half of the tooth on the posterolingual flank of the protoconid. A low ridge of enamel joins the protoconid to hypoconid. No posterior cingulid is present. Morphology of m2 is similar to that of m1 except as follows. The paraconid is directly anterior to the metaconid. The paracristid is longer. The distance between the metaconid and protoconid is greater but less than that between the paraconid and protoconid. In the trigonid, the smallest dimension is between the paraconid and the metaconid. The anterior cingulid is wide and continuous from the tip to the anterobuccal flank of the protoconid. The entoconid lies directly posterior to

5 156 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 FIGURE 2. Galadi grandis, sp. nov., lower premolars (holotype QM F23394). A, occlusal view; B, buccal view. Abbreviations: p1 p3, lower premolar 1 to premolar 3. Scale bar equals 2 mm. FIGURE 3. Galadi grandis, sp. nov., lower molars (holotype QM F23394). A, occlusal view; B, buccal view. Abbreviations: acd, anterior cingulid; co, crista obliqua; end, entoconid; hyd, hypoconid; hyld, hypoconulid; m1 m4, lower molar 1 to lower molar 4; mecd, metacristid; med, metaconid; pacd, paracristid; pad, paraconid; ped, preentocristid; phd, posthypocristid; prd, protoconid. Scale bar equals 2 mm. the metaconid. The cristid obliqua terminates midway between the metaconid and protoconid. The angle formed by crests of the hypoconid is reduced. Morphology of m3 is similar to that of m2 except as follows. The protoconid lies directly anterior to the hypoconid. The paracristid and metacristid are longer. The talonid and trigonid are equivalent in width. The cristid obliqua is concave at the anterior end and terminates on the lingual half of the crown. Morphology of m4 is similar to that of m3 except as follows. The trigonid basin is anteroposteriorly widened. The talonid is reduced, but two cusps are retained: hypoconid and entoconid. The crista obliqua and preentocristid meet just anteriorly to the midpoint between the entoconid and hypoconid. The posthypocristid meets the postentocristid posteriorly to the hypoconid. Upper Dentition Occlusal shape of M1 rounded to triangular (Fig. 4). The only specimen is worn. Stylar cusp D is the tallest cusp, followed by stylar cusp B, metacone, stylar cusp E, paracone, stylar cusp A (referred to as anterior cingulum tip in Travouillon et al., 2010), and protocone. The anterior cingu- lum forms a continuous shelf from the anterolabial corner of the tooth to the protocone via a connection with the preprotocrista. Stylar cusp B is posterobuccal to the paracone. In occlusal view, the preparacrista curves posteriorly to stylar cusp B from the paracone. There is no connection of stylar cusp A to the paracone. Stylar cusp A is lower than the parastylar shelf. The stylar shelf crest is continuous from the preparacrista, through the stylar cusp B to the posterobuccal corner of the crown. Stylar cusp D is longer than wide and directly buccal to the metacone. Stylar cusp E is small but prominent. The centrocrista does not breech the ectoloph. The postprotocrista runs parallel to the stylar shelf before curving around the base of the metaconular hypocone. It then thins along the posterior flank of the metacone to terminate as the posterior cingulum just prior the crown s posterior corner. The anterior cingulum continues through the protocone to the posterior cingulum rounding the lingual edge of crown. The buccal flank of the crown is bulbous and rounded.

6 TRAVOUILLON ET AL. FOSSIL BANDICOOTS FROM AUSTRALIA 157 FIGURE 4. Galadi grandis, sp. nov., upper molars (top to bottom, paratypes QM F23395, QM F23396, and QM F23397). A, occlusal view; B, lingual view. Abbreviations: ac, anterior cingulum; cc, centrocrista; ect, ectoloph; M1 M3, upper molar 1 to upper molar 3; mcl, metaconular hypocone; me, metacone; pa, paracone; pc, posterior cingulum; pmc, premetacrista; pomc, postmetacrista; popc, postparacrista; ppc, preparacrista; pr, protocone; prprcr, preprotocrista; psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, D, and E. Scale bar equals 2 mm. Morphology of M2 is similar to that of M1 except as follows. The occlusal outline of the crown is more angular. Stylar cusp B is as large as stylar cusp D. Stylar cusp A is on the parastylar shelf and has a crest joining to the preparacrista. The anterior cingulum is continuous from the base of stylar cusp A to the preprotocrista after forming a weak notch with the anterolabial corner of the tooth. An anterior crest links stylar cusp B to the preparacrista and in occlusal view bends towards stylar cusp A. No posterior crest links stylar cusp B to stylar cusp D. The posterior cingulum is thinner than on M1. The protocone is more angular. The postprotocrista does not run parallel to the stylar shelf; the protocone is the most lingual portion of the crown. M3 morphology is similar to that of M2 except as follows. The crown is much more triangular and angular in occlusal outline. The width is greater than length. Stylar cusp C is small and lies on the anterior flank of stylar cusp D. Stylar cusp E is absent. No continuous crest is present from stylar cusp D to the posterobuccal corner of the crown. The posterior crest of stylar cusp B terminates prior to stylar cusp C. The metaconular hypocone is present but small. The postprotocrista terminates at the posterolingual base of the metaconular hypocone but a ridge runs along the posterolingual flank to the posterior corner of the crown. Measurements of the upper dentition of G. grandis are presented in Table 1 and those of the lower dentition in Table 2. GALADI AMPLUS, sp. nov. (Figs. 5 9) Specific Diagnosis Galadi amplus differs from G. grandis and G. speciosus in the following combination of features: upper and lower third premolar are raised, exposing the anterior roots and reclining toward M1; P3 is wider with a wider lingual shelf extending more anteriorly; postparacrista and premetacrista do not connect on any upper molar to form the centrocrista; stylar cusp E, if present, only on M3; stylar cusp B absent on M4; small anterior cingulid present on m1; distance between paraconid and metaconid is relatively shorter on all molars; lower molar length is relatively shorter but wider at the talonid; large conical entoconid present on m4; infraorbital foramen lies above anterior rather than the posterior root of P3; sagittal crest extends further anteriorly, up to half the length of the frontal; wider and longer canine alveoli with bone thickening; posterior opening of the internal nares wider; and palate more fenestrated than in G. speciosus, with larger maxillopalatine fenestrae and additional pair of maxillary fenestrae. Specific Etymology The species epithet amplus, Latin, means large, in reference to large maxillopalatine fenestrae. Holotype QM F23398, partial skull including fragments of nasal, premaxilla, maxilla, frontal, jugal, palatine, lacrimal, alisphenoid, squamosal, parietal, and basisphenoid. Contains broken M1 and complete but worn M2 4 (Fig. 5). Paratypes QM F23399, right maxillary fragment with C, P3, M2 3 (Figs. 5, 7); QM F56072, right dentary with c, p1 3, m1 4 (Figs. 6 8); QMF23401, left maxillary fragment with M2 4; (Fig. 9); QM F56074, left maxilla with M1 2, M4 (Fig. 9). Referred Material Cleft of Ages Site: QM F56068, left dentary with m1 4. Gag Site: QMF23400, right maxillary fragment with M1 3; QMF23402, right dentary fragment with p3, m2; QMF23403, right dentary with p3 m3; QMF24226, right dentary fragment with p3, m3 4; QM F56069, right maxilla with m2 3; QM F56075, right dentary with p2 3, m1 2; QM F56064, left dentary with m3 4; QMF 23404, left dentary with c, p2, m2 3; QM F56070, right dentary with m3 4. Henk s Hollow Site: QM F56076, isolated left M3. Last Minute Site: QM F56071, right dentary with m2 3; QM F56073, left dentary with m3 4. Wang Site: QMF 36308, left maxilla with M3 4. Type Locality The holotype QM F23398 and paratypes QM F23399 and QMF23401 are from Gag Site, paratype QM F56072 TABLE 1. Measurements (in mm) of the upper dentition of type and referred material of Galadi grandis. P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QM F23395 Camel Sputum Site QM F23396 Camel Sputum Site QM F23397 Camel Sputum Site QM F54575 Camel Sputum Site QM F56062 Ross Scott-Orr Site QM F54574 Wayne s Wok Site QM F56063 Wayne s Wok Site Abbreviations: L, anteroposterior length; M, molar;p, premolar; W, lingual-buccal width.

7 158 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 TABLE 2. Measurements (in mm) of the lower dentition of type and referred material of Galadi grandis. p1 p2 p3 m1 m2 m3 m4 Specimen Riversleigh locality L W L W L W L AW PW L AW PW L AW PW L AW PW QM F23394 Camel Sputum Site QM F54576 Camel Sputum Site QM F54577 Camel Sputum Site QM F54578 Camel Sputum Site QM F54579 Camel Sputum Site QM F30788 Creaser s Rampart Site QM F56066 Mike s Menagerie Site QM F56065 Mike s Menagerie Site QM F56067 Mike s Menagerie Site Abbreviations: AW, anterior width; L, anteroposterior length; m,molar;p, premolar; PW, posterior width. is from Henk s Hollow Site, and paratype QM F56074 is from Last Minute Site. These sites are on Gag Plateau, Riversleigh WHA, northwestern Queensland, Australia. Age and Stratigraphy The sites are part of Riversleigh WHA, northwestern Queensland, and contain terrestrial carbonates that appear to represent tufaceous deposits (Arena, 2004). They are part of Faunal Zone C, estimated to be approximately middle Miocene in age (Travouillon et al., 2006, 2011). Description Facial Skeleton The skull is poorly preserved (Fig. 5). It has short, broad snout, comparable to G. speciosus. The snout is wider than tall and curves ventrally to the tip. The maxilla has strong curvature around the canine root, more so than in G. speciosus and the bone surrounding the canine alveoli is thicker. Anteriorly, in dorsal/ventral view, the rostral end of the nasal is rounded. Posteriorly, the nasal-frontal suture is saw-edged and is posterior to the orbit. A small pointed tongue of frontal extends between the nasal and maxilla. The premaxilla is thin, extending to above P2. The most posterior end of the premaxilla terminates within the maxilla and ends half way through the length of the nasal. The infraorbital foramen is located directly above the anterior root of P3, as opposed to the posterior root in G. speciosus. The lateral posterior surface of this foramen inclines obliquely. The distance between the infraorbital foramen and the lacrimal foramen is short, but longer than in G. speciosus. The nasolabial/antorbital fossa is shallow compared FIGURE 5. Galadi amplus, sp. nov., skull (holotype QM F23398) and maxillary (paratypes QM F23399). A, dorsal view of skull; B, ventral view of skull; C, right lateral view of skull; D, left lateral view of skull; E, right lateral view of maxillary. Abbreviations: aofo, anterorbital fossa; as, alisphenoid; C, upper canine; fr, frontal; fro, foramen rotundum; inf, incisive foramen; iof, infraorbital foramen; ju, jugal; lac, lacrimal; lacf, lacrimal foramen; M2 M4, upper molar 2 to upper molar 4; mxf, maxillary fenestra; mp, maxillopalatine fenestra; mx, maxilla; na, nasal; pa, parietal; pal, palatine; plpf, posterolateral palatal foramen; pmx, premaxilla; ps, presphenoid; pt, pterygoid; P3, upper premolar 3; sg, saggitalcrest; sof, sphenorbital fissure. Scale bar equals 2 cm.

8 TRAVOUILLON ET AL. FOSSIL BANDICOOTS FROM AUSTRALIA 159 FIGURE 6. Galadi amplus, sp. nov., mandible (paratype QM F56072). A, buccal view of right mandible; B, lingual view of right mandible. Abbreviations: c, lower canine; m1 m4, lower molar 1 to lower molar 4; maf, masseteric fossa; manf, mandibular foramen; mas, mandibular symphysis; mf, mental foramen; p1 p3, lower premolar 1 to premolar 3. Scale bar equals 2 cm. with modern peramelemorphians and despite the jugal being poorly preserved, the maxilla-jugal suture is visible as a narrow V-shaped suture. The maxilla drops ventrally in the region of the molars. Very little of the lacrimal extends onto the facial region and surrounds anteriorly the lacrimal foramen, just anterior to the orbit. No distinct lacrimal tubercle or crests are present. Neurocranium Lacrimal sutures are poorly preserved. In dorsal view, the frontal is widest anterior to the orbit, then narrows, and widens again at its posterior end (Fig. 5). A high sagittal crest is present in the posterior region of the frontal. Very little of the parietal is preserved. Contact with the alisphenoid is not visible. The maxillary foramen is large and wholly bounded by the maxilla. The sphenopalatine foramen is large and round. The posterolateral palatal foramen is poorly preserved but would have been larger than in G. speciosus. The sphenorbital fissure is preserved; the foramen rotundum is lacking only its lateral wall. Palate Only the posterior end of one incisive foramen is preserved (Fig. 5). The palate region of the premolars has a pair of small irregularly shaped maxillary fenestrae, extending from P1 to P2. In the molar region, a pair of large maxillopalatine fenestrae is present, oval in shape with a thick dividing septum. They extend from level with the posterior root of P3 to level with the anterior root of M4 Cranial Floor The posterior opening of the internal nares is large, wider than in G. speciosus. Contact between the presphenoid and pterygoid is unclear (Fig. 5). The basicranial region is absent. Dentary Two mental foramina are present, the first lies beneath the anterior root of p1, the second beneath the anterior root of m1 (Fig. 6). The dentary is overall curved anteroposteriorly. The posterior part of the dentary is poorly preserved. The base of the coronoid process extends from the ramus at 100 degrees. Lower Dentition No lower incisors are preserved. The incisor region has alveoli for three single-rooted incisors. As in G. speciosus, the alveolus of i2 is raised to a level above that of i1. The canine is large and laterally flattened (Fig. 6). The canine is tall, well above molars and premolars. The canine and p1 are separated by a short diastema. The morphology of p1, p2, and p3 is very similar, only differing in size, which increases posteriorly (Figs. 6 7A). All premolars have only two cusps, with the major cusp anteriorly positioned above the anterior root. The posterior cusp is directly positioned posterior to the major cusp, and a crest joins the two cusps. This crest on p2 and p3 is heavily worn posteriorly. No cusp or crest lies anterior to the main cusp. The premolar crowns are raised

9 160 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 FIGURE 7. Galadi amplus, sp. nov., lower premolars (paratype QM F56072) and upper third premolar (paratypes QM F23399). A, occlusal view of lower premolars; B, occlusal view of upper third premolar. Abbreviations: p1 p3, lower premolar 1 to premolar 3; P3, upper third premolar. Scale bar equals 2 mm. above the dentary by extension of the roots. The p1 leans anteriorly, whereas p2 and p3 lean posteriorly. The anterior cingulid of m1 is short (Fig. 8). The paraconid is low. The metaconid is taller than the protoconid. The metaconid and protoconid are closer to each other than either is to the paraconid. The metaconid is almost directly lingual to the protoconid. The entoconid is larger than the paraconid and taller than the hypoconid. The entoconid is directly posterior to the metaconid. The hypoconulid is low, posterobuccal to the entoconid, and connected to the hypoconid by the posthypocristid. The cristid obliqua is straight, terminating on the buccal side of the trigonid. The talonid is wider than the trigonid. The hypoconid is directly buccal to the entoconid. The posthypocristid is parallel to the metacristid and is almost perpendicular to the tooth row. The morphology of m2 (Fig. 8) is similar to that of m1 except as follows. The paraconid is closer to the metaconid than either is to the protoconid. The paraconid is thin compared with the metaconid. The metacristid is perpendicular to the tooth row and parallel to the posthypocristid. The anterior cingulid is long and FIGURE 8. Galadi amplus, sp. nov., lower molars (paratype QM F56072). A, occlusal view; B, buccal view. Abbreviations: acd, anterior cingulid; co, crista obliqua; end, entoconid; hyd, hypoconid; hyld, hypoconulid; m1 m4, lower molar 1 to lower molar 4; mecd, metacristid; med, metaconid; pacd, paracristid; pad, paraconid; ped, preentocristid; phd, posthypocristid; prd, protoconid. Scale bar equals 2 mm. widens at the base of the protoconid. The paraconid is lower than the protoconid. The cristid obliqua terminates below the center of the metacristid. There is no connection of the preentocristid to the trigonid. The preentocristid is short, anteriorly directed, terminating at the anterior base of the entoconid. The worn entoconid is wider posteriorly than anteriorly, almost tearshaped. The trigonid and talonid are wider than on m1 but the trigonid remains narrower than the talonid. Morphology of m3 is similar to that of m2 except as follows (Fig. 8). The protoconid is more buccally oriented, lengthening the paracristid and metacristid. The anterior portion of the anterior cingulid is curved laterally. The morphology of m4 is similar to that of m3 except as follows (Fig. 8). The anterior cingulid is wider and straighter. The

10 TRAVOUILLON ET AL. FOSSIL BANDICOOTS FROM AUSTRALIA 161 FIGURE 9. Galadi amplus, sp. nov., upper molars (top to bottom, paratypes QM F56074 and QM F23401). A, occlusal view; B, lingual view. Abbreviations: ac, anterior cingulum; cc, centrocrista; ect, ectoloph; M1 M3, upper molar 1 to upper molar 3; mcl, metaconular hypocone; me, metacone; pa, paracone; pc, posterior cingulum; pmc, premetacrista; pomc, postmetacrista; popc, postparacrista; ppc, preparacrista; pr, protocone; prprcr, preprotocrista; psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, D, and E. Scale bar equals 2 mm.

11 162 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 metaconid is larger and equidistant from paraconid and protoconid. The protoconid is the tallest cusp. The talonid is reduced in both width and length. The hypoconid is more lingually positioned. The entoconid is a conical cusp, taller than the hypoconid. No distinct hypoconulid is present. The cristid obliqua terminates on the lingual side of the trigonid. Upper Dentition The incisor region is not preserved. The canine is strongly curved and large (Fig. 5E). This broken specimen shows no indication of any immediate tapering, suggesting a long length. It is laterally flattened. A short diastema is present between canine and alveoli for P1. P1 or P2 are not preserved. The alveoli of P1 are aligned parallel to the remaining tooth row. A large diastema is present between P1 and P2. The alveoli of P2 are the same as for P1 except the posterior is slightly larger than the anterior. No diastema separates P2 from P3. The crown of P3 is raised well above the maxilla with the roots exposed. The anterior root is raised further than the posterior root so that the tooth reclines against M1. The posterior root is wider than the anterior root. The main cusp is centrally located on the crown (Fig. 7B). A prominent crest joins the main cusp to the posterior-most corner of the crown. No anterior cusp or crest is present. The anterior margin is rounded. A cingulum is present on the posterobuccal surface and is continuous posteriorly to the anterolingual surface. Lingual development of the cingulum is greater than buccally with increased width lingual to the main cusp. In occlusal view, M1 is rounded on all margins (Fig. 9). Its buccal length is greatest. Stylar cusp D is the tallest cusp, followed in decreasing height by the posterolabial corner of the tooth (referred to as the metastylar tip in Travouillon et al., 2010), stylar cusp B, metacone, stylar cusp A, paracone, metaconular hypocone, and protocone. The preparacrista runs buccally from the paracone, then curves posteriorly to join stylar cusp B, continues through stylar cusp D, and ends at the posterolabial corner of the tooth. The postparacrista is straight and obliquely orientated. It ends at the base of the lingual flank of stylar cusp B and does not connect to the premetacrista to form a centrocrista. The premetacrista ends at the anterior base of the flank of stylar cusp D. The postmetacrista is the longest crest and runs from the metacone to the posterolabial corner of the tooth. Stylar cusp A is low, and just anterior to stylar cusp B. The anterior cingulum is a continuous shelf from the anterolabial corner of the tooth to the protocone. The protocone is posterolingual to the paracone. The postprotocrista runs posteriorly to the metaconular hypocone and curves around the base of the metacone, parallel to the postmetacrista, forming a small posterior cingulum, terminating before the posterolabial corner of the tooth. Morphology of M2 is similar to that of M1 except as follows (Fig. 9). The buccal length and anterior width of the crown are subequal. Stylar cusps B and D are larger and conical in shape. No crest is present between stylar cusps A and B. Stylar cusp A is on the parastylar shelf. The preparacrista is long and joins stylar cusp A without connecting to stylar cusp B. The postparacrista and premetacrista terminate at the base of stylar cusp B and stylar cusp D, respectively. A continuous crest runs from the anterior base of stylar cusp D to the posterolabial corner of the tooth. The protocone and metaconular hypocone lie directly lingual to the paracone and metacone, respectively. The preprotocrista runs anteriorly past the base of the paracone to terminate at the base of anterolabial corner of the tooth, forming a complete anterior cingulum. The postprotocristid runs through the metaconular hypocone and bypasses the base of the metacone to terminate on the posterolingual flank of the tooth, forming a posterior cingulum. Morphology of M3 is similar to that of M2 except as follows (Fig. 9). Stylar cusp A is taller than on M2 but lower than stylar cusp B. Stylar cusp B is equal in height to stylar cusp D. Stylar cusp E is present on some specimens. The postprotocrista curves and diverges around the metaconular hypocone. The protocone is further lingual than the metaconular hypocone, which is FIGURE 10. Galadi adversus, sp. nov., upper molars (holotype QM F16911). A, occlusal view; B, lingual view. Abbreviations: ac, anterior cingulum; cc, centrocrista; ect, ectoloph; M1 M3, upper molar 1 to upper molar 3; mcl, metaconular hypocone; me, metacone; pa, paracone; pc, posterior cingulum; pmc, premetacrista; pomc, postmetacrista; popc, postparacrista; ppc, preparacrista; pr, protocone; prprcr, preprotocrista; psprcr, postprotocrista; St A, St B, St C, St D, St E, stylar cusps A, B, C, D, and E. Scale bar equals 2 mm. much smaller than on M2. The postprotocrista is therefore longer and more obliquely orientated than on M2. The preprotocrista is longer. The metastylar shelf has a weak continuous crest joining stylar cusp D to the posterolabial corner of the tooth. No posterior cingulum is present. The morphology of M4 is similar to that of M3 except as follows (Fig. 9). The anterior cingulum is further developed. The paracone is lingually level with the metacone of M3. No stylar cusp B is present and no cusp is present on the posterobuccal edge of the crown. There is no metastylar shelf. The postprotocrista and postparacrista both terminate at the most posterior point on the crown. No metaconular hypocone is present. Measurements of the upper dentition of G. amplus are given in Table 3 and those of the lower dentition in Table 4. GALADI ADVERSUS, sp. nov. (Fig. 10) Specific Diagnosis Galadi adversus differs from G. amplus, G. grandis, andg. speciosus in the following combination of features: infraorbital foramen lies above the posterior root of P3 (as in G. speciosus, above anterior root in G. amplus); postparacrista and premetacrista connect to form a complete centrocrista on M3; postparacrista and premetacrista do not connect to form a complete centrocrista on M1 2; stylar cusp E present on M1 3; stylar cusp B present on M4 (present in G. speciosus but absent in G. amplus); posterior cingulum present on M1 and M2, but not connected to postprotocrista; wider anterior cingulum on M3; very small metaconular hypocone; and wider metastylar shelf, making the overall upper molar shape more triangular in occlusal outline. Specific Etymology Adversus, Latin word meaning opposite, in reference to the centrocrista being complete on M3, and incomplete on M1 2, which is the other way around in Galadi speciosus.

12 TRAVOUILLON ET AL. FOSSIL BANDICOOTS FROM AUSTRALIA 163 TABLE 3. Measurements (in mm) of the upper dentition of type and referred material of Galadi amplus. P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QM F23398 Gag Site QM F23399 Gag Site QM F23400 Gag Site QM F23401 Gag Site QM F56069 Gag Site QM F56076 Henk s Hollow Site QM F56074 Last Minute Site QMF36308 Wang Site Abbreviations: L, anteroposterior length; M, molar;p, premolar; W, lingual-buccal width. Holotype QM F16911, maxillary fragment with M1 4 (Fig. 10). Type Locality Dirk s Towers Site, Riversleigh WHA, northwestern Queensland, Australia. Age and Stratigraphy This site is part of Riversleigh WHA, northwestern Queensland, and contains terrestrial carbonates that appear to represent a vadose cave deposits (Arena, 2004). It is part of Faunal Zone B, estimated to be approximately middle Miocene in age (Travouillon et al., 2006). Description Maxilla The maxilla is poorly preserved. The infraorbital foramen is located above the posterior alveolus of P3. The infraorbital canal is short and the maxillary foramen is wholly bounded by the maxilla. A heavily worn fragment of the jugal meets the maxilla in a narrow V-shaped maxilla-jugal suture. Upper Dentition M1 is longer than wide and its occlusal outline is rounded (Fig. 10). Stylar cusp D is the tallest cusp followed by posterolabial corner of the tooth, stylar cusp E, stylar cusp B, metacone, paracone, stylar cusp A, protocone, and metaconular hypocone. From the paracone, the preparacrista runs bucally toward stylar cusp B, then curves and continues to the posterolabial corner of the tooth through the ectoflexus, stylar cusp D, and stylar cusp E. The postparacrista and premetacrista do not join to form a centrocrista; instead, the postparacrista ends at the base of the posterolingual flank of stylar cusp B and the premetacrista ends at the base of the anterolingual flank of stylar cusp D. From the metacone, the postmetacrista runs to the posterolabial corner of the tooth as the crown s longest crest. Anterior to stylar cusp B, the anterolabial corner of the tooth lies below the parastylar shelf, with a crest running anteroposteriorly to a small stylar cusp A. The anterior cingulum runs from the anterolabial corner of the tooth to the protocone. The postprotocrista runs posteriorly only as far as a small metaconular hypocone. A small posterior cingulum occurs on the posterior flank of the metastylar shelf and runs from level with the metacone to two thirds of the way to the postmetacrista. Morphology of M2 is similar to that of M1 except as follows (Fig. 10). It is shorter wider. Stylar cusps B and D are larger and more conical. Stylar cusp A is on the parastylar shelf and connects to the preparacrista only. A crest runs posterolingually from the tip of stylar cusp B and connects with the postparacrista. The premetacrista connects to a crest at the base of stylar cusp D, which continues posteriorly through stylar cusp E and terminates at the posterolabial corner of the tooth. The metaconular hypocone lies directly lingual to the metacone. Morphology of M3 is similar to that of M2 except as follows (Fig. 9). Stylar cusp A is taller than on M2, but lower than stylar cusps B and D, which are equal in height. No crest runs between stylar cusp D, stylar cusp E, and the posterolabial corner of the tooth. The anterior cingulum is further developed. The metaconular hypocone is larger. No posterior cingulum is present. The postparacrista connects with the premetacrista to form a centrocrista. The morphology of M4 is similar to that of M3 except as follows (Fig. 9). No metastylar shelf is present. Stylar cusp B is present but no other cusp or crest is present on the posterobuccal edge of the crown. No metaconular hypocone is present. The postprotocrista and postparacrista both terminate at the most posterior point on the crown. Measurements of the upper dentition of G. adversus are given in Table 5. RESULTS In the first analysis of 74 craniodental characters included in our matrix, 71 were parsimony informative. Unconstrained TABLE 4. Measurements (in mm) of the lower dentition of type and referred material of Galadi amplus. p1 p2 p3 m1 m2 m3 m4 Specimen Riversleigh locality L W L W L W L AW PW L AW PW L AW PW L AW PW QM F56068 Cleft of Ages Site QM F23402 Gag Site QM F23403 Gag Site QM F23404 Gag Site QM F24226 Gag Site QM F56075 Gag Site QM F56064 Gag Site QM F56070 Gag Site QM F56072 Henk s Hollow Site QM F56071 Last Minute Site QM F56073 Last Minute Site Abbreviations: AW, anterior width; L, anteroposterior length; m, molar;p, premolar; PW, posterior width.

13 164 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 1, 2013 TABLE 5. Measurements (in mm) of the upper dentition of type and referred material of Galadi adversus. P1 P2 P3 M1 M2 M3 M4 Specimen Riversleigh locality L W L W L W L W L W L W L W QMF Dirk s Towers Site Abbreviations: L, anteroposterior length; M, molar;p, premolar; W, lingual-buccal width. parsimony analysis of the matrix recovered 483 most parsimonious trees (tree length = 315 steps; Fig. 11A). All Galadi and Yarala taxa are recovered outside the crown group Peramelemorphia, with the relationship of Peroryctes broadbenti unresolved. The two Yarala species form a weakly supported clade (bootstrap value of 61%), as does crown group Peramelemorphia (bootstrap = 58%). Within the crown group, there is strong support for a clade comprising species of Isoodon, Macrotis, Chaeropus, and the Pliocene taxa Ischnodon australis (bootstrap = 74%), and strong support for monophyly of Macrotis species (bootstrap = 94%). In the second analysis of 74 craniodental characters included in our matrix, 72 were parsimony informative. Unconstrained parsimony analysis of the matrix recovered 15 most parsimonious trees (tree length = 299 steps; Fig. 11B). All Galadi and Yarala taxa are recovered outside crown group Peramelemorphia and form a sister clade to the crown group. Yarala species form a strong clade (bootstrap = 81%), whereas three of the four FIGURE 11. Phylogenetic relationship of Galadi amplus, G. grandis, and G. adversus, spp. nov., based on a 74-character craniodental matrix. Fossil and recently extinct taxa are indicated by. Galadi amplus, G. grandis, and G. adversus, spp. nov., are highlighted in bold. Crown group Peramelemorphia is bracketed. A, Analysis 1 containing all taxa. Strict consensus of 483 most parsimonious trees (tree length = 315; consistency index excluding uninformative characters = ; retention index = ) from unconstrained maximum parsimony analysis of the matrix. Numbers above branches represent bootstrap values (100 replicates). Branch lengths are arbitrary. B, Analysis 2 containing all taxa except Galadi adversus, Perameles sobbei, Ischnodon australis, Numbigilga ernielundeliusi, cf. Peroryctes tedfordi, and cf. Peroryctes sp. Strict consensus of 15 most parsimonious trees (tree length = 299; consistency index excluding uninformative characters = ; retention index = ) from unconstrained maximum parsimony analysis of the matrix. Numbers above branches represent bootstrap values (100 replicates). Branch lengths are arbitrary.