Records of the Western Australial1 Museum Supplement No. 57: 299-306 (1999).

First Australian Pliocene species of (Microchiroptera: )

Suzanne J. Hand and Henk Godthelp School of Biological Science, University of New South Wales, Sydney, NSW 2052; email: [email protected]

Abstract A new Australian Pliocene hipposiderid is described on the basis of maxillary fragments representing at least four individuals recovered from Rackham's Roost Site, Riversleigh World Heritage property, Lawn Hill National Park, northwestern Queensland. Hipposideros wil1sburyorum sp. novo is distinguished from all other Hipposideros species (and all other hipposiderids) by its very large, round infraorbital foramen. Dentally, it shares most similarities with members of the diverse and widespread Hipposideros bicolor group of Hill (1963) as well as Hipposideros megalotis of East Africa and the extinct H. (Syl1desmotis) vetus lineage from the middle Miocene of Morocco and Pliocene of France.

INTRODUCTION only Pliocene deposit, Rackham's Roost Site Tertiary deposits in the Riversleigh World (Archer et al. 1994). A number of other distinctive Heritage property, Lawn Hill National Park, Oligo-Miocene hipposiderid taxa awaits northwestern Queensland are particularly rich in description. the remains of members of the family In this paper, the first species of Hipposideros from Hipposideridae (Sige et al. 1982; Hand 1993, 1997a, the Australian Pliocene is described, based on 1997b, 1997c, 1998a, 1998b). This Old World family maxillary fragments from Rackham's Roost Site. contains 65 living species, referred to the Skull terminology follows Hill (1963) and Hand Hipposideros (53 species) and eight other genera (of (1997a). Dental terminology follows Sige et al. one or two species each): , , (1982). The prefix QM F refers to specimens held in Paracoelops, , Cloeotis, Anthops, , and the fossil collections of the Queensland Museum, (Koopman 1994). Within Hipposideros, a Brisbane. number of species groups is generally recognized (e.g., Gray 1866; Peters 1871; Tate 1941; Hill 1963). SYSTEMATIC PALAEONTOLOGY Recent authors (e.g., Koopman 1994) generally accept Hill's (1963) seven species groups or Authorities for all scientific names not specifically subgroups (i.e., the H. megalotis, bicolor, cyclops, referenced in this paper can be found in Koopman speoris, pratti, armiger, and diadema subgroups), (1994). with some also recognizing the subgenus Suborder Microchiroptera Dobson, 1875 Syndesmotis for H. megalotis (e.g. Legendre 1982). Hill (1963) further grouped Hipposideros species into Superfamily Weber, 1928 three primary divisions (i.e., the bicolor, cyclops and diadema groups) interpreted to represent three Family Hipposideridae Miller, 1907 distinct evolutionary trends within the genus. Recent Australian hipposiderids include five Hipposideros Gray, 1831 species of Hipposideros (representing the bicolor, Hipposideros winsburyorum sp. novo cyclops and diadema groups) and the endemic Figure 1; Table 1 Rhinonicteris aurantius, all of which inhabit warm humid caves in tropical to subtropical latitudes. In Material Examined the Australian (i.e. Riversleigh) Tertiary record several hipposiderid genera or subgenera are Holotype represented: Brachipposideros, Hipposideros, QM F30571, a right maxilla fragment with P4_M2 Rhinonicteris, Miophyllorhina, Xenorhinos and and alveoli for Cl and M3. Riversleigha in Oligo-Miocene deposits (Sige et al. 1982; Hand 1997a, 1997b, 1997c, 1998a, 1998b), and Pa ra types Brachipposideros and Rhinonicteris in Riversleigh's QM F30572, a right maxilla fragment with P4_M2 300 S.J. Hand, H. Godthelp

Figure 1 Hipposideros winsburyorum sp. nov., from the Pliocene Rackham's Roost Site, Riversleigh World Heritage property, Lawn Hill National Park, northwestern Queensland. QM F30571, holotype, right maxilla contain­ ing p4, MI-2. A-A', oblique-occlusal view, stereopair; H, lateral view. Scale = 2 mm.

and alveoli for Cl and M3; QM F30573, a left maxilla because it contains a macropodid similar to fragment with MI and alveoli for p4 and M2-3; QM Protemnodon snewini Bartholomai, 1978 from the F30574, a right maxilla fragment with p4-MI and early Pliocene Bluff Downs Local Fauna of alveolus for Cl and M2; QM F30575, a left maxilla northeastern Queensland (Archer and Wade 1976) fragment with P4_MI; QM F30576, a right maxilla and abundant plesiomorphic murids, including fragment with P4_M2. Zyzomys rackhami Godthelp, 1997, several species of Pseudomys and a number of new genera. The Type Locality, Age and Associated Fauna Rackham's Roost deposit also contains the remains The type locality, Rackham's Roost Site of crustaceans, fish, frogs, lizards, small crocodiles, (Godthelp 1987, 1997; Archer et al. 1989, 1994; snakes, birds, dasyurids, peramelids, a Hand 1995, 1996), occurs at 19°02.09'5., pseudocheirid, an extinct potoroid, two 138°41.60'E. (Global Positioning Satellite device) in emballonurids (Taphozous spp.), at least four other the Riversleigh World Heritage property, Lawn hipposiderids, four vespertilionids, the small Hill National Park, northwestern Queensland. It megadermatid Megaderma richardsi and the large, appears to represent the indurated floor of a long, living Ghost Bat Macroderma gigas (Hand 1995, 1996; narrow cave developed in Cambrian Thorntonia Boles 1998). The very finely broken remains of the Limestone. The sediment is a breccia of tiny, small vertebrates and depressed fractures and mostly fragmented, bones and teeth set in a fine­ impressions in the bones characteristic of Recent grained, pink-coloured (presumably iron-stained) Macroderma gigas canines suggest that the deposit limestone. The deposit covers an area of largely represents the remains of prey accumulated approximately 200 m 2, with a maximum depth of by an early population of that species. Boids, the 0.5 to 1.0 m. emballonurids, hipposiderids and vespertilionids The deposit is interpreted to be Pliocene in age probably cohabited the megadermatid roost; the Australian Pliocene Hipposideros 301

Table 1 Measurements (in mm) of types of bar) that is straight, almost perpendicular to the nov. from the zygoma and of equal thickness throughout (i.e. it Pliocene Rackham's Roost Rlversleigh, northwestern Queensland. I maXinlUm lacks a wing). A foramen Ulacrimal) opens directly length; W wIdth; tooth absent. above the antenor attachment point of the antorbital bar at the edge of the circumorbital rim. Holotype Paratypes The lateroventral fossa, a deep and very broad QMF 30571 30572 30573 30574 30575 30576 fossa in the ventral part of the anterior recess of the orbital fossa, exposes the maxillary covering P' 0.91 086 0.80 0.81 082 the roots of the posterior cheek teeth; in this region, W 1.25 128 120 120 1.10 the maxillary is perforated by many foramina. In M1 1.36 1.34 1.39 1.30 122 128 lateral view, the ventral margin of the orbit extends W 1.34 1.30 1.38 1.35 1.29 1.37 to a point level with the posterior face of MI, its 2 anterolateral rim being low and rounded (rather M 1.25 121 1.18 than raised and sharp). w 1.39 1.39 1.34 The midline of the palate is preserved posteriorly but not anteriorly; it appears, however, to have been quite short. The posterior extension of the macropodid remains were recovered from what midline of the palate is level with the posterior face was possibly an entrance to the cave. of M2; it does not extend posteriorly beyond the toothrow. The palation (Hill 1963) appears to have Diagnosis been shallow but round with no postpalatal spine. The posterolateral indentation (incisura) extends This Hipposideros species differs from all others in anteriorly to at least the level of the posterior face the following combination of features: of M2. The palate's anterior border, for the junction exceptionally large, round infraorbital foramen, with the premaxillae, extends posteriorly at least to low on the face and dorsal to P'-MI; anteriorly the level of the metacone of MI, has a rounded convergent toothrows; anterior margin of palate U­ lateral margin and was evidently a broad U-shape. shaped; loss of p2; p4 wider than long, with shallow Assuming that the anterolateral margins of the lingual cingulum and poorly developed sphenorbital bridge were subparallel (as in most anterolingual cingular cusp; MI with small, hipposiderids), the toothrows would have posterolingually developed heel and posteriorly converged anteriorly. opening protofossa; M2 with little heel The alveolus for Cl is broken anteriorly in QM development; M3 reduced in length and width. F30571 but is complete in QM F30572 and QM Assignment of the new species to the genus F30574. These alveoli indicate that it was not a Hipposideros is discussed in Remarks (below). particularly long tooth, with a relatively rounded rather than flattened root. p2 is completely lacking, Etymology there being no sign of an alveolus for this tooth The species is named for Keith and Janet either within or extruded (lingually or buccally) Winsbury, long-term supporters of the Riversleigh from the toothrow. p3 is also absent, as in all fossil research. hipposiderids. p4 is wider than long, and only just narrower than MI and M2. The lingual cingulum is Description shallow but distinct and virtually complete. The The new taxon is known from six maxillary anterobuccal cingular cusp is poorly developed fragments: the holotype QM F30571 and the five and the anterolingual cingular cusp lacking. paratypes listed above. The material represents at MI-3 all have three roots. In MI2 the heels are least four individuals. All specimens preserve the poorly developed such that the teeth are infraorbital foramen and all clearly lack an alveolus conspicuously shorter lingually than buccally (i.e., for pl. The lower dentition is not yet known. they are not squared). MI has a small but distinct Several dentary fragments of appropriate size, and posterolingually-directed heel, while M2 virtually possibly dental morphology, have been recovered lacks a heel. In these teeth the postprotocrista from among hundreds of Rackham's Roost approaches but does not meet the base of the hipposiderid specimens, but they are not metacone so that the protofossa is open posteriorly, sufficiently complete to confidently assign to noticeably more so in MI than in M2. There is no Hipposideros willsbllryorum (see also Remarks crest issuing from the end point of the 2 below). postprotocrista in either MI or M • The lingual and The infarorbital foramen is extremely large and posterior cingula are continuous in MI2 and the round, with a minimum diameter of 0.5 mm. It is posterolingual cingulum surrounding the heel only situated low on the face and dorsal to P4_M l . It is slightly thickened. TI1e three alveoli for M 3 indicate enclosed by a narrow bar of bone (the antorbital it was reduced in both length and width. The 302 S.J. Hand, H. Godthelp paracone root alveolus occurs on the buccal margin Further, in Coelops species the postprotocrista of the toothrow, but that for the metacone is continues posterobuccally as a metacingulum lingually displaced such that it occurs on the thereby closing the protofossa in MI-3; in species of posterior border of the maxilla. The alveolus for Cloeotis the protofossa is also closed in MI-3, but the metacone root is damaged posteriorly but M3 here the postprotocrista meets the base of the appears to have been between one-half and two­ metacone (as it does also in species of the cyclops thirds the length of MI or M2. The alveolus for the group and Palaeophyllophora). protocone root (also damaged) occurs level with The anterior convergence of the toothrows the protofossa of MI-2, indicating that M3 was also observed in H. winsburyonlm is similar to that seen in signficantly narrower than the other cheekteeth. H. megalotis and members of the bicolor group as well as species of Coelops and Cloeotis. In Cloeotis species Remarks the palate extends posteriorly only to the posterior In lacking p2, the Rackham's Roost hipposiderid face of M2, as in H. winsburyonlm; in Coelops, H. differs from extinct species of Palaeophyllophora, megalotis and in most H. bicolor species it extends Pseudorhinolophus, Brachipposideros and Vaylatsia, further posteriorly. In species of Asellia, Aselliscus, and extant species of Coelops, Paracoelops, Triaenops, Triaenops, Anthops, Brachipposideros, Rhinonicteris and Rhinonicteris, Anthops and Aselliscus. (Paracoelops the H. cyclops and diadema groups the toothrows are megalotis, known only from the type specimen, was subparallel reflecting the typically broad rostrum in not examined in this study; its description suggests these groups. The anterior margin of the palate is U­ it is very similar to species of Coelops.) In this shaped in species of Asellia, Coelops, H. megalotis and feature H. winsburyorum resembles living and some members of the cyclops and diadema groups, as extinct species of Asellia, Cloeotis and Miophyllorhina it is in H. winsburyorum, but is more typically V­ as well as some species of Hipposideros, Le. H. shaped in species of the bicolor group and other megalotis, H. sabanus of the bicolor group, H. stenotis hipposiderid genera. of the cyclops group and the extinct H. (Syndesmotis) Cl morphology distinguishes various vetus lineage of the Miocene of North Africa and hipposiderid groups. Hipposideros megalotis, H. Pliocene of southern France (Lavocat 1961; Sige vetus, and species of Rhinonicteris, Triaenops, 1976; Legendre 1982). Coelops, Cloeotis, Anthops, Brachipposideros, The new hipposiderid further shares with species Xenorhinos and Riversleigha possess a well­ of Asellia and some species of Hipposideros a developed secondary posterior cusp on Cl. Species significantly reduced MJ. In A. tridens, H. megalotis of Aselliscus, Palaeophyllophora and members of the and H. vetus, M3 is significantly reduced such that cyclops group lack this cusp. In Pseudorhinolophus, the premetacrista is lost and the tooth is much Asellia and the H. diadema group, the cusp, if shorter Gust more than one-half the length of MI or present, is generally less well developed. In the M2) and much narrower than P"-M2. In H. sabanus bicolor group, a tall posterior cusp is variably and the French Miocene Asellia mariaetheresae Mein, present and/or developed. The morphology of the 1958, M3 is slightly less reduced, in this way perhaps Cl of H. winsburyorum is not yet known. Most more closely resembling H. winsburyorum. The isolated hipposiderid Cls from the Rackham's Rackham's Roost hipposiderid differs from Roost deposit have a discrete posterior secondary Riversleigh's Miocene Miophyllorhina riversleighensis cusp but some completely lack the secondary cusp. Hand, 1997c (the only other Riversleigh hipposiderid In hipposiderids, the relative length of the Cl lacking P2) in its broad p4 with indistinct alveolus does not necessarily reflect secondary anteroIingual cingular cusp, and poorly developed cusp development (e.g. Coelops and Cloeotis spp.). heel on M2. No dentary fragments have been referred to H. Of hipposiderids lacking p2 and with a reduced winsburyorum. The ascending ramus is M3, only H. megalotis and H. vetus further share taxonomically important in hipposiderids but is with H. winsburyorum a P" that is wider than long missing in all specimens of appropriate size and and lacks an anterolingual cingular cusp. dental morphology (e.g., P2 and M3 reduced) to Hipposideros winsburyorum resembles many potentially represent this species. This part of the species of the bicolor group, as well as H. megalotis dentary is diagnostic for H. megalotis and H. vetus and H. vetus, in its small molar heels, posteriorly (Le. low coronoid process and long and outwardly open protofossae and gracile nature of MI-2. These deflected angular process; Legendre 1982) and teeth differ markedly from the squared MI-2 appears to be characteristic for some species typically found in species of Brachipposideros, groups of Hipposideros (e.g., tall coronoid process Rhinonicteris, Triaenops, Xenorhinos, Riversleigha, in species of the diadema and cyclops groups). Its Asellia, Aselliscus and many (but not all) diadema height and shape are also diagnostic for species of group taxa. Coelops species differ from H. the Brachipposideros-Rhinonicteris lineage (i.e. low winsburyorum in their MI-2 with pronounced heels coronoid process and broad angular process; Sige and very tall posterolingual cingular cusps. et al. 1982; Legendre 1982). Australian Pliocene Hipposideros 303

Hipposidcros winsburyorulll is unique among their phylogenetic analysis of fossil and living hipposiderids in its exceptionally large, rounded hipposiderids, found no clear division between the infraorbital foramen. At least four individuals, all bicolor and megalo/is groups; in some analyses of similar morphology, are represented in the megalo/is appeared to be among the most Rackham's Roost sample, indicating that this is not plesiomorphic of Hipposidcros taxa (figure Ib) while an aberration. The foramen is unlike the typically in others it nested among bicolor taxa (figures la elongate infraorbital foramen characterizing most and 2b). Additionally, they found no special hipposiderids, including Il. mcgalotis and members relationship between H. mcgalotis and the of the bicolor group (notwithstanding the fact that Australian Brachipposideros- I<.h inonic/cris lineage. the fora men was described by Hill (1963) as Bogdanowicz and Owen (1998) also failed to somewhat rounded in H. mcgalotis, H. pygmacus, H. resolve the relationships of megalotis in their cur/us and H. bentus among others). The foramen is phylogenetic analysis of 57 living hipposiderid taxa, also relatively much larger and rounder than in the although in some trees (e.g. figure 3) megalotis spcoris, pratti, annigcr and diadcma groups. clustered with H. sabanlls (among other bicolor taxa). Hipposideros colllmersoni of the diadema group is a In no case, however, did it group with Rhinonicteris notable exception but otherwise differs strikingly allrantills. from H. winsburyorum in its very large size, P4_M2 Hill (1963: 15) described the megalotis-bicolor morphology, and in the position of the infraorbital group as a wide variety of loosely-allied species, foramen (high on the face and dorsal to M 23 rather few of them widespread, and many apparently than very low and dorsal to P4_Ml in H. representing independent lineages. He interpreted winsbllryonml). The foramen is perhaps more them to be the most primitive group in the genus similar to the foramen characterizing many and the group from which the other more members of the cyclops group, which otherwise specialized sections had been derived. Members of differ from H. winsbllryonml in, among other this group are generally smalL with broad, usually features, their distinctive molar morphology and rounded, ears, typically a simple noseleaf, an subparallel toothrows, as discussed above. elongate skull with moderately inflated braincase, a Based on comparisons of tooth size (e.g., Hill narrow rostrum and an unspecialized auditory 1963, table 2), Hipposideros winsbllryorum would be region. Hand and Kirsch (1998) concluded that the similar in size to Hipposideros a/er and H. mcgalo/is genus Hipposideros is probably paraphyletic (see both of which are small hipposiderids with also Hugueney 1965; Sige 1968; Legendre 1982; forearm lengths of 34-38 mm and 33-43 mm Bogdanowicz and Owen 1998) with species of respectively (e.g., Koopman 1994). Other bicolor Asel/ia, Palaeophyliophora and Pseudorhinolophlls species of similar size include H. cineracells (32-37 nesting within it (these three commonly being mm), H. pygmaeus (36-40 mm), H. sabanlls (37-48 associated with cyclops taxa). They agreed, however, mm), H. pomona (37-44 mm), and H. hallophyl/lIs that Hill's (1963) Division 1 taxa (the megalotis­ (35-39 mm). bicolor group) was probably the most plesiomorphic within Hipposideros. The new Pliocene hipposiderid from Rackham's DISCUSSION Roost appears to be a member of the megalotis­ On the basis of its dental formula and dental and bicolor group. However, although H. winsbllryorum palate morphology, the Rackham's Roost is perhaps closer in overall dental morphology to hipposiderid appears to be most similar to East H. megalotis and H. vetus than bicolor taxa such as Africa's Hipposideros megalotis, the Tertiary H. vc/us H. sabanlls, the lack of complete dentaries and/or lineage from North Africa and France, and associated upper canines precludes a more precise members of the H. bicolor group, 33 species of which placement of this Riversleigh hipposiderid within are distributed widely from Africa to northern the megalotis-bicolor group at this stage. Further Australia and Vanuatu. The relationship of megalotis complicating placement is the fact that this species to other Hipposideros species is far from clear. Hill exhibits a complex mix of plesiomorphic and (1963) interpreted mcgalotis to be an early but apomorphic features. For example, the small heel specialized offshoot of the Hipposideros lineage and size of M12 is probably a plesiomorphic feature, the sole living member of the megalotis group, a while the lack of a prominent anterolingual combined mcgalotis-bicolor group comprising his cingular cusp on the broad p 4 is possibly derived Division 1 of Hipposideros. Legendre (1 among hipposiderids (see Hand 1997a; Hand and interpreted megalotis to be a derived offshoot of the Kirsch 1998). The loss of p2 is a derived feature for Brachipposideros radiation, resurrecting the subgenus hipposiderids, evidently independently acquired in Syndesmotis for megalotis and the Miocene H. ve/us a number of separate lineages (e.g., in species of from North Africa and the Pliocene H. sp. cf. H. Asel/ia and Cloeotis) including perhaps the vetlls from southern France (Lavocat 1961; Sige winsbllryonml, megalotis and sabanlls lineages. The 1976; Legendre 1982). Hand and Kirsch (1998), in reduced M 3 is probably derived among 304 S.}. Hand, H. Godthelp hipposiderids (e.g., in Hipposideros species) but is Palaeophyllophora, Hipposideros species appear possibly plesiomorphic in some subgroups (for relatively recently in the world hipposiderid example, species of the H. cyclops group; see Hand record. However, this is possibly an artefact of the 1997a). The short palate with U-shaped anterior Tertiary fossil bat record. Hand and Kirsch margin and convergent toothrows are probably (1998) found Eocene Pseudorhinolophus and also derived features among hipposiderids but Palaeophyllophora to be relatively derived plesiomorphic among Division 1 taxa. The very hipposiderids nesting within Hipposideros. Basal to large infraorbital foramen appears to be an this Hipposideros-dominated clade was a group autapomorphy of this species, although the round containing widely distributed species of shape could be plesiomorphic (being found in Brachipposideros, Rhinonicteris, Coelops, Cloeotis and rhinolophid and megaderrnatid sister-groups). Triaenops among others. If the phylogenetic The unique combination of features exhibited by interpretation by Hand and Kirsch is correct, then H. winsburyorum precludes it from direct ancestry representatives of almost all hipposiderid lineages, of any living Australasian members of the bicolor including the bicolor group, might be expected to group (or indeed any hipposiderid living in the be found in the early Tertiary record, with their Australian Region) including H. ater, a common earliest members possibly in the Asian or cave-dwelling species in the Riversleigh region Australasian region. Sige (1977) and Sige and today. It is likely that skull material will be Legendre (1983) have suggested that the origin of required to more precisely determine the the family was outside Europe, probably in the relationships of H. winsburyorum. Hand and Kirsch Southern Hemisphere (Sige 1991). Unfortunately, (1998) found in their phylogenetic analyses of 37 the Asian fossil bat record is poor and there is a hipposiderid taxa that although dental features (20 profound gap in the Australian fossil characters) alone were insufficient to resolve record from the earliest Eocene to late Oligocene. relationships among hipposiderid genera and Nevertheless, the appearance of a member of the species groups, a combination of cranial and dental highly specialized H. cyclops group in the features (56 characters) provided greater Australian early Miocene (Hand 1997a) indicates resolution. Ongoing taxonomic revisions of that species of Hipposideros had already radiated by Indonesian Hipposideros by Kitchener and co­ that time. authors (e.g., Kitchener et al. 1992; Kitchener and Hipposideros almost certainly had its primary Maryanto 1993; Kitchener and Maharadatunkamsi radiation in Asia or Australasia. The bicolor and 1995; Kitchener et al. 1996) have highlighted the diadema groups are interpreted to be taxonomic problems still to be resolved in this predominantly Indo-Australian-centred (e.g. Hill genus. 1963), while most members of the cyclops group are Hipposideros winsburyorum is the second species of endemics of Australo-Papua (although the roots of Hipposideros described from the Australian Tertiary the latter specialized group possibly lie in Africa: and one of few recorded from the Tertiary Hill 1963; Flannery and Colgan 1993; Hand 1997a; worldwide. Hipposideros bernardsigei was described Hand and Kirsch 1998). The genus appears to have from Riversleigh's early Miocene Neville's Garden radiated rapidly and widely with many closely Site on the basis of near-complete skull material related species pairs living in Asia and Africa (Hill and is the first and only known fossil member of 1963). There appear to have been few barriers to the cyclops group (Hand 1997a). Hipposideros felix dispersal for hipposiderids in the middle and later has been described on the basis of an upper third Tertiary as indicated by the wide distribution of molar and lower first molar from early Miocene species of Brachipposideros and related taxa (Sige et deposits at Li Mae Long, Thailand (Mein and al. 1982, 1994; Hand 1997b, 1998a, 1998b). The long Ginsburg 1997). From East African Miocene history and wide distribution of the H. megalotis deposits, Butler (1969, 1984) recorded three lineage, from contemporary East Africa to middle unnamed hipposiderids, one of which was Miocene North Africa and Pliocene Europe (and reported to be very similar to the European perhaps even Australia), is further evidence of this Pseudorhinolophus bouziguensis. The living H. dispersal capacity. commersoni and Triaenops persicus, and a smaller, By Riversleigh standards, Hipposideros now locally extinct, unidentified hipposiderid, winsburyorum, known from only six maxillary have been reported from the Pliocene of fragments from a single deposit, is poorly Madagascar (Sabatier and Legendre 1985). North represented. Most hipposiderids recovered from Africa's H. (5yndesmotis) vetus and France's H. (5.) Riversleigh's Oligo-Miocene deposits are sp. cf. H. vetus (Lavocat 1961; Sige 1976; Legendre represented by several if not dozens of specimens, 1982) are the only other records of Hipposideros typically skulls. The remains of these older, better­ from the Tertiary. represented hipposiderids are interpreted to have Compared with the early Eocene appearance of gradually accumulated within or adjacent to the European species of Pseudorhinolophus and limestone caves in which they roosted. Hipposideros Australian Pliocene Hipposideros 305

willsbllryorllm, however, appears to have suffered Butler, P.M. (1969). Insectivores and from the the same fate as most Rackham's Roost vertebrates: Miocene of East Africa; new material. In Leakey, to become the finely comminuted prey of the Ghost LSB. (ed.), Fossil vertebrates of Africa, 1: 1-37, Bat Macroderma gigas. Academic Press, New York, N.Y., USA. Butler, P.M. (1984). Macroscelidea, Insectivora and Chiroptera from the Miocene of East Africa. ACKNOWLEDGEMENTS Palaeovertebrata 14: 117-200. Work at Riversleigh has been supported by the Flannery, T.F. and Colgan, D.). (1993). A new species Australian Research Council, the Department of and two new subspecies of Hipposideros (Chiroptera) the Environment, Sport and Territories, National from western Papua New Guinea. Records of the Estate Programme Grants (Queensland), Australian Museum 45: 43-57. Queensland National Parks and Wildlife Service, Godthelp, H. (1987). Riversleigh scene 4: Rackham's Roost the Australian Geographic Society, the Linnean - the beginnings of the modem world. In Hand, 5.). Society of New South Wales, lCI, the Queensland and Archer M. (eds), TIle Antipodean Ark: 81-83, Angus and Robertson, Sydney, New South Wales. Museum and the University of New South Wales. Study of the Riversleigh bat material would not Godthelp, H. (1997). Zyzomys rackhami sp. novo (Rodentia, Muridae), a rockrat from Pliocene Rackham's Roost have been possible without the support and Site, Riversleigh, northwestern Queensland. Memoirs encouragement of Professor Michael Archer of the of the Queensland Museum 41: 329-333. University of New South Wales. The following Gray, ).E. (1866). A revision of the genera of people kindly provided access to comparative Rhinolophidae, or horseshoe bats. Proceedings of the specimens in their institutions: B. Engesser, H. Zoological Society, London 1866: 81-83. Felten, T.F. Flannery, W. Fuchs, L. Gibson, J.E. 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