Journal of Vertebrate Paleontology l8(2)::130 '139. June l99lt O 1998 by the Society of Vertebrate Paleontology

XENORHINO.S, A NEW GENUS OF OLD WORLD LEAF-NOSED (MICROCHIROPTERA: ) FROM THE AUSTRALIAN MIOCENE

SUZANNE HAND School of Biological Scicnce. University of New South Wales, Sydney, New South Wales 2052, Australia

ABSTRACT-A new genus and spcciesol'hipposidcrid is describcd fl-om thc Bitesantenn.rrvSitc. Riversleigh,north w,esternQueensland, Austr:rlia. Xenorhino.s hulli. gen. ct sp. nov.. diff'erstionr all othcr hipposideridsin. alrtlttg tlther 1'eatures.its broad rostrum and interorbital rcgion. exceptionallv short palate.constrictccl sphenoidll bridge. and pro- nOuncedrotation of thc rostrunr.lts precisc phylogeneticrclatronships remain obscurc. but it lippearslo hc part ot an early hipposidcridradiation that includesspecics ol' (-oelt4ts.Clocoti.s.'l-riuenttp.s, ltcl Rhitrortt'ttt'ri.r.attd that is u'iclely distributedthroughout the Old World tropics. Fror-nanalogy with liring hipposidcrids.Lhe peculiar rcstral and palatal n.rorphologyol'X. lrulli is probably correlatedwith ultrasounclproduction anclentission. ancl. lt-ss certainly. with size and structureol thc noseleirt.

INTRODUCTION Museum, Brisbane. Stratigraphic nomenclature tor the River- sleigh region lbllows Archer et al. (1994). Acetic acid-processing of Tertiary freshwater limestones from the Riversleigh World Heritage property, Lawn Hill Na- SYSTEMATIC PALEONTOLOGY tional Park, northwestern Queensland, Australia, has produced a number of new late Oligocene of early Pliocene microchirop- Suborder MlcnocHrtt<.rp'nr.RADobson. 1875 teran species(Archer et al., 1994). These bats include hippos- Superfamily RHr^-or.opsotoEnBell, 1836 (Weber, 1928) iderids, megadermatids,molossids, vespertilionids, and embal- Family HtppostoentorEMiller, 1907 lonurids (Sig6 et al., 1982; Hand. 1985, 1990. 1995, 1996. XtNonutt'os,gen. nov. 1997a,b, in press).Hipposiderids are by far the most abundant and diverse chiropteran group represented. Type and Only Species-Xenorhinos hulli, gen. et sp. nov. Living hipposiderids.commonly known as Old World leaf- (Figs. l-2). nosed bats, are mostly cave-dwelling. They are referred to 65 Age and Distribution-Early Miocene of northern Austra- extant species(Koopman, 1994);the most speciosegenus, 11zp- lia. po.sideros, contains 53 species, with eight other genera (Rfti- Diagnosis-The genus diagnosis is the same as that fbr the nonycteri.s, , Paracoelops, , Cloeotis, An' type speciesuntil additional spccics are known. thops, , and Aselliscas) containing one to two species Etymology-The namc ref'ers t. the strange and unique each. The family has an Old World tropical to subtropical mod- stmcture of the rostrum and palate. ern distribution, and apparently similar fossil distribution. Ter- tiary hipposiderids are known fiom rnainly karstic sediments, XtxonutNosHAI-1-1. sD. nov. with the oldest representatives being fiom the middle Eocene Figures 1 2 of Europe, early Oligocene of Arabia. late Oligocene of Aus- Holotype-QM F229 18, skull with left C I. P4-M3 and right tralia, early Miocene of Africa, and Pleistoceneof Asia. Riv- P4-M3. ersleigh's fossil hipposiderids are ref-erable to the genera and Paratypes-QM F22919, right dentary with ml-m3; subgenera Rhinonyt'teris, Riversleigha, , and Bra' QM F2Z92O. left maxillary fragment with Cl-M3; F22921. chipposidero.r(Sig6 et al., l9t12: Hand, 1997a,b). QM right maxillary fiagment with Cl-M3; F22922, left den- The new hipposiderid described here is represented by many QM tary with cl, p4-m3; QM F22923. left dentary with cl-m-l: hundredsof well-preserved skulls, dozens of which have been QM F22924. skull with left and right periotics in situ. prepared.It is so lar known only fiom BitesantennarySite, a Diagnosis-Differing tiom all other hipposiderids in its to be of early Miocene age (Archer et al., deposit interpreted broad rostrum and interorbital region, exceptionally short pal- Riversleigh 1994).This site was the first of the many depositsat ate. constricted sphenoidal bridge, and pronounced rostral ro- (Hand to be interpreted as represcnting an ancient cave-fill et tation. al., 1989). It contains at least eight other hipposiderid species, Locality, Stratigraphic Position, and Age-The Bitesan- a megadermatid , many snails, and rarer marsupials,birds. tennary Site occurs on the northeasternedge of the D Site Pla- reptiles, and frogs. teau. in the Riversleigh World Heritage property, Lawn Hill The new species differs strikingly from all other hipposider- National Park. northwestern Queensland (Hand et al.. 1989; ids in, among other features, its broad rostrum and interorbital Archer et al.. 1994). The 150 square metre deposit is cut into region, its exceptionally shorl palate, pronounced rostral rota- an older, relatively non-fossilil'erouslimestone that may be con- tion, and narrow sphenoidal bridge that completely exposes the tiguous with lirnestonesunderlying or containing three adjacent optic foramen and sphenoidal {issure. Its many autapomorphies Oligo-Miocene fbssil deposits: VIP, Burnt Ofl'ering, and Ne- and probable phylogenetic relationships suppon its ref-erralto a ville's Garden sites.On the basis of stratigraphyand contained new genus. local faunas, the latter have been interpreted to be part of Riv- Skull terminology follows Hand (1993. and Fig. l); dental ersleigh'sSystem B depositsand. as such, early Miocene in age terminology follows Sigd et al. (1982). The prefix QM F ref'ers (Archer et al. 1994; Hand, 1997a, b). to specimensheld in the fossil collections ol- the Queensland Etymology-The species is named for Dr. Leslie S. Hall of 430 HAN D-MIOC ENE HI P P OS ID ERID 431

the University of Qucensland,Brisbane, in recognition of his (?lacrimal foramen; Pedersen (1995) has fbund that the lacrimal pioneering work on all aspects of Australian chiropteran biol- bone does not develop in at least some living hipposiderids).A ogy. smaller, circular foramen opens posteroventrally to this. Medi- Associated Fauna and Taphonomy-The Bitesantennary ally, at the most anteroventralpoint of the orbit, is the post- deposit contains thousandsof bat skulls, limb bones,and snails. palatal foramen. Almost all arc complete, suggesting fossilisation occurred at or Palate-The tooth rows are convergent anteriorly. The deep, very near the point of accumulation. The boundary or contact V- to U-shaped indentation that marks the junction of the palate between the fossiliferous fil| and remnants of the older, non- with the premaxillae extends posteriorly to the level of the mc- fbssiliferous cave wall has been identified at several points tacone of M1. The midline of the palate extends posteriorly around the perimeter of the deposit. For these reasons, the de- level with the posterior tace of M2, resulting in a strikingly posit is interpreted to be a cave-fill (Hand et al., 1989). The short palate, equal in length to 1-1.5 molar lengths. Two pos- many snails of the deposit and what seem to be extensive algal- terolateral indentations extend anteriorly to at least the level of like mats suggest that, for some period during its history, the the paracone of M3. Maximum width of the palate occurs at depositional area was open to light and under water. M3, and palatal foramina are well devcloped adjacent to the Xenorhinos halli is preserved in the deposit with at least cight paracone of M3. Paired, anteroposteriorly elongated foramina other hipposiderids (Hand, 199'7b) and a megadermatid. Less occur adjacent to the anterior face of Ml, close to the border well represented in the Bitesantennary deposit are frogs, Iizards, between the premaxillae and palate. a boid, a stort, a swift, peramelids, a dasyurid, and a bulun- Nasals-The inflated nasal cavities arc incomplctcly divided gamayine macropodid. by a median septum. The septum is fused with the anterior one- third of the dorsal palate, such that the left and right nasal DESCRIPTION passagesare incompletely separated. Further, there is little lat- eral development of the mesethmoid plates so that the choanae Xenorhinos halli is known from many well-preserved skulls or ventral respiratory passagesare not separated by bone from and dentaries. The description of the skull is based primarily the dorsal, olfactory chambers. Limitcd lateral development of on the holotype (QM F22918), but other refered specimens the plates occurs well posterior to the posterior limit of the hard (list available through the author and/or Museum) Queensland palate. These plates extend anteroventrally only fbr a short dis- provide additional matcrial. As in the skull of Brachipposidero,s tance (to a point that is still posterior to the level of the hard nooraleebu"^ (Hand, 1993) from Riversleigh, it has not been palate), but extend dorsally to reach the roof of the nasal cavity possible to determine the precise limits of each bone of Xeno- and anterodorsally branch to almost fill the nasal cavity. In an- rhinos halli due to fusion of elements during development (a terior view, an ethmoidal lattice partially divides each nasal cornmon feature of microchiropteran bats). However, as a guide cavity into an anterolateral rnaxillary chamber and a narrower for interpreting the approximate boundaries between bones, median passage.In this view, a large, paired hook-like swelling, severaljuvenile rhinolophoids were examined (i.e., specimens developed in the ethmoid lattice and protruding into the median of Hipposidero,s diadema, Rhinoktphus megaphyllus, and passageway of each nasal cavity, is conspicuous. Posterodor- Mat:roderma gigas). sally, the median septum meets the anteriorly inclined cribri- form plate. Ventrally, it extends further posteriorly and probably Skull fuses with the vomer, which protrudes very conspicuously be- General Outline and Proportions-The skull is relatively yond the palate. The median septum is only slightly swollen long and broad, with little interorbital constriction. Rostral throughout it length, suggesting that, as in other rhinolophoids, length is more than two-thirds braincase length, maximum ros- the vomeronasal organ was poorly devcloped. The septum car- tral width is almost as great as mastoid width, and interorbital ries a large duct that exits posteriorly at the posterior limit ol- width is more than half rostral (and mastoid) width; rostral the ?vomer. The cribriform plate is sparsely perforatcd dorsally height is subequal to braincase height. The braincase is mod- but not perfbrated at all ventrally. erately inflated, as long as wide, and broadest across the mas- Interorbital and Pterygoid Region-Between the orbital toids (at the level of the posttympanic processes);maximum fossae, the braincase narrows only moderately. The point at zygomatic width is subequal to mastoid width. In lateral view, which the two supraorbitalcrests unite 1i.e.,where thc sagittal rostral rotation (Pedersen, 1993, 1995) or flexure of the head crest originates) is clearly posterior to the point of nan'owest on the basicranial axis (as defined by Freeman, 1984) is pro- constriction. The palatines and pterygoids are evidently ex- nounced, i.e., greater than -25o. tremely reduced such that the roof and side walls of the most Rostrum The rostrum is slightly longer than wide; its lar posterior part of the nasal passage are poorly formed and the eral borders converge anteriorly. The anterior emargination of sphenoidal bridge is narrow, almost splint-like. In ventral view, the nasals, which is marked by a broad median spine, describes the sphenorbital fissure, which is large and circular, is almost a broad arc between the maxillae. Rostral inflations are poorly completely exposed. What are probably pterygoid processesoc- defined; an unpaired nasal foramen (for the noseleaf nerve) oc- cur at a point anterior to the cribriform plate, well anterior to curs in the shallow trough separating the inflations. A more the junction of the supraorbital ridges. These processes have a pronounced depression, pierced by one or two foramina, oceurs secondary flange directed posteroventrally. A nanow bony laterally at the junction of the ?nasal and maxilla (above the bridge separatesthe sphenorbital fissure liom the large, almost level of M1). Supraorbital ridges are poorly developed; they semicircular optic foramen. A splint of bone separates the meet the sagittal crest relatively posteriorly (i.e., at a point al- paired optic fbramina. Anterodorsally, in most specimens.are most level with the posterior roots of the zygoma). The zygo- closely paired fenestrae that lead into the nasal cavity. Imme- matic arch extends onto the face as an antorbital bar. which is diately anterolateral to the sphenorbital Iissure arc two foramina 'wing' roughly unifbrm in width but has a conspicuous devel- that drain the antcrior transverse sinus. In the lateral wall of the oped anterodorsally. The elongated infraorbital fbramen occurs interorbital region, at the level of the pterygoid procc'sscs.is a above M2 3: it is low on the face-level with the root of the large circular foramen ('/sphenopalatine) that leads into the pos- anteroventrallydirected alveolus of Cl. teroventral part of the nasal cavity; anterior to this, the post- In the lateral wall of the face, at the anterodorsal edge of the palatal foramen opens. circumorbital rim and under the anterior attachment point of the Zygornatic Arch-The zygoma has a tall projection that oc- antorbital bar, a large dorsoventrally elongated loramen opens cupies approximately half its lcngth. Thc vcntral margin of ll-,r +-'tz JOURNAL OF VERTEBRATEPALEONTOLOGY, VOL. 18, NO.2, ]998

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oc HAN D-M IOC EN E HI PPOSI D ERID +-1-1 zygomatic arch is only slightly arched and is dorsal to the tooth (vomer). The periotic is bordered posteromedially by the basi- row. The dorsal margin rises nearly vertically to form the tall occipital. Anteromedially, the periotic abuts the basisphenoid; projection that rises to the level of the dorsal edge of the ?lac- there is no lateral extension of the basisphenoid nor medial rimal foramen. Its maximum height is dorsal to the glenoid extension of the petrosal. surface. Its apex is rounded, converging slightly towards the Occipital-The most posterior point of the skull is posterior skull, and its posterior margin convex and vertical to backward- to the junction of the interparietal and supraoccipitals. In lateral sloping. A masseteric scar occurs on the anterolateral ventral view, the supraoccipital is not very convex posteriorly, but the surface of the arch. In dorsal view, the junction between the exoccipitals nonetheless overhang the posteroventrally directed zygomatic arch and antorbital bar is not greatly marked by a foramen magnum. The dorsal edge of the foramen magnum is change in direction, the antorbital bars not diverging posteriorly rounded and not thickened. Lambdoidal crests fbrm a distinct much more than the zygomatic arches proper. The latter diverge ridge marking the most dorsal junction of the interparietal and only slightly posteriorly. supraoccipitals. This parallels the dorsal edge of the fbramen Cranial Vault-The braincase is rather square in shape. It magnum, terminating anteriorly in the region dorsal and lateral is widest at the level of the posttympanic processes,narrowest to the paroccipital process. A supramastoid fbramen opens on in the interorbital area, and highest dorsal and posterior to the the occipital surface dorsal to the ventral condyloid fbssa. postglenoid processes. A sagittal crest is present but low, ex- In ventral view, the contour of the supraoccipital at the level tending anteriorly to the supraorbital ridges and posteriorly to of the Iambdoidal ridge is rounded. The foramen magnum is the lambdoidal crest; it is tallest in the parietal region. There is round, being approximately as deep as wide. Between the oc- moderate development of the lambdoidal crests, which rise pos- cipital condyle and paroccipital process is a deep condyloid teriorly to form a peak with the posterior extremity of the sag- fossa. The paroccipital process is well developed and forms the ittal crest. Approximately two-thirds along the length of the most lateral part of the occipital bone. The tip of the process braincase, within the parietals, is the posterior transverse sinus. articulates with the mastoid. The condyle projects most strongly It tracks laterally, then anterolaterally, before opening extra- ventrally at the posterior extremity, its thickened Iip partially cranially in the posterior part of the postglenoid foramen. Along concealing the condyloid foramen in the anterolateral face. the sinus several small foramina open to the exterior. Ear Region-The ectotympanics and auditory ossicles are Glenoid-The glenoid surface is anterodorsally-posteroven- missing. The anterior part of the middle ear cavity is unossified trally inclined, only slightly concave, and wider than long. The as a pyriform fenestra, a restricted opening between the periotic postglenoid process is equal in height to almost half the length and the squamosal, and the most posterolateral part of the al- of the glenoid surface; its anterior face is anteriorly recurved. isphenoid. There is moderate development of an epitympanic The postglenoid foramen is elongate and opens immediately recess and ectotympanic groove. Posteriorly, the large jugular posteriorly to the postglenoid process. Posteromedial to the gle- foramen and the more medial. narrow basicochlear fissure co- noid surface, the fbramen ovale opens in the alisphenoid. alesce. In the periotic, the cochlear labyrinth is only just visible Temporal Region-The posttympanic process of the squa- through the bone of the promontorium. Three turns of the lab- mosal is developed with a vertical process that forms a slim, yrinth can be seen in section. The mastoid wall is relatively posteromedially directed point. In lateral view, the posteroven- thick but through it can be seen the three semicircular canals. tral part of the lateral face of the periotic is exposed. In the otic The petrosal is slightly enlarged, its width being 2.5 to 3 times region, the squamosal swells slightly laterally and circumscribes the minimum basioccipital width. and indentation that would have accomodated the tympanic ring. The squamosal laterally and ?anterolaterally, the alisphen- Dentition oid anteriorly, and the basisphenoid medially, contribute to the anterior edge of the cavity occupied by the periotic. At the The dental formula of Xenorhinos halli is | ?1/2 Cl/l P2/2 anterolateral corner of the tympanic cavity, the pyriform fenes- M3/3. tra forms a broad gap separating the posterior face of the gle- Upper Dentition-The premaxillae and upper incisors are noid liom the anterolateral wall of the periotic. The posterolat unknown. C1 is tall, slendeq and anteroposteriorly shorI, with eral wall of the skull is deeply notched along an extent anterior a tall posterior accessory cusp and well-developed buccal cin- to the post-tympanic process of the squamosal and posterior to gulum. P2 is positioned in the tooth row such that Cl and P4 the paroccipital process. This notch is filled by the mastoid of are not in contact. P4 is approximately as long as wide, and the periotic. conspicuously smaller (although taller) than Ml. On P4, the Basicranium-The diamond-shaped basisphenoid is slightly anterior and lingual cingulum are sometimes continuous with a concave in its central region. Faint, anteriorly convergent ridges pronounced anterolingual cusp. In other specimens, the well- define the basisphenoid fbssa (and probably mark the boundary developed anterolingual cusp terminates the anterior cingulum, between the alisphenoid and basisphenoid posteriorly). The an- so that the lingual cingulum arises on the cusp as a crest linking terior margin of the basicranial area is constricted. Where and the apex of the cusp to the main cusp (?paracone). On M1 and how the presphenoid joins the vomer is not clear, but the junc- M2, the protofossa is open, and the postprotocrista makes an tion is probably dorsal to the duct that runs through the septum obtuse angle to form a crest that reaches the lingual cingulum.

FIGURE 1. Xenorhirto.shatti. sp. nov., BitesantennarySite, Riversleigh, northwesternQueensland. Holotype, QM F22918, skull with right left P4 M3 and left C1, P4 M3; lateral,dorsal and ventral views. Paratype,QM F22919, right jaw with m1 3; lateral view (revcrsed).Drawings by Jeanette Muirhcad. Bar indicatcs 5 mrn. Abbreviations: AB anteorbital bar, ABW anteorbital bar wing, AP angular process, BO basioccipital, BS basisphenoid.C1 upper canine, CF condyloid fbramen, CFO condyloid fossa, CON condyle, COR coronoid, FM fbrarnen magnum, FNL fbramen fbr noseleaf nervc, FO fbramen ovale, FOR fbramen, G glenoid, IF infraorbital foramen, LR lambdoidal ridge, LVF lateroventral fossa, Ml llrst upper molar, M3 third upper molar, MF mental forarnen, MFO mandibular fossa, NS nasal spine, OC occipital condyle, OF optic fbramen, P2A alveolus fbr second upper premolar, P4 lburth upper premolar, PGF postglenoid foramen, PGP postglenoid process, PLF palatal foramina, PMP prernaxillary process, POP paroccipital process, PP pterygoid process, PPF postpalatal foramcn, PTP post tympanic proccss, PI'SF fbramina of postcrior transversesinus, SB sphenorbitalbridge, SC sagittal crest, SF spenorbitalfissure, SMF supramastoidfbramen, SO supraoccipital, SR supraorbital ridge, VO vonter, ZP zygomatic pro.iection. 434 JOURNAL OF VERTEBRATEPALEONTOLOGY, VOL. ]8, NO.2, 1998

FIGURE 2 Xenorhinos ftalli. sp nov , BitesantennarySite. Riversleigh,northwestcrn Queensland Paratypcs A. QM F2292O.occlusal vicw B. QM F22921. occlusalview C-D'. QM F22922: C. lateral view: D-D'. occlusal view. stcreopair.Bar indicatcs lrnrn HAND-M I O CENE H I P P OS IDERI D 435

(r.nrn) On M1 and M2, a notch is presentin thc lingual margin of the TABLE, l. Skull, dcntary and tooth mcitsurernents of the holo- and paratypes of Xerutrhino.shalli, sp. nov., lrom the Mioccne tooth at the level of the posterior facc of the protofbssa, i.e., type BitcsantcnnarySite, Rivcrsleigh, northwcstern Abbrevia- The Ml Queensland. basal to the lingual edge of the base of the metacone. tions: max., maximum; min.. rninintutn:l. Icngth; w. width; T, trigonid; heel is lingually directed and is better developed than on M2. t. talonid; *. nrcirsuredpcrpcndicular to hard palatet #, measurcd per- Nonetheless,M2 has a well-developed hcel, which is postero- pendicularto basicranium.Nunrbcrs in parenthcscsindicate estimatcs. lingually developed and nanow. M3 is little reduced in length or width, with the prernetacrista being half to threc-quarters of Holo Para Para- Para- thc lcngth of thc paracrista. rype type type type Dentary and Lower Dentition-The dentary symphysis is QM F 22918 22924 22920 22921 vertically oriented, with a conspicuously developed chin. A Greatestskull length (dorsal) 23.9 23.6 beneath the anterior edge of p2. large mental foramen opens Rostral length 9.5 9.2 The coronoid is subequal in height to the tip of cl, and hence Braincasclengtl.t l3.l 13.4 is only about one molar crown height above the apices of m3. Rostral width (rnax.) 9.4 9.6 The condyle is low, equal in hcight to the hypoconids of ml Min. interorbital width .+.-1 ;1.6 3. The angular process is broad and splayed laterally. There are Zygornatic width ( 12.9) width ( l2.l alveoli fbr two pairs of lower incisors.The canine is relatively Mastoid ) Rostral height+' 7.1 p'1. tall and straight, being approximately twice the height of Braincaschcight (nrax.)# 8.2 The p2 is of similar length to pzl but half its height. Lower Pirlate lengtl.r l.-5 t.8 molar length decreasesfrom ml to m3, as does relative length Max. palatalwidth (baseof M3) ;+.tt of thc trigonid. Thc m3 is little rcduced and retainsan entoconid lnterperioticdistance t.6 1.5 and hypoconulid. On ml-3, the protoconid is the tallest cusp, L R followed by the metaconid, entoconid, paraconid, hypoconid, CI-M3 8.0-s 7.10 6.13 and hypoconulid. P4_M3 -5.8 (s.9) ,1.89 5.08 Measurements of the holotype and referred specimens of Xe- MI_M3 1.1 3.93 ,t.0C) norhinos halli are given in Table 1. A study of intraspecific cll 2.19 2.06 1.73 I.6t rnorphologicalvariation, including possiblesexual dimorphism, I .:14 1..19 10 1.34 in thc large samplc of X. halli frorn the Bitesantennary Site is Dt l t.25 | . )l 1.29 22 1.31 l.4t) 34 1.26 in progress. 36 Mll 98 89 t.'73 58 1.58 6'/ o: r.65 68 l.68 '79 '75 PHYLOGENETIC RELATIONSHIPS M2 I t.17 1.46 l.-50 t.11 1.12 t.'79 t.70 1.68 The phylogenetic intcrrelationships of genera and species M3 l t.21 1.28 l 07 1.17 groups in the family Hipposidcridae have recently been exarn- 1.69 | .15 l.6c) 1.60 ined by Hand and Kirsch (in prcss) and Bogdanowicz and Para- Para- Para- Owen (in press). In their PAUP 3.1.1 (Swofford, 1993) and rype rypc type Hennig-86 (Farris, 1988) analyscs,Hand and Kirsch (in press) QM F 229t9 22922 22923 included 30 extant hipposiderid speciesand seven fossil taxa, which they scored fbr 59 discrcte characters (36 cranial, 20 Max. dentary length 12.1 12.9 below rr2 1.8 1.61 1.62 dental, and three skeletal). ln Bogdanowicz and Owen's anal- Dcntary depth ?Condylc height 1.5 4.9 in 57 exlant ysis, 45 metrical and 3O discrete-state characters cl n.r3 I 8.09 8.2:l hipposiderid species (but no fbssil taxa) were exarnined. Al- p4-m3 I 5.92 6.23 though the results of these two studies differed signilicantly nr l-m3 I 1.99 1.81 5.17 from each other in many respects (e.g., in interpreted relation- cll l .3I 1.08 ships among Hipposideros species), they agreed in a number 1.21 l.l1 1.t2 probably p2 I of points, including that the family Hipposideridae is 0.92 monophyletic, that the genus Hipposident"^ is probably para- plI l.16 l.16 phyletic, and that the composition of previously proposed di- 1.02 0.92 visions within Hipposidero.s necds reexamination. Both studies ml I t.12 1.1| 1.85 suggcsted associations of geographically widely separatedtaxa, It 1.00 0.94 t.o2 including species of Coelops and Cloeotis, and Triaenops and wT l.l0 Ir o.72 0.85 Rhinonl,cte ris (see below). wt o.9r 1.lo Xerutrhino,s halli was included in the phylogcnetic study by m2 I 1.61 l.61 1.'71 Hand and Kirsch. The analysis could not unambiguously re- IT o.96 0.89 solve the relationshipsol X. halli. with two alternative,equally wT l .3l t.22 parsimonioushypotheses proposed (Fig. 3). In the first hypoth- Ir 0.61 0.83 wt l.3l 1.16 esis of relationship (Fig. 3A), X. halli appears to be a basal rn3 I 1.6 t.62 l.ll member of a clade that includes species of Anthops, Asellia, IT o.92 0.87 Hipposideros, Palaeophyllophora, and Pseudorhiru.tlophus, and wT 1.23 1.0,1 is not especially closcly related to other Australian fossil hip- Ir 0.12 0.80 posiderids, including species of Bra<:hipposideros, Rhinonyc' wt 1.0.1 1.00 teris, and River.sleig,ha.In the second hypothesis (Fig. 3B-C), X. halli appears to be most closely related to other extinct and/ or endemic Australian taxa, as well as to African species of Triaenops and, more distantly, Cloeoti"^ and Coelops. In both rica, and Cloeoti.spercivuli of southern and eastern Afiica fbrm hypotheses, X. halli lies outside (i.e., is basal to) the large l1ry.r- a group characterised principally by a number of common f'ea- posideros crown group. tures of the noseleaf.Gray (1866) earlier recognisedthe unique- HilI (1982) suggested Ihat Rhinonyc'teris aurantius of north- ness of the noseleaf of Rhinonycterrs, separating the genus from ern Australia, Triaerutp.sspccies of southwestern Asia and Af- other hipposiderids and rhinolophids as the sole member of a 436 TOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. ]8, NO. 2, ]998

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FIGURE 3. Phylogenetic hypothesesof hippctsicleridrelationships presented by Hand and Kirsch (in press)resulting from analysesconrluctcd - on 40 living and extinct taxa and 59 dental, cranial and skcletal characters:A, strict consensusof 4 PAUP trees, all unorderedcharacters (Cl 0.25:8':'.5%, resolution): I]. strict consensusof 8 PAUP trees, some ordered characters1CI = 0.231 82.57o resolution); C. Hennig86 Nelson consensus.unordered charactcrs. HAND-M IOC ENE H I P P OS I D ERI D +-) I distinct group, the Rhinonycterina.Gray's deiinition of the last and rhinolophids, X. halli exhibits pronounced rostral rotation, group, as pointed out by Hill (1982), would also include species the angle between the planes of the hard palate and basicranial of the more recently described genera Triaenops and Cloer.ttis. axis being greater than 25" and hence as large or larger than Hill (1982: 114-l'75) interpreted Triaenops as being possibly that observed in other bats (Freeman, 1984; Pedersen, 1993: closer to Rhinonycteri.s than to Cloeotis, sharing a similar struc- table 2). ture of the interorbital region, the architecture of the jugal The nasal cavities of rhinolophoid bats are radically modified 'resonators' prominence of the zygoma, and curously thickened premaxillae. into that amplify the echolocation call (Suthers et Koopman (1994) included the three genera in his subtribe Rhi- al., 1988). The rostrum morphology of X. halli appears to have nonycterina. been similarly modified. This fossil species has parlicularly vo- Xenorhinos hulli may belong to this group (Fig. 3B-C). Fea- luminous nasal cavities, with incomplete separation between the tures shared by species of Triaenop"^, Rhinonycterls, and Xe- nasal chambers and little bony separation of the buccal and norhinos include the accessorycusp on Cl, little reduction of nasal cavities. The ethmoturbinals are unlike those of all other M3, presence of P2, and apomorphies such as markedly ex- hipposiderids examined. A comparative study of the nasal cav- panded zygoma, and thickened and crested premaxillae. Furtheq ities of hipposiderids, including their structure/function rela- Xenorhinos halli may be more closely related to Triaenops per- tionships as a potentially rich source of phylogenetic informa- sicn.sthan to the living Australian R. aurantius (e.g., Fig. 3B). tion, is in progress. Rostral fontanels, fbund by Pedersen (1995) Apparent apomorphies shared by species of Xenorhinos and between the nasal, rnaxillary, and frontal bones in Old World Triaerutps, but not Rhirutnycteris, include a broad, flattened ros- nasal-emitters, are also large and extensive in X. httlli. Unlike tnrm nearly as deep as the braincase, very broad anterior nares other hipposiderids, X. halii has a broad, but flattened, inter- with a short but distinct median spine on the posterior border, orbital region. Nevertheless, the olf'actory lobe was probably and enlarged petrosals that in diameter are more than twice as not large since the cribiform plate is only sparsely perforated wide as the basioccipital-characters noted by Hill (1982) to and only so dorsally (Bhatnagar and Kallen, 1974), and the be unique to Triaenops. Features shared by species of Xenor- vomeronasal organ, as in other rhinolophoids (Cooper and hinos and Cloeotis include marked constriction of the basi- Bhatnagaq 1976), was probably not well developed. sphenoid, as well as the accessorycusp on Cl, little reduction The palate is extremely short in X. halli (less than l 5 molar of M3, and retention of P2. On the other hand, apparent apo- lengths) as is the ventral (but not dorsal) interorbital region, morphies shared by X. halli and species of Anthops, Asellia, with the pterygoid processes located less than a molar length H ipp o si de ro s, P a la eop hy I Io p ho ra, and P se trdo r hino kt p hus but posterior to the level of M3 and occuring in the rostral port:ion not other Australian fossil taxa (Fig. 3.A), include an elongate of the skull. The nasopharynx was thus also presumably very and medially positioned infraorbital foramen, a mesopterygoid short, probably radically affecting the echolocation call of this roof that lacks a groove, and small P4. ln this hypothesis (Fig. bat. This morphology is unique among bats, and quite unlike 3A), the subtribe Rhinonycterina is interpreted to be paraphy- that lbund in its possible relatives-species of Rhinonycteris, letic and shared features (see above) are intetpreted as symple- Triaenops, and Cloeotis. Marked constriction of the sphenor- siomorphies. bital bridge occurs in Cloeotis percivali, but the rostrum is not The precise relationships of the Australian Miocene Xeno- large nor the palate exceptionally shor1. rhiruts halli to other living and fbssil hipposiderids are not clear. The broad, deep rostrum of X. halli cannot be directly cor- However, X. halli evidently lies outside the large, relatively related with size or complexity of the noseleaf. For example, derived Hipposideros crown group, and within or close to a although living members of the Rhinonycterina (the hipposi- group of old endemic Afiican, Australian, and Asian taxa, in- derid subtribe to which X. halli is probably referable) may be cluding extant species of Triaenops, Rhinonycteris, Cloeotis, identified and clustered on the basis of their distinctive nose- and Coelops. The latter appear to be early but autapomorphi- leaves (HilI, 1982) and less clearly on features of the premax- cally specialized offshoots of the Old World hipposiderid ra- illa, zygoma, basicranium, and dentition (Hand and Kirsch, in diation. press), rostral morphology in this group varies widely, fiom the very small, poorly inflated rostrum of Cloeotis percivali to the DISCUSSION large, inflated rostrum of Rhinonyr:teris aurantius and Triaen- ops persicus. The relative size of the noseleaf also varies among At least some of the most strikingly distinctive characteristics living rhinonycterinae: in Rhinctny<:terls,the noseleaf is rela- of Xenorhinos halli, such as its broad, deep rostrum, volumi- tively large and completely covers the muzzle; in Cloeotis, it nous nasal cavities, broad interorbital region, extremely short is relatively small, and in Triaenops it is narrow, in neither case palate, splint-like sphenoidal bridge, and conspicuous angle in covering the muzzle (Hill, 1982). There are also conspicuous the basicranial axis, are probably specializations correlated with interspecific diff'erences in noseleaf and rostral morphology: echolocation. Pedersen (1993, 1995) found that head posture Triaenops furculus of Madagascar and islands of the western and skull form in bats is constrained by the demands of vocal- Indian Ocean has a much less complex noseleaf but more in- ization during echolocation, with profound differences in skull flated rostrum than Z. persicus of Madagascar, Afica, Iran, morphology between oral- and nasal-emitting bats occurring Oman, and Aden (Hill, 1982). In this hipposiderid subtribe, at early in embryonic development. Nasal emission radically af- least, it appears that size and shape of the rostrum are not di- fects those structures immediately adjacent to the vocal tract, rectly correlated with-nor an accurate predictor of-size and with strong selective forces acting on the form and function of complexity of the noseleaf. The morphology of the noseleaf has the pharynx and larynx during echolocation (Pedersen,1995). been used extensively in systematic studies of extant hipposi- Rotation of the rostrum is one of the most conspicuous cor- derids, particularly in alpha (e.g., Gray, 1866; Peters, nasal emission (Freeman, 1984; Pedersen,1993). In 1871; Tate, t94la, b; Hill, 1963, 1982; Flannery and Colgan, relates of .1986; nasal-emittingbats (hipposiderids,rhinolophids, phyllostomids, 1993; HilI et al., Kitchener et al. 1992), but structural mormoopids, nycterids, megadermatids, and possibly rhino- and functional relationships of the noseleaf are not well under- pomatids), the rostrum is rotated ventrally to align the naso- stood. Pye (1988) suggested that it possibly serves to beam pharynx with the direction of flight, the foramen magnum is ultrasonic calls and thus to contribute to the directionality of moved ventrally, and the inner ear is rotated posteriorly to com- the signal. The complex flaps and folds may also serve to shield pensate fbr the general rotation of the skull ventrally about the the ears from outbound, nasally-emitted signals and so increase cranio-cervical axis (Pedersen,1993). Like other hipposiderids overall sensitivity to returning echoes (Hill and Smith, 1984). 438 JO(IRNAL OF VERTEBRATEPALEONTOLOGY, VOL. ]8, NO. 2, 1998

The dentition of X. halli appears to be relatively unspecial- B. Engesser,H. Felten. T. Flannery, W. Fuchs, L. Gibson, J. E. ized. It is similar to that found in species of Rhinr.tnyc:teris, HiIl, M. Hugueney, P. Jenkins, D. Kitcheneq K. Koopman, P Triaenops, Brachipposidents, Riversleigha, and to a lesser ex- Mein, R. Rachl, B. Sig6, N. B. Simmons, G. Storch, M. Suter- Ient Cloeotis. Thcse bats exhibit a broad range of skull shapes meister, and S. Van Dyck. Two anonymous ref'ereesgreatly im- and morphology but preserve a dentition that is probably ple- proved the manuscript. The Riversleigh project has been sup- siomorphic among hipposiderids. Hipposiderids typically feed ported by the Australian Research Council, the Department of on moths and beetlescaught in the wing or tail membranewhile the Environment, Sport and Territories, National Estate Pro- foraging low over ground and bushes. The unusual structure oI' gramme Grants (Queensland), Queensland National Parks and the rostrum of X. halli, however, suggeststhal this speciesmay Wildlife Service, the Australian Geographic Society, the Lin- have had a unique feeding behavior. nean Society of New South Wales, ICI, the Queensland Mu- At least 12 hipposiderid species are known tiorn the Oligo- seum, and the University of New South Walcs. Miocenc fieshwater Iimestonc deposits at Riversleigh, norlh- western Queensland (Archer et al., 199'1). Xerutrhinos halli is LITERATURE CITED preserved at the early Miocene Bitesantennary Site along with fiogs, lizards, birds, marsupials, a megadermatid bat, and at least Arcl.rer,M., S. J. Hand,and H. Godthelp.1994. Riversleigh: The Story Rainlbrestsof Inland Australia.Reed eight other hipposiderids, representing at least four genera, Xe- of Anirnalsin the Ancient Books,Sydney, 256 pp. norhinos, Riversleigha, Rhinonycteris, and ?Bra<-hipposideros Bhatrragirr,K. P, and Fl C. Kallcn. 197'1.Morphology of thc nasalcav- (Hand, 199'7b, in press). The Bitesantennary hipposiderids, itics and :rssociatcd strllctures in Artilrcus.itrrttttit'ensis and Mlotl.s which appear to have coexisted, range in size from an estimated Iucifitgtts.American Journal ol Anatonry 139:167-190. 20 to 60 mm in fbrearm length (skull lengths 12-24 mm) and Bogdanowicz, W.. and R. D. Owcn. In press. In the Minotaur's laby show a wide range of skull shapes.They appear to be less diverse rinth: a phylogeny lbr the Hipposideridae;iri T. Kunz and P Racey in dental and postcranial morphology. The immediate deposi- (eds.).Proceedings of thc lOth Internati()nalBat RescarchConf'er- tional environment of the Bitesantennary Site is interpreted tcr cnce. Boston. SmithsonianInstitution, Washington. have been a cave developed in Tertiary limestone surrounded by Coopcr. J. Ci.. and K. P Bhatnagar.19-76. Comparative anatomy of the 122:-57| 601. lowland rainforest (Archer ct al. 1994, and see abovc). The clos- vorncronasalcomplcx in bats. Journal of Anatomy Farris. J. S. 1988. Hennig86, Version 1.5.Cornputer Program and Doc- est modern analogy to this environment is found in rainforested umentation.Port Jel-fersonStation. New York. regions o[ southcasternAsia and possibly New Guinea, limestone Flannery, T. F. 1990. of Ncw Cuinca. Robert Brown arnd as many as six hipposiderid species occur sympatrically where Associatcs,Carina, Quccnsland.'139 pp. (Payne et al., 1985; Flanncry, 1990). In many early to middle and D. J. Colgan. 1993.A ncw speciesand two new subspecies Terliary sites in Europe, up to five hipposiderid taxa. representing of Hippo,sidero,i(Chiroptera) fionr wcstern Papua New Cuinea. Re- three genera, are found together (Remy et al., 1987). Xenorhino.s cords of thc Australian Muscul.tt'15:43-57. halli has possible close relatives in other Riversleigh Oligo Mio- Frecrnan.ll 198,1.Functional analysisof largc animalivorous bats (Mi cene deposits (?X. sp. from RSO, CS, and Upper Sites on God- crochiroptera).Biological Journal o1'the Linncan Society 2l:3t37 ,108. thelp Hill), but there appears to be none among the five or six E. 1866. A revision o1'the getterao1'Rhinolophidire, or horsc- hipposiderid specics representedin Riversleigh's Pliocene Rack- Gray. J. shoe bats. Procccdings of thc Zoological Society, London 1866: ham's Roost Site deposit. 8l u3. is indica- The diverse Riversleigh hipposiderid assemblage Hand, S. J. 198-5.Ncw Miocenc mcgaderntatids(Megadermatidae, CI.ri tive of an earlier radiation. A minimum Eocene age for the early roptera) from Australia witl.t conrntcnts on nlcgadernratidphylo- radiation of the tamily Hipposideridae is provided by the age genetics.Australian Malnmalogy 8:-5 43. of relatively derived fbssil taxa from Europe (e.g., species of 1990. Australia's first Tertiary molossid (Microchiroptcra: Mo P se udn rhinolop hu.s and P a lae ophyllo pho ra). How ever, Hand lossidae):its phylogcnetic and biogeographic irnplications.Mem- and Kirsch (in press) have suggestedthat becausc many evi- oirs of thc QueenslandMuseum 28:175-192. (Bnu'hipposide- dently plesiomorphic hipposiderid taxa (e.g., speciesof Asellis- 1993. First skull of a spccies of Hippo.sidarrts ros) (Microchiroptera: Hipposidericlae),fionr Australian Miocenc <:us,Rhinonycteris, Brachippo.sideros, etc.; Fig. 3) occur in the sedimcnts.Memoirs of the Museum 3l l-/9 192. region, the early radiation of the lamily could have Quecnsland Australian 1995.First record of the genusMegadennu Gcol'li-oyl8l0 (Mi- that occurred there. Certainly bats were present in the region at crochiroptera: Megudennu) from Australia. Palaeovcrtcbrata24: time, as the early Eocenc archaeonycteridid Australonycteris 11 66. <:larkae from Murgon, southeastern Queensland, demonstrates 1996. Ncw Miocene and Pliocene tncgadert.natids(Microchi- (Hand et al., 199.1). roptcra: Megaderma) lront Austlalia. with brclaclercomlncnts on The closest relatives <>f Xenorhinos appear to be spread nrcgadcrmatidevolution. (ieobios 29:365 311. (Micro- tbroughout the Old World tropics (e.g., Cloeoti.s from Africa, 1997a. Hipposirleros benrurtlsigai, a new hipposiderid a reconsiderationo1' Coelops fiorn Asia, and Rhinonyr:/eris from Australia), and as chiroptera)frorn the Mioccne ol'Australia and monophyly of related spccies groups. Miinchner Gcowissen widely as the Palearctic and Ethiopian regions (e.g., Brachip' the schafilicheAbhandlungen A 31:13-92. posideros Thcre is evidence ofendemic radiation andTriaenops). 1997b. New Mioccne leaf'-noscdbats (Microchiroptera: Hip- in areas such as Europe and nofihern Australia of hipposiderids posideridae) from Riverslcigh. Queensland. Memoirs of the (Sig6 et a1., during the Paleogene and Neogene rcspectively QuccnslandMuscunr ztl:33.5 3219. 1994; Hand and Kirsch, in press), but less evidence of profound In prcss. Rit,ersleightrwilliansi. a ncw Miocenc hipposidcrid or long-lasting barriers to dispersal for hipposiderids throughout from Riversleigh.Queensland. Alcheringa. thc Tertiary. Thc wide distribution of hipposiderids in the Old and J. A. W. Kirsch. ln prcss.A southernorgin for thc Hippos World, coupled with their relatively good lbssil record, makes ideridac (Microchiroptcra)'?Evidencc l'rom the Australian tbssil (eds.). of the lOth these bats potentially very useful biocorrelators of intercontinen- record; ur T. Kunz and P Racey Proceedings InternationalBat ResearchConl'crence, Boston. Srnithsoniarrlnsti tal Tertiary sediments (Sig6 et al., 1982; Hand 1997b). tution. Washington. M. Archet and H. Ciodthelp. l9lt9. A fbssil hat-rich. Oligo- ACKNOWLEDGMENTS Miocene cavc-fill from Rivcrsleigh Station- northwcstcrn Queens- land. Conf-erenccon Australasian Vertebratc Evoltttion. Palaeon- I thank Michael Archer, Henk Godthelp and Bernard Sig6 for tology and Systematics.Sydney, March 1989. Abstracts:7. their help and encouragement, and the following people for M. Novacck. H. Godthelp. and M. Archcr. 199'1.First Eocenc kindly providing accessto comparativespecitnens in their care: bat from Australia.Journal of VcrtcbratePalcontology l'l:375-38 l. HAN D-M IOC EN E HI PPO SID ERID 439

Hill. J. E. 1963. A rcvision of the genus HipTtosideros.Bulletin of the Rcmy, J. A., J.-Y Crochet, B. Sig6, J. Sudre, L. de Bonis, M. Vianey- British Museum (Natural History), Zoology I l:l-129. Liaud. M. Godinot, J.-L. Hartenberger,B. Lange-Badre, and B. 1982. A review of the leaf-nosed bars Rhinortvcteris, Cloeotis Comte. 1987. Biochronologie des phosphoritesdu Quercy: mise ir and Triaenops (Chiroptcra: Hipposideridac). Bonncr zoologische jour des listes fhuniques et nouveaux gisements de mammifbres Beitriigc33:165-86. fossilcs. Mtinchner CeowissenschaftlicheAbhandlungen, A, 10: and J. D. Smith 19U,1.Bats: A Natural History. Rigby, Adelaide, 169-l88. 213 pp. Sigd. 8., S. J. Hand, and M. Archer. 1982 An Australian Miocene Bra- A. Zubaid. and G. W. H. Davison. 1986.The taxonomy of leafl chipposideros (Mammalia, Chiroptera) related to Miocene repre noseclbats of the Hippositleros bicolor group (Chiroptera: Hippos scntatives from France. Palaeovertebrata 12:149-17 l. ideridae)from southeasternAsia. Mammalia 50:535--5.10. H. Thomas, S. Sen. E. Gheerbrandt,J. Roger, and Z. Al-Su- Kitchener. D. J.. R. A. How. N. K. Cooper, and A. Suyanto. 1992. laimani. 1994.Lcs chiroptbresde Taquah (Oligocbne infdrieur,Sul- Ftipposideros diudcnut (Chiroptera Hipposideridae) in thc Lesser tanat d'Oman). Premier inventairc systdmatique.Mijnchner Geo- Sunda Islancls,Indonesia: taxonorny and geographicrnorphological wissenschlafilicheAbhandlungen A 26:35 48. vrriation. Records of thc Wcstcrn Australian Muscum l6:l-60. Suthers,R. A., Hartley, D- J.. and Wcnstrup, J. J., 1988. The acoustic Kooprnan. K. F: 199'1.Chiroptcra: systcmatics-handhook of zoology. role of trachcal chambers and nasal cavitics in the production of VIII. 60. Mamrnalia:l217. sonirr pulses by the horseshoe bat, Rhinolophus hildebrandti. Jour- Payne, J.. C. M. Francis,and K. Phillips. 198-5.A Field Guidc to the nal of Comparative Physiology A 162:799-813. Mammals of Borneo. Thc SabahSocietl, with World Wildlife Fund Swofford. D. L. 1993. PAUP: PhylogeneticAnalysis Using Parsirnony, Malaysia, Kuala l-umpuq 332 pp. Vcrsion 3.1. Computer Program Distributed by the Illinois Natural Peclersen,S. C. 1993. Cephalometriccorrelates of echolocation in the History Survey. Champaign. Illinois. Chir()ptera.Journal of Morphology 2l 8:85-98. Tate. Ci. H. H. l9,l1a. A revicw of the genus Hiltposideros with special 1995. Ccphalometric correlatesof ccholocation in thc Chirop- ref'erenceto Indo-Australianspecies. Bulletin of the American Mu- tera: II. Fctal developrnent.Journal of Morphology 225:l(\1-123. scum of Natural History 78:353-393. Pctcrs. W. 187l. Uber dic Gattungen und Arten der Huf'eiscnnasen. l94lb. Rcsults of the Archbold Expeditions. No. 36. Remarks preuss. Wiss. Rhinolophi. MonatsberichteK. Akad. 301-332. on some Old World lcaf-nosed bats. American Museum Novitates Noseleavesand bat pulses; p1.r.'791196 in P Nachtigall Pyc. J. 1988. llzl0:l-ll. and P Moore (ed.), Sonar:Proccsses and Performance-Pro- ceeclingsNATO Advanccd Study lnstitute on Animal Sonar Sys- tcrns.Plcnum Press.New York. ReceivedIl December 1996: accaptetl6 Junuarv 1998.