OldestAustralian vespertilionid(Microchiroptera) from the earty Miocene of Riversleigh,Queensland

HENRI MENU, SUZANNE HAND AND BERNARD SIGE

MnNU.H.. Hern. S. J. & Srr;E,8.,30.5.2002.oldest Australianvespertilionld (Microchiroptera) liorn the early Miocene of Rivcrsleigh, Queensland.Alcheringa 26, 3t9-331. ISSN 03 I I -55llt.

An upper molar of a srnall , here

llenri Menu.20 rue des Granrls Clos,5l220 Cauroy lis Hermonville,France; suzanne oJ' Nev South IIontl Is.hanrl(glunsw-etlu.au],Schctot o./' Birttogical Science, Unit'ersity Ittales, Sydnei, New South ll/oles 2052; Bernard Sigi, Centre de Pal6onto.logre stratigraphique et Palioicologie, IJnit'ersirt ()laude-Bernard,Lyon I' 27-43 Bouiet'ard ret'isetl du I I Not,embre,69622 Villeurbanne cetlex, F'ronce; receit'ed 12.2.2001, 21 11.200l.

Keywonls: Vespertilionidae,I'euconoe, bat, carly Miocene, Riversleigh' Australla

TERTIARY freshwaterlimestone deposits in the rhinolophids arethe only modernAustralian RiversleighWorld Heritagefossil properfy,Lawn not representedin Riversleigh'sTertiary deposits' Hill National Park, northwesternQueensland are A singletooth recoveredfrom Riversleigh's rich in the remainsof vertebratesand particularly MioceneRV S\te(Araheretal' 1989,1994;Creaser microchiropteranbats. Some 40 bat specieshave 1997)is referableto thefamily Vespertilionidae, now beenidentified from lateOligocene to middle making it the oldestknown vespertilionidfossil Miocenesediments and another13 speciesfrom from Australia.Vespertilionids form the largest thePliocene Rackham's Roost Site cave deposit. componentof living Australianbats, numbering Thesebat species include at least25 hipposiderid 40 of 90 species(Duncan et al. 1999),andare by species of the six genera Hipposideros, far themost speciose bat f-amilyin theworld (more Brachipposideros,Rhinonicteris, Riversleigha, than300 species). Their fossil record in Australia, M i op hyI I o r hin a andXe n o r hi no s (Sig6e t al. 19 82; however,is comparativelypoor. The four Pliocene Hand 1997a,b, c, 1998a,b); eightmegadermatids Rackham'sRoost vespertilionids and, possibly, representing at least two genera (Macroderma a singlelower molar from thePliocene Hamilton andMegaderma; Hand 1985,1995, 1996); five Local Faunaof Victoria (Turnbull & Lundelius molossidsof at leasttwo genera(Petramops and 1970) are the only other Australian Tertiary Roost Motmopterus lHydromopsl; Hand I 990, Hand e/ vespertilionid records.The Rackham's al. 1997)and severalrepresentatives ofthe New vespertilionidsare very poorly representedbut Zealandfamily Mystacinidae (Hand et al. 1998, at leasttwo of the four are probably referable to 2001).In thePliocene Rackham's Roost deposit, the endemic Australian generaScotorepens five hipposiderids,two megadermatids,two Troughton,| 94 4 andC h al inol ob us Peters, 1 866' emballonuridsand four vespertilionidshave been In this paper, the tooth from RV Site is reported(Archer et a|.1994;Hand 1995,1996; describedand is refened to the cosmopolitan Hand & Godthelp 1999). Pteropodidsand vespertilionineLeuconoe Boie. I 830.fwo species 031l/ssl8/2002/01319-13 $3.00 0 AAP of which occur today in easternAustralia 3r0 MENUETAL ALCHERINGA

(Kitcheneret al. 1995.Duncan et ol. 1999). Systematicsand dental terminology follow Menu (1985,1987, 1988). Riversleigh stratigraphy tbllowsArcher et al. (1989,1994)and Creaser 0997\.

Systematicpalaeontology

SuborderMICROCHIROPTERA Dobson. 1 875 SuperfarnilyVESPERTILIONOIDEA Gray, I 82l (Weber,1928) Family \GSPERTILIONIDAE Gray, I 82I SubfamilyVESPERTILIONINAE Grav. I 821 (Webea1928)

LeuconoeBoie, 1 830

Typespecies. Leuconoe daubentoni (Laisler in Kuhl1818-19)

Remarks. Originally described as a genusby Boie (1830),Leuconoe has been regardedas a subgenusof Myotis Kaup, 1829 by most bat taxonomists(e.g. Koopman 1993). However. on the basis of tooth morphology, regrettably neglectedby bat neontologists,Leuconoe was raisedagain to thelevel of genusby Menu ( I 985, 1987).When examinedin fine detail, as in the standardpalaeontological approach, tooth structuresprovide the best available common ground for studiesinvolving fossil and living bats. On this revisedbasis, Leuconoe includes many living speciesonce assigned to the supra- specificlaxa Leuconoe, Se/ysi r.rs Bonaparte, I 84I ; I so t us Kolenati, 18 5 6 ; P antmy o t i s Bianchi, I 9 I 6 ; c RickettiaBianchi,1916; CistugoThomas, 19 l2; ChrysopteronJentink, 1910; and Myotis (Menu 1987,p.83,emend. Menu 1988)and, hence, most fossil and living speciesattributed to Myotis sensulato within recentaccounts and revisions (e.g.Koopman 1993).The remaininggenera amongthe leuconoformes(as definedby Menu 1987) areMyotis sensustricto, perimyotisMenu, 1984and Pizo,nyx Miller, 1906. To reiterateMenu ( 1985,1987), and as a partial differential diagnosis,the dental charactersof Lcuconoeare as follows: ALCITERINGA ITARLYMIOCENE VESPERTILIONID FROM RIVERSI-EIGIi 321

Upper tooth row: Ir at various evolutionary thanSite D (SystemA assemblageof Archerct stages(see Menu 1987),trending to linkedbilidy a1.1989), close to the level of RSOSite (low System and narrowness;Ir trending to reducedcrown B), andpossibly belou' CS and Upper Sites (rnid width; Cr rvith oval cross section at base, SystemB) (Creaser1997). On the basisof its cingulumwith completeplane and without lateral stratigraphicposition and its containedmammal 'crowns projections:Pi oval.unicuspate and P' fauna,the RV depositis interpretedas early 'Ihis trendingto reduction;Pr with subtriangularbase Miocene. is apparentlyslightly younger and cusptip median:M r. al variousslages. than the South AustralianKutjamarpu Local morphologictypes trending to paralophmissing, Fauna(Woodbume e/ a/. 1985)wl.rich. c-rn the metaconuleregressed, diversifi ed postprotocrista basis of biostratigraphyand magneto- orientation.metaloph vanishing and protofossa stratigraphy.has been estimated at probably' latest openingposteriorly; Mr variousstages trending Oligocene(Woodbume ct ul.1993). to regressedmesostyle and metacone. Vertebrateremains presen'ed in theRV deposit Lowcr tooth row: incisorsoverlapping; I1-r aregenerally fiagmentary and cl-small to medium- long,narrow, pectiniform with 3 or4 denticles,12 sizedanimals. t heyinclude a skink.a madtsoiid with lingualbase of crown swollen;Ir thicker, snake.a smallcrocodile. a chelid.a peramelid.a i.l'ithrhomboid outline, buccal cusps:mesial pseudocheirid.the phascolarclidNimiokoulu strong.central isolated, distal regressed, lingual grc.vsttmesiBlack & Archer.1997. an undescribed cuspwell scparated:C1 with oval crown basc. molossid and at leasttwo hipposiderids, tip in medianaxis of thetooth, weak lateral cre sts. Rhinonitteri,stedf'ordi Hand. 1997aand at variousstages increasing size comparedto M iotrthy' II o ri nu r':'e rs I e i gheirsls Hand, 1997b. Pa;P2-3 very variable,with oval outline regular , -Ihe cingulum.tip well in medianaxis, P3 trending to Dc.scription. toothis thatof a smallspecies reduction,Pa outline varying fiom subrecta:-rgular of bat.It is 1.38mm longand 1.70 wide. to subtriangularwithout attainingthis stage;C 1- A number of featuresindicate that the P4 row with the tips of the cuspswell aligned; specimenis a firstmolar, Ml , ratherthan a sccond. M1-2strictly myotodont; M3 with correlatively M2. These f-eaturesinclude the follor',lng: the reducedentoconid and talonid. buccalflexus is slightlytilted on thetransverse axis; the posteriorV of the ectolophis deeper Leuconoesp. thanthe anterior one: although slightly damaged, themetacone is tallerthan the paracone; and the Referredmaterial. QM F30577.a subcomplete lengtir/widthratio x 100is high (c. 8l ). lefi MI The buccalflexus has rather rcgular shallolv slopeson both sidesof thebuccally con'u'ex and Loculity, strutigruphic position, age untl low-cuspedmesostyle. The sonrewhatslighter ctssociatedfctuna. RV Site occurs within the post-ectoflexushas a weak.flat cingulum At the 1'ertiarysequence of freshwaterlimestone anteriorcorner. the parastyleis a mesially depositsoutcropping on Godthelp'sHill, projecting,declining crest. It hasrather sn.rall fuversleighWorld tleritageproperty, rn Lawn Hill proportions.At theposterior corner. the metastyle NationalPark, nor-thwestem Queensland (Archer is the declining crestiform end of the et ul. 1989,i 994;Creaser I 997).It occursin the postmetacrista. SystemB sequenceas defined by Archeret ul. A wide regular separatesthe paracone "'alley (1989,1994) and discussed by Creaser(1997).lt and metaconecusps. It openslingually in the is interpretedas being stratigraphicallyhigher roundedand deepprotof'ossa. l here is nrt

World llentagc Flg. /. A-C, !.euconoesp., QM F30577,subcornplete LMr. lionr RV Sitc, GodthelpHill, Riversleigh C. lingual propety, Lawn Hill NationalPark, no(hwestcm Queensland, Australta. A, occlusalview; B. posteriorvieu'; vicw. Scalebar: I mm. L.M. del. 322 MENUETAL. ALCHEKIAIGA

continuousand regular,being taller at the lingual comer,where it isjoined by thedistal crest of the metaconule.

Comparisons

The single bat tooth describedand illustrated here exhibits a morphology which differs from those of all other bats reported from Australian Tertiary deposits.In contrast,the specimen exhibitsgeneral similarity to homologousteeth (M') of many Recentand fossil bats referredto the Vespertilionidae Thesebats constitutea very diversefamily which has been defined and examined in successiveaccounts (e.g. Boie 1830,Miller & Allen 1928,Tate l94l,Thomas 1904,1915) and also has been the subject of revision and re- classification(e.g. Dobson 1878,Miller 1907, Ognev 1928,Simpson 1945, Koopman 1993).' Fig 2 Leuconoesp, SEM photographof a cast of eM Whereassome of its subgroupshave been given- F30511 occlusal , view Max width I 70 mm familial rank (e.g miniopterids, kerivoulids), conslstentparaloph and metaloph, nor a others are consideredto be subfamilies (e.g. differentiated paraconule, although faint signs murinines, tomopeatinines).Whatever their are seen under oblique lighting. Thus the taxonomic rank, all these subgroups can be paraconeand metaconehave smooth linsual distinguishedon the basisof their dentitions. slopes and the protofossa freely op-ens The larger remaining group is the posteriorly. homogeneousvespertilionine subfamily. Studies Theprotocone is a strongcusp, almost as tall of the dentitionsofRecent vespertilionines(Menu as the paracone.From the protocone tip, the 1985, 1987)indicate that boundariesbetween preprotocrlstaextends first mesially then dentalmorphologies of componentgenera are continuouslyto thelingual base of theparastyle. blurred, with completetooth rows often required At this point a small notch marks the contact to discriminatebetween genera. Similar tooth point with the buccal crest of p4. From the morphologiesare sharedby somegenera which protocone tip, the postprotocristaextends, are clearly separatedon other evidence.Very decliningdistally, rising againto a metaconule often, distinctive patterns of particular dental cusp,and then decliningdistally once more to features are sharedby some genera,while they reach the posterior cingular border. The are associatedwith other dental featuresin very protocone gives the lingual region its abrupt varied combinations of patterns.This reflects a mesialslope. At the lurgualbase of the protocone, richly mosaicevolutionary process, as in all bats, the cingulum is briefly intemrpted. and in turn this greatly reducesthe power of The metaconuleis a distinct, transversely Cuvier's (1825, p. 95 et seq.) correlationof compressedcusp. It hasa roundedbase and it is charactersprinciple. Furthermore, intraspecific well separatedfrom the protocone by a lingual variation is not negligible. This makes valley ascendingto the postprotocristaand a assessmentof genericrelationships difficult on correspondingenlargement of the lingual cmgular the basis of an isolated tooth, such as the plane. The posterior cingulum border is Riversleigh upper molar, which is currently the AI,CHERINGA I:ARI-Y MIOCENEVESPERTII -IONID F I{OM RI\'ERSI.F.IGH 323

only specimenof this distinctivcbat. The M of speciesof sonle.but not all. Neverthelcss.comparisons of molar endemicAustralian vespertilioninc genera also morphologyreveal cl

broaderprotofossa, with the metaconuleless Whereasprimitive andother vespertiiionoid bats cuspidateand developedas a small but distinct suchas paleochiropterygids (including Stehlinia, lingualheel. There is significantvariation in Mr asrecently argued by Sig6 I 997) or philisidsare morphologybetween Vespadelus species: in reportedfrom the early Tertiary, leuconoforme somethe metaconule is low andcrestiform rather vespertilionineshave been describedmostly thancuspidate (e.9. V. pumilus fGray, 1841]), in from late Paleogeneand Neogenedeposits of some it is cuspidateand quite similar to the Europeand westemAsia. In Europe,lacustrine Riversleightooth in this respect(e.g. V. depositsor karsticfillings of variousages have durlingtonifAllen, 1933]), in someit is tiny and yielded remainsdescribed or quoted as Mycttis buccallysituated (e.9. V. yulturnusfThomas, or more recentlyLeuconoe or leuconoforme 19l4l), and in othersit is developedinto a species(e.g. Revilliod 1922, Mein 1964,Sig6 & posterolingualheel, as in V. regulus (Thomas, Menu I992^1995,Sig6 et al. 1997,Liegler2000). 1e06). Close relationshipsto Leuconoehave been Othernon-Australian vespertilionine groups presumed(Sig6 & Menu 1995)for a single (or some memberstherein) which have an Mr Oligoceneupper molar from centralKazakstan resemblingthe Riversleightooth include (Gabunia& Gabunia1987), and also for restricted EudiscopusConisbee, 1953; Hypsugo Kolenati, late Eocenematerial fiom Le Batut locality in 1856; L/espertilioLtnnaeus, 1758; Eptesicus southwesternFrance (Muratet et al.1985,pl. I Rafinesque,1820 Nyctalu.sBowdich, 1825; fig. 8).These are generally small to medium-sized Ci.stugoThomas, 1912; Chrysopteron and species.The distinctiveLeuconoe upper molar LaephotisThomas, I 90 I . Hor.vever,none in this morphologyis clearlyexhibited by thelate heterogeneousgroup resembles the Riversleigh speciesfiom Le Batut,as well asby the lateearly tooth as closelyas certainspecies of Leuconoe OligoceneZe uconoe lavocnli Sige & Menu, I 992 includingL. mystucinusand L. californicus. from Le Garouillaslocality in theFrench Quercy phosphorites(Sige & Menu 1995) and contemporaneousZ. salodorensis from Comparisons with Australian Tertiary vespertilionines Oensingenin theSwiss Molasse basin (Revilliod 1922). As discussedabove. the only other Australian The M' from the RiversleighRV site and the Tertiary vespertilionidrecords are Pliocene. homologoustooth from Le Batut locality share Theseare a single,un-named lower molar from several features(general proportions and theHamilton Local Faunaof Victoria (Tumbull & structure;occlusal outline; proportions and Lundelius 1970) and four taxa representedby opening of the ectoloph valleys; shapeof the dentaryand maxilla fragmentsand isolatedupper protofossa;proportions of the protocone, viz. and lower teethrecovered from the Rackham's its weak lingual extentand abrupt lingual wall; Roostdeposit (Archer et al. 1994).At leasttwo wide continuousanterior border made by the of the four Rackham'sRoost vespertilionids preprotocristaextending to become the appearto belong to the Australian endemic paracingulum;intemrpted lingual cingulum below generaScotorepens and Chalinolobus,but it is the protocone;presence ofa thin paraloph)and not yet clear whether they representextant very similar dimensions.Nevertheless many species.These Pliocene Mrspecimens dilfer from featuresdistinguish the Le Bafut species(a weak the N{iocenetooth in their lower, lesscuspidate, but almostcontinuous cingulum along the buccal often crestiformmetaconule and lack of deeo edge;a strongerand more projectedparastyle; a lurrowseparating the protocone and metaconule larger and bulbous mesostyle;the posterior (seealso above). buccaledge being more labially projected). Other significant featuresinvolve the postprotocrista Compari.sonsv,ith Tertiary leuconoformes and related structures:this crest extends in a ALCHEMNGA EARLYMIOCENEVESPERTILIONIDFROMRI\'ERSLEIGH 325

postero-buccaldirection, joining the basal to the protocone;the para- and metalophmore posteriorcingulum (metacingulum);there is no distinctand more oblique antero-lingually; these differentiatedmetaconule and the crest profile crestsand a protoconeslight median lingual keel has no notch posteriorto the protocone;a providing sliding guidesfor deep occlusionof metalophis presentand the protofbssais closed the hypoconidof the lower molar into the posteriorly;the lingual flank of the protoconels protofossa. not depressedposterior to the protocone;and In contrast,and of more immediateinterest, possesses the postcingulumis more distinct and regular. theLeuconoe sp.2 from Bouzigues-l These differencesrelate to a less derived an M' which is structurallyclose to that of the conditionin the Eocenespecies, consistent lvith Riversleighspecies. The most representative its olderage. specimenis figured(Sig6 1968, ftg.21) but the Leuconoespecies so f'ardescribed from the conspecificityof theM'(frg. 28) is doubtful.The Oligoceneof Europeshare some of the similarities Europeanand Australian forms share the and diff-erencesreported above betweenthe following characters:M without marked RiversleighRV and Le Batut species.Further transverseshape; buccal edge with slight pre- detailscan be noted by consideringthe best and postectoflexusnotches; proportions of the documentedspecies up to now, L. lavocati. ln ectoloph and protofossa;and above all the this species,the postprotocristaends lingual to postprotocristajoining a well differentiatedand thebase of themetacone or evenjoins it; if not, a inflated metaconule,well separatedfrom the metaloph is presentand in either case the protoconeby a marked furrow on the lingual protofossais closedposteriorly; there is no flank. Neverthelessthe Bouzigues-I species distinct metaconule.nevertheless this structure shows some differences,including the distinct is suggestedby a slight bulge of the posterior remainsof a buccal cingulum, the more flank and a slightdepression ofthe iingual flank difl'erentiatedparastyle, the stronger and bulbous posteriorto the protocone.This undoubtediy metaconule.the absenceof distinctcontinuation representsa transitionalevolutionary stage, of the postprotocristabeyond the metaconule, anticipatinga rvell differentiatedmetaconule. thelack of a metaloph,the wide posterior opening Among younger representatlvesare ofthe protofossa,and the lingual basal cingulum numerousspecies fiom Miocene,Pliocene and being not erasedposterior to the protocone. reflect relationships" Quaternarykarstic or openbasin deposits of the Theseconditions both close lar:geperi-Mediterranean area (see Sig6 & of presumablephylogenetic meaning and supra- Legendre1983 for a survey).Diversity is well specific (subgeneric'7)value, as weil as a establishedin theearly Miocene localities, where departure,the Bouzigues-1species being more severalspecies occur which aremore or lessclose conservativein someaspects (buccal and iingual to a generalizedLeuconoe tlpe (Sig6et al 1997). cingula and strongparasfyle preserved) and more Thesespecies are close in age to that fiom the derivedin otheraspects (stronger metaconule. Riversleigh RV site. In the single site ot protofossalargely open backwards). Here agatn, Bouzigues-1,some speciessuch as Leuconoe a mosaic pattern of characterdistribution is sp. I and 3 clearly difl'er from the Riversleigh apparent.Also significant is the shared speciesin thefollorving features: more transverse occurrenceofbat generain westernEurope and occlusalshape; more crestiformstructure; pre- Australia during the late Oligocene and early and postectoflexusshorter and deeperand Miocene.now includingLeuconoe as well as conelativelymore individualizedmesostyle; more Hydromops (Hand et al. 1991) and projectingparastyle; narrower valleys of the Brachipposidero.r(Sig6 et al.1982). Thesefacts ectoloph;postprotocrista ending lingual to the revealthe precocious. broad. Oid World dispersal metacone,as an incipient metaconulewithout of somebats belonging to a moremodem tropicaL traceof depressionof the linguai flank posterior bat fauna, clearly derived u'ith respectto the Jlo MENUETAL. ALCHERINGA

previous,late Eocene and early Oligocene fauna Australianvespertilionid relationships asknown fiom the fossilrecord. Ziegler(2000) described two palaeokarsticbat anddispersal faunasfrom Herrlingen8 & 9 localities,Germany, that are late Oligocene(MP 29 of the European Despite their poor Australian fbssil record, PaleogeneMammals scale). These faunas include vespertilionidscomprise almost half the extant three leuconoformevespertilionine species. In Australianbat species.Of the 1I vespertilionid consideringthe genericidentity ofsuch species, sen.sulato generanow occurring in Australia, the author attemptedto discriminatebetween approximatelyhalf (Leuconoe,Minioptarus then.ron thebasis of a frameworkof dentalfeatures Bonaparte,1837 , Kerivoula Gray, 1.842,Murinu adaptedfrom the one used in the presentstudy Gray, I 842 andPipi,strellus Kaup.l 829)have an (Menu1985, 1987). In additionto somebasic upper Asian or w'orld-widedistribution. The other genera(Nyct molar features,Ziegler's key used other tooth oph i Ius, C hal ino I ob us, Vesp ud e I us, characterssuch as relative height and crest Scotorepens,Scoteanax and Fulsistrellus)are sharpnessof the lower canine,and the length/ restrictedto Australia,New Zealand.NewGuinea width ratio of the last upper molar (a rather and nearby islands.Traditional understanding allometricelement). Ziegler's generic key isbased of'the latter taxa has been that eachgroup or on a study restrictedto eight recent European lineagehas its closestrelatives outside the species,whereas the problem involves a group Australianregion (e.g. Tate 194I , SimpsonI 961 , of 85 recent speciesof worldwide distribution Keast 1972"Kitchener & Capuri 1985),r'",ith (Koopman1993) and requires much Simpson(1961) describingthe Ausrralianbat so a broader 'moderately study. faunaas an only attenuated'santple Ziegler (2000) acceptedboth M1olis and of the southeasternAsian bat fauna. Leuconoeas valid genera,but nevertheless However,Volleth & Tidemann(1991) suggestedthat the use of Leuconcteas a genus suggested,on thebasis of karyologicaldata, that or a subgenusis more a matterof taste.thereby the Australianendemic vespertilionine taxa (together departingsharply from the positionsof various with the New Guinean Phurotis authorshistorically concernedby this prcblem Thomas,1914) form a monophyleticgroup. thc (seeMenu 1987. p. 84-85). ancestorof which possiblyreached Australia in After identifying the threeHerrlingen 8 & 9 the Oligocene.They also speculatedthat early speciesas new speciesof M1,o1i5,Zeigler then membersof the other vespertilioninegenera, reassessedvarious classicOligocene and includingLeuconoe, did not coloniseAustralia Miocene Europeanspecies, attributing them to until thePliocene. either Mttotis or Leuconoe, regardlessof The Riversleightooth testifies to thc presence evolutionary,stratigraphic" biochronological, and of vespertilionidsin Australiaat least20 Ma ago. palaeobiogeographicaspects of their dentitions Further, if our identificationof the taxon as a - a more or lessplesiomorphic (Leuconoe) or speciesof Leuconoais correct,it suggeststhat sharplyapomorphic (Myot is) tooth morphology. at least one cosmopolitanlineage of seenespecially in thebasic pattern ofthe upper vespertilionines ', hadcolonised northem Australia molarsM' asargued by Menu ( 1987). by the earlyMiocene. Certainly representatives ApplyingZiegler's (2000, table 5) key would of other vespertilionoidgroups with worldwide leadus to referthe Riversleighvespertihonine to distributions,such as molossids (e.g. species of Mv-otis,and the Herrlingen 8 &9 Myotisminorto Mormopterus;Hand et al. 1997),had reached Leuconoe.However, according to our criteria,at Australiaby thistime. leastM. minor belongs Io Leuc,onoe.whereas The oldest known vespertilionidbar, a the other two speciesmay require further Leuconoe-ltkespecies (Sig6 1995), appears in the assessmeni. fossii recordin the late Eoceneof southwestern 327 lLCHERINGA EARLYMIOCLNEVESPERTILIONIDFROMRI\ERSLEIGH

By the Pliocene the ProPortion of France(Muratet et ut.1985),whereasa nyctaloid taxa preseled in Riversleigh's (i.e. non-leuconoformeand non-M-yoli'tas so- vespertilionid kar.sticdeposits had changed'Of the 13 called) vespertilionid occurs in the early microchiropterantaxa identilled from thePliocene Oligoceneof Belgium(Quinet 1965;Menu 1987' Rackham'sRoost cave deposit,four are p. *;. Witti ttreexception of derivedfeatures such vespertilionids(see above). It is possible that as myotodonty,the dentalmorphology of early some of thesemay have been carried into the leuconoformesis interpretedto be generally as prey by the carnivorousghost bat plesiomorphicfor vespertilionines(Menu 1987) cave Macroclermagigas(Hand 1995),but at leastfour and close to the stock from which all vespertilionid speciesare commonly fbund vespertilioninegroups arose' As suchthe sudden roosting in Riversleighcaves today (Leuconoe appearancein thefossil record ofthis lateEocene moI ucc ctrum. Ch al inolob us nigrogri se u s (Gould' Leuconoe-\kebat was interpreted(Menu 1987) 1856),Scolorepens greyii (Gray, 1843) and asrepresenting an irnmigrationevent liom outside Vespadelus.finla1'soni Kitchener' Jones & Caputt' Europe,possibly from Asia.Species of Leuconoe le87). reachedNorth America possiblyas early as the The two extant Australian speclesoI Oligocene(Menu 1987,fig' 2) andindependent!1 Lt'utonttc and macroptts)are nom ttreOligomyotis occurrence (Galbreath 1962) lmolut'carum distributedin a narrow band around the coast aspreviously discussed by Sige& Menu (1995' from northwesternWestern Australia to nearthe p.if :). The rarity of vespertilionidsin the Victoria-SouthAustralia border' Ihe Australian Australianmiddle Tertiaryrecord possibly reflects speciesuse a range of roosts including caves' the relative number of vespertilionidtaxa tunnels,bridges. buildings and densefoliage' occurringin Australiaduring thatperiod (onein never far from bodies of freshwater(Richards cc 40 speciesfor the late Oligoceneto middle 1995).In the Riversleigharea' L' nroluccarum Miocene). However, it is more likely to be an roostsin deepcrevices close to the entranceof artefactof the Australianfbssll bat record:good limestonecaves near water' lt foragesfor insects fbr cave-drvellers,poorer for others' The vast andsmall fish over water,typically by raking the majority of Australia'sTertiary bats have been surfacewith its largehind feet' recoveredfiom karstic deposits,particularly The living AustralianLeuconoe specles are thoseof Riversleighwhich includeboth caveand generally regardedto representa southerly lacustrinedeposits (Archer er c/' 1994) Few non- of the L. adt'ersusgroup wnose karsticsites have produced identifiable Tertiary extension distributionand diversity centre on Sumatraand bat taxa in Australia,notable exceptions being Bomeo in theAsian Archipelago(Findley 1972' theearly Eocene Tingamarra deposit (preservtng Kitcheneret at.1995). The Miocene taxon tiom an archaeonycteridid;Hand ct al' 1994)and the Riversleighseems to belongnot to this modern' PlioceneChinchilla deposit (containing a Asian-basedgroup but perhapsto an earlier molossid;Hand ct a/. 1999). whose living descendantsmay include At Riversleigh.the Oligo-Miocenebat- lineage more widespreadspecies such as Leuconoe bearingdeposits are dominated by rhinolophoids andL. colifornicus' (hipposideridsand megadermatids),in both my'stacinus number of taxa and specimens'Most of these were probably cave-dwellers,like their living Acknowledgments relatives.Rarer molossids, mystacinids and the This collaborativeresearch by French and single vespertilionidbat recoveredfrom the ,Australianpalaeontologists on Riversleighfossil depositsmay haveroosted in eitheror both caves material has been supportedby the Australian ani fbrest.In Australiatoday most vespertilionids ResearchCouncil; the Department of the roostin hollow limbsof trees,forest rubble' loose Environment. Sport and Territories; National bark,crevices among rocks and, less often, caves' 328 MENUETAL. ALCHERTIVGA

EstateProgram Grants (Queensland); Queensland Cnr,rsrn,P., l997.Oligo-Miocenesedirnents of Rivenlcigh: National Parks and Wildlife Service: the the potential significanceof topography.Memoir.s oJ the Museum 41,303-314. Australian GeographicSociety; the Linnean Queensland Cuvrun,G., 1825.Discours surles ret,olutions de la sutfacc Societyof New SouthWales; ICI; theQueensland du globe et sur les changemen.sqtt'elle.s ont prodtrits Museum; the Australian Museum; and the dans le rigne . 3dne Edition, G. Dufbur & E. University of New South Wales. We thank M. d'Ocagne,Paris, 400 p. Doesox, G.E., 1875.On thc genusScotophilus, Archer, H. Godthelp and A. Gillespie for with a dcscriptionofa new genusand spcciesallied thereto. facilitating our study of the Riversleighbat Proceetlings of the Zoological Soc.iet.v(l,ondon1 material,L. Meslin (Montpellier) fbr the drawn 187-r,312. illurstration,as well asR. Smithand J. Cilis (Institut Dc;sscrN,G.E., 1878. Catalogue of the Chiroptera in the Collection of the llritish Museum (Ncttural ITi.storv) Royal desSciences Naturelles, Bruxelles) and J.- British Mussurn,London, 567 p. P. Agurlar for kindly processingand improving Durca:.r,A., Bnrln, C.B. & Moirlc;ctlrpny,N., eds, I99t). 'l'he the SEM illustration.We thank GerhardStorch aclion plan./br Au,stalian bats. Natural Herilage and Chris Beard for their constructivecriticism Tru:t. Canhcrra.I(r.1 n. Frr:rnr-ey,J.S., 1972. Pheneticrelationships of an earlierdraft of thispaper. arnongbats of the genus Mvotis. SystemaiicZoolctgv 2l, 3l-52. G,tst.xu.,L.K. & C,rnl:rre,V.J., 1987.On the llrst flnd ol Ref'erences lbssilbats (Chiroptera) in thc Paleogcncol'the USSR. Bulletin oJ the Academ.vScience ol the Georgiutt ArrEr, C.M., 1933.Two ncw batstiotn Aushalia.,lournal SctvietState Republic 126, 1, 191-200. oJ Ilamnalogy 14, 149-150. G,lr.Rae,urr,E.C., 1962. 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llisror.v,serit's 8, 11, 381-383. GustavFischer, xxiv + 898 p. THowr,q.s,O., 1915. Scientiflcrcsults fiom the Marnrnal Woonaunxe, M.O., TEopono, R.H., AncHEn, M., TttnNoulr, Suruey no. l0(A). The Indian bats assignedto genus W.D., Pr-.qNE,M.D. & Luxoprrus,8.L.,1985. trIvoti.s..Journal oJ' the: Bomhay Natural Historv Biochronology of the continental matnmal record Sor:iety23. 607-612. of Australia and New Cuinea. fpecial Publication of Tnouc;Hrox,E. le G., 1944.Furred Animals of Austroliu. the South Auslralian I)eporlment of Mines and 2nd edition.Angus & Roberlson,Sydncy. 374 p. I:ner91,5, 341-363. Tr;Rr'rlrL.r-,W.D. & Lurotlrus, E.L.Jr, 1970.The llarnilton Woooor.rnxr,M.O., M,qcF,qootx, B.J., Casr, J.A., Srttxcrn, fauna. A late Pliocenernatnmalian fauna fl'orn the M.S.,Prr.oce, N.S., Powen, J.D., WooosLrnxe, J.M. & Grangc Burn, Victoria, Australia. Fieldiunu Sparrcen,K.B., 1993.Land mammalbiostratigraphy ((ieolog.v)19, l-163. and magnctostratiglaphyof tlie Etadunna Fonration Vor.L-r,ln.M. & Trorverr. C.R., 1991.Thc origino1'the (latc Oligocene)of South Australia..lournal o.f Australian Vcspertilionincbats, as indicatedby I"ertebratePaleontolog.v /J, 483-515. chrornosorrral studies. .Z;,llrcf i/i ./irr Sci u ge t i er kund e Zrccr.rn,R., 2000. Thc bats (Chiroptera.Marnrnalia) liorn .56.321-330. the Late Oligocene FissureFillings Hcnlingen 8 and Wrrrr:n,1927-1928. Dit Saugelicre.2, Svstematisther Herrlingen 9 near Ulrn (Baden-Wiilttetnberg). 7'.il (1928.uith tolloborationof OthcntoAbel). lena, Senckenbergianalethoea 80, 641-683.