New Miocene and Pliocene Megadermatids (Mammalia, Micro Chiroptera) from Australia, with Comments on Broader Aspects of Megadermatid Evolution

NEW MIOCENE AND PLIOCENE MEGADERMATIDS (MAMMALIA, MICRO CHIROPTERA) FROM AUSTRALIA, WITH COMMENTS ON BROADER ASPECTS OF MEGADERMATID EVOLUTION

Suzar.nuJ.IIAND

HAND S.J. 1996. New Miocene and Pliocene megadermatids (Mammalia, Microchiroptera) from Australia, with comments on broader aspects of megadermatid evolution. [Nouveaux M6gadermatid6s miocdnes et pliocbnes (Mammgliq, Microchiroptera) d'Australie;commentaires sur les aspects g6n6raux de l'6volution des M6gadermatid6sl. GEOBIOS' 29,3 :365-377. Villeurbanne le 30.06.1996.

Manuscrit d6pos6 le 17.05.1993 ; accept6 d6finitivement le 10.01.1995.

ABSTRACT - Macroderma malugara nov. sp., a new false vampire from the middle Miocene Gotham City Site of Riversleigh Station in northwestern Queensland, is described and three others discussed. Seven megadermatids are now known from the Australian fossil record. Morphological changes in the Australian Macroderma lineage are traced from the Oligo-Miocene to the present. In some Riversleigh deposits, there is evidence for sy'rnpatry of very differently-sized megadermatids. Riversleigh megadermatids provide an opportunity to trace an apparent trend to shorten the face in the Mairoclerma lineage flom the Oligo-Miocene to the present and to fufther document a tendency to gigantism in inde- pendent megadermatid lineages.

KEY.WORDS : CHIROPTtrRA. NEW SPECIES, EVOLUTION, MIOCENE, PLIOCENE, AUSTRALIA

RESUME - Macroderma malugara nov. sp.) un nouveau faux vampire du Miocdne moyen de Gotham City, Riversleigh Station, Queensland NW, est d6crit, et trois autres formes sont discut6es. Sept m6gadermatid6s fossiles sont maintenant connus d'Australie. Les modifications morphologiques dans la lign6e australienne de Macroderma sont retrac6es depuis l'Oligo-Miocbne jusqu'ir lActuel. Dans certains d6pdts de Riversleigh, la sSnnpatrie de m6gadermatid6s de diff6rentes tailles est 6vidente. Les m6gadermatid6s de Riversleigh montrent une apparente tendance au raccourcissement du museau dans la iign6e des Macroderma de l'Oligo-Miocdne a l'Actuel et, de plus une tendance au gigantisme dans des lign6es de 1\{6gadermatid6s ind6pendantes.

MOTS-CLES : CHIROPTERES, NOUVELLE ESPECtr, EVOLUTION, MIOCENE, PLIOCENE, AUSTRALIE.

INTRODUCTION lophoid families, the Hipposideridae (minimum 12 species)and Megadermatidae (minimum 7 species). More than 33 new microchiropterans have been Riversleigh fossil hipposiderids (Sig6 e/ al. 1982 ; identified f?om Oligo-Miocene and Pliocene fresh- lJandet al. 1989 a,b, 1990 ; Hand 1993) are the sub- water limestone deposits of northwestern Queens- ject of an extensive study in progress. Two land's Riversleigh Station (Archer et al.1989,1991). Riversleigh megadermatids (Macroderma godthelpi Many of these species, including Hipposideros and the Dwornamor Variant) have been previously (Brachipposideros) nooraleebus Stcp, HaNl & described (Hand 1985). ARCHER, 1982, are represented by partial skulls, complete dentaries and most postcranial elements. In this paper, a new Tertiary megadermatid is des- Others are currently known from less complete but cribed and three others discussed. Morphological nonetheless distinctive remains G.g. Petrctmops changes in the Australian Macrodermo lineage are creaseri IIAND, 1990) that significantly contribute to traced from the Oligo-Miocene to the present and the picture of high bat species diversity emerging comments are made on broader aspects of meqa- from study of the Riversleigh fossil mammal faunas dermatid evolution. (Archer 1989,1991). et al. Dental terminology follows Hand (1985). The prefix Riversleigh Tertiary deposits are particularly rich QM F_ indicates the fossil collections of the in the remains of members of the Old World rhino- Queensland Museum, Brisbane. JOt)

SYSTEMATICS rently understood to outcrop over an area of approximately 16 square metres. Vertebrate fossils are concentrated in a 15 centi- Order CHIROPTERABlumenbach, 1779 metre-thick band within the limestone. The fossils are mostly Suborder MICROCHIROPTERA small, fragmentary bones and teeth but include occasional lar- Dobson, 1875 ger, often worn bone fragments. Superfamily RHINOLOPHOTDEAWeber, 1928 On the basis of its stratigraphic position and marnmal fauna Family MEGADERMATIoAPAllen, 1864 (Hand 1990), the Gotham City deposit is interpreted to be middle Miocene in age and slightly younger than the South Genus Macroderma Mu,LpR, 1907 Australian Kutjamarpu Local Fauna (Woodburne ef al. 1985) but older than the Riversleigh Macroderma ntalugara nov. sp. Jaw Junction and Northern Territory late Miocene Alcoota Local Faunas (Woodburne et al. 1g85 : Holotype (Pl 48) - Queensland Museum F23611, right maxilla- Archer ef rul. 1989, 1991). ry fragment containing F, M'and alveolus for P'. Associated fauna and taphonomy - The nature of the Etymology - The species name comes from the local Aboriginal C.otham City deposit and fauna has been discussed previously (Hand word (Wanyi language) malugara meaning a good killer. It 1990). Preserved in the deposit are the remains of small alludes to the apparent carnivorous habits of this large mega- teleost fish, tiny lungfish, frogs, lizards, small freshwater croco- dermatid; the fossil fauna with whtctr Macroderma malugara rs diles and turtles, a passerine bird, dasyurids, peramelids, petau- preserved appears to rnainly represent the accumulated remains rids, pseudocheirids, acrobatids, a phalangerid, a diminutive of its prey. phascolarctid, a potoroid, four hipposiderids and the molossid Petramops creaseri (HAND, 1990). Some of the mammal species Paratypes (Pl. 48) - QM F23612, right C' ; QM F23613, right P.; are shared with contemporaneous Riversleigh deposits, but QM F23614, left M, ; and QM F23615, right M,. others are unique to Gotham City and,judging by the size and fragmentation of the material, probably represent the remains of Referred material - F23616, a large periotic from the QM prey brought from adjacent habitats by Macrodcrma malugara. Henk's Hollow Local Fauna is tentatively referred to this taxon. Larger, worn bone fragments representing large turtles, a balun- By comparison with living and fossil species,this periotic repre- gamayine potoroid, a zygomaturine and a possible Wahaleo spe- sents a much larger megadermatid than the Henk's Hollow cies may have been washed into the limestone roost durins megadermatid described below as megadermatid indet. or the changes in the level ofthe water table. Boids and some ofthe hip- Oligo-Miocene Macroderma godtltelpi HAND, 19B5.It is compa- posiderids probably co-habited the megadermatid roost as they rable in size to, but morphologically different from, that of the commonly do today. living Macroderma gigas (DOBSON, 1880). Diagnosis - This fossil species differs from Recent Type locality, age and lithology - The type locality, Gotham populations of Australia's City Site (Hand 1990), occurs within the Tertiary sequence of only living megaderma- limestones outcropping on Riversleigh Station, northwestern tid, Macroderma gigas, in its retention of P2, its Queensland (Archer et al. 1989, 1991). It is situated near the nor- double infraorbital foramina, its less well-developed theastern edge of Ray's Amphitheatre (Archer et al. I98g, IggI) heels on P'and M' and parastyle on P4,its M1 with at approximately 19.00.88 N, 138.39.07 E (Global Positioning strongly-developed anterior cingulum and heel with Satellite Device, co-ordinates for Ringtail Site located aproxlma- lingual cingular cusp, its less posteriorly-recurved tely 30 metres southwest of Gotham).It is stratigraphically near anterior cutting crests on Pt, its more reduced and the top of the System C sequence as defrned by Archer et aL. buccally-situated (1989, 1991). This is interpreted to be stratigraphically higher Mz metaconid, its broader and more robust Mz cingulum, than Site D (System A containing the Riversleigh Local Fauna ; its very poorly-developed Archer ef al. 1997), Microsite (containing the Nooraleeba Local and less posteriorly-situated M, h;poconulid, its Fauna ; Sig6, Hand & Archer 1982 ; Hand 1993) and Gag Site less anteroposteriorly-compressed Mz talonid, its (near the basc of System C, containing the Dwornamor Local less lingually-open Mz talonid with better-developed Fauna ; Archer & Flannery 1984 ; Hand 1985 ; Archer et cl. pre-entocristid and its tall but much less postero- 1991), close to the level of T\vo (containing Trees Site the Two Iingually-extended Cr that lacks an anterolinzual Trees Local Fauna ; Flannery & Archer 1987) and below the cingular cusp. It differs ftom the Pliocene Welling- Henk's Hollow Site (Flannery & Archer 1987 ; Archer et al. \98g, 1991). ton Caves species, Macroderma koppa (Hand et el. The sediment is a horizontally-bedded, frne-grained argillaceous 1988), in its retention of P,, its smaller size, its less limestone representing an as yet unnamed formation. It is cur- well-developed heels on Pa and M1 and parastyle on

PLATE 48

Macroderma malugara nov. sp. from Gotham City Site, Riversleigh Station, northwestern Queensland. Holotype, QM F23611, a right maxillary fragment with P4,M' and alveolus for P'. A, buccal view ; B-B', stereopairs, occlusai-bltiq,r" view ; D-D', stereopairs, occusal view. Paratypes : QM F23672-23675. QM F23612, a rigltt C' : C-Cl, stereopairs, occiusal view E, lingual view. ; QM F23613, a right Pn : F, buccal view ; G, lingual view ; H-H', stereopairs, occlusal view. eM F23614, a left Mz : I-I', stereopairs, occlusal view ; J, buccal view. QM F23615, a right M, : K, buccai view ; L, Iingual view; M-Mf, stereopairs, occlusal view. Scale = 1mm. Fragment de maxillaire droit auec , Pt, M, et I'alu1ole de pt. A, uue buccale ; B-B', uues occlusales obliques en stAr1o.D-D', uues stdrdo de la surface occlusale. Paratypes : C, cJroite. C-C,, uues stdrdo de la surface occlusale. E, uue linguale. P' droite, uue buccale. G, uue linguale. H-H', uun iterdo d.e la surface occlusale Mz droite, K, uue buccdle. L, uue linguale. M-M', uue st1rAo de la surface occlusale. Geobios PI.48 no 29, fasc. 3 S.J. Hand 368

P', its M'heel with distinct lingual cingular cusp, its tooth that occurred on the lingual side ofthe tooth less posteriorly-recurved anterior cutting crest on row immediately anterolingual to the anterior cin- Pa,its much less well-developed P'posterior cmgu- gular cusp of P'. lum and lingual cingular cusp, its P4 with postero- The P' is represented by a complete tooth in lingual groove or depression only weakly developed, QM F23611. This tooth is remarkably similar in mor- its broader Mz anterior cingulum, its very poorly phology to its counterpart in Macroderma gigas but developed and less posteriorly-situated Mz hypoco- in contrast has an almost complete anterobuccal nulid, its better-developed pre-entocristid, its more and lingual cingulum. (The tooth is fractured such reduced and buccally-situated M2 metaconid and its that the posterobuccal cingulum is not preserved.) much less posterolingually-extended Cr that lacks The posterior cingulum merges smoothly with the an anterolingual cingular cusp. It differs from the extended heel rather than meeting its buccal edge Riversleigh middle Miocene species, Macroderma at an angle. This apparently reflects the more lin- godthelpi (HaNo, 1985), in its much larger size, its gual (rather than posterior) direction of heel deve- taller crowned teeth, its smaller and more lingual- lopment in M. malugara. The heel, although large, ly-situated P', its better-developed heel on Pnand is less well-developed than in M. gigas. A lingual Iingual cingular cusp on M', its more robust and cingular cusp is only very faintly developed and broad anterior cingulum on Mr, its more anteropos- there is little parastylar development. The anterior teriorly-compressed talonid and its complete cingu- cutting crest of the paracone is less posteriorly- lum on Cr. recurved. Description - The only maxillary fragment is the The M'is known from the nearly-complete RM'of holotype F23611 which preserves part of the QM the holotlpe F23611. It is very similar in mor- cheek region. TWo infraorbital foramina are exhibi- QM phology to that tooth in M. gigas (HaNn, 1985, frg. ted. The anterior foramen is large and opens onto 16) but differs in its shorter, broader and generally the cheek with a fossa developed above the anterior less well-developcd heel and its very pronounced end of Pn. A second, much smaller and more poste- lingual cusp. rior fossa is developed in a position above the posterior root of Pa. This appears to lead to a second The lower dentition is known from four teeth : a foramen that penetrates the maxilla, possibly for a lower canine, a right Pa and two specimens of Mr in lateral branch of the trigeminal or facial nerve. The dentary fragments. The lower canine is represented anterior fossa is directed anteriorly ; the smaller by an isolated, right C', QM F23612. It is similar to fossa appears to lead more anteroventrally. The Ct in M. gzgos but the posteriorly-extended heel is upper dentition is known from the holotype, QM less well-developed so that its basal crown area is F23611. The P' is represented by a single alveolus. smaller than that in M. gigas. The basal cingulum It appears to have been a very small, single-rooted is complete and well-defined. Unlike M. gigas,there

Holoboe Para-tvp€s

QM F23611 QMF23612 QMF23613 QMF2:]614 QMF23615

P4 buccal length 3.35 P4 lingual length 3.20 Pa width 3.15 M1 wjdth 4.I4

Cl length 2.99 C1 width 2.77 Cl heisht (He) /. -td C1 height (Hi) o.au TABLE 1 - Measurements (mm) of the holotype (QM F23611) and para- P4 length t1t types (QM F23612-236I5) of Macro- P4 anterior cingulum to protoconid length t.23 derma malugaro nov sp. from the Miocene Gotham City Site. River- P4 width 1..b5 sleigh Station, northwestern Queen- sland. Mensurations (mm) de I'holo- M2 length (in situ) 2 AA 3.60 paratypes type et des de Macroder- M2 trigonid length 1.99 2.t7 ma malugara nou. sp.du Miocine de I.46 Gotham City, Riuersleigh Station, M2 talonid length r.42 Queensland lflV. M2 trigonid width 2.27 2.6r M2 talonid width 2.2r 2.45 369 is no developmentof an anterolingual cingular cusp Comments - Specimensof Macroderma gigas so adjacent to where the outer lower incisor would far recovered from the Rackham's Roost deposit abut. (e.g.QM F23560-23562)are relatively small (Table 2), being much closerin size to small representa- The Pois represented by a complete,isolated tooth, tives from RecentM. gigas populations such as Pine QM F23613. This tooth is laterally compressedas Creek, Camooweal, the Kimberley Range, the in M. gigas, but is relatively smaller. Lower molars Pilbara and Groote Eylandt (Hand & York 1990) are known from a left and right M, (QM F23614 and than to large individuals from, for example,existing QM F23615 respectively).The morpholory of Mz in Cape York and Rockhampton populations. They M. malugara differs from that of M. gigos in its appear to be much smaller than large individuals of more reduced and buccally-situated metaconid and M. gigas collectedas subfossilsfrom the McDonnell its less anteroposteriorly-compressedtalonid. The and Flinders Ranges and fossils from Pleistocene hypoconulid is less well-developedand less poste- deposits in southwestern Western Australia (Hand rioily-situated. The talonid is less open lingually, & York 1990). its pre-entocristid extending further anteriorly and being better-developed. The cingulum is broader Small size in the Rackham's Roost Macrodermcr and more robust, particularly anteriorly. Fragments population is perhaps most evident in skull and of dentaries associated with these teeth confirm dentary characterssuch as palate length and den- that they are Mz'sbut provide little additional infor- mation. Measurementsof the holotype and paratypes of Macroderma malugara nov.sp. are given in Table 1. Macrodermagigas (DOBSON,1880) Fig I

Referred material - QM F23560, a partial skull with P, M', M' and M'' and single infraorbital foramen ; QM F23561, a dentary containing Pn,Mrr and alveoli for Lz and Pr ; QM F23562, maxillary fragment with P, M' and M'and two infraorbital foramina; QM F23563, maxillary fragment with F-Mi ;QM F23578, distal end of humerus ; QM F23579, proximal end of ulnar ; QM F23580, partially erupted M, ;QM F23590, periotic.

Locality, lithology and age - Rackham's Roost Site (Godthelp 1988 ;Archeret aI. 1989,1991) occurs at19.02 09 N, 138.41 60 E (Global Positioning Satellite Device) on Riversleigh Station, northwestern Queensland. It appears to represent the indurated floor of a long, narrow cave developed in Cambrian Thorntonia Limestone. At one end, the cave evidently opened onto a vertical cliff flanking what is now the Gregory River. More than 50 metres away in the opposite direction, another larger, lower ope- ning appears to have been developed. The sediment is a breccia oftiny, mostly fragmented bones and teeth set in a fine-grained, pink-coloured (presumably iron-stained) lirnestone The deposit covers an area of approximately 200 square metres, with a maximum depth of 0 5 to 1.0 metre. On the basis of its mammal fauna, the Rackham's Roost deposit is interpreted to be early to middle Pliocene in age. It contains abundant plesiomorphic murids (at least 12 species) and a macropodid sirnilar to Proternnodon snewini BARTHOLOMAI, 1978 frorn the early Pliocene Bluff Downs Local Fauna of northeastern Queensland (Archer & Wade 1976)

Associated fauna and taphonomy - This species occurs in the Rackham's Roost deposit with fragmentary remains of crusta- ceans, frogs, lizards, small crocodiles, birds, dasyurids, peramelids, a pseudocheirid, an extinct macropodid and potoroid, at FIGIBE 1 - Specimens of Macrodcrnta gigas recovered from least 12 murids, an emballonurid, four hipposiderids, four ves- Rackham's Roost Site, Riversleigh Station A, QM F23560, a par- pertilionids and a second,tiny megadermatid (Archer ef ol. 1991; tial skull with F, M', M'and M'and single infraorbital foramen: Hand 1987, 1995) The very finely broken remains of the small lateral view. QM F23562, maxillary fragment with F, M' and M' vertebrates and depressed fractures and impressions in the and two infraorbital foramina : B, lateral view ; C-C', stereopairs, bones characteristic of Recent Macrodcrma gigas canines sug- occlusal view. Scale = 5 mm. A, crdnc incompLet auec P, M', Met gest that the deposit represents the remains of prey accumula- M'' et un forarnen infraorbitaire unique : uttc Latirale QM, fras- infraorhitaires. ted by this species Boids, the emballonurid, hipposiderids and ment de rnaxillaire auec F, MI et M'ct 2lbramens uues stdrdo de La surlacc occlusale. vespertilionids probably co-habited the megadermatid roost. B, uue latdrale. C-C', 370 tary length and depth (compare Table 2 and Hand oMF23560 QMF23562 & York 1990, table 2), but too few individuals have been collected to determine whether they are signi- Palatal length 8.80 p4-rr.t3 10.75 ficantly smaller in these features than Quaternary specimens. Preliminary study suggests that mea- It1-lt3 8.01 surable characters that are most variable in F buccal length 3.00 3.05 M1 buccallength 3.36 e<9 Quaternary specimens of M. gigas (e.g. lower tooth widths and lengths) are less variable in the M2 buccallength 3.27 Rackham's Roost population. At least one character M3 buccallength 1.81 a^n that appears to be fixed in Quaternary populations P4 lingual length 3.21 of M. gigas, however, seems to be variable in the M1 lingual length J.DY 3.85 Rackham's Roost population. While in most M2 lingual len6h a aa a7a Rackham's Roost specimens (n=5), as in # width 2.51 2.60 M1 Quaternary M. gigas, a single infraorbital foramen width 3.94 on the facial side of the maxilla is exhibited (Fig. M2 width 2

FIGURE 2 - Macrod.ermo sp. from Henk's Hollow Site, Riversleigh Station, QM F23617, a right M' : A'A', stereopairs, occlusal view Megadermatid indet from i{enk's Hollow, QM F23618, b left M' : B-R_',sterelpairs, occlusal view ; C, buccal view ;D, lingual view. Scale = 1"rnm Mt droite : A-A', uues stdrdo de la surface occlusale. Mt gauche :B-B., uues std.rdode la surface occlusale C, uue buccale D, uue linpuale.

Macrodermo. Features of the Mr shared with other malugara nov. sp. and Henk's Hollow Macroderma megadermatids include : nyctalodonty (as defrned sp., and the Pliocene Wellington Caves M. hoppa. by Menu & Sig6 1971) ; Iaterally-compressed trigo- The genus Megaderma GEoFFRoY, 1810 is repre- nid and anteroposteriorly-compressed talonid ; sented in Australia for the first time by a tiny new reduced paraconid, the paracristid being much lon- species from the Pliocene Rackham's Roost deposit ger than the metacristid ; and twinned entoconid (Hand 1995). Another new megadermatid whose and hypoconulid. Features that appear to exclude it dental morphology appears to place it outside both from the Australian genus Macroderma include its the Macroderma an;.dMegaderma lineages, is repre- gracile cingulum and the unique development of its sented by an isolated M' from the later middle pre-entocristid which lingually walls the talonid Miocene Henk's Hollow Local Fauna. in a manner not unlike that in the living basin A phylogenetic hypothesis based on shared-derived African megadermatid Lauia (see Hand 1985, flons character states involving all adequately-represen- fig. 18). Features that distinguish the Henk's ted fossil and living megadermatids was given by Hollow M' from its counterpart in L. include frons Hand (1985 and see Hand el ol. 1988). In that its markedly compressed trigonid and talonid. ; hypothesis, species of Macroderma were interpreted to be the most derived megadermatids and Lauia frons the most plesiomorphous of living DISCUSSION megadermatids. The French fossil Necromantis adichaster was considered to be the most plesiomor- PHYI,O GENETIC REI"NIIONSHIPS phous member of the Megadermatidae. Macro- derma species formed a clade vrit}l. I'yroderma spe- Seven megadermatids are now known from the cies, Megaderma species formed its sister-group, Riversleigh Tertiary deposits (Table 3). The oldest and Cardioderma cor was the sister-group of a are the Oligo-Miocene Macroderma godthelpi from M egaderma -I4troderma-M acrodermo clade. the Nooraleeba and Dwornamor Local Faunas and the DwornamorVariant from the Dwornamor Local Sig6 (1976) also regarded the dentition of Lauia Fauna (Hand 1985 ; Archer et al. 1989, 1991). frons to be "peu 6volu6e" with respect to other living Representatives of the Macroderma lineage now megadermatids and excluded it from a group in appear to also include (apart from the living Ghost which he placed Macroderma gigas, Megaderma Bat M. gigas) the later middle Miocene Gotham M. spasma, M. lyra and Cardiodermo cor megaderma- Dr70

tids he described as having "denture morphologi- quement 6volu6e". Oligo-Miocene Pliocene Quar Griffiths et al. (1992), however, reached contradicto- t ry conclusions on the basis ofa cladistic analysis of r\ hyoid morphology in living megadermatids. In that analysis, Macroderma gigas is interpreted to be the U) .= I (h most plesiomorphous and Lctuia frorus the most apo- ,a o morphous of living megadermatids. The phylogene- >i E tic positions of species of Cardioderma and o U 6) J E E Megaderma (including Lyroderma) are, as pointed ./) 3 ts J U) J out by Griffiths et al., not very different in the a 6 0 models of Hand and Griffiths et al. In both, species z ra of Megadermo are considered relatively derived and Cardioderma cor intern'rediate in the cladog.ram. Macroderma gigas x X

If Griffrths et al.'s internretation is correct. then the Macrodenna lcoppa x polarity of dental charaiters used by, among others, Sig6 (1976) and Hand (1985) is in doubt. However, Megaderma richardsi x for the dental characters to concord with Griffrth e/ ol.'s cladogram, an unacceptable amount of homo- Megadermatid indet. X plasy would be required. For example, if transverse compression of the M' trigonid is considered to be Macroderma sp. X the result of homoplasy rather than a sJmapomor- Macroderma malugara X phy for most megadermatids then it must be hypo- thesised that its absence in Lauia frons renresents Dwornamor Variant a reversal event, or that it has been independently acquired in megadermatids several times. Thrs Macroderma godthelpi would be true also for a number of other characters that are diagnostic for most megadermatids except Lauia frons (e.g. reduction in the pre-entocristid to TABLE 3 - Distribution of megadermatids in Australian fossil open the talonid basin in M' s) and are not found in deposits. Abbreviations : Riv, Riversieigh Station, Queensland ; Wellington C., Wellington Caves, New South Wales ; Quat, the potential sister-groups nycterids, rhinolophids, Pleistocene and Recent. Sites are shown (left to right) in probable emballonurids or rhinopomatids. This is by no stratigraphic sequence (i.e decreasing age) following Archer et al. means impossible but it is less parsimonious. (199I). Rdpartition des migadermatidds dans les sddiments f'os- siles d'Australie. Les gisements sont rangds de la gauclrc uers la, On the basis of fossil material and characters avai- droite dans I'ordre stratigraphique probable. lable for study, the majority of Australian Tertiary tures exhibited to varying degrees in other mega- (Fig. megadermatids appear referable to a clade 3) dermatids (e.g.very enlarged heels on P'-M', a very which also includes Macroderma gigas, and are tall, prominent paracone on Cl, and robust cingula referred here to that genus. A combination of fea- on the lower molars). It is possible that such cha- tures appears to define this clade : a massively-roo- racters may be the result of allometry rather than ted, posteriorly-recurved C' with tall, slender, domi- phylogeny, but they are not all present to the same posterior nant main cusp and large accessory cusp degree in large non-Australian taxa such as the well-separated from the paracone but occurring Miocene European and North Aliican Megaderma within the cingular rim and a well-developed ante- gaillardi (Tnounssenr, 1B9B) and M. brailloni rolingual cingular cusp ; enlarged heels on P'-M' ; SrcE, 1968. only moderately elongate postmetacristae on M'" ; lower molars with laterally-compressed trigonid, Within the Australian radiation, Macroderma anteroposteriorly-compressed talonid, reduced malugara shares a number of apparent sJ,ryrapomor- paraconid with the paracristid longer than the phies with the Pliocene species M. lzoppa and metacristid (although both crests are relatively Pliocene to Recent M. gigas that are not shared transversely oriented), twinned entoconid and with the older Dwornamor M. godthelpi (Fig. 3). hypoconulid, reduced pre-entocristid such that the These include large size, taller-crowned teeth, well- talonid is open lingually, reduced metaconid contri- developed P' heel and more anteroposteriorly-com- bution to the cristid obliqua, and robust, continuous pressed talonid. There is nothing known about M. anterior, buccal and posterior cingula. godthelpi that would exclude it from the ancestry of M. malugara. It is noteworthy that species of Macroderma tend to be large in absolute size and apparent sJmapomor- The evolutionary relationships of the poorly-known phies for the clade include an exaggeration of fea- middle Miocene Macrodermo sp. and megaderma- D '.) tid indet. from Henk's Hollow Site are unclear. mediate Dwornamor M. godthelpl and the large However, rvith species of Megaderma, I'yroderma Gotham M. malguaro. Known from the Pliocene are and Macrodenna, megadermatid indet. shares a a tiny megadermatid (Hand 1987 ; 1995), the larger markedly laterally-compressed trigonid while lac- Rackham's Roost Macroderma glgczs and the very king the extreme degree of anteroposterior com- large M. hoppa ftom Wellington Caves, New South pression of the talonid exhibited by species of Wales. Lyrodernta and Macroderma. In the development of Sig6 (1976) has suggested that in megadermatids a its pre-entocristid, it more closely resembles the pri- trend to gigantism may have occur-red in a number mitive African megadermatid Lauia frons (but it is of separate lineages (e.g. the middle Tertiary possible that the derived condition of this character may have been autapomorphically lost in the Megadermatidindet. Henk's Hollow animal). Although this megadermatid appears to lie outside the Macroderma lineage, Megadermaspp. neither can it be positively referred to the lineage represented in Australia by the middle Miocene Dwornamor Variant. DwornamorVariant

MORPHOLOGICAL TRENDS IN TERTiARY Macrodermagodthelpi MEGADERMATIDS

In the Australian megadermatid radiation, three Macroderma sp. distinctly differently-sized species arc now known both the Oligo-Miocenc and the Pliocene. from ,r. Macrodcrma nnlugara Represented in the Australian Oligo-Miocene are the small Henk's Hollow Macrodermo sp., the inter-

Ilypothises plLylogdnfuu.es sur lcs relations cLentdgadermatid1.s fossiles de Riuersleigh ouec Les outres migaderntatidds d partir des caractires d,iriu6s partagis ; Ies hipposiddrldis (T{ipposideros ater) sont pris comme extra-groupe (uoir aussi Sigtl 1976 ; Hand FIGURE 3 - Hypothesis of phylogenetic relationships of based 1985; Hand. ct ol. 1988 ; Hand. 1995).Apomorphies possihles : 7, Riversleigh fossil megadermatids to other megadermatids longueur de M, pLuscou.rte ou 6gale d celLede M"; trigonide de M' on shared-derived dental character states using hipposiderids con'tprintd Latiralement, paraconide rdduit ouec paracristide (e.g.Hipposidcros ater) as an outgroup (see also Sig6 1976 ; Hand beaucoup plus ktng r1u.emitacristide ; prd-entocristide de M" 1985 : Hand et al. 79BB; Hand 1995). Potential apomorphies court (interron.pu) si bien que le hassin du talonide est ottuert lin- include : 1, M' shorter than or equal in length to M,, M' trigonid paracristid guaLement ; taktnide ccnnprimi antdro-postdrieurement ; C' auec laterally compressed, paraconid reduced with much (disrupted) utt tubercule acccssoirepctstirieu.r lLaut, gtndraLement en fbrme de longer than metacristid, Mr 3 pre-entocristid short so anteroposteriorly lame ; M" d.paracone riduit, mitctcorte massif, mdsostyle diplact that talonid basin drains lingually, talonid Lingualcncn.t et post-metacristo allon.gdc.2, C' auec un tuberculc compressed, C'wiih tall, usually bladelike posterior accessory accessoircposti.rieur. coniqu.erelotiuement court, bien sdpari. du cusp, M" with reduced paracone and massive metacone, lingual- paracone mais inclui dans la bord,u.recingulairc et ouec un tuber- ly displaced mesostyle and elongated postmetacrista ; 2, C' with cule cingulaire antiro-linguol bicn ddueloppi ; P' et M'" auec un relatively short, conical posterior accessory cusp well-separated talon aLLongd; P' haute auec tubercule principal en position huc- from the paracone but occurring within the cingular rim and P' and N{"with calc ; M" auec lorge hordure huccalc et mdsostyle nettement with well-developed anterolingual cingular cusp, ddplact lingualement ntnis post metacristae moddrdment elongated heel, tall P'with buccally situated main cusp, Mt" with mesostyle allongics ; compression antiro-postdrieure du talonide de M" broad buccal shelf and markedly lingually displaced antero- plus poussde ; parocristide plu.s long que mdtacristide mais les but only moderately elongate postmetacristae, further dcux crAfus or ien.tdesrelatiuement transuersalement, contrihution posterior compression of M" talonid, paracristid longer than riduite du mdtaconide d La crista obliqua ; cinqulum antiricu4 metacristid but both crests relatively transversely oriented, huccaLet post6rieur robustes at continlts.3, dents de grand.e taille, reduced metaconid contribution to the cristid obliqua, robust, d couronne haute ; P' plu.s petitc et situic liLgualement ; talon de continuous anterior, buccal and posterior cingula ;3, large size, hecl lar- P plus largc ; C' a,uaccingulum compLet ; compression ant6ro- tall crowned teeth, P'srnaller and lingually situated, Po cingulttm antd' ger, C' with complete cingulum, M', further anteroposterior com- post6rieure suppl|men.taire dcs taLonidesde M' " ; rieur robu.staet lorge.4, P'disparuc : talons de P' et M' encore pression of talonid, robust and broad anterior cingulum ; 4, P' plus dducloppts ; P' auec parastyle et crAfu antdrieure couponte lost, Po and M' heels further expanded, P' with parastyle and plus recourbdepostdrieurement ; Cr nettement 1tirde postdro-lin- more posteriorly-recurwed anterior cutting crest, C' markedly guaLemcnt et auec un tubercule cinguLaire antdro-lingual ; M' ' posterolingually-extended and with anterolingual cingular cusp, plus grande, aucc un hypoconuLkle situt. plus postdrieurement et M,, larger and more posteriorly-situated hypoconulid and pre- le prderLtocristide encore plu.s rdduit. 5, foramen infra-orbitaire entocristid further reduced ; 5, single infraorbital foramen uniqu.e (populations cluaternaires) ; boucLier frontal bien ddue' (Quaternary populations), frontal shield well developed, C'with Loppi ; C' auec une flexu.red.e conpression antdro-postdrieurepro- pronounced anteroposterior compressional flexure, P' shorter noncde; P' pLus courte et it cotrronne plus haute ; tolonides de and higher crowned, M', talonids further compressed anteropos- Mr r ctLcoreplu.s comprimds ant6ro-postirieurement. Les reLotions teriorly. The phylogenetic relationships of Riversleigh's plrylogdniqucs du Megaderrnatidi indet. de. Riuersleis4h, du Megadermatid indet., Dwornamor Variant and Macroderma sp. uariant de Dwornamor ef de Macroderma sp.restent LncertaLnes. remai.n unclear an^

M egaderma gaillar di and Meg aderma lug dunensi s Although carnivorous bats have been described (e.g. (DnenRnr, 1892) species groups). Ifthe phylogene- Freeman 1984) as having longer faces than their tic hypothesis based on the character analysis of insectivorous relatives, megadermatids are in fact Hand (1985) reflects actual phylogeny, then large shorter-faced than most other rhinolophoids, the size has also been acquired independently in shorter face in this case perhaps being correlated Australian megadermatids. At least some with a more powerful bite by bringing the canines Australian taxa increased in size (e.g.Macroderma closer to the fulcrum of the jaw. In megadermatids, koppa) while others such as the tiny Rackham's the mandibular condyle is relatively low and the Roost megadermatid became very small. coronoid process high, a condition typical in carni- vores whereby the temporalis (the primary prey- In the later middle Miocene Henk's Hollow and the seizing muscle) is developed at the expense of the Pliocene Rackham's Roost deoosits. there is eviden- masseter (Maynard Smith & Savage 1959). This ce for syrnpatry of very diffelently-sized megader- allows less transverse activity but greater vertical matids (i.e. greater than 30Vodifference). A compa- movement and slicing ability (Freeman 1984). The rable difference in size in contemporaneous fossil direction of pull of the temporalis in megadermatids megadermatids is seen in the middle Miocene Beni has been found to be much more vertical than in Mellal Local Fauna of Morocco with the large other rhinolophoids (Freeman 1984, fig. 7), thus Megaderma gaillardi and very small M. jaegeri presumably providing a more powerful bite force (Srco, 1976). Presumably, this difference in size of (see also Emerson & Radinsky 1980). However, as slrnpatric carnivorous megadermatids relates to Freeman (1984) points out, a more vertically-orien- competition for resources. While there is little ted temporalis also tends to be negatively correla- obvious increase in carnassialization in very large ted with the tilting of the head on the basicranial forms, an evident increase in the crushing compo- axis. The latter appears to bc controlled at least in nent of the dentition may reflect an important shift part by the functional requirements of echolocation in diet. In the Riversleigh Dwornamor Local Fauna, such that orally emitting echolocators have positi- two similarly-sized s1'rnpatric megadermatids (M. vely tilted heads with respect to the basicranial axis godthelpi and the Dwornamor Variant) are refe- while nasal emitters (e.g.megadermatids and other rable to two separate lineages. Living non- rhinolophoids) have negatively tilted heads Australian megadermatids that exist sympatrically (Freeman 1984, fig. 6). over part of their range are either significantly different in size (e.g. the Asian Megaderma spasma In the Macrodermrz lineage, the trend to shorten the and Lyroderma lyra ; av.25g vs 50g, Lekagul & Mc face and thereby apparently increase the power of Neely 1977) or represent two distinct lineages (e.g. the canines, can be traced from the Oligo-Miocene the African Lauia frons and Cardioderma cor ; M. godthelpl to the middle Miocene M. malugara, Hand 1985 ; also Griffiths et al.I992). The latter in the early Pliocene M. koppa and Pliocene and each of these species pairs consumes significantly Quaternary specimens of M. gigas. Macroderma more vertebrate prey than the other (Brosset 1962; godthelpi has a relatively elongate face (P' being Wickler & tlhrig 1969 ; Kingdon 1974 ; Vaughan large and in the tooth row), a plesiomorphic double 1976 :Advani 1981 ;Advani & Makwana 1981). infraorbital foramen, an unmodified frontal nose- shield and a discrete and well-separated posterolin- Another general trend in megadermatid lineages, gual accessorycusp on C'. The later middle Miocene evident also in the Australian Macrodermalineage, M. malugaro retains the double infraorbital fora- involves marked shortening of the face and tooth men but has reduced the P' to a rudimentary tooth row Brachycephaly of this kind is even evident in that lies lingual to the tooth row. The early Pliocene the oldest known megadermatid, the French early M. hoppa has lost the P' but retains the double Tertiary short-faced Necromantis adichaster infraorbital foramen and isolated cusD on C' and REVILIIoD, \922. Megadermatids are distinguished exhibits little modifrcation of the frontal nose- from other bats by the following combination of cra- shield. specimens of M. gigas are more nial and dental characters associatedwith a shorte- Quaternary short-faced exhibitine a lack of P'. reduction of P' ning of the face : almost complete loss of the pre- from a more sectoriaf to a higher crowned stabbing maxillaries of the upper incisors of ;loss ;reduction tooth, a C'with less discrete accessorycusp and pro- the lower incisors anteroposterior compression of ; nounced anteroposterior compressional flexure Mr such that it is shorter than or equal in length to (that is most obvibus in the lingual cingulum) and a M2 anteroposterior compression of the talonids of ; modif,rednose-shield. These animals exhibit a single Mr.: reduction of M: in the num- ;marked ;reduction infraorbital foramen, but at least some Pliocene ber of premolars to a maximum of two in the upper specimens of M. gigas retain two inftaorbital fora- and lower tooth rows (N. adichaster retains a tiny mlna. P,r);and posterior extension of the palate lingual to M' presumably for compensatory support (see Of Macrodermo species, specimens of the Pliocene below). M. hoppa and Pliocene to Recent M. gigas are best 375 preserved. At some point in the evolutionary histo- the Gotham City deposit, but missing from ry of these two species,a shortening in the anterior Rackham's Roost, are acrobatids, burramyids, part ofthe face appears to have resulted in a relati- petauroids, small phalangcrids, peroryctids and a ve decrease (from hoppa to gigas) in the size of inci- new family of tiny perameloids (all small enough to sors, canines and premolars, possibly a modification be megadermatid prey). Although some marsupial of the frontal nose-shield and a fusion of the plesio- families are known flom Rackham's Roost (e.g. morphic double infraorbital foramen into the auta- dasy'r-rrids,peramelids, potoroids, etc.) most are pomorphic single foramen of Quaternary M. gigas represented by one or two specimens each. (there being little concomitent measurable differen- Dominating the assemblage is a single mammal ce in the molar region between the two species). family, the Muridae, represented by at least 12 spe- Speciation events associated with changes of shape cies, many of which are represented by hundreds of linked with size are often described in terms of specimens each (Archer et al. 1991). heterochrony and allometry. A change in the rate of growth and development of a feature due to a minor Arboreal mammals make up a large proportion of genetic change may have a large impact upon the the prey collected by Macroderma malugoro most adult phenotype. Such a change in gene regulation of which are missing from the Rackham's Roost may have occurred during the phylogenetic history assemblage collected by M. gigas. The obsewed of the lineage spawning M. koppa to produce the shift in prey assemblages between the sites might shorter-faced M. gigas. be due to a change in prey availability in the imme- diate area, probably as the result of a gradual Useful gene combinations may be preserved in a drying in climate in northern Australia at the end species to adapt local populations in minor ways to fluctuating environments. That at least the premo- of the Tertiary. The middle Miocene Gotham City 'plastic' lar region has remained in M. gigas is sug- Local Fauna appears to have been culled from a gested by the fact that these characters are among rainforest environment, the Pliocene Rackham's those that appear to distinguish geographic popula- Roost Local Fauna ftom a dry sclerophyll or wood- land, probably not unlike the Riversleigh area today tions of Quaternary M. gigas (Hand & York 1990) ; premolar characters are the only ones found in M. (Archer et al. I99I). Douglas (1967), Toop (1985) gigas to be highly correlated with each other and to and Schulz (1986) have studied the prey of modern covary relatively independently of other g.roups of M. gigas which inhabits a wide range of environ- characters (Hand & York 1990). It is also in this ments in northern Australia, from semi-arid to region that Pliocene representatives of M. glgos dif- rainforest habitats. In the forested coastal environ- fer from Quaternary specimens by way of variation ment around Rockhampton, central Queensland, in the number of infraorbital foramina. the prey chosen by M.glgos was found by Toop (1985) to be predominantly arboreal. Rodent The short face and exnansion of the frontal nose- remains were collected, but only in roosts adjacent shield in megadermatids may also correlate with to cleared and ploughed agricultural land where the the very large nose-leaf.Following Freeman's (1984) hunting of ground-dwelling animals was possible. argument, the head would need to be less negative- In northern ly tilted on the basicranial axis for the nose-leafto semi-arid and desert areas of Australia, be directed forward. The large eyes, large nose-leafi the most common mammalian remains in the fee- large erect ears and powerful canines could thus be ding roosts of M. gigas have been found to be, like in trained simultaneously on their prey, a procedure the Rackham's Roost deposit, rodents, dasyrrrids (Douglas that typifies the hunting behaviour of living mega- and small bats 1967 ; Schulz 1986). dermatids who inspect their surroundings from a Acknowledgements - Work at Riversleigh has been supported perch before ambushing and killing their prey by the Australian Research Council, the Department of the Arts, usually with one swift bite to the back of the head Sport, the Environment, Tourism and Territories, National (Kulzer et al. I9B4). Estate Programme Grants (Queensland), the Australian Geographic Society, ICI, the Museum and the PAI,AEOECOLOGY Queensland University of New South Wales. Access to comparative speci- Both the Gotham City and Rackham's Roost depo- mens was generously provided by B. Engesser, H. Felten, T Flannery, L. Gibson, M. Hugueney, P Mein, B. and sits are interpreted to Iargely represent the Sig6 G. Storch. The photos were taken by R. Arnett, University of New South remains of prey accumulated by, respectively, Wales and the SEMs in the Microscopy Unit, Biological Sciences, gigas, Macroderma malugara nov. sp. and M. mega- Macquarie University. I am particularly grateful to my col- dermatids of approximately equal size. However, in leagues Michael Archer and Henk Godthelp at the University of terms of vertebrate diversit5r, the far more extensi- New South Wales for their assistance and advice. During this ve Rackham's Roost deposit appears depauperate work, I was supported by a Special Research Grant from the with respect to the Gotham City deposit. For University of New South Wales. I thank M. Hugueney and B. example, among marsupial families represented in Sig6 for constructively criticizing a draft ofthis paper. 376 REFERENCES comments on megadermatid phylogenetics. Australian Mommalogy,8 : 5-43. - ADVANI R. 1981 - Seasonal fluctuations in the feeding TIAND S.J. 1987 Phylogenetic studies of Australian ecology of the Indian false vampire Megaderma lyra Tertiary bats : a summary of PhD thesis. Macrodertna, lyro (Chiroptera : Megadermatidae) in Rajasthan, 3 :9-12. - India. Zeit scltrift fiir S augetierkunde, 46 : 90 -93. HAND S.J. 1990 Australia's first Tertiary molossid ADVANI R. & MAKWANA S.C. 1981 - Composition and (Microchiroptera : Molossidae) : its phylogenetic and seasonal occurrence of animal remains in the roosting biogeographic implications. Memoirs of tlrc Queens- habitat of Megaderma lyra lyra in Rajasthan, India. land Museum. 28 : 175-192. Angewandt Zoologisclt, 68 : 175-182. HAND S.J. 1993 - First skull of a species of Hipposideros ARCHER M. & FLANNERY T.F. 1984 - Revision of the (Brachipposidaros) (Microchiroptera : Hipposideridae), extinct gigantic rat kangaroos (Potoroidae: Marsu-pia- from Australian Oligo-Miocene sediments. Mernoirs of Iia), with description of a new Miocene genus and spe- tlrc QueenslandMuseum, 3l :179-192. cies, anrd a new Pliestocene species of Propleopus. IIAND S.J. 1995 - First record of the genus Megaderma Jountal of Paleontology,89 : 1131-1149. GEOFFROY,1810 (Microchiroptera : Megadermatidae) -66. ARCHER M., GODTHELP H., IIAND S.J. & MEGIRIAN D. flom Australia. Palaeouertebrata, 24 : 47 1989 - Preliminary over-view of mammalian diversity, HAND S.J..ARCHER M. & GODTHELP H. 1989a - A fossil biostratigraphy, correlation and environmental change bat-rich, Oligo-Miocene cave-frll from Riversleigh evidenced by the fossil deposits of Riversleigh, north- Station, northwestern Queensland. Confbrence on western Queensland. Australian. Zoologist, 25 : 35-69. Australasian Vertebrate Eu olutiort, Pal aeontol ogy and ARCI-IER M., }IAND S.J. & GODTI]ELP H. 1991 - RiUCTS- Systematics,Abstracts'. 7 . leiglt. Reed Books, Sydney : 264 p. I{AND S.J,, ARCHER N{. & GODTHELP H. 1989b - ThC ARCHER M. & WADE M. 1976 - Results of the Ray E. Australian bat fauna : a 30 million-year history? Lemley Expeditions, Part 1. The Allingham Formation Australian Marnmol Society Conference, April 1989, and a new Pliocene vertebrate fauna from northern Abstracts : p.26. '. - Queensland. Mernoirs of the Queensland Museum,lT HAND S.J., ARCHER M. & GODTTIELP H. 1990 379-397. Australian fossil hipposiderids : new evidence about BARTHOLOMAI A. 1978 - The Macropodidae (Marsu-pia- the evolutionary history and radiation of Old World lia) from the Allingham Formation, northern leaf-nosed bats- Macroderma, 5 : 12. - Queensland. Results of the Ray E. Lemley expeditions, HAND S.J.,DAWSON L. & AUGEE M. 19BB MACTOdCTMA part 2. Metnoirs of tlrc Queensland Museum,18 : 127- koppa, a new Pliocene species of false vampire bat r43. (Microchiroptera: Megadermatidae) fiom Wellington BROSSETA. 1962 - The bats of central and western India. Caves, New South Wales. Records of tlte Australian Paft, 2. Jountal of tlrc Bombay Natural History Society, Museurn,40 :343-351. 59 :583-624. IIAND S.J. & YORK A. 1990 - Morphological variation in DOUGLAS A.1\{. 1967 - The natural history of the Ghost the dentition of the Australian Ghost Tiat, Macro- B at, M acrorlerma gig as (Microchiroptera, Megaderma- denna gigas (Microchiroptera : Megadermatidae). Aus- tidae), in Western Australia. \Yestern Australian. tralion Journol of Zoology,33 : 263-286. Naturalist, 10 : 125-138. KINGDON J. 1974 - East African Marnmals. Volume II, EMtrRSON S.B. & RADINSKY L. 1980 - Functional analy- Part A, Insectiuoresand Bats. Academic Press, London: sis of sabeftooth cranial morphology. Paleobiology, 6 : 341 p. 295-312. KULZER E., NELSON J.E., MCKEON J.L. & MOEHRES FLANNERYTF. & ARCHERM. 1987- Triclnsurusdickso- F.P 1984 - Prey-catching behaviour and echolocation ni and Strigocusus gymnotoides, new middle Miocene in the Australian Ghost Bat, Macroderma gigas phalangerids (Marsupialia) from northwestern (Microchiroptera : Megadermatidae). Australian '. Queensland. 1z ARCFIER N{. (ed.) Possums and Mcrmmalogy,T :37-50. Opossums : Studies in Euolution. Surrey Beatty and LEKAGUL B. & MCNEELY J.A. 7977 - Mammals of Sons and the Royal Zoological Society of New South Thailand. Association for the Conserwation of Wildlife, Wales: 527-536. Bangkok : 758 p. FREEMAN P.W. 1984 - Functional cranial analysis of MA\T{ARD SMITH J. & SAVAGE R.J.G. 1959 - The mecha- large animalivorous bats (Microchiroptera). Biological nics of mammalian jaws. School Science Reuiew,4O : Journal of tlrc Linnean Society, 2l : 387-408. 289-301. GODTHELP H. 1988 - Riversleigh scene 4 : Rackham's MENU H. & SIGE B.I97I - Nyctalodontie et myotodontie, Roost--the beginnings of the modern world. In IIAND importants caractdres de grades 6volutifs chez les chi- S.J. & ARCHER M. (eds) : The Antipodean Ark. Angus roptbres entomophages. Cotnptes Rendus de I'Aca- and Robertson :81-83. ddmie des Sciences,Paris,272: 1735-1738. GRIFFITFIS T.A., TRUCKENBROD A, & SPONHOLTZ P.J. SCHULZ M. 1986 - Vertebrate prey of the Ghost Bat, 1992 - Systematics of megadermatid bats (Chiroptera, Macroderma gigas, at Pine Creek, Northern Territory. Megadermatidae), based on hyoid morphology. Macroderma,2 :59-62. American Museum Nouitates,SO4l : L-21. SIGE B. 1976 - Les Megadermatidae (Chiroptera, HAND S.J. 1985 - New Miocene megadermatids Mammalia) miocdnes de Beni Mellal, Maroc. Gdologie (Chiroptera : Megadermatidae) I|om Australia with mdditerrandenne, 3 : 7 7-86. a,7n

SIGE 8., HAND S.J. & ARCHER M. 1982 - Arr Australian (Geoffioy) (Chiroptera, Megadermatidae). Zeit schrift Miocene Brachipposideros (Mammalia, Chiroptera) fi)r Tierpsychologie, 26 : 726-7 36. related to Miocene representatives from France. WOODBURNE M.O., TEDFORD R.H., ARCHER M,, TURN- - Palaeouertebrata, L2 : 149-171. BULL W.D., PLANE M.D. & LUNDtrLIUS E.L. 1985 TOOP J. 1985 - Habitat requirements, sur-vival strategies Biochronology of the continental mammal record of and ecology of the Ghost Bat Dobson (Microchiroptera, Australia and New Guinea. SpecictlPublication, Soutlt Megadermatidae), in central coastal Queensland. Australian Department of Mines and Energy, 5 :347- Macroderma, | :37-41. JO,l. VAUGIIAN T.A. 1976 - Nocturnal behaviour of the African (Cardioderma false vampire bat cor). Journal of S.J. HAND Mammalogy, 57 : 227-248. School of Biological Science WICKLER W. & UHRIG G. 1969 - Verhalten und okolo- University of New South Wales gische Nische der Gelbflugelfledermaus, Lavia ffons Sydney, NSW 2052, Australia