Microchiroptera: Mystacinidae) from Australia, with a Revised Diagnosis of the Genus

New Miocene Icarops material (Microchiroptera: Mystacinidae) from Australia, with a revised diagnosis of the genus

SUZANNE HAND, MICHAEL ARCHER & HENK GODTHELP

HAND, S.l., ARCHER, M. & GODTHELP, H., 2001:12:20. New Miocene lcarops material (Microchiroptera: Mystacinidae) from Australia, with a revised diagnosis of the genus. Memoirs of the Association of Australasian Palaeontologists 25,139-146. ISSN 0810-8889

New fossil material referable to Icarops paradox Hand et al., 1998 is described from the early Miocene Judith's Horizontalis Site in the Riversleigh World Heritage Property of northwestern Queensland. Fused dentaries contain the partial lower dentition of I. paradox. The diagnosis of the genus Icarops is revised. The new material confirms the identity of Icarops species as mystacinids and enablesre-examination of interrelationships between extinct and extant members of this Gondwanan bat family.

S.J: Hand, M. Archer* & H. Godthelp, School of Biological Science, University of New South Wales,New South Wales, 2052; * also Australian Museum, 6-8 College St, Sydney, New South Wales,2000. Received ]4 December 2000

Keywords: Mystacinidae, Icarops, Mystacina, bat, lower dentition, Miocene, Riversleigh

THE FIRST pre-Pleistocene record for the QMF refers to specimens held in the fossil Mystacinidae and first record of this bat family collections of the QueenslandMuseum, Brisbane. from outside New Zealand were reported by Hand et al. ( 1998) from Miocene sedimentsin Australia. SYSTEMAllC PALAEONTOLOGY Three species of the new mystacinid genus Icarops were described: Icarops breviceps from OrderCIllROPTERAB1wnenbach, 1779 the middle Miocene Bullock Creek deposit of the SuborderMICROCIllROPTERA Dobson, 1875 Northern Territory; I. aenae from the early SuperfamilyNocmIoNoIDEA Van Va1en, Miocene Wayne's Wok deposit, D Site Plateau, 1979 Riversleigh, northwestern Queensland; and I. Family MYSTACINIDAE (Gray, 1843) paradox from the early Miocene Neville's Garden Site, D Site Plateau, Riversleigh. The Australian Icarops Hand et al., 1998 mystacinids were described mainly on the basis of dentaries; only I. breviceps was represented Type species. Icarops breviceps Hand, Murray, also by teeth: an M2 and MJ preserved in the Megirian, Archer & Godthelp, 1998 holotype. Additional mystacinid material has since been Revised diagnosis. Speciesof Icarops differ from collected from another early Miocene deposit on Mystacina species in the following combination Riversleigh 's D Site Plateau,Judith 's Horizontalis of features:It width and length approximately equal Site. This material includes a mandible with fused (i.e. not especially wide); p 4 less than half MI dentaries containing the partial lower dentition length, with two roots oriented longitudinally or for a small species of Icarops. Based on only slightly obliquely with respect to the comparisonsof size and alveolar patterns,the new toothrow; all cusps of lower molars including material is referableto I. paradox. Icarops paradox entoconids taller and more distinct, with MI-2 is one of 44 species of fossil microchiropterans talonid wider than trigonid. recorded from Riversleigh's Oligo-Miocene and Pliocene limestone deposits (Archer et al. 1994, Other species. Icarops aenae Hand et al., 1998; Hand 1999). Icarops paradox Hand et al., 1998 In this paper, the new Icarops material is described and the diagnosis of the genus revised. Icarops paradox (Fig. lA-D) Taxonomy and dental terminology follows Miller (1907) and Hand et al. (1998). Stratigraphic Holotype. QMF20808, partial mandible preserving nomenclature for the Riversleigh region follows fragments of left and right dentaries with alveoli Archer et al. (1994) and Creaser(1997). The prefix forLI"C, andP.,andRC1,P,4andM,. Type localitY~ Neville's Garden Site, D Site evidence of crowding of teeth with overlapping Plateau, Riversleigh World Heritage Property, of the alveoli. Most of the crown ofP 4is broken Lawn Hill National Park, northwestern offbut it has two roots oriented slightly obliquely Queensland (Archer et al. 1994). with respect to the toothrow; the anterior alveolus is significantly smaller than the posterior one and New material. QMF31561, a partial, fused displaced towards the buccal margin of the mandible with left dentary preserving C1 and MI-3 toothrow. The anterior alveolus for MI is also and alveoli for I1 and p 24' and right dentary with slightly buccally displaced. broken p 4' MI-3 and associated RIl. The lower dental formula is 11'CI' P2.4' M123. The alveoli for the canines (but not the mciso"rs; Locality, age and lithology. Judith's Horizontalis contra Hand et at. 1998) are ventrally displaced Site, D Site Plateau,Riversleigh (seeCreaser 1997), with respect to the toothrow. The margins of the a freshwater limestone deposit interpreted as early canine alveoli are smooth and complete. The Miocene, based on stratigraphy and contained posterior alveolus of P4 is compressed by the local fauna {Archer et al. 1994: Creaser 1997). anteriorly inclined alveolus for the anterior root ofMI. The posterior alveolus for MI is larger than Associatedfauna. Other vertebrates recovered so the anterior one. far from the Judith 's Horizontalis Site include: RI1 was found within the jaw but was removed frogs, lizards, turtles, passerinebirds, bandicoots, during preparation of the mandible. It is relatively a marsupial mole, yalkaparidontid, burramyid large, and deeply and evenly trifid. The crown is possum, balbarid kangaroo, carnivorous not markedly extended backwards; its width and kangaroo, and a megadermatid bat (A. Gillespie, length are approximately equal. pers. comm.). Cl is simple in form; its posterior base has a small, rounded median cusp, marked by a small Revised diagnosis. Smaller than I. breviceps and notch for the anterior edge of P2. The basal I. aenae, and with P 2 larger and P4 more cingulum is almost complete except at its most transversely oriented than in I. aenae (Hand et al. anterolingual point; its anterolingual edge has a 1998). Additionally, it differs from I. breviceps in distinct convexity just above the level of the MJ being less reduced (with trigonid and talonid incisors; here the cingulum is broken and of subequal width). connected to a short crest on the lower part of the crown. Description. The left and right dentaries of Most of the crown ofRP 4 is missing; the roots QMF31561 are completely fused, with no sign of remain. Judging from its alveoli and remaining the symphysis, and meet at an angle of crown, P4 was longer than P2 and less than half approximately 45°. The mandible's anterior the length ofMJ .Its two roots are oriented slightly margin, in lateral profile, is receding ventrally, obliquely with respect to the toothrow; the rounded and without a chin process. A ventral anterior alveolus is smaller than the posterior one mandibular shelf extends posteriorly to the and is close to the buccal margin of the dentary. alveolus for the anterior root of P4. A small but MI has two roots and five distinct cusps, the deep invagination in the shelf's posteroventral hypoconulid being a small cingular cusp. face marks the attachment point for the digastric Although individualised, the cusps appear muscle/s (in the holotype of I. paradox, relatively low and inclined rather than tall and QMF20808, the site for attachmentof the digastric upright. The trigonid is conspicuously narrower, muscles was not clear). The deI!tary decreasesin but approximately the same length as the talonid. depth only slightly from P2 to below the posterior The protoconid and hypoconid are the dominant root of MJ. The ascending ramus, condyle and cusps in height and volume, but they are not angular process are not preserved. massive. The protoconid is only just taller than Unlike other Icarops specimens(including the the hypoconid which is taller than the metaconid; holotype of I. paradox), there is no mental foramen the entoconid and paraconid are subequal in beneath P2 nor further posteriorly. There is, height; all are much taller than the hypoconulid. however, a small, dorsally directed foramen below The protoconid shows conspicuously more wear and betweenthe alveoli forC1 and 11.This foramen than the other cusps. The paracristid is just longer appearsto be homologous with that found in this than the metacristid; the protoconid contributions region in other Icarops specimens. The incisor to the paracristid and metacristid are longer than alveolar region is well preserved and contained a the paraconid and metaconid contributions; the single pair of incisors. The canine alveolus is large metaconid and protoconid contributions of the and oval. The single alveolus for Pis smaller than metacristid aremore equal and meet at a more acute the canine alveolus. As in the holotype, there is angle. The cristid obliqua, in occlusal view, is AAP Memoir 2

Fig. l.lcaropsparadox Hand, Murray, Megirian, Archer & Godthelp, 1998. QMF31561, from the early Miocene Judith's Horizontalis Site, Riversleigh World Heritage Property, Lawn Hill National Park, northwestern Queensland. A, lateral view; B, oblique occlusal view; C, stereopair, occlusal view. Scale bar indicates 2 mrn. v= ventral mandibular shelf. uncurved and contacts the trigonid at a point between the hypoconid and entoconid. A pre- directly below the junction of the components of entocristid, straight and steeply dipping, links the the metacristid. In lateral view there is an inflexion entoconid to the trigonid at the base of the along the cristid obliqua close to the trigonid. The metaconid (making an angle with the metaconid hypocristid extends from the hypoconid directly of just less than 900). The angle between the para- to the entoconid, almost perpendicular to the axis and metacristids is relatively broad, at of the toothrow, isolating the small hypoconulid approximately 750. The cristid obliqua and and thereby exhibiting the myotodont condition paracristid are almost parallel to each other. There (Menu & Sige 1971 ). The greatly bowed inflexion is a uniform, non-sinuous, continuous anterior, in the hypocristid occurs approximately midway buccal and posterior cingulum, terminated at its posterolingual end well short of the hypoconulid molars, including entoconids, taller and more providing a notch for the anterior cingulum of distinct, MI-2 talonid wider than trigonid. MrArelatively strong but short lingual cingulum occurs at the base of the trigonid. Comparisons with molossids. Icarops species M2 is described insofar as it differs from MI. share with many molossids (Hand et al. 1998): a M2 is narrower and shorter in length but taller deep and robust dentary with mental foramen than MI. The trigonid and talonid are wider but occurring beneath p 2; a single pair of lower slightly shorter, the protoconid is taller (and not incisors; two anterior premolars overlapping so as worn) but the hypoconid is of similar size, and that their roots are oblique in the toothrow; general the metaconid and protoconid contributions of appearance of MI and M2, notably myotodonty, the metacristid are more equal and meet at a more talonid wider than trigonid, tall hypoconid, acute angle. The angle made between the paraconid development similar to metaconid (i.e. paracristid and metacristid is approximately 600. without strong reduction; except in the molossid The short lingual cingulum at the base of the Cheiromeles torquatus), cristid obliqua complete trigonidisweakerinM (andM ). (Sige 1985, p. 170-1). Additional dental features M3 is described insofar as it differs from M1-r now found to be similar to molossids include C( It is a slightly narrower, shorter tooth. The trigonid being simple in form without a long heel, and p 4 is similar in width to the talonid, and the talonid is being smaller than MI. longer than the trigonid. The protoconid is the Dental features of Icarops paradox now known tallest and most massive cusp, the paracristid is to differ from molossids include: a trifid rather than longer than the metacristid, the protoconid bifid single incisor; CI with posterobasal cusp; p 2 contribution to the paracristid being particularly less reduced than in most molossids (although in long. All cusps are present, including the molossids it is double-rooted even when very hypoconulid. reduced e.g. molossines, cheiromeles etc.); no sinuous buccal cingulum in MI-J' talonid not much Measurements. Maximum dentary depth (below wider than trigonid, without pronounced buccal alveolus for PJ 1.30 mm, CI-M3length 5.65 mm, bulging of the protoconid and hypoconid; less MI-31ength3.83 mm. hypsodont. Also unlike Icarops species, many molossids (e.g. Cheiromeles) have nyctalodont COMPARISONS or submyotodont lower molars, and in Comparisons with other mystacinids. Icarops Cheiromeles and many molossines MJ is more paradox differs from the Australian Miocene I. reduced. breviceps and I. aenae in being smaller, with M} lessreduced (with trigonid and talonid of subequaJ Comparisons with noctilionoids. Icarops species width) than in at least I. breviceps; p 21argerand p 4 share with South America 's noctilionids a deep more transversely oriented than in I. aenae. As in and robust dentary with tall ascending ramus (as other Icarops species, the area of attachment for in I. breviceps) and fused dentary symphysis, a digastric muscle(s) appearsto be well developed single pair of lower incisors, two anterior in this specimen of I. paradox (see Hand et al. premolars, and myotodont lower molars (Hand et 1998).Unlike the holotype QMF20808 (andan other al. 1998). However, noctilonids differ strikingly extinct and extant bats examined), QMF31561 has from Icarops (and Mystacina) species in having no mental foramen; the condition is interpreted to M2 with extremely wide talonid, wall-like pre- be an aberrant condition in this individual. entocristid and cristid obliqua extending to the Icarops paradox and New Zealand's lingual margin of the crown, more obliquely Quaternary Mystacina species (i.e., M. oriented P4' and small, lingually displaced P .The tuberculata and the recently extinct M. robusta ) new Icarops paradox material additionally Jiffers share the following dental and mandibular in its C1with posterobasal cusp, and the single features:a fused dentary symphysis; a single trifid lower incisor being trifid rather than bifid. lower incisor; CI with posterior heel and The speciose bat family Phyllostomidae posterobasal cusp; and myotodont lower molars. exhibits an enormous range of dental Icarops paradox differs from Mystacina species morphologies which reflects a great variety of in the following features: II width and length diets (including blood, nectar, fruit, flesh and approximately equal (i.e. not especially wide); far insects)making phyllostomids much more difficult less massive canine with shorter heel, less to characterise than other bat groups. However, developed posterobasal cusp and better Icarops species appear to differ from developed buccal cingulum; p 41essthan half MI phyllostomids (and mormoopids ) in the following length with its two roots oriented slightly combination of features: reduction of the lower obliquely to the toothrow; all cusps of lower incisors to one pair, fusion of dentary symphysis, t\AP Mernoi relatively large p 2 with small p 4' and myotodont lower than the metaconid. The talonid ofM3 may lower molars. Icarops species share with many be greatly or little reduced. phyllostomids and mormoopids trifid rather than bifid lower incisors. Comparisons with extinct bat families (archaeonycteridids, palaeochiropterygids and Comparisons with vespertilionids. Vesper- hassianycteridids). The most archaic of early and tilionids lack a fused dentary symphysis, retain middle Eocene bats are referred to a number of two or more pairs of lower incisors, and tend to extinct families and lack most of the derived have lower molars with talonid wider than trigonid. features that distinguish bats of modern families Most groups are dominated by taxa with (seeRussell & Sige 1970,Russell et al. 1973,Smith nyctalodont lower molars but some ( e.g. Myotis, & Storch 1981, Habersetzer & Storch 1987, Plecotus, Scoteanax, Chalinolobus) have Simmons & Geisler 1998). Icarops speciesdiffer myotodont molars like Icarops species. from these early bats as follows. In Similarities with the new Icarops paradox material archaeonycteridids, palaeochiropterygids and include: trifid lower incisors; p 2and p 4relatively hassianycteridids (including species of small and with p 2singly rooted. No vespertilionids Archaeonycteris, Ageina, Australonycteris, have reduced the lower incisors to one pair, nor Icaronycteris, Palaeochiropteryx, Cecilio- have a fused dentary or an extended heel on C,. nycteris, Matthesia, Stehlinia and Hassia- nycteris), unlike species of Icarops, the dentary Comparisons with nataloids s. I. Natalids, symphysis is not fused, at least two pairs of lower thyropterids, furipterids and myzopodids are incisors are present, p 3 is retained, the trigonid grouped together here in the superfamily cuspsof the lower molars are typically much taller Nataloidea following Simmons (1998). Also than the talonid cusps, the talonid is unreduced grouped here are the kerivoulines (species of and hypoconulid more buccally situated, and all Kerivoula and Phoniscus) which are generally cusps are more individualised. The lower molars included in either the Natalidae or Vespertilionidae. of most of these archaic bats exhibit the All taxa lack a fused dentary symphysis, retain nyctalodont pattern ( or pre-nyctalodont pattern two or more pairs of lower incisors, and tend to with the hypoconulid more buccally situated) but have lower molars with talonid wider than trigonid. some palaeochiropterygids ( e.g. in the genera The lower canine is generally exceptionally Palaeochiropteryx and Stehlinia; see Sige 1997 slender with a subterete shaft. Most have on the reassignment of Stehlinia to this family) nyctalodont lower molars but kerivoulines have exhibit the myotodont pattern. In Icarops species myotodont molars like Icarops species. the toothrow is relatively much shorter Similarities to the new Icarops paradox material (foreshortened) than in archaic bats. include: the presence of trifid incisors (although they retain three pairs); p 2and p 4being relatively DISCUSSION small; and p 2 singly rooted. Comparisons of the lower dentition of Icarops paradox with those of other microchiropterans Comparisons with rhinolophoids. Icarops confirm its identity as a member of the family species differ from rhinolophoids in their fused Mystacinidae. Icarops paradox and New dentary symphysis, single pair of lower incisors Zealand's Quaternary Mystacina tuberculata and and in their talonid morphology. Three pairs of M. robusta share a unique combination of dental lower incisors are retained by nycteridids and two and dentary features including a fused dentary by megadermatids and hipposideriQs; these are symphysis; presence of a single trifid incisor; CI commonly trifid. In rhinolophoids p 2is small, p 3is with posterior heel and posterobasal cusps; an<1 retained by rhinolophids and some archaic myotodontlowermolars. hipposiderids, the talonid of the lower molars The new material also enables further tends to be rather low, narrow and short and examination of the relationships within this displays the nyctalodont pattern. Australasian bat family. Hand et al. (1998) suggested,on the basis of the fossil material then Comparisons with emballonuroids. Three pairs known, that Icarops species were probably of lower incisors are retained by Emballonura plesiomorphic with respect to Quaternary and New World emballonuroid taxa, and two pairs mystacinids, i.e. M. tuberculata and M. robusta. by Taphozous and Saccolaimus; these are Study of more of the lower dentition supports typically trifid and the outer incisor is sometimes this hypothesis. Icarops paradox differs from separated from the canine by a small gap. The Mystacina species in: ~ width and length being molars are usually but not always nyctalodont, approximately equal; L. far less massive with and the paraconid is reduced and conspicuously shorter heel, less developed posterobasal cusps and better developedbuccal cingulum; P41ess than archaeonycteridid from Australia (Hand et al. half Mi length with its two roots oriented slightly 1994), to which the much younger and more obliquely to the toothrow; all cusps of lower derived mystacinids bear little resemblance. The molars, including entoconids, taller and more oldest fossil bats in South America are a late distinct, M talonid wider than trigonid. All of Oligocene Brazilian molossid and middle Miocene these dentalteatures appear to be less specialised Colombian taxa, including the phyllostomids than in New Zealand's Mystacina species, Notonycteris magdalenensis and Tonatia sp., a although some ( e.g. loss of two pairs of lower glossophagine, phyllostomine and Noctilio incisors, fused dentary symphysis) are more albiventris (Czaplewski 1997). derived than found in other microchiropteran Isolated teeth of mystacinids (yet to be families including, for example, rhinolophoids, described) have now been recovered from late emballonuroids, vespertilionids, archaeo- Oligocene deposits at Riversleigh (and possibly nycteridids, palaeochiropterygids and hassia- South Australia) confirming that the family nycteridids. Mystacinidae was representedin Australia at least The relationships of mystacinids to other 25 million years ago. It survived here until at least families of bats are less clear. Mystacina 12 million years ago, occurring in middle Miocene tuberculata, a semi-terrestrial microchiropteran deposits at Bullock Creek, Northern Territory, but endemic to New Zealand, is the sole surviving had disappearedfrom at least the Riversleigh area member of the Mystacinidae. The oldest record by 3-5 million years ago as evidenced by their of the family in New Zealand is 18,000 years bp absencefrom the early Pliocene Rackham's Roost (Mystacina spp.) from Hermit's Cave, near deposit (Archer et al. 1994). The decline and, Charleston, South Island, NZ (Worthy & ultimately, extinction of this lineage in Australia Holdaway 1994). Mystacina was interpreted by may reflect the intense drying of Australia since Piersonet al. ( 1986),on the basisofimmunological the late Miocene which resulted in the replacement distance, to be a member of the South American ofwet forests by open woodlands and grasslands superfamily Noctilionoidea, evidently dispersing over much of the continent. The sole surviving from South America to New Zealand some 35 member of the mystacinid lineage is now restricted million years ago. Subsequently,Simmons ( 1996) to New Zealand's endemic Gondwanan-type found. using a 'total evidence' approach in which forests dominated by Nothofagus, Podocarpus, morphological, reproductive, behavioural, DNA Dacrydium and kauri trees (Daniel & Williams and other data are included in a single data set, 1984). that mystacinids are probably basal to a large Hand et al. (1998) have argued that the group of bats including the cosmopolitan presence of plesiomorphic mystacinids in the Vespertilionidae, Molossidae and SouthAmerican Australian Tertiary record (the only pre- Nataloidea, implying that ancestral mystacinids Pleistocene record for mystacinids) most diverged from other bats at least 55 million years parsimoniously suggests that Australia was the ago. DNA-DNA hybridisation studies by Kirsch immediate source of New Zealand's Quaternary et al. (1998) confirmed the affinity of Mystacina mystacinids. New Zealand separatedfrom the rest with noctilionoids as suggested by earlier of Gondwana 80-90 million years ago (Fleming serological studies (contra Simmons 1996) but 1979), long before the world's first bats had suggested separation from its South American evolved. Australia, Antarctica and South America relatives occurred between 54 and 66 million years were connectedas the last fragments ofGondwana ago. DNA sequencing studies (Kennedy et al. until about 35 million years ago (Veevers 1991), 1999) also indicate a mystacinid-noctilionoid throughout the period when most modern bat relationship,but differ from other molecular results lineages first appeared in the fossil record (Sige in placement of Mystacina within the superfamily 1991, Simmons 1996, 1998). The southern closer to phyllostomids and mormoopids; they supercontinent possibly shared a number of bat calculatethe time of divergenceof Mystacina from families, but precisely when and where the curious other bat taxa to be anywhere from 45 to 68 million mystacinids originated is yet to be discovered. years ago. Hershkovitz (1972) and Pierson (1986) have A CKN OWLEDGEMENffl argued for a Southern Hemisphere origin for the Work at Riversleigh has been supported by world's extant bat radiation on the basis of the Australian Research Council, Department of distributions of endemic bat families. Sige ( 1991) the Environment, Sport and Territories, National proposed that modern bat groups evolved from Estate Programme Grants (Queensland), isolated immigrant archaic groups in the Southern Queensland National Parks and Wildlife Service, Hemisphere in the early Eocene. The oldest bat in the Australian Geographic Society, Linnean the Southern Hemisphere is a 55-million-year-old Society of New South Wales. ICI. Oueensland AAP Memoir :

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