This article was downloaded by: [University of Regina] On: 15 May 2013, At: 02:58 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Alcheringa: An Australasian Journal of Palaeontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/talc20 New isolated pterodactyloid bones from the Albian Toolebuc Formation (western Queensland, Australia) with comments on the Australian pterosaur fauna Alexander W.A. Kellner a , Thomas H. Rich b , Fabiana R. Costa a , Patricia Vickers-Rich c , Benjamin P. Kear d , Mary Walters c & Lesley Kool c a Departamento de Geologia e Paleontologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil b Museum Victoria, PO Box 666, Melbourne, Victoria, 3001, Australia c School of Geosciences, Monash University, Melbourne, Victoria, 3800, Australia d Palaeobiology Programme, Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36, Uppsala, Sweden Published online: 01 Jun 2010. To cite this article: Alexander W.A. Kellner , Thomas H. Rich , Fabiana R. Costa , Patricia Vickers- Rich , Benjamin P. Kear , Mary Walters & Lesley Kool (2010): New isolated pterodactyloid bones from the Albian Toolebuc Formation (western Queensland, Australia) with comments on the Australian pterosaur fauna, Alcheringa: An Australasian Journal of Palaeontology, 34:3, 219-230 To link to this article: http://dx.doi.org/10.1080/03115511003656552 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and- conditions
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This article was downloaded by: [University of Regina]On: 15 May 2013, At: 02:58Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registeredoffice: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK
Alcheringa: An Australasian Journal ofPalaeontologyPublication details, including instructions for authors andsubscription information:http://www.tandfonline.com/loi/talc20
New isolated pterodactyloid bonesfrom the Albian Toolebuc Formation(western Queensland, Australia) withcomments on the Australian pterosaurfaunaAlexander W.A. Kellner a , Thomas H. Rich b , Fabiana R. Costa a
, Patricia Vickers-Rich c , Benjamin P. Kear d , Mary Walters c &Lesley Kool ca Departamento de Geologia e Paleontologia, Museu Nacional,Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, SãoCristóvão, 20940-040 Rio de Janeiro, RJ, Brazilb Museum Victoria, PO Box 666, Melbourne, Victoria, 3001,Australiac School of Geosciences, Monash University, Melbourne, Victoria,3800, Australiad Palaeobiology Programme, Department of Earth Sciences,Uppsala University, Villavägen 16, SE-752 36, Uppsala, SwedenPublished online: 01 Jun 2010.
To cite this article: Alexander W.A. Kellner , Thomas H. Rich , Fabiana R. Costa , Patricia Vickers-Rich , Benjamin P. Kear , Mary Walters & Lesley Kool (2010): New isolated pterodactyloid bonesfrom the Albian Toolebuc Formation (western Queensland, Australia) with comments on theAustralian pterosaur fauna, Alcheringa: An Australasian Journal of Palaeontology, 34:3, 219-230
To link to this article: http://dx.doi.org/10.1080/03115511003656552
PLEASE SCROLL DOWN FOR ARTICLE
Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions
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ALEXANDER W.A. KELLNER, THOMAS H. RICH, FABIANA R. COSTA,PATRICIA VICKERS-RICH, BENJAMIN P. KEAR, MARY WALTERS ANDLESLEY KOOL
KELLNER, A.W.A., RICH, T.H., COSTA, F.R., VICKERS-RICH, P., KEAR, B.P., WALTERS, M. & KOOL, L., September,2010. New isolated pterodactyloid bones from the Albian Toolebuc Formation (western Queensland, Australia) withcomments on the Australian pterosaur fauna. Alcheringa 34, 219–230. ISSN 0311-5518.
New isolated pterodactyloid bones from the Toolebuc Formation are described. The first one consists of a completewing metacarpal 212 mm long, representing an individual with an estimated wing span of 4 m. Small depressions onthe anterior surface are present and represent tooth marks showing that this specimen was subjected to scavengingprior to fossilization. The other bone consists of a three-dimensionally preserved cervical vertebra lacking most of theneural arch. The specimens are clearly referable to the derived pterosaur clade Pterodactyloidea. Based on severalfeatures such as the position of the pneumatic foramen and the particular shape and proportions of those elements,they possibly are members of, or closely related to, the Anhangueridae. The record of the Australian pterosaurs isreviewed here and represents the known southern distributional limit for Cretaceous pterosaurs, arguing againstsome older ideas of a more geographically restricted range for these flying reptiles.
Alexander W.A. Kellner [[email protected]] and Fabiana R. Costa [[email protected]], Departamento deGeologia e Paleontologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, SaoCristovao, 20940-040 Rio de Janeiro, RJ, Brazil; Thomas H. Rich [[email protected]], Museum Victoria, POBox 666, Melbourne, Victoria 3001, Australia; Patricia Vickers-Rich [[email protected]], Mary Walters[[email protected]] and Lesley Kool [[email protected]], School of Geosciences, MonashUniversity, Melbourne, Victoria 3800, Australia; Benjamin P. Kear [[email protected]], PalaeobiologyProgramme, Department of Earth Sciences, Uppsala University, Villavagen 16, SE-752 36 Uppsala, Sweden (addressfor correspondence) and Department of Genetics, La Trobe University, Melbourne, Victoria 3086, Australia. Received11.5.2009, revised 21.7.2009, accepted 3.8.2009.
PTEROSAUR remains are exceedinglysparse in Australia and limited to essentiallyisolated Cretaceous occurrences (Molnar &Thulborn 2007). The first remains of theseflying reptiles from this region were found inthe Lower Cretaceous Toolebuc Formation,near Boulia, western Queensland. Thismaterial includes a scapulocoracoid that,according to the original description, sharessome similarities with Pteranodon Marsh,
1876a (and Nyctosaurus Marsh, 1876b), oneincomplete vertebra, and a section of a lowerjaw, similar to those of the Ornithocheiruscomplex from the Cambridge Greensand(Molnar & Thulborn 1980).
Since then a few more pterosaur remainshave been found, mostly isolated specimensfrom the Toolebuc Formation, including anincomplete pelvis also regarded as repre-senting a taxon related to Pteranodon(Molnar 1987) and a partial skull and lowerjaw recently named Mythunga camaraby Molnar & Thulborn (2007). Another
ISSN 0311-5518 (print)/ISSN 1752-0754 (online)� 2010 Association of Australasian PalaeontologistsDOI: 10.1080/03115511003656552
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occurrence of these volant archosaurs is asmall crushed element from the EarlyCretaceous Otway Group (Otway Basin)of Victoria, first identified as a tibiotarsus ofa small pterosaur (Rich & Rich 1989), butlater reinterpreted as a metatarsal of a largepterosaur (Bennett & Long 1991). Theproximal end of an ulna from the MiriaFormation (upper Maastrichtian; Carnar-von Basin) of Western Australia was tenta-tively referred to an azhdarchid (Bennett &Long 1991). Kear et al. (2010) tentativelyascribe a jaw fragment to the Ornithocheir-idae from the Molecap Greensand (Ceno-manian–Coniacian; northern Perth Basin),Western Australia, completing the pub-lished pterosaur record from this region.There are more isolated specimens awaitingdetailed description (Fletcher et al. 2007).
Herein we describe two new specimensfrom the Toolebuc Formation. They consistof a complete wing metacarpal and a cervicalvertebra. We also review the current knowl-edge of the Australian pterosaur fauna.
Institutional abbreviations. AMNH, Amer-ican Museum of Natural History, NewYork, USA; MN, Museu Nacional/UFRJ,Rio de Janeiro, Brazil; NMV P, NationalMuseum of Victoria, Melbourne, Australia;QM, Queensland Museum, Brisbane,Australia; SAMP, South AustralianMuseum, Adelaide, Australia; YPM, YalePeabody Museum, New Haven, USA.
Geological settingSenior et al. (1975), Senior et al. (1978) andMoore et al. (1986) have provided detailedaccounts of the lithology, stratigraphicalrelationships and distribution (includingmaps) of the Toolebuc Formation. In brief,this unit comprises calcareous and carbonac-eous mudstones with abundant coquinite(composed mainly of Inoceramus Sowerby,1814) and bituminous shales. Stratigraphi-cally, it constitutes a medial subdivision
within the Rolling Downs Group (EromangaBasin) and has a conformable lower bound-ary with the Wallumbilla Formation and anupper conformity with the Allaru Mudstone(Moore et al. 1986). Age determinations forthe Toolebuc Formation based on dinofla-gellates and miospores indicate a latestmiddle to late Albian range: Pseudoceratiumludbrookiae dinoflagellate Zone and Coptos-pora paradoxa spore-pollen Zone (McMinn& Burger 1986, Moore et al. 1986).
The Toolebuc Formation exposuresaround Boulia incorporate around 25 –36 m (Boulia A type section: Senior et al.1975) of carbonaceousmudstone, with fossil-rich concretionary limestone; the latter iscommonly exposed and broken up duringexfoliation of the laminated parent rock.Invertebrate (ammonites and bivalves: Day1969) and vertebrate macrofossils (marinereptiles andfish:Kear 2003,Kear&Lee 2006,Kear 2007) are common and typically pre-served as complete/articulated specimensconsistent with accumulation in low-energyshallow-water conditions. The presence ofdark organic-rich shale layers indicates stra-tification within the water column includingdistinct dysaerobic bottom-waters (Mooreet al. 1986). Depositional interpretationssuggest that the Toolebuc Formation wasdeposited slowly in a restricted marineenvironment (Morgan 1980, McMinn &Burger 1986). This has been linked to theonset of a maximum transgressive episodeextending southwest into the Great ArtesianBasin complex from the southern TethysOcean (Morgan 1980; Fig. 1).
DescriptionWing metacarpalThe specimen is housed at Museum Victor-ia, Melbourne (NMV P197962) and a cast isdeposited at the Museu Nacional/UFRJ(MN 4721-V). It was collected from ex-posures of the Toolebuc Formation onSlashers Creek Station a few kilometres
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west of the homestead, which was locatedat 228520S, 1408430E. Slashers CreekStation is located east of Boulia, Queens-land, Australia.
NMV P197962 is a 212 mm long,complete right wing metacarpal (Fig. 2;Table 1). As has been observed previously inderived pterosaurs, despite this bone being acomparatively robust element, it is pneu-matic, with a very thin cortex from 0.7 to0.8 mm thick on the anterior and posteriorsurface to ca 1.4 mm on the dorsal and
ventral margins. Except for the distalarticulation (three-dimensionally pre-served), the main part of the shaft and theproximal articulation are flattened antero-posteriorly, a common feature in pterosaurspecimens (e.g. Wang et al. 2007). The bonesurface is remarkably well preserved, andthe internal surface, exposed in a few areas,indicates the presence of a trabecularsystem.
The proximal end is more expanded thanthe distal end, a feature typical of pterosaur
Fig. 1. Map of Jurassic–Cretaceous sedimentary basins in Australia showing the distribution of pterosaur fossils. (A)Giralia Range (Miria Formation) aff. Azhdarchidae: incomplete ulna (WAM 60.57); (B) Boulia (ToolebucFormation) aff. Anhangueridae: metacarpal (MNV P197162), vertebrae (QM F10614, SAM P41968), scapula-coracoid (QM F10612), lower jaw (QM F10613), incomplete pelvis (QM F12982); (C) Hughenden (ToolebucFormation) aff. Anhangueridae: Mythunga camara jaws (QM F18896); (D) Otway Ranges (Eumeralla Formation)Pterodactyloidea indet.: metatarsal (NM V185816); (E) Gingin (Molecap Greensand) ?Ornithocheiridae: jawfragment (see Kear et al. 2010—this volume). Base map after Turner et al. (2009).
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wing metacarpals. Despite the bone beingcrushed, it shows that the articulation withthe distal carpal series is formed by twoarticular surfaces and tuberculum. Thedorsally placed articular surface is slightlyconvex and separated from the secondarticular surface by a marked sulcus. Thesecond articular surface occupies most ofthe proximal end of the metacarpal and isalso slightly convex. The tuberculum is
blunt, projects slightly and is located ante-rior to the articular surfaces. This config-uration allows some rotation between thedistal syncarpal and the wing metacarpal.Some rugosities appear to be muscle scars.
Despite it being compressed, the trans-verse section of the shaft must have beenoval, with the longer axis oriented dorso-ventrally. In an anteroposterior view theventral margin is more concave than thedorsal one.
The distal end shows the typical doublecondyle articulation for the first wingphalanx. Both condyles are convex andsubequal in size, with the ventral oneprojecting more anteriorly. The distal endis preserved in three dimensions, whichsuggests that the bone in this region isdenser than elsewhere. On the anteriorsurface, proximal to the condyles, there isa well-marked concavity that does not end
Fig. 2. Wing metacarpal (NMV P197962) from dorsal (A, B), ventral (C, D), and proximal (E) views.dep¼depression; fo¼ foramen. Scale bar¼ 50 mm.
Bone characters Measurements
Length 212Maximum width of proximalarticulation
53.3
Maximum width of distalarticulation
34.3
Table 1. Measurements of metacarpal characters(in millimetres) of NMV P197962.
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in a pneumatic foramen. On the posteriorsurface, close to the more dorsally placedcondyle, a well-developed oval pneumaticforamen is evident (9.86 3.3 mm).
Another very interesting feature of thisspecimen is the presence of small depres-sions on the anterior surface (Fig. 3). Fourof them were identified, all subcircular.Three are located in the anterior dorsal halfand positioned in a longitudinal sequence,with a diameter of 4.4 mm, 4.1 mm and2.6 mm, respectively. The fourth lessmarked one, with a diameter of about3.3 mm, is positioned close to the ventralmargin. No such marks are clearly distin-guishable in other parts of this bone. Theentire bone surface is slightly depressedinwards.
These depressions are neither artefacts ofpreparation or preservation, nor insectborings, also reported in one pterosaur(Kellner & Langston 1996), since they donot pierce the bone. Furthermore, insectborings do not push the bone surface insidethe depression (or hole). These depressionsare not the result of plant roots that affectthe bone in a quite distinct manner, neitherdo they reflect attachment scars. Accordingto Bennett (1991), some specimens ofPteranodon (e.g. YPM 2452) have variableirregular scarring on the distal third to half
of the anterior surface of the shaft ofmetacarpal IV that may reflect intermeta-carpal ligaments. In the Australian meta-carpal IV all depressions are subcircular andaligned in a longitudinal sequence to forman arch. Moreover, they are locally placedat the anterior dorsal half of the bonesurface, slightly shifted to the margin of thebone. In Pteranodon, the scars are widelydispersed at the diaphysis of the metacarpal,being irregularly distributed and of distinctshapes. Furthermore, attachment scars lackthe collapsed bone particles that character-ize tooth puncture marks.
Cervical vertebraThe specimen SAMP41968 from the samedeposit is housed in the South AustralianMuseum. It consists of a three-dimension-ally preserved cervical vertebra with most ofits neural arch eroded away (Fig. 4; Table 2).As in all pterosaurs, this vertebra isprocoelous (Andres & Norell 2005) andhas a length/width ratio of52.5 (Unwin &Junchang 1997). The neural arch is shearedoff obliquely across the zygapophyses,clearly exposing the fine internal bone‘strut-work’ typical of pterosaurs; thus theneural spine height can not be determined.However, there is a very obvious excavation
Fig. 3. Tooth marks (tm) in dorsal view. Scale bar¼ 100 mm.
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conforming to the ventral margin of theforamen pneumaticum.
At the anterior end, a concave cotyle withan inverted subtriangular shape is flankedlaterally by prezygapophyses with subcircu-lar articular surfaces. It has a medianhypapophysis projecting from its ventralmargin. Remnants of the transverse processare also found at the anterior end of thevertebra. The centrum is strongly com-pressed dorsoventrally, near elliptical inarticular outline and preserving both post-exapophyses and a convex condyle at itsposterior end. It bears well-developed lateral
pneumatic foramina. In general form, thisbone closely resembles the cervical centra ofthe anhanguerid Anhanguera (Wellnhofer1991, Kellner & Tomida 2000) and thepteranodontid Pteranodon (Bennett 1991).
Regarding this similarity and consider-ing the close morphological resemblance ofcervicals 3 to 7 in Anhangueridae, Istiodac-tylidae and Pteranodon, it is difficult todetermine precisely the position of thisvertebra. However, based on the generalshape and compared with the above-men-tioned taxa, this element clearly belongs tothe middle part of the neck. It should be
noted that the presence of a bone bardividing the lateral pneumatic foramen, afeature that can be used to identify the 7thcervical in Anhanguera Campos & Kellner1985 (Kellner & Tomida 2000), is notevident in this specimen.
DiscussionPterosaur fossils are typically rare. Exceptfor a few deposits where complete specimensare found in abundance such as the Jurassicdeposits of Solnhofen (e.g. Wellnhofer 1970,1991), the Crato and Romualdo lagerstattenof the Lower Cretaceous Santana Group(e.g. Kellner & Campos 1999), the LowerCretaceous Yixian and Jiufotang formations(e.g. Wang et al. 2005) and the UpperCretaceous Niobrara Chalk in North Amer-ica (e.g. Bennett 2001), most pterosaurremains tend to be fragmentary. In themajority of localities, the record of ptero-saurs is limited to isolated bones (e.g. Kellner& Mader 1997, Kellner et al. 2003, Calvoet al. 2007, Costa & Kellner 2009). Even insome places where pterosaurs are abundant,such as in the Upper Cretaceous CambridgeGreensand deposits, they tend to be foundfragmented and incomplete (e.g. Unwin2001).
The Australian deposits with pterosaurremains are no exception. All of thepreviously found specimens are incompleteisolated bones. The wing metacarpal (NMVP197962) and the cervical vertebra(SAMP41968) here described are clearlyreferable to the Pterodactyloidea that unitesall more derived pterosaurs. They differfrom the non-pterodactyloids by beingmuch longer and larger elements (e.g.Kellner 2003, Unwin 2003).
The Pterodactyloidea is divided into twomain clades: the Archaeopterodactyloideaand the Dsungaripteroidea (Kellner 2003).The Archaeopterodactyloidea comprisessmall to middle-sized forms that overalltend to lack pneumatic foramina in theskeleton. NMV P197962 differs from all theknown archaeopterodactyloids by its largersize and the presence of a well-developedpneumatic foramen on the anterior marginclose to the distal articulation.
Comparing this specimen with the Dsun-garipteroidea, the Australian metacarpal isnot extremely elongated as in the Nycto-sauridae (Bennett 2001, Kellner 2003) andalso is much shorter than the conditionobserved in the Tapejaroidea, a clade thatunites Dsungaripteridae (Young 1973), Ta-pejaridae and Azhdarchidae (Unwin & Lu1997, Kellner 2004). NMV P197962 showsthe basic morphology and is within the sizerange of typical wing metacarpals of thePteranodontoidea and is, therefore, confi-dently allocated to this group. The Pter-anodontoidea is composed of three maingroups, namely Pteranodontidae, Istiodac-tylidae and Anhangueridae (Kellner 2003).According to Bennett (2001), the twoarticular surfaces of the proximal end ofthe metacarpal in Pteranodon are subequalin size, differing from the condition ob-served in NMV P197962 (and in theAnhangueridae and Istiodactylus Howse,Milner & Martill 2001), in which the dorsalsurface is significantly smaller than theventral one. Furthermore, Pteranodon has a
Bone characters Measurements
Maximum anteroposteriorlength
40.8
Maximum transverse width(across prezygapophyses)
37.8
Maximum anteroposteriorlength of centrum midline
32.4
Maximum transverse widthacross the posteriorexapophyses
20.9
Maximum dorsoventral heightof centrum anterior articularface (including hypapophysis)
9.5
Table 2. Measurements of cervical vertebracharacters (in millimetres) of SAMP41968.
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pneumatic foramen on the posterior surfacebetween the condyles of the distal articula-tion, whereas a pneumatic foramen is locatedon the anterior surface, close to the dorsalcondyle on the Australian specimen.
Compared with Istiodactylidae, the basicmorphology is the same (including theproximal articulation). Members of thisclade include Istiodactylus latidens Hooley,1913 and some Chinese species (e.g. Nurha-chius ignaciobritoi Wang et al. 2005, andIstiodactylus sinensis Andres & Ji, 2006),some lacking postcranial elements (Wanget al. 2008). A pneumatic foramen has notbeen reported on any of them, but this couldbe a result of these fossils having beencrushed.
Compared with all pteranodontoids,NMV P197962 shares more features withthe Anhangueridae (Wellnhofer 1985,Kellner & Tomida 2000), including theposition of the pneumatic foramen. There-fore, this metacarpal likely represents ananhanguerid from Australia.
Although it is difficult to estimate the sizeof a pterosaur from a single limb element,comparisons of NMV P197962 with Anhan-guera piscatorKellner & Tomida 2000, and acomplete wing from the Romualdo Forma-tion of the Santana Group in the AraripeBasin of Brazil (AMNH 22552 see Well-nhofer 1991) suggest that the Australianmetacarpal represents a pterosaur with awing span of about 4 m.
Because of its length/width ratioof 52.5, the specimen SAMP41968 is notconsistent with an archaeopterodactyloid oran azhdarchid pterosaur. The lateral pneu-matic foramina are larger than those ofAnhanguera, but smaller than those ofIstiodactylus. Primitively pterosaurian cer-vical vertebrae lack any lateral pneumaticforamen on the centrum (Wang et al. 2009).In Rhamphorhynchus von Meyer, 1846 thereare some pneumatic foramina beneath thediaphysis for the cervical ribs, but not onthe centrum (pers. obser.). The lateral sides
of the centrum laterally excavated to housepneumatic foramina is a condition observedin the Ornithocheiroidea excepting theAzhdarchidae; thus, it seems to be asynapomorphy of the former that wassecondarily lost in the latter (Kellner 2003).
Remnants of cervical ribs are found insome ornithocheiroids generally being com-pletely fused to the vertebra and withouttraceable limits. Taking the form of a smallbony arch spanning the gap between theprezygapophysial process and the centrum,they can appear in some azhdarchids, whoseouter wall is probably formed from aremnant of the cervical rib, now completelyfused to the vertebra (Unwin 2003).
The presence of postexapophyses oncervical vertebrae is a condition observedin the Dsungaripteroidea (Kellner 2003).Thus, regarding the pneumatic foraminaand the presence of postexapophyses as themost relevant diagnostic characters to theidentification of this specimen, this vertebrawould be assigned to a non-azhdarchidornithocheiroid, possibly an anhanguerid.
Other pterosaur material from AustraliaDespite being fragmentary, a few otherpterosaur specimens from Australia havebeen described. Molnar & Thulborn (1980)described a vertebra, a scapulocoracoid anda mandibular symphysis from marine Low-er Cretaceous sediments of the EromangaBasin, western Queensland. The vertebrawas regarded as being a dorsal vertebra, butmay be a last cervical vertebra of a largeanimal. Besides a thick and anteroposter-iorly elongated neural spine and a largepneumatic opening on the posterior surfaceof the transverse process, this specimen(QM F10614, cast MN 4721-V) shows noparticular diagnostic feature and couldrepresent a large and derived non-archae-opterodactyloid pterosaur.
The scapula and coracoid (QM F10612,cast MN 4721-V) are fused, representing
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an adult individual first regarded as apteranodontid (Molnar & Thulborn 1980)and latter as aff. Anhanguera (Molnar &Thulborn 2007). Based on the size of thecoracoid, which is larger than the scapula,this specimen can be assigned to thePteranodontoidea (Kellner 2003). Since thetips of both elements are missing, it is notclear whether the scapula is substantiallyshorter than the coracoid (e.g. sca/cor50.8), which is regarded as an anhan-guerid synapomorphy (Kellner 2004).Nevertheless, the particular shape of thescapula, being stout and having a con-stricted shaft, suggests that this specimen isclosely related to the Istiodactylidae andAnhangueridae and not to the Pteranodon-tidae. It could potentially represent a speciesclosely related to the Anhangueridae asproposed by Molnar & Thulborn (2007).
The lower jaw (QM F10613, cast MN4721-V) belongs to a toothed pterosaurtentatively regarded as closely related toOrnithocheirus (Molnar & Thulborn 1980)or Lonchodectes Hooley, 1914 (Molnar &Thulborn 2007). It is clear that this materiallacks the expanded distal tip diagnostic ofthe Anhangueridae and can not, therefore,be referred to this clade. The taxonomicstatus of Lonchodectes and Ornithocheirus isstill disputed, but this specimen lacks anydiagnostic feature of the Archaeopterodac-tyloidea. Nevertheless, it shows a well-developed mandibular groove suggestingthat it is closely related to the Anhanguer-idae (but not being a member of that group).
A partial pelvis (QMF 12982, cast MN4721-V) was regarded by Molnar (1987) asrepresenting an animal closely related toPteranodon. Later, Molnar & Thulborn(2007) noticed some similarities of thismaterial with Anhanguera. Unfortunately,the material is not sufficiently diagnostic toallow a positive identification.
The incomplete ulna described by Ben-nett & Long (1991) from the MaastrichtianMiria Formation of Western Australia was
originally thought to represent an azhdarch-id pterosaur. Since publication, additionalpterosaur specimens have been found, in-cluding tapejarid ulnae from the RomualdoFormation (Santana Group) that are similarto the Australian material. The ulnae ofDsungaripterus Young, 1964 also has simila-rities with this specimen. Based on thepresent geological record of pterosaurs, weagree with Bennett & Long (1991) that thismaterial might represent an animal moreclosely related to the Azhdarchidae sincetapejarids and dsungaripterids have not beenrecorded in Maastrichtian deposits thus far.
Kear et al. (2010) described a toothlessjaw fragment from the Cenomanian–Con-iacian Molecap Greensand, Perth Basin,Western Australia. The presence of labio-lingually compressed alveoli that are ante-rolaterally oriented, variable in shape, andwidely spaced is reminiscent of the Ornitho-cheiridae—a widespread clade of predomi-nantly Early Cretaceous pterodactyloids.
The last pterosaur specimen from Aus-tralia to be mentioned here is Mythungacamara, regarded by Molnar & Thulborn(2007) as a member of the Archaeopterodac-tyloidea (Early Cretaceous). It is known froma partial skull found in marine rocks of theToolebuc Formation (Albian) near Hughen-den, Queensland. However, based on pub-lished pictures, this important pterosaurspecimen shows strong teeth, particularlythe first preserved ones (Molnar & Thulborn2007, fig. 4). It is inconsistent with thedentition reported thus far for any archae-opterodactyloid (e.g.Wang et al. 2005, 2007).Furthermore, the size estimated for thispterosaur, over 4.5 m (Molnar & Thulborn2007), is much larger than the size of anyarchaeopterodactloid known to date (Wanget al. 2005). On the other hand, the size anddentition of Mythunga camara are moreconsistent with the Anhangueridae or Or-nithocheiridae (Kellner & Tomida 2000,Kellner 2003, Unwin 2001, 2003). Since thedistal end of the skull andmandible ismissing
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and the dorsal margin is broken where someof the most significant cranial synapomor-phies of the Anhangueridae are found (Kell-ner 2003), it is not certain whetherMythungais a member of this clade.
SummaryDespite the fragmentary nature of thepterosaur material from Australia, it is clearthat moderate diversity is present. There is apotential record of an azhdarchid in theUpper Cretaceous deposits of the WesternAustralia (Bennett & Long 1991), but thespecimens from the Albian Toolebuc For-mation are most important. The specimensstudied so far point to the presence oftoothed pterosaurs, one of which is a non-anhanguerid but belongs to a closely relatedtaxon (the lower jaw—QM F10613).Mythunga and the scapulocoracoid arepotential anhanguerids, but the best evidenceof this clade in Australia is the material heredescribed. No evidence of the Pteranodonti-dae, a clade of toothless pterosaurs, wasconclusively reported from these strata.
An interesting aspect of NMV P197962 isthe tooth marks. It is possible that thesemarks were made by some scavenger duringthe dismembering of the carcass. However,the metacarpal was not gnawed by thescavenger because there are just clean punc-tures at the bone surface rather thanscratches or grooves. This might be due tothe small amount of muscle associated withthis part of the pterosaur body, detractingfrom it being targeted as a good food source.
Lastly, the record of the Australianpterosaurs represents the southern limit ofdistribution for Cretaceous pterosaurs(Molnar & Thulborn 1980), arguing againstsome older ideas of restricted geographicrange. The occurrence of pterosaurs inAntarctica briefly mentioned by Hammer& Hickerson (1999) is another argumentagainst any particular geographic restrictionfor these volant archosaurs.
AcknowledgementsWe thank Mr and Mrs Pip Prince ofWestward Ho Station, Boulia, Queensland,for not only allowing access to SlashersCreek Station but also for permitting us toutilize the homestead on that property fora base of operations when the fossil wascollected in June. We are grateful to ChrisBennett (Fort Hays State University) andan anonymous reviewer for their commentson the manuscript. We would like also toacknowledge Steve McLoughlin for themap, Vanessa Machado (Museu Nacio-nal/UFRJ) for the drawings and IvanNunes (Museu Nacional/UFRJ) for help-ing with the figures. This project waspartially funded by Conselho Nacional deDesenvolvimento Cientıfico e Tecnologico(CNPq) (fellowship to FRC) with addi-tional support by the Fundacao CarlosChagas Filho de Amparo a Pesquisa doRio de Janeiro (FAPERJ, n8. E-26/152.885/2006 to AWAK) and ConselhoNacional de Desenvolvimento Cientıfico eTecnologico (CNPq, no. 304965/2006 – 5 toAWAK).
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