The first (Testudines: ) from the Cenozoic of


FITZGERALD, E.M.G. & KOOL, L., XX.XX.2015. The first fossil sea turtles (Testudines: Cheloniidae) from the Cenozoic of Australia. Alcheringa 39, xxx–xxx. ISSN 0311-5518

An isolated dentary and costal identified as cf. Pacifichelys and Cheloniidae indet., respectively, are described from the upper –lower Plio- cene Black Rock Sandstone of Beaumaris, Victoria, Australia. These remains represent the first fossil evidence of sea turtles from the Cenozoic of Australia. Neither of the can be referred to living genera, indicating that extinct cheloniids occurred in southeast Australian coastal waters for at least part of the late . Thus, the taxonomic composition of the current sea fauna of Australia was apparently established within the last five to six million .

Erich M. G. Fitzgerald [efi[email protected]] and Lesley Kool [[email protected]], Geosciences, Museum Victoria, GPO Box 666, Melbourne, Victoria, 3001, Australia. Received 26.6.2014; revised 9.8.2014; accepted 14.8.2014.

Key words: Pacifichelys, Neogene, Miocene, Pliocene, Victoria, marine, .

AUSTRALIAN are inhabited by six of the seven liv- 2012), sirenians (Fitzgerald et al. 2013), phocid seals ing species of the families Cheloniidae and (Fordyce & Flannery 1983), baleen whales (Fitzgerald , including one endemic species 2004, 2012), odontocetes (Chapman 1912, 1917) and (Márquez 1990). marine basins of northern rare remains of terrestrial dromornithid birds (Park & Australia have additionally produced an abundance of sea Fitzgerald 2012b) and marsupials (Piper et al. 2006). turtle fossils, including the extinct genera , To this rich local fauna we now add cheloniid turtles, Cratochelone, Bouliachelys and other indeterminate which constitute the first fossil evidence of sea turtles forms (Kear 2003, Kear & Lee 2006). Yet, a nearly from the Cenozoic of Australia. 66-million- gap in the Australian sea turtle record separates an upper fossil from Western Australia (Kear & Siverson 2010) and the Materials and methods diversity of sea turtles. Despite reports of alleged ‘turtle Institutional Abbreviations. LACM: Natural History ’ from the marine lower Pliocene Grange Burn Museum of Los Angeles County, Los Angeles, USA. Formation of Victoria (Chapman 1914, p. 47), Australia NMV D: Museum Victoria and Amphibians

Downloaded by [Monash University Library] at 16:56 18 October 2014 has remained the only continental mainland without sub- Collection, Melbourne, Australia. NMV P: Museum stantiated Cenozoic fossil sea turtles. Proximal to the Victoria Palaeontology Collection, Melbourne, Australia. Australian mainland, fossil sea turtles have been described from the Quaternary of New Guinea (Vis Comparative Material. Dermochelyidae (Dermochelys 1905), and the Paleocene (Fordyce 1979, Buchanan et al. coriacea): NMV D54542, D57420 and D74912. Chelo- 2007), (Fordyce 1979, Köhler 1995, Karl & niidae (Chelonia mydas): NMV D5828, D58526 and Tichy 2007, Grant-Mackie et al. 2011) and Miocene D74919. Cheloniidae (Caretta caretta): NMV D54543, (Buckeridge 1981) of New Zealand. D65879 and D74935. Cranial osteological terminology – The shallow marine upper Miocene lower Pliocene follows Gaffney (1979). Black Rock Sandstone of Beaumaris, Victoria (southeast Australia; Fig. 1), has produced the Beaumaris Local Fauna, including and rays (Kemp 1991), Geological setting fi osteichthyan sh (Chapman & Pritchard 1907), The coastal section at Beaumaris occurs onshore in the penguins (Park & Fitzgerald 2012a), diomedeid and cliffs, shore platform and shingle from pelagornithid seabirds (Wilkinson 1969, Fitzgerald et al. Table Rock about 1.6 km northeast to ‘Dog Tooth Rock’ (approximately opposite the intersection of Beach Road and Cliff Grove), and approximately 100 m out to © 2014 Museum Victoria sea as submarine outcrop (Gill 1957). The rock is 2 ERICH M. G. FITZGERALD AND LESLEY KOOL ALCHERINGA

144° 148° ferruginous sandstone containing burrows but no

carbonate (Abele et al. 1988, Wallace et al. 2005). The

urr M ay Rive r 36° Beaumaris section of the Black Rock Sandstone repre- Australia Victoria sents a shallow marine sandy lower shoreface facies 27° (Wallace et al. 2005). The nodule bed and overlying 6.7 m of the area shown below Bass Strait Black Rock Sandstone at Beaumaris constitute the type area shown at right 40° 0 1000 km 0 100 km section of the Cheltenhamian southeast Australian mar- Tasmania ine stage, which was originally correlated with the N 145°06´ upper Miocene (Singleton 1941). More recently, the Melbourne microfossil assemblage from the Black Rock Sandstone at Beaumaris has indicated its deposition during plank- tonic foraminiferal zones N17b–N18 (Mallett 1977), Beaumaris late Messinian–early , or about 4.5–6.5 Ma (McGowran et al. 2004, Hilgen et al. 2012). 87Sr/86Sr 38°03´ Beaumaris Bay 38°03´ dates from the basal nodule bed and overlying 5 m of Black Rock Sandstone at Beaumaris range between 6.2 Port Phillip Ma (within basal nodule bed) to 4.9 Ma (at 4.3 m high; Bay Dickinson et al. 2002, Wallace et al. 2005, Dickinson & Wallace 2009). These data indicate that the fossilifer- ous Black Rock Sandstone at Beaumaris was deposited between the latest Miocene and earliest Pliocene. Bass Strait The cheloniid fossils described here (NMV P240713, NMV P232865) were collected as float in the 0 10 km 145°06´ intertidal zone at Beaumaris; however their stratigraphic Fig. 1. Locality of Beaumaris in Victoria, southeastern Australia. provenance is not in doubt. Fine-grained, yellow, calcar- eous sandy matrix was found adhering to the fossils: exposed parallel (W–E) to the shoreline by a shallow this, and the relatively fine preservation of the bones asymmetrical anticline with its axis occurring at approx- themselves, suggests that the fossils were derived from imately the level (horizontally) of the intersection of the lower 6.7 m of the Black Rock Sandstone overlying Beach Road and Banksia Avenue (near 37°59ʹS, the basal nodule bed. Dickinson & Wallace (2009) 145°02ʹE; Gill 1957). From here, the strata dip eastward reported a spread of 87Sr/86Sr dates between 4.9 and 6.0 along strike at a shallow angle of ≤2° (Gill 1957). The Ma from this part of the section at Beaumaris. cliffs along the shoreline are parallel with the eroded Beaumaris Monocline, which has a seaward (SE) aver- Systematic palaeontology age dip angle of 10–20° (Gill 1957, Kenley 1967). The base of the sequence at Beaumaris consists of Class REPTILIA Laurenti, 1768 Fyansford Formation, which is discon- TESTUDINES Batsch, 1788 formably overlain by a thin (ca 20 cm thick) phosphatic Suborder Cope, 1868 nodule bed at the base of the Black Rock Sandstone, Superfamily CHELONIOIDEA Baur, 1893 Downloaded by [Monash University Library] at 16:56 18 October 2014 which has a maximum thickness of about 15 m (Kenley Family CHELONIIDAE Oppel, 1811 1967, Abele et al. 1988). The clayey limestone of the Pacifichelys Fyansford Formation is not exposed in onshore outcrop, cf. Parham & Pyenson, 2010 and only in limited areas on the sea bed close to shore where it is covered by beach sand (Gill 1957). The Material. NMV P240713, nearly complete left dentary, phosphatic nodule bed at the base of the Black Rock collected by Timothy Flannery ca January 1982 – Sandstone consists of phosphatic and limonitic intra- (Figs 2 3). clasts and mollusc shells, together with resistant and usually isolated and abraded vertebrate elements (e.g., Locality. Found as float below high tide level on the teeth, vertebrae, ribs, cetacean ear bones) within a western shore of Beaumaris Bay at Beaumaris, north- quartz-rich sandy matrix (Singleton 1941, Gill 1957, east side of Port Phillip Bay, central coastal Victoria, Wallace et al. 2005). The nodule bed is only exposed at southeast Australia, near 37°59ʹ34ʺS, 145°02ʹ32ʺE. low tide. The succeeding ca 6.7 of Black Rock – Sandstone consists of fossiliferous fine calcareous sands Stratigraphic unit. Upper Miocene lower Pliocene and silts, commonly burrowed and containing mollusc- Black Rock Sandstone. and echinoid- (especially Lovenia) rich layers (Gill 1957, Abele et al. 1988). The top 8.5 m of the Black Description. The dentary is nearly complete, relatively Rock Sandstone comprises sparsely fossiliferous broad and dorsoventrally flattened (Fig. 2A, C). The ALCHERINGA CENOZOIC SEA TURTLES FROM AUSTRALIA 3

Fig. 2. cf. Pacifichelys, NMV P240713, isolated left dentary in: A, dorsal; B, ventral; C, lateral; and D, medial views. Specimen whitened with ammonium chloride.

symphyseal length (20.0 mm) is 50.7% of the preserved of the dentary is smoothly rounded in the transverse dentary length (39.4 mm), similar in size to the juvenile plane. mandibles of Pacifichelys urbinai (Parham & Pyenson 2010, fig. 7). The triturating surface is expanded and flat to slightly concave transversely, with no accessory Comparisons. NMV P240713 compares most closely to fi ridge; its posterior edge is at a level just anterior to the the dentaries of Paci chelys urbinai Parham & position of the foramen dentofaciale majus. The broad Pyenson, 2010 and P. hutchisoni Lynch & Parham, and flattened triturating surface suggests specialization 2003 by having the following combination of fi fl for durophagous feeding (sensu Parham & Pyenson characters: low pro le in lateral view; wide and at 2010). Viewed dorsally, the labial and lingual edges of triturating surface; a relatively low labial ridge and the triturating surface converge posteriorly. In medial negligible development of a lingual ridge; sagittal view, the labial ridge curves dorsally towards the posi- swelling on the posterodorsal surface of the symphysis; tion of the coronoid and is higher than the lingual edge. a sulcus cartilaginis meckelii that does not reach the The labial ridge lacks development of any serrations or symphysis (Fig. 3); and a lateral surface that does not cusps. The width of the dentary at the level of the fora- expose an anterior extension of the surangular. Downloaded by [Monash University Library] at 16:56 18 October 2014 men dentofaciale majus is 26.3 mm, indicating that the The dentaries of dermochelyids and the cheloniids width of a hypothetically complete pair of mandibles Chelonia, Eretmochelys, Natator and Syllomus differ was approximately 52.6 mm at the level of the foramen from NMV P240713 by being transversely narrow and dentofaciale majus. At the posterior edge of the sym- high and by having a convex triturating surface with physis, there is a slight sagittal swelling. The sulcus significant lingual ridges (Zangerl et al. 1988, Zug cartilaginis meckelii is present on the medial aspect of 2001, Hasegawa et al. 2005, Parham & Pyenson 2010). the dentary; however, it does not extend anteriorly to The Dermochelys dentary differs from NMV P240713 the symphysis (Fig. 3). The foramen alveolare inferius by having a dorsally projecting anterior apex of the occurs adjacent to the posterodorsal edge of the sulcus symphysis, a strongly salient labial ridge, an anteropos- cartilaginis meckelii. Ventral to the posterior half of the teriorly short and transversely narrow triturating surface, sulcus cartilaginis meckelii is a region of rugose a strongly keeled ventral edge, a sulcus cartilaginis that represents the facet for the angular (Figs 2D, 3A). meckelii that extends anteriorly to the symphysis, and a Laterally, the external surface of the dentary is pitted. A lingual ridge that overhangs the ventromedial edge of distinct foramen dentofaciale majus occurs immediately the sulcus cartilaginis meckelii. The dentary of anterior to a small ovoid fossa on the posterolateral Chelonia further differs from NMV P240713 by expos- surface of the dentary that represents part of the ing laterally a V-shaped wedge of the surangular that insertion area for the m. adductor mandibulae externus extends forward to the level of the foramen dentofaciale pars superficialis (Jones et al. 2012). The ventral edge majus; having a distinct sagittal ridge along the dorsal 4 ERICH M. G. FITZGERALD AND LESLEY KOOL ALCHERINGA

majus (Zangerl et al. 1988). Dentaries of Carettini (Caretta + Lepidochelys) differ from NMV P240713 by being of higher profile in lateral view, and having a sulcus cartilaginis meckelii that extends anteriorly to the symphysis (Jones et al. 2012). The dentary of Lepidochelys further differs from NMV P240713 by exposing laterally a V-shaped wedge of the surangular that extends forward to the level of the foramen dentof- aciale majus (Zangerl et al. 1988, Jones et al. 2012).

Cheloniidae indet.

Referred material. NMV P232865, proximal end of a costal bone, collected by Brian Crichton on 17 June 1976 (Fig. 4).

Locality. Found as float below high tide level near ‘Dog Tooth Rock’ on the western shore of Beaumaris

Fig. 3. Cheloniid left mandibles in posteromedial view. A,cf.Pacifichelys, NMV P240713, isolated left dentary; B, Pacifichelys hutchisoni, LACM 103351, fused left and right mandibles; C, Caretta caretta, NMV D54543, fused left and right mandibles. The shaded region outlined by a broken line in A represents the facet for the angular. The shaded region in B and C outlined by a thin solid line represents the angular. The thick solid line in B and C represents the posterior margin of the external surface of the dentary. Specimen in A is whitened with ammonium chloride.

surface of the symphysis, a salient longitudinal acces- sory ridge between the labial and lingual ridges, and a sulcus cartilaginis meckelii that extends anteriorly to the Downloaded by [Monash University Library] at 16:56 18 October 2014 symphysis. The dentary of Erquelinnesia differs from NMV P240713 by having a symphysis with a posterior edge located posterior to the level (anteroposteriorly) of the foramen dentofaciale majus, and an enlarged fossa for the m. adductor mandibulae externus pars superficia- lis on the lateral surface of the dentary (Gaffney 1979, Hirayama 1995). The dentary of Natator further differs from NMV P240713 by having a distinct sagittal ridge along the dorsal surface of the symphysis, a much lar- ger foramen dentofaciale majus, a sulcus cartilaginis meckelii that extends anteriorly to the symphysis and a lingual ridge that overhangs the ventromedial edge of the sulcus cartilaginis meckelii (Zangerl et al. 1988). The dentary of Eretmochelys further differs from NMV P240713 by being transversely narrow with straight lat- eral edges in dorsal view (Wyneken 2001), and expos- Fig. 4. Cheloniidae gen. et sp. indet., NMV P232865, isolated proxi- ing laterally a V-shaped wedge of the surangular that mal end of costal in: A, dorsal; B, visceral; and C, proximal views. extends forward to the level of the foramen dentofaciale Specimen whitened with ammonium chloride. ALCHERINGA CENOZOIC SEA TURTLES FROM AUSTRALIA 5

Bay at Beaumaris, northeast side of Port Phillip Bay, Described costals of Pacifichelys are similar to NMV central coastal Victoria, southeast Australia, near P232865 in having a dorsal surface that lacks sculptur- 37°59ʹ15ʺS, 145°03ʹ00ʺE. ing and scale sulci (Lynch & Parham 2003, Parham & Pyenson 2010). The preserved features of NMV Stratigraphic unit. Upper Miocene–lower Pliocene P232865, combined with its incomplete preservation, Black Rock Sandstone. prevent referral to any known cheloniid .

Description. NMV P232865 represents the proximal end Discussion of a costal bone measuring 47.8 mm wide and 56.8 mm The identification of two late Miocene–early Pliocene long. All margins are abraded, but the longitudinal sea turtle fossils from Australia, one similar to the stem margins appear to approximate their original extent, apart cheloniid Pacifichelys and the other an indeterminate from a conchoidal break halfway down one side. The cheloniid, contributes to: (1) the post-Mesozoic history longitudinal margins are parallel to each other, and of sea turtles in and around Australia; (2) the palaeobi- the proximal margin is both straight and perpendicular to ogeography of Pacifichelys; and (3) the timing of the the longitudinal margins. This suggests that the costal ‘modernization’ of sea turtle faunas. Until now, the evo- was located in the mid-region of the , perhaps lutionary history of sea turtles in Australia could be representing the second, fourth or sixth costal. The summarized as extinct protostegids and possible derm- dorsoventral thickness of the costal averages between ochelyids from ca 66–100 Ma during the Cretaceous 3.5 mm and 4.6 mm. Dorsally, the bone is flat, and the (Kear 2003, Kear & Lee 2006, Kear & Siverson 2010), texture on the dorsal surface is finely vascular with a followed by the living fauna of cheloniids and Derm- subtle radiating pattern. There is no distinct sculpturing ochelys during the Holocene. The intervening ca 66 or scute sulci. The visceral surface is smooth and slightly million years of the Cenozoic has hitherto yielded no polished by abrasion. The obliquely abraded rib head is fossil evidence on the of the Australian sea narrow and oval in cross-section and is situated close to turtle fauna. The late Miocene–early Pliocene fossils the proximal margin of the costal. The exposed internal reported here, therefore, show that: (1) extinct cheloni- structure of the costal reflects histological adaptation to ids occurred in southeast Australian coastal waters for an aquatic environment by increased vascularization of at least part of the late Neogene; (2) at least one of the the outer cortical layer and an increase in the extinct taxa (cf. Pacifichelys) was durophagous; and (3) homogeneity of both the cortical and cancellous bone the taxonomic composition of the current sea turtle (Scheyer & Sander 2007). fauna of Australia was apparently established within the last five to six million years. Comparisons. NMV P232865 possesses the following The stem cheloniid Pacifichelys has hitherto been combination of features unique to Cheloniidae costals: a recorded from the middle Miocene (ca 11–16 Ma) of relatively broad and flat dorsal surface with no distinct the eastern North Pacific (California: Lynch & Parham sculpturing; a thin dorsoventral diameter; parallel longi- 2003) and the eastern South Pacific (: Parham & tudinal margins; thin external cortical bone; and Pyenson 2010). The recovery of remains from Australia strongly vascularized internal cortical/cancellous bone. that are generically compatible with this genus might, The costals of Dermochelyidae differ from NMV therefore, extend its geochronologic range by at P232865 by being narrow and strap-like, tapering least five million years into the Miocene–Pliocene Downloaded by [Monash University Library] at 16:56 18 October 2014 distally, and having a lozenge-shaped cross-section. (ca 4.9–6.2 Ma) and evince a trans-Pacific distribution. Among Cheloniidae, the costals of Syllomus and Published records of sea turtles from the late Natator differ from NMV P232865 by having an ‘irreg- Miocene worldwide are few, consisting only of: indeter- ular vermiculation’ sculpture on their dorsal surface minate Cheloniidae from the Tortonian Gatun Formation (Zangerl et al. 1988, p. 26; Lynch & Parham 2003, of Panama (Cadena et al. 2012); and ?Cheloniidae from Hasegawa et al. 2005); and one or more scute sulci on the Tortonian Diest Formation of Belgium (Bosselaers the dorsal surface of their proximal end (Zangerl et al. et al. 2004). Therefore, the potential late Messinian 1988, Hasegawa et al. 2005). The costals of the occurrences from Beaumaris, despite being fragmentary, cheloniids Chelonia, Eretmochelys, Lepidochelys and add useful evidence for interpreting the late Neogene Caretta differ from NMV P232865 by having one or evolution of sea turtle faunas. Specifically, the ‘modern- more scute sulci on the dorsal surface of their proximal ization’ of cheloniid assemblages globally from the end (Zangerl & Turnbull 1955, Witzell 1983). The cos- middle Miocene to early Pliocene followed a pattern of tals of the early carettinin grandaeva stem and aberrant crown cheloniid in paral- Leidy, 1851 differ from NMV P232865 by having one lel with the emergence of extant cheloniid genera. or more scute sulci on the dorsal surface of their proxi- Certainly, the geologically earliest crown cheloniid is mal end (although the sulci are not deep: Zangerl & probably Procolpochelys grandaeva from the early Turnbull 1955, p. 354), and a proximal suture edge Miocene of the western North Atlantic (Zangerl & formed by three facets (Zangerl & Turnbull 1955). Turnbull 1955, Sugarman et al. 1993), whereas the 6 ERICH M. G. FITZGERALD AND LESLEY KOOL ALCHERINGA

earliest known appearances of the living Chelonia, CHAPMAN,F.&PRITCHARD, G.B., 1907. Fossil fish remains from the – Tertiaries of Australia. Part II. Proceedings of the Royal Society Lepidochelys and Caretta are from the mid late – – of Victoria 20,59 75. Miocene (ca 10.3 13.6 Ma) of the western North COPE, E.D., 1868. On the origin of genera. Proceedings of the Atlantic (Dodd & Morgan 1992). The divergent extinct Academy of Natural Sciences of Philadelphia 20, 242–300. Syllomus survived into the early Pliocene of the western DICKINSON, J.A. & WALLACE, M.W., 2009. Phosphate-rich deposits North Atlantic (Zug 2001), implying that the taxonomic associated with the Mio-Pliocene unconformity in south-east Australia. Sedimentology 56, 547–565. composition of the modern sea turtle fauna was estab- DICKINSON, J.A., WALLACE, M.W., HOLDGATE, G.R., GALLAGHER, S.J. & lished within the last four million years. This report of THOMAS, L., 2002. Origin and timing of the Miocene-Pliocene Pacifichelys-like fossils from the late Miocene–early unconformity in southeast Australia. Journal of Sedimentary – Pliocene thus hints at a longer and more recent Research 72, 288 303. DODD, C.K. Jr & MORGAN, G.S., 1992. Fossil sea turtles from the early temporal overlap between stem- and crown-group Pliocene Formation, central . Journal of cheloniids, including living genera. Herpetology 26,1–8. FITZGERALD, E.M.G., 2004. A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia. Memoirs of Museum Victoria 61, 183–208. Acknowledgements FITZGERALD, E.M.G., 2012. Possible neobalaenid from the Miocene of Australia implies a long evolutionary history for the pygmy right The authors thank W. Joyce and J. Parham for advice whale Caperea marginata (Cetacea, Mysticeti). Journal of and provision of materials, and K. Roberts and K. Smith Vertebrate Paleontology 32, 976–980. for access to Museum Victoria’s Reptiles and Amphibi- FITZGERALD, E.M.G., PARK,T.&WORTHY, T.H., 2012. First giant ans Collection. J. Parham and M. Rabi provided helpful bony-toothed bird (Pelagornithidae) from Australia. Journal of Vertebrate Paleontology 32, 971–974. reviews. S. McLoughlin and B. Kear assisted with FITZGERALD, E.M.G., VELEZ-JUARBE,J.&WELLS, R.T., 2013. Miocene editorial comments. J. Velez-Juarbe supplied the photo- sea cow (Sirenia) from Papua New Guinea sheds light on sirenian graph of LACM 103351 used in Fig. 3. Brian Crichton evolution in the Indo-Pacific. Journal of Vertebrate Paleontology – is thanked for his collection and donation of NMV 33, 956 963. FORDYCE, R.E., 1979. Records of two Paleogene turtles and notes on P232865 to Museum Victoria. Astrid Werner is thanked other Tertiary reptilian remains from New Zealand. New Zealand for her skilful preparation of NMV P232865 and other Journal of Geology and Geophysics 22, 737–741. significant fossils. FORDYCE, R.E. & FLANNERY, T.F., 1983. Fossil phocid seals from the late Tertiary of Victoria. Proceedings of the Royal Society of Victoria 95,99–100. GAFFNEY, E.S., 1979. Comparative cranial morphology of Recent and References fossil turtles. Bulletin of the American Museum of Natural History ABELE, C., GLOE, C.S., HOCKING, J.B., HOLDGATE, G., KENLEY, P.R., 164,65–376. LAWRENCE, C.R., RIPPER, D., THRELFALL, W.F. & BOLGER, P.F., GILL, E.D., 1957. The stratigraphical occurrence and palaeoecology of 1988. Tertiary. In Geology of Victoria.DOUGLAS, J.G. & some Australian Tertiary marsupials. Memoirs of the National FERGUSON, J.A., eds, Victorian Division, Geological Society of Museum of Victoria 21, 135–203. Australia, Melbourne, 251–350. GRANT-MACKIE, J.A., HILL,J.&GILL, B.J., 2011. Two Eocene cheloni- BATSCH, A.J.G.C., 1788. Versuch einer Anleitung, zur Kenntniß und oid turtles from Northland, New Zealand. New Zealand Journal Geschichte der Thiere und Mineralien, für akademische Vorlesun- of Geology and Geophysics 54, 181–194. gen entworfen, und mit den nöthigsten Abbildungen versehen. HASEGAWA, Y., HIRAYAMA, R., KIMURA, T., TAKAKUWA, Y., NAKAJIMA,H. Erster Theil. Allgemeine Geschichte der Natur; besondre der &KENKYUKAI,GUNMA KOSEIBUTSU, 2005. Skeletal restoration of a Säugthiere, Vögel, Amphibien und Fische. Akademische fossil sea turtle, Syllomus, from the middle Miocene Haratajino Buchhandlung, Jena, 528 pp. Formation, Tomioka Group, Gunma Prefecture, central Japan. BAUR, G., 1893. Notes on the classification of the Cryptodira. Bulletin of Gunma Museum of Natural History 9,29–64. American Naturalist 27, 672–674. (in Japanese). BOSSELAERS, M., HERMAN, J., HOEDEMAKERS, K., LAMBERT, O., MARQUET, ILGEN OURENS AN AM EU OYES

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