The first fossil sea turtles (Testudines: Cheloniidae) from the Cenozoic of Australia

The first fossil sea turtles (Testudines: Cheloniidae) from the Cenozoic of Australia ERICH M. G. FITZGERALD and LESLEY KOOL 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 Miocene lower Pliocene 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 fossils can be referred to living genera, indicating that extinct cheloniids occurred in southeast Australian coastal waters for at least part of the late Neogene. Thus, the taxonomic composition of the current sea turtle fauna of Australia was apparently established within the last five to six million years. Erich M. G. Fitzgerald [efitzgerald@museum.vic.gov.au] and Lesley Kool [koollesley@gmail.com], 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, vertebrate. AUSTRALIAN seas are inhabited by six of the seven living sea turtle species of the families Cheloniidae and Dermochelyidae, including one endemic species (Márquez 1990). Cretaceous marine basins of northern Australia have additionally produced an abundance of sea turtle fossils, including the extinct genera Notochelone, Cratochelone, Bouliachelys and other indeterminate forms (Kear 2003, Kear & Lee 2006). Yet, a nearly 66-million-year gap in the Australian sea turtle record separates an upper Maastrichtian fossil from Western Australia (Kear & Siverson 2010) and the Holocene diversity of sea turtles. Despite reports of alleged turtle bones from the marine lower Pliocene Grange Burn Formation of Victoria (Chapman 1914, p. 47), Australia has remained the only continental mainland without substantiated Cenozoic fossil sea turtles. Proximal to the Australian mainland, fossil sea turtles have been described from the Quaternary of New Guinea (Vis 1905), and the Paleocene (Fordyce 1979, Buchanan et al. 2007), Eocene (Fordyce 1979, Köhler 1995, Karl & Tichy 2007, Grant-Mackie et al. 2011) and Miocene (Buckeridge 1981) of New Zealand. The shallow marine upper Miocene lower Pliocene Black Rock Sandstone of Beaumaris, Victoria (southeast Australia; Fig. 1), has produced the Beaumaris Local Fauna, including sharks and rays (Kemp 1991), osteichthyan fish (Chapman & Pritchard 1907), penguins (Park & Fitzgerald 2012a), diomedeid and pelagornithid seabirds (Wilkinson 1969, Fitzgerald et al. 2014 Museum Victoria http://dx.doi.org/10.1080/03115518.2015.964047 2012), sirenians (Fitzgerald et al. 2013), phocid seals (Fordyce & Flannery 1983), baleen whales (Fitzgerald 2004, 2012), odontocetes (Chapman 1912, 1917) and rare remains of terrestrial dromornithid birds (Park & Fitzgerald 2012b) and marsupials (Piper et al. 2006). To this rich local fauna we now add cheloniid turtles, which constitute the first fossil evidence of sea turtles from the Cenozoic of Australia. Materials and methods Institutional Abbreviations. LACM: Natural History Museum of Los Angeles County, Los Angeles, USA. NMV D: Museum Victoria Reptiles and Amphibians Collection, Melbourne, Australia. NMV P: Museum Victoria Palaeontology Collection, Melbourne, Australia. Comparative Material. Dermochelyidae (Dermochelys coriacea): NMV D54542, D57420 and D74912. Cheloniidae (Chelonia mydas): NMV D5828, D58526 and D74919. Cheloniidae (Caretta caretta): NMV D54543, D65879 and D74935. Cranial osteological terminology follows Gaffney (1979). Geological setting The coastal section at Beaumaris occurs onshore in the cliffs, shore platform and beach shingle from 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 sea as submarine outcrop (Gill 1957). The rock is

2 ERICH M. G. FITZGERALD AND LESLEY KOOL ALCHERINGA 27 area shown at right 0 1000 km 0 N 38 03 Bass Strait 10 km Australia 0 100 km Melbourne Beaumaris Bay area shown below Port Phillip Bay 145 06 145 06 144 148 Murray Victoria Bass Strait Tasmania Beaumaris exposed parallel (W E) to the shoreline by a shallow asymmetrical anticline with its axis occurring at approximately the level (horizontally) of the intersection of Beach Road and Banksia Avenue (near 37 59ʹS, 145 02ʹE; Gill 1957). From here, the strata dip eastward along strike at a shallow angle of 2 (Gill 1957). The cliffs along the shoreline are parallel with the eroded Beaumaris Monocline, which has a seaward (SE) average dip angle of 10 20 (Gill 1957, Kenley 1967). The base of the sequence at Beaumaris consists of middle Miocene Fyansford Formation, which is disconformably overlain by a thin (ca 20 cm thick) phosphatic nodule bed at the base of the Black Rock Sandstone, which has a maximum thickness of about 15 m (Kenley 1967, Abele et al. 1988). The clayey limestone of the Fyansford Formation is not exposed in onshore outcrop, and only in limited areas on the sea bed close to shore where it is covered by beach sand (Gill 1957). The phosphatic nodule bed at the base of the Black Rock Sandstone consists of phosphatic and limonitic intraclasts and mollusc shells, together with resistant and usually isolated and abraded vertebrate elements (e.g., teeth, vertebrae, ribs, cetacean ear bones) within a quartz-rich sandy matrix (Singleton 1941, Gill 1957, Wallace et al. 2005). The nodule bed is only exposed at low tide. The succeeding ca 6.7 metres of Black Rock Sandstone consists of fossiliferous fine calcareous sands and silts, commonly burrowed and containing molluscand echinoid- (especially Lovenia) rich layers (Gill 1957, Abele et al. 1988). The top 8.5 m of the Black Rock Sandstone comprises sparsely fossiliferous River 36 40 38 03 Fig. 1. Locality of Beaumaris in Victoria, southeastern Australia. ferruginous sandstone containing burrows but no carbonate (Abele et al. 1988, Wallace et al. 2005). The Beaumaris section of the Black Rock Sandstone represents a shallow marine sandy lower shoreface facies (Wallace et al. 2005). The basal nodule bed and overlying 6.7 m of the Black Rock Sandstone at Beaumaris constitute the type section of the Cheltenhamian southeast Australian marine stage, which was originally correlated with the upper Miocene (Singleton 1941). More recently, the microfossil assemblage from the Black Rock Sandstone at Beaumaris has indicated its deposition during planktonic foraminiferal zones N17b N18 (Mallett 1977), late Messinian early Zanclean, or about 4.5 6.5 Ma (McGowran et al. 2004, Hilgen et al. 2012). 87 Sr/ 86 Sr dates from the basal nodule bed and overlying 5 m of Black Rock Sandstone at Beaumaris range between 6.2 Ma (within basal nodule bed) to 4.9 Ma (at 4.3 m high; Dickinson et al. 2002, Wallace et al. 2005, Dickinson & Wallace 2009). These data indicate that the fossiliferous Black Rock Sandstone at Beaumaris was deposited between the latest Miocene and earliest Pliocene. The cheloniid fossils described here (NMV P240713, NMV P232865) were collected as float in the intertidal zone at Beaumaris; however their stratigraphic provenance is not in doubt. Fine-grained, yellow, calcareous sandy matrix was found adhering to the fossils: this, and the relatively fine preservation of the bones themselves, suggests that the fossils were derived from the lower 6.7 m of the Black Rock Sandstone overlying the basal nodule bed. Dickinson & Wallace (2009) reported a spread of 87 Sr/ 86 Sr dates between 4.9 and 6.0 Ma from this part of the section at Beaumaris. Systematic palaeontology Class REPTILIA Laurenti, 1768 Order TESTUDINES Batsch, 1788 Suborder CRYPTODIRA Cope, 1868 Superfamily CHELONIOIDEA Baur, 1893 Family CHELONIIDAE Oppel, 1811 cf. Pacifichelys Parham & Pyenson, 2010 Material. NMV P240713, nearly complete left dentary, collected by Timothy Flannery ca January 1982 (Figs 2 3). Locality. Found as float below high tide level on the western shore of Beaumaris Bay at Beaumaris, northeast side of Port Phillip Bay, central coastal Victoria, southeast Australia, near 37 59ʹ34ʺS, 145 02ʹ32ʺE. Stratigraphic unit. Upper Miocene lower Pliocene Black Rock Sandstone. Description. The dentary is nearly complete, relatively 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 dentary length (39.4 mm), similar in size to the juvenile mandibles of Pacifichelys urbinai (Parham & Pyenson 2010, fig. 7). The triturating surface is expanded and flat to slightly concave transversely, with no accessory ridge; its posterior edge is at a level just anterior to the position of the foramen dentofaciale majus. The broad and flattened triturating surface suggests specialization for durophagous feeding (sensu Parham & Pyenson 2010). Viewed dorsally, the labial and lingual edges of the triturating surface converge posteriorly. In medial view, the labial ridge curves dorsally towards the position of the coronoid and is higher than the lingual edge. The labial ridge lacks development of any serrations or cusps. The width of the dentary at the level of the foramen dentofaciale majus is 26.3 mm, indicating that the width of a hypothetically complete pair of mandibles was approximately 52.6 mm at the level of the foramen dentofaciale majus. At the posterior edge of the symphysis, there is a slight sagittal swelling. The sulcus cartilaginis meckelii is present on the medial aspect of the dentary; however, it does not extend anteriorly to the symphysis (Fig. 3). The foramen alveolare inferius occurs adjacent to the posterodorsal edge of the sulcus cartilaginis meckelii. Ventral to the posterior half of the sulcus cartilaginis meckelii is a region of rugose bone that represents the facet for the angular (Figs 2D, 3A). Laterally, the external surface of the dentary is pitted. A distinct foramen dentofaciale majus occurs immediately anterior to a small ovoid fossa on the posterolateral surface of the dentary that represents part of the insertion area for the m. adductor mandibulae externus pars superficialis (Jones et al. 2012). The ventral edge of the dentary is smoothly rounded in the transverse plane. Comparisons. NMV P240713 compares most closely to the dentaries of Pacifichelys urbinai Parham & Pyenson, 2010 and P. hutchisoni Lynch & Parham, 2003 by having the following combination of characters: low profile in lateral view; wide and flat triturating surface; a relatively low labial ridge and negligible development of a lingual ridge; sagittal swelling on the posterodorsal surface of the symphysis; a sulcus cartilaginis meckelii that does not reach the symphysis (Fig. 3); and a lateral surface that does not expose an anterior extension of the surangular. The dentaries of dermochelyids and the cheloniids Chelonia, Eretmochelys, Natator and Syllomus differ from NMV P240713 by being transversely narrow and high and by having a convex triturating surface with significant lingual ridges (Zangerl et al. 1988, Zug 2001, Hasegawa et al. 2005, Parham & Pyenson 2010). The Dermochelys dentary differs from NMV P240713 by having a dorsally projecting anterior apex of the symphysis, a strongly salient labial ridge, an anteroposteriorly short and transversely narrow triturating surface, a strongly keeled ventral edge, a sulcus cartilaginis meckelii that extends anteriorly to the symphysis, and a lingual ridge that overhangs the ventromedial edge of the sulcus cartilaginis meckelii. The dentary of Chelonia further differs from NMV P240713 by exposing laterally a V-shaped wedge of the surangular that extends forward to the level of the foramen dentofaciale 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 dentofaciale 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). 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 accessory ridge between the labial and lingual ridges, and a sulcus cartilaginis meckelii that extends anteriorly to the 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 superficialis 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 larger 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 lateral edges in dorsal view (Wyneken 2001), and exposing laterally a V-shaped wedge of the surangular that extends forward to the level of the foramen dentofaciale Locality. Found as float below high tide level near Dog Tooth Rock on the western shore of Beaumaris Fig. 4. Cheloniidae gen. et sp. indet., NMV P232865, isolated proximal end of costal in: A, dorsal; B, visceral; and C, proximal views. Specimen whitened with ammonium chloride.

ALCHERINGA CENOZOIC SEA TURTLES FROM AUSTRALIA 5 Bay at Beaumaris, northeast side of Port Phillip Bay, central coastal Victoria, southeast Australia, near 37 59ʹ15ʺS, 145 03ʹ00ʺE. Stratigraphic unit. Upper Miocene lower Pliocene Black Rock Sandstone. Described costals of Pacifichelys are similar to NMV P232865 in having a dorsal surface that lacks sculpturing and scale sulci (Lynch & Parham 2003, Parham & Pyenson 2010). The preserved features of NMV P232865, combined with its incomplete preservation, prevent referral to any known cheloniid genus. Description. NMV P232865 represents the proximal end of a costal bone measuring 47.8 mm wide and 56.8 mm long. All margins are abraded, but the longitudinal margins appear to approximate their original extent, apart from a conchoidal break halfway down one side. The longitudinal margins are parallel to each other, and the proximal margin is both straight and perpendicular to the longitudinal margins. This suggests that the costal was located in the mid-region of the carapace, perhaps representing the second, fourth or sixth costal. The dorsoventral thickness of the costal averages between 3.5 mm and 4.6 mm. Dorsally, the bone is flat, and the texture on the dorsal surface is finely vascular with a subtle radiating pattern. There is no distinct sculpturing or scute sulci. The visceral surface is smooth and slightly polished by abrasion. The obliquely abraded rib head is narrow and oval in cross-section and is situated close to the proximal margin of the costal. The exposed internal structure of the costal reflects histological adaptation to an aquatic environment by increased vascularization of the outer cortical layer and an increase in the homogeneity of both the cortical and cancellous bone (Scheyer & Sander 2007). Comparisons. NMV P232865 possesses the following combination of features unique to Cheloniidae costals: a relatively broad and flat dorsal surface with no distinct sculpturing; a thin dorsoventral diameter; parallel longitudinal margins; thin external cortical bone; and strongly vascularized internal cortical/cancellous bone. The costals of Dermochelyidae differ from NMV P232865 by being narrow and strap-like, tapering distally, and having a lozenge-shaped cross-section. Among Cheloniidae, the costals of Syllomus and Natator differ from NMV P232865 by having an irregular vermiculation sculpture on their dorsal surface (Zangerl et al. 1988, p. 26; Lynch & Parham 2003, Hasegawa et al. 2005); and one or more scute sulci on the dorsal surface of their proximal end (Zangerl et al. 1988, Hasegawa et al. 2005). The costals of the cheloniids Chelonia, Eretmochelys, Lepidochelys and Caretta differ from NMV P232865 by having one or more scute sulci on the dorsal surface of their proximal end (Zangerl & Turnbull 1955, Witzell 1983). The costals of the early carettinin Procolpochelys grandaeva Leidy, 1851 differ from NMV P232865 by having one or more scute sulci on the dorsal surface of their proximal end (although the sulci are not deep: Zangerl & Turnbull 1955, p. 354), and a proximal suture edge formed by three facets (Zangerl & Turnbull 1955). Discussion The identification of two late Miocene early Pliocene sea turtle fossils from Australia, one similar to the stem cheloniid Pacifichelys and the other an indeterminate cheloniid, contributes to: (1) the post-mesozoic history of sea turtles in and around Australia; (2) the palaeobiogeography of Pacifichelys; and (3) the timing of the modernization of sea turtle faunas. Until now, the evolutionary history of sea turtles in Australia could be summarized as extinct protostegids and possible dermochelyids from ca 66 100 Ma during the Cretaceous (Kear 2003, Kear & Lee 2006, Kear & Siverson 2010), followed by the living fauna of cheloniids and Dermochelys during the Holocene. The intervening ca 66 million years of the Cenozoic has hitherto yielded no fossil evidence on the evolution of the Australian sea turtle fauna. The late Miocene early Pliocene fossils reported here, therefore, show that: (1) extinct cheloniids occurred in southeast Australian coastal waters for at least part of the late Neogene; (2) at least one of the extinct taxa (cf. Pacifichelys) was durophagous; and (3) the taxonomic composition of the current sea turtle fauna of Australia was apparently established within the last five to six million years. The stem cheloniid Pacifichelys has hitherto been recorded from the middle Miocene (ca 11 16 Ma) of the eastern North Pacific (California: Lynch & Parham 2003) and the eastern South Pacific (Peru: Parham & Pyenson 2010). The recovery of remains from Australia that are generically compatible with this genus might, therefore, extend its geochronologic range by at least five million years into the Miocene Pliocene (ca 4.9 6.2 Ma) and evince a trans-pacific distribution. Published records of sea turtles from the late Miocene worldwide are few, consisting only of: indeterminate Cheloniidae from the Tortonian Gatun Formation of Panama (Cadena et al. 2012); and?cheloniidae from the Tortonian Diest Formation of Belgium (Bosselaers et al. 2004). Therefore, the potential late Messinian occurrences from Beaumaris, despite being fragmentary, add useful evidence for interpreting the late Neogene evolution of sea turtle faunas. Specifically, the modernization of cheloniid assemblages globally from the middle Miocene to early Pliocene followed a pattern of stem and aberrant crown cheloniid extinctions in parallel with the emergence of extant cheloniid genera. Certainly, the geologically earliest crown cheloniid is probably Procolpochelys grandaeva from the early Miocene of the western North Atlantic (Zangerl & Turnbull 1955, Sugarman et al. 1993), whereas the

6 ERICH M. G. FITZGERALD AND LESLEY KOOL ALCHERINGA earliest known appearances of the living Chelonia, Lepidochelys and Caretta are from the mid late Miocene (ca 10.3 13.6 Ma) of the western North Atlantic (Dodd & Morgan 1992). The divergent extinct Syllomus survived into the early Pliocene of the western North Atlantic (Zug 2001), implying that the taxonomic composition of the modern sea turtle fauna was established within the last four million years. This report of Pacifichelys-like fossils from the late Miocene early Pliocene thus hints at a longer and more recent temporal overlap between stem- and crown-group cheloniids, including living genera. Acknowledgements The authors thank W. Joyce and J. Parham for advice and provision of materials, and K. Roberts and K. Smith for access to Museum Victoria s Reptiles and Amphibians Collection. J. Parham and M. Rabi provided helpful reviews. S. McLoughlin and B. Kear assisted with editorial comments. J. Velez-Juarbe supplied the photograph of LACM 103351 used in Fig. 3. Brian Crichton is thanked for his collection and donation of NMV P232865 to Museum Victoria. Astrid Werner is thanked for her skilful preparation of NMV P232865 and other significant fossils. References ABELE, C., GLOE, C.S., HOCKING, J.B., HOLDGATE, G., KENLEY, P.R., LAWRENCE, C.R., RIPPER, D., THRELFALL, W.F. & BOLGER, P.F., 1988. Tertiary. In Geology of Victoria. DOUGLAS, J.G. & FERGUSON, J.A., eds, Victorian Division, Geological Society of Australia, Melbourne, 251 350. BATSCH, A.J.G.C., 1788. Versuch einer Anleitung, zur Kenntniß und Geschichte der Thiere und Mineralien, für akademische Vorlesungen entworfen, und mit den nöthigsten Abbildungen versehen. Erster Theil. Allgemeine Geschichte der Natur; besondre der Säugthiere, Vögel, Amphibien und Fische. Akademische Buchhandlung, Jena, 528 pp. BAUR, G., 1893. Notes on the classification of the Cryptodira. American Naturalist 27, 672 674. BOSSELAERS, M., HERMAN, J., HOEDEMAKERS, K., LAMBERT, O., MARQUET, R. & WOUTERS, K., 2004. Geology and palaeontology of a temporary exposure of the late Miocene Deurne Sand Member in Antwerpen (N. Belgium). Geological Belgica 7,27 39. BUCHANAN, L.A., CONSOLI, C.P. & STILWELL, J.D., 2007. Early Paleocene marine vertebrates from the Wangaloa Formation, South Island, New Zealand. New Zealand Journal of Geology and Geophysics 50, 33 37. BUCKERIDGE, J.S., 1981. A marine turtle (Cheloniidae) from the lower Miocene of Port Waikato, New Zealand. New Zealand Journal of Geology and Geophysics 24, 435 437. CADENA, E., BOURQUE, J.R., RINCON, A.F., BLOCH, J.I., JARAMILLO, C.A. & MACFADDEN, B.J., 2012. New turtles (Chelonia) from the late Eocene through late Miocene of the Panama Canal Basin. Journal of Paleontology 86, 539 557. CHAPMAN, F., 1912. On the occurrence of Scaldicetus in Victoria. Records of the Geological Survey of Victoria 3, 236 238. CHAPMAN, F., 1914. On the succession and homotaxial relationships of the Australian Cainozoic system. Memoirs of the National Museum, Melbourne 5, 5 52. CHAPMAN, F., 1917. New or little-known Victorian fossils in the National Museum. Part XXI. Some Tertiary cetacean remains. Proceedings of the Royal Society of Victoria 30, 32 43. CHAPMAN, F.& PRITCHARD, G.B., 1907. Fossil fish remains from the Tertiaries of Australia. Part II. Proceedings of the Royal Society of Victoria 20, 59 75. COPE, E.D., 1868. On the origin of genera. Proceedings of the Academy of Natural Sciences of Philadelphia 20, 242 300. DICKINSON, J.A. & WALLACE, M.W., 2009. Phosphate-rich deposits associated with the Mio-Pliocene unconformity in south-east Australia. Sedimentology 56, 547 565. DICKINSON, J.A., WALLACE, M.W., HOLDGATE, G.R., GALLAGHER, S.J. & THOMAS, L., 2002. Origin and timing of the Miocene-Pliocene unconformity in southeast Australia. Journal of Sedimentary Research 72, 288 303. DODD, C.K. Jr & MORGAN, G.S., 1992. Fossil sea turtles from the early Pliocene Bone Valley Formation, central Florida. Journal of 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. FITZGERALD, E.M.G., 2012. Possible neobalaenid from the Miocene of Australia implies a long evolutionary history for the pygmy right whale Caperea marginata (Cetacea, Mysticeti). Journal of Vertebrate Paleontology 32, 976 980. FITZGERALD, E.M.G., PARK, T. & WORTHY, T.H., 2012. First giant bony-toothed bird (Pelagornithidae) from Australia. Journal of Vertebrate Paleontology 32, 971 974. FITZGERALD, E.M.G., VELEZ-JUARBE, J.& WELLS, R.T., 2013. Miocene sea cow (Sirenia) from Papua New Guinea sheds light on sirenian evolution in the Indo-Pacific. Journal of Vertebrate Paleontology 33, 956 963. FORDYCE, R.E., 1979. Records of two Paleogene turtles and notes on other Tertiary reptilian remains from New Zealand. New Zealand Journal of Geology and Geophysics 22, 737 741. 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 fossil turtles. Bulletin of the American Museum of Natural History 164, 65 376. GILL, E.D., 1957. The stratigraphical occurrence and palaeoecology of some Australian Tertiary marsupials. Memoirs of the National Museum of Victoria 21, 135 203. GRANT-MACKIE, J.A., HILL, J.& GILL, B.J., 2011. Two Eocene chelonioid turtles from Northland, New Zealand. New Zealand Journal of Geology and Geophysics 54, 181 194. HASEGAWA, Y., HIRAYAMA, R., KIMURA, T., TAKAKUWA, Y., NAKAJIMA, H. & KENKYUKAI, GUNMA KOSEIBUTSU, 2005. Skeletal restoration of a fossil sea turtle, Syllomus, from the middle Miocene Haratajino Formation, Tomioka Group, Gunma Prefecture, central Japan. Bulletin of Gunma Museum of Natural History 9, 29 64. (in Japanese). HILGEN, F.J., LOURENS, L.J., VAN DAM, J.A., BEU, A.G., BOYES, A.F., COOPER, R.A., KRIJGSMAN, W., OGG, J.G., PILLER, W.E. & WILSON, D.S., 2012. The Neogene Period. In The Geologic Time Scale 2012. GRADSTEIN, F.M., OGG, J.G., SCHMITZ, M.D. & OGG, G.M., eds, Elsevier, Amsterdam, 923 978. HIRAYAMA, R., 1995. Phylogenetic systematics of chelonioid sea turtles. The Island Arc 3, 270 284. [For 1994] JONES, M.E.H., WERNEBURG, I., CURTIS, N., PENROSE, R., O HIGGINS, P., FAGAN, M.J. & EVANS, S.E., 2012. The head and neck anatomy of sea turtles (Cryptodira: Chelonioidea) and skull shape in Testudines. PLoS ONE 7(11), e47852. doi:10.1371/journal. pone.0047852. KARL, H.-V. & TICHY, G., 2007. Maorichelys wiffeni n. gen. n. sp., a new sea turtle from the Eocene of New Zealand (Chelonii: Dermochelyidae). Studia Geologica Salmanticensia 43, 11 24. KEAR, B.P., 2003. Cretaceous marine reptiles of Australia: a review of taxonomy and distribution. Cretaceous Research 24, 277 303. KEAR, B.P. & LEE, M.S.Y., 2006. A primitive protostegid from Australia and early sea turtle evolution. Biology Letters 2,116 119. KEAR, B.P. & SIVERSON, M., 2010. First evidence of a Late Cretaceous sea turtle from Australia. Alcheringa 34, 265 272. KEMP, N.R., 1991. Chondrichthyans in the Cretaceous and Tertiary of Australia. In Vertebrate Palaeontology of Australasia. VICKERS-RICH,

ALCHERINGA CENOZOIC SEA TURTLES FROM AUSTRALIA 7 P., MONAGHAN, J.M., BAIRD, R.F.& RICH, T.H., eds, Pioneer Design Studio in cooperation with the Monash University Publications Committee, Melbourne, 497 568. KENLEY, P.R., 1967. Tertiary. In Geology of the Melbourne District. Victoria. Geological Survey of Victoria Bulletin 59, 31 46. KÖHLER, R., 1995. A new species of the fossil turtle Psephophorus (order Testudines) from the Eocene of the South Island. Journal of the Royal Society of New Zealand 25, 371 384. LAURENTI, J.N., 1768. Specimen medicum, exhibens synopsis Reptilium emendatam cum experimentis circa venena et antidota Reptilium Austriacorum. Trattnern, Vienna, 214 pp. LYNCH, S.C. & PARHAM, J.F., 2003. The first report of hard-shelled sea turtles (Cheloniidae sensu lato) from the Miocene of California, including a new species (Euclastes hutchisoni) with unusually plesiomorphic characters. PaleoBios 23, 21 35. MALLETT, C.W., 1977. Studies in Victorian Tertiary Foraminifera Neogene Planktonic Faunas. PhD dissertation, University of Melbourne, Melbourne, 646 pp. (unpublished) MÁRQUEZ, M.R., 1990. FAO Species Catalogue. Vol. 11: Sea Turtles of the World. An Annotated and Illustrated Catalogue of Sea Turtle Species Known to Date. Food and Agriculture Organization of the United Nations, Rome, 81 pp. MCGOWRAN, B., HOLDGATE, G.R., LI, Q. & GALLAGHER, S.J., 2004. Cenozoic stratigraphic succession in southeastern Australia. Australian Journal of Earth Sciences 51, 459 496. OPPEL, M., 1811. Die Ordnungen. Familien und Gattungen der Reptilien als Prodromeiner Naturgeschichte derselben. J. LINDAUER, München, 86 pp. PARHAM, J.F. & PYENSON, N.D., 2010. New sea turtle from the Miocene of Peru and the iterative evolution of feeding ecomorphologies since the Cretaceous. Journal of Paleontology 84, 231 247. PARK, T.& FITZGERALD, E.M.G., 2012a. A review of Australian fossil penguins (Aves: Sphenisciformes). Memoirs of Museum Victoria 69, 309 325. PARK, T. & FITZGERALD, E.M.G., 2012b. A late Miocene early Pliocene Mihirung bird (Aves: Dromornithidae) from Victoria, southeast Australia. Alcheringa 36, 419 422. PIPER, K.J., FITZGERALD, E.M.G. & RICH, T.H., 2006. Mesozoic to early Quaternary mammal faunas of Victoria, south-east Australia. Palaeontology 49, 1237 1262. SCHEYER, T.M. & SANDER, P.M., 2007. Shell bone histology indicates terrestrial palaeoecology of basal turtles. Proceedings of the Royal Society B: Biological Sciences 274, 1885 1893. SINGLETON, F.A., 1941. The Tertiary geology of Australia. Proceedings of the Royal Society of Victoria 53, 1 125. SUGARMAN, P.J., MILLER, K.G., OWENS, J.P. & FEIGENSON, M.D., 1993. Strontium-isotope and sequence stratigraphy of the Miocene Kirkwood Formation, southern New Jersey. Geological Society of America Bulletin 105, 423 436. VIS, C.W. DE., 1905. Fossil vertebrates from New Guinea. Annals of the Queensland Museum 6, 26 31. WALLACE, M.W., DICKINSON, J.A., MOORE, D.H. & SANDIFORD, M., 2005. Late Neogene strandlines of southern Victoria: a unique record of eustasy and tectonics in southeast Australia. Australian Journal of Earth Sciences 52, 277 295. WILKINSON, H.E., 1969. Description of an upper Miocene albatross from Beaumaris, Victoria, Australia, and a review of fossil Diomedeidae. Memoirs of the National Museum of Victoria 29, 41 51. WITZELL, W.N., 1983. FAO Fisheries Synopsis No. 137: Synopsis of Biological Data on the Hawksbill Turtle Eretmochelys imbricata (Linnaeus, 1766). Food and Agriculture Organization of the United Nations, Rome, 78. WYNEKEN, J., 2001. The anatomy of sea turtles. US Department of Commerce NOAA Technical Memorandum NMFS-SEFSC-470, 1 172. ZANGERL, R. & TURNBULL, W.D., 1955. Procolpochelys grandaeva (Leidy), an early carettine sea turtle. Fieldiana. Zoology 37, 345 382. ZANGERL, R., HENDRICKSON, L.P. & HENDRICKSON, J.R., 1988. A redescription of the Australian flatback sea turtle, Natator depressus. Bishop Museum Bulletin in Zoology 1, 1 69. ZUG, G.R., 2001. Turtles of the Lee Creek Mine (Pliocene: North Carolina). Smithsonian Contributions to Paleobiology 90, 203 218.