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18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

John M. Legler & Arthur Georges

1 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

2 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

The earliest known , Proganochelys, had unspecialised cervical vertebrae (Gaffney 1990) and presumably were unable to retract the head beneath the shell. This innovation was later achieved in two ways. One lineage, the suborder , evolved a mechanism whereby the neck flexes in the vertical plane when retracted beneath the shell. In these forms, the head is still pointed in a forward direction when retracted. The suborder comprises nine extant families, and most of the living species. The majority occur in the temperate region of the northern hemisphere, and in the tropics. The only Australian representatives are the marine turtles, families and , and the freshwater pig-nosed , Carettochelys insculpta. In the second lineage, the suborder , the neck flexes in the horizontal plane so that the head points towards the left or right when retracted. The suborder comprises two families. The includes five genera Erymnochelys, Pelomedusa, Peltocephalus, and . Though presently restricted to the South American and African continents, they were once widely distributed across the northern hemisphere. The includes five genera , , , Pseudemydura and currently found in , , the island of Roti (Lesser Sunda Islands, ), and South America. There are reports of fossil chelid turtles from Europe, North Africa and India, but the diagnosis of these fossils as chelids is either incorrect or has been brought into serious question (Williams 1953, 1954b). There is no well accepted evidence that fossil chelids occur outside the range of extant taxa (Pritchard 1979b; Pritchard & Trebbau 1984). As such, chelid turtles are the only group with clear Gondwanan origins.

DISTRIBUTION Freshwater turtles are cosmopolitan with breeding populations occurring in suitable habitats in temperate and tropical regions of major continents, large islands, and some oceanic islands. There are epicentres of diversity in the eastern United States of America and southern Asia. The five extant genera of marine turtles have worldwide distributions in tropical and temperate waters, except for Natator depressus which is endemic to the Australian region.

Figure 18.1 Worldwide distribution of the freshwater chelonians. Numbers of genera are indicated for each of eight regions: North, Central and South America, sub- saharan Africa, with Madagascar as a subset, Europe and North Africa, India, South- East Asia, and Australia. [[W. Mumford]]

3 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

Cryptodires are now the only turtles of the Northern Hemisphere, but they occur also in the Southern Hemisphere. Pleurodires now occur only in the Southern Hemisphere. There is a substantial fossil record for pelomedusid turtles in the Northern Hemisphere, but there is no evidence that chelids ever occurred there. Cryptodires exhibit extremes in geographic range. Painted turtles (Chrysemys picta) and snapping turtles ( serpentina) of Canada have the most northerly distribution (about 52°). These turtles probably occur at or near the northern limits for oviparous . This distribution is equalled only by a few oviparous (Opheodrys, Pituophis and Storeria) (Conant 1975). chiliensis, a terrestrial has the most southerly distribution, and occurs at 43°S in Argentina. The emyidid, scripta, ranges from 42°N to 36°S, a total of 78° of latitude, possibly the widest breeding range of any freshwater turtle (Legler 1990). Sea turtles may range widely to the north and south of their breeding ranges. Dermochelys coriacea has the largest latitudinal range of any known turtle, from the north coast of Iceland to 47°S, a total latitudinal range of 113°. Most chelid turtles occur in temperate or tropical latitudes. There is a varied of pelomedusids and testudinid at or near the southern tip of Africa (ca. 35°S). geoffroanus is the southern-most South American chelid (30°S). Australian chelids occur to the latitudinal extremes of the continent, but have epicentres of diversity between 23° and 25°S in eastern (see Chapter 21).

Limiting Factors All extant Australian chelids are aquatic. Their geographic distribution is generally limited by the presence of permanent water or its seasonal yet predictable occurrence in natural drainage systems (Anon. 1967; Leeper 1970) as discussed in Chapter 21. A notable exception is Chelodina steindachneri, which occurs in water courses that fill unpredictably and which may be dry for well over one year. However, in well-watered temperate regions, temperature is probably a limiting factor in its effect on the incubation of eggs and the digging of nests. Turtles are oviparous and must rely on a warm season long enough for nesting and at least partial embryonic development. Overwintering of eggs and hatchlings is known (Carr 1952; Ernst & Barbour 1972; Ewert 1979). Turtles seem to prefer microhabitats in which they can voluntarily move into different temperatures. Temperatures of approximately 25°C are probably close to optimum, but there is evidence to suggest that temperate aquatic turtles (including chelids) can move about comfortably, eat, mate, and nest at body temperatures of 15°C or lower (see Chapter 16). Southern Hemisphere turtles in general, and chelids in particular, never experience the vicissitudes of temperature of a boreal winter. Water seems to be a far greater limiting factor in the Southern Hemisphere. Substrata which do not permit the digging of nests may limit distribution in various areas of otherwise favourable habitat. In streams draining to the Murray- Darling system from the New England Tableland, there is usually a distinct fauna above falls or cascades and another below. The intervening zone contains few or no turtles and, although the water in this zone is fast, it is probably the lack of nesting sites that excludes the turtles.

PHYLOGENETIC ORIGIN The oldest known turtle, Proganochelys quenstedti from the Upper of Bavaria, lived approximately 200 million years ago (Gaffney 1990). The species was first described by Jaekel (1914, 1918). More recently, Gaffney (1990) prepared a monograph on a series of six specimens, including some that were

4 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

Figure 18.2 Phylogenetic tree showing the relationships of living and fossil turtles. (After Gafney & Meylan 1988) [[D. Wahl]] virtually complete skeletons, such that this species is now one of the best known skeletally. Gaffney’s restorations and illustrations of P. quenstedti suggest an with habits, habitat, body form, and general mass similar to the large North American Snapping Turtle, Chelydra serpentina. Proganochelys has some primitive characters that are absent in living chelonians, but is otherwise quite clearly a bona fide turtle. By Triassic times, Proganochelys already possessed most of the adaptive modifications that are diagnostic of turtles. In particular, the shell had evolved fully and surrounded the limb girdles. Unfortunately, P. q ue n st ed t i provides little or no information about the evolutionary steps leading from the generalised tetrapod body plan to the unique turtle body plan.

5 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

The search for turtle ancestry relies heavily on osteological comparisons (usually of the skull) between bona fide turtles and groups of reptiles at or near the base of the amniote stock, usually referred to as Stem Reptiles. The two best supported hypotheses of chelonian origin identify different fossil groups of reptiles as the sister group of the Chelonia. Gaffney & Meylan (1988) used the Captorhinidae, a group of that would necessitate an origin for turtles as early as the early or even late . This hypothetical ancestor would require a gap of approximately 100 million years between the origin of turtles and the oldest material of P. q ue n st ed t i Gaffney & Meylan (1988) built their hypothesis on four sets of shared, derived characters (synapomorphies) of the cranium, not all of which are unique to the chelonians and the captorhinids. Reisz & Laurin (1991) invoked the procolophonids, a group of small tetrapod reptiles (or ‘parareptiles') from the Permian of South Africa, as the ancestors of turtles. They supported this hypothesis on one postcranial and nine cranial synapomorphies, all of which are unique to the two sister groups. The hypothesis implies an origin for turtles as late as the late Permian, necessitating a much shorter gap in evolutionary history. Although these hypotheses are of great interest and importance, neither really addresses the matter of the transition from a generalised tetrapod body plan to that of a turtle. Nor does either have an impact on an understanding of chelonian since the Triassic, or of relationships between existing groups within the order Chelonia. To date, Deraniyagala (1939) has been the only author to suggest and illustrate a body plan (a whole hypothetical organism) that constitutes a stage intermediate between a generalised tetrapod and a pre-chelonian. His ‘saurotestudinate’ ancestor from the Permian or Carboniferous, although somewhat fanciful, combines many of the characters that most current turtle biologists would predict in an ancestral group (see Chapter 16).

AFFINITIES WITHIN THE CHELIDAE Morphology has played an important part and, until recently, the only part in the classification of Australian chelids. However, Burbidge, Kirsch & Main (1974) utilised a serological approach to the classification of the Chelidae, and Georges & Adams (1992) recently presented an analysis of relationships within the Australian chelids, based on biochemical data. There is a general congruence between their findings and those of Legler (1981), based on a phenetic analysis of morphological characters (see Chapter 21). Evidence from both studies, together with that from the recent cladistic analysis of Gaffney & Meylan (1988) strongly support the monophyly of the family Chelidae, and the common ancestry of South American and Australian chelids. The simplest biogeographic scenario to explain monophyly involves the presence of chelids on the large southern continent, . The land mass split into South America and Australia-Antarctica during the Palaeocene or , 50 to 65 million years ago. Following the separation of Australia and Antarctica, about 45 to 49 million years ago, the Australian plate was completely isolated as it drifted 15° northward, until it collided with the Asian plate about 10 to 15 million years ago (Runcorn 1962; Irving 1964; Dietz & Holden 1970; Doutch 1972; Raven & Axelrod 1972; Tarling & Tarling 1975; Tyler 1979) . The origin of the longnecked freshwater turtles, Chelodina and from a common longnecked ancestor is well supported. Hydromedusa is known from the early Eocene of South America (Wood & Moody 1976), probably before the isolation of that continent from Gondwana. The longnecked chelids are, therefore, more likely to be monophyletic (Gaffney & Meylan 1988) than

6 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

Figure 18.3 Cladogram of relationships within the Chelidae. (After Georges & Adams 1992) [[D. Wahl]] convergent (Pritchard 1984). It seems logical that the longnecked condition is derived; Hydromedusa is, therefore, regarded as already too specialised to have been ancestral to both longnecked and shortnecked groups (see also Chapter 21). The evidence for shortneck lineages is not so clear and is not supported by fossils. Cann & Legler (1993) placed Australian shortnecked chelids in three distinct groups. Pseudemydura, which is not closely related to any Australian chelid and might be related to Platemys of South America (see also Chapter 21); Emydura; and the ‘Elseya complex', containing the Elseya dentata group, the Elseya latisternum group, Rheodytes, and shortneck alpha (see Chapter 21). Although Emydura and the Elseya complex almost certainly have a common ancestry, the present authors do not regard the relationship between these two groups to be as close as do some other authors (for example, Gaffney & Meylan 1988; MacDowell 1983), who place them all in the Emydura.

BIOGEOGRAPHY The modern Australian chelonian fauna can be considered in three categories. The ancient autochthonous fauna includes the chelids, which constitute the dominant continental turtle fauna of Australia. They are the only modern family of turtles common to Australia and South America. An Asian element of Tertiary origin has entered Australia via the Indo- Australian Archipelago since the Australian and Asiatic plates collided 10 to 15 million years ago. The Australian reptile fauna is predominantly oriental (Tyler 1979), with a large component derived after Australia collided with the Asiatic plate. Carettochelys insculpta is probably a part of this oriental group and occurs

7 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA only in a few river systems in northern Australia and in southern New Guinea. The cryptodiran genus belongs in this group if southern New Guinea is considered as a part of the Australian continental plate. The cosmopolitan element includes the marine turtles of the families Cheloniidae and Dermochelyidae. These occur worldwide and are subject to few or no distributional barriers, with the exception of Natator depressus, known only from Australian and New Guinean coastal waters.

FOSSIL RECORD The Australian fossil record has yielded sparse but identifiable specimens of the Chelidae and Carettochelyidae, two taxa of Early marine turtles included in the Chelonioidea and specimens of the extinct cryptodiran families Meiolaniidae and (Gaffney 1981). The chelonioid fossils are some 110 million years old. Fossils presently not clearly assigned to any higher taxon, such as Chelycarapookus of the Lower Cretaceous of (Chapman 1919), provide tantalising examples of what may yet be discovered in the fossil record. Warren (1969) was unable to determine the suborder, although the species was assigned recently to a new family, the Chelycarapookidae.

Meiolaniidae Meiolaniids are considered to be the only real Australian novelty provided by the fossil record. They occur from the to the Pleistocene on , in the Pleistocene of Lord Howe Island and New Caledonia, and in the Cretaceous and Eocene of South America (Gaffney 1981). The best known fossil specimens are of Meiolania platyceps of Lord Howe Island. They were large terrestrial turtles with cranial horns and frills, cervical ribs, a bony club at the end of their and numerous epidermal ossicles (Gaffney 1983, 1985a). Their bizarre appearance led to much confusion when early palaeontologists endeavoured to identify the fossil remains. Their ecology is unknown, but it is thought likely that they had habits similar to the testudinids of the Galapagos and Aldabra Islands (Gaffney 1985b).

Trionychidae The trionychids are soft-shelled turtles which occur today in Africa, Asia and North America but not in South America or Australia. Pelochelys bibroni of New Guinea probably reached the Australasian region from Asia (Darlington 1957).

Figure 18.4 Fossil skeleton of Meiolania platyceps. [[Photo ©]] 8 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA

Figure 18.5 Fossil plastron and carapace of Elseya. [[Photo by J. Cann]]

Fossil material from Australian deposits is fragmentary, but has been diagnosed as Trionychidae on the basis of surface texture pattern and a natural distal edge to the costals, rather than a suture, suggesting that peripheral bones were absent (Gaffney & Batholomai 1979). Trionychids were widespread through Queensland in the late Cainozoic and possibly as early as the Miocene.

Carettochelyidae Carettochelys insculpta is found today in the southern flowing rivers of New Guinea and the larger rivers of the in Australia (Cogger 1992). It is the sole surviving species of a family that was widespread in the late Tertiary.The genus is represented in the Australasian fossil record by a specimen from the Miocene of New Guinea (Glaessner 1942). A fossil from was misidentified as Carettochelys by Goster & McNicoll (1978) (Gaffney 1981).

Chelonioidea The oldest turtles from Australia with definitive stratigraphic data are the marine chelonioids Cratochelone and Notochelone from the Toolebuc Limestones of Julia Creek (Gaffney 1981, 1991).

Chelidae The Chelidae dominate the extant freshwater turtle fauna of Australia, and are also the most common and widespread fossil turtles. Chelids are restricted in distribution to Australasia and South America, both as living and fossil forms. They range in age from Miocene to Recent in Australian deposits. Chelids are distinguished from other turtles by the presence of a pelvis that is fused to the carapace and plastron, but in the absence of fossil evidence of this character, Australian material often is assigned to the family if neural bones are absent (Gaffney 1981). Fossil specimens have been assigned to the genera Chelodina, Emydura, Elseya and Pseudemydura on the basis of diagnostic scute arrangements of the carapace and plastron (Fig. 18.5; Gaffney 1977, 1981; Gaffney, Archer & White 1989). New insight into the evolution of the chelid fauna awaits discovery. Four of the six extant genera of Australian freshwater turtles are monotypic (Pseudemydura, Rheodytes, shortnecked alpha, and Carettochelys), represented by species that are distinct both morphologically (Ramsay 1886; Siebenrock 1901; Legler & Cann 1980; Cann & Legler 1993) and biochemically (Georges & Adams 1992). Each of these genera arises from lineages of considerable antiquity, as Chelodina and Pseudemydura may have their closest living relatives among South American forms (Gaffney 1977; Legler 1981). Carettochelys insculpta represents an independent lineage of some 40 million

9 18. BIOGEOGRAPHY AND PHYLOGENY OF THE CHELONIA years standing (Chen, Mao & Ling 1980). Although today Australia is the driest continent after Antarctica (see Chapter 21), this was not always so. The extant freshwater turtle fauna probably should be considered a relict of a much more diverse fauna of wetter times. This assertion is not well supported by the fossil record, in part because of little attention having been paid to the turtle elements of Australian fossil collections until recently and, in part, because the morphology of extant forms is poorly known in detail. As Gaffney (1981) pointed out, it is not possible to deal realistically with fossil chelid material at the species level because of the paucity of skeletal material and descriptive work for extant forms. The new taxa described by de Vis around the turn of the century can be diagnosed to family only, and no extinct species can be substantiated at present (Gaffney 1981).

LITERATURE CITED

Anon. (1967). Atlas of Australian Resources. 2nd series. Surface Water Resources. Department of Natural Development : Canberra 26 pp. Cann, J. & Legler, J.M. (1993). The Mary River Tortoise: a new genus and species of the short-necked chelid from Queensland, Australia (Testudines: Pleurodira). Records of the Australian Museum (in press) Carr, A.R. (1952). Handbook of Turtles. The turtles of the United States, Canada and Baja California. Comstock : Ithaca 542 pp. Chapman, F. (1919). New or little known Victorian fossils in the National Museum. Part XXIV. On a fossil tortoise in ironstone from Carrapook, near Casterton. Proceedings of the Royal Society of Victoria 32: 31-32 Chen, B.-Y., Mao, S.-H. & Ling, Y.-H. (1980). Evolutionary relationships among turtles suggested by immunological cross-reactivity of albumins. Comparative Biochemistry and Physiology 66B: 421-425 Cogger, H.G. (1992). Reptiles and of Australia. 5th edition. Reed Books : Chatswood 775 pp. Conant, R. (1975). A field guide to reptiles and amphibians of eastern and central North America. 2nd edition. Houghton Mifflin Co. : Boston 429 pp. Darlington, P.J. (1957). Zoogeography: The Geographical Distribution of . John Wiley & Sons : New York 675 pp. Deraniyagala, P.E.P. (1939). The tetrapod reptiles of Ceylon. Vol. 1. Testudinates and Crocodilians. Colombo Museum : London 412 pp. Dietz, R.S. & Holden, J.C. (1970). The breakup of Pangaea. Scientific American 223: 30-41 Doutch, H.F. (1972). The paleography of northern Australia and New Guinea and its relevance to the Torres Strait area. Pp. 1-10 in Walker, D. (ed.) Bridge and Barrier: The Natural and Cultural History of Torres Strait. Publication BG/3. Research School of Pacific Studies, Department of Biogeography and Geomorphology, Australian National University : Canberra Ernst, C.H. & Barbour, R.W. (1972). Turtles of the United States. University Press : Lexington 347 pp. Ewert, M.A. (1979). The embryo and its egg: development and natural history. Pp. 333-413 in Harless, M. & Morlock, H. (eds) Turtles: Perspectives and Research. John Wiley & Sons : New York Gaffney, E.S. (1977). The side-necked turtle family Chelidae: a theory of relationships using shared derived characters. American Museum Novitates 2620: 1-28

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Gaffney, E.S. (1981). A review of the fossil of turtles of Australia. American Museum Novitates 2720: 1-38 Gaffney, E.S. (1983). The cranial morphology of the extinct horned turtle, Meiolania platyceps , from the Pleistocene of Lord Howe Island. Bulletin of the American Museum of Natural History 175: 361-479 Gaffney, E.S. (1985a). The cervical and caudal vertebrae of the cryptodiran turtle, Meiolania platyceps , from the Pleistocene of Lord Howe Island, Australia. American Museum Novitates 2805: 1-29 Gaffney, E.S. (1985b). Meiolania platyceps Owen, 1886. The Lord Howe Island Horned Turtle. Pp. 132-136 in Rich, P.V. & van Tets, G.F. (eds) Kadimakara. Extinct Vertebrates of Australia. Pioneer Design Studio : Lilydale Gaffney, E.S. (1990). The comparative osteology of the Triassic turtle Proganochelys. Bulletin of the American Museum of Natural History No. 194: 1-263 Gaffney, E.S. (1991). The fossil turtles of Australia. Pp. 704-720 in Vickers- Rich, P., Monaghan, R.F. & Rich, T.H. (eds) Vertebrate Palaeontology of Australasia. Pioneer Design Studio, for Monash University Publications : Lilydale, Victoria Gaffney, E.S. & Bartholomai, A. (1979). Fossil trionychids of Australia. Journal of Paleontology 53: 1354-1360 Gaffney, E.S. & Meylan, P.A. (1988). A phylogeny of turtles. Pp. 157-219 in Brenton, M.J. (ed.) The Phylogeny and Classification of the Tetrapods. Vol. 1 Amphibians, Reptiles, Birds. Systematic Association Special Volume 35A. Clarendon Press : Oxford Gaffney, E.S., Archer, M. & White, A. (1989). Chelid turtles from the Miocene freshwater limestones of Riversleigh Station. American Museum Novitates 2959: 1-10 Georges, A. & Adams, M. (1992). A phylogeny for Australian chelid turtles based on allozyme electrophoresis. Australian Journal of Zoology 40: 453-476 Glaessner, M.F. (1942). The occurrence of the New Guinea turtle ( Carettochelys) in the Miocene of Papua. Records of the Australian Museum 21: 106-109 Gorter, J.D. & Nicoll, R.S. (1978). Reptilian fossils from Windjana Gorge, Western Australia. Journal of the Royal Society of Western Australia 60: 97-104 Irving, E. (1964). Paleomagnetism. Wiley : New York Jaekel, O. (1914). Ueber die Wirbeltierfunde in der oberen Trais von Halberstadt. Palaeontology Zoology 1: 155-215 Jaekel, O. (1918). Die Wirbeltierfunde aus dem Keuper von Halberstadt. Serie II. Testudinata. Tiel 1. Stegochelys dux , n.g., n.sp. Palaeontology Zoology 2: 88-214 Leeper, G.W. (ed.) (1970). The Australian Environment. 4th edition. 163 pp. CSIRO : Melbourne Legler, J.M. (1981). The , distribution, and ecology of Australian freshwater turtles (Testudines: Pleurodira: Chelidae). National Geographic Society Research Report 13: 391-404 Legler, J.M. (1990). The genus Pseudemys in Mesoamerica: taxonomy, distribution, and origins. Pp. 82-105 in Gibbons, J.W. (ed.) Life History and Ecology of the Slider Turtle. Smithsonian Institution Press : Washington D.C. Legler, J.M. & Cann, J. (1980). A new genus and species of chelid turtle from Queensland, Australia. Contributions to Science of the Natural History Museum, Los Angeles County 324: 1-18

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McDowell, S.B. (1983). The genus Emydura (Testudines: Chelidae) in New Guinea with notes on the penial morphology of Pleurodira. Pp. 169-189 in Rhodin, A.G.J. & Miyata, K. (eds) Advances in Herpetology and Evolutionary Biology. Museum of Comparative Zoology : Cambridge Pritchard, P.C.H. (1979b). Taxonomy, evolution, and zoogeography. Pp. 1-42 in Harless, M. & Morlock H. (eds) Turtles. Perspectives and Research. John Wiley & Sons : New York Pritchard, P.C.H. (1984). Piscivory in turtle and evolution of the long-necked Chelidae. Pp. 87-110 in Ferguson, M.W.J. (ed.) The Structure, Development and Evolution of Reptiles. Symposia of the Zoological Society of London No 52. Academic Press : London Pritchard, P.C.H. & Trebbau, P. (1984). The Turtles of Venezuela. Society for the Study of Amphibians and Reptiles : Oxford, Ohio 403 pp. Ramsay, E.P. (1886). On a new genus and species of freshwater tortoise from River, New Guinea. Proceedings of the Linnean Society of (2) 1: 158-162 Raven, P.H. & Axelrod, D.I. (1972). and Australasian paleobiogeography. Science 176: 1379-1386 Reisz, R.R. & Laurin, M. (1991). Owenetta and the origin of turtles. 349: 324-326 Runcorn, S.K. (ed.) (1962). . Academic Press : New York 338 pp. Siebenrock, F. (1901). Beschreibung einer neuen Schildkrotengattung aus der Familie Chelydidae von Australien: Pseudemydura. Anzeiger der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissen schaftliche Classe, Wein 38: 248-250 Tarling, D. & Tarling, M. (1975). Continental Drift. Revised Edition Books. Bill & Sons, Ltd 142 pp. Tyler, M.J. (1979). Herpetofaunal relationships of South America with Australia, Pp. 73-106 in W.E. Duellman (ed.) The South American Herpetofauna: its Origin, Evolution and Dispersal. Museum of Natural History, University of Kansas Monograph No. 7 Warren, J.W. (1969). A fossil chelonian of probable lower Cretaceous age from Victoria. Memoirs of the National Museum, Victoria 29: 23-28 Williams, E.E. (1953). Fossils and the distribution of Chelyid turtles. 1. Hydraspis leithii (Carter) in the Eocene of India is a Pelomedusid. Breviora 1953: 1-8 Williams, E.E. (1954). Fossils and the distribution of chelyid turtles. 2. Breviora 1954: 1-6 Wood, R.C. & Moody, R.J.J. (1976). Unique arrangement of carapace bones in the South American chelid turtle Hydromedusa maximiliani (Mikan). Zoological Journal of the Linnean Society (London) 59: 69-78

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