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Thematic Article Phylogenetic Systematics of Chelonioid Sea Turtles

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Thematic Article Phylogenetic Systematics of Chelonioid Sea Turtles The Island Arc (1994) 8, 270-284 Thematic Article Phylogenetic systematics of chelonioid sea turtles REN HIRAYAMA Faculty of Information, Teik:yo Heisei University of Technology, 2289, Uruido, Ichihara, Chiba 290-01, Japan Abstract Chelonioid sea turtles (Order Testudinata; Superfamily Chelonioidea) first appear in the early Cretaceous (Aptian-Albian). Their long fossil record is excellent compared to most marine tetrapods. Nonetheless, there has been no inclusive attempt to provide an analysis of cladistic relationships among this group. In this paper, the following chelonioid classification is proposed, based on 76 characters among 20 fossil and six living well- represented genera: (1) Family Cheloniidae (Coniacian to Recent); ·roxochelys, Ctenochelys, Osteopygis, Erquelinnesia, Allopleuron, Argillochelys, Puppigerus, Eochelone, Syllomus, Natator, Eretmochelys, Chelonia, Ca·retta, Lepidochelys (2) Family Protostegidae (Aptian to Maastrichtian?); 'Santana new protostegid', Rhino­ chelys, Notochelone, Desmatochelys, Chelosphargis, Protostega, Archelon (3) Family Dermochelyidae (Santonian? to Recent); Corsochelys, 'HMG new dermo­ chelyid', Eosphargis, Psephophorus, Dermochelys. Key words: Cheloniidae, Chelonioidea, classification, Dermochelyidae, osteology, Protostegidae, Reptilia, Testudinata. INTRODUCTION The chelonioid sea turtles have a long history from METHODS the early Cretaceous (Aptian- Albian; Hirayama 1993) as marine tetrapods. Their fossils are com­ During the course of this study the author exam­ mon in shallow water marine sediments. Although ined over 500 fossil and Recent chelonioid speci­ their morphological diversity in the past has been mens. A list of specimens examined is available well documented by many students (summarized in upon request. Zanger! 1980), the phylogenetic relationships of Although numerous fossil genera and species chelonioids based on detailed morphology has not have been allocated to the Chelonioidea (Kuhn been published, as reviewed by Gaffney and Maylan 1964), many of them were based on poor materials (1988). More recently, Hirayama (1992a) pub­ or poor descriptions. Thus, I have selected 26 gen­ lished the tentative results of a phylogenetic analy­ era, including 20 plesions, as terminal basic taxa sis at the generic level among the chelonioids; among chelonioids. Of them, two new unnamed ple­ however, the study used only humeral morphology. sions, the 'Santana protostegid' (TUTg1386) and After the IGC symposium (Hirayama 1992d), a the Japanese 'HMG dermochelyid' (Hirayama & beautifuJly preserved complete skeleton of the old­ Chitoku 1994) are included here because of their est known chelonioid from the Santana Formation importance to chelonioid phylogeny. of Brazil became available to the author and was A survey of 76 osteological characters was carefully prepared with acid (Hirayama 1993), and developed for these 26 genera. Character polarities t he previous conclusions about chelonioid relation­ were determined primarily using out-group com­ ships were necessarily changed after this new parisons. After hypotheses of polarity were formu­ discovery. In this paper the character analysis of lated, these decisions were summarized and ex­ chelonioids is summarized and conclusions are pressed in cladograms. My choice of the final made about the relationships and the systematics cladogram was made by an a posteriori weighting of this spectacular group of marine tetrapods. of characters. Phylogeny of chelonioid sea turtles 271 OBSERVATIONS AND RESULTS (13) Anterior vomer-palatine contact; aper­ tura narium interna entirely formed by vomer The derived characters used in analysis of the and palatines. phylogeny of chelonioids are as follows: (14} Primary palate involving vomer. Cranial and lower jaw elements (Figs 1-5) (15) Palatines medially meeting; reduction of (1) Cranial scute sulci present on dermal roof­ posterior portion of vomer. ing elements. (16} Foramen palatinum posterius reduced (2) Loss of nasal bone. and lost. (3) Loss of nasal-frontal contact; medial sep­ (17) Latellay open foramen palatinum poste­ aration of prefrontals. rius. ( 4) Prefrontal-postorbital contact; exclusion (18} Loss of processus pterygoideus externus. of frontal from orbital rim. (19) Pterygoid extending onto mandibular ar- (5) Parietal-squamosal contact. ticulating surface of quadrate. (6) Shallow superior temporal emargination. (20) Narrow middle portion of pterygoids. (7) Jugal-quadrate contact; exclusion of quad- (21) Large processus trochlearis oticum. ratojugal from lower cheek margin. (22) Short crista supraoccipitalis, not project­ (8) Jugal-squamosal contact; loss of post­ ing beyond parietal. orbital-quadratojugal contact. (23) Foramen posterius canalis carotici in­ (9) Medial process of jugal lost. terni between pterygoid and basisphenoid. (10} Loss of foramen praepalatinum. (24) Ventral surface of basisphenoid with (11) Secondary palate involving palatines. prominent V-shaped crest. (12) Secondary palate involving vomer in addi­ (25) Large basisphenoid, lying beneath the sig­ tion to palatines. nificant portion of pterygoids. (a) (e) (f) (g) (h) Fig. 1 Skulls of chelonioids, left lateral view; (a) Toxochelys latiremis (largely based on AMNH 51 18, condyle-premaxilla length 123 mm, with additions from YPM3602, modified from Gaffney 1979; Senonian of USA); (b) A/lopleuron hoffmanni (NHMM 9017, 258 mm; Upper Maastrichtian of Holland); (c) Osteopygis emarginatus (NJSM 11872, 122 mm; Upper Maastrichtian? of USA); (d) Chelonia mydas (after easier 1968; Recent). (e) 'Santana protostegid' (TUTg 1386, 37 mm; Upper Aptian or Lower Albian of Brazil); (f) Rhinochelys pu/chriceps (largely based on BM(NH) R2226. 62 mm, reversed , with additions from BM(NH) 46371 and 46371a; Upper Albian of England); (g) Desmatoche/ys lowi (primarily based on KUVP 1200, 191 mm as estimated; Lower Turonian of USA; with additions from TUTg 262; Middle Turonian of Japan); (h) Protostega gigas(based on FMNH PR65, PR170, PR197 and P27314. about350 mm; lower Campanian of USA); (i) Dermochelys coriacea (after Gaffney 1979, reversed; Recent). 272 R. Hirayama (a) (f) (g) (h) •• Fig. 2 Skulls of chelonioids, dorsal view. The dotted line shows lhe scule sulci on lhe dermal roofing elements; (a) Toxoche/ys /atiremis (largely based on AMNH 5118, with additions from YPM 3602, modified from Gaffney 1979); (b) Ctenochelys stenoporus (largely based on USNM 11639, 128 mm; Senonian of USA) . (c) Allopleuron hoffmanni(NHMM 9017); (d) Osteopygis emarginatus (NJSM 11872); (e) Chelonia mydas (after easier, 1966); (f) 'Santana prolostegid' (TUTg 1386); (g) Rhinochelys pulchriceps (largely based on BM (NH) R2226); (h) Oesmatochelys /owi(KUVP1200); (i) Protostega gigas (based on FMNH PR65, PR170, PR197 & P27314); (j) Dermochelys coriacea (after Gaffney 1979). (26) High dorsum sellae, not concealing poste­ (39) Double cervical central articulation be­ rior portion of sella turcica. tween 7th and 8th centra. (27) Rod-like ossification of rostrum basisphen­ (40) Platycoelous cervical central articulation oidale. between 6th and 7th centra. (28) Reduced ossification of rostrum basisphen­ ( 41) First thoracic vertebra with anterior artic­ oidale. ulation facing anteroventrally. (29) Foramen caroticum laterale larger than Appendicular skeleton (Figs 6- 10) foramen anterius canalis carotici interni. (30)Foramen caroticum laterale confluent with (42) Elongate 3rd to 5th digits of manus and ( canalis cavernosum. pes, without movable articulation; carpal and tarsal (31) Broad and flat lower triturating surface, elements flattened, with restricted joint surfaces; with enlarged, deep lateral concavity of dentary humerus as long as femur, with nearly straight where adductor mandibular muscle attaches. shaft, and lateral process is located distal to caput (32) High lingual and symphyseal ridge of den- humerus. tary, visible from lateral view. ( 43) Movable articulations of first and second (33) Denticulated labial ridge of dentary. digits lost. (34) Denticulated lingual ridge of the dentary. ( 44) Coracoid longer than humerus, often with (36) Surangular process extending anteriorly a high dorso-median keel for the M supracora­ onto dentary. coideus. (37) Loss of coronoid and articular bones. ( 45) Elongate coracoid thickened, showing Vertebrae column-like appearance. (38) Longer cervical vertebrae with much ( 46) Scapular angle, formed by scapular prong wider central articulations. and acromion, larger than 110 o. Phylogeny of chelonioid sea turtles 273 (a) ,. (b) .. Fig. 3 Skulls of chelonioids, ventra l view; (a) Toxochelys /aliremis (largely based on AMNH 5118, with additions from AMNH 1024, modified from Gaffney 1979); (b) Ctenoche/ys stenoporus (largely based on USNM 11639; Senonian of USA); (c) Allopleuron hoffmanni (based on NHMM 9017 and IRScNB, an uncatalogued specimen); (d) Osteopygis emarginatus (NJSM 11872); (e) Chelonia mydas (alter easier 1966); (f) 'Santana protostegid' (TUTg 1386); (g) Rhinochelys pu/chriceps (largely based on BM(NH) 1606, 36 mm, with additions from BM(NH) R2224, R2276, 46371 and R8339; Middle to Upper Albian of England); (h) Desmatochelys lowi (KUVP 1200); (i) Protostega gigas (based on FMNH PR 170, PR65. P27385 and CM 26593). (j) Dermochelys coriacea (after Gaffney 1979). (4 7) 'Cheloniid' humerus; the V-shaped struc­ (59) Tibia pit forM pubotibialis and M flexor ture of lateral process. tibialis internus. (48) 'Protostegid' humerus; lateral process of Shell (Figs 8-10) humerus restricted onto the anterior portion of (60)
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