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The Phylogeny of Cheloniid Sea Turtles Revisited. James F. Parham And

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The Phylogeny of Cheloniid Sea Turtles Revisited. James F. Parham And o, ee7 by ch",J;Xl?:i::1,';;lii'll,:''"- The Phylogeny of Cheloniid Sea Turtles Revisited Jauns F. Pnnnlur2 AND Dlvrn E. Fmrovsxvr tDepartmentof Geologt, Universitl,of Rhode Island, Kingston, Rhode Island0288l USA; 2Present Address: Department of Integrative Biolog1,, Universi4' of Califumia, Berkeley, Califurnia 94720 USA I F ax : 5 1 0 -642 - I 822 ; E-mail : P arham@ s oc rate s.be rke let'.e dn ] Ansrnacr. - Thirteen chelonioid taxa are examined cladistically using 24 cranial and post-cranial osteological characters. Shell characters are found to be useful in elucidating phylogenetic relation- ships within the Cheloniidae. The results of this study support a monophyletic Osteopyginae. The mutual affinities of the Eocene stem cheloniines are not resolved. Syllomus is the sister taxon to all the Cheloniinae (not just to Natator) and, Eretmochelys is the sister taxon to the Carettini. Kny Wonos.-Reptilia; Testudines; Cheloniidae; Toxochelys;CtenochelyslOsteopygis;Erquelinnesia; Argillochelysi Eochelone; Puppigerus; Syllomus; Natatorl Chelonia; Eretmochelysl Caretta; Lepidochely s; systematics ; paleontology; phylogeny Cryptodiran turtles have invaded the marine realm Eretmochelys, Caretta, Lepidochelys) and numel ous extinct twice: once in the Jurassic and once in the early Cretaceous taxa (see Pritchard and Trebbau, 1984). Excluding the prob- (Gaffney andMeylan, 1988;Hirayaffia, 1994). By Campanian lematical extinct generaToxochelv s and Ctenochebs (whose time, the Cretaceous radiation (Chelonioidea) had already affinities are discussed below), the Cheloniidae can be diversified into three major clades: Cheloniidae, subdivided into three, not necessarily monophyletic, groups: Dermochelyidae, and Protostegidae. The latter two families 1) Osteopygin ae (Zangerl, 1953) ;2)Eocene stem cheloniines form the monophyletic Dermochelyoidea (Gaffney and ("Eochelyinae" of Moody, 1968); and 3) Cheloniinae (de- Meylan, 1988), the sister-taxon to Cheloniidae. Pelagic fined here as the crown-group of the Cheloniidae, i.e., those adaptations such as the loss of moveable articulations in the taxa that share the most recent common ancestor of the manus to form a paddle, straightening of the humerus, and extant cheloniids; see de Quetroz and Gauthier, 1990). reduction of the plastral bones were all apparently indepen- Toxochelys and Ctenochelvs (Late Cretaceous) are dently acquired in dermochelyids, protostegids, and among the earliest chelonioids represented by ample cranial cheloniids (Hirayaffi&, 1994). Of the three clades, Cheloniidae and post-cranial material. They were considered to be mem- seems to have retained the primitive condition the longest; bers of the Toxochelyidae (Zangerl, 1953) until Fastovsky even Recent cheloniids (Caretta, Chelonia, Eretmochelys, ( 1985) demonstrated that Ctenochelys (referred to as FMNH Lepiclochelys,, Ncrtcrtor) have not developed the level of PR444; Gaffney and Meylan [ 1988] have since referred this pelagic specialization (e.g., broad flat humeri, extremely specimen to Ctenochelys) is more closely related to Recent reduced shells, greatly elongated forelimbs) exhibited by cheloniids than to Toxocltelys. As a result, the family dernrochelyoids (epitomized in Recent Dennochelys). Toxochelyidae (sensu Zangerl,, 1953) was rendered Classification within the Cheloniidae has traditionally paraphyletic. been based on the abundant shell material (Zangerl, 1953, These findings have been incorporated into two hypoth- 1958). While the appendicular skeleton was given some eses. Gaffney and Meylan ( 1988) suggested that Ctenochelys consideration, cranial material was essentially ignored. More should be considered a sister taxon to all chelonioids except recently, de scriptions of new and neglected cranial material Toxochelys (Fig. I a). Hirayama (1994), however, argued (Moody, 1974; Gaffney, 1979; Foster, 1980; Fastovsky, that Toxochelys and Ctenochelys should be considered as I 985) combined with cladistic methodology have resulted in members of an expanded Cheloniidae (Fig. I b). The latter hypotheses based almost entirely on cranial characters, hypothesis is based on an analysis comprised of more taxa particularly from the basicranium (Fastovsky, 1985; Gaffney and more characters and is adopted here. However, we have and Meylan, 1988). Hirayama (1994) was the first to attempt included neither Dermochelyidae nor Protostegidae in our data to treat cranial and post-cranial characters equally. set and therefore cannot comment further on this problem. This study pools the data of previous workers who have We use a stem-based definition (de QueirozandGauthier, studied the evolutionary relationships of cheloniids and 1990) of Cheloniidae as those turtles that share a more recent expands them with additional shell characters. Combination common ancestor with extant marine turtles (exclusive of of all the available data should produce the most defendable Dermochelvs) than with Dermochelys or Protostegct This hypothesis of cheloniid phylogeny to date. definition agrees with Hirayama (1994) and differs only slightly from Gaffney and Meylan ( 1988) who recognized SYSTEMATICS an Osteopygidae (Osteopyginae here and rn Zangerl, 1 953,, l9l l, Fastovsky, 1985). The three major groupings of Chel,oniidae. This is a well-known family of marine Cheloniidae (Osteopyginae, stem cheloniines, Cheloniinae) turtles comprised -of five living genera (Natator,, Chelonia, are discussed below in detail. Pnnunla AND FasrovsKy Cheloniid Phylogeny - 549 a.) (6 H Perhaps the most interesting attribute of these stem qJ OJ€ OJ o.o 6 c6 6 "r{ cheloniines is the disparate palatal and dentary morpholo- (D 0.) rd q) (6 .35 (6 !^E-} 'r, }: .r< >, 3- 'n qJ \H' ro bD 'Fo bo gies that they exhibit. Prior --r q) : to the appearance of these taxa ^H!F\ x 91 u X \.t t"E" { t P-,. tr O* s -f)Ll in the fossil record, all well-known (To.uochelt,s, - '-r O o R-baU cheloniids H o OF o .v* OJ .t-r >( 0.) S d9 E 4, q) G) .L' /^ L< OJ *HXk *.i V) Ctertochells, and the osteopygines) exhibit flat palates and Su si A &A 5 dentaries of varying lengths. Each of the Eocene stem cheloniines possess a different morphology. The symphysis Cheloniidae of Puppigerus does not extend beyond the forarnina dentofaciale majus, as it does in osteopygines (Zangerl,, (a) (b) l9l l), but Puppigerus does have a secondary palate equal to Figure 1. Hypotheses of the phylogenetic relationships of sea the extent of that in osteopygines . E,oclrelorte. however. turtles (Chelonioidea) showing alternative relationships of lacks a secondary palate (Casier, 1968), and this has been (a) To.rochelys and Ctenocltelys to Cheloniidae: Gaffney and interpreted as a reversal (Gaffney, 1979; Hirayamzl, 1994). Meylan, 1988: (b) Hirayama,, 1994. Finally . Argillochelys retains the secondary palate, but with a Osteoplginae This subfamily is composed of two groove to receive a markedly ridged dentary (the ridge occur- taxa, Osteopl,gis (Late Cretaceous) and Ercluelinnesicr (Early ring along the syrnphysis). A ridged dentary is not restricted to Eocene)., both known from cranial and post-cranial material. Argillrtchehs; some lowerjaws from the Late Cretaceous of The skulls of osteopygines are easily identifiable due to their New Jersey (AMNH 14205), not referable to any known extremely long secondary palates and broad, flat, dentaries. taxon, also have an elevated symphysis. Fr"rrthermore, some Like Toxocltel_r,s and Ctenochelts, osteopygines were once cheloniine taxa have a symphyseal ridge on the dentary. considered to be members of the Toxochelyidae (Zangerl, Chelortiinae This subfamily is defined here as those 1953,l97l). This assignment was based on the presence of turtles that share the most recent common ancestor of the wide plastra and curved humeri, characters that are now living cheloniids. S.r'//o t't't"Lt,s, a Miocene cheloniid with considered to be prirnitive for cheloniids (Gaffney and pseudodont dentition, has recently been assigned to this Meylan, 1988). Hirayama (1994) inferred that, like group by Hirayama (1994). He hypothesized that Syllomus Toxocltelys and Ctenochells,, osteopygines retain movable and Natator form a monophyletic sister group to the rest ol' articulations in the first and second digits (suggesting that the cheloniines. they lack the pelagic adaptation of a well-developed paddle); Some morphological analyses have divided the however, osteopygine specimens which retain these ele- Cheloniinae into two groups, Chelonini (Cheloniu and ments are not known to us. Eretmochely,s; Zangerl, 1958) and Carettini (Carerra and Fastovsky ( 1985) used cranial characters, such as a Lepiclochelys; Gray, 1825; as Carettidae). A rnorrophyletic basioccipital depression and the presence of a secondary Chelonini has been based primarily on characters which are palate, to ally the osteopygines with Recent cheloniids now considered to be primitive for cheloniines (e.g.', Al1 (Cheloniinae). Because the taxa included both have long elevated symphysis, a wide angle fonned by the scapular palates and the coinciding extensive symphyses, the processes, and the number of peripherals and pleural scutes; monophyly of osteopygines has never been challenged. Zangerl, 1958; Gaffney and Meylan, 1988). However, a However, the dentaries of turtles are notoriously plastic (Gaffney and Meylan, 1988) and the shells of the two taxa are quite different; Osteop.ygis possesses an ex- tremely ankylosed shell with no post-nuchal or costo- peripheral fontanelles whereas Ercluelinnesia has both (Fig. 2). For these reasons
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