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A Primitive Protostegid from Australia and Early Sea Turtle Evolution

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A Primitive Protostegid from Australia and Early Sea Turtle Evolution Downloaded from http://rsbl.royalsocietypublishing.org/ on May 18, 2015 Biol. Lett. (2006) 2, 116–119 Toxochelys and Ctenochelys (Gaffney & Meylan 1988) doi:10.1098/rsbl.2005.0406 as stem chelonioids; these taxa are currently placed Published online 15 November 2005 within crown-chelonioids, along the lineage leading to living cheloniids (see Hirayama 1994, 1997, 1998). A primitive protostegid 2. MATERIAL AND METHODS Bouliachelys suteri gen. et sp. nov. (figure 1a–g and electronic from Australia and early supplementary material) was added to the most comprehensive published phylogenetic data set of sea turtles (Hirayama 1998), sea turtle evolution which was revised as follows. The chimera taxon Osteopygis was separated into two taxa (the cranial Euclastes and postcranial 1,2, 1,2 Benjamin P. Kear * and Michael S. Y. Lee Osteopygis; Parham 2005), and additional postcranial characters for Euclastes scored from E.(Erquelinnesia) gosseleti (Lynch & Parham 1School of Earth and Environmental Sciences, University of Adelaide, 2003; Hirayama 1994). Modifications were also made to the Adelaide 5005, Australia codings and/or character state definitions of 12 characters. A full 2Earth Sciences Section, South Australian Museum, North Terrace, matrix and character list with annotations discussing these points is Adelaide 5000, Australia provided in electronic supplementary material. Maximum parsi- *Author and address for correspondence: Earth Sciences Section, mony trees, bootstrap frequencies (1000 replicates), and Bremer South Australian Museum, North Terrace, Adelaide 5000, Australia support were calculated using heuristic searches in phylogenetic ([email protected]). analysis using parsimony (PAUP; Swofford 2000) employing 1000 Sea turtles (Chelonioidea) are a prominent random-addition replicates. Analyses performed with multi-state group of modern marine reptiles whose early characters ordered or unordered (see electronic supplementary material) yielded similar results; the optimal tree (LZ211) from the history is poorly understood. Analysis of excep- unordered analysis is shown in figure 2. tionally well preserved fossils of Bouliachelys suteri gen. et sp. nov., a large-bodied basal protostegid (primitive chelonioid) from the 3. PHYLOGENETIC TAXONOMY Early Cretaceous (Albian) of Australia, indi- All higher taxon names sensu Joyce et al. 2004. cates that early sea turtles were both larger and Testudines Batsch, 1788; Chelonioidea Baur, 1893; more diverse than previously thought. The anal- Pandermochelys Joyce et al.(2004); Protostegidae ysis implies at least five distinct sea turtle Cope, 1872 lineages existed around 100 million years ago. B. suteri gen. et sp. nov. Currently, the postcranially primitive Cteno- chelys and Toxochelys are interpreted as crown- group sea turtles closely related to living chelo- (a) Etymology niids (e.g. Chelonia); in contrast, the new phylo- After Boulia township, the most productive locality geny suggests that they are transitional for this taxon (see below); and Richard and John (intermediate stem-taxa) between continental Suter who discovered many of the specimens. testudines and derived, pelagic chelonioids. Keywords: sea turtles; protostegidae; (b) Holotype, locality and horizon Early Cretaceous; Australia; stem chelonioids Holotype (figure 1a–c) QM F31669 (Queensland Museum, Brisbane, Australia) from Dunraven Station, near Hughenden, central-northern Queens- land, Australia; referred specimen (figure 1d–g)SAM 1. INTRODUCTION P41106 (South Australian Museum, Adelaide, Aus- Sea turtles (Chelonioidea) have a long fossil record tralia) from Boulia region, western Queensland, stretching back to the late Early Cretaceous (late Australia. Both localities are Toolebuc Formation Aptian/early Albian, ca 105 Myr ago; see Hirayama (Rolling Downs Group), Eromanga Basin, latest 1998). However, their early evolutionary history middle to late Albian, P. l u d b r o o k i a e Zone/upper remains largely unknown, as recent phylogenies do C. paradoxa–P. pannosus Zone (McMinn & Burger not recognize any definitive stem-group taxa. At 1986; Alexander & Sansome 1996). present, all fossil sea turtles are assigned to one of two extant lineages within crown chelonioids (c) Diagnosis (Hirayama 1994, 1997, 1998; Joyce et al.2004; Identical for genus and species due to monotypy. Lehman & Tomlinson 2004): one leads to living Bouliachelys possesses the unique derived features cheloniids (Pancheloniidae: Joyce et al. 2004), and (within chelonioids) of highly sculpted skull roof the other to living Dermochelys (Pandermochelys: bones, a rugose boss anterior to each orbit, and Joyce et al. 2004). The latter group includes the double longitudinal keels (formed by the basisphe- extinct Protostegidae, a diverse Cretaceous clade noid and pterygoids) on the ventral surface of the containing some spectacular Late Cretaceous giants basicranium and palate. Bouliachelys differs from (e.g. Archelon, possibly up to 4 m maximum length; Notochelone (the only other Australian fossil sea turtle Wieland 1896). The cranial morphology of protoste- known from cranial remains) in several additional gids is poorly known (Hooks 1998). However, here traits: it is larger, lacks a jugal-quadrate contact9, we report on several exceptionally well preserved possesses hooked premaxillae13, and an upper tritur- skulls belonging to one of the stratigraphically oldest ating surface incorporating the palatine15 but not and most primitive protostegids yet found. A re- vomer16, a pterygoid excluded from the mandibular evaluation of sea turtle phylogeny incorporating these condyle24, and a basiphenoid-pterygoid with a new fossils (re)establishes the primitive, shallow-water V-shaped crest31 (character subscript numbering follows the data matrix in the electronic supplemen- The electronic supplementary material is available at http://dx.doi. org/10.1098/rsbl.2005.0406 or via http://www.journals.royalsoc.ac. tary material, where distributions are scored across all uk. chelonioids). Bouliachelys is distinguished from all Received 21 July 2005 116 q 2005 The Royal Society Accepted 10 October 2005 Downloaded from http://rsbl.royalsocietypublishing.org/ on May 18, 2015 Primitive sea turtle B. P. Kear & M. S. Y. Lee 117 (a) (c) 20mm (b) (d) pm pa fr ( f ) (g) pm pf na po mx so pf vo q fr 40mm pm qj pt pal ju mx pa ju cor po pr (e) pr q qj q pt art den op bs sur bo ang so ex so Figure 1. Skulls of Bouliachelys suteri gen. et sp. nov. (a) Holotype (QM F31669) skull, (b) parietal section, and (c) mandible in dorsal view. Referred (SAM P41106) (d ) skull and (e) mandible in lateral, ( f ) dorsal and (g) ventral views (sutures highlighted). Additional photos can be found in the electronic supplementary material. Abbreviations: ang, angular; art, articular; bo, basioccipital; bs, basisphenoid; cor, coronoid; den, dentary; ex, exoccipital; fr, fontal; ju, jugal; mx, maxilla; na, nasal; op, opisthotic; pa, parietal; pal, palatine; pf, postfrontal; pm, premaxilla; po, postorbital; pr, prootic; pt, pterygoid; q, quadrate; qj, quadratojugal; so, supraoccipital; sur, surangular; vo, vomer. other chelonioids in displaying a unique mosaic of (an arrangement proposed elsewhere on braincase primitive and derived features that suggest it is a basal traits; see Gaffney and Meylan 1988; Hooks 1998). protostegid (figure 2). All protostegids can be diag- This hypothesis is further supported here by limb and nosed by presence of nasal bones2, palatines meeting girdle morphology, with Toxochelys and Ctenochelys medially20,foramenpalatinum posterius open retaining primitive postcranial features lost in crown posteriorly21, biconvex second or third cervical chelonioids (figure 2): ischium with a well developed vertebra50, lateral process restricted to anterior sur- lateral process64, humerus with a capitellum that is face of humerus shaft71, and a curved radius75 (c.f. upturned and shouldered68, first and second digits Hirayama 1998). Santanachelys is the most basal incorporated into paddle78, and femur with distinct known protostegid; Bouliachelys and later forms share trochanters79. The derived states of these postcranial the loss of the ventral cheek emargination12, and a characters—functionally related to pelagic natatory strong lingual ridge of the maxilla19. Finally, proto- habits—thus appear only once and were present in stegids above Santanchelys and Bouliachelys are united the ancestor of crown chelonioids; consequently they in having a triturating surface that excludes the do not need to evolve convergently in the lineages palatine15 but includes the vomer16. leading to living cheloniids and dermochelyids. The dorsally oriented orbits5 of Toxochelys and Ctenochelys also suggest that they retained the primitive condition 4. EVOLUTIONARY IMPLICATIONS of shallow-water and/or benthic habits (and poten- A phylogenetic analysis of major sea turtle lineages, tially a less pelagic existence; c.f. Hirayama 1997); in including Bouliachelys, was conducted (see electronic contrast, the orbits of most crown chelonioids face supplementary material). The traits discussed above laterally, implying a more ‘pelagic-adapted’ skull. robustly place Bouliachelys within crown (living) This new data clarifies other evolutionary trends Chelonioidea, Pandermochelys and Protostegidae; within sea turtles. (i) The Australian fossil taxa are and (with less certainty) suggest it is the most basal amongst the largest known before the Late Cretac- protostegid after Santanachelys (figure 2). While eous; the skull of Bouliachelys
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