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Bulletin of the Peabody Museum of Natural History 60(2): 129–155, 2019" Which Should Be Cited to Refer to This Work

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Bulletin of the Peabody Museum of Natural History 60(2): 129–155, 2019 Published in "Bulletin of the Peabody Museum of Natural History 60(2): 129–155, 2019" which should be cited to refer to this work. A Review of the Fossil Record of Nonbaenid Turtles of the Clade Paracryptodira Walter G. Joyce1 and Jérémy Anquetin2 1 Corresponding author: Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland —email: [email protected] 2 JURASSICA Museum, 2900 Porrentruy, Switzerland, and Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland —email: [email protected] ABSTRACT The fossil record of nonbaenid paracryptodires ranges from the Late Jurassic (Kimmeridgian) to the Paleocene of North America and Europe only. Earlier remains may be present as early as the Middle Jurassic (Bathonian). Only a single dispersal event is documented between the two conti- nents after their breakup during the Cretaceous in the form of the appearance of the Compsemys lineage in the Paleocene of France. Nonbaenid paracryptodires were restricted to freshwater aquatic environments but display adaptations to diverse feeding strategies consistent with gener- alist, gape-and-suction, and hypercarnivorous feeding. Current phylogenies recognize two species-rich subclades within Paracryptodira, Baenidae and Pleurosternidae, which jointly form the clade Baenoidea. A taxonomic review of nonbaenid paracryptodires concludes that of 34 named taxa, 11 are nomina valida, 15 are nomina invalida, and 8 are nomina dubia. KEYWORDS Phylogeny, Biogeography, Paleoecology, Paracryptodira, Baenoidea, Pleurosternidae Introduction (see Joyce et al. [2011] compared with Joyce [2017]), future work will need to clarify if all or The term Paracryptodira is herein used to refer to parts of the latter, at least as currently understood, the most inclusive clade of turtles that includes the are nested within the other. The fossil record of Eocene baenid Baena arenosa Leidy, 1870 and the Baenidae, one of the primary clades of Paracryp- Late Jurassic pleurosternid Pleurosternon bullockii todira, was previously summarized by Joyce and Owen, 1842, but no extant turtle (Lyson and Joyce Lyson (2015). We therefore here summarize the http://doc.rero.ch 2011). The name was originally introduced by fossil record of the remainder of the group. Gaffney (1975) to unite all turtles that possess a The idea that European pleurosternids and foramen posterius canalis carotici interni located North American baenids form a group had midway along the contact of the basisphenoid with already been proposed by Dollo (1886), who the pterygoid. Although Evans and Kemp (1976) united these turtles in his classification. This soon after questioned the utility of this character, arrangement was soon after supported by Lydekker as it reasonably may be interpreted as a symple- (1889a), who proposed the redundant names siomorphy, more recent analyses of turtle relation- Amphichelydia and Pleurosternidae based on a ships that densely sample at the species level series of characters that now must be viewed as typically retrieve the group of turtles with the plesiomorphic (Gaffney 1975). The grouping of paracryptodiran condition as monophyletic (e.g., Lydekker (1889a) does not fully correspond to the Joyce 2007; Lyson and Joyce 2011; Joyce et al. 2016). current understanding of Paracryptodira, how- As it is somewhat difficult to rigorously differenti- ever, as it included the stem pleurodire Platychelys ate paracryptodires from sympatric helochelydrids oberndorferi Wagner, 1853 and the helochelydrid 1 Helochelys danubina Meyer, 1854. Over the course The skulls of nonbaenid paracryptodires gen- of the succeeding decades, the meanings of the erally show poorly developed lower and upper names Amphichelydia and Pleurosternidae were temporal fenestrae (fully absent in Compsemys further removed from the currently hypothesized victa), but they range from flat and elongate with content of Paracryptodira, as increasing numbers dorsally oriented orbits (e.g., Glyptops ornatus), to of stem turtles were united with the former (e.g., short and high with dorsolaterally oriented orbits Williston 1925; Nopcsa 1928), while the latter was (e.g., Uluops uluops), to triangular and massive increasingly circumscribed to the exclusion of with laterally oriented orbits (e.g., Compsemys baenids (e.g., Baur 1891; Hay 1908b; Gilmore victa). All forms possess relatively large nasals 1919; Williams 1950). Although Gaffney (1975) with a midline contact, relatively small prefrontals could have used either name for the clade of tur- that lack a midline contact, frontals that con- tles he recognized, we here conform to his choice tribute to the orbits (absent in Compsemys victa), of the new name Paracryptodira, as this has been parietals that contact the squamosals (unclear in firmly entrenched in the literature in the last Compsemys victa), jugals that contribute to the decades. orbits (absent in Glyptops ornatus), and open For institutional abbreviations, see Appendix 1. incisurae columella auris (superficially enclosed Named nonbaenid paracryptodiran genera are in Compsemys victa). The pterygoids have been listed in Appendix 2. reported as lacking a midline contact in Glyptops ornatus and Pleurosternon bullockii, but this is Skeletal Morphology probably a preservational artifact, especially as a midline contact is clearly present in all taxa where Cranium this region is sufficiently preserved. The foramen At present, the skull is only known for five non- posterius canalis carotici interni enters the skull baenid paracryptodires: Compsemys victa from midway along the contact of the pterygoid with the Paleocene of Colorado, USA (see Lyson and the basisphenoid, but it is unclear if the palatal Joyce 2011); Dorsetochelys typocardium from the branch of the internal carotid is present, consider- Early Cretaceous of the United Kingdom (see ing that it has recently been shown to be absent in Dorsetochelys delairi of Evans and Kemp 1976); some baenids (Lipka et al. 2006; Rollot et al. 2018) Pleurosternon bullockii from the Early Cretaceous and that the internal anatomy of no skull has of of the United Kingdom (see Mesochelys durlsto- yet been studied. The basicranial region resem- nensis of Evans and Kemp 1975 and Pleurosternon bles that of most turtles by having an elongate bullockii of Sterli et al. 2010); Glyptops ornatus basisphenoid and broad pterygoids, with the from the Late Jurassic of Wyoming, USA (see exception of Compsemys victa, which displays an Glyptops plicatulus of Gaffney 1979); and Uluops extremely shortened basisphenoid and narrow uluops from the Late Jurassic of Wyoming, USA pterygoids. (Carpenter and Bakker 1990). A well-preserved Much variation is apparent in the palate of http://doc.rero.ch skull represents the holotype of Dorsetochelys buz- nonbaenid paracryptodires, which highlights zops (Bakker 1998), but as this specimen was likely differences in feeding ecology. In Glyptops never figured (only idealized sketches are avail- ornatus and Pleurosternon bullockii, the labial able) and is now lost, it is disregarded herein. ridges are low, and the triturating surfaces narrow. Although the available nonbaenid paracryptodire The labial ridges are higher and the triturating sur- skulls are relatively complete, most are either faces are expanded in Dorsetochelys typocardium heavily crushed (e.g., Dorsetochelys typocardium, and Uluops uluops, particularly through contribu- Glyptops ornatus, and Mesochelys durlstonensis) or tions from the palatines. In contrast, the labial poorly documented (e.g., Dorsetochelys typo - ridges of Compsemys victa are marked by a tooth- cardium and Uluops uluops). The only remaining like median projection, and the broad secondary skull, that of Compsemys victa, is highly apomor- palate elaborated by low accessory ridges is formed phic and not representative for the group. We by the maxillae, palatines, and vomer. These therefore do not figure any material herein and morphologies are broadly consistent with gape- keep our summary to a minimum based on the and-suction feeding, generalist feeding, and macro- sources cited previously. carnivory, respectively (see “Paleoecology”). 2 FIGURE 1. Shell morphology of basal Paracryptodira as exemplified by three species. A, Glyptops ornatus (redrawn from Gaffney 1979). B, Dorsetochelys typocardium (redrawn from Pérez-García 2014). C, Compsemys victa (redrawn from Gilmore 1919). Abbreviations: Ab, abdominal scute; An, anal scute; Ce, cervical scute; co, costal; ent, entoplastron; epi, epiplastron; Ex, extragular scute; Fe, femoral scute; Gu, gular scute; Hu, humeral scute; hyo, hyoplastron; hyp, hypoplastron; IM, inframarginal scute; Ma, marginal scute; mes, mesoplastron; ne, neural; nu, nuchal; Pe, pectoral scute; per, peripheral; Pl, pleural scute; py, pygal; spy, suprapygal; Ve, vertebral scute; xi, http://doc.rero.ch xiphiplastron. Scale bar approximates 5 cm. Shell Ortega 2011), and Toremys cassiopeia (Pérez- Most of the herein recognized species are known García, Espílez, et al. 2015). from well-preserved shell material, but not all are The carapace of most nonbaenid paracryp- sufficiently described. Informative descriptions todires resembles that of other basal turtles by are nevertheless available for Compsemys victa consisting of a nuchal, 8 neurals that fully sepa- (Gilmore 1919; Figure 1C), Dinochelys whitei rate the costals along the midline, 8 pairs of (Gaffney 1979), Dorsetochelys typocardium costals, 11 pairs of peripherals, 2 suprapygals, and (Pérez-García 2014; Figure
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