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An Alternative Interpretation of Peltochelys Duchastelii As a Paracryptodire

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An Alternative Interpretation of Peltochelys Duchastelii As a Paracryptodire Foss. Rec., 23, 83–93, 2020 https://doi.org/10.5194/fr-23-83-2020 © Author(s) 2020. This work is distributed under the Creative Commons Attribution 4.0 License. An alternative interpretation of Peltochelys duchastelii as a paracryptodire Walter G. Joyce and Yann Rollot Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland Correspondence: Walter G. Joyce ([email protected]) Received: 16 December 2019 – Revised: 3 April 2020 – Accepted: 13 April 2020 – Published: 14 May 2020 Abstract. Peltochelys duchastelii is an enigmatic turtle from mines of Bernissart, located in Hainault Province, Belgium. Early Cretaceous (middle Barremian–early Aptian) Sainte- Dollo (1884) only provided an extremely brief description of Barbe Formation of Bernissart, Belgium. In more recent liter- his new species that highlights the presence of a single gular ature, there has been strong support for the trionychian affini- scute (his plaque intergulaire), the absence of mesoplastra, ties of this taxon, but this interpretation is less consistent with and a neck inconsistent with cryptodiran neck retraction. external data, in particular biogeography and temporal con- Lydekker and Boulenger (1887) concluded that the type siderations. We provide a reinterpretation here of the lecto- material of Peltochelys duchastelii is referable to the Early type of Peltochelys duchastelii that differs from more recent Cretaceous (Valanginian) taxon Tretosternon (their Tre- ones in key features. Although we agree that this turtle lacks tosternum) bakewelli (Mantell, 1833), together with a small mesoplastra, we conclude that it has 11 pairs of peripherals assortment of specimens from the Early Cretaceous (Berri- (not 10), likely had 12 pairs of marginals (not 11), lacks a asian) of Sussex, United Kingdom, and the Early Cretaceous contact between peripheral 1 and costal 1, and that a midline (Barremian) of the Isle of Wight, United Kingdom. It is diffi- contact of peripherals 1 hinders the nuchal from contributing cult to parse out the rationale of these authors (Joyce, 2017), to the margin of the shell. This unusual set of characters is as the available material shows little anatomical or temporal otherwise known from roughly coeval taxa from Europe and overlap. This synonymy was nevertheless accepted by sub- North America. Phylogenetic analysis yields a previously un- sequent authors (e.g., Lydekker, 1889; Dollo, 1909; Wegner, recognized clade of basal paracryptodiran turtles from the 1911), and Tretosternon bakewelli was typically character- Late Jurassic to Paleogene of Euramerica united by a nuchal ized by reference to Peltochelys duchastelii (e.g., Lydekker that is withdrawn from the anterior margin of the shell. The and Boulenger, 1887; Lydekker, 1889). Nopcsa (1928a, b) name Compsemydidae is referred to this clade. The novel concluded that the abovementioned English material can be hypothesis suggests that Peltochelys duchastelii did not im- united with other material from the Cretaceous of North migrate from Asia but instead originated in Europe. America and Europe into a previously unrecognized group, Helochelydridae (his Helochelydrinae, also known as Sole- mydidae), based on the presence of a distinct shell tex- ture consisting of raised tubercles, and that Peltochelys 1 Introduction duchastelii can be distinguished from helochelydrids by its subdued shell sculpturing and the absence of mesoplastra. Peltochelys duchastelii Dollo, 1884, is a fossil turtle from Even though this conclusion was further cemented by the in- the Early Cretaceous of Belgium with a particularly com- fluential work of Hummel (1929, 1932), the synonymy of plex history of research. The species was originally named Tretosternon bakewelli with Peltochelys duchastelii was re- by Dollo (1884) based on three individuals, in particu- vived once again many decades later by Młynarski (1976, lar the shell of an adult and two partial skeletons of ju- fig. 60.1), who presented the shell of Peltochelys duchastelii veniles. All specimens were collected in Early Cretaceous under the name Tretosternon bakewelli. (middle Barremian–early Aptian) exposures of the Sainte- Barbe Formation (Yans et al., 2005) in the underground coal Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 84 W. G. Joyce and Y. Rollot: Interpretation of Peltochelys duchastelii as a paracryptodire Dollo (1884) originally based Peltochelys duchastelii on istics that are otherwise only known to occur in combina- three specimens, of which two are juveniles. A century later, tion in carettochelyids. All subsequent analyses agree that Meylan (1988) designated the adult specimen as the lecto- Peltochelys duchastelii is a pan-trionychian, but there is mi- type, as he noted that a previously undescribed third juvenile nor disagreement as to whether it is the most basal represen- differs from the adult in the number of peripherals. In particu- tative of the lineage (e.g., Brinkman, 1998), the immediate lar, he noted that the third juvenile has 11 pairs of peripherals, sister to Trionychia (Meylan and Gaffney, 1989; Brinkman of which the most anterior closely approximate each other, and Peng, 1996; Joyce, 2007; Perez-García, 2011), or placed while the adult appears to have only 10 pairs of peripherals. within Carettochelyidae (e.g., Lapparent de Broin, 2001). As Pérez-García (2011) agreed with this conclusion but noted the basal evolution of Pan-Trionychia otherwise mostly oc- that the unfigured juvenile of Meylan (1988) appears to be curred in Asia, either placement implies independent disper- lost. Pérez-García (2015) soon after described two additional sal of the Peltochelys duchastelii lineage from Asia to Europe juvenile specimens but concluded that all available juvenile (Danilov, 2005). Finally, Danilov et al. (2017) summarized material, to the exclusion of the now lost specimen of Mey- this uncertainty by classifying Peltochelys duchastelii as a lan (1988), may reasonably represent Peltochelys duchastelii eucryptodire of uncertain affinities. after all. Peltochelys duchastelii is arguably the best figured Early Karl et al. (2012) expanded the known stratigraphic range Cretaceous turtle worldwide. Dollo (1884, pl. 2) furnished of Peltochelys duchastelii by referring new and previously photographs of the full type series in dorsal and ventral described specimens from the Late Jurassic (Kimmeridgian– view. Hummel (1929, fig. 2) provided interpretive line draw- Tithonian) of Guimarota and Lourinha, Portugal, and the ings of the lectotype in dorsal and ventral view, which were Early Cretaceous (Berriasian) of Schaumburg-Lippe and redrawn by Młynarski (1976, fig. 60.1), though under the Bückeberg, northern Germany. These referrals, however, name Tretosternon bakewelli. Meylan (1988, figs. 3, 4) cre- were convincingly rebuffed by Pérez-García (2014) and ated new photographs and interpretative line drawings of Pérez-García et al. (2015b), who noted that the Portuguese the lectotype, of which the line drawings were redrawn by material is only diagnostic of an indeterminate pleurosternid, Danilov (2005) and Karl et al. (2007). The full type series while the German material represents the pleurosternid was once again rephotographed and re-illustrated by Pérez- Dorsetochelys typocardium. Peltochelys duchastelii is there- García (2011, figs. 1–3). Although all available illustrations fore still only known from its type locality. of the lectotype greatly resemble one another, they differ in The phylogenetic relationships of Peltochelys duchastelii regard to some key features of great systematic value. The have been under debate ever since it was discovered. purpose of this contribution is to provide yet another inter- Dollo (1884) referred Peltochelys duchastelii to Pleurodira pretation of the morphology of Peltochelys duchastelii and (his Chelydes), likely based on the purported presence to propose the novel hypothesis that this enigmatic turtle is of a single gular (his plaque intergulaire) and the per- an unusual paracryptodire, not a representative of the pan- ceived lack of cryptodiran neck retraction. Lydekker and trionychian lineage. Boulenger (1887), in contrast, concluded that Tretosternon bakewelli (to the inclusion of Peltochelys duchastelii) has affinities with Chelydridae, but, as outlined above, their char- 2 Systematic paleontology acter evidence was drawn from what is now understood to be Testudinata Batsch, 1788 a chimera (Joyce et al., 2014; Joyce, 2017). Nopcsa (1928a, b) referred Peltochelys duchastelii to Dermatemydidae (his Paracryptodira Gaffney, 1975 Dermatemydinae) but did not discuss character evidence that would support this assessment. Hummel (1929, 1932) Compsemydidae Pérez-García et al., 2015b noted that Peltochelys duchastelii shows dermatemydid char- acters (i.e., presence of scutes and two suprapygals) and Peltochelys duchastelii Dollo, 1884 carettochelyid characters (i.e., overall size and shape of the shell, development of a preneural, and the presence Type specimen of a textured shell) and therefore suggested that it might be the Early Cretaceous dermatemydid ancestor of caret- IRSNB Ct.R.16, lectotype, a near complete shell (Dollo, tochelyids. Chkhikvadze (1975) referred this turtle to Adoci- 1884, pl. 2.1–2; Hummel, 1929, fig. 2; Meylan, 1988, figs. 3, dae, while Nessov (1977) referred it to Nanhsiungchelyidae. 4; Pérez-García, 2011, fig. 1; Fig. 1). Broin (1977) noted that Peltochelys duchastelii occurs too late in the fossil record to be considered an ancestral
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