A large non-marine turtle from the Upper Cretaceous of Alabama and a review of North American "Macrobaenids".

The abundant record of marine turtles from the Late Cretaceous of Appalachia contrasts with the relatively poor record of contemporaneous terrestrial and freshwater species. A new taxon from the Santonian-Campanian of Alabama, Appalachemys ebersolei gen. et sp. nov., is described here and assigned to a grade of freshwater turtles known as "macrobaenids." Appalachemys can be differentiated from other "macrobaenids" by the presence of a nearly round carapace, deep nuchal emargination, and nine pairs of costals. With a carapace more than 80 cm in length, Appalachemys is among the largest freshwater turtles to ever inhabit North America. The absence of pre-Campanian "macrobaenids" from Laramidia indicates that the North American distribution of this grade may have been restricted to Appalachia prior to the recession of the Western Interior Seaway. Phylogenetic analysis places Appalachemys as the sister taxon to all post-Santonian "macrobaenids." Although the phylogeny lacks statistical support, it demonstrates morphological similarities between the K/Pg boundary species Osteopygis emarginatus and Maastrichtian-Danian species referred to Judithemys. We, therefore, refer all but the Campanian type species of Judithemys to the genus Osteopygis. A review of all North American "macrobaenid" occurrences reveals that despite originating in Asia, the record of the grade (as defined here) is predominantly North American. Future studies can test whether late Paleocene records in Asia and Europe resulted from dispersal from North America.

Asmodochelys parhami, a new fossil marine turtle from the Campanian Demopolis Chalk and the stratigraphic congruence of competing marine turtle phylogenies.

Resolving the phylogeny of sea turtles is uniquely challenging given the high potential for the unification of convergent lineages due to systematic homoplasy. Equivocal reconstructions of marine turtle evolution subsequently inhibit efforts to establish fossil calibrations for molecular divergence estimates and prevent the accurate reconciliation of biogeographic or palaeoclimatic data with phylogenetic hypotheses. Here we describe a new genus and species of marine turtle, Asmodochelys parhami, from the Upper Campanian Demopolis Chalk of Alabama and Mississippi, USA represented by three partial shells. Phylogenetic analysis shows that A. parhami belongs to the ctenochelyids, an extinct group that shares characteristics with both pan-chelonioids and pan-cheloniids. In addition to supporting Ctenochelyidae as a sister taxon of Chelonioidea, our analysis places Protostegidae outside of the Chelonioidea crown group and recovers Allopleuron hofmanni as a stem dermochelyid. Gap excess ratio (GER) results indicate a strong stratigraphic congruence of our phylogenetic hypothesis; however, the highest GER value is associated with the phylogenetic hypothesis of marine turtles which excludes Protostegidae from the Cryptodira crown group. Ancestral range estimations derived from our phylogeny imply a European or North American origin of Chelonioidea in the middle-to-late Campanian, approximately 20 Myr earlier than current molecular divergence studies suggest.

Prionochelys matutina Zangerl, 1953 (Testudines: Pan-Cheloniidae) from the Late Cretaceous of the United States and the evolution of epithecal ossifications in marine turtles.

BACKGROUND: Many neritic to nearshore species of marine adapted turtle from the Late Cretaceous of North America are thought to represent the stem lineage of Cheloniidae but due to fragmentary holotypes, low total specimen counts, and resultantly incomplete morphological character suites, are routinely placed either within or outside of crown group Chelonioidea leaving their precise cladistic affinities uncertain. Despite this systematic ambiguity, the referral of these species to either the stem of Cheloniidae or Chelonioidea belies the critical importance of these taxa in any investigation into the origins of extant marine turtles. The adequate incorporation of these species into phylogenetic studies requires the formal description of relatively complete specimens, particularly those possessing associated cranial and post-cranial material. METHODS: Remarkably complete fossil specimens of several adult and juvenile marine turtles from the Mooreville Chalk and Eutaw Formations (Alabama, USA) are formally described and assigned to Prionochelys matutina. This material provides new information into the anatomy, ontogeny, and cladistic affinities of the species. A phylogenetic hypothesis for Late Cretaceous marine turtles is then generated through the consilience of stratigraphic, morphological, and molecular data. RESULTS: Phylogenetic analysis places Prionochelys matutina on the stem of Cheloniidae as a member of a monophyletic clade with other putative pan-cheloniids, including Ctenochelys stenoporus, Ctenochelys acris, Peritresius martini, and Peritresius ornatus. The members of this clade possess incipient secondary palates, pronounced carapacial and plastral fontanelles at all stages of development, and are characterized by the presence of superficial ossifications at the apices of the neural keel elevations along the dorsal midline of the carapace. DISCUSSION: The epithecal osteoderms dorsal to the neural series (epineurals) found in Ctenochelyidae are unique among turtles. The presence of epineurals in ctenochelyid turtles shows that epithecal ossifications arose independently in both leatherback (Dermochelyidae) and hard-shelled (Cheloniidae) marine turtles. Whether or not the epineurals of Ctenochelyidae are homologous with the dermal ossicles comprising the carapace of Dermochelys coriacea remains untested however, histological thin sectioning of dermochelyid and ctenochelyd epithecal elements may reveal meaningful information in future studies.

A new species of Peritresius Leidy, 1856 (Testudines: Pan-Cheloniidae) from the Late Cretaceous (Campanian) of Alabama, USA, and the occurrence of the genus within the Mississippi Embayment of North America.

Late Cretaceous members of Peritresius belong to a diverse clade of marine adapted turtles currently thought to be some of the earliest representatives of the lineage leading to modern hard-shelled sea turtles (Pan-Cheloniidae). Prior studies have suggested that Peritresius was monospecific, with a distribution restricted to Maastrichtian deposits in North America. However, new Peritresius specimens identified from Alabama and Mississippi, USA, show that the genus contains two taxa, Peritresius ornatus, and a new species Peritresius martini sp. nov. These two taxa are characterized by the presence of a generally cordiform carapace with moderately serrated peripherals, well-developed ventral flanges beginning at the third peripheral, squarish umbilical and lateral plastral fontanelles, and a narrow bridge formed by the contact between the hyoplastron and hypoplastron. Peritresius martini sp. nov. can be distinguished by its lack of dermal ornamentation and the presence of a 'rib-free' 10th peripheral. These new specimens represent the first occurrences of Peritresius from the Late Cretaceous Mississippi Embayment and extend the temporal range of this genus back to the early Campanian. When tested within a global phylogenetic context, Peritresius is placed on the stem of Cheloniidae (Pan-Cheloniidae) along with Ctenochelys and Allopleuron hofmanni. The heavily vascularized and uniquely sculptured dermal elements of P. ornatus are interpreted here as potentially relating to thermoregulation and therefore may have been one of the key factors contributing to the survival of Peritresius into the Maastrichtian, a period of cooling when other lineages of Campanian marine turtles (e.g., Protostegids, Toxochelys, and Ctenochelys) went extinct.