Turtles have been known since the Upper Triassic (210 Myr old); however, fossils recording the first steps of turtle evolution are scarce and often fragmentary. As a consequence, one of the main questions is whether living turtles (Testudines) originated during the Late Triassic (210 Myr old) or during the Middle to Late Jurassic (
ca
160 Myr old). The discovery of the new fossil turtle,
Condorchelys antiqua
gen. et sp. nov., from the Middle to Upper Jurassic (
ca
160–146 Myr old) of South America (Patagonia, Argentina), presented here sheds new light on early turtle evolution. An updated cladistic analysis of turtles shows that
C. antiqua
and other fossil turtles are not crown turtles, but stem turtles. This cladistic analysis also shows that stem turtles were more diverse than previously thought, and that until the Middle to Upper Jurassic there were turtles without the modern jaw closure mechanism.
Methods improving the performance of molecular dating of divergence time of clades have improved dramatically in recent years. The calibration of molecular dating using the first appearance of a clade in the fossil record is a crucial step towards inferring the minimal diversification time of various groups and the choice of extinct taxa can strongly influence the molecular dates. Here, we evaluate the uncertainty on the phylogenetic position of extinct taxa through non-parametric bootstrapping. The recognition of phylogenetic uncertainty resulted in the definition of the Bootstrap Uncertainty Range (BUR) for the age of first appearance of a given clade. The BUR is calculated as the interval of geological time in which the diversification of a given clade can be inferred to have occurred, based on the temporal information of the fossil record and the topologies of the bootstrap trees. Divergence times based on BUR analyses were calculated for three clades of turtles: Testudines, Pleurodira and Cryptodira. This resulted in extensive uncertainty ranges of topology-dependent minimal divergence dates for these clades.
Pleurodires or side-necked turtles are today restricted to freshwater environments of South America, Africa–Madagascar and Australia, but in the past they were distributed much more broadly, being found also on Eurasia, India and North America, and marine environments. Two hypotheses were proposed to explain this distribution; in the first, vicariance would have shaped the current geographical distribution and, in the second, extinctions constrained a previously widespread distribution. Here, we aim to reconstruct pleurodiran biogeographic history and diversification patterns based on a new phylogenetic hypothesis recovered from the analysis of the largest morphological dataset yet compiled for the lineage, testing which biogeographical process prevailed during its evolutionary history. The resulting topology generally agrees with previous hypotheses of the group and shows that most diversification shifts were related to the exploration of new niches, e.g. littoral or marine radiations. In addition, as other turtles, pleurodires do not seem to have been much affected by either the Cretaceous–Palaeogene or the Eocene–Oligocene mass extinctions. The biogeographic analyses highlight the predominance of both anagenetic and cladogenetic dispersal events and support the importance of transoceanic dispersals as a more common driver of area changes than previously thought, agreeing with previous studies with other non-turtle lineages.
ABSTRACT-The fossil record of solemydid turtles is primarily based on isolated fragments collected from Late Jurassic to Late Cretaceous sediments throughout North America and Europe and little is therefore known about the morphology and evolutionary history of the group. We here provide a detailed description of the only known near-complete solemydid skeleton, which was collected from the Lower Cretaceous (Aptian-Albian) Antlers Formation of Texas during the midtwentieth century, but essentially remains undescribed to date. Though comparison is limited, the skeleton is referred to Naomichelys speciosa, which is based on an isolated entoplastron from the Lower Cretaceous (Aptian-Albian) Kootenai (Cloverly) Formation of Montana. The absence of temporal emarginations, contribution of the jugals to the orbits, and a clear subdivision of the middle and inner cavities, and the presence of elongate postorbitals, posteriorly expanded squamosals, a triangular fossa at the posterior margin of the squamosals, an additional pair of tubercula basioccipitale that is formed by the pterygoids, foramina pro ramo nervi vidiani (VII) that are visible in ventral view, shell sculpturing consisting of high tubercles, a large entoplastron with entoplastral scute, V-shaped anterior peripherals, and limb osteoderms with tubercular sculpture diagnose Naomichelys speciosa as a representative of Solemydidae. The full visibility of the parabasisphenoid complex in ventral view, the presence of an expanded symphyseal shelf, and the unusual ventromedial folding of the coronoid process are the primary characteristics that distinguish Naomichelys speciosa from the near-coeval European taxon Helochelydra nopcsai.
The origin and evolution of the crown-group of turtles (Cryptodira + Pleurodira) is one of the most interesting topics in turtle evolution, second perhaps only to the phylogenetic position of turtles among amniotes. The present contribution focuses on the former problem, exploring the phylogenetic relationships of extant and extinct turtles based on the most comprehensive phylogenetic dataset of morphological and molecular data analyzed to date. Parsimony analyses were conducted for different partitions of data (molecular and morphological) and for the combined dataset. In the present analysis, separate analyses of the molecular data always retrieve Pleurodira allied to Trionychia. Separate analysis of the morphological dataset, by contrast, depicts a more traditional arrangement of taxa, with Pleurodira as the sister group of Cryptodira, being Chelonioidea the most basal cryptodiran clade. The simultaneous analysis of all available data retrieves all major extant clades as monophyletic, except for Cryptodira given that Pleurodira is retrieved as the sister group of Trionychia. The paraphyly of Cryptodira is an unorthodox result, and is mainly caused by the combination of two factors. First, the molecular signal allies Pleurodira and Trionychia. Second, the morphological data with extinct taxa locates the position of the root of crown-group Testudines in the branch leading to Chelonioidea. This study highlights major but poorly explored topics of turtle evolution: the alternate position of Pleurodira and the root of crown turtles. The diversification of crown turtles is characterized by the presence of long external branches and short internal branches (with low support for the internal nodes separating the major clades of crown turtles), suggesting a rapid radiation of this clade. This rapid radiation is also supported by the fossil record, because soon after the appearance of the oldest crown-group turtles (Middle-Late Jurassic of Asia) the number and diversity of turtles increases remarkably. This evolutionary scenario of a rapid diversification of modern turtles into the major modern lineages is likely the reason for the difficulty in determining the interrelationships and the position of the root of crown-group turtles.
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