Clade Of Turtles Research Articles

A systematic comparative description of extant turtle humeri, with comments on humerus disparity and evolution based on fossil comparisons.

The humerus is central for locomotion in turtles as quadrupedal animals. Osteological variation across testudine clades remains poorly documented. Here, we systematically describe the humerus anatomy for all major extant turtle clades based on 38 species representing the phylogenetic and ecological diversity of crown turtles. Three Late Triassic species of shelled stem turtles (Testudindata) are included to establish the plesiomorphic humerus morphology. Our work is based on 3D models, establishing a publicly available digital database. Previously defined terms for anatomical sides of the humerus (e.g., dorsal, ventral) are often not aligned with the respective body sides in turtles and other quadrupedal animals with sprawling gait. We propose alternative anatomical directional terms to simplify communication: radial and ulnar (the sides articulating with the radius/ulna), capitular (the side bearing the humeral head), and intertubercular (opposite to capitular surface). Turtle humeri show low morphological variation with exceptions concentrated in locomotory specialists. We propose 15 discrete characters to summarize osteological variation for future phylogenetic studies. Disparity analyses comparing non-shelled and shelled turtles indicate that the presence of the shell constrains humerus variation. Flippered aquatic turtles are released from this constraint and significantly increase overall disparity. Ontogenetic changes of turtle humeri are related to increased ossification and pronunciation of the proximal processes, the distal articulation areas, and the closure of the ectepicondylar groove to a foramen. Some turtle species retain juvenile features into adulthood and provide evidence for paedomorphic evolution. We review major changes of turtle humerus morphology throughout the evolution of its stem group.

Two Major Extinction Events in the Evolutionary History of Turtles: One Caused by an Asteroid, the Other by Hominins.

AbstractWe live in a time of accelerated biological extinctions that has the potential to mirror past mass extinction events. However, the rarity of mass extinctions and the restructuring of diversity they cause complicate direct comparisons between the current extinction crisis and earlier events. Among animals, turtles (Testudinata) are one of few groups that have both a rich fossil record and sufficiently stable ecological and functional roles to enable meaningful comparisons between the end-Cretaceous mass extinction (∼66 Ma) and the ongoing wave of extinctions. Here we analyze the fossil record of the entire turtle clade and identify two peaks in extinction rates over their evolutionary history. The first coincides with the Cretaceous-Paleogene transition, reflecting patterns previously reported for other taxa. The second major extinction event started in the Pliocene and continues until now. This peak is detectable only for terrestrial turtles and started much earlier in Africa and Eurasia than elsewhere. On the basis of the timing, geography, and functional group of this extinction event, we postulate a link to co-occurring hominins rather than climate change as the cause. These results lend further support to the view that negative biodiversity impacts were already incurred by our ancestors and related lineages and demonstrate the severity of this continued impact through human activities.

Skull osteology, neuroanatomy, and jaw-related myology of the pig-nosed turtle Carettochelys insculpta (Cryptodira, Trionychia).

The osteology, neuroanatomy, and musculature are known for most primary clades of turtles (i.e., "families"), but knowledge is still lacking for one particular clade, the Carettochelyidae. Carettochelyids are represented by only one living taxon, the pig-nosed turtle Carettochelys insculpta. Here, we use micro-computed tomography of osteological and contrast-enhanced stained specimens to describe the cranial osteology, neuroanatomy, circulatory system, and jaw musculature of Carettochelys insculpta. The jaw-related myology is described in detail for the first time for this taxon, including m. zygomaticomandibularis, a muscular unit only found in trionychians. We also document a unique arterial pattern for the internal carotid artery and its subordinate branches and provide an extensive list of osteological ontogenetic differences. The present work provides new insights into the craniomandibular anatomy of turtles and will allow a better understanding of the evolutionary history of the circulatory system of trionychians and intraspecific variation among turtles.

Morphological Diversity of Turtle Hyoid Apparatus is Linked to Feeding Behavior.

The hyoid apparatus of tetrapods is highly diverse in its morphology. It plays an important role in feeding, breathing, sound production, and various other behaviors. Among turtles, the diversity of the hyoid apparatus has been recurrently linked to their habitat. The ossification of the hyoid corpus is often the main trait used in correlations with "niche" occupancy, an ossified corpus being associated with aquatic environments and a cartilaginous corpus with terrestrial life. Most studies conducted so far have focused on species belonging to Testudinoidea, the clade that occupies the biggest diversity of habitats (i.e., terrestrial, semi-terrestrial, and aquatic animals), while other turtle lineages have been largely understudied. We assessed the adult anatomy of the hyoid apparatus of 92 turtle species from all "families", together with ossification sequences from embryological series of 11 species, some described for the first time here. Using nearly 40 different discrete anatomical characters, we discuss the evolutionary patterns and the biological significance of morphological transformations in the turtle hyoid elements. Morphological changes are strongly associated to feeding modes, with several instances of convergent evolution within and outside the Testudines clade, and are not as strongly connected to habitat as previously thought. Some of the hyoid character states we describe are diagnostic of specific turtle clades, thus providing phylogenetically relevant information.

A description of a Denazinemys nodosa specimen (Testudinata, Baenidae) from the Late Cretaceous Kaiparowits Formation of southern Utah

Denazinemys nodosa is a Late Cretaceous representative of the North American turtle clade Baenidae diagnosed, among others, by a shell surface texture consisting of raised welts. We provide a detailed description of a partial skeleton from the late Campanian Kaiparowits Formation of Utah, USA, including bone-by-bone analysis of its cranium based on images obtained using micro-computed tomography. A revised phylogenetic analysis confirms placement of Denazinemys nodosa close to Eubaena cephalica and Boremys spp. within the clade Eubaeninae. Comparison with a second skull from the Kaiparowits Formation previously assigned to Denazinemys nodosa questions its referral to this taxon. An assortment of specimens from the Early to Late Campanian of Mexico and the USA had previously been referred to Denazinemys nodosa based on shell surface texture alone, even though this characteristic is known to occur in other baenids. Our review of all available material concludes that Denazinemys nodosa is currently only known from the Late Campanian of New Mexico and Utah.

Variation and development of the turtle chondrocranium, with a description of the common musk turtle (Sternotherus odoratus, Kinosternidae, Cryptodira, Testudines)

Based on histological cross-sections, the chondrocranium of the common musk turtle (Sternotherus odoratus) was reconstructed, described, and compared with other turtles. It differs from that of other turtle chondrocrania by possessing elongated, slightly dorsally orientated nasal capsules with three dorsolateral foramina, which might be homologous to the foramen epiphaniale, and by having an enlarged crista parotica. Additionally, the posterior part of the palatoquadrate is more elongated and more slender than in other turtles, while its ascending process is connected to the otic capsule by appositional bone. The proportions of the chondrocranium were also compared with those of "mature" chondrocrania of other turtle species in a Principal Component Analysis (PCA). Other than expected, the S. odoratus chondrocranium is not similar in proportions to those of chelydrids, the closest related species in the sample. The results indicate to differences in the proportions among larger turtle clades (e.g., Durocryptodira, Pleurodira, and Trionychia). S. odoratus is an exception to this pattern since it shows elongated nasal capsules similar to the trionychid Pelodiscus sinensis. A second PCA comparing the chondrocranial proportions of multiple developmental "stages" mostly shows differences between trionychids and all other turtles. S. odoratus is again similar to trionychids along PC1, but its proportions are the most similar along PC2 and PC3 to older "stages" of americhelydians, including the chelydrid Chelydra serpentina, which is related to chondrocranium height and quadrate width. We discuss potential ecological correlations of our findings mirrored in late embryonic stages.

Comparative biogeography of North American turtle faunas: Neogene regionalization

North America harbors substantial species diversity in nonmarineturtles (includes tortoises and terrapins), much of which arose in the Neogene Period (Miocene and Pliocene epochs) within Kinosternon, Emydinae, Trachemys, Pseudemys, Graptemys, and Gopherus. This diversity is distributed among 16 biogeographical provinces, but natural, hierarchical relations among provinces are unresolved. We used three-item analysis to identify such relations among provinces for these clades, following a recent, relatively complete phylogenetic reconstruction. The final three-item analysis identified 53,353 taxon‐area cladograms — free of paralogy and multiple areas on a single branch. The final intersection tree has a retention index of 73.6% and a completeness index of 75.4%, both indicating moderately strong congruence in patterns among turtle clades. All six turtle clades support some nodes on the intersection tree, which is divided into eastern and western forks. The crown group on the eastern fork includes four provinces almost entirely east of the Mississippi River drainage, whereas the western fork is split into wet tropics and aridlands sub-forks. The east-west transition zone spans the Mississippi River drainage and Great Plains. Our summary of divergence estimates and geological history suggests that although phylogenetic synchrony existed between select taxa, there was general asynchrony because provincial turtle faunas developed over an extended period (Neogene). Temperature-sensitive distributions of various taxa responded to climatic cycles by expanding during periods of warming, but contracting during periods of cooling. Grassland expansion and geomorphic change in the Neogene created provincial boundaries that, although sometimes crossed under favorable climates, more commonly acted as dispersal barriers. Despite asynchrony in faunal assembly, existence of shared patterns reveals natural relations among provinces, producing a regionalization for North American turtles, useful for understanding their evolution as well as the biogeography of North America.

Genetic variation among sea turtle life stages and species suggests connectivity among ocean basins.

Regional genetic differentiation of mitochondrial lineages occurs in migratory species with natal philopatry such as sea turtles. However, early juvenile dispersal represents a key opportunity for gene flow and colonization of new regions through founder events, making it an important yet under-studied life stage. To assess connectivity among sea turtle life stages and ocean basins, we sequenced mitochondrial DNA (mtDNA) fragments from 35 juveniles sampled in the Gulf of Mexico from the rarely observed dispersal stage across three species: green turtles (Chelonia mydas; n= 30), hawksbills (Eretmochelys imbricata; n= 3), and loggerheads (Caretta caretta; n= 2). We estimated green turtle rookery contributions using a many-to-many Bayesian mixed stock analysis that incorporated dispersal probabilities based on rookery size and transport via ocean currents. We assembled a gene tree including 709 distinct mtDNA control region haplotypes from the literature for all seven extant sea turtle species to assess gaps in life-stage data across ocean basins, as well as contextualize the lineages we sampled from dispersing juveniles. Our results indicate a high likelihood that green turtles sampled in the Gulf of Mexico originated from rookeries along the coast of Mexico, with smaller contributions from Costa Rica and Suriname. The gene tree analysis yielded species-level relationships consistent with those presented previously, while intra-species relationships between lineages and ocean basins differed, particularly within loggerhead and green turtle clades. Our results highlight the lack of genetic data from juvenile sea turtles, especially the early dispersal stage, and the potential for these data to answer broader questions of connectivity and diversification across species and lineages.

The evolution of reproductive strategies in turtles.

Optimal egg size theory assumes that changes in the egg and clutch are driven by selection, resulting in adjustments for the largest possible production of offspring with the highest fitness. Evidence supports the idea that large-bodied turtles tend to produce larger clutches with small and round eggs, while smaller species produce small clutches with large and elongated eggs. Our goals were to investigate whether egg and clutch size follow the predictions of egg size theory, if there are convergent reproductive strategies, and identify ecological factors that influence clutch and egg traits across all clades of living turtles. Using phylogenetic methods, we tested the covariance among reproductive traits, if they are convergent among different turtle lineages, and which ecological factors influence these traits. We found that both egg shape and size inversely correlate with clutch size, although with different evolutionary rates, following the predictions of the egg size theory. We also present compelling evidence for convergence among different turtle clades, over at least two reproductive strategies. Furthermore, climatic zone is the only ecological predictor to influence both egg size and fecundity, while diet only influences egg size. We conclude that egg and clutch traits in Testudines evolved independently several times across non-directly related clades that converged to similar reproductive strategies. Egg and clutch characteristics follow the trade-offs predicted by egg size theory and are influenced by ecological factors. Climatic zone and diet play an important role in the distribution of reproductive characteristics among turtles.

Iberoccitanemys atlanticum (Lapparent de Broin & Murelaga, 1996) n. comb.: new data on the diversity and paleobiogeographic distributions of the Campanian-Maastrichtian bothremydid turtles of Europe

Bothremydidae is the most abundant clade of turtles in the Campanian and Maastrichtian (Upper Cretaceous) fossil record of southwestern Europe. Several members of Foxemydina Gaffney, Tong & Meylan, 2006 are known in an area that includes Southern France and the North-Eastern half of Spain. The problematic ‘Polysternon’ atlanticum is the worst characterized, lacking a diagnosis that allows its specific validity to be confirmed, and whose generic attribution has been recognized as doubtful. Its presence was exclusively proposed in its type locality, the upper Campanian quarry of Laño, in Treviño County (Burgos Province, North of Spain). Despite the fact that knowledge about Bothremydidae has markedly increased after the description of ‘Polysternon’ atlanticum Lapparent de Broin & Murelaga, 1996, no new information about this species has been published since the 1990s. The analysis of abundant unpublished material of the bothremydid from Laño allows us to confirm the validity of this species. As a consequence of this study, it is not only identified in its type locality, but also in other Spanish regions and in the south of France. The diversity of Bothremydidae Baur, 1891 in the Upper Cretaceous of Europe is lower than previously considered. Thus, the species ‘Iberoccitanemys convenarum’ (Laurent, Tong & Claude, 2002), originally defined for the French record, and subsequently also identified in Spain, is identified here as a synonym of the species described in Laño. An emended diagnosis for the upper Campanian to upper Maastrichtian, Iberoccitanemys atlanticum (Lapparent de Broin & Murelaga, 1996) n. comb., is proposed.

Evolutionary history of mental glands in turtles reveals a single origin in an aquatic ancestor and recurrent losses independent of macrohabitat

Despite the relevance of chemical communication in vertebrates, comparative examinations of macroevolutionary trends in chemical signaling systems are scarce. Many turtle and tortoise species are reliant on chemical signals to communicate in aquatic and terrestrial macrohabitats, and many of these species possess specialized integumentary organs, termed mental glands (MGs), involved in the production of chemosignals. We inferred the evolutionary history of MGs and tested the impact of macrohabitat on their evolution. Inference of ancestral states along a time-calibrated phylogeny revealed a single origin in the ancestor of the subclade Testudinoidea. Thus, MGs represent homologous structures in all descending lineages. We also inferred multiple independent losses of MGs in both terrestrial and aquatic clades. Although MGs first appeared in an aquatic turtle (the testudinoid ancestor), macrohabitat seems to have had little effect on MG presence or absence in descendants. Instead, we find clade-specific evolutionary trends, with some clades showing increased gland size and morphological complexity, whereas others exhibiting reduction or MG loss. In sister clades inhabiting similar ecological niches, contrasting patterns (loss vs. maintenance) may occur. We conclude that the multiple losses of MGs in turtle clades have not been influenced by macrohabitat and that other factors have affected MG evolution.

A new pelomedusoid turtle, Sahonachelys mailakavava, from the Late Cretaceous of Madagascar provides evidence for convergent evolution of specialized suction feeding among pleurodires.

The Maevarano Formation in northwestern Madagascar has yielded a series of exceptional fossils over the course of the last three decades that provide important insights into the evolution of insular ecosystems during the latest Cretaceous (Maastrichtian). We here describe a new genus and species of pelomedusoid turtle from this formation, Sahonachelys mailakavava, based on a nearly complete skeleton. A phylogenetic analysis suggests close affinities of Sahonachelys mailakavava with the coeval Madagascan Sokatra antitra. These two taxa are the only known representatives of the newly recognized clade Sahonachelyidae, which is sister to the speciose clade formed by Bothremydidae and Podocnemidoidae. A close relationship with coeval Indian turtles of the clade Kurmademydini is notably absent. A functional assessment suggests that Sahonachelys mailakavava was a specialized suction feeder that preyed upon small-bodied invertebrates and vertebrates. This is a unique feeding strategy among crown pelomedusoids that is convergent upon that documented in numerous other clades of turtles and that highlights the distinct evolutionary pathways taken by Madagascan vertebrates.

A new baenid turtle, Palatobaena knellerorum sp. nov., from the lower Paleocene (Danian) Denver Formation of south-central Colorado, U.S.A.

ABSTRACT Baenidae are the most diverse clade of turtles from the latest Cretaceous and earliest Paleogene of North America. Palatobaena species have a distinctive cranial bauplan within baenid turtles and we herein describe a new species, P. knellerorum, based on a complete cranium from the lower Paleocene Denver Formation in the Denver Basin, Colorado. Palatobaena knellerorum differs from other Palatobaena species in the extreme degree of its dorsally oriented orbits, as well as the presence of a broad pentagonal midline crest formed by the supraoccipital and parietals. It can be further diagnosed from other species of Palatobaena by the following unique combination of characters: having a broadly rounded cranium, presence of upper temporal emargination that extends well anterior to the otic chamber, a broad exposure of the supraoccipital on the dorsal skull roof, a broadly rounded ‘tongue groove’ between the maxillae and premaxillae, and a deeply emarginated nasal region that exposes the underlying premaxillae in dorsal view. Palatobaena knellerorum most closely resembles the Maastrichtian/Danian taxon Palatobaena cohen and the Danian taxon P. bairdi. A phylogenetic analysis places P. knellerorum as sister to P. bairdi, and these taxa are sister to P. cohen. Palatobaena knellerorum overlaps in time with P. cohen, but the two are separated geographically with P. knellerorum restricted to the Denver Basin and P. cohen restricted to the Williston Basin. Cranial anatomical data and sedimentological data suggest Palatobaena spp. were bottom dwelling turtles living in shallow water environments that used a putative proboscis to prey upon freshwater-shelled invertebrates.

A nomenclature for fossil and living turtles using phylogenetically defined clade names

Over the last 25 years, researchers, mostly paleontologists, have developed a system of rank-free, phylogenetically defined names for the primary clades of turtles. As these names are not considered established by the PhyloCode, the newly created nomenclatural system that governs the naming of clades, we take the opportunity to convert the vast majority of previously defined clade names for extinct and extant turtles into this new nomenclatural framework. Some previously defined names are converted with minor adjustments. We also define a number of new clade names to close apparent nomenclatural gaps. In total, we establish 113 clade names, of which 79 had already received phylogenetic definitions and 34 are new.

A review of the carotid artery and facial nerve canal systems in extant turtles.

The cranial circulation and innervation systems of turtles have been studied for more than two centuries and extensively used to understand turtle systematics. Although a significant number of studies related to these structures exists, a broader comprehension of variation across the tree has been hindered by poor sampling and a lack of synthetic studies that addressed both systems together. We here provide new insights regarding the carotid circulation and facial nerve innervation systems in a broad set of extant turtles using CT (computed tomography) scans, which allow us to trace the canals these structures form in bone and understand the interaction between both systems. We document that the palatine artery, including the lateral carotid canal, is absent in all pleurodires and carettochelyids and was likely reduced or lost several times independently within Testudinoidea. We also highlight osteological correlates for the location of the mandibular artery. We finally summarize variation regarding the placement of the mandibular artery, location of the geniculate ganglion, placement of the hyomandibular and vidian nerves, and situations where we recommend caution when assessing canals in fossils. A morphometric study confirms that the relative sizes of the carotid canals are correlated with one another. Our results have the potential for building new phylogenetic characters and investigating the circulation systems of fossil taxa, which are expected to shed light on the evolution of the circulation system of turtles and clarify some unresolved relationships between fossil turtle clades.

CONTRIBUCIÓN A LA DIVERSIDAD DEL REGISTRO FÓSIL DE TORTUGAS (TESTUDINATA) DE LA PROVINCIA DE CHUBUT (ARGENTINA) Y SU SIGNIFICADO EN LA COMPRENSIÓN DE LA EVOLUCIÓN DE LAS TORTUGAS EN EL SUR DE SUDAMÉRICA

In this work we focus on the fossil record of turtles and tortoises from the Chubut Province, in Patagonia, Argentina. This record is the richest, most diverse, the longest and continuous in the country and one of the most important in the continent. In this work, we present and study new fossils from all known clades of turtles from the province, coming from targeted field campaigns, as well as from past investigations, placed in a comprehensive and detailed chronostratigraphic context. In Chubut, more than 241 occurrences of turtles from at least 223 collection points are known, spanning from the Toarcian (Jurassic) to the Tortonian (Miocene). we manage to fill some of the gaps in the fossil record, complete the anatomical knowledge of many taxa, explore new geographical areas, and present some important highlights. Among these, the record of the oldest pan-chelid turtles indicates the significant potential of the sedimentary deposits of Chubut. Furthermore, we discuss the changes in the diversity and faunal turnovers of the various turtle clades in this region during the last 180 Ma and across important events during the Cretaceous/Paleocene and Oligocene/Miocene boundaries.

Turtle remains from the late Miocene of the Cessaniti area, southern Italy—insights for a probable Tortonian chelonian dispersal from Europe to Africa

We here describe turtle remains from the late Miocene (Tortonian) of Cessaniti (Calabria, southern Italy), an area that recently has been palaeogeographically reconstructed as being, at that time of the Neogene, directly connected (or at least rather proximate) to northern Africa, instead of Europe. The material pertains to three different turtle clades, i.e., pan-trionychids, pan-cheloniids, and pan-geoemydids. Although the material is incomplete, it nevertheless permits a more precise identification for the pan-trionychid specimens, which are referred to the species Trionyx pliocenicus, as well as the pan-geoemydid, which is attributed to the genus Mauremys. Especially for the case of T. pliocenicus, the new Cessaniti specimens expand its geographic and stratigraphic distribution and further comprise the sole existing material known for this species, considering that its holotype and so far only known material is currently lost. Overall, besides its taxonomic significance, the Cessaniti chelonian assemblage affords the potential for important biogeographic implications, attesting that the lineages of Trionyx and Mauremys could have potentially used the Sicily–Calabria arch for their dispersal from Europe to Africa during the Tortonian. The new turtle specimens further complement the associated mammal remains in envisaging the Cessaniti assemblage as a mosaic of both African and Eurasian (Pikermian) faunal elements.

Turtle biogeography: Global regionalization and conservation priorities

Defaunation in the Anthropocene has created a need to focus limited conservation resources on geographically-explicit areas with high conservation significance. Priority conservation areas are often defined as those with high biodiversity – hotspots. While these conservation areas are critical to securing global biodiversity, prevailing approaches for their delineation are often qualitative. Here, we demonstrate the benefits of a clade-specific approach that improves conservation. We use the distinct, imperiled clade of turtles (tortoises and freshwater turtles) to delineate biogeographical regions and characterize their comparative levels of biodiversity and conservation values. We produce a quantitative, revisable map of 63 global turtle regions and identify several turtle regions, mostly in the Indomalayan turtle realm, that are high-priority conservation regions. While our high-priority turtle regions include those previously reported in the literature as turtle hotspots, we also describe a new priority conservation region in Southeast Asia (Yangtze-Huang He-Xi Yiang) which has both high species and endemism richness. Although not considered a high or intermediate conservation priority, our analyses delineated another previously unidentified turtle hotspot – the Kalahari Basin-Rift Valley. Additionally, we identify several turtle regions, largely in Central America, with intermediate conservation priority. Our results reveal that many turtle hotspots represent complex biogeographical areas with high inter-regional β-diversity, and several of these turtle-hotspots occur in transition zones with high biogeographical complexity. In these cases, inter-regional conservation efforts will be necessary to ensure turtle biodiversity conservation.

Effects of Sutured Pelvic Elements on Turtle Shell Strength: A Comparison of Pleurodire and Cryptodire Shell Mechanics

Abstract The two extant clades of turtles, Pleurodira and Cryptodira, can be distinguished by the fact that the pelvic elements of pleurodire turtles are extensively sutured to both the carapace an...

A new testudinoid turtle from the middle to late Eocene of Vietnam.

BackgroundTestudinoidea is a major clade of turtles that has colonized different ecological environments across the globe throughout the Tertiary. Aquatic testudinoids have a particularly rich fossil record in the Tertiary of the northern hemisphere, but little is known about the evolutionary history of the group, as the phylogenetic relationships of most fossils have not been established with confidence, in part due to high levels of homoplasy and polymorphism.MethodsWe here focus on describing a sample of 30 testudinoid shells, belonging to a single population that was collected from lake sediments from the middle to late Eocene (35–39 Ma) Na Duong Formation in Vietnam. The phylogenetic placement of this new material is investigated by integrating it and 11 other species of putative geoemydids from the Eocene and Oligocene to a recently published matrix of geoemydid turtles, that embraces the use of polymorphic characters, and then running a total-evidence analysis.ResultsThe new material is highly polymorphic, but can be inferred with confidence to be a new taxon, Banhxeochelys trani gen. et sp. nov. It shares morphological similarities with other southeastern Asian testudinoids, Isometremys lacuna and Guangdongemys pingi, but is placed phylogenetically at the base of Pan-Testuguria when fossils are included in the analysis, or as a stem geoemydid when other fossils are deactivated from the matrix. The vast majority of other putative fossil geoemydids are placed at the base of Pan-Testuguria as well.DiscussionThe phylogenetic placement of fossil testudinoids used in the analysis is discussed individually and each species compared to Banhxeochelys trani gen. et sp. nov. The high levels of polymorphism observed in the new taxon is discussed in terms of ontogenetic and random variability. This is the first time that a large sample of fossil testudinoids has its morphological variation described in detail.