Terrestrial Turtles Research Articles

A synopsis of the Early Miocene St Bathans Fauna of New Zealand

The St Bathans Fauna, from sites near the village of St Bathans, Central Otago, South Island, is the first substantive pre-Quaternary terrestrial vertebrate fossil fauna discovered in New Zealand. This fauna derives from 33 sites or discrete sedimentary beds located in the lower 50 m of the lacustrine Bannockburn Formation, Manuherikia Group, and is generally accepted as local stage Altonian (19–15.9 Ma; Burdigalian, Early Miocene) in age. Investigations since 2001 have revealed an abundant and diverse fauna from over 9000 catalogued lots that is herein reviewed. Invertebrates notably include eight genera and species of terrestrial molluscs. Among vertebrates, freshwater fish remains dominate with 17 species evidenced by 16,500 analysed otoliths (genera Neochanna, Galaxias, Prototroctes, and Mataichthys) and many thousands of bones. Birds (minimally 45 species, several thousand bones) are the most common non-fish vertebrates, among which waterfowls dominate all assemblages (10 species). Co-occurring with these was a diverse herpetofauna, including undetermined crocodylians and a terrestrial turtle, both absent in Recent faunas. Significantly, the St Bathans Fauna evidences that Zealandia already had all of New Zealand’s ‘old’ endemic Recent taxa (sphenodontids, leiopelmatids, dinornithiforms, apterygids, aptornithids, strigopoid parrots, acanthisittids, and mystacinids) during the Early Miocene. Furthermore, it includes Australasia’s oldest ardeids, two flightless rallids, a novel higher landbird family, a greater diversity of bats, and terrestrial mammals. All sites reflect a single fauna, except that the ducks Manuherikia lacustrina (stratigraphically lower in section) and M. primadividua (higher) have a mutually exclusive distribution that is not yet correlated with any other biotic distribution differences.

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.

Primeras huellas de tortugas terrestres gigantes del Cretácico Inferior (Soria, España): Testudopodus iberiae, nov. icnogen. nov. icnoesp.

En este trabajo se analizan las icnitas del yacimiento de “Valdelalosa III”, perteneciente a la subcuenca oriental de la Cuenca de Cameros (Cordillera Ibérica), Formación Huérteles, Berriasiense medio-superior (Cretácico basal), situado en los Valles de Valdelalosa, en el municipio de San Pedro Manrique (provincia de Soria, España). Las huellas de manos y pies son de diferente tamaño, tienen forma circular u ovalada y ocasionalmente marcan tres-cuatro uñas cortas y redondeadas. En el yacimiento se contabilizan 106 icnitas de las que 76 son huellas aisladas, 11 configuran un rastro, 15 son subhuellas y 4 son huellas parciales. En el rastro las impresiones de las extremidades del lado derecho del animal son paralelas a las del lado izquierdo y mantienen una distancia constante entre sí, carácter diagnóstico de los rastros de las tortugas. Las icnitas se han comparado morfológicamente y morfométricamente con otras icnitas de tortugas, con huellas de tortugas actuales, y con el pie y la mano de varios ejemplares de tortugas terrestres gigantes. Los resultados obtenidos de los análisis de las icnitas del yacimiento de “Valdelalosa III” permiten atribuirlas al paso de unas tortugas terrestres de gran tamaño y, debido a sus características se propone la creación del icnotaxon Testudopodus iberiae, nov. icnogen. nov icnosp.

Ecological and Fitness Correlates of Personality in a Long-lived Terrestrial Turtle.

An individual's behavioral tendencies (i.e., personality or temperament) can influence its interactions with the environment and thus have important ecological and evolutionary consequences for animal populations. Boldness, defined as an individual's tendency to engage in risk-taking activities, is a phenotypically variable trait linked with numerous behavioral and fitness outcomes in free-ranging animals. We examined variation and repeatability of boldness and other behavioral characteristics in two wild Eastern Box Turtle (Terrapene carolina carolina) populations using radiotelemetry, and assessed fitness correlates of boldness over multiple years. We observed large amounts of among-individual variation and within-individual consistency (i.e., repeatability) of boldness as measured by their head emergence latency following a standardized confinement assay. Individuals were also consistent in several in-field behaviors including movement rate, home range size, and date of emergence from overwintering refuges. Individuals with shorter head emergence latencies (i.e., bolder turtles) had larger home ranges, emerged earlier from overwintering dormancy, and experienced moderately lower survival compared with shy individuals. Boldness did not affect time spent within the thermal preference range, somatic growth rates, or the frequency of mating or same-sex aggressive encounters. Boldness and its effects on in-field behaviors differed between sexes and populations, and the relationship between boldness and survival was temporally variable. Our results suggest possible intrinsic behavioral types in T. c. carolina and highlight the importance of long-term and multipopulation studies when examining ecological and evolutionary processes that shape personality phenotypes in turtles.

(Not) far from home: No sex bias in dispersal, but limited genetic patch size, in an endangered species, the Spotted Turtle (Clemmys guttata).

Sex-biased dispersal is common in many animals, with male-biased dispersal often found in studies of mammals and reptiles, including interpretations of spatial genetic structure, ostensibly as a result of male-male competition and a lack of male parental care. Few studies of sex-biased dispersal have been conducted in turtles, but a handful of studies, in saltwater turtles and in terrestrial turtles, have detected male-biased dispersal as expected. We tested for sex-biased dispersal in the endangered freshwater turtle, the spotted turtle (Clemmys guttata) by investigating fine-scale genetic spatial structure of males and females. We found significant spatial genetic structure in both sexes, but the patterns mimicked each other. Both males and females typically had higher than expected relatedness at distances <25 km, and in many distance classes greater than 25 km, less than expected relatedness. Similar patterns were apparent whether we used only loci in Hardy-Weinberg equilibrium (n=7) or also included loci with potential null alleles (n=5). We conclude that, contrary to expectations, sex-biased dispersal is not occurring in this species, possibly related to the reverse sexual dimorphism in this species, with females having brighter colors. We did, however, detect significant spatial genetic structure in males and females, separate and combined, showing philopatry within a genetic patch size of <25 km in C.guttata, which is concerning for an endangered species whose populations are often separated by distances greater than the genetic patch size.

Comparative postcranial osteohistology and bone histovariability of aquatic and terrestrial turtles: the case of the South American Phrynops hilarii, Hydromedusa tectifera (Pleurodira, Chelidae), and Chelonoidis chilensis (Cryptodira, Testudinidae).

This article presents a detailed comparative analysis of the bone microstructure of three extant species of South American turtles. The main histological characteristics of postcranial bones are identified, as well as the intraskeletal, ontogenetic and interspecific variation between aquatic and terrestrial species. For this purpose, thin sections of postcranial bones (seventh cervical vertebra, coracoid, scapula, humerus, radius, ulna, ischium, ilium, pubis, femur, tibia, and fibula) of juvenile and adult specimens of aquatic (Phrynops hilarii and Hydromedusa tectifera) and terrestrial (Chelonoidis chilensis) turtles were analyzed. Bone histology revealed an intraskeletal variation of the microanatomical and microstructural organization in these turtles. The cortical bone is composed of poorly vascularized lamellar and parallel-fibered bone tissue interrupted with lines of arrested growth (LAGs), reflecting a cyclical slow growth rate throughout these turtles' life. Although in the adult specimens a growth rate decrease was observed, none of them have reached somatic maturity. The juvenile and the adult of Chelonoidis chilensis, unlike the aquatic species studied, presented a higher vascularization in their bones, which could imply a faster growth rate in this land specie. The number of LAGs was higher in the stylopodial and zeugopodial bones, which would make these elements suitable for approximate age estimations. Pectoral and pelvic girdle bones also exhibited a good record of LAGs. The information here obtained on extant species represents a powerful tool for the interpretation of paleobiological traits present in closely related fossil forms.

Oropharyngeal cavity floor morphology in Eretmochelys imbricata (Testudines: Cheloniidae) hatchlings and evolutionary implications.

Morphological studies of the oropharyngeal cavity of chelonians have become an interesting tool in the understanding of evolutionary processes associated with feeding habits in aquatic animals and the transition from aquatic to terrestrial forms. In this context, the aim of the present study was to describe the oropharyngeal cavity floor morphology of hawksbill sea turtle (Eretmochelys imbricata) hatchlings. Ten dead hatchlings of undefined sex were obtained from nests hatched on the coast of the state of Rio Grande do Norte, Brazil. The heads of each specimen were fixed, dissected, and analyzed at the macroscopic and microscopic levels. The oropharyngeal cavity floor of the hawksbill sea turtle hatchlings is formed by the tongue, pharynx, floor muscles, and hyolingual skeleton, delimited in the rostral and lateral directions by a keratinized beak, called the rhamphotheca, and in the caudal region at the limit between the pharynx and the esophagus. The tongue muscles and the muscles that support the floor of the oral cavity comprise the following: m. hypoglossohyoideus, m. hypoglossoglossus, m. hyoglossus, m. genioglossus, m. constrictor laryngis, m. geniohyoideus pars lateralis, and m. intermandibularis. The oropharyngeal cavity floor mucosa is formed by keratinized stratified squamous epithelium and the lamina propria is formed by loose connective tissue. The floor mucosa is devoid of taste buds. We believe that the basic oropharyngeal cavity floor characteristics in hawksbill sea turtle hatchlings may comprise indications that these animals are plesiomorphic and that semiaquatic and terrestrial turtles may have undergone adaptations to feed out of water.

A comparison of non-surgical methods for sexing young gopher tortoises (Gopherus polyphemus).

Many turtle species have temperature-dependent sex determination (TSD), raising the prospect that climate change could impact population dynamics by altering sex ratios. Understanding how climate change will affect populations of animals with TSD requires a reliable and minimally invasive method of identifying the sexes of young individuals. This determination is challenging in many turtles, which often lack conspicuous external sexual dimorphism until years after hatching. Here, we explore four alternatives for sexing three age classes of captive-reared young gopher tortoises (Gopherus polyphemus), a terrestrial turtle of conservation concern native to the southeastern United States: (1) naive testosterone levels, (2) testosterone levels following a follicle stimulating hormone (FSH) challenge, (3) linear morphological measurements, and (4) geometric morphometrics. Unlike some other turtle species, male and female neonatal gopher tortoises have overlapping naive testosterone concentration distributions, justifying more complicated methods. We found that sex of neonates (<7 days old) is best predicted by a “random forest” machine learning model with naive testosterone levels and morphological measurements (8% out-of-bag error). Sex of hatchlings (4–8 months old) was predicted with 11% error using a simple threshold on naive testosterone levels, or with 4% error using a simple threshold on post-FSH testosterone levels. Sex of juveniles (approximately 3.5 years old) was perfectly predicted using a simple threshold on naive testosterone levels. Sexing hatchlings at >4 months of age is the easiest and most reliable non-surgical method for sex identification. Given access to a rearing facility and equipment to perform hormone assays, these methods have the potential to supplant laparoscopic surgery as the method of choice for sexing young gopher tortoises.

Developing Collegial Relationships to Address Hurdles in Ex Situ Turtle Conservation on an Indonesian University Campus

Building collegial international relationships is an important aspect of conservation and conservation education. The relationship relies on researchers understanding the local context of the conservation project and the sociocultural perspectives of all partners. Therefore, we situated our case study within sociocultural theory. Our case study focuses on the relationships and project work which developed between a biologist/science educator at the University of Bengkulu in Bengkulu, Sumatra, Indonesia, and a science education professor at the University of North Carolina Greensboro, Greensboro, NC, USA, who shared longtime interests in herpetology. Employing the sociocultural perspective, we describe the background of each University group leader and how their previous experiences led to the development of a partnership focused on turtle conservation. In this case study, we seek to understand the social and cultural development of the researchers’ relationship as they teamed to address the ex situ conservation challenges of working with terrestrial and semiaquatic turtles. An examination of the researchers’ partnership provides a framework for the analysis of the work accomplished and remaining and offers insights to others interested in collaborative international conservation projects.

Chromosome-level genome assembly of the Chinese three-keeled pond turtle (Mauremys reevesii) provides insights into freshwater adaptation.

Mauremys reevesii is an endangered freshwater turtle that symbolizes longevity in Chinese culture. Despite its importance, genetic studies of this species remain limited, with no genomic sequence reported to date. Here, we report a high-quality, chromosome-level genomic sequence of M. reevesii obtained using a combination of Nanopore and Hi-C sequencing technologies. The 2.37Gb M. reevesii genome was assembled from a total of ~226.80Gb of Nanopore sequencing data. The M. reevesii genome contig N50 is 34.73Mb, the highest value in published turtle genomes. In total, 18,238genes were functionally annotated. The contigs were clustered and ordered onto 27pseudochromosomes covering ~96.55% of the genome assembled with Hi-C data. To explore genome evolution, synteny analysis was performed between M. reevesii (freshwater turtle) and Gopherus evgoodei (terrestrial turtle) genomes. In general, each chromosome of M. reevesii corresponded to one chromosome of Gopherus evgoodei, but some interchromosomal rearrangements occurred between the two species based on the assembled genomes. These interchromosomal rearrangements were further confirmed by mapping of the long-read nanopore data to the assembly. The reconstructed demographic history showed varied effective population size among freshwater, marine and terrestrial turtles. We also discovered expansion of genes related to the innate immune system in M. reevesii that may provide defence against freshwater pathogens. The high-quality genomic sequence provides a valuable genetic resource for further studies of genetics and genome evolution in turtles.

A large and unusually thick-shelled turtle egg with embryonic remains from the Upper Cretaceous of China.

Turtle eggs containing embryos are exceedingly rare in the fossil record. Here, we provide the first description and taxonomic identification, to our knowledge, of a fossilized embryonic turtle preserved in an egg, a fossil recovered from the Upper Cretaceous Xiaguan Formation of Henan Province, China. The specimen is attributed to the Nanhsiungchelyidae (Pan-Trionychia), an extinct group of large terrestrial turtles (possibly the species Yuchelys nanyangensis). The egg is rigid, spherical, and is one of the largest and thickest shelled Mesozoic turtle eggs known. Importantly, this specimen allowed identification of other nanhsiungchelyid egg clutches and comparison to those of Adocidae, as Nanhsiungchelyidae and Adocidae form the basal extinct clade Adocusia of the Pan-Trionychia (includes living soft-shelled turtles). Despite the differences in habitat adaptations, nanhsiungchelyids (terrestrial) and adocids (aquatic) shared several reproductive traits, including relatively thick eggshells, medium size clutches and relatively large eggs, which may be primitive for trionychoids (including Adocusia and Carrettochelyidae). The unusually thick calcareous eggshell of nanhsiungchelyids compared to those of all other turtles (including adocids) may be related to a nesting style adaptation to an extremely harsh environment.

Salmonella Infection in Turtles: A Risk for Staff Involved in Wildlife Management?

Simple SummaryThe aim of this study was to investigate the occurrence of non-typhoidal Salmonella in the turtles housed in a regional wildlife rescue centre of Apulia, in southern Italy, to assess the presence of Salmonella serovars that may represent a risk for operators involved in wildlife management. Sixty-nine tortoises, of which 36 were males and 33 were females, belonging to different species (Testudo hermanni hermanni, T. h. boettgeri, T. graeca, and T. marginata) were tested. All the turtles were adults (34 between 6 and 10 years of age and 35 more than 10 years of age). Salmonella was statistically detected more frequently in T. hermanni hermanni. No differences of the infection prevalence related to animal gender or age were found. Two different species, S. enterica and S. bongori, three S. enterica subspecies (enterica, diarizonae, salamae), and five different serovars (Hermannswerder, Abony, Ferruch, Richmond, Vancouver) within the group S. enterica subspecies enterica were identified. Two Salmonella types with different combinations were simultaneously found in specimens of T. h. hermanni. Most of the detected Salmonella types may represent a potential risk for operators in wildlife rescue centres.Monitoring of infections that may be transmitted to humans by animals in wildlife rescue centres is very important in order to protect the staff engaged in rehabilitation practices. Salmonella may be a natural inhabitant of the intestinal tract of turtles, rarely causing disease. This may represent a potential risk for humans, increasing the sanitary risk for operators in wildlife rescue centres. In this paper, the occurrence of non-typhoidal Salmonella among terrestrial turtles housed in a wildlife rescue centre in Southern Italy was investigated, in order to assess the serovars more frequently carried by turtles and identify those that may represent a risk for operators involved in wildlife management. Sixty-nine adult turtles (Testudo hermanni hermanni, T. h. boettgeri, T. graeca, and T. marginata) were tested. Detection and serotyping of Salmonella strains were performed according to ISO 6579-1 and ISO/TR 6579-3:2013, respectively. The distribution of Salmonella spp. was significantly higher in T. hermanni hermanni than in other species, independent of the age and gender of the animals. Two different Salmonella species, S. enterica and S. bongori, three S. enterica subspecies (enterica, diarizonae, salamae), and five different serovars (Hermannswerder, Abony, Ferruch, Richmond, Vancouver) within the group S. enterica subspecies enterica were identified. Different combinations of Salmonella types were simultaneously found in specimens of T. h. hermanni. Most of detected Salmonella types may represent a potential risk for public health. Adopting correct animal husbandry procedures and informing on potential sanitary risks may be useful for minimising the risk of transmission of Salmonella to workers involved in wildlife management.

A Pliocene\u2013Pleistocene continental biota from Venezuela

The Pliocene–Pleistocene transition in the Neotropics is poorly understood despite the major climatic changes that occurred at the onset of the Quaternary. The San Gregorio Formation, the younger unit of the Urumaco Sequence, preserves a fauna that documents this critical transition. We report stingrays, freshwater bony fishes, amphibians, crocodiles, lizards, snakes, aquatic and terrestrial turtles, and mammals. A total of 49 taxa are reported from the Vergel Member (late Pliocene) and nine taxa from the Cocuiza Member (Early Pleistocene), with 28 and 18 taxa reported for the first time in the Urumaco sequence and Venezuela, respectively. Our findings include the first fossil record of the freshwater fishes Megaleporinus, Schizodon, Amblydoras, Scorpiodoras, and the pipesnake Anilius scytale, all from Pliocene strata. The late Pliocene and Early Pleistocene ages proposed here for the Vergel and Cocuiza members, respectively, are supported by their stratigraphic position, palynology, nannoplankton, and 86Sr/88Sr dating. Mammals from the Vergel Member are associated with the first major pulse of the Great American Biotic Interchange. In contrast to the dry conditions prevailing today, the San Gregorio Formation documents mixed open grassland/forest areas surrounding permanent freshwater systems, following the isolation of the northern South American basin from western Amazonia. These findings support the hypothesis that range contraction of many taxa to their current distribution in northern South America occurred rapidly during at least the last 1.5 million years.

Overwintering behavior reduces mortality for a terrestrial turtle in forests managed with prescribed fire

Prescribed fire is an essential management practice in pyrogenic ecosystems, but fire can also be a significant disturbance and source of mortality for both target and non-target species. Seasonal periods of animal inactivity may provide opportunities to design burn plans that minimize negative impacts to species of conservation concern, but few studies have rigorously examined this possibility. Using radiotelemetry, we studied overwintering behavior and interactions with fire in a forest-dwelling terrestrial turtle, the Eastern Box Turtle (Terrapene carolina carolina), over an eight-year period at two sites that use prescribed fire in forest management. Turtles at both sites selected predominantly hardwood forests and mesic habitats and avoided upland pine forests. Turtles buried deepest (2.9 – 3.2 cm) below the soil-litter interface in late February and then moved gradually shallower until emergence in early April. Emergence timing varied over a 58-day period, but was consistent within individuals from year to year. Turtles also maintained fidelity to refuge locations, but those overwintering in burned areas selected sites over twice as far from refuges used in previous years compared to those in unburned areas. The areas and habitats selected by turtles during winter served as refugia from fire, and those whose refuges did burn remained buffered from lethal temperatures even at shallow burial depths. The only fire-related injury or mortality occurred during seasons of surface activity. Timing burning and other forest management practices during periods of winter dormancy may thus minimize threats to turtle populations, but modifications to prescribed fire regimes must also be balanced with other management objectives.

The freshwater and terrestrial turtles from Monte Pila and Fuenmayor (La Rioja, northern Spain): new data on the lower Miocene turtle diversity of the Ebro Basin

Abundant turtle fossils from two lower Miocene localities in which this lineage had not previously been studied are described herein. The localities are Monte Pila and Fuenmayor, which are 10 km apart, both located in the Autonomous Community of La Rioja (northern Spain), in the Ebro Basin. Information about the turtles from the lower Miocene levels in this basin was until now restricted to the finds made in the MN3 biozone of the Bardenas Reales of Navarre. Several turtle taxa had been recognized there. However, the material attributed to each of them was scarce. Monte Pila and Fuenmayor are located in biozone MN2. The remains of turtles are abundant in both sites. A terrestrial lineage, recognized at Monte Pila, is not represented in the Bardenas Reales fauna of Navarre. It represents the first record of TitanochelonPérez-García & Vlachos, 2014 in La Rioja and is one of the oldest member of this genus, being assigned to Titanochelon cf. bolivari (Hernández-Pacheco, 1917). The only lineage of turtles identified in Fuenmayor, and the most abundant in Monte Pila, is Ptychogasterinae. All material of this clade of freshwater turtles identified there is compatible with a single form, for which a high range of intraspecific variability is recognized. It is attributed to Ptychogaster (Temnoclemmys) cf. bardenensis Murelaga, Lapparent de Broin, Pereda Suberbiola & Astibia, 1999, this species having been recognized, until now, as exclusive to the Bardenas Reales of Navarre.

The ecomorphology of the shell of extant turtles and its applications for fossil turtles.

Turtles are a successful clade of reptiles that originated in the Late Triassic. The group adapted during its evolution to different types of environments, ranging from dry land to ponds, rivers, and the open ocean, and survived all Mesozoic and Cenozoic extinction events. The body of turtles is characterized by a shell, which has been hypothesized to have several biological roles, like protection, thermal and pH regulation, but also to be adapted in its shape to the ecology of the animal. However, only few studies have investigated the relationships between shell shape and ecology in a global context or clarified if shape can be used to diagnose habitat preferences in fossil representatives. Here, we assembled a three-dimensional dataset of 69 extant turtles and three fossils, in particular, the Late Triassic Proganochelys quenstedtii and Proterochersis robusta and the Late Jurassic Plesiochelys bigleri to test explicitly for a relationship between shell shape and ecology. 3D models were obtained using surface scanning and photogrammetry. The general shape of the shells was captured using geometric morphometrics. The habitat ecology of extant turtles was classified using the webbing of their forelimbs as a proxy. Principal component analysis (PCA) highlights much overlap between habitat groups. Discriminant analyses suggests significant differences between extant terrestrial turtles, extant fully aquatic (i.e., marine and riverine) turtles, and an unspecialized assemblage that includes extant turtles from all habitats, mostly freshwater aquatic forms. The paleoecology of the three fossil species cannot be determined with confidence, as all three fall within the unspecialized category, even if Plesiochelys bigleri plots closer to fully aquatic turtles, while the two Triassic species group closer to extant terrestrial forms. Although the shape of the shell of turtles indeed contains an ecological signal, it is overall too weak to uncover using shell shape in paleoecological studies, at least with the methods we selected.

Neuroanatomy of the European uppermost Cretaceous stem turtle Kallokibotion bajazidi

Kallokibotion bajazidi is one of the few stem turtles identified in the Upper Cretaceous of Europe. This form, exclusive to Maastrichtian outcrops of the Hațeg Basin (Romania), was described almost a century ago by the famous paleontologist Franz Nopcsa. However, it was identified as an enigmatic taxon until the end of the 20th century due to the limited available information. Recent studies have significantly improved knowledge about this taxon, confirming its attribution to a stem turtle, and being proposed as the sister taxon of the crown Testudines. The first three-dimensional reconstruction of the skull of this bizarre turtle is presented here. It has been made from the best-preserved skull so far known, the information provided by this specimen being complemented by that of an unpublished basicranium, which is analyzed in detail. This study allows to improve knowledge about the cranial osseous anatomy of the species, its phylogenetical identification as form closely related with the crown Testudines being supported. Furthermore, its neuronanatomical study is carried out. Detailed neuroanatomical analyzes on a very limited number of stem turtles have so far been performed. Considering the phylogenetic position of Kallokibotion bajazidi, the comparison of characters linked to the endocast, cranial nerves, nasal cavities, inner ears and circulatory system with those of these forms, as well as with those of members of the stem group, is carried out. Implications relative to the sensory capabilities interpreted for Kallokibotion bajazidi are discussed, the hitherto poorly supported hypothesis regarding its identification as a terrestrial turtle being supported.

The mid holocene occupation of the Pearl Islands: A case of unusual insular adaptations on the Pacific Coast of Panama

A survey of the Pearl Island archipelago discovered a Preceramic shell-bearing midden on the island of Pedro González located in Panama Bay in the eastern tropical Pacific Ocean. This occupation dates between ca. 6200 and 5600 cal yr BP, and revealed a mixed marine and terrestrial economy including the exploitation of sea and terrestrial turtles, snakes, iguanas, dolphins, and land mammals such as dwarf deer. Test excavations in 2007–2010 provided well-preserved invertebrate and vertebrate samples, which were considerably more diverse taxonomically for fish than for other vertebrate categories. At that time, only boa among a large snake sample was identified to species. A 7 × 4 m pit dug in 2015 increased the faunal sample but added little to diversity. Development of the ophidian comparative collection, augmented by on-line research, enhanced snake diversity and highlighted the dietary importance of snakes. Two marine turtle species – hawksbill and Pacific green– remain the only ones exploited. Improvements in stratigraphic sectioning have fine-tuned chronological fluctuations of faunal populations as in the case of green iguana and terrestrial mammals. Birds have experienced the greatest increase in diversity, but not as much as expected on the basis of the present-day island avifauna. Only two terrestrial mammal genera – squirrel, and rabbit – were added to the inventory inferred during 2007–2010, which comprised deer, opossum, monkey, agouti, paca, spiny rat, furry spiny rat, and probable capybara. The occupants of Pedro González took advantage of the abundant agate nodules eroding from the island's volcanic bedrock. This latest study reveals the presence of a substantial microlithic component that relied on the specialized heat treatment of whole nodules of lithic raw materials. As an idea to be explored further, we suggest that these microliths may have been produced by women for the manufacture and maintenance of composite cutting and/or grating tools used for the preparation of various foods. Ongoing analyses from the 2015 excavations are providing new information on the impact of these early colonists on the local fauna and have demonstrated how the physical properties of the island's lithic raw materials influenced their reduction strategies.

Main airway throughout the nasal cavity of green sea turtles is lined by keratinized stratified squamous epithelium

Structural and histological features of the nasal cavity of sea turtles are largely different from those of other Testudines species. The sea turtle nasal cavity is a pair of tubular structures with three diverticula and an excavation in the center, and three types of sensory epithelium are present in these four significant structures. To more clarify the adaptation of the nasal cavity to marine life style in sea turtles, non-sensory epithelium in the nasal cavity of green sea turtles (Chelonia mydas) were histologically determined from nostril to choanae in this study. Unlike many other animals including terrestrial turtles, the vestibular area and nasopharyngeal duct were all lined by keratinized stratified squamous epithelium. In the main nasal cavity, the margins of each sensory epithelium turned into respiratory epithelium with goblet cells, followed by keratinized stratified squamous epithelium. Keratinized epithelium appears more appropriate in sea turtle upper airway to protect against osmotic pressure when they release seawater through the nostrils, and thus this histological feature of upper airway might reflect adaptation to marine life style.

Surviving the Cretaceous-Paleogene mass extinction event: A terrestrial stem turtle in the Cenozoic of Laurasia

Findings of terrestrial stem turtles are not uncommon at Mesozoic continental sites in Laurasia, especially during the Upper Cretaceous. Thus, the record of several lineages is known in uppermost Cretaceous ecosystems in North America (Helochelydridae), Europe (Helochelydridae and Kallokibotion) and Asia (Sichuanchelyidae). No terrestrial stem turtle had been described in Laurasia after the Cretaceous-Paleogene mass extinction event. Thus, the only representatives described in the Cenozoic record worldwide corresponded to forms from southern Gondwana, where some of them survived until the Holocene. A bizarre terrestrial stem turtle from the upper Thanetian (upper Paleocene) of Europe is described here: Laurasichersis relicta gen. et sp. nov. Despite its discovery in France, in Mont de Berru (Marne), this Laurasian taxon is not recognized as a member of a European clade that survived the Cretaceous-Paleogene extinction event. It belongs to Sichuanchelyidae, a hitherto exclusively Asian Mesozoic group, known from the Middle Jurassic. Finds at the Belgian site of Hainin (Hainaut) show that this dispersion from Asia and the occupation of some niches previously dominated by European Mesozoic terrestrial stem forms had already taken place a few million years after the mass extinction event, at the end of the lower Paleocene.