Natural external plastron mold of the Triassic turtle Proterochersis: An unusual mode of preservation.

Impressions of vertebrate bodies or their parts, such as trace fossils and natural molds of bones, are a valuable source of information about ancient faunas which may supplement the standard fossil record based on skeletal elements. Whereas trace fossils of animal activity are relatively common and actively studied within the field of ichnology, and natural impressions of internal or external surfaces are a frequent preservation mode in fossil invertebrates, natural molds of bones are comparatively rare and less extensively documented and discussed. Among them, internal molds (steinkerns) of turtle shells are a relatively well-known form of preservation, but the mechanisms and taphonomic prerequisites leading to their formation are poorly studied. External shell molds are even less represented in the literature. Herein, we describe a historic specimen of a natural external turtle plastron mold from the Triassic (Norian) Löwenstein Formation of Germany-a formation which also yielded a number of turtle steinkerns. The specimen is significant not only because it represents an unusual form of preservation, but also due to its remarkably large size and the presence of a potential shell pathology. Although it was initially interpreted as Proterochersis sp., the recent progress in the knowledge of proterochersid turtles leading to an increase in the number of known taxa within that group allows us to verify that assessment. We confirm that the specimen is morphologically consistent with the genus and tentatively identify it as Proterochersis robusta, the only representative of that genus from the Löwenstein Formation. We note, however, that its size exceeds the size observed thus far in Proterochersis robusta and fits within the range of Proterochersis porebensis from the Grabowa Formation of Poland. The marks interpreted as shell pathology are morphologically consistent with Karethraichnus lakkos-an ichnotaxon interpreted as a trace of ectoparasites, such as leeches. This may support the previously proposed interpretation of Proterochersis spp. as a semiaquatic turtle. Moreover, if the identification is correct, the specimen may represent a very rare case of a negative preservation of a named ichnotaxon. Finally, we discuss the taphonomy of the Löwenstein Formation turtles in comparison with other Triassic turtle-yielding formations which show no potential for the preservation of internal or external shell molds and propose a taphonomic model for the formation of such fossils.

Digital image processing: A new tool for morphological measurements of freshwater turtles under rehabilitation.

Freshwater fauna is facing an uphill task for survival in the Ganga Basin, India, due to a range of factors causing habitat degradation and fragmentation, necessitating conservation interventions. As part of the ongoing efforts to conserve the freshwater fauna of the Basin, we are working on rehabilitating rescued freshwater chelonians. We carry out various interventions to restore rescued individuals to an apparent state of fitness for their release in suitable natural habitats. Morphometric measurements are crucial to managing captive wild animals for assessing their growth and well-being. Measurements are made using manual methods like vernier caliper that are prone to observer error experience and require handling the specimens for extended periods. Digital imaging technology is rapidly progressing at a fast pace and with the advancement of technology. We acquired images of turtles using smartphones along with manual morphometric measurements using vernier calipers of the straight carapace length and straight carapace width. The images were subsequently processed using ImageJ, a freeware and compared with manual morphometric measurements. A significant decrease in the time spent in carrying out morphometric measurements was observed in our study. The difference in error in measurements was, however, not significant. A probable cause for this may have been the extensive experience of the personnel carrying out the measurements using vernier caliper. Digital image processing technology can cause a significant reduction in the stress of the animals exposed to handling during measurements, thereby improving their welfare. Additionally, this can be used in the field to carry out morphometric measurements of free-ranging individuals, where it is often difficult to capture individuals, and challenges are faced in obtaining permission to capture specimens.

Shell shape does not accurately predict self-righting ability in hatchling freshwater turtles.

Flat hydrodynamic shells likely represent an evolutionary trade-off between adaptation to an aquatic lifestyle and the instability of more rounded shells, thought beneficial for self-righting. Trade-offs often result in compromises, this is particularly true when freshwater turtles, with flatter shells, must self-right to avoid the negative effects of inverting. These turtles, theoretically, invest more biomechanical effort to achieve successful and timely self-righting when compared to turtles with rounded carapaces. This increase in effort places these hatchlings in a precarious position; prone to inversion and predation and with shells seemingly maladapted to the act of self-righting. Here, we examine hatchling self-righting performance in three morphologically distinct freshwater turtle species (Apalone spinifera, Chelydra serpentina and Trachemys scripta scripta) that inhabit similar environmental niches. We demonstrate that these hatchlings were capable of rapid self-righting and used considerably less biomechanical effort relative to adult turtles. Despite differences in shell morphology the energetic efficiency of self-righting remained remarkably low and uniform between the three species. Our results confound theoretical predictions of self-righting ability based on shell shape metrics and indicate that other morphological characteristics like neck or tail morphology and shell material properties must be considered to better understand the biomechanical nuances of Testudine self-righting.

Turtle skull development unveils a molecular basis for amniote cranial diversity.

Amniote skulls display diverse architectural patterns including remarkable variations in the number of temporal arches surrounding the upper and lower temporal fenestrae. However, the cellular and molecular basis underlying this diversification remains elusive. Turtles are a useful model to understand skull diversity due to the presence of secondarily closed temporal fenestrae and different extents of temporal emarginations (marginal reduction of dermal bones). Here, we analyzed embryos of three turtle species with varying degrees of temporal emargination and identified shared widespread coexpression of upstream osteogenic genes Msx2 and Runx2 and species-specific expression of more downstream osteogenic genes Sp7 and Sparc in the head. Further analysis of representative amniote embryos revealed differential expression patterns of osteogenic genes in the temporal region, suggesting that the spatiotemporal regulation of Msx2, Runx2, and Sp7 distinguishes the temporal skull morphology among amniotes. Moreover, the presence of Msx2- and/or Runx2-positive temporal mesenchyme with osteogenic potential may have contributed to their extremely diverse cranial morphology in reptiles.

A new specimen of Solnhofia parsonsi from the Upper Jurassic (Kimmeridgian) Plattenkalk deposits of Painten (Bavaria, Germany) and comments on the relationship between limb taphonomy and habitat ecology in fossil turtles.

The limestones of the Solnhofen area in southern Germany are one of the most important fossil Lagerstätten from the entire Mesozoic era, especially famous for the exquisitely preserved vertebrates. The turtles from the Solnhofen Limestone have been always of special interest because they include some of the best-preserved specimens from the Mesozoic. Here, we describe a new turtle specimen from the Torleite Formation (Kimmeridgian) of Painten and refer it to the thalassochelydian turtle Solnhofia parsonsi based on the presence of a unique combination of characters. The far majority of morphological differences from previously published specimens can be explained by ontogeny as the new specimen represents a larger, more ossified, and presumably older individual. Additionally, the specimen from Painten is the first described specimen of S. parsonsi preserving the largely complete and articulated limbs, the preservation of which indicates that the taxon did not possess stiffened paddles present in more pelagic marine turtles and is consistent with a previously inferred nearshore marine lifestyle. Contrary to previous inferences, we argue that taphonomic preservation of digits in articulated fossil turtles from laminated deposits cannot be used alone to infer marine or freshwater habitat. Finally, the new specimen from Painten is only the second, for which detailed information on its stratigraphic position and locality of origin are known.

Soft-robotic green sea turtle (Chelonia mydas) developed to replace animal experimentation provides new insight into their propulsive strategies.

Green sea turtles (Chelonia mydas) can swim up to 50 km per day while only consuming seagrass or microalgae. How the animal accomplishes this vast journey on such low energy intake points to the effectiveness of their swimming technique and is a testament to the power of evolution. Understanding the green sea turtle's ability to accomplish these journeys requires insight into their propulsive strategies. Conducting animal testing to uncover their propulsive strategies brings significant challenges: firstly, the ethical issues of conducting experiments on an endangered animal, and secondly, the animal may not even swim with its regular routine during the experiments. In this work, we develop a new soft-robotic sea turtle that reproduces the real animal's form and function to provide biomechanical insights without the need for invasive experimentation. We found that the green sea turtle may only produce propulsion for approximately 30% of the limb beat cycle, with the remaining 70% exploiting a power-preserving low-drag glide. Due to the animal's large mass and relatively low drag coefficient, losses in swim speed are minimal during the gliding stage. These findings may lead to the creation of a new generation of robotic systems for ocean exploration that use an optimised derivative of the sea turtle propulsive strategy.

A new species of Nanhsiungchelys (Testudines: Cryptodira: Nanhsiungchelyidae) from the Upper Cretaceous of Nanxiong Basin, China.

Nanhsiungchelyidae are a group of large turtles that lived in Asia and North America during the Cretaceous. Here we report a new species of nanhsiungchelyid, Nanhsiungchelys yangi sp. nov., from the Upper Cretaceous of Nanxiong Basin, China. The specimen consists of a well-preserved skull and lower jaw, as well as the anterior parts of the carapace and plastron. The diagnostic features of Nanhsiungchelys include a large entire carapace length (∼55.5 cm), a network of sculptures consisting of pits and ridges on the surface of the skull and shell, shallow cheek emargination and temporal emargination, deep nuchal emargination, and a pair of anterolateral processes on the carapace. However, Nanhsiungchelys yangi differs from the other species of Nanhsiungchelys mainly in having a triangular-shaped snout (in dorsal view) and wide anterolateral processes on the carapace. Additionally, some other characteristics (e.g., the premaxilla is higher than wide, the maxilla is unseen in dorsal views, a small portion of the maxilla extends posterior and ventral of the orbit, and the parietal is bigger than the frontal) are strong evidence to distinguish Nanhsiungchelys yangi from Nanhsiungchelys wuchingensis. A phylogenetic analysis of nanhsiungchelyids places Nanhsiungchelys yangi and Nanhsiungchelys wuchingensis as sister taxa. Nanhsiungchelys yangi and some other nanhsiungchelyids bear distinct anterolateral processes on the carapace, which have not been reported in any extant turtles and may have played a role in protecting the head. The Nanxiong Basin was extremely hot during the Late Cretaceous, and so we suggest that nanhsiungchelyids might have immersed themselves in mud or water to avoid the heat, similar to some extant tortoises. If they were capable of swimming, our computer simulations of fluid flow suggest the anterolateral processes could have reduced drag during locomotion.

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.

New findings on the evolution of turtles: A symposium in honor of Marcelo S. de la Fuente.

With their particular body plan within amniotes and their amazing fossil record, turtles represent a great interest for both neontologists and paleontologists with a strong anatomical background. The Turtle Evolution Symposia are regular international meetings that gather scientists working with different aspects related to the evolutionary history of turtles, from their origin and early evolution until recent times. The latest edition of the Turtle Evolution Symposium was organized in 2021 amidst the COVID-19 outbreak and held virtually from the facilities of the Museo Paleontológico Egidio Feruglio in Trelew (Patagonia, Chubut, Argentina). More than 75 scientists from 25 countries presented their latest advances on topics related to turtle evolution, some of which are published in this Special Volume of The Anatomical Record. Both the Turtle Evolution Symposium 2021 and this Special Volume are dedicated to Marcelo S. de la Fuente who was the first researcher who specialized in the study of extinct turtles in South America, and his studies have an important regional and international impact.

Histological features and Gαolf expression patterns in the nasal cavity of sea turtles.

Sea turtles use olfaction to detect volatile and water-soluble substances. The nasal cavity of green turtles (Chelonia mydas) comprises morphologically defined the anterodorsal, anteroventral, and posterodorsal diverticula, as well as a single posteroventral fossa. Here, we detailed the histological features of the nasal cavity of a mature female green turtle. The posterodorsal diverticulum contained spongy-like venous sinuses and a wave-shaped sensory epithelium that favored ventilation. Secretory structures that were significant in sensory and non-sensory epithelia were probably involved in protection against seawater. These findings suggested that green turtles efficiently intake airborne substances and dissolve water-soluble substances in mucous, while suppressing the effects of salts. In addition, positive staining of Gαs/olf that couples with olfactory, but not vomeronasal, receptors was predominant in all three types of sensory epithelium in the nasal cavity. Both of airborne and water-soluble odorants seemed to be detected in cells expressing Gαolf and olfactory receptors.

A well-preserved cranium from the Judith River Formation (Montana, USA) reveals the inner ear and neuroanatomy of a Campanian baenid turtle.

A cranium belonging to a baenid turtle was recently recovered from the lower half of the Judith River Formation, Montana. Badlands Dinosaur Museum (BDM) 004 is a well-preserved partial cranium that includes the posterior cranial vault, cranial base, and otic capsules. Based on diagnostic characters, the skull can be attributed to Plesiobaena antiqua, which has been previously reported from the Judith River Formation. It also shares with palatobaenines projecting posterior processes of the tubercula basioccipitale and a prominent condylus occipitalis with a deep central pit, demonstrating variation within the Pl. antiqua hypodigm. In a phylogenetic analysis, an operational taxonomic unit of BDM 004 was positioned within Baenodda in an unresolved polytomy with Pl. antiqua, Edowa zuniensis, Palatobaeninae, and Eubaeninae. Microcomputed tomographic (µCT) scans revealed morphology of the middle and inner ear and endocast that are largely unknown in baenids. Semicircular canals of BDM 004 are virtually identical to those of Eubaena cephalica and consistent in dimensions to those of other turtle taxa, including anterior and posterior semicircular canals that are robust and taller than the common crus and diverge from each other at an angle of approximately 90°. The digital endocast reveals a moderately flexed brain with rounded cerebral hemispheres and minimal separation between the metencephalon and myelencephalon. Its well-preserved columella auris (stapes) is gracile with a posterodorsally flared basis columella. It arcs across the middle ear and flattens near its terminus. This study adds to the understanding of baenid middle and inner ear and neuroanatomical morphology and expands the morphological understanding of Pl. antiqua.

Morphological tongue and palate characterizations in Trachemys adiutrix (Vanzolini, 1995) turtles.

Morphological studies on the oropharyngeal cavity of turtles are an interesting tool in understanding evolutionary processes associated with feeding habits. There is paucity of morphological information on the oropharyngeal cavity of the Trachemys adiutrix turtle. Thus, the aim of this study was to describe the tongue and palate morphology of T. adiutrix to establish a standard model for the species, providing information that may improve knowledge on the species feeding habits. Gross dissection, light microscopy and scanning electron microscopy assessments of 11 specimens were carried out. The tongue of T. adiutrix is triangular with a slightly rounded apex and broad base and lingual papillae widely distributed throughout the entire dorsal surface. The palate is composed of bony structures, displaying a triangular apex with one pair of choanae and palatine raphe that begin in the middle region, dividing the two openings. The body was elongated and smooth. The root was wrinkled and opened into the oesophagus. The tongue was lined by non-keratinized stratified squamous epithelium, rich in mucous cells and a large number of different-shaped papillae and taste buds. The palate was mainly lined by a pseudostratified cylindrical ciliated epithelium rich in mucous cells. In conclusion, the tongue and palate characteristics of T. adiutrix include evidence that these animals are semiaquatic, displaying morphological characteristics associated with aquatic and terrestrial trophic ecology, similar to that observed in other semiaquatic and terrestrial chelonian species.

A systematic compendium of turtle mandibular anatomy using digital dissections of soft tissue and osteology.

Turtles are a charismatic reptile group with a peculiar body plan, which most notably includes the shell. Anatomists have often focused descriptive efforts on the shell and other strongly derived body parts, such as the akinetic skull, or the cervical vertebrae. Other parts of turtle osteology, like the girdles, limbs, and mandibles, are documented with less rigor and detail. The mandible is the primary skeletal element involved in food acquisition and initial food processing of turtles, and its features are thus likely linked to feeding ecology. In addition, the mandible of turtles is composed of up to seven bones (sometimes fused to as little as three) and has thus anatomical complexity that may be insightful for systematic purposes and phylogenetic research. Despite apparent complexity and diversity to the mandible of turtles, this anatomical system has not been systematically studied, not even in search of characters that might improve phylogenetic resolution. Here, we describe the mandibular osteology for all major subclades of extant turtles with the help of digitally dissected 3D models derived from high-resolution computed tomography (µCT) scans of 70 extant species. We provide 31 fully segmented mandibles, as well as 3D models of the mandibular musculature, innervation, and arterial circulation of the cryptodire Dermatemys mawii. We synthesize observed variation into 51 morphological characters, which we optimize onto a molecular phylogeny. This analysis shows some mandibular characters to have high systematic value, whereas others are highly homoplastic and may underlie ecological influences or other factors invoking variation.

Skeletal repatterning enhances the protective capacity of the shell in African hinge-back tortoises (Kinixys).

Changes in the structural association of skeletal traits are crucial to the evolution of novel forms and functions. In vertebrates, such rearrangements often occur gradually and may precede or coincide with the functional activation of skeletal traits. To illustrate this process, we examined the ontogeny of African hinge-back tortoises (Kinixys spp.). Kinixys species feature a moveable "hinge" on the dorsal shell (carapace) that enables shell closure (kinesis) when the hind limbs are withdrawn. This hinge, however, is absent in juveniles. Herein, we describe how this unusual phenotype arises via alterations in the tissue configuration and shape of the carapace. The ontogenetic repatterning of osseous and keratinous tissue coincided with shifts in morphological integration and the establishment of anterior (static) and posterior (kinetic) carapacial modules. Based on ex vivo skeletal movement and raw anatomy, we propose that Kinixys employs a "sliding hinge" shell-closing system that overcomes thoracic rigidity and enhances the protective capacity of the carapace. Universal properties of the vertebrate skeleton, such as plasticity, modularity, and secondary maturation processes, contributed to adaptive evolutionary change in Kinixys. We discuss a hypothetical model to explain the delayed emergence of skeletal traits and its relevance to the origins of novel form-to-function relationships.

A new large-sized species of Chelonoidis (Testudinidae) without gibbosities from the middle Miocene of Aguada Escondida (NW Chubut, Patagonia, Argentina).

Previously, only one small-sized species of Testudinidae (Chelonoidis gringorum) was named from Lower-Middle Miocene of Central Patagonia. In this short article, we describe a new large Testudinidae, here named Chelonoidis meridiana sp. nov. This large new species (carapace up to 80 cm) differs from other large species of the southern South American fossil record by the absence of gibbosities on neural and costal carapacial bones, a short and wide entoplastron with a humeropectoral sulcus that was placed well behind its posterior end, and pectoral scutes that are medially longer, approximately one-third of the medial length of the hyoplastra. The discovery of this new species in the Middle Miocene increases the raw turtle diversity in Chubut, permitting to reduce the gap between the raw and phylogenetic diversity previously proposed. This occurrence is also important from both a temporal and geographic point of view. It permits fixing the presence of large-sized tortoises in the continent since ~15 million years ago (Middle Miocene). This occurrence predates all other large and giant tortoises from Argentina and South America and it is older than the giant Chelonoidis from the Langhian-Tortonian of Colombia. This is also the southernmost occurrence of a large-sized testudinid in the world.

The latest Miocene to Pliocene small-sized fossil turtles, Testudinidae and Geoemydidae, from Makrygialos, Thermaikos Gulf, Northern Greece.

Although the existence of fossil turtles in Makrygialos and the surrounding areas, on the west side of the Thermaikos Gulf in Northern Greece, has been known for almost two decades, it has not been studied so far in detail. Over the last 5 years, we have conducted systematic fieldwork and exploration, focusing on the discovery and recovery of fossils in the coastal broader area of Makrygialos. Eleven turtle specimens have been found so far, eight of which have been collected, on the outcrops of the beaches of Makrygialos and Agiannis. Their age is estimated between the latest Miocene and Pliocene. Four of these specimens are terrestrial giant tortoises, two of them are small tortoises which we identify as the species Chersine hermanni sensu lato, and the last two are freshwater turtles which we attribute to the genus Mauremys sensu lato. Here, we present our studies on the small-sized turtle species from this region. The fossil turtle fauna of Makrygialos is among the richest and most diverse in Greece, as in only a few other Greek localities at least three species of fossil turtles have been found. The C. hermanni specimens found in Makrygialos are probably the oldest fossils of the species found in Europe. The small-turtle faunal composition during the Neogene along the east and west sides of the Thermaikos Gulf was completely different.

Morphological changes in the digestive tract of the Chinese soft-shelled turtle (Pelodiscus sinensis) during embryonic development.

The digestive tract development of the Pelodiscus sinensis embryo is described through the observation of the embryonic morphology on hematoxylin and eosin stained tissue sections. During the first 9 days of embryonic development, the anterior intestine of the embryo divides into the oral cavity, pharyngeal cavity, esophagus, stomach, and small intestine, while the caudal intestine differentiates into the cloaca, the anterior and caudal tubes of the large intestine. Between days 10-24, the wall of the digestive tract forms a two-layer structure consisting of mucosa and submucosa. The endoderm evolves into epithelial tissue in each part of the digestive tract, the mesoderm goes from a dense cluster of cells to looser mesenchymal tissue then divides into loose connective tissue, mesothelium, and muscle tissue. There is no clear temporal boundary between development of mesenchymal tissue and the early loose connective tissue, which is a gradual process.

Ludwig Heinrich Bojanus and the anatomy of the European pond turtle: Facts, fiction, and future.

Detailed anatomical knowledge is a prerequisite for studying a variety of biological questions. This holds true particularly for the evolutionary realm, in which fossils play an important role. Putting paleontological data in perspective consequently requires a profound knowledge of the corresponding structures of the extant representatives of a given lineage as well. Numerous morphological studies have contributed to our understanding of chelonian anatomy over the past centuries, but one of the earliest ones-whose completion celebrated its bicentenary with this symposium-still stands out among all of them: Anatome Testudinis Europaeae, published by Ludwig Heinrich Bojanus (1776-1827) in two installments in 1819 and 1821. Numerous myths around this publication have evolved ever since. The intent of this article is to set the record straight for some of them, and, based on personal inspection of multiple copies, to discuss currently unrecognized aspects about the existence of exceptional colored variant editions. This ends in a proposal for how this work could be used even more effectively in its upcoming third century of existence. Last but not least and despite the controversies related to its bibliographical aspects, one fact remains an unchallenged reality: Anatome Testudinis Europaeae, even 200 years after its first publication, still ranks very high among the most detailed and accurate anatomical works on a single chelonian-and for that matter reptilian-species ever produced and has lost absolutely none of its value for modern-day science.

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.