Vincent Fernandez

Ontogeny of the Early Triassic Cynodont Thrinaxodon liorhinus (Therapsida): Dental Morphology and Replacement

ABSTRACT Here for the first time we use micro-computed tomography (µCT) scanning techniques to study dental replacement in the Early Triassic cynodont Thrinaxodon liorhinus.We analyzed five specimens ranging 37–87 mm in skull length using µCT scanning, which were supplemented by detailed anatomical analysis of 48 specimens with a basal skull length of 30–96 mm. Our results indicate that lower postcanines are more numerous and present amore complex morphology than the upper postcanines, even in the same individual; only the lower postcanines have more than three sectorial cusps and a cingular collar on the lingual margin. Complexity of the postcanines increases from the smallest individual to specimens with a skull length of 75 mm, but complexity decreases in larger specimens. Our results confirm the alternate replacement of the postcanines and the posterior migration of the postcanine series (including the loss without replacement of the anterior-most postcanines). Observations point to a posterior-to-anterior replacement wave in lower postcanines, but the evidence is not clear-cut for the upper series. The virtual extraction of functional and replacement teeth permitted us to conclude that in most of the cases the upper canines were replaced anteriorly, whereas lower canines were replaced posteriorly. The presence of two simultaneous replacements of the upper canine tooth was observed in two small juveniles, suggesting a higher rate of canine replacement at a younger age. Incisors also had a sequential replacement pattern, and more replacement teeth were present in medium-sized individuals.

Heart fossilization is possible and informs the evolution of cardiac outflow tract in vertebrates

Elucidating cardiac evolution has been frustrated by lack of fossils. One celebrated enigma in cardiac evolution involves the transition from a cardiac outflow tract dominated by a multi-valved conus arteriosus in basal actinopterygians, to an outflow tract commanded by the non-valved, elastic, bulbus arteriosus in higher actinopterygians. We demonstrate that cardiac preservation is possible in the extinct fish Rhacolepis buccalis from the Brazilian Cretaceous. Using X-ray synchrotron microtomography, we show that Rhacolepis fossils display hearts with a conus arteriosus containing at least five valve rows. This represents a transitional morphology between the primitive, multivalvar, conal condition and the derived, monovalvar, bulbar state of the outflow tract in modern actinopterygians. Our data rescue a long-lost cardiac phenotype (119-113 Ma) and suggest that outflow tract simplification in actinopterygians is compatible with a gradual, rather than a drastic saltation event. Overall, our results demonstrate the feasibility of studying cardiac evolution in fossils. DOI: http://dx.doi.org/10.7554/eLife.14698.001

Ontogeny of the Early Triassic Cynodont Thrinaxodon liorhinus (Therapsida): Cranial Morphology

The cranial morphology of 68 Thrinaxodon liorhinus specimens, ranging in size from 30 to 96 mm in basal skull length, is investigated using both qualitative and quantitative analyses. From this comprehensive survey, we determined that nine cranial features, including five in the temporal region, separated the sample into four ontogenetic stages. A bivariate analysis of 60 specimens indicated that the skull generally increased in size isometrically, with the exception of four regions. The orbit had negative allometry, a result consistent with other ontogenetic studies of tetrapods, whereas the length of the snout, palate, and temporal region showed positive allometry. The last trend had strong positive allometry indicating that during ontogeny the length of the sagittal crest increased at a much faster rate than the rest of the skull. The large number of changes in the temporal region of the skull of Thrinaxodon may indicate a greater development of the posterior fibres of the temporalis musculature from an early ontogenetic stage. For example, the posterior sagittal crest developed much earlier in ontogeny than the anterior crest that formed in adults, and bone was deposited dorsally creating a unified posterior sagittal crest rather than having a suture that spanned the entire depth of the skull roof. In combination with the isometric height of the zygomatic arch and the almost complete absence of the zygomatic arch angulation, these ontogenetic changes suggest that there was greater development of the temporalis relative to the masseter muscles, indicating a strong posterodorsal movement of the mandible in Thrinaxodon. Anat Rec, 298:1440–1464, 2015.

Homogenization of sample absorption for the imaging of large and dense fossils with synchrotron microtomography

Propagation phase-contrast synchrotron radiation microtomography (PPC-SRμCT) has proved to be very successful for examining fossils. Because fossils range widely in taphonomic preservation, size, shape and density, X-ray computed tomography protocols are constantly being developed and refined. Here we present a 1-h procedure that combines a filtered high-energy polychromatic beam with long-distance PPC-SRμCT (sample to detector: 4–16 m) and an attenuation protocol normalizing the absorption profile (tested on 13-cm-thick and 5.242 g cm−3 locally dense samples but applicable to 20-cm-thick samples). This approach provides high-quality imaging results, which show marked improvement relative to results from images obtained without the attenuation protocol in apparent transmission, contrast and signal-to-noise ratio. The attenuation protocol involves immersing samples in a tube filled with aluminum or glass balls in association with a U-shaped aluminum profiler. This technique therefore provides access to a larger dynamic range of the detector used for tomographic reconstruction. This protocol homogenizes beam-hardening artifacts, thereby rendering it effective for use with conventional μCT scanners.

Ecomorphological analysis of aerial performance in a non-specialized lacertid lizard, Holaspis guentheri.

SUMMARY Controlled aerial descent has evolved at least 30 times independently in different vertebrate and invertebrate lineages. A whole suite of morphological modifications, such as patagia, lateral skin folds and webbed feet, have been suggested to enhance descending ability. In this study, we compare aerial performance (i.e. vertical and horizontal velocity, horizontal distance covered, duration of descent) and morphology (body mass, body width, inter limb distance, surface area and wing loading) among three species of lizards, representing a range of aerial descenders present within the clade. Our performance measurements show that the lacertid Holaspis guentheri performs intermediately to the specialized gekkonid Ptychozoon kuhli and the rock-dwelling lizard Podarcis muralis. The small relative body mass of H. guentheri results in a low wing loading similar to that of P. kuhli thus enhancing its aerial performance. Whereas the latter generates great lift forces and is able to cover great horizontal distances, H. guentheris low wing loading seems to be responsible for a slow descent and low impact forces upon landing. Our results show that very small morphological changes may result in noticeable and ecologically relevant changes in performance.

Oxygen isotope composition of continental vertebrate apatites from Mesozoic formations of Thailand; environmental and ecological significance

Abstract Phosphatic remains (tooth enamel, turtle shell fragments and fish scales) of continental vertebrates (freshwater fish, crocodilians, turtles, and theropod and sauropod dinosaurs) recovered from eight localities of NE Thailand ranging in age from the Late Jurassic to the late Early Cretaceous have been analysed for their oxygen isotopic compositions (δ18Op). From these preliminary data, local meteoric water δ18Ow values estimated using δ18Op values of crocodilians and turtles range from −4.1±2‰ at the end of the Jurassic to −8.3±2‰ during the Early Cretaceous, suggesting a transition from dry to wetter climates with increasing amount of seasonal precipitation from several hundred millimetres per year to several thousand millimetres. Measurable offsets in δ18Op values observed between dinosaur taxa (the spinosaurid theropod Siamosaurus, other theropods and nemegtosaurid sauropods) are interpreted in terms of differences in water strategies, and suggest that Siamosaurus had habits similar to those of semi-aquatic vertebrates such as crocodilians or freshwater turtles.

A Re-Description of ‘Mycterosaurus’ smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids

‘Mycterosaurus’ smithae, from the Cisuralian (early Permian) of Colorado, was first described in 1965 as a second species of the genus Mycterosaurus. While the type species of this genus, M. longiceps, has been shown by multiple cladistic analyses to belong to the basal synapsid family Varanopidae, ‘M.’ smithae has been largely ignored since its original description. Additional preparation and synchrotron scanning has revealed new significant information that supports the assignment of this species to a new genus: Vaughnictis gen. nov. Vaughnictis lacks many of the characteristics of mycterosaurines and varanopids in general: it lacks the slender femur, the linguo-labially compressed and strongly recurved teeth, and the lateral boss on the postorbital characteristic of this family. Instead, it possesses coronoid teeth, a large supratemporal, and a large pineal foramen positioned midway along the length of the parietal, features that support its assignment to Eothyrididae. Moreover, the postcranium shares many characters with the eothyridid Oedaleops. An expanded version of a recently published phylogenetic analysis of pelycosaurian-grade synapsids positions Vaughnictis as the sister taxon of Eothyris within the clade Eothyrididae. The addition of data on the postcranium of eothyridids and the inclusion of the recently-described basal caseid Eocasea confirms the recently-disputed position of caseasaurs as the most basal synapsids. As the parsimony analysis produced low support values and a lack of resolution due to missing data, additional analyses were undertaken using Bayesian and Implied Weights methods, which produced better resolution and relationships with higher support values. While the results are similar, alternative positions for the enigmatic Moscovian age (Carboniferous) synapsid Echinerpeton are suggested by Bayesian analysis; the parsimony analysis found it to be an ophiacodontid, while the Bayesian and Implied Weights analysis found it to be the sister to the Sphenacomorpha.

Cranial bosses of choerosaurus dejageri (therapsida, therocephalia): Earliest evidence of cranial display structures in eutheriodonts.

Choerosaurus dejageri, a non-mammalian eutheriodont therapsid from the South African late Permian (~259 Ma), has conspicuous hemispheric cranial bosses on the maxilla and the mandible. These bosses, the earliest of this nature in a eutheriodont, potentially make C. dejageri a key species for understanding the evolutionary origins of sexually selective behaviours (intraspecific competition, ritualized sexual and intimidation displays) associated with cranial outgrowths at the root of the clade that eventually led to extant mammals. Comparison with the tapinocephalid dinocephalian Moschops capensis, a therapsid in which head butting is strongly supported, shows that the delicate structure of the cranial bosses and the gracile structure of the skull of Choerosaurus would be more suitable for display and low energy combat than vigorous head butting. Thus, despite the fact that Choerosaurus is represented by only one skull (which makes it impossible to address the question of sexual dimorphism), its cranial bosses are better interpreted as structures involved in intraspecific selection, i.e. low-energy fighting or display. Display structures, such as enlarged canines and cranial bosses, are widespread among basal therapsid clades and are also present in the putative basal therapsid Tetraceratops insignis. This suggests that sexual selection may have played a more important role in the distant origin and evolution of mammals earlier than previously thought. Sexual selection may explain the subsequent independent evolution of cranial outgrowths and pachyostosis in different therapsid lineages (Biarmosuchia, Dinocephalia, Gorgonopsia and Dicynodontia).

Synchrotron scanning reveals the palaeoneurology of the head-butting Moschops capensis (Therapsida, Dinocephalia)

Dinocephalian therapsids are renowned for their massive, pachyostotic and ornamented skulls adapted for head-to-head fighting during intraspecific combat. Synchrotron scanning of the tapinocephalid Moschops capensis reveals, for the first time, numerous anatomical adaptations of the central nervous system related to this combative behaviour. Many neural structures (such as the brain, inner ear and ophthalmic branch of the trigeminal nerve) were completely enclosed and protected by bones, which is unusual for non-mammaliaform therapsids. The nearly complete ossification of the braincase enables precise determination of the brain cavity volume and encephalization quotient, which appears greater than expected for such a large and early herbivore. The practice of head butting is often associated with complex social behaviours and gregariousness in extant species, which are known to influence brain size evolution. Additionally, the plane of the lateral (horizontal) semicircular canal of the bony labyrinth is oriented nearly vertically if the skull is held horizontally, which suggests that the natural position of the head was inclined about 60–65°to the horizontal. This is consistent with the fighting position inferred from osteology, as well as ground-level browsing. Finally, the unusually large parietal tube may have been filled with thick conjunctive tissue to protect the delicate pineal eye from injury sustained during head butting.

Hyperspecialization in Some South American Endemic Ungulates Revealed by Long Bone Microstructure

Adaptation to aquatic or semi-aquatic habits has been demonstrated in several distantly related mammalian clades worldwide during the Cenozoic, but curiously none has been conclusively evidenced inland for South America although a few South American native ungulates (SANU) have been proposed as possible subaquatic taxa. These taxa mostly correspond to large-sized herbivorous forms among astrapotheres, pyrotheres, and some toxodontid notoungulates, found at the end of the Paleogene and/or beginning of the Neogene. As no clear argument was provided for these ecological hypotheses, an analysis of the microanatomical features of long bones of some of these taxa in a comparative context was conducted in order to address the question of the paleoecology of these organisms. Our study highlights a variety of osseous specializations in the stylopod bones of SANU, and notably that Parastrapotherium, Pyrotherium, and Nesodon are affected by bone mass increase. If the microanatomical features of Parastrapotherium and Nesodon evoke what is observed in some terrestrial or semi-aquatic graviportal taxa, the very high compactness of Pyrotherium suggests extreme functional requirements in this taxon such as a graviportal hyperspecialization. This study thus evidences the occurrence of several convergent adaptations to graviportality and/or subaquatic habits within SANU and contributes to an important step towards a better integration of South American endemic mammals in large-scaled paleoecological studies.