Reassessing trauma in a Late Neolithic cranium from Megalitho da Capella (Figueira da Foz, Portugal).

The Megalitho da Capella (Figueira da Foz, Coimbra, Portugal) is one of at least 21 dolmens in a megalithic complex explored by António dos Santos Rocha between 1880 and 1909. Among the human remains from Megalitho da Capella is an incomplete and fragmented cranium. Santos Rocha interpreted a groove on the parietal bone as an incision of traumatic origin with signs of remodeling that was suggestive of prolonged survival after an intentional intervention. This study provides a new examination of the groove using microcomputed tomographic (microCT) imaging, microscopy, and macroscopic observations of the groove in addition to the direct dating the skeletal remains. Results indicate that the human remains are dated to the Late Neolithic and that the "incision" is a normal anatomical variant corresponding to impressions from vascular tissue and temporal projections of the squamosal. We conclude that studies of bone surface modifications should consider normal anatomical variants (e.g., sulci, grooves, and furrows) when reporting results. Paleoimaging, microscopy, and comparative observations can assist in the identification bone modifications versus anatomical variants.

A conserved tooth resorption mechanism in modern and fossil snakes.

Whether snakes evolved their elongated, limbless bodies or their specialized skulls and teeth first is a central question in squamate evolution. Identifying features shared between extant and fossil snakes is therefore key to unraveling the early evolution of this iconic reptile group. One promising candidate is their unusual mode of tooth replacement, whereby teeth are replaced without signs of external tooth resorption. We reveal through histological analysis that the lack of resorption pits in snakes is due to the unusual action of odontoclasts, which resorb dentine from within the pulp of the tooth. Internal tooth resorption is widespread in extant snakes, differs from replacement in other reptiles, and is even detectable via non-destructive µCT scanning, providing a method for identifying fossil snakes. We then detected internal tooth resorption in the fossil snake Yurlunggur, and one of the oldest snake fossils, Portugalophis, suggesting that it is one of the earliest innovations in Pan-Serpentes, likely preceding limb loss.

A new Early Cretaceous lizard in Myanmar amber with exceptionally preserved integument.

We here report on a well-preserved juvenile lizard specimen in Albian amber (ca. 110 mya) from the Hkamti site (Myanmar). This new taxon is represented by an articulated skull and the anterior portion of the trunk, including the pectoral girdle and forelimbs. The scleral ossicles and eyelid are also visible, and the specimen exhibits pristine detail of the integument (of both head and body). In a combined molecular and morphological analysis, it was consistently recovered as a scincoid lizard (Scinciformata), as sister to Tepexisaurus + Xantusiidae. However, the phylogenetic position of the new taxon should be interpreted with caution as the holotype is an immature individual. We explored the possibility of miscoding ontogenetically variable characters by running alternative analyses in which these characters were scored as missing data for our taxon. With the exception of one tree, in which it was sister to Amphisbaenia, the specimen was recovered as a Pan-xantusiid. Moreover, we cannot rule out the possibility that it represents a separate lineage of uncertain phylogenetic position, as it is the case for many Jurassic and Cretaceous taxa. Nonetheless, this fossil offers a rare opportunity to glimpse the external appearance of one group of lizards during the Early Cretaceous.

Neuroanatomy of the nodosaurid Struthiosaurus austriacus (Dinosauria: Thyreophora) supports potential ecological differentiations within Ankylosauria.

Nodosauridae is a group of thyreophoran dinosaurs characterized by a collar of prominent osteoderms. In comparison to its sister group, the often club-tailed ankylosaurids, a different lifestyle of nodosaurids could be assumed based on their neuroanatomy and weaponry, e.g., regarding applied defensive strategies. The holotype of the nodosaurid Struthiosaurus austriacus consists of a single partial braincase from the Late Cretaceous of Austria. Since neuroanatomy is considered to be associated with ecological tendencies, we created digital models of the braincase based on micro-CT data. The cranial endocast of S. austriacus generally resembles those of its relatives. A network of vascular canals surrounding the brain cavity further supports special thermoregulatory adaptations within Ankylosauria. The horizontal orientation of the lateral semicircular canal independently confirms previous appraisals of head posture for S. austriacus and, hence, strengthens the usage of the LSC as proxy for habitual head posture in fossil tetrapods. The short anterior and angular lateral semicircular canals, combined with the relatively shortest dinosaurian cochlear duct known so far and the lack of a floccular recess suggest a rather inert lifestyle without the necessity of sophisticated senses for equilibrium and hearing in S. austriacus. These observations agree with an animal that adapted to a comparatively inactive lifestyle with limited social interactions.

The neuroanatomy of Zulmasuchus querejazus (Crocodylomorpha, Sebecidae) and its implications for the paleoecology of sebecosuchians.

The endocranial structures of the sebecid crocodylomorph Zulmasuchus querejazus (MHNC 6672) from the Lower Paleocene of Bolivia are described in this article. Using computed tomography scanning, the cranial endocast, associated nerves and arteries, endosseous labyrinths, and cranial pneumatization are reconstructed and compared with those of extant and fossil crocodylomorphs, representative of different ecomorphological adaptations. Z. querejazus exhibits an unusual flexure of the brain, pericerebral spines, semicircular canals with a narrow diameter, as well as enlarged pharyngotympanic sinuses. First, those structures allow to estimate the alert head posture and hearing capabilities of Zulmasuchus. Then, functional comparisons are proposed between this purportedly terrestrial taxon, semi-aquatic, and aquatic forms (extant crocodylians, thalattosuchians, and dyrosaurids). The narrow diameter of the semicircular canals but expanded morphology of the endosseous labyrinths and the enlarged pneumatization of the skull compared to other forms indeed tend to indicate a terrestrial lifestyle for Zulmasuchus. Our results highlight the need to gather new data, especially from altirostral forms in order to further our understanding of the evolution of endocranial structures in crocodylomorphs with different ecomorphological adaptations.

Significant Asymmetry of the Bilateral Upper Extremities of a Skeleton Excavated from the Mashiki-Azamabaru Site, Okinawa Island, Japan.

The human skeleton of a young adult male with marked asymmetry of the bilateral upper extremities was excavated from the Mashiki-Azamabaru site (3000-2000 BCE) on the main island of Okinawa in the southwestern archipelago of Japan. The skeleton was buried alone in a corner of the cemetery. In this study, morphological and radiographic observations were made on this skeleton, and the pathogenesis of the bone growth disorder observed in the left upper limb was discussed. The maximum diameter of the midshaft of the humerus was 13.8 mm on the left and 21.2 mm on the right. The long bones comprising the left upper extremity lost the structure of the muscle attachments except for the deltoid tubercle of the humerus. The bone morphology of the right upper extremity and the bilateral lower extremities was maintained and was close to the mean value of females from the Ohtomo site in northwestern Kyushu, Japan, during the Yayoi period. It is assumed that the anomalous bone morphology confined to the left upper extremity was secondary to the prolonged loss of function of the muscles attached to left extremity bones. In this case, birth palsy, brachial plexus injury in childhood, and acute grey matter myelitis were diagnosed. It was suggested that this person had survived into young adulthood with severe paralysis of the left upper extremity due to injury or disease at an early age.

Dental replacement in Mesozoic birds: evidence from newly discovered Brazilian enantiornithines.

Polyphyodonty-multiple tooth generations-in Mesozoic birds has been confirmed since the nineteenth century. Their dental cycle had been assessed through sparse data from tooth roots revealed through broken jawbones and disattached teeth. However, detailed descriptions of their tooth cycling are lacking, and the specifics of their replacement patterns remain largely unknown. Here we present unprecedented µCT data from three enantiornithine specimens from the Upper Cretaceous of southeastern Brazil. The high resolution µCT data show an alternating dental replacement pattern in the premaxillae, consistent with the widespread pattern amongst extinct and extant reptiles. The dentary also reveals dental replacement at different stages. These results strongly suggest that an alternating pattern was typical of enantiornithine birds. µCT data show that new teeth start lingually within the alveoli, resorb roots of functional teeth and migrate labially into their pulp cavities at an early stage, similar to modern crocodilians. Our results imply that the control mechanism for tooth cycling is conserved during the transition between non-avian reptiles and birds. These first 3D reconstructions of enantiornithine dental replacement demonstrate that 3D data are essential to understand the evolution and deep homology of archosaurian tooth cycling.

Arothron: An R package for geometric morphometric methods and virtual anthropology applications.

OBJECTIVES: The statistical analysis of fossil remains is essential to understand the evolution of the genus Homo. Unfortunately, the human fossil record is straight away scarce and plagued with severe loss of information caused by taphonomic processes. The recently developed field of Virtual Anthropology helps to ameliorate this situation by using digital techniques to restore damaged and incomplete fossils. MATERIALS AND METHODS: We present the package Arothron, an R software suite meant to process and analyze digital models of skeletal elements. Arothron includes tools to digitally extract virtual cavities such as cranial endocasts, to statistically align disarticulated or broken bony elements, and to visualize local variations between surface meshes and landmark configurations. RESULTS: We describe the main functionalities of Arothron and illustrate their usage through reproducible case studies. We describe a tool for segmentation of skeletal cavities by showing its application on a malleus bone, a Neanderthal tooth, and a modern human cranium, reproducing their shape and calculating their volume. We illustrate how to digitally align a disarticulated model of a modern human cranium, and how to combine piecemeal shape information on individual specimens into one. In addition, we present useful visualization tools by comparing the morphological differences between the right hemisphere of the Neanderthal and the modern human brain. CONCLUSIONS: The Arothron R package is designed to study digital models of fossil specimens. By using Arothron, scientists can handle digital models with ease, investigate the inner morphology of 3D skeletal models, gain a full representation of the original shapes of damaged specimens, and compare shapes across specimens.

The impact of locomotion on the brain evolution of squirrels and close relatives.

How do brain size and proportions relate to ecology and evolutionary history? Here, we use virtual endocasts from 38 extinct and extant rodent species spanning 50+ million years of evolution to assess the impact of locomotion, body mass, and phylogeny on the size of the brain, olfactory bulbs, petrosal lobules, and neocortex. We find that body mass and phylogeny are highly correlated with relative brain and brain component size, and that locomotion strongly influences brain, petrosal lobule, and neocortical sizes. Notably, species living in trees have greater relative overall brain, petrosal lobule, and neocortical sizes compared to other locomotor categories, especially fossorial taxa. Across millions of years of Eocene-Recent environmental change, arboreality played a major role in the early evolution of squirrels and closely related aplodontiids, promoting the expansion of the neocortex and petrosal lobules. Fossoriality in aplodontiids had an opposing effect by reducing the need for large brains.

24.0 kyr cal BP stone artefact from Vale da Pedra Furada, Piauí, Brazil: Techno-functional analysis.

Current archaeological paradigm proposes that the first peopling of the Americas does not exceed the Last Glacial Maximum period. In this context, the acceptance of the anthropogenic character of the earliest stone artefacts generally rests on the presence of projectile points considered no more as typocentric but as typognomonic, since it allows, by itself, to certify the human character of the other associated artefacts. In other words, without this presence, nothing is certain. Archaeological research at Piauí (Brazil) attests to a Pleistocene human presence between 41 and 14 cal kyr BP, without any record of lithic projectile points. Here, we report the discovery and interpretation of an unusual stone artefact in the Vale da Pedra Furada site, in a context dating back to 24 cal kyr BP. The knapping stigmata and macroscopic use-wear traces reveal a conception centred on the configuration of double bevels and the production in the same specimen of at least two successive artefacts with probably different functions. This piece unambiguously presents an anthropic character and reveals a technical novelty during the Pleistocene occupation of South America.

Preliminary paleohistological observations of the StW 573 ('Little Foot') skull.

Numerous aspects of early hominin biology remain debated or simply unknown. However, recent developments in high-resolution imaging techniques have opened new avenues in the field of paleoanthropology. More specifically, X-ray synchrotron-based analytical imaging techniques have the potential to provide crucial details on the ontogeny, physiology, biomechanics, and biological identity of fossil specimens. Here we present preliminary results of our X-ray synchrotron-based investigation of the skull of the 3.67-million-year-old Australopithecus specimen StW 573 ('Little Foot') at the I12 beamline of the Diamond Light Source (United Kingdom). Besides showing fine details of the enamel (i.e., hypoplasias) and cementum (i.e., incremental lines), as well as of the cranial bone microarchitecture (e.g., diploic channels), our synchrotron-based investigation reveals for the first time the 3D spatial organization of the Haversian systems in the mandibular symphysis of an early hominin.

First rebbachisaurid sauropod dinosaur from Asia.

Dzharatitanis kingi gen. et sp. nov. is based on an isolated anterior caudal vertebra (USNM 538127) from the Upper Cretaceous (Turonian) Bissekty Formation at Dzharakuduk, Uzbekistan. Phylogenetic analysis places the new taxon within the diplodocoid clade Rebbachisauridae. This is the first rebbachisaurid reported from Asia and one of the youngest rebbachisaurids in the known fossil record. The caudal is characterized by a slightly opisthocoelous centrum, 'wing-like' transverse processes with large but shallow PRCDF and POCDF, and the absence of a hyposphenal ridge and of TPRL and TPOL. The neural spine has high SPRL, SPDL, SPOL, and POSL and is pneumatized. The apex of neural spine is transversely expanded and bears triangular lateral processes. The new taxon shares with Demandasaurus and the Wessex rebbachisaurid a high SPDL on the lateral side of the neural spine, separated from SPRL and SPOL. This possibly suggests derivation of Dzharatitanis from European rebbachisaurids. This is the second sauropod group identified in the assemblage of non-avian dinosaurs from the Bissekty Formation, in addition to a previously identified indeterminate titanosaurian.

Distinguishing Discoid and Centripetal Levallois methods through machine learning.

In this paper, we apply Machine Learning (ML) algorithms to study the differences between Discoid and Centripetal Levallois methods. For this purpose, we have used experimentally knapped flint flakes, measuring several parameters that have been analyzed by seven ML algorithms. From these analyses, it has been possible to demonstrate the existence of statistically significant differences between Discoid products and Centripetal Levallois products, thus contributing with new data and a new method to this traditional debate. The new approach enabled differentiating the blanks created by both knapping methods with an accuracy >80% using only ten typometric variables. The most relevant variables were maximum length, width to the 25%, 50% and 75% of the flake length, external and internal platform angles, maximum width and number of dorsal scars. This study also demonstrates the advantages of the application of multivariate ML methods to lithic studies.

A comprehensive diagnostic approach combining phylogenetic disease bracketing and CT imaging reveals osteomyelitis in a Tyrannosaurus rex.

Traditional palaeontological techniques of disease characterisation are limited to the analysis of osseous fossils, requiring several lines of evidence to support diagnoses. This study presents a novel stepwise concept for comprehensive diagnosis of pathologies in fossils by computed tomography imaging for morphological assessment combined with likelihood estimation based on systematic phylogenetic disease bracketing. This approach was applied to characterise pathologies of the left fibula and fused caudal vertebrae of the non-avian dinosaur Tyrannosaurus rex. Initial morphological assessment narrowed the differential diagnosis to neoplasia or infection. Subsequent data review from phylogenetically closely related species at the clade level revealed neoplasia rates as low as 3.1% and 1.8%, while infectious-disease rates were 32.0% and 53.9% in extant dinosaurs (birds) and non-avian reptiles, respectively. Furthermore, the survey of literature revealed that within the phylogenetic disease bracket the oldest case of bone infection (osteomyelitis) was identified in the mandible of a 275-million-year-old captorhinid eureptile Labidosaurus. These findings demonstrate low probability of a neoplastic aetiology of the examined pathologies in the Tyrannosaurus rex and in turn, suggest that they correspond to multiple foci of osteomyelitis.

Comparative isotope ecology of western Amazonian rainforest mammals.

Closed-canopy rainforests are important for climate (influencing atmospheric circulation, albedo, carbon storage, etc.) and ecology (harboring the highest biodiversity of continental regions). Of all rainforests, Amazonia is the world's most diverse, including the highest mammalian species richness. However, little is known about niche structure, ecological roles, and food resource partitioning of Amazonian mammalian communities over time. Through analyses of δ13Cbioapatite, δ13Chair, and δ15Nhair, we isotopically characterized aspects of feeding ecology in a modern western Amazonian mammalian community in Peru, serving as a baseline for understanding the evolution of Neotropical rainforest ecosystems. By comparing these results with data from equatorial Africa, we evaluated the potential influences of distinct phylogenetic and biogeographic histories on the isotopic niches occupied by mammals in analogous tropical ecosystems. Our results indicate that, despite their geographical and taxonomic differences, median δ13Cdiet values from closed-canopy rainforests in Amazonia (-27.4‰) and equatorial Africa (-26.9‰) are not significantly different, and that the median δ13Cdiet expected for mammalian herbivores in any closed-canopy rainforest is -27.2‰. Amazonian mammals seem to exploit a narrower spectrum of dietary resources than equatorial African mammals, however, as depicted by the absence of highly negative δ13Cdiet values previously proposed as indicative of rainforests (<-31‰). Finally, results of keratin and bioapatite δ13C indicate that the predictive power of trophic relationships, and traditional dietary ecological classifications in bioapatite-protein isotopic offset expectations, must be reconsidered.

Aenigmachannidae, a new family of snakehead fishes (Teleostei: Channoidei) from subterranean waters of South India.

Pronounced organism-wide morphological stasis in evolution has resulted in taxa with unusually high numbers of primitive characters. These 'living fossils' hold a prominent role for our understanding of the diversification of the group in question. Here we provide the first detailed osteological analysis of Aenigmachanna gollum based on high-resolution nano-CT scans and one cleared and stained specimen of this recently described snakehead fish from subterranean waters of Kerala in South India. In addition to a number of derived and unique features, Aenigmachanna has several characters that exhibit putatively primitive conditions not encountered in the family Channidae. Our morphological analysis provides evidence for the phylogenetic position of Aenigmachanna as the sister group to Channidae. Molecular analyses further emphasize the uniqueness of Aenigmachanna and indicate that it is a separate lineage of snakeheads, estimated to have split from its sister group at least 34 or 109 million years ago depending on the fossil calibration employed. This may indicate that Aenigmachanna is a Gondwanan lineage, which has survived break-up of the supercontinent, with India separating from Africa at around 120 mya. The surprising morphological disparity of Aenigmachanna from members of the Channidae lead us to erect a new family of snakehead fishes, Aenigmachannidae, sister group to Channidae, to accommodate these unique snakehead fishes.

Air Breathing in an Exceptionally Preserved 340-Million-Year-Old Sea Scorpion.

Arachnids are the second most successful terrestrial animal group after insects [1] and were one of the first arthropod clades to successfully invade land [2]. Fossil evidence for this transition is limited, with the majority of arachnid clades first appearing in the terrestrial fossil record. Furthermore, molecular clock dating has suggested a Cambrian-Ordovician terrestrialization event for arachnids [3], some 60 Ma before their first fossils in the Silurian, although these estimates assume that arachnids evolved from a fully aquatic ancestor. Eurypterids, the sister clade to terrestrial arachnids [4-6], are known to have undergone major macroecological shifts in transitioning from marine to freshwater environments during the Devonian [7, 8]. Discoveries of apparently subaerial eurypterid trackways [9, 10] have led to the suggestion that eurypterids were even able to venture on land and possibly breathe air [11]. However, modern horseshoe crabs undertake amphibious excursions onto land to reproduce [12], rendering trace fossil evidence alone inconclusive. Here, we present details of the respiratory organs of Adelophthalmus pyrrhae sp. nov. from the Carboniferous of Montagne Noire, France [13], revealed through micro computed tomography (µ-CT) imaging. Pillar-like trabeculae on the dorsal surface of each gill lamella indicate eurypterids were capable of subaerial breathing, suggesting that book gills are the direct precursors to book lungs while vascular ancillary respiratory structures known as Kiemenplatten represent novel air-breathing structures. The discovery of air-breathing structures in eurypterids indicates that characters permitting terrestrialization accrued in the arachnid stem lineage and suggests the Cambrian-Ordovician ancestor of arachnids would also have been semi-terrestrial.

Techno-functional and 3D shape analysis applied for investigating the variability of backed tools in the Late Middle Paleolithic of Central Europe.

In the Late Middle Paleolithic of Central Europe, two main cultural complexes have been distinguished: the Micoquian or Keilmessergruppe (KMG), and the Mousterian. Their differences mainly consist in the frequence of some retouched tools and the presence of bifacial technology. When these industries coexist, one element of discussion is the application of different concepts to manufacture tools with the same techno-functionality. This is particularly true for backed artifacts, such as Keilmesser (backed, asymmetrical bifacially-shaped knives) opposed to flake-tools equipped with a natural or knapped back. We conducted a techno-functional analysis of the backed tools from the G-Layer-Complex of Sesselfelsgrotte, one of the main Late Middle Paleolithic sequences in Central Europe, characterized by a combination of KMG and Mousterian aspects. In order to better understand the morpho-metrical data, 3D scans were used for recording technical features and performing semi-automatic geometric morphometrics. Results indicate that the techno-functional schemes of Keilmesser show a moderate variability and often overlap with the schemes of other typological groups. Within bifacial backed knives, a process of imitation of unifacial flake tools' functionaly was recognized particularly in the cutting edge manufacturing. Keilmesser proved to be the long-life, versatile version of backed flake-tools, also due to the recurrent valence as both tool and core. This is why Keilmesser represent an ideal strategic blank when a mobile and multi-functional tool is needed. Based on these data, it is assumed that the relationship between Mousterian and KMG is deeply rooted and the emergence of KMG aspects could be related to constrained situations characterizing the long cold stages of the Early Weichselian. A higher regional mobility caused by the comparably low predictability of resources characterized the subsistence tactics of Neanderthal groups especially at the borders of their overall distribution. For this reason, Keilmesser could have represented an ecological answer before possibly becoming a marker of cultural identity.

Radiographic analysis and virtual cleaning of a bioarchaeological remain enclosed in mineral deposits from a limestone cave.

In limestone caves, environmental processes often cause alterations of human or animal skeletal remains, complicating classical analytical methods. Exemplary, a proximal femoral skeletal fragment, enclosed by a thick layer of speleothemic calcite deposits, was discovered during the exploration of the Bedara cave in Zumberak, Croatia. An examination without removal of the surrounding mineral deposits, possibly leading to damage of the specimen, was, therefore, desirable.We describe and discuss the applied techniques, including clinical computed tomography, virtual cleaning by a specially developed segmentation protocol using an open-source DICOM viewer, and virtual visualisation and dimensioning using computer-aided design software, so that this "hidden" specimen could be non-invasively examined in great detail. We also report on the circumstances and origin of the find, the results of radiocarbon dating, and its anatomical and taxonomic identification, according to which, the bone fragment belonged to a wild boar (Sus scrofa) from the timeframe of the Middle Eneolithic Retz-Gajary culture in the region (4,781 ± 35 years before present). This study provides a reference for future paleontological and anthropological analyses, seeking to unlock the enormous potential of anatomical studies of comparable skeletal remains that are either petrified or enclosed in speleothemic deposits.

Cranial morphology of the tanystropheid Macrocnemus bassanii unveiled using synchrotron microtomography.

The genus Macrocnemus is a member of the Tanystropheidae, a clade of non-archosauriform archosauromorphs well known for their very characteristic, elongated cervical vertebrae. Articulated specimens are known from the Middle Triassic of Alpine Europe and China. Although multiple articulated specimens are known, description of the cranial morphology has proven challenging due to the crushed preservation of the specimens. Here we use synchrotron micro computed tomography to analyse the cranial morphology of a specimen of the type species Macrocnemus bassanii from the Besano Formation of Monte San Giorgio, Ticino, Switzerland. The skull is virtually complete and we identify and describe the braincase and palatal elements as well the atlas-axis complex for the first time. Moreover, we add to the knowledge of the morphology of the skull roof, rostrum and hemimandible, and reconstruct the cranium of M. bassanii in 3D using the rendered models of the elements. The circumorbital bones were found to be similar in morphology to those of the archosauromorphs Prolacerta broomi and Protorosaurus speneri. In addition, we confirm the palatine, vomer and pterygoid to be tooth-bearing palatal bones, but also observed heterodonty on the pterygoid and the palatine.