A proxy for brain-to-endocranial cavity index in non-neornithean dinosaurs and other extinct archosaurs.

Although the brain fills nearly the entire cranial cavity in birds, it can occupy a small portion of it in crocodilians. The lack of data regarding the volumetric correspondence between the brain and the cranial cavity hampers thorough assessments of the degree of encephalization in non-neornithean dinosaurs and other extinct archosaurs and, consequently, informed inferences regarding their cognitive capacities. Existing data suggest that, across extant archosaurs, the degree of endocranial doming and the volume of intracranial nonneural components are inversely related. We build upon this information to develop an equation relating these two anatomical features in non-neornithean dinosaurs and other extinct archosaurs. We rely on measurements of the endocast doming and brain-to-endocranial cavity (BEC) index in extant relatives of non-neornithean dinosaurs, namely, the crurotarsans Caiman crocodilus, Crocodylus niloticus, and Crocodylus porosus; the paleognaths Struthio camelus and Apteryx mantelli; and the fowl Macrocephalon maleo, Gallus gallus, Meleagris gallopavo, Phasianus colchicus, and Anas platyrhynchos. Applying the equation to representative endocasts from major clades of dinosaurs, we found that BEC varies from about 0.6 in ceratopsians and thyreophorans to around 0.7 in ornithopods, pachycephalosaurians, sauropods, and theropods. We, therefore, warn against the use of a catch-all value, like 0.5, and instead encourage refinement in the adoption of BEC across archosaurs.

Palaeoneurology of the early cretaceous iguanodont Proa valdearinnoensis and its bearing on the parallel developments of cognitive abilities in theropod and ornithopod dinosaurs.

Proa valdearinnoensis is a relatively large-headed and stocky iguanodontian dinosaur from the latest Early Cretaceous of Spain. Its braincase is known from three specimens. Similar to that of other dinosaurs, it shows a mosaic ossification pattern in which most of the bones seem to have fused together indistinguishably while a few (frontoparietal, basioccipital) might have remained loosely attached. The endocasts of the three specimens are described based on CT data and digital reconstructions. They show unmistakable morphological similarities with the endocast of closely related taxa, such as Sirindhorna khoratensis (which is close in age but from Thailand). This supports a high conservatism of the endocranial cavity. The issue of volumetric correspondence between endocranial cavity and brain in dinosaurs is analyzed. Although a brain-to-endocranial cavity (BEC) index of 0.50 has been traditionally used, we employ instead 0.73. This is indeed the mid-value between the situation in adults of Alligator mississippiensis and Gallus gallus, which are members of the extant bracketing taxa of dinosaurs (Crocodilia and Aves). We thence gauge the level of encephalization of P. valdearinnoensis through the calculation of the encephalization quotient (EQ), which remains valuable as a metric for assessing the degree of cognitive function in extinct taxa, especially those with fully ossified braincases like dinosaurs and other archosaurs. The EQ obtained for P. valdearinnoensis (3.611) suggests that this species was significantly more encephalized than most if not all extant nonavian, nonmammalian amniotes. Our work adds to the growing body of data concerning theoretical cognitive capabilities in dinosaurs and supports the idea that an increasing encephalization was fostered not only in theropods but also in parallel in the shorter-lived lineage of ornithopods. P. valdearinnoensis was ill-equipped to respond to theropod dinosaurs and possibly lived in groups as a strategy to mitigate the risk of being predated upon. We hypothesize that group-living and protracted caring of juveniles in this and possibly many other iguanodontian ornithopods favored a degree of encephalization that was outstanding by reptile standards.

Avian palaeoneurology: Reflections on the eve of its 200th anniversary.

In birds, the brain (especially the telencephalon) is remarkably developed, both in relative volume and complexity. Unlike in most early-branching sauropsids, the adults of birds and other archosaurs have a well-ossified neurocranium. In contrast to the situation in most of their reptilian relatives but similar to what can be seen in mammals, the brains of birds fit closely to the endocranial cavity so that their major external features are reflected in the endocasts. This makes birds a highly suitable group for palaeoneurological investigations. The first observation about the brain in a long-extinct bird was made in the first quarter of the 19th century. However, it was not until the 2000s and the application of modern imaging technologies that avian palaeoneurology really took off. Understanding how the mode of life is reflected in the external morphology of the brains of birds is but one of several future directions in which avian palaeoneurological research may extend. Although the number of fossil specimens suitable for palaeoneurological explorations is considerably smaller in birds than in mammals and will very likely remain so, the coming years will certainly witness a momentous strengthening of this rapidly growing field of research at the overlap between ornithology, palaeontology, evolutionary biology and neurosciences.

First palaeoneurological study of a sauropod dinosaur from France and its phylogenetic significance.

Despite continuous improvements, our knowledge of the palaeoneurology of sauropod dinosaurs is still deficient. This holds true even for Titanosauria, which is a particularly speciose clade of sauropods with representatives known from numerous Cretaceous sites in many countries on all continents. The data currently available regarding the palaeoneurology of titanosaurs is strongly biased towards Gondwanan forms (Argentina above all, but also India, Malawi and Australia). In contrast, the palaeoneurology of Laurasian titanosaurs is known only from a few taxa from Spain and Uzbekistan, despite the discovery in other countries of Laurasia of a number of neurocranial remains that would lend themselves well to investigations of this kind. To fill in this gap in our knowledge, we subjected a titanosaurian braincase from the uppermost Upper Cretaceous of southern France to X-ray computed tomographic scanning, allowing the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves and blood vessels, as well as the labyrinth of the inner ear. These reconstructions are used to clarify the phylogenetic position of the specimen from the Fox-Amphoux-Métisson site. A combination of characters, including the presence of two hypoglossal rami on the endocast, the average degree of development of the dorsal-head/caudal-middle-cerebral vein system and the relatively short and subequal lengths of the ipsilateral semicircular canals of the labyrinth, are particularly revealing in this respect. They suggest that, compared with the few other Laurasian titanosaurs for which in-depth palaeoneurological data are available, the French taxon is more derived than the distinctly more ancient, possibly non-lithostrotian titanosaur from the Uzbek site of Dzharakuduk but more basal than derived saltasaurids, such as the coeval or slightly more recent forms from the Spanish locality of Lo Hueco.

First levantine fossil murines shed new light on the earliest intercontinental dispersal of mice.

Recent extensive field prospecting conducted in the Upper Miocene of Lebanon resulted in the discovery of several new fossiliferous localities. One of these, situated in the Zahleh area (Bekaa Valley, central Lebanon) has yielded a particularly diverse vertebrate fauna. Micromammals constitute an important part of this assemblage because not only do they represent the first Neogene rodents and insectivores from Lebanon, but they are also the only ones from the early Late Miocene of the Arabian Peninsula and circumambient areas. Analyses of the murines from Zahleh reveal that they belong to a small-sized early Progonomys, which cannot be assigned to any of the species of the genus hitherto described. They are, thereby, shown to represent a new species: Progonomys manolo. Morphometric analyses of the outline of the first upper molars of this species suggest a generalist and omnivorous diet. This record sheds new light onto a major phenomenon in the evolutionary history of rodents, which is the earliest dispersal of mice. It suggests that the arrival of murines in Africa got under way through the Levant rather than via southern Europe and was monitored by the ecological requirements of Progonomys.

Pheomelanin pigment remnants mapped in fossils of an extinct mammal.

Recent progress has been made in paleontology with respect to resolving pigmentation in fossil material. Morphological identification of fossilized melanosomes has been one approach, while a second methodology using chemical imaging and spectroscopy has also provided critical information particularly concerning eumelanin (black pigment) residue. In this work we develop the chemical imaging methodology to show that organosulfur-Zn complexes are indicators of pheomelanin (red pigment) in extant and fossil soft tissue and that the mapping of these residual biochemical compounds can be used to restore melanin pigment distribution in a 3 million year old extinct mammal species (Apodemus atavus). Synchotron Rapid Scanning X-ray Fluorescence imaging showed that the distributions of Zn and organic S are correlated within this fossil fur just as in pheomelanin-rich modern integument. Furthermore, Zn coordination chemistry within this fossil fur is closely comparable to that determined from pheomelanin-rich fur and hair standards. The non-destructive methods presented here provide a protocol for detecting residual pheomelanin in precious specimens.

Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards.

Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.

A diminutive perinate European Enantiornithes reveals an asynchronous ossification pattern in early birds.

Fossils of juvenile Mesozoic birds provide insight into the early evolution of avian development, however such fossils are rare. The analysis of the ossification sequence in these early-branching birds has the potential to address important questions about their comparative developmental biology and to help understand their morphological evolution and ecological differentiation. Here we report on an early juvenile enantiornithine specimen from the Early Cretaceous of Europe, which sheds new light on the osteogenesis in this most species-rich clade of Mesozoic birds. Consisting of a nearly complete skeleton, it is amongst the smallest known Mesozoic avian fossils representing post-hatching stages of development. Comparisons between this new specimen and other known early juvenile enantiornithines support a clade-wide asynchronous pattern of osteogenesis in the sternum and the vertebral column, and strongly indicate that the hatchlings of these phylogenetically basal birds varied greatly in size and tempo of skeletal maturation.

Digital reconstruction of the mandible of an adult Lesothosaurus diagnosticus with insight into the tooth replacement process and diet.

Fragmentary caudal ends of the left and right mandible assigned to Lesothosaurus diagnosticus, an early ornithischian, was recently discovered in the continental red bed succession of the upper Elliot Formation (Lower Jurassic) at Likhoele Mountain (Mafeteng District) in Lesotho. Using micro-CT scanning, this mandible could be digitally reconstructed in 3D. The replacement teeth within the better preserved (left) dentary were visualised. The computed tomography dataset suggests asynchronous tooth replacement in an individual identified as an adult on the basis of bone histology. Clear evidence for systematic wear facets created by attrition is lacking. The two most heavily worn teeth are only apically truncated. Our observations of this specimen as well as others do not support the high level of dental wear expected from the semi-arid palaeoenvironment in which Lesothosaurus diagnosticus lived. Accordingly, a facultative omnivorous lifestyle, where seasonality determined the availability, quality, and abundance of food is suggested. This would have allowed for adaptability to episodes of increased environmental stress.

A New Titanosaurian Braincase from the Cretaceous "Lo Hueco" Locality in Spain Sheds Light on Neuroanatomical Evolution within Titanosauria.

Despite continuous improvements, our knowledge of the neurocranial anatomy of sauropod dinosaurs as a whole is still poor, which is especially true for titanosaurians even though their postcranial remains are common in many Upper Cretaceous sites worldwide. Here we describe a braincase from the uppermost Cretaceous locality of ''Lo Hueco" in Spain that is one of the most complete titanosaurian braincases found so far in Europe. Although the titanosaurian Ampelosaurus sp. is known from the same locality, this specimen is clearly a distinct taxon and presents a number of occipital characters found in Antarctosaurus and Jainosaurus, which are approximately coeval taxa from southern Gondwana. The specimen was subjected to X-ray computed tomographic (CT) scanning, allowing the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves, and blood vessels, as well as the labyrinth of the inner ear. These findings add considerable knowledge to the field of sauropod paleoneuroanatomy in general and titanosaurian endocast diversity in particular. Compared with that of many sauropodomorphs, the endocast appears only slightly flexed in lateral view and bears similarities (e.g., reduction of the rostral dural expansion) with Gondwanan titanosaurians such as Jainosaurus, Bonatitan, and Antarctosaurus. The vestibular system of the inner ear is somewhat contracted (i.e., the radius of the semicircular canals is small), but less so than expected in derived titanosaurians. However, as far as the new specimen and Jainosaurus can be contrasted, and with the necessary caution due to the small sample of comparative data currently available, the two taxa appear more similar to one another in endocast morphology than to other titanosaurians. Recent phylogenetic analyses of titanosaurians have not included virtually any of the taxa under consideration here, and thus the phylogenetic position of the new Spanish titanosaurian--even its generic, let alone specific, identification--is not possible at the moment. Nevertheless, both the braincase osteology and the endocast morphology suggest that the specimen represents a derived titanosaurian that presumably branched further from the base of Lithostrotia, potentially even near Saltasauridae, comparable in evolutionary terms with Jainosaurus.

First Miocene rodent from Lebanon provides the 'missing link' between Asian and African gundis (Rodentia: Ctenodactylidae).

Ctenodactylinae (gundis) is a clade of rodents that experienced, in Miocene time, their greatest diversification and widest distribution. They expanded from the Far East, their area of origin, to Africa, which they entered from what would become the Arabian Peninsula. Questions concerning the origin of African Ctenodactylinae persist essentially because of a poor fossil record from the Miocene of Afro-Arabia. However, recent excavations in the Late Miocene of Lebanon have yielded a key taxon for our understanding of these issues. Proafricanomys libanensis nov. gen. nov. sp. shares a variety of dental characters with both the most primitive and derived members of the subfamily. A cladistic analysis demonstrates that this species is the sister taxon to a clade encompassing all but one of the African ctenodactylines, plus a southern European species of obvious African extraction. As such, Proafricanomys provides the 'missing link' between the Asian and African gundis.

Causal evidence between monsoon and evolution of rhizomyine rodents.

The modern Asian monsoonal systems are currently believed to have originated around the end of the Oligocene following a crucial step of uplift of the Tibetan-Himalayan highlands. Although monsoon possibly drove the evolution of many mammal lineages during the Neogene, no evidence thereof has been provided so far. We examined the evolutionary history of a clade of rodents, the Rhizomyinae, in conjunction with our current knowledge of monsoon fluctuations over time. The macroevolutionary dynamics of rhizomyines were analyzed within a well-constrained phylogenetic framework coupled with biogeographic and evolutionary rate studies. The evolutionary novelties developed by these rodents were surveyed in parallel with the fluctuations of the Indian monsoon so as to evaluate synchroneity and postulate causal relationships. We showed the existence of three drops in biodiversity during the evolution of rhizomyines, all of which reflected elevated extinction rates. Our results demonstrated linkage of monsoon variations with the evolution and biogeography of rhizomyines. Paradoxically, the evolution of rhizomyines was accelerated during the phases of weakening of the monsoons, not of strengthening, most probably because at those intervals forest habitats declined, which triggered extinction and progressive specialization toward a burrowing existence.

Neurocranial osteology and neuroanatomy of a late Cretaceous titanosaurian sauropod from Spain (Ampelosaurus sp.).

Titanosaurians were a flourishing group of sauropod dinosaurs during Cretaceous times. Fossils of titanosaurians have been found on all continents and their remains are abundant in a number of Late Cretaceous sites. Nonetheless, the cranial anatomy of titanosaurians is still very poorly known. The Spanish latest Cretaceous locality of "Lo Hueco" yielded a relatively well preserved, titanosaurian braincase, which shares a number of phylogenetically restricted characters with Ampelosaurus atacis from France such as a flat occipital region. However, it appears to differ from A. atacis in some traits such as the greater degree of dorsoventral compression and the presence of proatlas facets. The specimen is, therefore, provisionally identified as Ampelosaurus sp. It was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. Our investigation highlights that, although titanosaurs were derived sauropods with a successful evolutionary history, they present a remarkably modest level of paleoneurological organization. Compared with the condition in the basal titanosauriform Giraffatitan brancai, the labyrinth of Ampelosaurus sp. shows a reduced morphology. The latter feature is possibly related to a restricted range of head-turning movements.

A comprehensive phylogeny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals.

The subfamily Rhizomyinae is known from the Late Oligocene up to the present. Today this group comprises six species, which live in southern Asia and eastern Africa. Despite the current moderate diversity of the rhizomyines, they had a greater diversification and wider distribution in the past: from Asia, their land of origin, to Africa, which they entered during the Early Miocene. So far 33 fossil species can be referred to this group. A cladistic analysis involving fossil and living species has been carried out. Prokanisamys spp. turned out to be the most basal taxa of the ingroup. This analysis calls into question the monophyly of several genera, and allows the proposal of a phylogenetic definition of the tribes Tachyoryctini and Rhizomyini. It also provides information about the origin of the African rhizomyines and allows inferring multiple dispersal phenomena from Asia to Africa in Early and Late Miocene times.

The braincase of the basal sauropod dinosaur Spinophorosaurus and 3D reconstructions of the cranial endocast and inner ear.

BACKGROUND: Sauropod dinosaurs were the largest animals ever to walk on land, and, as a result, the evolution of their remarkable adaptations has been of great interest. The braincase is of particular interest because it houses the brain and inner ear. However, only a few studies of these structures in sauropods are available to date. Because of the phylogenetic position of Spinophorosaurus nigerensis as a basal eusauropod, the braincase has the potential to provide key evidence on the evolutionary transition relative to other dinosaurs. METHODOLOGY/PRINCIPAL FINDINGS: The only known braincase of Spinophorosaurus ('Argiles de l'Irhazer', Irhazer Group; Agadez region, Niger) differs significantly from those of the Jurassic sauropods examined, except potentially for Atlasaurus imelakei (Tilougguit Formation, Morocco). The basisphenoids of Spinophorosaurus and Atlasaurus bear basipterygoid processes that are comparable in being directed strongly caudally. The Spinophorosaurus specimen was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. The endocast resembles that of most other sauropods in having well-marked pontine and cerebral flexures, a large and oblong pituitary fossa, and in having the brain structure obscured by the former existence of relatively thick meninges and dural venous sinuses. The labyrinth is characterized by long and proportionally slender semicircular canals. This condition recalls, in particular, that of the basal non-sauropod sauropodomorph Massospondylus and the basal titanosauriform Giraffatitan. CONCLUSIONS/SIGNIFICANCE: Spinophorosaurus has a moderately derived paleoneuroanatomical pattern. In contrast to what might be expected early within a lineage leading to plant-eating graviportal quadrupeds, Spinophorosaurus and other (but not all) sauropodomorphs show no reduction of the vestibular apparatus of the inner ear. This character-state is possibly a primitive retention in Spinophorosaurus, but due the scarcity of data it remains unclear whether it is also the case in the various later sauropods in which it is present or whether it has developed homoplastically in these taxa. Any interpretations remain tentative pending the more comprehensive quantitative analysis underway, but the size and morphology of the labyrinth of sauropodomorphs may be related to neck length and mobility, among other factors.