The endocast of the insular and extinct Sylviornis neocaledoniae (Aves, Galliformes), reveals insights into its sensory specializations and its twilight ecology.

Sylviornis neocaledoniae (Galliformes, Sylviornithidae), a recently extinct bird of New-Caledonia (Galliformes, Sylviornithidae) is the largest galliform that ever lived and one of the most enigmatic birds in the world. Herein, for the first time, we analyze its neuroanatomy that sheds light on its lifestyle, its brain shape and patterns being correlated to neurological functions. Using morphometric methods, we quantified the endocranial morphology of S. neocaledoniae and compared it with extinct and extant birds in order to obtain ecological and behavioral information about fossil birds. Sylviornis neocaledoniae exhibited reduced optic lobes, a condition also observed in nocturnal taxa endemic to predator-depauperate islands, such as Elephant birds. Functional interpretations suggest that S. neocaledoniae possessed a well-developed somatosensorial system and a good sense of smell in addition to its specialized visual ability for low light conditions, presumably for locating its food. We interpret these results as evidence for a crepuscular lifestyle in S. neocaledoniae.

Isotopic systematics point to wild origin of mummified birds in Ancient Egypt.

Millions of mummified birds serving for religious purpose have been discovered from archeological sites along the Nile Valley of Egypt, in majority ibises. Whether these birds were industrially raised or massively hunted is a matter of heavy debate as it would have a significant impact on the economy related to their supply and cult, and if hunted it would have represented an ecological burden on the birds populations. Here we have measured and analysed the stable oxygen, carbon and radiogenic strontium isotope compositions as well as calcium and barium content of bones along with the stable carbon, nitrogen and sulfur isotope composition of feathers from 20 mummified ibises and birds of prey recovered from various archeological sites of Ancient Egypt. If these migratory birds were locally bred, their stable oxygen, radiogenic strontium and stable sulfur isotopic compositions would be similar to that of coexisting Egyptians, and their stable carbon, nitrogen and oxygen isotope variance would be close, or lower than that of Egyptians. On one hand, isotopic values show that ibises ingested food from the Nile valley but with a higher isotopic scattering than observed for the diet of ancient Egyptians. On the other hand, birds of prey have exotic isotopic values compatible with their migratory behaviour. We therefore propose that most mummified ibises and all the birds of prey analysed here were wild animals hunted for religious practice.

Cranial evolution in the extinct Rodrigues Island owl Otus murivorus (Strigidae), associated with unexpected ecological adaptations.

Island birds that were victims of anthropic extinctions were often more specialist species, having evolved their most distinctive features in isolation, making the study of fossil insular birds most interesting. Here we studied a fossil cranium of the 'giant' extinct scops owl Otus murivorus from Rodrigues Island (Mascarene Islands, southwestern Indian Ocean), to determine any potential unique characters. The fossil and extant strigids were imaged through X-ray microtomography, providing 3D views of external and internal (endocast, inner ear) cranial structures. Geometric morphometrics and analyses of traditional measurements yielded new information about the Rodrigues owl's evolution and ecology. Otus murivorus exhibits a 2-tier "lag behind" phenomenon for cranium and brain evolution, both being proportionately small relative to increased body size. It also had a much more developed olfactory bulb than congeners, indicating an unexpectedly developed olfactory sense, suggesting a partial food scavenging habit. In addition, O. murivorus had the eyes placed more laterally than O. sunia, the species from which it was derived, probably a side effect of a small brain; rather terrestrial habits; probably relatively fearless behavior; and a less vertical posture (head less upright) than other owls (this in part an allometric effect of size increase). These evolutionary features, added to gigantism and wing reduction, make the extinct Rodrigues owl's evolution remarkable, and with multiple causes.

The earliest Tyrannida (Aves, Passeriformes), from the Oligocene of France.

Passeriformes is the most diverse bird order. Nevertheless, passerines have a remarkably poor early fossil record. In addition, high osteological homoplasy across passerines makes partial specimens difficult to systematically assign precisely. Here we describe one of the few earliest fossil passerines, from the early Oligocene (ca 30 Ma) of southern France, and one of the best preserved and most complete. This fossil can be conservatively assigned to Tyrannida, a subclade of the New World Tyranni (Suboscines), i.e. of the Tyrannides. A most probably stem-representative of Tyrannida, the new fossil bears strong resemblance with some manakins (Pipridae), possibly due to plesiomorphy. Furthermore, it yields a new point of calibration for molecular phylogenies, already consistent with the age of the fossil. Tyrannida, and the more inclusive Tyrannides, are today confined to the New World. Therefore, the new fossil calls for scenarios of transatlantic crossing during or near the Oligocene. Later, the European part of the distribution of the Tyrannida disappeared, leading to a relictual modern New World distribution of this clade, a pattern known in other avian clades. The history of Tyrannida somehow mirrors that of the enigmatic Sapayoa aenigma, sole New World representative of the Eurylaimides (Old World Tyranni), with transatlantic crossing probably caused by similar events.

Bony pseudoteeth of extinct pelagic birds (Aves, Odontopterygiformes) formed through a response of bone cells to tooth-specific epithelial signals under unique conditions.

Modern birds (crown group birds, called Neornithes) are toothless; however, the extinct neornithine Odontopterygiformes possessed bone excrescences (pseudoteeth) which resembled teeth, distributed sequentially by size along jaws. The origin of pseudoteeth is enigmatic, but based on recent evidence, including microanatomical and histological analyses, we propose that conserved odontogenetic pathways most probably regulated the development of pseudodentition. The delayed pseudoteeth growth and epithelium keratinization allowed for the existence of a temporal window during which competent osteoblasts could respond to oral epithelial signaling, in place of the no longer present odontoblasts; thus, bony pseudoteeth developed instead of true teeth. Dynamic morphogenetic fields can explain the particular, sequential size distribution of pseudoteeth along the jaws of these birds. Hence, this appears as a new kind of deep homology, by which ancient odontogenetic developmental processes would have controlled the evolution of pseudodentition, structurally different from a true dentition, but morphologically and functionally similar.

Synchrotron imaging of dentition provides insights into the biology of Hesperornis and Ichthyornis, the "last" toothed birds.

BACKGROUND: The dentitions of extinct organisms can provide pivotal information regarding their phylogenetic position, as well as paleobiology, diet, development, and growth. Extant birds are edentulous (toothless), but their closest relatives among stem birds, the Cretaceous Hesperornithiformes and Ichthyornithiformes, retained teeth. Despite their significant phylogenetic position immediately outside the avian crown group, the dentitions of these taxa have never been studied in detail. To obtain new insight into the biology of these 'last' toothed birds, we use cutting-edge visualisation techniques to describe their dentitions at unprecedented levels of detail, in particular propagation phase contrast x-ray synchrotron microtomography at high-resolution. RESULTS: Among other characteristics of tooth shape, growth, attachment, implantation, replacement, and dental tissue microstructures, revealed by these analyses, we find that tooth morphology and ornamentation differ greatly between the Hesperornithiformes and Ichthyornithiformes. We also highlight the first Old World, and youngest record of the major Mesozoic clade Ichthyornithiformes. Both taxa exhibit extremely thin and simple enamel. The extension rate of Hesperornis tooth dentine appears relatively high compared to non-avian dinosaurs. Root attachment is found for the first time to be fully thecodont via gomphosis in both taxa, but in Hesperornis secondary evolution led to teeth implantation in a groove, at least locally without a periodontal ligament. Dental replacement is shown to be lingual via a resorption pit in the root, in both taxa. CONCLUSIONS: Our results allow comparison with other archosaurs and also mammals, with implications regarding dental character evolution across amniotes. Some dental features of the 'last' toothed birds can be interpreted as functional adaptations related to diet and mode of predation, while others appear to be products of their peculiar phylogenetic heritage. The autapomorphic Hesperornis groove might have favoured firmer root attachment. These observations highlight complexity in the evolutionary history of tooth reduction in the avian lineage and also clarify alleged avian dental characteristics in the frame of a long-standing debate on bird origins. Finally, new hypotheses emerge that will possibly be tested by further analyses of avian teeth, for instance regarding dental replacement rates, or simplification and thinning of enamel throughout the course of early avian evolution.

Structure and growth pattern of pseudoteeth in Pelagornis mauretanicus (Aves, Odontopterygiformes, Pelagornithidae).

The extinct Odontopterygiformes are the sole birds known to possess strong and sharp bony pseudoteeth, the shape and location of which are closely mimetic of real teeth. The structure of the pseudoteeth is investigated here in a late Pliocene/early Pleistocene species, Pelagornis mauretanicus, using X-ray microtomography and thin sections. The results are interpreted with regard to the pseudotooth mode of growth, and have implications concerning aspects of Pelagornis ecology. The larger pseudoteeth are hollow and approximately cone-shaped, and the smaller ones are rostro-caudally constricted. The walls of pseudoteeth are composed of bone tissue of the fibro-lamellar type, which is intensively remodeled by Haversian substitution. The jaw bones display the same structure as the pseudoteeth, but their vascular canals are oriented parallel to the long axis of the bones, whereas they are perpendicular to this direction in the pseudoteeth. There is no hiatus or evidence of a fusion between the pseudoteeth and the jaw bones. Two possible models for pseudotooth growth are derived from the histological data. The most plausible model is that pseudotooth growth began after the completion of jaw bone growth, as a simple local protraction of periosteal osteogenic activity. Pseudotooth development thus occurred relatively late during ontogeny. The highly vascularized structure and the relative abundance of parallel-fibered bone tissue in the pseudoteeth suggest poor mechanical capabilities. The pseudoteeth were most likely covered and protected by the hardened, keratinized rhamphotheca in the adult during life. The late development of the pseudoteeth would involve a similarly late and/or partial hardening of the rhamphotheca, as displayed by extant Anseriformes, Apterygiformes and some Charadriiformes. This would add support to the hypothesis of a close phylogenetic relationship between Odontopterygiformes and Anseriformes. The late maturation of the Pelagornis feeding apparatus, and hence the delayed capability for efficient prey catching, suggests that Pelagornis was altricial.

Nomenclatural and taxonomic problems related to the electronic publication of new nomina and nomenclatural acts in zoology, with brief comments on optical discs and on the situation in botany.

In zoological nomenclature, to be potentially valid, nomenclatural novelties (i.e., new nomina and nomenclatural acts) need first to be made available, that is, published in works qualifying as publications as defined by the International Code of zoological Nomenclature ("the Code"). In September 2012, the Code was amended in order to allow the recognition of works electronically published online after 2011 as publications available for the purpose of zoological nomenclature, provided they meet several conditions, notably a preregistration of the work in ZooBank. Despite these new Rules, several of the long-discussed problems concerning the electronic publication of new nomina and nomenclatural acts have not been resolved. The publication of this amendment provides an opportunity to discuss some of these in detail. It is important to note that: (1) all works published only online before 2012 are nomenclaturally unavailable; (2) printed copies of the PDFs of works which do not have their own ISSN or ISBN, and which are not obtainable free of charge or by purchase, do not qualify as publications but must be seen as facsimiles of unavailable works and are unable to provide nomenclatural availability to any nomenclatural novelties they may contain; (3) prepublications online of later released online publications are unavailable, i.e., they do not advance the date of publication; (4) the publication dates of works for which online prepublications had been released are not those of these prepublications and it is critical that the real release date of such works appear on the actual final electronic publication, but this is not currently the case in electronic periodicals that distribute such online prepublications and which still indicate on their websites and PDFs the date of release of prepublication as that of publication of the work; (5) supplementary online materials and subsequent formal corrections of either paper or electronic publications distributed only online are nomenclaturally unavailable; (6) nomenclatural information provided on online websites that do not have a fixed content and format, with ISSN or ISBN, is unavailable. We give precise examples of many of these nomenclatural problems. Several of them, when they arise, are due to the fact that the availability of nomenclatural novelties now depends on information that will have to be sought not from the work itself but from extrinsic evidence. As shown by several examples discussed here, an electronic document can be modified while keeping the same DOI and publication date, which is not compatible with the requirements of zoological nomenclature. Therefore, another system of registration of electronic documents as permanent and inalterable will have to be devised. ZooBank also clearly needs to be improved in several respects. Mention in a work of its registration number (LSID) in ZooBank would seem to be possible only if this registration has occurred previously, but some works that have purportedly been registered in ZooBank are in fact missing on this web application. In conclusion, we offer recommendations to authors, referees, editors, publishers, libraries and the International Commission on Zoological Nomenclature, in the hope that such problems can be limited along with the potential chaos in zoological nomenclature that could result, if careful attention is not paid to the problems we highlight here, from a somewhat misplaced, and perhaps now widespread, understanding that electronic publication of nomenclatural novelties is now allowed and straightforward. We suggest that, as long as the problematic points linked to the new amendment and to electronic publication as a whole are not resolved, nomenclatural novelties continue to be published in paper-printed journals that have so far shown editorial competence regarding taxonomy and nomenclature, which is not the case of several recent electronic-only published journals.

From snout to beak: the loss of teeth in birds.

All living birds are toothless, constituting by far the most diverse toothless vertebrate clade, and are striking examples of evolutionary success following tooth loss. In recent years, an unprecedented number of Mesozoic birds have been described, illustrating the evolution of dentition reductions. Simultaneously, major advances in experimental embryology have yielded new results concerning avian edentulism. Reviewing these lines of evidence, we propose hypotheses for its causes, with a prominent role for the horny beak during development. A horny beak and a muscular gizzard functionally 'replaced' dentition for food acquisition and processing, respectively. Together with edentulism itself, these features and others contributed to the later success of birds, as a result of their high performance or additional functionality working in concert in these complex organisms.

Taphonomic, avian, and small-vertebrate indicators of Ardipithecus ramidus habitat.

Thousands of vertebrate specimens were systematically collected from the stratigraphic interval containing Ardipithecus ramidus. The carcasses of larger mammals were heavily ravaged by carnivores. Nearly 10,000 small-mammal remains appear to be derived primarily from decomposed owl pellets. The rich avifauna includes at least 29 species, mostly nonaquatic forms. Modern analogs of the most abundant birds and of a variety of rodents are associated with mesic woodland environments distant from large water bodies. These findings support inferences from associated geological, isotopic, invertebrate, and large-vertebrate assemblages. The combined results suggest that Ar. ramidus occupied a wooded Pliocene habitat.

Emergence of long distance bird migrations: a new model integrating global climate changes.

During modern birds history, climatic and environmental conditions have evolved on wide scales. In a continuously changing world, landbirds annual migrations emerged and developed. However, models accounting for the origins of these avian migrations were formulated with static ecogeographic perspectives. Here I reviewed Cenozoic paleoclimatic and paleontological data relative to the palearctic-paleotropical long distance (LD) migration system. This led to propose a new model for the origin of LD migrations, the 'shifting home' model (SHM). It is based on a dynamic perspective of climate evolution and may apply to the origins of most modern migrations. Non-migrant tropical African bird taxa were present at European latitudes during most of the Cenozoic. Their distribution limits shifted progressively toward modern tropical latitudes during periods of global cooling and increasing seasonality. In parallel, decreasing winter temperatures in the western Palearctic drove shifts of population winter ranges toward the equator. I propose that this induced the emergence of most short distance migrations, and in turn LD migrations. This model reconciliates ecologically tropical ancestry of most LD migrants with predominant winter range shifts, in accordance with requirements for heritable homing. In addition, it is more parsimonious than other non-exclusive models. Greater intrinsic plasticity of winter ranges implied by the SHM is supported by recently observed impacts of the present global warming on migrating birds. This may induce particular threats to some LD migrants. The ancestral, breeding homes of LD migrants were not 'northern' or 'southern' but shifted across high and middle latitudes while migrations emerged through winter range shifts themselves.

Fossil birds of the Kibish Formation.

The Kibish Formation has yielded a small collection of bird fossils, which are identified here as belonging to five species in four different families: Pelecanidae (pelicans), Anhingidae (darters), Ardeidae (herons) and Phasianidae (gamefowl). Two species of pelicans are identified: Pelecanus cf. P. onocrotalus, and P. aff. P. rufescens. The darter is referrable to Anhinga melanogaster. The heron is identifiable as Ardea sp., and the gamefowl as Numidinae indet. (guineafowl). Pelecanus cf. P. onocrotalus is represented by, among other remains, a well-preserved partial skull. Four of the birds are thus referrable to extant taxa that provide some paleoenvironmental clues for Member I of the Kibish Formation. The two species of pelican, the darter, and the heron indicate the presence of local freshwater bodies, a lake or a slow river, supporting resources of fish. The guineafowl is poorly informative ecologically, but probably excludes the notion that the local terrestrial landscape was treeless.

Hummingbird with modern feathering: an exceptionally well-preserved Oligocene fossil from southern France.

Hummingbirds (Trochilidae) today have an exclusively New World distribution, but their pre-Pleistocene fossil record comes from Europe only. In this study, we describe an exceptionally preserved fossil hummingbird from the early Oligocene of southeastern France. The specimen is articulated, with a completely preserved beak and feathering. Osteological characters allow to identify it as Eurotrochilus sp. This genus is a stem group representative of Trochilidae and was recently described from the early Oligocene of southern Germany. The new fossil reveals that these European Trochilidae were remarkably modern in size, skeletal proportions and the shape of the wing, tail and beak and hyoid bones. These features confirm the early acquisition of the abilities of hovering and nectarivory in hummingbirds, probably before the Oligocene. In several morphological characteristics, they resemble members of the 'true hummingbirds' (subfamily Trochilinae) and differ from hermits (Phaethornithinae). These features, which include a short and square tail and a moderately long, almost straight beak, appear to be primitive within the family Trochilidae.

Asa Issie, Aramis and the origin of Australopithecus.

The origin of Australopithecus, the genus widely interpreted as ancestral to Homo, is a central problem in human evolutionary studies. Australopithecus species differ markedly from extant African apes and candidate ancestral hominids such as Ardipithecus, Orrorin and Sahelanthropus. The earliest described Australopithecus species is Au. anamensis, the probable chronospecies ancestor of Au. afarensis. Here we describe newly discovered fossils from the Middle Awash study area that extend the known Au. anamensis range into northeastern Ethiopia. The new fossils are from chronometrically controlled stratigraphic sequences and date to about 4.1-4.2 million years ago. They include diagnostic craniodental remains, the largest hominid canine yet recovered, and the earliest Australopithecus femur. These new fossils are sampled from a woodland context. Temporal and anatomical intermediacy between Ar. ramidus and Au. afarensis suggest a relatively rapid shift from Ardipithecus to Australopithecus in this region of Africa, involving either replacement or accelerated phyletic evolution.

A new hominid from the Upper Miocene of Chad, Central Africa.

The search for the earliest fossil evidence of the human lineage has been concentrated in East Africa. Here we report the discovery of six hominid specimens from Chad, central Africa, 2,500 km from the East African Rift Valley. The fossils include a nearly complete cranium and fragmentary lower jaws. The associated fauna suggest the fossils are between 6 and 7 million years old. The fossils display a unique mosaic of primitive and derived characters, and constitute a new genus and species of hominid. The distance from the Rift Valley, and the great antiquity of the fossils, suggest that the earliest members of the hominid clade were more widely distributed than has been thought, and that the divergence between the human and chimpanzee lineages was earlier than indicated by most molecular studies.