The last known freshwater coelacanths: New Late Cretaceous mawsoniid remains (Osteichthyes: Actinistia) from Southern France.

Coelacanths are iconic fishes represented today by a single marine genus. The group was a little bit more diversified in the Mesozoic, with representatives in marine and continental environments in the Late Cretaceous. Here we describe isolated skull bones of the last know freshwater coelacanths found in several fossil sites from the Early Campanian to the Early Maastrichtian of Southern France (in the Departments of Aude, Bouches-du-Rhône, Hérault, and Var). The sample does not allow distinguishing different species, and all material is referred to Axelrodichthys megadromos Cavin, Valentin, Garcia originally described from the locality of Ventabren in Southern France. A reconstruction of the skull is proposed. Previously unrecognized features are described, including parts of the postparietal portion of the skull, of the suspensorium and of the mandible. The new data confirm the assignation of the species to the mawsoniids, and more specifically to Axelrodichthys. A cladistic analysis scoring new character states provides a similar topology than a previous analysis, i.e. A. megadromos is placed in a polytomy with Axelrodichthys araripensis and Lualabaea lerichei, two species from the Early Cretaceous of Brazil and from the Late Jurassic of the Democratic Republic of the Congo, respectively. A. megadromos appears to have been restricted to freshwater environments, to the contrary of oldest Western Gondwanan representatives of the family that were able to live in brackish and marine waters. A. megadromos is the last representative of the mawsoniids and its occurrence in Europe is probably the result of a dispersal event from Western Gondwana that happened somewhen in the Cretaceous. Based on the available data, the mawsoniids went extinct in the mid-Maastrichthian, i.e. before the end-Cretaceous mass extinction. But it is possible that the fossil record of this family, which has been only recently recognized in Late Cretaceous European deposits, will geographically and stratigraphically widen with further discoveries.

Oxygen isotope fractionation between bird bone phosphate and drinking water.

Oxygen isotope compositions of bone phosphate (δ18Op) were measured in broiler chickens reared in 21 farms worldwide characterized by contrasted latitudes and local climates. These sedentary birds were raised during an approximately 3 to 4-month period, and local precipitation was the ultimate source of their drinking water. This sampling strategy allowed the relationship to be determined between the bone phosphate δ18Op values (from 9.8 to 22.5‰ V-SMOW) and the local rainfall δ18Ow values estimated from nearby IAEA/WMO stations (from -16.0 to -1.0‰ V-SMOW). Linear least square fitting of data provided the following isotopic fractionation equation: δ18Ow = 1.119 (±0.040) δ18Op - 24.222 (±0.644); R 2 = 0.98. The δ18Op-δ18Ow couples of five extant mallard ducks, a common buzzard, a European herring gull, a common ostrich, and a greater rhea fall within the predicted range of the equation, indicating that the relationship established for extant chickens can also be applied to birds of various ecologies and body masses. Applied to published oxygen isotope compositions of Miocene and Pliocene penguins from Peru, this new equation computes estimates of local seawater similar to those previously calculated. Applied to the basal bird Confuciusornis from the Early Cretaceous of Northeastern China, our equation gives a slightly higher δ18Ow value compared to the previously estimated one, possibly as a result of lower body temperature. These data indicate that caution should be exercised when the relationship estimated for modern birds is applied to their basal counterparts that likely had a metabolism intermediate between that of their theropod dinosaur ancestors and that of advanced ornithurines.

Oxygen isotope fractionation between bird eggshell calcite and body water: application to fossil eggs from Lanzarote (Canary Islands).

Oxygen and carbon isotope compositions of fossil bird eggshell calcite (δ(18)Ocalc and δ(13)Ccalc) are regularly used to reconstruct paleoenvironmental conditions. However, the interpretation of δ(18)Ocalc values of fossil eggshells has been limited to qualitative variations in local climatic conditions as oxygen isotope fractionations between calcite, body fluids, and drinking water have not been determined yet. For this purpose, eggshell, albumen water, and drinking water of extant birds have been analyzed for their oxygen and carbon isotope compositions. Relative enrichments in (18)O relative to (16)O between body fluids and drinking water of +1.6 ± 0.9 ‰ for semi-aquatic birds and of +4.4 ± 1.9 ‰ for terrestrial birds are observed. Surprisingly, no significant dependence to body temperature on the oxygen isotope fractionation between eggshell calcite and body fluids is observed, suggesting that bird eggshells precipitate out of equilibrium. Two empirical equations relating the δ(18)Ocalc value of eggshell calcite to the δ(18)Ow value of ingested water have been established for terrestrial and semi-aquatic birds. These equations have been applied to fossil eggshells from Lanzarote in order to infer the ecologies of the Pleistocene marine bird Puffinus sp. and of the enigmatic giant birds from the Pliocene. Both δ(13)Ccalc and δ(18)Ocalc values of Puffinus eggshells point to a semi-aquatic marine bird ingesting mostly seawater, whereas low δ(13)Ccalc and high δ(18)Ocalc values of eggshells from the Pliocene giant bird suggest a terrestrial lifestyle. This set of equations can help to quantitatively estimate the origin of waters ingested by extinct birds as well as to infer either local environmental or climatic conditions.

Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

Hadrosauroid Dinosaurs from the Late Cretaceous of the Sultanate of Oman.

Fragmentary post-cranial remains (femora, tibia, vertebrae) of ornithischian dinosaurs from the Late Cretaceous of the Sultanate of Oman are described and referred to hadrosauroids. The specimens come from the Al-Khod Conglomerate, of latest Campanian to Maastrichtian age, in the north-eastern part of the country. Although the fragmentary condition of the fossils precludes a precise identification, various characters, including the shape of the fourth trochanter of the femur and the morphology of its distal end, support an attribution to hadrosauroids. With the possible exception of a possible phalanx from Angola, this group of ornithopod dinosaurs, which apparently originated in Laurasia, was hitherto unreported from the Afro-Arabian plate. From a paleobiogeographical point of view, the presence of hadrosauroids in Oman in all likelihood is a result of trans-Tethys dispersal from Asia or Europe, probably by way of islands in the Tethys shown on all recent paleogeographical maps of that area. Whether hadrosauroids were widespread on the Afro-Arabian landmass in the latest Cretaceous, or where restricted to the « Oman island ¼ shown on some paleogeographical maps, remains to be determined.

Evidence of Egg Diversity in Squamate Evolution from Cretaceous Anguimorph Embryos.

Lizards are remarkable amongst amniotes, for they display a unique mosaic of reproduction modes ranging from egg-laying to live-bearing. Within this patchwork, geckoes are believed to represent the only group to ever have produced fully calcified rigid-shelled eggs, contrasting with the ubiquitous parchment shelled-eggs observed in other lineages. However, this hypothesis relies only on observations of modern taxa and fossilised gecko-like eggshells which have never been found in association with any embryonic or parental remains. We report here the first attested fossil eggs of lizards from the Early Cretaceous of Thailand, combining hard eggshells with exquisitely preserved embryos of anguimoph (e.g. Komodo dragons, mosasaurs). These fossils shed light on an apparently rare reproduction strategy of squamates, demonstrate that the evolution of rigid-shelled eggs are not an exclusive specialization of geckoes, and suggest a high plasticity in the reproductive organs mineralizing eggshells.

Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago.

The Late Cretaceous was a time of tremendous global change, as the final stages of the Age of Dinosaurs were shaped by climate and sea level fluctuations and witness to marked paleogeographic and faunal changes, before the end-Cretaceous bolide impact. The terrestrial fossil record of Late Cretaceous Europe is becoming increasingly better understood, based largely on intensive fieldwork over the past two decades, promising new insights into latest Cretaceous faunal evolution. We review the terrestrial Late Cretaceous record from Europe and discuss its importance for understanding the paleogeography, ecology, evolution, and extinction of land-dwelling vertebrates. We review the major Late Cretaceous faunas from Austria, Hungary, France, Spain, Portugal, and Romania, as well as more fragmentary records from elsewhere in Europe. We discuss the paleogeographic background and history of assembly of these faunas, and argue that they are comprised of an endemic 'core' supplemented with various immigration waves. These faunas lived on an island archipelago, and we describe how this insular setting led to ecological peculiarities such as low diversity, a preponderance of primitive taxa, and marked changes in morphology (particularly body size dwarfing). We conclude by discussing the importance of the European record in understanding the end-Cretaceous extinction and show that there is no clear evidence that dinosaurs or other groups were undergoing long-term declines in Europe prior to the bolide impact.

Insight into the growth dynamics and systematic affinities of the Late Cretaceous Gargantuavis from bone microstructure.

Enigmatic avialan remains of Gargantuavis philoinos from the Ibero-Armorican island of the Late Cretaceous European archipelago (Southern France) led to a debate concerning its taxonomic affinities. Here, we show that the bone microstructure of Gargantuavis resembles that of Apteryx, the extinct emeids and Megalapteryx from New Zealand, and indicates that like these slow-growing terrestrial birds, it took several years to attain skeletal maturity. Our findings suggest that the protracted cyclical growth in these ornithurines may have been in response to insular evolution.

First Clarkforkian equivalent Land Mammal Age in the latest Paleocene basal Sparnacian facies of Europe: fauna, flora, paleoenvironment and (bio)stratigraphy.

The Paleocene-Eocene Thermal Maximum (PETM) is correlated with the first occurrences of earliest modern mammals in the Northern Hemisphere. The latest Paleocene Clarkforkian North American Land Mammal Age, that has yielded rodents and carnivorans, is the only exception to this rule. However, until now no pre-PETM localities have yielded modern mammals in Europe or Asia. We report the first Clarkforkian equivalent Land Mammal Age in the latest Paleocene deposits of the basal Sparnacian facies at Rivecourt, in the north-central part of the Paris Basin. The new terrestrial vertebrate and macroflora assemblages are analyzed through a multidisciplinary study including sedimentologic, stratigraphic, isotopic, and palynological aspects in order to reconstruct the paleoenvironment and to evaluate biochronologic and paleogeographic implications. The mammals are moderately diverse and not abundant, contrary to turtles and champsosaurs. The macroflora is exceptional in preservation and diversity with numerous angiosperms represented by flowers, fruits, seeds and wood preserved as lignite material, revealing an abundance of Arecaceae, Betulaceae, Icacinaceae, Menispermaceae, Vitaceae and probably Cornaceae. Results indicate a Late Paleocene age based on carbon isotope data, palynology and vertebrate occurrences such as the choristoderan Champsosaurus, the arctocyonid Arctocyon, and the plesiadapid Plesiadapis tricuspidens. However, several mammal species compare better with the earliest Eocene. Among these, the particular louisinid Teilhardimys musculus, also recorded from the latest Paleocene of the Spanish Pyrenees, suggests a younger age than the typical MP6 reference level. Nevertheless, the most important aspect of the Rivecourt fauna is the presence of dental remains of a rodent and a "miacid" carnivoran, attesting to the presence of two modern mammalian orders in the latest Paleocene of Europe. Interestingly, these two groups are also the only modern groups recorded from the latest Paleocene of North America, making Rivecourt the first direct equivalent to the Clarkforkian Land Mammal Age outside of North America.

"Terror birds" (Phorusrhacidae) from the Eocene of Europe imply trans-Tethys dispersal.

BACKGROUND: Phorusrhacidae was a clade including middle-sized to giant terrestrial carnivorous birds, known mainly from the Cenozoic of South America, but also occurring in the Plio-Pleistocene of North America and the Eocene of Africa. Previous reports of small phorusrhacids in the Paleogene of Europe have been dismissed as based on non-phorusrhacid material. METHODOLOGY: we have re-examined specimens of large terrestrial birds from the Eocene (late Lutetian) of France and Switzerland previously referred to gastornithids and ratites and have identified them as belonging to a phorusrhacid for which the name Eleutherornis cotei should be used. CONCLUSIONS/SIGNIFICANCE: The occurrence of a phorusrhacid in the late Lutetian of Europe indicates that these flightless birds had a wider geographical distribution than previously recognized. The likeliest interpretation is that they dispersed from Africa, where the group is known in the Eocene, which implies crossing the Tethys Sea. The Early Tertiary distribution of phorusrhacids can be best explained by transoceanic dispersal, across both the South Atlantic and the Tethys.

Gavialis from the Pleistocene of Thailand and its relevance for drainage connections from India to Java.

BACKGROUND: The genus Gavialis comprises a single living but endangered species, G. gangeticus, as well as fossil species recorded in the Miocene to Pleistocene deposits of the Indian subcontinent. The genus is also represented in the Pleistocene deposits of Java by the species G. bengawanicus, which was recently recognized to be valid. Surprisingly, no detailed report of the genus exists between these two provinces and the recent evolutionary history of Gavialis is not understood. METHODOLOGY/PRINCIPAL FINDINGS: We report new material consisting of skull and mandibular remains of Gavialis from the Early Pleistocene of Khok Sung, Nakhon Ratchasima Province, northeastern Thailand. The Gavialis material described herein is attributed to Gavialis cf. bengawanicus and sheds new light on the occurrence of the genus in mainland SE Asia. CONCLUSIONS/SIGNIFICANCE: Comparison of this new material with other species referred to the genus Gavialis led us to preliminary restrict the content of the genus to three species, namely G. gangeticus Gmelin, G. bengawanicus Dubois and G. lewisi Lull. The occurrence of G. cf. bengawanicus in Thailand allows us to propose a scenario for the dispersal of Gavialis from Indo-Pakistan to Indonesia, thus bridging a geographical gap between these two provinces. Dispersal by sea appears a less likely possibility than dispersal through fluvial drainages.

Modified laminar bone in Ampelosaurus atacis and other Titanosaurs (Sauropoda): implications for life history and physiology.

BACKGROUND: Long bone histology of the most derived Sauropoda, the Titanosauria suggests that titanosaurian long bone histology differs from the uniform bone histology of basal Sauropoda. Here we describe the long bone histology of the titanosaur Ampelosaurus atacis and compare it to that of basal neosauropods and other titanosaurs to clarify if a special titanosaur bone histology exists. METHODOLOGY/PRINCIPAL FINDINGS: Ampelosaurus retains the laminar vascular organization of basal Sauropoda, but throughout most of cortical growth, the scaffolding of the fibrolamellar bone, which usually is laid down as matrix of woven bone, is laid down as parallel-fibered or lamellar bone matrix instead. The remodeling process by secondary osteons is very extensive and overruns the periosteal bone deposition before skeletal maturity is reached. Thus, no EFS is identifiable. Compared to the atypical bone histology of Ampelosaurus, the large titanosaur Alamosaurus shows typical laminar fibrolamellar bone. The titanosaurs Phuwiangosaurus, Lirainosaurus, and Magyarosaurus, although differing in certain features, all show this same low amount or absence of woven bone from the scaffolding of the fibrolamellar bone, indicating a clear reduction in growth rate resulting in a higher bone tissue organization. To describe the peculiar primary cortical bone tissue of Phuwiangosaurus, Ampelosaurus, Lirainosaurus, and Magyarosaurus, we here introduce a new term, "modified laminar bone" (MLB). CONCLUSIONS/SIGNIFICANCE: Importantly, MLB is as yet not known from extant animals. At least in Lirainosaurus and Magyarosaurus the reduction of growth rate indicated by MLB is coupled with a drastic body size reduction and maybe also a reduction in metabolic rate, interpreted as a result of dwarfing on the European islands during the Late Cretaceous. Phuwiangosaurus and Ampelosaurus both show a similar reduction in growth rate but not in body size, possibly indicating also a reduced metabolic rate. The large titanosaur Alamosaurus, on the other hand, retained the plesiomorphic bone histology of basal neosauropods.

Phase contrast synchrotron microtomography: improving noninvasive investigations of fossil embryos in ovo.

Fossil embryos are paramount for our understanding of the development of extinct species. However, although thousands of fossil amniote eggs are known, very few embryos in ovo have been described. First reports of fossil embryos were based on broken eggs, where the embryonic remains were already exposed, because destructive methods on complete eggs were avoided. Investigations of complete eggs therefore required nondestructive approaches, such as X-ray microtomography (µCT). However, due to the general low density contrast between fossilized bones and infilling matrix, only a few specimens have been reported using these techniques. Using propagation phase contrast X-ray synchrotron microtomography (PPC-SR-µCT), we report here the discovery of three well-preserved embryos in Early Cretaceous eggs from Thailand. By scanning these eggs using different imaging techniques, we show that vastly different interpretations can be made regarding the preservation state and/or the developmental stage of these embryos. PPC-SR-µCT also revealed differential contrast between bone categories, presumably reflecting the ossification pattern of these embryos. Applying such an approach to large-scale studies of fossil eggs could lead to more discoveries and detailed studies of fossil embryos, providing important developmental and phylogenetic information on extinct and extant amniotes.

Oxygen isotopes of East Asian dinosaurs reveal exceptionally cold Early Cretaceous climates.

Early Cretaceous vertebrate assemblages from East Asia and particularly the Jehol Biota of northeastern China flourished during a period of highly debated climatic history. While the unique characters of these continental faunas have been the subject of various speculations about their biogeographic history, little attention has been paid to their possible climatic causes. Here we address this question using the oxygen isotope composition of apatite phosphate (δ ) from various reptile remains recovered from China, Thailand, and Japan. δ values indicate that cold terrestrial climates prevailed at least in this part of Asia during the Barremian-early Albian interval. Estimated mean air temperatures of about 10 ± 4 °C at midlatitudes (∼ 42 °N) correspond to present day cool temperate climatic conditions. Such low temperatures are in agreement with previous reports of cold marine temperatures during this part of the Early Cretaceous, as well as with the widespread occurrence of the temperate fossil wood genus Xenoxylon and the absence of thermophilic reptiles such as crocodilians in northeastern China. The unique character of the Jehol Biota is thus not only the result of its evolutionary and biogeographical history but is also due to rather cold local climatic conditions linked to the paleolatitudinal position of northeastern China and global icehouse climates that prevailed during this part of the Early Cretaceous.

The end of the fat dodo? A new mass estimate for Raphus cucullatus.

A new mass estimate for the dodo (Raphus cucullatus), based on the lengths of the femur, tibiotarsus and tarsometatarsus, is attempted. The obtained mean mass is 10.2 kg, which is less than previous estimates based on other methods, which ranged from 10.6 to 21.1 kg, and much lower than the 50 lbs reported by a seventeenth-century eyewitness. The new estimated mass, which is similar to that of a large wild turkey, seems more realistic than previous ones and supports the hypothesis that contemporary illustrations of extremely fat dodos were either exaggerations, or based on overfed specimens. Pictures of "fat" dodos may also have been based on individuals exhibiting a display behaviour with puffed out feathers.

Regulation of body temperature by some Mesozoic marine reptiles.

What the body temperature and thermoregulation processes of extinct vertebrates were are central questions for understanding their ecology and evolution. The thermophysiologic status of the great marine reptiles is still unknown, even though some studies have suggested that thermoregulation may have contributed to their exceptional evolutionary success as apex predators of Mesozoic aquatic ecosystems. We tested the thermal status of ichthyosaurs, plesiosaurs, and mosasaurs by comparing the oxygen isotope compositions of their tooth phosphate to those of coexisting fish. Data distribution reveals that these large marine reptiles were able to maintain a constant and high body temperature in oceanic environments ranging from tropical to cold temperate. Their estimated body temperatures, in the range from 35 degrees +/- 2 degrees C to 39 degrees +/- 2 degrees C, suggest high metabolic rates required for predation and fast swimming over large distances offshore.

A two-headed reptile from the Cretaceous of China.

A malformed embryonic or neonate choristoderan reptile from the Lower Cretaceous Yixian Formation of northeastern China is described. The tiny skeleton exhibits two heads and two necks, with bifurcation at the level of the pectoral girdle. In a fossil, this is the first occurrence of the malformation known as axial bifurcation, which is well known in living reptiles.

First nonavian dinosaur from Lebanon: a brachiosaurid sauropod from the Lower Cretaceous of the Jezzine District.

Two sauropod teeth from an Early Cretaceous (Neocomian) fluviodeltaic sandstone near Jezzine (Southern Lebanon) are the first nonavian dinosaur remains to be reported from Lebanon. Their distinctive character places them within Brachiosauridae. The sauropod teeth from Lebanon are a significant addition to the very scanty dinosaur record from the Levant, which hitherto consisted mainly of very poorly preserved and not easily identifiable specimens. The Basal Cretaceous Sandstone of Lebanon, thus, appears to be a potentially important source of fossil vertebrate material.

Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition.

We report on very small fossil eggs from the Lower Cretaceous of Thailand, one of them containing a theropod embryo, which display a remarkable mosaic of characters. While the surficial ornamentation is typical of non-avian saurischian dinosaurs, the three-layered prismatic structure of the eggshell is currently known only in extant and fossil eggs associated with birds. These eggs, about the size of a goldfinch's, mirror at the reproductive level the retention of small body size that was paramount in the transition from non-avian theropods to birds. The egg-layer may have been a small feathered theropod similar to those recently found in China.