A gigantic bizarre marine turtle (Testudines: Chelonioidea) from the Middle Campanian (Late Cretaceous) of South-western Europe.

Marine turtles were common in the subtropical Upper Cretaceous epi-continental seas that once washed the coasts of the ancient European archipelago. But unlike its contemporaneous faunas from North America, in Europe no taxon surpassed the 1.5 m shell-length. Here, the remains of a new large marine turtle, Leviathanochelys aenigmatica gen. et sp. nov., from the Middle Campanian of the Southern Pyrenees are described. Anatomical and histological evidence concur in identifying the specimen as a basal chelonioid. The new taxon autapomorphically differs from other marine turtles by possessing an additional process on the anteromedial side of the pelvis, and an acetabulum directed strongly ventrally. Based on the pelvis size, it is likely that Leviathanochelys was as large as Archelon, thus becoming one of the largest marine turtles found to ever exist. The large body size of the new taxon could have evolved as a response to the unique habitat conditions of the European Cretaceous archipelago seas. The presence of the accessory pubic process further suggests the occurrence of an additional insertion point of the Musculus rectus abdominis, which together with the paleohistologic evidences support the hypothesis that the new taxon had an open marine pelagic lifestyle.

A titanosaurian sauropod with Gondwanan affinities in the latest Cretaceous of Europe.

The origin of the last sauropod dinosaur communities in Europe and their evolution during the final 15 million years of the Cretaceous have become a complex phylogenetic and palaeobiogeographic puzzle characterized by the controversy on the alleged coexistence of immigrant, Gondwana-related taxa alongside relictual and insular clades. In this context, we describe a new titanosaurian sauropod dinosaur, Abditosaurus kuehnei gen. et sp. nov., from the Late Cretaceous (Maastrichtian) Tremp Group of Catalonia (Spain). Phylogenetic analyses recover Abditosaurus separately from other European titanosaurs, within a clade of otherwise South American and African saltasaurines. The affinity of the new taxon with southern landmasses is reinforced by spatiotemporal co-occurrence with Gondwanan titanosaurian oospecies in southern Europe. The large size and the lack of osteohistological features potentially related to insular dwarfism or size reduction support the idea that Abditosaurus belongs to an immigrant lineage, unequivocally distinct from some of the island dwarfs of the European archipelago. The arrival of the Abditosaurus lineage to the Ibero-Armorican Island is hypothesized to have occurred during the earliest Maastrichtian (70.6 Ma), probably as a result of a global and regional sea-level drop that reactivated ancient dispersal routes between Africa and Europe. The arrival of large-bodied titanosaurs to the European archipelago produced dramatic changes in its insular ecosystems and important evolutionary changes in its dinosaur faunas, especially with respect to the 'island rule' effect.

A fast-growing basal troodontid (Dinosauria: Theropoda) from the latest Cretaceous of Europe.

A characteristic fauna of dinosaurs and other vertebrates inhabited the end-Cretaceous European archipelago, some of which were dwarves or had other unusual features likely related to their insular habitats. Little is known, however, about the contemporary theropod dinosaurs, as they are represented mostly by teeth or other fragmentary fossils. A new isolated theropod metatarsal II, from the latest Maastrichtian of Spain (within 200,000 years of the mass extinction) may represent a jinfengopterygine troodontid, the first reported from Europe. Comparisons with other theropods and phylogenetic analyses reveal an autapomorphic foramen that distinguishes it from all other troodontids, supporting its identification as a new genus and species, Tamarro insperatus. Bone histology shows that it was an actively growing subadult when it died but may have had a growth pattern in which it grew rapidly in early ontogeny and attained a subadult size quickly. We hypothesize that it could have migrated from Asia to reach the Ibero-Armorican island no later than Cenomanian or during the Maastrichtian dispersal events.

A small Cretaceous crocodyliform in a dinosaur nesting ground and the origin of sebecids.

Sebecosuchia was a group of highly specialized cursorial crocodyliforms that diversified during the Cretaceous and persist until the end of the Miocene. Their unique combination of cranial and post-cranial features indicates that they were active terrestrial predators that occupied the apex of the Late Cretaceous terrestrial ecosystems, even competing with theropod dinosaurs. Here, we report the discovery of the earliest sebecid worldwide, and the first from Eurasia, Ogresuchus furatus gen. et sp. nov., based on a semi-articulate specimen located in a titanosaurian sauropod nesting ground. The new taxon challenges current biogeographical models about the early dispersal and radiation of sebecid crocodylomorphs, and suggests an origin of the group much earlier than previously expected. Moreover, the new taxon suggests a potential convergent evolution between linages geographically isolated. Taphonomic evidences suggest that Ogresuchus died almost in the same place where fossilized, in a dinosaur nesting area. Biometric and morphologic observations lead to speculate that Ogresuchus could easily predate on sauropod hatchlings.

Correction: Dynamic Similarity in Titanosaur Sauropods: Ichnological Evidence from the Fumanya Dinosaur Tracksite (Southern Pyrenees).

[This corrects the article DOI: 10.1371/journal.pone.0057408.].

Ontogeny and taxonomy of the hadrosaur (Dinosauria, Ornithopoda) remains from Basturs Poble bonebed (late early Maastrichtian, Tremp Syncline, Spain).

The lower Maastrichtian site of Basturs Poble (southern Pyrenees, Spain) is the first hadrosaur bonebed reported from Europe. It is an accumulation of disarticulated lambeosaurine skeletal elements, possibly belonging to Pararhabdodon isonensis. The sample shows high intraspecific morphological variability among many skeletal elements, suggesting the need for caution in choosing characters for phylogenetic analyses. Juvenile to adult individuals are represented in the sample, while hatchling remains are absent. Bone histology reveals that juveniles are over-represented and that the youngest individuals represented by tibia specimens were two years old. Adult individuals, with tibiae 550-600 mm long, were 14-15 years old when they died. However, individual variation in tibia length at skeletal maturity occurs within the sample, so individual maturity cannot be assumed on the basis of bone size alone. The Basturs Poble bonebed occurs within the upper part of the C31r magnetochron. Thus, lambeosaurine hadrosaurids were already present and abundant in the Ibero-Armorica Island at the end of the early Maastrichtian and P. isonensis spans the upper part of the lower Maastrichtian to the upper part of the upper Maastrichtian (upper part of C31r-lower part of C29r).

Evidence of Reproductive Stress in Titanosaurian Sauropods Triggered by an Increase in Ecological Competition.

The occurrence of dinosaur pathologic eggs in the Late Cretaceous of Europe is well known, but their origin remains unclear. Here we expose the results of a detailed sampling of the conspicuous fossil record of Late Cretaceous titanosaurian eggs (oogenus Megaloolithius) from several southwestern Europe basins. After examining more than 450 samples, we observed a remarkable and statistically supported occurrence of multiple pathologic eggs in a relatively short stratigraphic range at the end of the early Maastrichtian, circa 71-70 Ma. All pathologic specimens exhibit multi-layered eggshell condition, a characteristic related to dystocia, or egg retention within the female uterus for an abnormal prolonged period of time. After exploring various scenarios, the occurrence of pathologic eggs is strongly correlated with an intense dinosaur faunal replacement that occurred during the early Maastrichtian in the Ibero-Armorican Island. Given that inter-species competiveness is proved to produce major affects in ecological communities, our results suggest that pathologies in the eggs of European titanosaurians could be a consequence of an increase in reproductive stress triggered by direct ecological competition between different dinosaurs. Thus, the present study provides a new perspective of how dinosaurs might have been affected by ecological/environmental disturbance.

Ichnological evidence of Megalosaurid Dinosaurs Crossing Middle Jurassic Tidal Flats.

A new dinosaur tracksite in the Vale de Meios quarry (Serra de Aire Formation, Bathonian, Portugal)preserves more than 700 theropod tracks. They are organized in at least 80 unidirectional trackways arranged in a bimodal orientation pattern (W/NW and E/SE). Quantitative and qualitative comparisons reveal that the large tridactyl, elongated and asymmetric tracks resemble the typical Late Jurassic-Early Cretaceous Megalosauripus ichnogenus in all morphometric parameters. Few of the numerous tracks are preserved as elite tracks while the rest are preserved as different gradients of modified true tracks according to water content, erosive factors, radial fractures and internal overtrack formations. Taphonomical determinations are consistent with paleoenvironmental observations that indicate an inter-tidal flat located at the margin of a coastal barrier. The Megalosauripus tracks represent the oldest occurrence of this ichnotaxon and are attributed to large megalosaurid dinosaurs. Their occurrence in Vale de Meios tidal flat represents the unique paleoethological evidence of megalosaurids moving towards the lagoon, most likley during the low tide periods with feeding purposes.

3D computational mechanics elucidate the evolutionary implications of orbit position and size diversity of early amphibians.

For the first time in vertebrate palaeontology, the potential of joining Finite Element Analysis (FEA) and Parametrical Analysis (PA) is used to shed new light on two different cranial parameters from the orbits to evaluate their biomechanical role and evolutionary patterns. The early tetrapod group of Stereospondyls, one of the largest groups of Temnospondyls is used as a case study because its orbits position and size vary hugely within the members of this group. An adult skull of Edingerella madagascariensis was analysed using two different cases of boundary and loading conditions in order to quantify stress and deformation response under a bilateral bite and during skull raising. Firstly, the variation of the original geometry of its orbits was introduced in the models producing new FEA results, allowing the exploration of the ecomorphology, feeding strategy and evolutionary patterns of these top predators. Secondly, the quantitative results were analysed in order to check if the orbit size and position were correlated with different stress patterns. These results revealed that in most of the cases the stress distribution is not affected by changes in the size and position of the orbit. This finding supports the high mechanical plasticity of this group during the Triassic period. The absence of mechanical constraints regarding the orbit probably promoted the ecomorphological diversity acknowledged for this group, as well as its ecological niche differentiation in the terrestrial Triassic ecosystems in clades as lydekkerinids, trematosaurs, capitosaurs or metoposaurs.

3D bite modeling and feeding mechanics of the largest living amphibian, the Chinese giant salamander Andrias davidianus (Amphibia:Urodela).

Biting is an integral feature of the feeding mechanism for aquatic and terrestrial salamanders to capture, fix or immobilize elusive or struggling prey. However, little information is available on how it works and the functional implications of this biting system in amphibians although such approaches might be essential to understand feeding systems performed by early tetrapods. Herein, the skull biomechanics of the Chinese giant salamander, Andrias davidianus is investigated using 3D finite element analysis. The results reveal that the prey contact position is crucial for the structural performance of the skull, which is probably related to the lack of a bony bridge between the posterior end of the maxilla and the anterior quadrato-squamosal region. Giant salamanders perform asymmetrical strikes. These strikes are unusual and specialized behavior but might indeed be beneficial in such sit-and-wait or ambush-predators to capture laterally approaching prey. However, once captured by an asymmetrical strike, large, elusive and struggling prey have to be brought to the anterior jaw region to be subdued by a strong bite. Given their basal position within extant salamanders and their "conservative" morphology, cryptobranchids may be useful models to reconstruct the feeding ecology and biomechanics of different members of early tetrapods and amphibians, with similar osteological and myological constraints.

Intra-trackway morphological variations due to substrate consistency: the El Frontal dinosaur tracksite (Lower Cretaceous, Spain).

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes in footprint parameters such as the pace length, the track length, depth, and height of displacement rims. Sedimentological analyses suggest that the original substrate was non-homogenous due to lateral changes in adjoining microfacies. Multidata analyses indicate that morphological differences in these deep and shallow tracks represent a part of a continuum of track morphologies and geometries produced by a gradient of substrate consistencies across the site. This implies that the large range of track morphologies at this site resulted from similar trackmakers crossing variable facies. The trackways at the El Frontal site present an exemplary case of how track morphology, and consequently potential ichnotaxa, can vary, even when produced by a single trackmaker.

The latest succession of dinosaur tracksites in Europe: Hadrosaur ichnology, track production and palaeoenvironments.

A comprehensive review and study of the rich dinosaur track record of the Tremp Formation in the southern Pyrenees of Spain (Southwestern Europe) shows a unique succession of footprint localities prior to the end-Cretaceous mass extinction event. A description of some 30 new tracksites and data on sedimentary environments, track occurrence and preservation, ichnology and chronostratigraphy are provided. These new track localities represent various facies types within a diverse set of fluvial environments. The footprint discoveries mostly represent hadrosaurian and, less abundantly, to sauropod dinosaurs. The hadrosaur tracks are significantly smaller in size than, but morphologically similar to, those of North America and Asia and are attributable to the ichnogenus Hadrosauropodus. The track succession, with more than 40 distinct track levels, indicates that hadrosaur footprints in the Ibero-Armorican region occur predominantly in the late Maaastrichtian (at least above the early Maastrichtian-late Maastrichtian boundary). The highest abundance is found noticeably found in the late Maastrichtian, with tracks occurring in the C29r magnetochron, within about the latest 300,000 years of the Cretaceous.

Diversity, relationships, and biogeography of the lambeosaurine dinosaurs from the European Archipelago, with description of the New Aralosaurin Canardia garonnensis.

We provide a thorough re-evaluation of the taxonomic diversity, phylogenetic relationships, and historical biogeography of the lambeosaurine hadrosaurids from the European Archipelago. Previously published occurrences of European Lambeosaurinae are reviewed and new specimens collected from upper Maastrichtian strata of the south-central Pyrenees are described. No support is found for the recognition of European saurolophines in the available hadrosaurid materials recovered so far from this area. A new genus and species of basal lambeosaurine, Canardia garonnensis, is described on the basis of cranial and appendicular elements collected from upper Maastrichtian strata of southern France. C. garonnensis differs from all other hadrosaurids, except Aralosaurus tuberiferus, in having maxilla with prominent subrectangular rostrodorsal flange; it differs from A. tuberiferus in a few maxillary and prefrontal characters. Together with A. tuberiferus, C. garonnensis integrates the newly recognized tribe Aralosaurini. Inference of lambeosaurine interrelationships via maximum parsimony analysis indicates that the other three known European lambeosaurines are representatives of two additional subclades (tribes) of these hadrosaurids: Tsintaosaurini (Pararhabdodon isonensis) and Lambeosaurini (the Arenysaurus ardevoli-Blasisaurus canudoi clade). The tribes Aralosaurini, Tsintaosaurini, Lambeosaurini, and Parasaurolophini are formally defined and diagnosed for the first time. Three event-based quantitative methods of ancestral range reconstruction were implemented to infer the historical biogeography of European lambeosaurines: Dispersal-Vicariance Analysis, Bayesian Binary MCMC, and Dispersal-Extinction-Cladogenesis. The results of these analyses, coupled with the absence of pre-Maastrichtian lambeosaurines in the Mesozoic vertebrate fossil record of Europe, favor the hypothesis that aralosaurins and tsintaosaurins were Asian immigrants that reached the Ibero-Armorican island via dispersal events sometime during the Maastrichtian. Less conclusive is the biogeographical history of European lambeosaurins; several scenarios, occurring sometime during the Maastrichtian, are possible, from vicariance leading to the splitting of Asian or North American from European ranges to a dispersal event from North America to the European Archipelago.

Dynamic similarity in titanosaur sauropods: ichnological evidence from the Fumanya dinosaur tracksite (southern Pyrenees).

The study of a small sauropod trackway from the Late Cretaceous Fumanya tracksite (southern Pyrenees, Catalonia) and further comparisons with larger trackways from the same locality suggest a causative relationship between gait, gauge, and body proportions of the respective titanosaur trackmakers. This analysis, conducted in the context of scaling predictions and using geometric similarity and dynamic similarity hypotheses, reveals similar Froude numbers and relative stride lengths for both small and large trackmakers from Fumanya. Evidence for geometric similarity in these trackways suggests that titanosaurs of different sizes moved in a dynamically similar way, probably using an amble gait. The wide gauge condition reported in trackways of small and large titanosaurs implies that they possessed similar body (trunk and limbs) proportions despite large differences in body size. These results strengthen the hypothesis that titanosaurs possessed a distinctive suite of anatomical characteristics that are well reflected in their tracks and trackways.

Manus track preservation bias as a key factor for assessing trackmaker identity and quadrupedalism in basal ornithopods.

BACKGROUND: The Las Cerradicas site (Tithonian-Berriasian), Teruel, Spain, preserves at least seventeen dinosaur trackways, some of them formerly attributed to quadrupedal ornithopods, sauropods and theropods. The exposure of new track evidence allows a more detailed interpretation of the controversial tridactyl trackways as well as the modes of locomotion and taxonomic affinities of the trackmakers. METHODOLOGY/PRINCIPAL FINDINGS: Detailed stratigraphic analysis reveals four different levels where footprints have been preserved in different modes. Within the tridactyl trackways, manus tracks are mainly present in a specific horizon relative to surface tracks. The presence of manus tracks is interpreted as evidence of an ornithopod trackmaker. Cross-sections produced from photogrammetric digital models show different depths of the pes and manus, suggesting covariance in loading between the forelimbs and the hindlimbs. CONCLUSIONS/SIGNIFICANCE: Several features (digital pads, length/width ratio, claw marks) of some ornithopod pes tracks from Las Cerradicas are reminiscent of theropod pedal morphology. This morphological convergence, combined with the shallow nature of the manus tracks, which reduces preservation potential, opens a new window into the interpretation of these tridactyl tracks. Thus, trackmaker assignations during the Jurassic-Cretaceous interval of purported theropod trackways may potentially represent ornithopods. Moreover, the Las Cerradicas trackways are further evidence for quadrupedalism among some basal small- to medium-sized ornithopods from this time interval.

Skull mechanics and the evolutionary patterns of the otic notch closure in capitosaurs (Amphibia: Temnospondyli).

Capitosaurs were among the largest amphibians that have ever lived. Their members displayed an amphibious lifestyle. We provide new information on functional morphology data, using finite element analysis (FEA) which has palaeoecological implications for the group. Our analyses included 17 taxa using (2D) plate models to test four loading cases (bilateral, unilateral and lateral bitings and skull raising system simulation). Our results demonstrates that, when feeding, capitosaurs concentrated the stress at the circumorbital region of the capitosaur skull and cranial sutures probably played a key role in dissipating and absorbing the stress generated during biting. Basal members (as Wetlugasaurus) were probably less specialized forms, while during Middle- and Late Triassic the group radiated into different ecomorphotypes with closed otic notch forms (as Cyclotosaurus) resulting in the strongest skulls during biting. Previous interpretations discussed a trend from an open to closed otic notch associated with lateral repositioning of the tabular horns, but the analysis of the skull-raising system reveals that taxa exhibiting posteriorly directed tabular horns display similar results during skull raising to those of closed otic notch taxa. Our results suggest that various constraints besides otic notch morphology, such as the elongation of the tabular horns, snout length, skull width and position, and size of the orbits affect the function of the skull. On the light of our results, capitosaur skull showed a trend to reduce the stresses and deformation during biting. Capitosaurs could be considered crocodilian analogues as they were top-level predators in fluvial and brackish Triassic ecosystems.

3-D modelling of megaloolithid clutches: insights about nest construction and dinosaur behaviour.

BACKGROUND: Megaloolithid eggs have long been associated with sauropod dinosaurs. Despite their extensive and worldwide fossil record, interpretations of egg size and shape, clutch morphology, and incubation strategy vary. The Pinyes locality in the Upper Cretaceous Tremp Formation in the southern Pyrenees, Catalonia provides new information for addressing these issues. Nine horizons containing Megaloolithus siruguei clutches are exposed near the village of Coll de Nargó. Tectonic deformation in the study area strongly influenced egg size and shape, which could potentially lead to misinterpretation of reproductive biology if 2D and 3D maps are not corrected for bed dip that results from tectonism. METHODOLOGY/FINDINGS: Detailed taphonomic study and three-dimensional modelling of fossil eggs show that intact M. siruguei clutches contained 20-28 eggs, which is substantially larger than commonly reported from Europe and India. Linear and grouped eggs occur in three superimposed levels and form an asymmetric, elongate, bowl-shaped profile in lateral view. Computed tomography data support previous interpretations that the eggs hatched within the substrate. Megaloolithid clutch sizes reported from other European and Indian localities are typically less than 15 eggs; however, these clutches often include linear or grouped eggs that resemble those of the larger Pinyes clutches and may reflect preservation of incomplete clutches. CONCLUSIONS/SIGNIFICANCE: We propose that 25 eggs represent a typical megaloolithid clutch size and smaller egg clusters that display linear or grouped egg arrangements reported at Pinyes and other localities may represent eroded remnants of larger clutches. The similarity of megaloolithid clutch morphology from localities worldwide strongly suggests common reproductive behaviour. The distinct clutch geometry at Pinyes and other localities likely resulted from the asymmetrical, inclined, and laterally compressed titanosaur pes unguals of the female, using the hind foot for scratch-digging during nest excavation.