Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

Atoposaurids are a group of small-bodied, extinct crocodyliforms, regarded as an important component of Jurassic and Cretaceous Laurasian semi-aquatic ecosystems. Despite the group being known for over 150 years, the taxonomic composition of Atoposauridae and its position within Crocodyliformes are unresolved. Uncertainty revolves around their placement within Neosuchia, in which they have been found to occupy a range of positions from the most basal neosuchian clade to more crownward eusuchians. This problem stems from a lack of adequate taxonomic treatment of specimens assigned to Atoposauridae, and key taxa such as Theriosuchus have become taxonomic ‘waste baskets’. Here, we incorporate all putative atoposaurid species into a new phylogenetic data matrix comprising 24 taxa scored for 329 characters. Many of our characters are heavily revised or novel to this study, and several ingroup taxa have never previously been included in a phylogenetic analysis. Parsimony and Bayesian approaches both recover Atoposauridae as a basal clade within Neosuchia, more stemward than coelognathosuchians, bernissartiids, and paralligatorids. Atoposauridae is a much more exclusive clade than previously recognized, comprising just three genera (Alligatorellus, Alligatorium, and Atoposaurus) that were restricted to the Late Jurassic of western Europe, and went extinct at the Jurassic/Cretaceous boundary. A putative Gondwanan atoposaurid (Brillanceausuchus) is recovered as a paralligatorid. Our results exclude both Montsecosuchus and Theriosuchus from Atoposauridae. Theriosuchus is polyphyletic, forming two groupings of advanced neosuchians. Theriosuchus (restricted to Theriosuchus pusillus, Theriosuchus guimarotae, and Theriosuchus grandinaris) spanned the Middle Jurassic to early Late Cretaceous, and is known from Eurasia and North Africa. Two Cretaceous species previously assigned to Theriosuchus (‘Theriosuchus’ ibericus and ‘Theriosuchus’ sympiestodon) are shown to be nested within Paralligatoridae, and we assign them to the new genus Sabresuchus. The revised phylogenetic placement of Theriosuchus has several implications for our understanding of eusuchian evolution. Firstly, the presence of fully pterygoidean choanae, previously regarded as a defining characteristic of Eusuchia, is not found in some basal members of Eusuchia. However, eusuchians can be distinguished from Theriosuchus and other basal neosuchians in that their choanae are posteriorly positioned, with an anterior margin medial to the posterior edge of the suborbital fenestra. This feature distinguishes eusuchians from Theriosuchus and more basal neosuchians. Secondly, our refined understanding of Theriosuchus implies that this taxon possessed only amphicoelous presacral vertebrae, and therefore fully developed vertebral procoely is likely to have evolved only once in Crocodylomorpha, on the lineage leading to Eusuchia. These and other findings presented herein will provide an important framework for understanding the neosuchian–eusuchian transition

The first “Protosuchian” (Archosauria: Crocodyliformes) from the Cretaceous (Santonian) of Gondwana

El primer “protosuquio” (Archosauria: Crocodyliformes) del Cretácico (Santoniano) de Gondwana. Los restos de “protosuquios” del Cretácico provienen, a excepción del “crocodyliforme de Las Hoyas” del Cretácico Inferior de España, exclusivamente de Asia Central: Zaraasuchus, Gobiosuchus, Zosuchus y Artzosuchus del Cretácico Superior de Mongolia; Tagarosuchus del Cretácico Inferior del sur de Siberia; Edentosuchus, Sichuanosuchus y Shantungosuchus del Cretácico Inferior de China. Aquí reportamos un nuevo taxón de crocodyliforme basal, Neuquensuchus universitas gen.nov., sp.nov., de la provincia de Neuquén, Argentina, correspondiente a la Formación Bajo de la Carpa, representando el primer y único “protosuquio” del Cretácico de Gondwana. Los materiales fragmentarios y articulados corresponden a una especie esbelta y delgada, con extremidades largas y delgadas. Al igual que en Shantungosuchus, los centros cervicales son alargados, con una quilla ventral prominente y parapófisis anteroventrales bien desarrolladas. Como en los crocodyliformes basales, Neuquensuchus posee dos vértebras sacras. Además, una hoja escapular muy expandida, con un puente acromial bien desarrollado y el borde posterior similar a Sichuanosuchus. La cresta deltopectoral pronunciada en el húmero es equivalente a la de Sichuanosuchus y al igual que este, la diáfisis es circular, alargada y delgada con el cóndilo medial mayor que el lateral. Asimismo, las proporciones del radio y la úlna son similares a Sichuanosuchus. Como este, el pubis es alargado, muy delgado en su sección media y poco expandido distalmente. El fémur, tibia y fíbula son alargados y similares a otros crocodyliformes no derivados. Además de representar el primer “protosuquio” cretácico de Gondwana, su presencia fuera de Asia y Europa durante el Cretácico ofrece nueva evidencia de un evento de dispersión pre-Albiano entre Gondwana y Asia Central a través de Europa.The remains of “protosuchians” from the Cretaceous come, to exception of “Las Hoyas crocodyliform” from the Lower Cretaceous of Spain, exclusively of Central Asia: Zaraasuchus, Gobiosuchus, Zosuchus, and Artzosuchus from the Upper Cretaceous of Mongolia; Tagarosuchus from Lower Cretaceous of Southern Siberia; Edentosuchus, Sichuanosuchus, and Shantungosuchus from Lower Cretaceous of China. We report a new basal crocodyliform taxon, Neuquensuchus universitas gen.nov., sp.nov., from Neuquén Province, Argentina, belonging to Bajo de la Carpa Formation, representing the first and only “protosuchian” from the Cretaceous of Gondwana. The articulated and fragmentary materials belonged to a willowy, slender species, with very long and thin extremities. As in Shantungosuchus, the cervical centers are lengthened, with prominent ventral keel and well developed anteroventral parapophyses. As in basal crocodylomorphs, it possesses two sacral vertebrae. Also, a much enlarged scapular blade, with well developed acromial ridge and the posterior edge similar to Sichuanosuchus. The pronounced deltopectoral crest in the complete humerus is equivalent to Sichuanosuchus and as this, a circular, elongated and thin shaft with the medial condyle longer than the lateral one. Also, the complete ulna and radius is similar in their proportions to Sichuanosuchus. As this, the pubis is lengthened, very thin in the half section and not very expanded distally. The femur, tibia and fibula are elongated and similar to other non-derivated crocodyliforms. Besides representing the first Cretaceous “protosuchian” of Gondwana, the occurrence of these outside of Asia and Europe during the Cretaceous offers new evidence of pre-Albian dispersion between Gondwana and Central Asia through Europe.Fil: Fiorelli, Lucas Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de Catamarca. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Secretaría de Industria y Minería. Servicio Geológico Minero Argentino. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Provincia de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja; ArgentinaFil: Calvo, Jorge Orlando. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad Nacional del Comahue. Centro Paleontológico Lago Barreales; Argentin

Novos restos de Notosuchus terrestris Woodward, 1896 (Crocodyliformes: Mesoeucrocodylia) do Cretáceo Superior de Neuquén, Patagônia, Argentina

New materials of Notosuchus terrestris are here described. They were found on Bajo de la Carpa Formation outcrops, near the Universidad Nacional del Comahue, Neuquén City. Descriptions were based on five specimens, one of them the only specimen of Notosuchus with postcranial remains articulated to the skull. As in Sphagesaurus, it presents triangular teeth in cross-section and oblique molariforms with worn facet surface. As in Mariliasuchus, it possesses procumbent mandibular incisiform teeth and, like in other notosuchians and basal crocodyliforms, it was able of proal mandibular movement. The centra of cervical vertebrae possess ventral keel as in Chimaerasuchus. Elongated cervical neural spines and suprapostzygapophyseal laminae in cervicodorsal vertebrae are observed. The scapular dorsal end is greatly enlarged, while the coracoid ventromedial process end is moderately developed. The dorsal surface of the ilium is lateromedially wide with a greatly expanded acetabular roof and a prominent anteromedial process in the femoral shaft. Based on diverse cranial and postcranial characters, we infer that Notosuchus possessed facial and perioral musculature well developed and an herbivore diet, confirming the suggestions of previous authors. Notosuchus represents, based on phylogenetic studies, the sister taxon of Mariliasuchus and the monophyly of Notosuchia is demonstrated. Paleobiogeographycally, the occurrence of Chimaerasuchus in China evidences the faunistic interchange between Gondwana and Central Asia during the Early Cretaceous.Novos materiais de Notosuchus terrestris são aqui descritos. Eles provêm de afloramentos da Formação Bajo de la Carpa, localizados próximos à Universidad Nacional del Comahue, na cidade de Neuquén. As descrições foram baseadas em cinco exemplares, um deles o único espécime de Notosuchus com restos pós-cranianos articulados ao crânio. Como em Sphagesaurus, N. terrestris apresenta dentes triangulares em seção cruzada e molariformes oblíquos com superfície da faceta com desgaste. Como em Mariliasuchus, a espécie possui dentes mandibulares incisiformes procumbentes e, como em outros notossúquios e crocodiliformes basais, era possível realizar o movimento antero-posterior mandibular. Os centros das vértebras cervicais possuem uma quilha ventral como em Chimaerasuchus. Espinhos neurais cervicais alongados e lâminas suprapószigapofiseais em vértebras cérvico-dorsais são observados. A extremidade escapular dorsal é amplamente alargada, enquanto a extremidade do processo ventro-medial do coracóide é pouco desenvolvida. A superfície dorsal do ilium é larga látero-medialmente com um teto acetabular amplamente expandido e processo ântero-medial proeminente na diáfise femoral. Baseado em diversos caracteres cranianos e pós-cranianos, infere-se que Notosuchus apresentava grande desenvolvimento da musculatura facial e perioral e tinha uma dieta herbívora, confirmando o que foi sugerido anteriormente por outros autores. Notosuchus representa, baseado em estudos filogenéticos, o táxon irmão de Mariliasuchus e a monofilia de Notosuchia é demonstrada. Paleobiogeograficamente, a ocorrência de Chimaerasuchus na China evidencia o intercâmbio faunístico entre o Gondwana e a Ásia Central durante o Cretáceo Inferior.Fil: Fiorelli, Lucas Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Universidad Nacional de Catamarca. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Secretaría de Industria y Minería. Servicio Geológico Minero Argentino. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja. - Provincia de La Rioja. Centro Regional de Investigaciones Científicas y Transferencia Tecnológica de La Rioja; Argentina. Universidad Nacional del Comahue. Centro Paleontológico Lago Barreales; ArgentinaFil: Calvo, Jorge. Universidad Nacional del Comahue. Centro Paleontológico Lago Barreales; Argentin

The first definitive Middle Jurassic atoposaurid (Crocodylomorpha, Neosuchia), and a discussion on the genus Theriosuchus

Atoposaurids were a clade of semiaquatic crocodyliforms known from the Late Jurassic to the latest Cretaceous. Tentative remains from Europe, Morocco, and Madagascar may extend their range into the Middle Jurassic. Here we report the first unambiguous Middle Jurassic (late Bajocian–Bathonian) atoposaurid: an anterior dentary from the Isle of Skye, Scotland, UK. A comprehensive review of atoposaurid specimens demonstrates that this dentary can be referred to Theriosuchus based on several derived characters, and differs from the five previously recognized species within this genus. Despite several diagnostic features, we conservatively refer it to Theriosuchus sp., pending the discovery of more complete material. As the oldest known definitively diagnostic atoposaurid, this discovery indicates that the oldest members of this group were small-bodied, had heterodont dentition, and were most likely widespread components of European faunas. Our review of mandibular and dental features in atoposaurids not only allows us to present a revised diagnosis of Theriosuchus, but also reveals a great amount of variability within this genus, and indicates that there are currently five valid species that can be differentiated by unique combinations of dental characteristics. This variability can be included in future broad-scale cladistics analyses of atoposaurids and closely related crocodyliforms, which promise to help untangle the complicated taxonomy and evolutionary history of Atoposauridae

The locomotion pattern of Baurusuchus salgadoensis Carvalho, Nobre & Campos, 2005 and the distribution of Baurusuchidae in Gondwanaland

Baurusuchus salgadoensis Carvalho, Nobre & Campos, 2005 is a ziphodont mesoeucrocodylian closely related to the Sebecosuchia. Cranial features indicate that this species had terrestrial predatory habits. The postcranial skeleton of B. salgadoensis show distinctive characteristics when compared to extant Crocodylomorpha. The dorsal vertebrae are close articulated with short and craniocaudally expanded neural spines, specially the posterior ones. The sacral vertebrae are robust and have the lateral processes highly fused to the ilium. The anterior caudal vertebrae are robust as the sacral ones but their neural spines are more stout, anteriorlly expanded and higher. The close articulation of the dorsal vertebrae may indicate a less flexible dorsal spine, ideal to limb-driven methods of locomotion. The high and expanded neural spines of the posterior dorsal and sacral vertebrae are associated to muscle attachment from the osteoderms and the pelvic musculature, all used in the limb-driven locomotion. The appendicular bones of B. salgadoensis are long and stout. They show a straight aspect of their diaphysis and very well- developed epiphysis. The femur is long when compared to extant crocodilians, showing a straight aspect in lateral view and a slight sigmoid aspect in frontal view. There is a small axial torsion at its proximal end. The fourth trochanter is pronouced and posteriorlly oriented. The proximal end possess a mesial projection that articulates itself to the illium, similar to those of Protosuchia and thecodont archosaurs as the Rauisuchia. The illium of B. salgadoensis present a lateral and posteriorlly expanded postaccetabular crest, similar to those observed in Rauisuchia and Protosuchia, referred as overhanging ilium. This pattern of articulation is observed in the rauisuchian thecodonts and interpreted as a characteristic trait of those able erect-posture and limb-driven predators of Triassic environments. B. salgadoensis exoskeleton has only two dorsal osteoderm rows that run from the neck to the tip of the tail. Most of the osteoderms are wider than longer, with a round lateral portion that does not articulate to any flank osteoderm. The only morphological difference appears at the caudalmost portion of the tail, where the osteoderms are craniocaudally elongated. The medial portion has little variation along the scutes rows and may bear medial lamellar dorsoventral structures of articulation to the adjacent bony plate. The anterior articular facet is discreet and the osteoderms are not as imbricated as occur in other crocodyliforms and even may not be imbricated at all. The pelvic region osteoderms have the tallest keel of the row that runs from the anterior articular facet to posteriosmost portion of the osteoderm. B. salgadoensis has a light exoskeleton and then is less encumbranced by it. Therefore, becaming more agile. The imbrication of dorsal armor assists the limb-driven locomotion of many crocodyliform by reducing the flexibility of the dorsal spine during the high-walk (erect stance). Therefore the long and stout limb bones, overhanging crest of the illium and the light weighted armor, also allowed to B. salgadoensis to have the limb-driven locomotion without this pattern of osteoderms. These anatomical data may indicate how the Baurusuchidae had a wide distribution in Gondwanaland as they were able to move across large distances

Functional aspects of titanosaur osteoderms

Though titanosaur osteoderms are not common findings, these elements are recorded widely in Gondwana and part of Laurasia. This assembly known by the date offers few resources for studies on the ecology of this group of dinosaurs. Recently, some eggs bearing titanosaur embryos with preserved skin, from Patagonia, Argentina, may shed some light on the function and disposition pattern of these dermic bones. Some of the skin patches associated with the titanosaur embryos show two distinct patterns of tuberosities: a longitudinal row and rosettes, both composed of closely attached tuberosities. These tuberosities do not seem to be ossified, but this might be due to the ontogenetic stage of the titanosaurs. Here we propose that these tuberosities might have been ossified in later ontogenetic stages, and then turning into real osteoderms providing physical defence for the juvenile titanosaurs. Amongst the remains of large titanosaurs like _Mendozasaurus neguyelap_ some osteoderms were recovered, but it is contrasting small to the animal when compared to other dinosaurs. These bones would not provide real defensive advantage to an adult titanosaur as they are small and also have a very spongy internal structure. This apparent fragility also may be the reason that few titanosaur osteoderms have been preserved. In comparison, the tuberosity of a young titanosaur is much larger than an adult osteoderm and its body armor would be much more effective against small predators like notosuchian crocodyliforms and small theropods. And if the titanosaur osteoderms are originated from the embryo’s tuberosities, the disposition of these elements in an adult animal would be very distant to each other. The calcium reserve may be considered as functional for an adult titanosaur though

A New Eusuchian Crocodyliform with Novel Cranial Integument and Its Significance for the Origin and Evolution of Crocodylia

Crocodyliforms were one of the most successful groups of Mesozoic tetrapods, radiating into terrestrial, semiaquatic and marine environments, while occupying numerous trophic niches, including carnivorous, insectivorous, herbivorous, and piscivorous species. Among these taxa were the enigmatic, poorly represented flat-headed crocodyliforms from the late Cretaceous of northern Africa. Here we report a new, giant crocodyliform from the early Late Cretaceous (Cenomanian) Kem Kem Formation of Morocco. Represented by a partial braincase, the taxon has an extremely long, flat skull with large jaw and craniocervical muscles. The skull roof is ridged and ornamented with a broad, rough boss surrounded by significant vascular impressions, likely forming an integumentary structure unique among crocodyliforms. Size estimates using endocranial volume indicate the specimen was very large. The taxon possesses robust laterosphenoids with laterally oriented capitate processes and isolated epipterygoids, features allying it with derived eusuchians. Phylogenetic analysis finds the taxon to be a derived eusuchian and sister taxon to Aegyptosuchus, a poorly understood, early Late Cretaceous taxon from the Bahariya formation. This clade forms the sister clade of crown-group Crocodylia, making these taxa the earliest eusuchian crocodyliforms known from Africa. These results shift phylogenetic and biogeographical hypotheses on the origin of modern crocodylians towards the circum-Tethyean region and provide important new data on eusuchian morphology and evolution

Dinosaur footprints and other Ichnofauna from the Cretaceous Kem Kem Beds of Morocco

We describe an extensive ichnofossil assemblage from the likely Cenomanian-age 'lower' and 'upper' units of the 'Kem Kem beds' in southeastern Morocco. In the lower unit, trace fossils include narrow vertical burrows in cross-bedded sandstones and borings in dinosaur bone, with the latter identified as the insect ichnotaxon Cubiculum ornatus. In the upper unit, several horizons preserve abundant footprints from theropod dinosaurs. Sauropod and ornithischian footprints are much rarer, similar to the record for fossil bone and teeth in the Kem Kem assemblage. The upper unit also preserves a variety of invertebrate traces including Conichnus (the resting trace of a sea-anemone), Scolicia (a gastropod trace), Beaconites (a probable annelid burrow), and subvertical burrows likely created by crabs for residence and detrital feeding on a tidal flat. The ichnofossil assemblage from the Upper Cretaceous Kem Kem beds contributes evidence for a transition from predominantly terrestrial to marine deposition. Body fossil and ichnofossil records together provide a detailed view of faunal diversity and local conditions within a fluvial and deltaic depositional setting on the northwestern coast of Africa toward the end of the Cretaceous

Complex macroevolutionary dynamics underly the evolution of the crocodyliform skull

All modern crocodyliforms (alligators, crocodiles and the gharial) are semi-aquatic generalist carnivores that are relatively similar in cranial form and function. However, this homogeneity represents just a fraction of the variation that once existed in the clade, which includes extinct herbivorous and marine forms with divergent skull structure and function. Here, we use high-dimensional three-dimensional geometric morphometrics to quantify whole-skull morphology across modern and fossil crocodyliforms to untangle the factors that shaped the macroevolutionary history and relatively low phenotypic variation of this clade through time. Evolutionary modelling demonstrates that the pace of crocodyliform cranial evolution is initially high, particularly in the extinct Notosuchia, but slows near the base of Neosuchia, with a late burst of rapid evolution in crown-group crocodiles. Surprisingly, modern crocodiles, especially Australian, southeast Asian, Indo-Pacific species, have high rates of evolution, despite exhibiting low variation. Thus, extant lineages are not in evolutionary stasis but rather have rapidly fluctuated within a limited region of morphospace, resulting in significant convergence. The structures related to jaw closing and bite force production (e.g. pterygoid flange and quadrate) are highly variable, reinforcing the importance of function in driving phenotypic variation. Together, these findings illustrate that the apparent conservativeness of crocodyliform skulls betrays unappreciated complexity in their macroevolutionary dynamics.Fil: Felice, Ryan N.. Colegio Universitario de Londres; Reino Unido. Natural History Museum; Reino UnidoFil: Pol, Diego. Museo Paleontológico Egidio Feruglio; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico; ArgentinaFil: Goswami, Anjali. Natural History Museum; Reino Unid

An enigmatic crocodyliform tooth from the bauxites of western Hungary suggests hidden mesoeucrocodylian diversity in the Early Cretaceous European archipelago

Background. The Cretaceous of southern Europe was characterized by an archipelago setting with faunas of mixed composition of endemic, Laurasian and Gondwanan elements. However, little is known about the relative timing of these faunal influences. The Lower Cretaceous of East-Central Europe holds a great promise for understanding the biogeographic history of Cretaceous European biotas because of the former proximity of the area to Gondwana (as part of the Apulian microcontinent). However, East-Central European vertebrates are typically poorly known from this time period. Here, we report on a ziphodont crocodyliform tooth discovered in the Lower Cretaceous (Albian) Alsopere Bauxite Formation of Olaszfalu, western Hungary. Methods. The morphology of the tooth is described and compared with that of other similar Cretaceous crocodyliforms. Results. Based on the triangular, slightly distally curved, constricted and labiolingually flattened crown, the small, subequal-sized true serrations on the carinae mesially and distally, the longitudinal fluting labially, and the extended shelves along the carinae lingually the tooth is most similar to some peirosaurid, non-baurusuchian sebecosuchian, and uruguaysuchid notosuchians. In addition, the paralligatorid Wannchampsus also possesses similar anterior teeth, thus the Hungarian tooth is referred here to Mesoeucrocodylia indet. Discussion. Supposing a notosuchian affinity, this tooth is the earliest occurrence of the group in Europe and one of the earliest in Laurasia. In case of a paralligatorid relationship the Hungarian tooth would represent their first European record, further expanding their cosmopolitan distribution. In any case, the ziphodont tooth from the Albian bauxite deposit of western Hungary belongs to a group still unknown from the Early Cretaceous European archipelago and therefore implies a hidden diversity of crocodyliforms in the area