A new species of Baurusuchus (Crocodyliformes, Mesoeucrocodylia) from the Upper Cretaceous of Brazil, with the first complete postcranial skeleton described for the family Baurusuchidae

The present work describes a new species of Baurusuchidae from Upper Cretaceous sediments of the Bauru Basin, and provides the first complete postcranial description for the family. Many postcranial features observed in the new species are also present in other notosuchian taxa, and are thus considered plesiomorphic for the genus. These are: long cervical neural spines; robust deltopectoral crest of the humerus; large proximal portion in the radiale that contacts the ulna; ulnare anterior distal projection; supra-acetabular crest well developed laterally; post-acetabular process posterodorsally deflected; presence of an anteromedial crest in the femur; fourth trocanter of femur posteriorly positioned; tibia with a laterally curved shaft; calcaneum tuber posteroventrally oriented; osteoderms ornamented with grooves and imbricated in the tail. On the other hand, we found the following sacral and carpal features to be unique among all mesoeucrocodylians analyzed: transverse processes of sacral vertebrae dorsolaterally deflected; presence of a longitudinal crest in the lateral surface of sacral vertebrae; pisiform carpal with a condyle-like surface. The majority of these cited features corroborates a cursorial locomotion for the new species described in the present study, suggesting that members of the family Baurusuchidae were also cursorial species.

A New Baurusuchid (Crocodyliformes, Mesoeucrocodylia) from the Late Cretaceous of Brazil and the Phylogeny of Baurusuchidae

Background: Baurusuchidae is a group of extinct Crocodyliformes with peculiar, dog-faced skulls, hypertrophied canines, and terrestrial, cursorial limb morphologies. Their importance for crocodyliform evolution and biogeography is widely recognized, and many new taxa have been recently described. In most phylogenetic analyses of Mesoeucrocodylia, the entire clade is represented only by Baurusuchus pachecoi, and no work has attempted to study the internal relationships of the group or diagnose the clade and its members. Methodology/Principal Findings: Based on a nearly complete skull and a referred partial skull and lower jaw, we describe a new baurusuchid from the Vale do Rio do Peixe Formation (Bauru Group), Late Cretaceous of Brazil. The taxon is diagnosed by a suite of characters that include: four maxillary teeth, supratemporal fenestra with equally developed medial and anterior rims, four laterally visible quadrate fenestrae, lateral Eustachian foramina larger than medial Eustachian foramen, deep depression on the dorsal surface of pterygoid wing. The new taxon was compared to all other baurusuchids and their internal relationships were examined based on the maximum parsimony analysis of a discrete morphological data matrix. Conclusion: The monophyly of Baurusuchidae is supported by a large number of unique characters implying an equally large morphological gap between the clade and its immediate outgroups. A complex phylogeny of baurusuchids was recovered. The internal branch pattern suggests two main lineages, one with a relatively broad geographical range between Argentina and Brazil (Pissarrachampsinae), which includes the new taxon, and an endemic clade of the Bauru Group in Brazil (Baurusuchinae).

Dois novos crocodilos (Crocodyliformes, Mesoeucrocodylia) do Mato Grosso, Bacia dos Parecis : descrição e relações filogenéticas com os "Notossúquios"

São descritos dois novos táxons de Crocodyliformes do Cretáceo do Mato Grosso, que apresentam: crânios curtos; dentição heterodonte composta por dentes com lado labial mais convexo que o lado lingual e serrilhados; dentes anteriores do dentário procumbentes; presença de uma superfície distintamente lisa, acima da margem alveolar, na superfície lateral da maxila, acima da qual ocorrem forames neurovasculares; e vértebras anficélicas. Estes novos materiais foram informalmente chamados Crocodilo I e Crocodilo II. O Crocodilo I apresenta as seguintes características: fenestra pré-orbitária pequena; barra palatina posterolateral em forma de bastão; dente caniniforme hipertrofiado na pré-maxila; 6 dentes maxilares; dentes anteriores do dentário levemente voltados para trás; junta quadrado-articular localizada ventralmente à fileira dentária. O Crocodilo II, por sua vez, apresenta: focinho tubular em seção; amplo palato secundário; espleniais fusionados; 5 dentes maxilares; mandíbula com região sinfiseal alongada e espatulada. Em ambos os casos, a morfologia geral aponta muitas semelhanças com os “notossúquios”: postura elevada, regiões diferenciadas na coluna, dentição heterodonte, palato secundário relativamente amplo e cavidade nasal relativamente ampla. Para testar as relações filogenéticas dos dois táxons, bem como o próprio monofiletismo dos “notossúquios”, foi construída uma matriz de dados com 27 táxons (incluindo, tanto quanto possível, todos aqueles que já foram ou são considerados “notossúquios”) e 60 caracteres. Com base nos mesmos, foi efetuada uma análise de parcimônia utilizando Paup 3.1.1 (“stepwise addition” - “random”, com 100 repetições e “tree-bisection-reconnection”) e Nona 2.0 (algoritmo heurístico, com 10 replicações e TBR - mult*max*) que apontou uma relação dos dois novos táxons como grupos irmão sucessivos de Notosuchus e Mariliasuchus (Crocodilo II (Crocodilo I (Notosuchus, Mariliasuchus))). Em comparação com outros crocodiliformes do Cretáceo da América do Sul, África, Madagascar e China, é possível assumir que ambos são notossúquios (sensu Pol & Apesteguia, 2005). Em nossa análise, os notossúquios formam um grupo, à exceção de A. buitreraensis, mais proximamente relacionado aos Eusuchia. Na análise ordenada, A. buitreraensis, A. patagonicus e Anatosuchus aparecem mais proximamente relacionados a Eusuchia e Lybicosuchus aloca-se junto a Baurusuchus e Sebecus. Paralelamente, são discutidas análises filogenéticas prévias envolvendo notossúquios, nas quais foram introduzidas modificações, levando a diferentes topologias. A partir disso, propõe-se uma discussão acerca da influência das escolhas e procedimentos de cada autor no resultado final de análises filogenéticas.

A new Sphagesaurus (Mesoeucrocodylia: Notosuchia) from the Upper Cretaceous of Monte Alto City (Bauru Group, Brazil), and a revision of the Sphagesauridae

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Morphological and anatomical observations about Mariliasuchus amarali and Notosuchus terrestris (Mesoeucrocodylia) and their relationships with other South American notosuchians.

The phylogenetic relationship of the notosuchians Mariliasuchus amarali (Campanian; Bauru Group) and Notosuchus terrestris (Santonian; Neuquen Group) is revised. Morpho-anatomical evaluation of Mariliasuchus in the current bibliography indicate close relationship with Notosuchus, while cladistic analysis either related Mariliasuchus to Candidodon itapecuruense (Albian/eo-Cenomanian; Sao Luis-Grajau Basin), as part of the phylotaxon Candidodontidae, or to Comahuesuchus brachybuccalis (Santonian; Neuquen Group). Comparative study of specimens shows similarities on the palate, choanae, dentition, retroarticular process, and other structures from Mariliasuchus and Notosuchus supporting the original classification as a Notosuchidae. Preliminary phylogenetic analysis sets these taxa as sister-groups. Reevaluation of a previously published phylogenetic analysis from other authors provides further support for the Mariliasuchus + Notosuchus clade. The current work indicates that Mariliasuchus is a Notosuchidae, refuting its allocation as a Candidodontidae. The influence of character construction and the definition of Notosuchia are discussed.

Morphology of the dental carinae in Mariliasuchus amarali (Crocodylomorpha, Notosuchia) and the pattern of tooth serration among basal Mesoeucrocodylia.

Carinated teeth are common in Mesoeucrocodylia, and the occurrence of denticles over the carinae is related to high predacious species, often referred as ziphodont. This characteristic is broadly recognized as homoplastic. Carinae morphology is cryptic, difficult to be studied under common techniques, and Scanning Electronic Microscopy (SEM) allows the access to detailed information, offering a higher degree of confidence. Previous SEM study allowed the recognition of true/false ziphodont patterns, according to the morphology of the denticles, but such studies on gondwanan mesoeucrocodyles are uncommon. Mariliasuchus amarali is an Upper Cretaceous notosuchian mesoeucrocodyle from South America (Bauru Group, Brazil), with carinated teeth and specialized dentition. Its geological and biochronological distribution are reappraised. SEM study of two teeth shows carinae composed of isolated tuberous anisomorphic true denticles, supporting previous study. Enamel ornamentation does not develop over the carinae, and fabric becomes anastomosed in middle and posterior teeth. Carinae only occur in posterior molariform teeth, related to food processing. Morphological variability of Mariliasuchus is commented, focusing on dentition. Overall characteristics, molariform morphology and wear planes support a non-predacious habit for Matiliasuchus. Matiliasuchus pattern could not be related to true/false ziphodont patterns, either by morphology or function, and is defined as ziphomorph. Ziphomorph pattern is evaluated within the range of mesoeucrocodyles. The detailed study of homoplastic characteristics, such as dental carinae, may provide useful apomorphic information for cladistic analysis.

Reconhecimento geológico (Formações Santo Anastácio e Adamantna) e paleobiológico (Mesoeucrocodylia Baurusuchidae e Sphagesauridae) na região do estado de São Paulo

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Analise cladística - filogenética e paleobiogeográfica dos Mesoeucrocodylia (Crocodylomorpha; Crocodyliformes) do Grupo Bauru (Cretáceo Superior) do Sudeste do Brasil

Pós-graduação em Geociências e Meio Ambiente - IGCE

Anatomy, phylogeny, and palaeoecology of the basal thalattosuchians (Mesoeucrocodylia) from the Liassic of Central Europe

In the marine Jurassic deposits of Europe, a group of marine crocodilians, the Thalattosuchia, belongs to the frequently found reptiles. Thalattosuchia are widely spread in Central Europe from the Jurassic to Lower Cretaceous, and some taxa are also distributed worldwide. The task of the work was to examine all taxa known from the Liassic of Europe. The most frequently known taxa Steneosaurus bollensis and Pelagosaurus typus are anatomically revised. New discoveries at the skull of Pelagosaurus typus e.g., the fact of a partly paired frontal are described by means of computed tomography investigations. In addition, juvenile specimens of this taxon are studied in detail for the first time. The rarely occurring taxon Platysuchus multiscrobiculatus is anatomically described in detail for the first time. It shows both in the skull and in the postcranial material morphological differences to Steneosaurus bollensis and Pelagosaurus typus. Thus Pl. multiscrobiculatus possesses, e.g., an ilium with a deeper acetabulum and a femur with a distinctly flexed femoral head. A juvenile specimen of Pl. multiscrobiculatus is now discovered and is described in parts for the first time, too. Furthermore, Steneosaurus gracilirostris and Steneosaurus brevior known from Lower Jurassic deposits of England are examined and in parts revised. In this work, Steneosaurs brevior is discovered with one specimen from the Upper Liassic of Holzmaden, Germany for the first time, and provides new evidence for the palaeobiogeographical distribution of the taxon. Because of the high number of investigated specimens, it is possible to study ontogenetic development from juvenile to adult stage in Steneosaurus bollensis, Pelagosaurus typus, and Platysuchus multiscrobiculatus. Biometric data are collected from thalattosuchians and extant crocodilians (e.g. Gavialis gangeticus) to investigate intraspecific variation, ontogenetic development, and taxa differentiation. The skulls of Platysuchus multiscrobiculatus and Steneosaurus bollensis are reconstructed three-dimensionally as wax models. The skull reconstructions form the basis of the jaw muscle restoration of Steneosaurus bollensis in connection with comparative studies at extant crocodilians. By means of functional morphologic analysis of the jaw musculature, the dentition, and the locomotor system of S. bollensis, possible conclusions are drawn for the prey options and the hunting behaviour. To clarify the relationships within the Thalattosuchia, a computer-based cladistic phylogenetic in-group analyse of 25 Thalattosuchia taxa is performed. For the analysis, following Thalattosuchia taxa are studied likewise at original material for comparisons: Metriorhynchus superciliosus, Metriorhynchus hastifer, Metriorhynchus leedsi, Geosaurus gracilis, Geosaurus giganteus, Teleidosaurus calvadosi, Teleidosaurus gaudryi, Teleosaurus cadomensis, Teleosaurus geoffroyi, Steneosaurus priscus, Steneosaurus edwardsi, Steneosaurus heberti, Steneosaurus leedsi, Steneosaurus boutilieri, Steneosaurus megarhinus, Steneosaurus obtusidens, and Machimosaurus hugii. The phylogenetic in-group analyse based on 115 characters, reveals a sister-group relationship of the monophyletic Teleosauridae and monophyletic Metriorhynchidae. Within the groups, some taxa are probably paraphyletic. The taxon Pelagosaurus typus is nested inside the Teleosauridae and not outside or within the Metriorhynchidae, as many authors suggested it so far. Based on these results, a tentative palaeobiogeographical-evolutionary scenario is developed.

Multicusped crocodyliform teeth from the Upper Cretaceous (Sao Jose do Rio Preto Formation, Bauru Group) of Sao Paulo, Brazil

The six peculiar multicusped teeth described here were collected from sediments of the Upper Cretaceous of Sao Jose do Rio Preto Formation, near Ibira (northeastern Sao Paulo, Brazil). Their bulbous crowns are slightly labio-lingual compressed, and bear a main plus two accessory cusps, which conceal a well developed cingulum. Wear facets are seen on the main and distal accessory cusps. Comparison to the known Crocodyliformes with multicusped teeth show that the new material is not referable to ""protosuchians"" or eusuchians, nor related to two unnamed forms from Morocco and ""notosuchians"" such as Uruguaysuchus, Chiamaerasuchus, and Simosuchus. On the other hand, possible affinities with Candidodon and Malawisuchus were maintained based on shared traits. This includes teeth with the main cusp and some accessory cusps arranged in more than one axis, a previously defined unambiguous apomorphy of the putative clade composed of Candidodon plus Malawisuchus. The term Candidodontidae can be applied to this group, and defined as all taxa closer to Candidodon itapecuruensis than to Notosuchus terrestris, Uruguaysuchus aznarezi, Comahuesuchus brachybuccalis, Sphagesaurus huenei, Baurusuchus pachecoi, and Crocodylus niloticus. (C) 2009 Elsevier Ltd. All rights reserved.

An Additional Baurusuchid from the Cretaceous of Brazil with Evidence of Interspecific Predation among Crocodyliformes

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

A species-level supertree of Crocodyliformes

With fossils found worldwide, Crocodyliformes stands as one of the best documented vertebrates over the Mesozoic and Cenozoic. The multiple phylogenetic hypotheses of relationship proposed for the group allow plenty of space for contentious results, partially due to the small overlapping of taxa and disagreeing homology statements among studies. We present two supertrees of Crocodyliformes, based on different protocols of source tree selection, summarising phylogenetic data for the group into a 'synthetic consensus'. The consensus of the most parsimonious trees, containing 184 terminal taxa, has a remarkably well-resolved branching structure, which may serve as a framework for further macroevolutionary studies. In addition, the IterPCR script was for the first time used in the supertree context to build a reduced consensus tree with the pruning of unstable taxa.