A new fossil frog (Lissamphibia: Anura) from the Late Cretaceous of Brazil and the early evolution of neobatrachians

SummaryBlind snakes (Scolecophidia) are minute cryptic snakes that diverged at the base of the evolutionary radiation of modern snakes. They have a scant fossil record, which dates back to the Upper Paleocene-Lower Eocene (∼56 Ma); this late appearance conflicts with molecular evidence, which suggests a much older origin for the group (during the Mesozoic: 160–125 Ma). Here we report a typhlopoid blind snake from the Late Cretaceous of Brazil, Boipeba tayasuensis gen. et sp. nov, which extends the scolecophidian fossil record into the Mesozoic and reduces the fossil gap predicted by molecular data. The new species is estimated to have been over 1 m long, much larger than typical modern scolecophidians (<30 cm). This finding sheds light on the early evolution of blind snakes, supports the hypothesis of a Gondwanan origin for the Typhlopoidea, and indicates that early scolecophidians had large body size, and only later underwent miniaturization.

A new notosuchian crocodyliform from the Late Cretaceous Bauru Group found in the southeastern State of São Paulo (Brazil) is described here. The new taxon, Caipirasuchus stenognathus, is referred as a new species of the recently erected genus Caipirasuchus within the clade Sphagesauridae based on a phylogenetic analysis of basal mesoeucrocodylians. Caipirasuchus stenognathus is represented by an almost complete skull and lower jaw that has autapomorphic characters that distinguish it from other species of Sphagesauridae. These autapomorphies include: maxilla forming part of the orbital margin (absence of lacrimal-jugal contact), nasal with smooth depressions on the posterior region close to the contact with the maxilla and lacrimal, postorbital with posterior palpebral facet that extends posteriorly underneath the ear-flap groove, and a distinct anterior process of the medial flange of the retroarticular process. Additionally, the new taxon lacks autapomorphic features described in other sphagesaurids. The phylogenetic analysis results in a monophyletic genus Caipirasuchus, that is the sister group of a clade fomed by Sphagesaurus huenei, Caryonosuchus pricei, and Armadillosuchus arrudai. Sphagesaurids also include a basal clade formed by Adamantinasuchus navae and Yacarerani boliviensis. Other notosuchian taxa, such as Mariliasuchus amarali, Labidiosuchus amicum, Notosuchus terrestris, and Morrinhosuchus luziae are successive sister taxa of Sphagesauridae, forming a clade of advanced notosuchians that are restricted to the Late Cretaceous of South America. These results contrast with most previous phylogenetic hypotheses of the group that depicted some members of Sphagesauridae as more closely related to baurusuchids, or found Asian (e.g., Chimaerasuchus) or African (Malawisuchus, Pakasuchus) forms nested within advanced notosuchians that are, according to our analysis, endemic of the Late Cretaceous of South America.

Sphagesauridae is a group of notosuchian crocodyliforms from the Late Cretaceous of South America characterized by highly specialized jaws and dentition. Here, we describe a new sphagesaurid from the Santo Anastácio Formation (Caiuá Group, Bauru Basin), south-east Brazil. The specimen is composed of a partial palate, neurocranium, mandible and fragmentary teeth. It represents a new species that can be assigned to Caipirasuchus due to the presence of a lateromedially narrow and anteroposteriorly long mandibular symphyseal region, apicobasal ridges on the posterior teeth, a diastema between D5 and D6, and a linear row of large neurovascular foramina on the lateral surface of the dentary. It differs from previously described Caipirasuchus species based on a ventrolaterally inclined surface of the dentaries posterior to the tooth row, a connection between the anteroventral margin of the external mandibular fenestra and the floor of the Meckelian canal, and the anterior process of the angular forming a ‘V’-shaped suture in its contact with the splenial. The results of a phylogenetic analysis of notosuchians recovered the so-called ‘advanced notosuchians’ in a clade, for which we propose a phylogenetic definition and erect the name Sphagesauria. We also recovered two more inclusive clades encompassing an array of Mesozoic notosuchians, here named Xenodontosuchia, which includes Sphagesauria + Baurusuchia, and Eunotosuchia, including Xenodontosuchia, Uruguaysuchidae and other Cretaceous forms. http://zoobank.org/urn:lsid:zoobank.org:pub:FD8D706F-9E01-4C09-BE4E-DC3BEDE151DC

Dryolestoids are an extinct mammalian group belonging to the lineage leading to modern marsupials and placentals. Dryolestoids are known by teeth and jaws from the Jurassic period of North America and Europe, but they thrived in South America up to the end of the Mesozoic era and survived to the beginnings of the Cenozoic. Isolated teeth and jaws from the latest Cretaceous of South America provide mounting evidence that, at least in western Gondwana, dryolestoids developed into strongly endemic groups by the Late Cretaceous. However, the lack of pre-Late Cretaceous dryolestoid remains made study of their origin and early diversification intractable. Here we describe the first mammalian remains from the early Late Cretaceous of South America, including two partial skulls and jaws of a derived dryolestoid showing dental and cranial features unknown among any other group of Mesozoic mammals, such as single-rooted molars preceded by double-rooted premolars, combined with a very long muzzle, exceedingly long canines and evidence of highly specialized masticatory musculature. On one hand, the new mammal shares derived features of dryolestoids with forms from the Jurassic of Laurasia, whereas on the other hand, it is very specialized and highlights the endemic, diverse dryolestoid fauna from the Cretaceous of South America. Our specimens include only the second mammalian skull known for the Cretaceous of Gondwana, bridging a previous 60-million-year gap in the fossil record, and document the whole cranial morphology of a dryolestoid, revealing an unsuspected morphological and ecological diversity for non-tribosphenic mammals.

O registro de titanossauros no Cretaceo do Brasil, associado ao genero patagonico Aeolosaurus Powell (Dinosauria, Sauropoda, Titanosauria), e discutido. A natureza fragmentaria e isolada de muitos dos materiais estudados e o conhecimento ainda incipiente da real diversidade e das relacoes filogeneticas dos titanossauros brasileiros tem gerado incertezas nas afinidades taxonomicas propostas. A revisao dos especimes brasileiros atribuidos a Aeolosaurus sugere que nao ha caracteres para sustentar a ocorrencia deste genero patagonico no Cretaceo do Brasil. A constatacao de que estes materiais isolados e fragmentarios foram inadequadamente reconhecidos em suas relacoes alerta para a necessidade de procedimentos taxonomicos mais adequados. Para tanto, identificacoes bem fundamentadas, por exemplo, na presenca de apomorfias, e indispensavel para a correta designacao taxonomica, e as consequentes inferencias paleobiogeograficas, paleobiostratigraficas e no estabelecimento de biocronos.Palavras-chave: Titanosauria, Aeolosaurini, Aeolosaurus, Cretaceo Superior, Grupo Bauru, Brasil.

A new Late Cretaceous crocodyliform from the western margin of Gondwana (La Rioja Province, Argentina)

The first bony theropod record from the Campanian Uberaba Formation (Bauru Group) is described. It consists of an isolated caudal centrum (CPPLIP 1324) found in the city of Uberaba, Minas Gerais State, Brazil. The amphicoelous centrum possesses a length to height ratio of 1.74, deep elliptical lateral pneumatic foramen representing 26% of centrum length with three main sub-circular air chambers, andcamellate internal structure.This combination of features is shared with Aerosteon, Megaraptor, and Orkoraptor from the Late Cretaceous of Argentinaand with the Megaraptora indet. fromthe São José do Rio Preto Formation (Bauru Group), São Paulo State,allowing us to refer it to the Megaraptora clade (Tetanurae, Neovenatoridae). As such, the new specimen represents the second megaraptoran from the Late Cretaceous of Brazil and provides new information on tail anatomy on this bizarre group.

First record of Siluriformes from the northernmost portion of the Bauru Group (Upper Cretaceous) in the Center-West region of Brazil

Titanosaurs were successful herbivorous dinosaurs widely distributed in all continents during the Cretaceous, with the major diversity in South America. The success of titanosaurs was probably due to several physiological and ecological factors, in addition to a series of morphological traits they achieved during their evolutionary history. However, the generalist nesting behaviour using different palaeoenvironments and strategies was key to accomplish that success. Titanosaur nesting sites have been found extensively around the world, with notable records in Spain, France, Romania, India, and, especially, Argentina. Here, we describe the first titanosaur nesting site from the Late Cretaceous of Brazil that represents the most boreal nesting site for South America. Several egg-clutches, partially preserved, isolated eggs and many eggshell fragments were discovered in an Inceptisol palaeosol profile of the mining Lafarge Quarry, at the Ponte Alta District (Uberaba Municipality, Minas Gerais State), corresponding to the Serra da Galga Formation (Bauru Group, Bauru Basin). Although classical mechanical preparation and CT scans have not revealed embryonic remains in ovo, the eggs and eggshell features match those eggs containing titanosaurian embryos found worldwide. The morphology of the egg-clutches and observations of the sedimentary characteristics bolster the hypothesis that these sauropods were burrow-nester dinosaurs, as was already suggested for the group based on other nesting sites. The egg-clutches distributed in two levels along the Lafarge outcrops, together with the geopalaeontological data collected, provide clear evidence for the first colonial nesting and breeding area of titanosaur dinosaurs in Brazil.

The Crocodyliformes from the Late Cretaceous of South America were one of the main components of the tetrapod faunas found in rocks of the Bauru Group at Goiás, Minas Gerais and São Paulo states, Brazil. The presence of osteoderms has been reported in articulated specimens or found isolated in the palaeontological record. In the present work, we describe the first osteoderm of Mesoeucrocodylia from the Campanian-Maastrichtian of the Marília Formation (Bauru Group) of the state of Goiás. Paleo-UFG-114 has a quadrilateral shape, a rough external texture with radially oriented corrosion, an internal facet with the presence of intersecting grooves, foramina in the internal and external region and the presence of a poorly developed keel. These features differentiate them from the osteoderms of titanosaur dinosaurs found in the rocks of this geological unit. The presence of Paleo-UFG-114 in the state of Goiás reaffirms that mesoeucrocodylians were important components of the reptile fauna, and contributes to increasing knowledge about the group’s geographic distribution in this poorly-explored region of Central Brazil

With nearly 30 living species of relatively similar ecological traits, Crocodylomorpha is represented today by only a small fraction of its past diversity. The well-documented crocodylomorph fossil record has revealed more than 500 taxa, with much higher ecological and morphological diversity than their extant counterparts. An example of such astonishing diversity is the Late Cretaceous rocks of the Bauru Group (southeast Brazil), from which numerous taxa are known, belonging to the clade Notosuchia. These were predominantly terrestrial taxa, some of which exhibited traits associated with omnivorous or even herbivorous feeding behaviors, such as Sphagesauridae, whereas others were adapted to a carnivore diet, such as Baurusuchidae and Peirosauridae. Among these is Barreirosuchus franciscoi, originally described as a neosuchian (Trematochampsidae) but later interpreted as a peirosaurid notosuchian. Even though included in recent morphological and phylogenetic analyses, B. franciscoi still lacked a more detailed description. Here, we provide an in-depth description of the cranial elements of B. franciscoi, using data from computed tomography and a broad sample of comparative material, including living and fossil crocodylomorphs. Also, the neuro-cavities, including the endocast, nasopharyngeal duct, and the olfactory region, were digitally reconstructed. Finally, a new phylogenetic analysis recovered B. franciscoi nested within Peirosauria, forming the Itasuchidae clade with other potentially semiaquatic species: Rukwasuchus yajabalajekundu, Pepesuchus deiseae, and Itasuchus jesuinoi. The morphological and phylogenetic reassessment of B. franciscoi indicates a semiaquatic form, highlighting the ecological diversity of notosuchians from the Bauru Group as well as the capacity of notosuchians to explore a myriad of environments.

A new Pissarrachampsinae specimen from the Bauru Basin, Brazil, adds data to the understanding of the Baurusuchidae (Mesoeucrocodylia, Notosuchia) distribution in the Late Cretaceous of South America

Caipirasuchus is a genus of sphagesaurid notosuchian from the Bauru Basin (Upper Cretaceous), southeast Brazil. They were small-sized herbivorous/omnivorous animals, that measured around 1 metre in length, and had a wide diversity and geographical dispersion in the basin. Here, we diagnosed the 6th species of the genus, originating from sandstones of the Adamantina Formation in the municipality of Catanduva, São Paulo State. Caipirasuchus presents a significant variation in the morphology of structures in the choana region. In particular, the new species has chambers in the wings of the pterygoids that connect with an extension of the airways, suggesting a region of resonance used in vocalisation. The different palatal structures in Caipirasuchus may be related to distinct bioacoustic signatures and indicate apossible improvement in the social organisation of Caipirasuchus.

The records of Cretaceous Brazilian titanosaurs assigned to the Patagonian genus Aeolosaurus Powell (Dinosauria, Sauropoda, Titanosauria) are discussed. The fragmentary and isolated nature of many of the specimens and the incomplete knowledge of the diversity and phylogenetic relationships of Brazilian titanosaurs generate uncertainty regarding the taxonomic assignments. Revision of the Brazilian specimens attributed to Aeolosaurus suggests that there are no characters indicating the presence of this Patagonian genus in the Late Cretaceous of Brazil. The conclusion that the relationships of these specimens were incorrectly recognized, points out that better taxonomic procedures, using consistent data, such as the presence of well-defined apomorphies, are needed for correct taxonomic designations, paleogeographic and paleobiostratigraphic inferences and the proposal of biochrons.Key words: Titanosauria, Aeolosaurini, Aeolosaurus, Late Cretaceous, Bauru Group, Brazil.

The application of sequence stratigraphy concepts to continental deposits lacking the referece provided sea level has been a challenge, mainly because the temporal relationships between stratigraphic surfaces and systems tracts depend on the tectonic and climatic evolution of the area. Using the concept of accommodation space (A) and sediment supply (S), we identify specific stacking patterns of aeolian, lacustrine, fluvial and alluvial systems that correspond to the particular tectonic and climatic evolution of the southeastern portion of South America. With the end of the Early Cretaceous volcanism (133 Ma), the southeastern portion of South America underwent tectonic restructuring, which generated basins that encompassed continental sedimentary sequences. The tectonic events responsible for the accumulation of these sequences occurred during two primary phases. The first phase is related to Early Cretaceous thermal subsidence, which was more pronounced in the regions where the thickest Serra Geral Formation basaltic successions are found, resulting in the formation of Bauru Basin. The second phase was related to the Late Cretaceous uplift in southeastern Brazil as a result of magmatic/volcanic activity associated with the Trindade Mantle Plume. Stratigraphic analysis based on well‐logs and outcrops and aided by petrographic studies identified three sequences that are bounded by regional unconformities that record important changes in the Bauru Basin's tectonic and paleoenvironmental conditions. The unconformity K‐0 is related to the origin of the Bauru Basin in the Early Cretaceous. The Early Cretaceous Sequence 1 (Caiuá Group) is interpreted as a second‐ order sequence, formed by aeolian and fluvial deposits and constituting a Fluvial‐Aeolian Systems Tract. Unconformity K‐1 that was generated in the Late Cretaceous (Cenomanian – Campanian?) is related to the tectonic evolution of the basin and source area. Overlying Unconformity K‐1, lacustrine, fluvial and alluvial deposits display progradational characteristics of the two‐third‐ order sequences: Sequences 2A and 2B, constituted by the Fluvial‐Lacustrine and Alluvial Systems Tracts, respectively, and separated by the Unconformity K‐1A. Sedimentological characteristics, paleosols and stratigraphic architecture, suggest that A/S ratio was neutral in the late stage of the Sequence 1, whereas in the Sequence 2 there was an increase (Sequence 2A) followed by a decrease in the A/S ratio (Sequence 2B). Aeolian facies and paleosol P1 (Sequence 1), fluvial‐lacustrine facies and hydromorphic soils (Sequence 2A), and alluvial facies and Paleosol P2 (Sequence 2B), indicate climatic changes in the South American during the Cretaceous. The stratigraphic framework, subaerial unconformities and paleosols provide key elements for subdividing of the Brazilian continental sequence into third‐order sequences and systems tracts, for identification of allocyclic and autocyclic patterns in time and space.