New reptile shows dinosaurs and pterosaurs evolved among diverse precursors.

Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1-3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6-10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the 'success' of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.

Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria.

Pterosaurs were the first vertebrates to evolve powered flight1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252-66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century2-4. These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives2-8, and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids-a group of cursorial, non-volant dinosaur precursors-are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution.

Fossil evidence of the avian vocal organ from the Mesozoic.

From complex songs to simple honks, birds produce sounds using a unique vocal organ called the syrinx. Located close to the heart at the tracheobronchial junction, vocal folds or membranes attached to modified mineralized rings vibrate to produce sound. Syringeal components were not thought to commonly enter the fossil record, and the few reported fossilized parts of the syrinx are geologically young (from the Pleistocene and Holocene (approximately 2.5 million years ago to the present)). The only known older syrinx is an Eocene specimen that was not described or illustrated. Data on the relationship between soft tissue structures and syringeal three-dimensional geometry are also exceptionally limited. Here we describe the first remains, to our knowledge, of a fossil syrinx from the Mesozoic Era, which are preserved in three dimensions in a specimen from the Late Cretaceous (approximately 66 to 69 million years ago) of Antarctica. With both cranial and postcranial remains, the new Vegavis iaai specimen is the most complete to be recovered from a part of the radiation of living birds (Aves). Enhanced-contrast X-ray computed tomography (CT) of syrinx structure in twelve extant non-passerine birds, as well as CT imaging of the Vegavis and Eocene syrinxes, informs both the reconstruction of ancestral states in birds and properties of the vocal organ in the extinct species. Fused rings in Vegavis form a well-mineralized pessulus, a derived neognath bird feature, proposed to anchor enlarged vocal folds or labia. Left-right bronchial asymmetry, as seen in Vegavis, is only known in extant birds with two sets of vocal fold sound sources. The new data show the fossilization potential of the avian vocal organ and beg the question why these remains have not been found in other dinosaurs. The lack of other Mesozoic tracheobronchial remains, and the poorly mineralized condition in archosaurian taxa without a syrinx, may indicate that a complex syrinx was a late arising feature in the evolution of birds, well after the origin of flight and respiratory innovations.

An enigmatic plant-eating theropod from the Late Jurassic period of Chile.

Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

Fossil fishes and anurans from the Miocene of Rio Chico and Cerro Zeballos, Chubut Province, Argentina.

The fossil record of freshwater fishes and anurans from the Miocene in Patagonia is relatively patchy, a large number of specimens remaining undescribed. The aim of the present contribution is to describe a fossil association of percomorphacean fishes and calyptocephalellid anurans from the early to late Miocene Collón Curá Formation, at Chubut province, Patagonia, Argentina. In spite of being represented by several specimens, both anurans and fishes show a very low taxonomic diversity. This pattern matches with other fossil sites from the Cenozoic of Patagonia, as well as with the extant Patagonian batrachofaunas and ichthyofaunas. The fossil record of frogs and fishes in Patagonia is represented by few lineages that have a large evolutionary history in the area, and occasionally can be traced up to the Late Mesozoic.

A rostral neurovascular system in the mosasaur Taniwhasaurus antarcticus.

Mosasaurs were a cosmopolitan group of marine squamate reptiles that lived during the Late Cretaceous period. Tylosaurinae mosasaurs were characterized for having an edentulous rostrum anterior to the premaxillary teeth. External morphology of the snout of the tylosaurine Taniwhasaurus antarcticus from the Upper Cretaceous beds at James Ross Island (Antarctic Peninsula) shows a complex anatomy with diverse large foramina and bone sculpture. A computed tomography scan of the Taniwhasaurus rostrum revealed a complex internal neurovascular system of branched channels in the anteriormost part of the snout. Systems like this are present in extant aquatic vertebrates such as cetaceans and crocodiles to aid them with prey detection, and are inferred to have functioned in a similar manner for several extinct reptile clades such as plesiosaurs and ichthyosaurs. Thus, it is probable that Taniwhasaurus also was able to detect prey with an enhanced neural system located in its rostrum. This condition may be more widespread than previously thought among mosasaurs and other marine reptiles.

New theropod dinosaur from the Upper Cretaceous of Patagonia sheds light on the paravian radiation in Gondwana.

The fossil record of basal paravians in Gondwana is still poorly known, being limited to the Cretaceous unenlagiids from South America and the problematic Rahonavis from Madagascar. Here we report on a new paravian from the Cenomanian-Turonian (Late Cretaceous) of Río Negro province, NW Patagonia, Argentina. The new taxon exhibits a derived bird-like morphology of the forelimbs (e.g., robust ulna with prominent, anteriorly oriented, and proximally saddle-shaped radial cotyle and wide medial flange on metacarpal I) and a plesiomorphic foot with a raptorial pedal digit II. Phylogenetic analysis recovers the new taxon in a monophyletic clade with Rahonavis, being the sister group of the remaining Avialae and more derived than other non-avian dinosaurs. Both exhibit derived forelimb traits in opposition with their plesiomorphic hind limbs. The position of the new taxon and Rahonavis as stem avialans indicates that Gondwanan basal paravians are represented by two different clades, at least. The new taxon probably constitutes a previously unknown grade in the avian-line theropods in which some flight-related adaptations of the forelimbs are present in cursorial taxa. The present discovery sheds light on the acquisition of flight-related traits in non-avian dinosaurs and on the still poorly known paravian radiation in Gondwana.

New record of a Mesozoic gondwanatherian mammaliaform from Southern Patagonia.

Gondwanatheria is an enigmatic mammaliaform clade distributed in the Cretaceous and Paleogene of South America, Africa, Madagascar, India and Antarctica. The Mesozoic record in South America is restricted to the Latest Cretaceous of Río Negro and Chubut provinces, Argentina and Magallanes Region of southern Chile. The aim of the present contribution is to describe a new specimen of gondwanatherian mammaliaforms from beds belonging to the Maastrichtian Chorrillo Formation, cropping out 30 km SW of El Calafate, Santa Cruz Province, Argentina. It is represented by a single molariform referable to the species Magallanodon baikashkenke with which it shares a unique combination of characters. Analysis of the unique combination of characters exhibited by Magallanodon shed doubts on the monophyly of Ferugliotheriidae and suggest that South American taxa may be closely related to each other. The wide geographical distribution and occurrence of gondwanatherians on geological units of diverse origins suggest that they were capable of facing disparate environmental conditions.

Vegaviidae, a new clade of southern diving birds that survived the K/T boundary.

The fossil record of Late Cretaceous-Paleogene modern birds in the Southern Hemisphere includes the Maastrichtian Neogaeornis wetzeli from Chile, Polarornis gregorii and Vegavis iaai from Antarctica, and Australornis lovei from the Paleogene of New Zealand. The recent finding of a new and nearly complete Vegavis skeleton constitutes the most informative source for anatomical comparisons among Australornis, Polarornis, and Vegavis. The present contribution includes, for the first time, Vegavis, Polarornis, and Australornis in a comprehensive phylogenetic analysis. This analysis resulted in the recognition of these taxa as a clade of basal Anseriformes that we call Vegaviidae. Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft. Isolated bones coming from the Cretaceous and Paleogene of South America, Antarctica, and New Zealand are also referred to here to Vegaviidae and support the view that these basal anseriforms were abundant and diverse at high southern latitudes. Moreover, vegaviids represent the first avian lineage to have definitely crossed the K-Pg boundary, supporting the idea that some avian clades were not affected by the end Mesozoic mass extinction event, countering previous interpretations. Recognition of Vegaviidae indicates that modern birds were diversified in southern continents by the Cretaceous and reinforces the hypothesis indicating the important role of Gondwana for the evolutionary history of Anseriformes and Neornithes as a whole.

A large therian mammal from the Late Cretaceous of South America.

Theria represent an extant clade that comprises placental and marsupial mammals. Here we report on the discovery of a new Late Cretaceous mammal from southern Patagonia, Patagomaia chainko gen. et sp. nov., represented by hindlimb and pelvic elements with unambiguous therian features. We estimate Patagomaia chainko attained a body mass of 14 kg, which is considerably greater than the 5 kg maximum body mass of coeval Laurasian therians. This new discovery demonstrates that Gondwanan therian mammals acquired large body size by the Late Cretaceous, preceding their Laurasian relatives, which remained small-bodied until the beginning of the Cenozoic. Patagomaia supports the view that the Southern Hemisphere was a cradle for the evolution of modern mammalian clades, alongside non-therian extinct groups such as meridiolestidans, gondwanatherians and monotremes.

Comments on the pelvic girdle anatomy of Lagerpeton chanarensis Romer, 1971 (Archosauria) and its implications on the posture and gait of early pterosauromorphs.

Lagerpeton chanarensis is an early avemetatarsalian from the lower Carnian (lowermost Upper Triassic) levels of the Chañares Formation, La Rioja Province, Argentina. Lagerpeton and its kin were traditionally interpreted as dinosaur precursors of cursorial habits, with a bipedal posture and parasagittal gait. Some authors also speculated saltatorial capabilities for this genus. Recent analyses indicate that lagerpetids are early-diverging pterosauromorphs, a hypothesis that invites a review of most aspects of their anatomy and function. A revision of available specimens and additional preparation of previously known individuals indicate that Lagerpeton lacked a parasagittal gait and was probably a sprawling archosaur. This latter inference is based on the femoral head articulation with the acetabulum. The acetabular rim has a strongly laterally projected posteroventral antitrochanteric corner, which results in a position of the legs that recalls that of sprawling living reptiles, such as lizards, and departs from the parasagittally positioned limbs of dinosaurs. This may indicate that early pterosauromorphs had a sprawling posture of their hindlegs, casting doubts on the significance of bipedal posture and parasagittal gait for the radiation of early ornithodirans, given that both traits have been regarded as key features that triggered the ecological and evolutionary success of the clade. Our results bolster recent claims of a high ecomorphological diversity among early avemetatarsalians.

A new global palaeobiogeographical model for the late Mesozoic and early Tertiary.

Late Mesozoic palaeobiogeography has been characterized by a distinction between the northern territories of Laurasia and the southern landmasses of Gondwana. The repeated discovery of Gondwanan lineages in Laurasia has led to the proposal of alternative scenarios to explain these anomalous occurrences. A new biogeographical model for late Mesozoic terrestrial ecosystems is here proposed in which Europe and "Gondwanan" territories possessed a common Eurogondwanan fauna during the earliest Cretaceous. Subsequently, following the Hauterivian, the European territories severed from Africa and then connected to Asiamerica resulting in a faunal interchange. This model explains the presence of Gondwanan taxa in Laurasia and the absence of Laurasian forms in the southern territories during the Cretaceous. In order to test this new palaeobiogeographical model, tree reconciliation analyses (TRAs) were performed based on biogeographical signals provided by a supertree of late Mesozoic archosaurs. The TRAs found significant evidence for the presence of an earliest Cretaceous Eurogondwanan fauna followed by a relatively short-term Gondwana-Laurasia dichotomy. The analysis recovered evidence for a biogeographical reconnection of the European territories with Africa and South America-Antarctica during the Campanian to Maastrichtian time-slice. This biogeographical scenario appears to continue through the early Tertiary and sheds light on the trans-Atlantic disjunct distributions of several extant plant and animal groups.

How many species of Sturisoma (Siluriformes: Loricariinae) inhabit the La Plata Basin?

The aim of the present contribution is to review the taxonomy of the loricariid Sturisoma in the La Plata basin. The original description of the species S. barbatum is analyzed and compared to Regans later description of S. robustum. We noticed that Kners illustrations may have influenced the description by Regan and other later authors, leading to confusion in the taxonomy of Sturisoma. The diagnostic characteristics and molecular data of S. barbatum and S. robustum are discussed, and we concluded that the differences observed by previous authors are part of the intraspecific variation within a single species. This indicates that S. barbatum is a senior synonym of S. robustum. Sturisoma barbatum is compared to other Sturisoma species.

First monotreme from the Late Cretaceous of South America.

Monotremata is a clade of egg-lying mammals, represented by the living platypus and echidnas, which is endemic to Australia, and adjacent islands. Occurrence of basal monotremes in the Early Cretaceous of Australia has led to the consensus that this clade originated on that continent, arriving later to South America. Here we report on the discovery of a Late Cretaceous monotreme from southern Argentina, demonstrating that monotremes were present in circumpolar regions by the end of the Mesozoic, and that their distinctive anatomical features were probably present in these ancient forms as well.

A Maastrichtian insect assemblage from Patagonia sheds light on arthropod diversity previous to the K/Pg event.

Insect faunas from the latest Cretaceous are poorly known worldwide. Particularly, in the Southern Hemisphere, there is a gap regarding insect assemblages in the Campanian-Maastrichtian interval. Here we present an insect assemblage from the Maastrichtian Chorrillo Formation, southern Argentina, represented by well-preserved and non-deformed, chitinous microscopic remains including head capsules, wings and scales. Identified clades include Chironomidae dipterans, Coelolepida lepidopterans, and Ephemeroptera. The assemblage taxonomically resembles those of Cenozoic age, rather than other Mesozoic assemblages, in being composed by diverse chironomids and lepidopterans. To the best of our knowledge, present discovery constitutes the first insect body fossils for the Maastrichtian in the Southern Hemisphere, thus filling the gap between well-known Early Cretaceous entomofaunas and those of Paleogene age. The presented evidence shows that modern clades of chironomids were already dominant and diversified by the end of the Cretaceous, in concert with the parallel radiation of aquatic angiosperms which became dominant in freshwater habitats. This exceptional finding encourages the active search of microscopic remains of fossil arthropods in other geological units, which could provide a unique way of enhancing our knowledge on the past diversity of the clade.

Wing osteology, myology, and function of Rhea americana (Aves, Rheidae).

The Greater Rhea (Rhea americana, Rheidae) is a flightless paleognath with a wide geographical distribution in South America. The morphology of its shoulder girdle and wings are different from those of flying birds and some characteristics are similar to basal birds and paravian theropods. We present a detailed osteological, myological, and functional study of the shoulder and the wing of the Greater Rhea. Particular features of the anatomy of the pectoral girdle and wing of Rhea include the lack of triosseal canal, reduced origin area of the mm. pectoralis p. thoracica and supracoracoideus and the lack of a propatagium. The wing muscle mass is markedly reduced, reaching only 0.89% of total body mass (BM). Forelimb muscles mass values are low compared to those of flying birds and are congruent with the non-use of wings for active locomotion movements. R. americana does not flap the wings dorso-ventral as typical for flying birds, but predominantly in cranio-caudal direction, following a craniolateral to caudomedial abduction-adduction arc. When the wings are fully abducted, they are inverted L-shaped, with the inner surface caudally faced, and when the wings are folded against the body, they do not perform the complete automatic wing folding nor the circumduction of the manus, a movement performed by extant volant birds. This study complements our knowledge of the axial musculature of the flightless paleognaths and highlights the use of the Greater Rhea as a model, which may help understand the evolution of Palaeognathae, as well as the origin of flapping flight among paravian theropods.

New specimens provide insights into the anatomy of the dinosauriform Lewisuchus admixtus Romer, 1972 from the upper Triassic levels of the Chañares Formation, NW Argentina.

Lewisuchus admixtus is an early dinosauriform described by Alfred Romer in 1972 on the basis of a single, incomplete skeleton, collected in lower Upper Triassic rocks of the renowned Chañares Formation, at the Los Chañares type-locality, La Rioja Province, north-western Argentina. Recent field explorations to the type-locality resulted in the discovery of two partial articulated skeletons, which provide significant novel information. The cranial bones, presacral series, femur, tibia, and proximal tarsals of the new specimens match the preserved overlapping anatomy of the holotype and previously referred specimens of L. admixtus, including the presence of unique combination of character states among dinosauriforms (anterior presacral column with additional ossification on the top of neural spines, dorsal neural spines fan-shaped, anterior surface of the astragalus with a dorsally curved groove, and an inflated area on the anterior portion of the medial surface of this bone). This new information improves our understanding of the anatomy and taxonomy of early dinosauriforms and reinforces the role of Argentinean beds on the study of the origin of dinosaurs.

A large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous (Maastrichtian) of Patagonia, Argentina.

Megaraptora is a theropod clade known from former Gondwana landmasses and Asia. Most members of the clade are known from the Early to Late Cretaceous (Barremian-Santonian), with Maastrichtian megaraptorans known only from isolated and poorly informative remains. The aim of the present contribution is to describe a partial skeleton of a megaraptorid from Maastrichtian beds in Santa Cruz Province, Argentina. This new specimen is the most informative megaraptoran known from Maastrichtian age, and is herein described as a new taxon. Phylogenetic analysis nested the new taxon together with other South American megaraptorans in a monophyletic clade, whereas Australian and Asian members constitute successive stem groups. South American forms differ from more basal megaraptorans in several anatomical features and in being much larger and more robustly built.

The earliest dromaeosaurid theropod from South America.

The evolutionary history of Maniraptora, the clade of carnivorous dinosaurs that includes birds and the sickle-clawed Dromaeosauridae, has hitherto been largely restricted to Late Jurassic and Cretaceous deposits on northern continents. The stunning Early Cretaceous diversity of maniraptorans from Liaoning, China, coupled with a longevity implied by derived Late Jurassic forms such as Archaeopteryx, pushes the origins of maniraptoran lineages back to Pangaean times and engenders the possibility that such lineages existed in Gondwana. A few intriguing, but incomplete, maniraptoran specimens have been reported from South America, Africa and Madagascar. Their affinities remain contested, however, and they have been interpreted as biogeographic anomalies relative to other faunal components of these land-masses. Here we describe a near-complete, small dromaeosaurid that is both the most complete and the earliest member of the Maniraptora from South America, and which provides new evidence for a unique Gondwanan lineage of Dromaeosauridae with an origin predating the separation between northern and southern landmasses.

Unenlagiid theropods: are they members of the Dromaeosauridae (Theropoda, Maniraptora)?

In the present paper we analyze the phylogenetic position of the derived Gondwanan theropod clade Unenlagiidae. Although this group has been frequently considered as deeply nested within Deinonychosauria and Dromaeosauridae, most of the features supporting this interpretation are conflictive, at least. Modification of integrative databases, such as that recently published by Hu et al. (2009), produces significant changes in the topological distribution of taxa within Deinonychosauria, depicting unenlagiids outside this clade. Our analysis retrieves, in contrast, a monophyletic Avialae formed by Unenlagiidae plus Aves.