A new Cretaceous thyreophoran from Patagonia supports a South American lineage of armoured dinosaurs.

The early evolution of thyreophoran dinosaurs is thought to have occurred primarily in northern continents since most evidence comes from the Lower and Middle Jurassic of Europe and North America. The diversification into stegosaurs and ankylosaurs is obscured by a patchy fossil record comprising only a handful of fragmentary fossils, most with uncertain phylogenetic affinities. Here we report the discovery of a new armoured dinosaur from the early Late Cretaceous of Argentina, recovered phylogenetically using various datasets either as a basal thyreophoran or a stem ankylosaur, closely related to Scelidosaurus. It bears unusual anatomical features showing that several traits traditionally associated with the heavy Cretaceous thyreophorans did not occur universally. Jakapil kaniukura gen. et sp. nov. is the first definitive thyreophoran species from the Argentinian Patagonia. Unlike most thyreophorans, it seems to show a bipedal stance, as in Scutellosaurus. Jakapil also shows that early thyreophorans had a much broader geographic distribution than previously thought. It is a member of an ancient basal thyreophoran lineage that survived until the Late Cretaceous in South America.

A Triassic stem lepidosaur illuminates the origin of lizard-like reptiles.

The early evolution of diapsid reptiles is marked by a deep contrast between our knowledge of the origin and early evolution of archosauromorphs (crocodiles, avian and non-avian dinosaurs) to that of lepidosauromorphs (squamates (lizards, snakes) and sphenodontians (tuataras)). Whereas the former include hundreds of fossil species across various lineages during the Triassic period1, the latter are represented by an extremely patchy early fossil record comprising only a handful of fragmentary fossils, most of which have uncertain phylogenetic affinities and are confined to Europe1-3. Here we report the discovery of a three-dimensionally preserved reptile skull, assigned as Taytalura alcoberi gen. et sp. nov., from the Late Triassic epoch of Argentina that is robustly inferred phylogenetically as the earliest evolving lepidosauromorph, using various data types and optimality criteria. Micro-computed tomography scans of this skull reveal details about the origin of the lepidosaurian skull from early diapsids, suggesting that several traits traditionally associated with sphenodontians in fact originated much earlier in lepidosauromorph evolution. Taytalura suggests that the strongly evolutionarily conserved skull architecture of sphenodontians represents the plesiomorphic condition for all lepidosaurs, that stem and crown lepidosaurs were contemporaries for at least ten million years during the Triassic, and that early lepidosauromorphs had a much broader geographical distribution than has previously been thought.

Unique Tooth Morphology and Prismatic Enamel in Late Cretaceous Sphenodontians from Argentina.

Mammals and reptiles have evolved divergent adaptations for processing abrasive foods. Mammals have occluding, diphyodont dentitions with taller teeth (hypsodonty), more complex occlusal surfaces, continuous tooth eruption, and forms of prismatic enamel that prolong the functional life of each tooth [1, 2]. The evolution of prismatic enamel in particular was a key innovation that made individual teeth more resilient to abrasion in early mammals [2-4]. In contrast, reptiles typically have thin, non-prismatic enamel, and shearing, polyphyodont dentitions with multi-cusped or serrated tooth crowns, multiple tooth rows, rapid tooth replacement rates, or batteries made of hundreds of teeth [5-9]. However, there are rare cases where reptiles have evolved alternative solutions to cope with abrasive diets. Here, we show that the combined effects of herbivory and an ancestral loss of tooth replacement in a lineage of extinct herbivorous sphenodontians, distant relatives of the modern tuatara (Sphenodon punctatus) [10], are associated with the evolution of wear-resistant and highly complex teeth. Priosphenodon avelasi, an extinct sphenodontian from the Cretaceous of Argentina, possesses a unique cone-in-cone dentition with overlapping generations of teeth forming a densely packed tooth file. Each tooth is anchored to its predecessor via a rearrangement of dental tissues that results in a novel enamel-to-bone tooth attachment. Furthermore, the compound occlusal surfaces, thickened enamel, and the first report of prismatic enamel in a sphenodontian are convergent strategies with those in some mammals, challenging the perceived simplicity of acrodont dentitions [11-15] and showcasing the reptilian capacity to produce complex and unusual dentitions.

New skulls and skeletons of the Cretaceous legged snake Najash, and the evolution of the modern snake body plan.

Snakes represent one of the most dramatic examples of the evolutionary versatility of the vertebrate body plan, including body elongation, limb loss, and skull kinesis. However, understanding the earliest steps toward the acquisition of these remarkable adaptations is hampered by the very limited fossil record of early snakes. Here, we shed light on the acquisition of the snake body plan using micro-computed tomography scans of the first three-dimensionally preserved skulls of the legged snake Najash and a new phylogenetic hypothesis. These findings elucidate the initial sequence of bone loss that gave origin to the modern snake skull. Morphological and molecular analyses including the new cranial data provide robust support for an extensive basal radiation of early snakes with hindlimbs and pelves, demonstrating that this intermediate morphology was not merely a transient phase between limbed and limbless body plans.

The evolution of the axial skeleton intercentrum system in snakes revealed by new data from the Cretaceous snakes Dinilysia and Najash.

Snakes are an extremely modified and long-lived clade of lizards that have either lost or highly altered many of the synapomorphies that would clearly link them to their closest sister-group among squamates. We focus here on one postcranial morphological complex, the intercentrum system which in most non-ophidian squamates is limited to the cervical and caudal regions. The Cervical Intercentrum System (CeIS) is composed of a single intercentral element that sometimes articulates with a ventral projection (hypapophyses) of the centrum; the Caudal Intercentrum System (CaIS) is formed by an intercentral element, the haemal arch/chevron bone, and paired ventral projections of the centrum, the haemapophyses. In modern snakes, the intercentrum element of the CeIS is considered lost or fused to the hypaphophysis, and the chevron bone in CaIS is considered lost. Here, we describe new specimens of the early snake Dinilysia patagonica, and reinterpret previously known specimens of Dinilysia and Najash rionegrina, that do not show the expected snake morphology. The anatomy of these fossil taxa unambiguously shows that free cervical and caudal intercentra attached to distinct downgrowths (hypapophyses and haemapophyses) of the centra, are present in basal fossil snakes, and agrees with the proposed loss of post atlas-axis intercentra in later evolving snakes.

A new long-spined dinosaur from Patagonia sheds light on sauropod defense system.

Dicraeosaurids are a group of sauropod dinosaurs characterized by a distinctive vertebral column with paired, long, neural spines, present in an extreme fashion in the South American form Amargasaurus cazaui. This distinctive morphology has been interpreted as a support structure for a thermoregulatory sail, a padded crest for display, a dorsal hump acting as fat reservoir, and even as inner cores for dorsal horns. Other inferred functions (if any) of this structure were related to sexual display and/or defense strategies. Here we describe a new dicraeosaurid sauropod, Bajadasaurus pronuspinax gen. et sp. nov., from Patagonia which preserves the most complete skull of the group and has extremely elongate bifid cervical neural spines that point permanently forward, irrespective of the neck position. Although much shorter versions of this neural spine configuration were already recorded for other dicraeosaurid taxa, the long, anteriorly bent spines of this new dinosaur support the hypothesis that these elongate spines of dicraeosaurid sauropods served as passive defense structures.

Postcranial skeletal anatomy of the holotype and referred specimens of Buitreraptor gonzalezorum Makovicky, Apesteguía and Agnolín 2005 (Theropoda, Dromaeosauridae), from the Late Cretaceous of Patagonia.

Here we provide a detailed description of the postcranial skeleton of the holotype and referred specimens of Buitreraptor gonzalezorum. This taxon was recovered as an unenlagiine dromaeosaurid in several recent phylogenetic studies and is the best represented Gondwanan dromaeosaurid discovered to date. It was preliminarily described in a brief article, but a detailed account of its osteology is emerging in recent works. The holotype is the most complete specimen yet found, so an exhaustive description of it provides much valuable anatomical information. The holotype and referred specimens preserve the axial skeleton, pectoral and pelvic girdles, and both fore- and hindlimbs. Diagnostic postcranial characters of this taxon include: anterior cervical centra exceeding the posterior limit of neural arch; eighth and ninth cervical vertebral centra with lateroventral tubercles; pneumatic foramina only in anteriormost dorsals; middle and posterior caudal centra with a complex of shallow ridges on lateral surfaces; pneumatic furcula with two pneumatic foramina on the ventral surface; scapular blade transversely expanded at mid-length; well-projected flexor process on distal end of the humerus; dorsal rim of the ilium laterally everted; and concave dorsal rim of the postacetabular iliac blade. A paleohistological study of limb bones shows that the holotype represents an earlier ontogenetic stage than one of the referred specimens (MPCA 238), which correlates with the fusion of the last sacral vertebra to the rest of the sacrum in MPCA 238. A revised phylogenetic analysis recovered Buitreraptor as an unenlagiine dromaeosaurid, in agreement with previous works. The phylogenetic implications of the unenlagiine synapomorphies and other characters, such as the specialized pedal digit II and the distal ginglymus on metatarsal II, are discussed within the evolutionary framework of Paraves.

The first iguanian lizard from the Mesozoic of Africa.

The fossil record shows that iguanian lizards were widely distributed during the Late Cretaceous. However, the biogeographic history and early evolution of one of its most diverse and peculiar clades (acrodontans) remain poorly known. Here, we present the first Mesozoic acrodontan from Africa, which also represents the oldest iguanian lizard from that continent. The new taxon comes from the Kem Kem Beds in Morocco (Cenomanian, Late Cretaceous) and is based on a partial lower jaw. The new taxon presents a number of features that are found only among acrodontan lizards and shares greatest similarities with uromastycines, specifically. In a combined evidence phylogenetic dataset comprehensive of all major acrodontan lineages using multiple tree inference methods (traditional and implied weighting maximum-parsimony, and Bayesian inference), we found support for the placement of the new species within uromastycines, along with Gueragama sulamericana (Late Cretaceous of Brazil). The new fossil supports the previously hypothesized widespread geographical distribution of acrodontans in Gondwana during the Mesozoic. Additionally, it provides the first fossil evidence of uromastycines in the Cretaceous, and the ancestry of acrodontan iguanians in Africa.

An Unusual New Theropod with a Didactyl Manus from the Upper Cretaceous of Patagonia, Argentina.

BACKGROUND: Late Cretaceous terrestrial strata of the Neuquén Basin, northern Patagonia, Argentina have yielded a rich fauna of dinosaurs and other vertebrates. The diversity of saurischian dinosaurs is particularly high, especially in the late Cenomanian-early Turonian Huincul Formation, which has yielded specimens of rebacchisaurid and titanosaurian sauropods, and abelisaurid and carcharodontosaurid theropods. Continued sampling is adding to the known vertebrate diversity of this unit. METHODOLOGY/ PRINCIPAL FINDINGS: A new, partially articulated mid-sized theropod was found in rocks from the Huincul Formation. It exhibits a unique combination of traits that distinguish it from other known theropods justifying erection of a new taxon, Gualicho shinyae gen. et sp. nov. Gualicho possesses a didactyl manus with the third digit reduced to a metacarpal splint reminiscent of tyrannosaurids, but both phylogenetic and multivariate analyses indicate that didactyly is convergent in these groups. Derived characters of the scapula, femur, and fibula supports the new theropod as the sister taxon of the nearly coeval African theropod Deltadromeus and as a neovenatorid carcharodontosaurian. A number of these features are independently present in ceratosaurs, and Gualicho exhibits an unusual mosaic of ceratosaurian and tetanuran synapomorphies distributed throughout the skeleton. CONCLUSIONS/ SIGNIFICANCE: Gualicho shinyae gen. et sp. nov. increases the known theropod diversity of the Huincul Formation and also represents the first likely neovenatorid from this unit. It is the most basal tetatanuran to exhibit common patterns of digit III reduction that evolved independently in a number of other tetanuran lineages. A close relationship with Deltadromaeus from the Kem Kem beds of Niger adds to the already considerable biogeographic similarity between the Huincul Formation and coeval rock units in North Africa.

The youngest South American rhynchocephalian, a survivor of the K/Pg extinction.

Rhynchocephalian lepidosaurs, though once widespread worldwide, are represented today only by the tuatara (Sphenodon) of New Zealand. After their apparent early Cretaceous extinction in Laurasia, they survived in southern continents. In South America, they are represented by different lineages of Late Cretaceous eupropalinal forms until their disappearance by the Cretaceous/Palaeogene (K/Pg) boundary. We describe here the only unambiguous Palaeogene rhynchocephalian from South America; this new taxon is a younger species of the otherwise Late Cretaceous genus Kawasphenodon. Phylogenetic analysis confirms the allocation of the genus to the clade Opisthodontia. The new form from the Palaeogene of Central Patagonia is much smaller than Kawasphenodon expectatus from the Late Cretaceous of Northern Patagonia. The new species shows that at least one group of rhynchocephalians not related to the extant Sphenodon survived in South America beyond the K/Pg extinction event. Furthermore, it adds to other trans-K/Pg ectotherm tetrapod taxa, suggesting that the end-Cretaceous extinction affected Patagonia more benignly than the Laurasian landmasses.

A diplodocid sauropod survivor from the early cretaceous of South America.

Diplodocids are by far the most emblematic sauropod dinosaurs. They are part of Diplodocoidea, a vast clade whose other members are well-known from Jurassic and Cretaceous strata in Africa, Europe, North and South America. However, Diplodocids were never certainly recognized from the Cretaceous or in any other southern land mass besides Africa. Here we report a new sauropod, Leikupal laticauda gen. et sp. nov., from the early Lower Cretaceous (Bajada Colorada Formation) of Neuquén Province, Patagonia, Argentina. This taxon differs from any other sauropod by the presence of anterior caudal transverse process extremely developed with lateroventral expansions reinforced by robust dorsal and ventral bars, very robust centroprezygapophyseal lamina in anterior caudal vertebra and paired pneumatic fossae on the postzygapophyses in anterior-most caudal vertebra. The phylogenetic analyses support its position not only within Diplodocidae but also as a member of Diplodocinae, clustering together with the African form Tornieria, pushing the origin of Diplodocoidea to the Middle Jurassic or even earlier. The new discovery represents the first record of a diplodocid for South America and the stratigraphically youngest record of this clade anywhere.

The Miocene mammal Necrolestes demonstrates the survival of a Mesozoic nontherian lineage into the late Cenozoic of South America.

The early Miocene mammal Necrolestes patagonensis from Patagonia, Argentina, was described in 1891 as the only known extinct placental "insectivore" from South America (SA). Since then, and despite the discovery of additional well-preserved material, the systematic status of Necrolestes has remained in flux, with earlier studies leaning toward placental affinities and more recent ones endorsing either therian or specifically metatherian relationships. We have further prepared the best-preserved specimens of Necrolestes and compared them with newly discovered nontribosphenic Mesozoic mammals from Argentina; based on this, we conclude that Necrolestes is related neither to marsupials nor placentals but is a late-surviving member of the recently recognized nontherian clade Meridiolestida, which is currently known only from SA. This conclusion is supported by a morphological phylogenetic analysis that includes a broad sampling of therian and nontherian taxa and that places Necrolestes within Meridiolestida. Thus, Necrolestes is a remnant of the highly endemic Mesozoic fauna of nontribosphenic mammals in SA and extends the known record of meridiolestidans by almost 45 million years. Together with other likely relictual mammals from earlier in the Cenozoic of SA and Antarctica, Necrolestes demonstrates the ecological diversity of mammals and the mosaic pattern of fauna replacement in SA during the Cenozoic. In contrast to northern continents, the Cenozoic faunal history of SA was characterized by a long period of interaction between endemic mammalian lineages of Mesozoic origin and metatherian and eutherian lineages that probably dispersed to SA during the latest Cretaceous or earliest Paleocene.

Cretaceous small scavengers: feeding traces in tetrapod bones from Patagonia, Argentina.

Ecological relationships among fossil vertebrate groups are interpreted based on evidence of modification features and paleopathologies on fossil bones. Here we describe an ichnological assemblage composed of trace fossils on reptile bones, mainly sphenodontids, crocodyliforms and maniraptoran theropods. They all come from La Buitrera, an early Late Cretaceous locality in the Candeleros Formation of northwestern Patagonia, Argentina. This locality is significant because of the abundance of small to medium-sized vertebrates. The abundant ichnological record includes traces on bones, most of them attributable to tetrapods. These latter traces include tooth marks that provde evidence of feeding activities made during the sub-aerial exposure of tetrapod carcasses. Other traces are attributable to arthropods or roots. The totality of evidence provides an uncommon insight into paleoecological aspects of a Late Cretaceous southern ecosystem.

Highly specialized mammalian skulls from the Late Cretaceous of South America.

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.

Unenlagiinae revisited: dromaeosaurid theropods from South America.

Over the past two decades, the record of South American unenlagiine dromaeosaurids was substantially increased both in quantity as well as in quality of specimens. Here is presented a summary review of the South American record for these theropods.Unenlagia comahuensis, Unenlagia paynemili, and Neuquenraptor argentinus come from the Portezuelo Formation, the former genus being the most complete and with putative avian features. Neuquenraptor is more incomplete and exhibits pedal features resembling those of Unenlagia. The earliest and most complete South American dromaeosaurid is Buitreraptor gonzalezoru, whose preserved cranial remains, provides important data in the characterization of unenlagiines. The most recently described,Austroraptor cabazai also with cranial remains, allows further comparisons with Laurasian lineages and a better characterization of unenlagiines. The possible synonymy between nenlagia and Neuquenraptor is discussed. Additional evidences from Brazil and Colombia show that dinosaurs with similar dentition to that of unenlagiines were present in the whole South America. However, it is not possible to discard that these remains may belong to other unknown maniraptoran lineages, considering the increasing number of taxa of this group found in South America.

Tunasniyoj, a dinosaur tracksite from the Jurassic-Cretaceous boundary of Bolivia.

Here we report a superbly preserved and profusely represented five-ichnotaxa dinosaur track assemblage near Icla village, 100 km southeast of Sucre, Bolivia. As preserved in reddish Jurassic-Cretaceous boundary aeolian sandstones, this rich and uncommon assemblage is, additionally, the oldest dinosaur tracksite for Bolivia. Four trackmakers were identified in the area: three quadrupedal and one bipedal, all of them with tracks of around 35 cm in lenght. One of the quadrupedals is represented by no less than five adult individuals (ichnotaxon A), and four purported juveniles (ichnotaxon B) walking in association. The other two quadrupedals (ichnotaxa C and D) involve four trackways, and the last, the bipedal trackmaker (ichnotaxon E), is represented by one trackway. The five ichnotaxa represented in the "Palmar de Tunasniyoj" could be tentatively assigned to the following trackmakers: Ichnotaxa A and B are assigned to basal stegosaurians; ichnotaxon C to a basal tyreophoran, perhaps related to the ankylosaur lineage; ichnotaxon D to the Ankylosauria, and ichnotaxon E to Theropoda. The Tunasniyoj assemblage, the oldest dinosaur tracksite for Bolivia, includes the oldest known evidence assigned to ankylosaurs and stegosaurs for South America.

Unenlagiinae revisited: dromaeosaurid theropods from South América

Over the past two decades, the record of South American unenlagiine dromaeosaurids was substantially increased both in quantity as well as in quality of specimens. Here is presented a summary review of the South American record for these theropods.Unenlagia comahuensis, Unenlagia paynemili, and Neuquenraptor argentinus come from the Portezuelo Formation, the former genus being the most complete and with putative avian features. Neuquenraptor is more incomplete and exhibits pedal features resembling those of Unenlagia. The earliest and most complete South American dromaeosaurid is Buitreraptor gonzalezoru, whose preserved cranial remains, provides important data in the characterization of unenlagiines. The most recently described,Austroraptor cabazai also with cranial remains, allows further comparisons with Laurasian lineages and a better characterization of unenlagiines. The possible synonymy between nenlagia and Neuquenraptor is discussed. Additional evidences from Brazil and Colombia show that dinosaurs with similar dentition to that of unenlagiines were present in the whole South America. However, it is not possible to discart that these remains may belong to other unknown maniraptoran lineages, considering the increasing number of taxa of this group found in South America.

Nas últimas duas décadas, o registro do dromaeossaurídeos Unenlagiinae da América do Sul aumentou substancialmente tanto em quantidade, assim como na qualidade dos espécimes. Uma revisão sumária do registro sul americano para estes terópodos é apresentada aqui. Unenlagia comahuensis, Unenlagia paynemili e Neuquenraptor argentinus provenientes da Formação Portezuelo, sendo o primeiro o mais completo e com características avianas putativas. Neuquenraptor, muito incompleto, demonstra as características do pé que assemelhamse àquelas de Unenlagia. O mais velho, e mais completo dromaeossaurídeo americano é Buitreraptor gonzalezorum, cujo possui preservado ossos cranianos, fornece implicações importantes na caracterização dos unenlagiines. Descrito recentemente, Austroraptor cabazai também com ossos cranianos, permite mais comparações com linhagens provenientes da Laurasia e uma melhor caracterização dos unenlagiineos. A possível sinonímia entre Unenlagia e Neuquenraptor é discutido. As evidências adicionais do Brasil e da Colômbia mostram que dinossauros com dentição semlhante à dos unenlagiineos eram representados em toda América do Sul; entretanto, estas evidências podem igualmente ser relacionadas a outros maniraptores desconhecidos, considerando a grande diversidade do grupo na América do Sul.

Tunasniyoj, a dinosaur tracksite from the Jurassic-Cretaceous boundary of Bolivia

Here we report a superbly preserved and profusely represented five-ichnotaxa dinosaur track assemblage near Icla village, 100 km southeast of Sucre, Bolivia. As preserved in reddish Jurassic-Cretaceous boundary aeolian sandstones, this rich and uncommon assemblage is, additionally, the oldest dinosaur tracksite for Bolivia. Four trackmakers were identified in the area: three quadrupedal and one bipedal, all of them with tracks of around 35 cm in lenght. One of the quadrupedals is represented by no less than five adult individuals (ichnotaxon A), and four purported juveniles (ichnotaxon B) walking in association. The other two quadrupedals (ichnotaxa C and D) involve four trackways, and the last, the bipedal trackmaker (ichnotaxon E), is represented by one trackway. The five ichnotaxa represented in the "Palmar de Tunasniyoj" could be tentatively assigned to the following trackmakers: Ichnotaxa A and B are assigned to basal stegosaurians; ichnotaxon C to a basal tyreophoran, perhaps related to the ankylosaur lineage; ichnotaxon D to the Ankylosauria, and ichnotaxon E to Theropoda. The Tunasniyoj assemblage, the oldest dinosaur tracksite for Bolivia, includes the oldest known evidence assigned to ankylosaurs and stegosaurs for South America.

Neste estudo é relatado um conjunto magnificamente preservado de pistas de dinossauros representados por cinco táxons distintos, situado próximo à Vila de Icla, 100 quilômetros a sudeste do Sucre, Bolívia. Preservados em arenitos eólicos avermelhados do limite Jurássico-Cretáceo. Quatro formas geradoras foram identificadas na área, sendo três quadrúpedes e um bípede, todos com as pegadas com cerca de 35 cm de comprimento. Um dos quadrúpedes (trilhas tipos A e B) é representado por pelo menos seis indivíduos em dois grupos de três, com adultos e juvenis juntos. Os outros dois quadrúpedes (trilhas tipos C e D) são geradores de quatro trilhas, e um gerador bípede (trilha E), é representado por uma trilha. Os quatro icnotáxons representados em "Palmar de Tunasniyoj", podem provisoriamente ser atribuídos aos seguintes geradores: Icnotáxon A e B são atribuídos a estegossauros basais. Icnotáxon C a um tireóforo basal, talvez relacionado à linhagem dos anquilossauros. Icnotáxon D aos Ankylosauria; e icnotáxon E a Theropoda. Este raro e belo conjunto de trilhas de dinossauros de Tunasniyoj, representa ainda a icnocenose mais antiga de dinossauros da Bolívia. Além disso, inclui a mais antiga evidência conhecida atribuída a anquilossauros e estegossauros para América do sul.

A Cretaceous terrestrial snake with robust hindlimbs and a sacrum.

It has commonly been thought that snakes underwent progressive loss of their limbs by gradual diminution of their use. However, recent developmental and palaeontological discoveries suggest a more complex scenario of limb reduction, still poorly documented in the fossil record. Here we report a fossil snake with a sacrum supporting a pelvic girdle and robust, functional legs outside the ribcage. The new fossil, from the Upper Cretaceous period of Patagonia, fills an important gap in the evolutionary progression towards limblessness because other known fossil snakes with developed hindlimbs, the marine Haasiophis, Pachyrhachis and Eupodophis, lack a sacral region. Phylogenetic analysis shows that the new fossil is the most primitive (basal) snake known and that all other limbed fossil snakes are closer to the more advanced macrostomatan snakes, a group including boas, pythons and colubroids. The new fossil retains several features associated with a subterranean or surface dwelling life that are also present in primitive extant snake lineages, supporting the hypothesis of a terrestrial rather than marine origin of snakes.

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.