A Complete Skull of an Early Cretaceous Sauropod and the Evolution of Advanced Titanosaurians

Advanced titanosaurian sauropods, such as nemegtosaurids and saltasaurids, were diverse and one of the most important groups of herbivores in the terrestrial biotas of the Late Cretaceous. However, little is known about their rise and diversification prior to the Late Cretaceous. Furthermore, the evolution of their highly-modified skull anatomy has been largely hindered by the scarcity of well-preserved cranial remains. A new sauropod dinosaur from the Early Cretaceous of Brazil represents the earliest advanced titanosaurian known to date, demonstrating that the initial diversification of advanced titanosaurians was well under way at least 30 million years before their known radiation in the latest Cretaceous. The new taxon also preserves the most complete skull among titanosaurians, further revealing that their low and elongated diplodocid-like skull morphology appeared much earlier than previously thought.

Evolution of Diplodocid Sauropod dinosaurs with emphasis on specimens from Howe Ranch, Wyoming (USA)

Diplodocidae are among the best known sauropod dinosaurs. Several species were described in the late 1800s or early 1900s. Since then, numerous additional specimens were recovered in the USA, Tanzania, Portugal, as well as possibly Spain, England, and Asia. To date, the clade includes about 12 to 15 different species, some of them with questionable taxonomic status (e.g. ‘Diplodocus’ hayi or Dyslocosaurus polyonychius). However, intrageneric relationships of the multi-species, iconic genera Apatosaurus and Diplodocus are still poorly known. The way to resolve this issue is a specimen-based phylogenetic analysis, which was done for Apatosaurus, but is here performed for the first time for the entire clade of Diplodocidae. New material from different localities and stratigraphic levels on the Howe Ranch (Shell,Wyoming, USA) sheds additional light on the evolution of Diplodocidae. Three new specimens are described herein, considerably increasing our knowledge of the anatomy of the group. The new specimens (SMA 0004, SMA 0011, and SMA 0087) represent two, to possibly three new diplodocid species. They preserve material from all parts of the skeleton, including two nearly complete skulls, as well as fairly complete manus and pedes, material which is generally rare in diplodocids. Thereby, they considerably increase anatomical overlap between the sometimes fragmentary holotype specimens of the earlier described diplodocid species, allowing for significant results in a specimenbased phylogenetic analysis. Furthermore, clavicles and interclavicles are identified, the latter for the first time in dinosaurs. Their presence seems restricted to early sauropods, flagellicaudatans, and early Macronaria, and might thus be a retained plesiomorphy, with the loss of these bones being synapomorphic for Titanosauriformes and possibly Rebbachisauridae. The new material allows to test previous hypotheses of diplodocid phylogeny. In order to do so, any type specimen previously proposed to belong to Diplodocidae was included in the study, as are relatively complete referred specimens, in order to increase the degree of overlapping material. For specimens subsequently suggested to be non-diplodocid sauropods, their hypothesized sister taxa were included as outgroups. The current phylogenetic analysis thus includes 76 operational taxonomic units, 45 of which belong to Diplodocidae. The specimens were scored for 477 morphological characters, representing one of the most extensive phylogenetic analyses done within sauropod dinosaurs. The resulting cladogram recovers the classical arrangement of diplodocid relationships. Basing on a newly developed numerical approach to reduce subjectivity in the decision of specific or generic separation, species that have historically been included into well-known genera like Apatosaurus or Diplodocus, were detected to be actually generically different. Thereby, the famous genus Brontosaurus is resuscitated, and evidence further suggests that also Elosaurus parvus (previously referred to Apatosaurus) or ‘Diplodocus’ hayi represent unique genera. The study increases our knowledge about individual variation, and helps to decide how to score multi-species genera. Such a specimen-based phylogenetic analysis thus proves a valuable tool to validate historic species in sauropods, and in paleontology as a whole.

Dinossáurios Eocretácicos de Lagosteiros

According to an ancient folkloric legend, Our Lady, stepping down from the sea, would have rided on a mule to the platform above the cliffs named Pedra da Mua at Lagosteiros'bay, near Espichel cape. Mule's footprints, regarded by fishermen as evidence, would be clearly recognizable on exposed surfaces of the rocks. Indeed there are footprints but from Dinosaurs of latest Jurassic, Portlandian age, this spectacular locality being specially rich in giant Sauropod tracks (that have seldom been found elsewhere in Europe). As we proceeded to its study, another locality with Dinosaur footprints, Lower Cretaceous (Hauterivian) in age, was found on the northern cliffs at Lagosteiros. It is probably the richest one in european Lower Cretaceous and the only of this age known in Portugal, so we decided to give priority to its study. Dinosaur tracks have been printed on calciclastic sands in a lagoonal environment protected by fringing coral reefs. There have been emersion episodes; beaches were frequented by Dinosaurs. Later on, the marine barremian ingression restablished a gulf and such animals could not come here any more. Under a paleogeographical viewpoint, the evidence of a marine regression near the end of Hauterivian is to be remarked. Five types of tracks and footprints have been recognized: - Neosauropus lagosteirensis, new morphogenus and species, tracks from a giant Sauropod, perhaps from Camarasaurus; with its proportions the total length of the author would be about 15,5 m. These are the only Sauropod tracks known till now in Europe's Lower Cretaceous. - tracks from a not so big quadruped, maybe a Sauropod (young individual?); however it is not impossible that they were produced by Stegosaurians or Ankylosaurians. -Megalosauropus (?Eutynichnium) gomesi new morphospecies, four Theropod tracks most probably produced by megalosaurs. - Iguanodon sp., represented by some footprints and specially by a set corresponding to the feet and tail from an individual standing in a rest position. - problematical, quite small-sized biped (maybe an Ornithopod related to Camptosaurus). Evidence points to a richer fauna than that known in barremian "Dinosaur sandstones" from a nearby locality, Boca do Chapim. Lagosteiros' association clearly indicates the predominance of herbivores, which required large amounts of vegetable food in the neighbourhood. This is an indirect evidence of the vegetal wealth, also suggested by associations of plant macrofossils, polen and spores found in early Cretaceous sediments at the same region. The relatively high proportion of Theropoda is related to the wealth of the whole fauna, which comprised a lot of the prey needed by such powerful flesh-eaters. The evidence, as a whole, points out to a warm and moist climate. All the tracks whose direction could be measured are directed to the southern quadrants, this being confirmed by the approximative direction of other footprints. Massive displacements (migration?) could take place during a brief emersion episode. This may result from the ingression of barremian seas, flooding the region and restablishing here a small gulf. Even if the arrival of the waters damaged certain footprints it has not destroyed them completely, thus allowing the preservation of such evidence from a remote past.

Recerca paleontològica als jaciments maastrichtians continentals de l'àrea de Fumanya (Alt Berguedà)

Memòria de la recerca realitzada a Vallcebre durant el 2006. S’ha estudiat el registre fòssil de l'àrea del sinclinal d’aquesta població de vertebrats i paleobotànic, concretament dels materials maastrichtians d'origen continental que hi afloren. S'han realitzat excavacions per recuperar restes òssies de vertebrats (per exemple dinosaures sauròpodes i cocodrils eosuquis) i cartografies d'icnites i rastres de titanosaure i de restes vegetals als megajaciments de Fumanya sud i Mina Esquirol. També s'ha avançat en l'estudi de postes de titanosaure excavades en campanyes d'anys anteriors. El treball realitzat està permeten acumular dades que seran de gran utilitat i rellevança científica per entendre com vivien els darrers dinosaures pirinencs.La recerca del projecte s'està realitzant en col•laboració amb l'Institut Català de Paleontologia, el Departament d'Estratigrafia de la Universitat Autònoma de Barcelona, el Departament d'Estratigrafia, Paleontologia i Geociències Marines de la Universitat de Barcelona i el Departament de Geociències de la Universitat de Rennes (França).

Stratigraphy and palaeoenvironment of Early Jurassic dinosaur-bearing volcaniclastic deposits from Toundoute (Province of Ouarzazate, Haut-Atlas, Morocco)

The Early Jurassic dinosaur site of Toundoute which yielded the basal sauropod Tazoudasaurus naimi is examined in the light of its stratigraphic, sedimentological and palaeoenvironmental context. A thin succession of Early Liassic marine carbonates (probably Hettangian-Sinemurian in age) is continuously overlain by continental beds with dinosaurs. These latter are assumed to be of Middle to Late Liassic age. The continental deposits include a large part of volcanoclastics, different from the Triassic basalts. The Jurassic volcanoclastics originated from an unknown but obviously close eruption centre. The continental sediments (channels and flood plain) were deposited under tropical climate conditions with alternating humid and dry episodes. The bones occur as isolated or partly articulated elements (parts of carcasses). The two bone-beds are related to typical mud-flows. This type of transport protected the bones from erosion, and favoured their burying and fossilization.

Paleobiology and skeletochronology of Jurassic dinosaurs: implications from the histology and oxygen isotope compositions of bones

Fossil biogenic phosphate of fast-growing primary bone tissue of dinosaurs can preserve a histologic and isotopic time-series of annual seasonality in temperature variations, similar to tooth enamel and other accretionary skeletal phases such as corals or wood. On two bone fragments from sympatric dinosaurs with different histologic patterns of bone growth, high-resolution oxygen isotope profiles were analyzed along the radial direction of bone growth. The investigated specimens are from the Jurassic Shishugou Formation in the Junggar Basin, NW China and have distinct patterns of compositional variation. A fibrolamellar dinosaur bone with multiple lines of arrested growth (LAGs) and periodic growth cycles of decreasing bone laminae thickness displays a cyclic intra-bone variation in delta(18)O values of about 2parts per thousand corresponding with the LAGs. These growth cycles in fast-growing fibrolamellar bone provide evidence for seasonal growth of dinosaurs in lower latitudes ( similar to 45degreesN), possibly influenced by a monsoon-type paleoclimate. Seasonal changes in temperature and water supply are consistent with the oxygen isotope composition measured in dinosaur bone phosphate as well as with growth rings in contemporaneous fossil conifer wood from the same locality. In contrast, a plexiform sympatric sauropod bone displays continuous growth, free of LAGs and has a lower intra-bone variation of less than or equal to 0.8parts per thousand. Differences in bone histology are also reflected in the oxygen isotopic composition and its intra-bone variability, indicating different physiological responses to external climatic stress between sympatric dinosaur species. Seasonal intra-bone oxygen isotope variations combined with bone histology may thus yield new insights into species-specific response to climatic stress and its influence on dinosaur growth, formation of growth marks, growth rates, as welt as dinosaur thermophysiology. (C) 2004 Elsevier B.V All rights reserved.

Trace fossils and pseudofossils from the Wealden strata (non-marine Lower Cretaceous) of southern England

The trace fossils of the Wealden (non-marine Lower Cretaceous) of southern England are described. Sixteen invertebrate ichnotaxa include Agrichnium fimbriatus, Beaconites antarcticus, B. barretti, Cochlichnus anguineus, Diplichnites triassicus, Diplocraterion parallelum, Lockeia siliquaria, L. serialis, Monocraterion cf. tentaculum, Palaeophycus striatus, P. tubularis, Planolites montanus, Protovirgularia rugosa, Rhizocorallium isp., Scoyenia cf. gracilis, Unisulcus minutus, insect and root traces. Tetrapod tracks and trackways include tridactyl Iguanodontipus burreyi and other ornithopods, theropod, and tetradactyl sauropod (or possibly ankylosaur), together with extensive dinosaur tramplings. Coprolites are referred to two broad types: spiral, with or without included fish scales (attributable to sharks), and elongate and irregular (possibly produced by reptiles). A skinprint and two types of pseudofossil are also included. Five environmental associations are recognised: (1) lacustrine/lagoonal; (2) brackish incursions (flooding events) into the lacustrine/lagoonal environment; (3) a marginal lacustrine association with fluvial input; (4) a fluvial (lacustrine delta) association; (5) floodplain sediments (seasonal wetlands). These associations are assigned to the fluvial-lacustrine Scoyenia Ichnofacies and the incursions to Glossifungites lchnofacies. (c) 2005 Elsevier Ltd. All rights reserved.

Registro de dinossauros na região da cidade de Coroatá, EO/Mesoalbiano da Bacia de São Luís-Grajaú, Estado do Maranhão

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

WATER VAPOR CONDUCTANCE OF THE LOWER CRETACEOUS DINOSAURIAN EGGS FROM SANAGASTA, LA RIOJA, ARGENTINA: PALEOBIOLOGICAL AND PALEOECOLOGICAL IMPLICATIONS FOR SOUTH AMERICAN FAVEOLOOLITHID AND MEGALOOLITHID EGGS

The water vapor conductance (G(H20)) of the neosauropod eggs from the Lower Cretaceous Sanagasta nesting site in La Rioja Province, Argentina, was examined and compared with other Cretaceous Argentinean oological material. The 2900 mgH(2)O/day.Torr G(H2O) of the Sanagasta eggshells confirms an extremely moist nesting environment and supports field observations of dug-out nests in a geothermal setting. The observed thinning of the outer eggshell surface during incubation increases gas conductance and concomitantly decreases eggshell mechanical resistance during the late ontogenetic stages, thus facilitating embryonic development and hatching. The Sanagasta and Entre Rios Province faveoloolithid eggs display the highest and comparable 61120 values and share several morphological and diagenetic characters, indicating comparable nesting strategy in geothermal settings. However, the faveoloolithid Yamintie and La Pampa Province specimens cluster together with lower G(H20) values closer to the megaloolithid eggs. The Gnu) of the megaloolithid egg Megaloolithus patagonicus was reconsidered and new results are now congruent with other reported megaloolithid GH2O values. Additionally, we hypothesize that V-shaped pore canals of M. patagonicus, which upper sections reach only the top third or half eggshell thickness and, a wider section in the middle would not compromise the overall egg mechanical resistance like vertical pores connecting directly the outer to the inner eggshell surfaces. Such pore spatial arrangement and geometry would enhance, as the eggshell thins during incubation, a greater G(H2O), G(O2) and G(CO2) and facilitate embryonic development in high moisture nesting contents. Overall, data suggests that neosauropod nesting and brooding behaviors were dependent on elevated moisture nesting environments.

Die Reproduktion von Dinosauriern, speziell der Sauropoden, und deren Bedeutung für ihren Gigantismus

Es wurde ein Teil der life-history, die Reproduktion, von Dinosauriern, speziell der Sauropoden, den größten bekannten jemals auf der Erde existierenden Landtieren, untersucht, um unter anderem den Zusammenhang zwischen Gigantismus und Reproduktion zu erforschen. Hierzu wurde eine mögliche life-history für Sauropoden, auf Grundlage des heutigen Forschungsstands in der Biologie und der Paläontologie, anhand einer Literaturrecherche erstellt. Des Weiteren wurde ein Modell zur Reproduktion bei ausgestorbenen oviparen Amnioten, basierend auf bestehenden Zusammenhängen zwischen Körpergröße und verschiedenen masse-spezifischen Reproduktionsmerkmalen (Eigewicht, Gelegegewicht, jähr. Gelegegewicht) bei rezenten oviparen Amnioten, erarbeitet. Mit Hilfe dieses Modells und Informationen aus Fossilfunden wurde der Frage nachgegangen, wie diese Reproduktionsmerkmale bei Dinosauriern wahrscheinlich ausgesehen haben. Weiterhin erfolgte die Überprüfung der Hypothese, dass Dinosaurier, insbesondere Sauropoden, eine höhere Reproduktionskapazität hatten als gleich große landlebende Säugetiere, was ersteren im Vergleich zu letzteren ermöglicht haben soll so viel größer zu werden (Janis und Carrano 1992). rnDie Untersuchungen der Zusammenhänge zwischen Körpergewicht und den masse-spezifischen Reproduktionsmerkmalen ergaben, dass das Körpergewicht immer stark mit den untersuchten Reproduktionsmerkmalen korreliert war. Große Vögel und große Reptilien unterscheiden sich in ihrem relativen Eigewicht (Eigewicht/Körpergewicht). Vögel haben relativ größere Eier. Betrachtet man das relative Gelegegewicht oder das relative jährliche Gelegegewicht so wird der Unterschied kleiner bzw. ist zwischen manchen Reptilien- und Vogelgruppen nicht mehr vorhanden. Dinosaurier hatten relative Eigewichte, die zwischen denen von Reptilien und Vögel liegen. Basale Dinosaurier, wie Prosauropoden, waren in ihrer Reproduktion eher reptilien-ähnlich, während vogel-ähnliche Theropoden eine Reproduktion hatten, die sich besser durch ein Vogelmodel beschreiben lässt. Die Reproduktion anderer Dinosaurier, wie Sauropoden und Hadrosaurier, lässt sich nicht eindeutig durch eines der beiden Modelle beschreiben und/oder die Modelle variierten in Abhängigkeit des betrachteten Merkmals. Trotzdem war es möglich für alle untersuchten Dinosaurier eine Abschätzung zur Gelegegröße und der Anzahl der jährlich gelegten Eier zu machen. Diese Schätzungen ergaben, dass die vermutete hohe Reproduktionskapazität von mehreren hundert Eiern pro Jahr nur für extrem große Sauropoden (70 t) haltbar ist. rnMit Ausnahme der Nagetiere fand ich die Unterschiede in der Reproduktionskapazität von Vögeln und Säugetieren, die Janis und Carrano (1992) postulierten, sogar auf der Ebene von Ordnungen. Dinosauriergelege waren größer als die Würfe von gleichgroßen (extrapolierten) Säugetieren während die Gelegegröße von gleichgroßen (extrapolierten) Vögeln ähnlich der von Sauropoden war. Da das Aussterberisiko häufig mit niedriger Reproduktionskapazität korreliert ist, impliziert dies ein geringeres Aussterberisiko großer Dinosaurier im Vergleich zu großen Säugetieren. Populationen sehr großer Dinosaurier, wie der Sauropoden, konnten vermutlich daher, über evolutionäre Zeiträume betrachtet, sehr viel länger existieren als Populationen großer Säugetiere.rn

El icnogénero Iguanodontipus en el yacimiento de “Las Cuestas I” (Santa Cruz de Yanguas, Soria, España)= The Iguanodontipus ichnogener in “Las Cuestas I” tracksite (Santa Cruz de Yanguas, Soria. Spain)]

Se define este icnogénero en el yacimiento paleontológico citado perteneciente al grupo Oncala de la facies Weald de la Cuenca de Cameros. El yacimiento de Las Cuestas I (Soria, España) es uno de los mayores del Grupo Oncala. Hasta el momento, se han catalogado en él casi 600 pisadas de ornitópodos, terópodos y, sobre todo, de saurópodos. Las huellas de ornitópodos son similares a las utilizadas para definir el icnogénero Iguanodontipus (Sarjeant et al., 1998) del Berriasiense de Dorset (Inglaterra), por lo que se incluyen en el mismo. Huellas semejantes a las descritas en este yacimiento se pueden ver en muchos otros del Grupo Oncala, tanto conocidos como inéditos. El análisis de los posibles icnopoyetas, autores de las huellas, indica que pudieron pertenecer a la familia Camptosauridae (Camptosaurus, Draconyx) o superfamilia Iguanodontoidea, de pequeño tamaño. Las Cuestas I (Soria, Spain) is one of the biggest tracksite in Oncala Group. Until the moment, it has been catalogued almost 600, ornithopod, theropod and mainly sauropod tracks. The ornithopod tracks are similar to those used to define the Iguanodontipus ichnogener (Sarjeant et al., 1998) from the Berriasian in Dorset (England), that is why they are also included.Very similar footprints to those described in this tracksite can be seen a lot of tracksites in Oncala Group, both known and unknown. They analysis of the potential trackmakers the tracks owners, reveals that they could belong to the family of Camptosauridae (Camptosaurus, Draconyx) or superfamily Iguanodontoidea, of small size.

On the tracks of the earliest dinosaurs: implications for the hypothesis of dinosaurian monophyly

From the record of dinosaurian skeletal remains it has been inferred that the origin and initial diversification of dinosaurs were rapid events, occupying an interval of about 5 million years in the Late Triassic. By contrast numerous reports of dinosauroid tracks imply that the emergence of dinosaurs was a more protracted affair extending through much of the Early and Middle Triassic. This study finds no convincing evidence of dinosaur tracks before the late Ladinian. Each of the three dinosaurian clades - Theropoda, Sauropodomorpha, Ornithischia - produced a unique track morphotype that appears to be an independent modification of the chirotherioid pattern attributed to stem-group archosaurs (thecodontian reptiles). The existence of three divergent track morphotypes is consistent with the concept of dinosaurian polyphyly but can be reconciled with the hypothesis of dinosaurian monophyly only by invoking many and rapid reversals in the locomotor anatomy of early dinosaurs. The origin of dinosaurs was not the correlate or consequence of any single event or process, be it global change, competitive replacement, or opportunism in the wake of mass extinction. Instead the origin of dinosaurs is envisaged as a series of three cladogenetic events over an interval of at least 10 million years and possibly as much as 25 million years. This scenario of dinosaurian polyphyly is as well-supported by fossil evidence as is the currently favoured view of dinosaurian monophyly.

Revisiting the Late Jurassic-Early Cretaceous of the NW South Iberian Basin: new ages and sedimentary environments

The study of the Upper Jurassic-Lower Cretaceous deposits (Higueruelas, Villar del Arzobispo and Aldea de Cortés Formations) of the South Iberian Basin (NW Valencia, Spain) reveals new stratigraphic and sedimentological data, which have significant implications on the stratigraphic framework, depositional environments and age of these units. The Higueruelas Fm was deposited in a mid-inner carbonate platform where oncolitic bars migrated by the action of storms and where oncoid production progressively decreased towards the uppermost part of the unit. The overlying Villar del Arzobispo Fm has been traditionally interpreted as an inner platform-lagoon evolving into a tidal-flat. Here it is interpreted as an inner-carbonate platform affected by storms, where oolitic shoals protected a lagoon, which had siliciclastic inputs from the continent. The Aldea de Cortés Fm has been previously interpreted as a lagoon surrounded by tidal-flats and fluvial-deltaic plains. Here it is reinterpreted as a coastal wetland where siliciclastic muddy deposits interacted with shallow fresh to marine water bodies, aeolian dunes and continental siliciclastic inputs. The contact between the Higueruelas and Villar del Arzobispo Fms, classically defined as gradual, is also interpreted here as rapid. More importantly, the contact between the Villar del Arzobispo and Aldea de Cortés Fms, previously considered as unconformable, is here interpreted as gradual. The presence of Alveosepta in the Villar del Arzobispo Fm suggests that at least part of this unit is Kimmeridgian, unlike the previously assigned Late Tithonian-Middle Berriasian age. Consequently, the underlying Higueruelas Fm, previously considered Tithonian, should not be younger than Kimmeridgian. Accordingly, sedimentation of the Aldea de Cortés Fm, previously considered Valangian-Hauterivian, probably started during the Tithonian and it may be considered part of the regressive trend of the Late Jurassic-Early Cretaceous cycle. This is consistent with the dinosaur faunas, typically Jurassic, described in the Villar del Arzobispo and Aldea de Cortés Fms.

Systematic review of Late Jurassic sauropods from the Museu Geológico collections (Lisboa, Portugal)

The Museu Geológico collections house some of the first sauropod references of the Lusitanian Basin Upper Jurassic record, including the Lourinhasaurus alenquerensis and Lusotitan atalaiensis lectotypes, previously considered as new species of the Apatosaurus and Brachiosaurus genera, respectively. Several fragmentary specimens have been classical referred to those taxa, but the most part of these systematic attributions are not supported herein, excluding a caudal vertebra from Maceira (MG 8804) considered as cf. Lusotitan atalaiensis. From the material housed in the Museu Geológico were identified basal eusauropods (indeterminate eusauropods and turiasaurs) and neosauropods (indeterminate neosauropods, diplodods and camarasaurids and basal titanosauriforms). Middle caudal vertebrae with lateral fossae, ventral hollow border by pronounced ventrolateral crests and quadrangular cross-section suggest for the presence of diplodocine diplodocids in north area of the Lusitanian Basin Central Sector during the Late Jurassic. A humerus collected from Praia dos Frades (MG 4976) is attributed to cf. Duriatitan humerocristatus suggesting the presence of shared sauropod forms between the Portugal and United Kingdom during the Late Jurassic. Duriatitan is an indeterminate member of Eusauropoda and the discovery of new material in both territories is necessary to confirm this systematic approach. The studied material is in according with the previous recorded paleobiodiversity for the sauropod clade during the Portuguese Late Jurassic, which includes basal eusauropods (including turiasaurs), diplodocids and macronarians (including camarasaurids and basal titanosauriforms).