New information on ornithopod dinosaurs from the Late Jurassic of Portugal

Funding Information: We would like to thank the Museu da Lourinhã for providing specimens and facilities, especially: Carla-Alexandra Tomás, Bruno Pereira, Alexandre Audigane, and Carla Abreu. To Jesper Milàn (Geomuseum Faxe, Denmark) and Micael Martinho (ML) who found and prepared some of the specimens. We also thank the Sociedade de Historia Natural (Torres Vedras, Portugal) and in particular Bruno Camilo Silva and Joana Ferreira for providing access and assistance to the holotype of Eousdryosaurus nanohallucis. Cristiano Dal Sasso (Museu di Storia Naturale di Milano, Italy) provided comparison material and William Harcourt-Smith (American Museum of Natural History, New York City, USA) for helping with the morphometric analysis. We also thank all the members of the Lourinhã Paleoteam for their inputs in earlier versions of this manuscript. We thank the editor Stephen Brusatte (School of GeoSciences, The University of Edinburgh, Edinburgh, UK) and the reviewers Tom Hübner (Stiftung Schloss Friedenstein Gotha, Germany) and Peter Galton (University of Bridgeport, Connecticut, USA) for their comments, which greatly improved the quality of this manuscript. A special thanks to Vincent J. Cheng (ML) who proof-read this manuscript. FMR is supported by the Fundação para a Ciência e a Tecnologia, Portugal (Grant SFRH/BD/146230/2019). MMA is supported by the Fundação para a Ciência e a Tecnologia, Portugal (Grant SFRH/BPD/113130/2015). The research here presented has been supported by grants (GeoBioTec-NOVA and UIDB/04035/2020).Ornithopods are one of the most speciose group of herbivorous dinosaurs, rising during the Jurassic and getting extinct at the Cretaceous Paleogene boundary. However, most of the attention has been given to derived forms (hadrosaurids). Herein, cranial and post-cranial ornithopod material from the Upper Jurassic Lourinhã Formation and housed at Museu da Lourinhã is described and discussed. Comparison and phylogenetic analyses has allowed the attribution of the material either to Dryosauridae or to Ankylopollexia. The large-sized taxa conservatively ascribed to Ankylopollexia, resemble more closely Early Cretaceous styracosternans than Late Jurassic taxa. Due to the lack of autapomorphic characters, it was not possible to assign the material to any of the two valid Jurassic ornithopod Portuguese species, Draconyx loureiroi and Eousdryosaurus nanohallucis, although phylogenetic analyses hint a close relationship between the Lourinhã dryosaurid material and E. nanohallucis. Principal Component Analysis plotting limb bones proportions indicates a not fully mature ontogenetic stage for the Portuguese specimens. Comparing the Portuguese ornithopod fauna with the one in Morrison Formation and Kimmeridge Clay Formation, it is remarked the key-role of Portugal to understand biogeographic patterns in the distribution of iguanodontians.publishersversionpublishe

Avian eggshell remains in the human bearing level TD6 of the Gran Dolina site (Early Pleistocene, Atapuerca, Spain)

The site of Gran Dolina (Atapuerca, Spain) has a 19-metre-thick Lower and Middle Pleistocene infilling, divided into eleven levels. This work is focused on the level TD6 (0.8–0.9Ma), consisting of a succession of debris flow and fluvial facies with a high diversity of vertebrates, including Homo antecessor. Here we describe for first time eggshell fragments from Atapuerca, recovered by screen washing of sediments from the excavations. The 22 eggshell fragments recovered are small and poorly preserved, with all material showing signs of abrasion. All eggshells have smooth outer surfaces, densely packed mammillae, and are multi-layered. Thus, they can be identified as avian eggshells. Most fragments were relatively well intact, with no signs of recrystallisation, as evidenced by cathodoluminescence analysis, however the degree of abrasion of the fragments hindered the identification of some ultrastructural characters. Three taxa have been identified: Charadriiformes, Gruidae and Anseriformes (all recorded in the TD6 assemblage by osteological remains). The association supports the presence of water bodies close to the site. The relatively high diversity of ootaxa in the small sample size analysed suggest that a wide range of the nesting species inhabited surrounding areas of Gran Dolina at the moment of accumulation of the TD6 level

Neurocranial bones are key to untangling the sea cow evolutionary tree: osteology of the skull of Sobrarbesiren cardieli (Mammalia: Pan-Sirenia)

The pan-sirenian Bauplan is conservative, probably owing to the constraints of adaptation to an aquatic lifestyle. Gathering morphological data from extinct forms is complex, resulting in poorly resolved phylogenies for stem pan-sirenians. Extant sirenians ossify the falx cerebri and the tentorium cerebelli, membranes of the dura mater of the brain attached to the parietal bone. Nevertheless, these ossifications are not present in some pan-sirenians. The basioccipital bone has received little attention in the literature except for establishing the relative age of individuals. Here, we present new cranial elements and a detailed description of the skull of Sobrarbesiren cardieli, a stem pan-sirenian from the Lutetian of Spain represented by eight individuals; we study its intraspecific variation and palaeoecological implications and explore the evolution of the endocranial structures and the basioccipital bone in pan-sirenians. Six new phylogenetic characters are added to the latest pan-sirenian dataset, resulting in a well-resolved topology where Sobrarbesiren is recovered close to the root, in a clade with Prototherium and Eotheroides aegyptiacum. The basioccipital bone and the ossified endocranial membranes have a phylogenetic signal, and the absence of such endocranial structures represents the plesiomorphic condition for pan-sirenians and is not diagnostic for the family Protosirenidae as previously believed

Plasticity in the morphology of the fused frontals of Albanerpetontidae (Lissamphibia; Allocaudata)

Albanerpetontidae form an enigmatic extinct group of lissamphibians, ranging from the early Bathonian to the early Pleistocene. The Upper Jurassic outcrops of Portugal yield a large collection of material, suitable for addressing the intraspecific variation in and diagnostic potential of the characteristic fused frontals. We revise 58 specimens from the Guimarota beds of the Kimmeridgian Alcobaca Formation and describe 62 new frontal bones from the Kimmeridgian - Tithonian Lourinha Formation. Smaller specimens exhibit a vermicular dorsal ornamentation, while it is polygonal in larger specimens and other albanerpetontids. Compared to small specimens, larger specimens display: (1) larger ventrolateral crests extending posteriorly after the parietal margin; (2) a relatively shorter internasal process; (3) a frontal width across posterior edges relatively smaller; and (4) a ventromedian crest less pronounced. Morphometric analyses suggest a single species with different ontogenetic stages. Specimens are attributed to aff. Celtedens sp., based on a bell-shaped outline with a curved orbital margin (although variable in Portuguese specimens), and a flabellate, bulbous-shaped internasal process. The species is more similar to C. megacephalus than C. ibericus, but its phylogenetic position comprises an unresolved trichotomy. Our results show that intraspecific variation and homoplasy render the fused frontal non-diagnostic below the generic level

New contributions to the phylogenetic position of the sauropod galvesaurus herreroi from the late kimmeridgian-early tithonian (jurassic) of teruel (spain)

Galvesaurus herreroiis a sauropod from the Villar del Arzobispo Formation (late Kimmeridgian-earlyTithonian), from the municipality of Galve (Teruel). Its phylogenetic relations have been long debated, so wecarried out a phylogenetic analysis, using a new data matrix recently published by Carballido et al.(2017). Thecharacters of Galvesauruswere coded on the basis of the redescription of the published remains and thedescription of two unpublished fossils: a right coracoid and a fragment of the right pubis. The results of theanalysis suggest the inclusion of Galvesaurusin the clade Titanosauriformes, as a sister taxon to Lusotitan,these two taxa form part of the Brachiosauridae clade. Likewise, a stratigraphic study was undertaken, plac-ing the Galvesaurussite in the lower part of the Villar del Arzobispo Formation, thus assigning the sauropoda late Kimmeridgian-early Tithonian age. Galvesaurus herreroies un saurópodo recuperado en la Formación Villar del Arzobispo (Kimmeridgiensesuperior-Titoniense inferior), del término municipal de Galve (Teruel). Sus relaciones filogenéticas han sidolargamente discutidas, para dar solución a este problema se ha llevado a cabo un análisis filogenético, connuevos datos y usando una moderna propuesta filogenética (Carballido et al., 2017). Los caracteres deGalvesaurusse han codificado basándose en la redescripción de los restos publicados y en la descripción dedos fósiles inéditos: el coracoides derecho y un fragmento del pubis derecho. Los resultados del análisis indu-cen a proponer la inclusión de Galvesaurusen el clado Titanosauriformes, como taxón hermano de Lusotitan,y siendo incluidos ambos dentro de Brachiosauridae. Así mismo, se ha realizado un estudio estratigráfico quesitúa el yacimiento de Galvesaurusen la parte baja de la Formación Villar del Arzobispo, asignándole al sau-rópodo una edad Kimmeridgiense superior-Titoniense inferior

Dryosaurid ornithopods from the Late Jurassic of Portugal: an overview

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PALEOECOLOGÍA Y SEDIMENTOLOGÍA DE UN YACIMIENTO DE LA SECUENCIA SUPERIOR DE LA FORMACIÓN BLESA (SUB-CUENCA DE OLIETE, BARREMIENSE, CRETÁCICO INFERIOR) EN ESTERCUEL (TERUEL)

La Formación Blesa presente en el Barremiense de la subcuenca de Oliete, que se encuentra en la parte oriental de la Cuenca del Maestrazgo (Teruel), se ha descrito como una formación geológica de espesor variable a lo largo de la subcuenca y que está formada por tres secuencias sedimentarias en las que se observa sedimentación de diferentes facies en distintos ambientes, tanto continental como con influencia marina (Aurell et al., 2018).En ella se ha estudiado el yacimiento Barranco del Hocino 3 realizándose un perfil estratigráfico y un estudio micropaleontológico en el que se han caracterizado taxones de dinosaurios, de crocodilomorfos y de peces del barremiense. <br /

Testing the usefulness of the surface collecting method in a vertebrate microfossil site from the barremian of Spain (Los Menires, Mirambel formation)

Surface collecting is the first and sometimes the only type of sampling carried out in many fossiliferous localities, including vertebrate microfossil assemblages. Nevertheless, it is rare to test how representative these surface-collected samples are of the palaeobiocoenosis. A first approach to the palaeontological analysis of Los Menires, a Barremian vertebrate microfossil assemblage, is here performed while testing the usefulness of the surface collection method. New fossil material obtained by surface collection is described, and the resulting data are compared with those obtained by screen-washing. The fossil assemblage of Los Menires is dominated by parautochthonous remains of freshwater aquatic and semiaquatic organisms—i.e., charophytes, ostracods, bivalves, gastropods, testudinatans, crocodylomorphs, lissamphibians, and osteichthyans—although parautochthonous remains of terrestrial vertebrates—i.e., dinosaurs, lacertilians, and mammaliforms—are also present. The accumulation of vertebrate hard parts in Los Menires took place in a low-energy, shallow-water, depositional environment within the alluvial-lacustrine system represented by the Mirambel Formation. Sampling test results indicate that surface collection is effective in recognizing the main fossil groups present in an assemblage. Yet, it is not suitable for capturing delicate and tiny fossils nor for recognizing the abundance of eggshells. In contrast, it can generate an overrepresentation of other hard components such as coprolites or ornithopod teeth