Pedro Mocho

A new dryosaurid ornithopod (Dinosauria, Ornithischia) from the Late Jurassic of Portugal

ABSTRACT A new dryosaurid ornithopod, Eousdryosaurus nanohallucis, gen. et sp. nov., is described here based on a single specimen from the Late Jurassic Alcobaça Formation of Portugal. Eousdryosaurus nanohallucis is distinguished from all other dryosaurids by eight autapomorphic features and an unique combination of characters, some of which are also shared by other dryosaurids. Eousdryosaurus is linked with Dryosauridae, because the fourth trochanter is proximally placed and widely separated from the scar for the insertion of the M. caudifemoralis longus, which is restricted to the medial surface of the femoral shaft. Phylogenetic analysis nests Eousdryosaurus in an unresolved polytomy at the base of Dryosauridae together with Callovosaurus, Dryosaurus, and Kangnasaurus. The complete pes of Eousdryosaurus, which has a phalangeal formula of 1-3-4-5-0, supports the putative autapomorphic reduction of the dryosaurid pes that also occurs in parallel in more derived ornithopods.

New data of the Portuguese brachiosaurid Lusotitan atalaiensis (Sobral Formation, Upper Jurassic)

Abstract Lusotitan atalaiensis was one of the first sauropod taxa established for the Upper Jurassic of the Lusitanian Basin (Portugal). The lectotype of L. atalaiensis found in Peralta (Lourinhã) was firstly considered as new species of Brachiosaurus, considering the similarities with Brachiosaurus and Giraffatitan. This specimen was originally considered as a brachiosaurid, and more recently, a cladistics analysis, suggested Lusotitan as a basal macronarian and a possible member of Brachiosauridae. In this study, new information is provided about the Lusotitan lectotype, with reinterpretation and description of previously described and undescribed elements (e.g. sacral vertebrae, chevrons, ulna, ischium, pubis or fibula). The validity of this taxon is confirmed by a revised diagnosis. The present comparative study and the phylogenetic analyses suggest the assignation of Lusotitan to Brachiosauridae supported by the presence of pronounced and dorsoventrally short deltopectoral crest, short ischiatic contribution to the acetabulum and radius length/tibia length ratio >1. The presence of brachiosaurids in the Portuguese Upper Jurassic record is confirmed, supporting the presence of a wide paleobiogeographic distribution for Brachiosauridae during the Late Jurassic, being present in Europe, North America and Africa.

Turiasauria-like teeth from the Upper Jurassic of the Lusitanian Basin, Portugal

Turiasauria is a clade of eusauropods with a wide stratigraphic range that could extend from the Bathonian to the lower Aptian including Turiasaurus, Losillasaurus, Zby and putatively, Galveosaurus, Atlasaurus and isolated remains from Middle Jurassic-to-Lower Cretaceous. Some are characterised by the presence of heart-shaped teeth. Several tooth occurrences from the Portuguese Upper Jurassic with this type of morphology (SI: 1.1–1.8) are reported and discussed. If this morphology is regarded as synapomorphic of Turiasauria, the teeth will be tentatively related to this clade. From a sample of 43 teeth, three main morphotypes are described. Three hypotheses might explain the morphological variation: (1) the range of tooth morphologies indicates variation in the jaw, (2) the range of tooth morphologies indicates taxonomic variation or (3) a combination of both. The general wear pattern in morphotypes I and II starts with a distal facet, then the appearance of mesial/apical facet and finally a ‘V’-shaped facet. In morphotype III, the wear begins with a mesial facet. The variability observed for Portuguese Upper Jurassic specimens is congruent with the morphological variability along the tooth row shown by other sauropods with spatulate/spoon-shaped teeth and it is considered the most parsimonious hypothesis to explain it.

New data on the anatomy of Torvosaurus and other remains of megalosauroid (Dinosauria, Theropoda) from the Upper Jurassic of Portugal

A set of cranial and postcranial specimens, including two partial maxillae, several isolated teeth, vertebrae and appendicular elements of theropod dinosaurs is described. These specimens were collected in different Upper Jurassic sites from the Lusitanian Basin (Portugal) and show several characters that allow its identification as belonging to megalosauroid tetanurans. Some of these elements have a combination of features exclusively shared with other megalosaurids known in the Portuguese record, in particular with the recently described species Torvosaurus gurneyi from synchronic sedimentary levels. The geographic and stratigraphic distribution of the specimens of megalosauroids currently known in the Portuguese record indicates that members of this clade were relatively abundant from the upper Kimmeridgian to the lowermost Tithonian of the Lusitanian Basin, especially in the Consolação Sub-basin. The analysis of the different specimens described allows testing hypotheses about the distribution and variability of some characters among the Portuguese forms related to Torvosaurus.ResumenSe describen varios elementos craneales y postcraneales, incluyendo dos fragmentos maxilares, varios dientes aislados, vértebras y elementos apendiculares de dinosaurios terópodos provenientes de diferentes localidades del Jurásico Superior de la cuenca lusitánica. Este conjunto de evidencias osteológicas presenta una combinación de caracteres que permite identificarlos como pertenecientes a tetanuros megalosauroideos. Algunos de estos elementos presentan características compartidas con Torvosaurus gurneyi, especie recientemente descrita en niveles sedimentarios sincrónicos. La distribución geográfica y estratigráfica de los ejemplares de megalosauroideos conocidos en la actualidad en el registro portugués sugiere que este clado era relativamente abundante en el Kimmeridgiense superior y Tithoniense inferior de la cuenca lusitánica, sobre todo en la subcuenca de Consolação. El análisis de los diferentes ejemplares descritos ha permitido comprobar algunas hipótesis sobre la distribución y variabilidad de algunos caracteres en las formas portuguesas afines a Torvosaurus.

A juvenile allosauroid theropod (Dinosauria, Saurischia) from the Upper Jurassic of Portugal

Abstract A new specimen of a theropod dinosaur found in Upper Jurassic sedimentary levels of the Lusitanian Basin (Portugal) is described. The specimen includes axial (cervical, dorsal, and caudal vertebrae and ribs) and pelvic elements, corresponding to a small-sized and juvenile individual. This specimen is one of the most complete theropod dinosaur from the Upper Jurassic of Portugal, and the only evidence of a post-hatchling juvenile theropod individual currently recognized in this record. The phylogenetic analysis recovered the new specimen as a basal Allosauroidea. It presents a combination of characters shared with other allosauroids already known in the Upper Jurassic of the Lusitanian Basin, Allosaurus and Lourinhanosaurus, but also some differences relative to both taxa. Some of these differences may be related to the juvenile condition of the specimen, but other unusual features cannot be properly explained by ontogeny, and are interpreted as having taxonomic significance. This combination of features might justify the description of a new theropod taxon for the Portuguese Upper Jurassic. Nevertheless, the presence of three sympatric and almost synchronic, closely related basal allosauroids requires further exploration of their intra- or interspecific variability.

New data on Late Jurassic sauropods of central and northern sectors of the Bombarral Sub-basin (Lusitanian Basin, Portugal)

Abstract The Upper Jurassic’s central and northern sectors of the Bombarral Sub-basin are relatively poor in sauropod material, highlighting the specimens (mainly teeth) found in the Guimarota mine (Leiria) and the Andrés (Pombal) fossil site. The study of published and the unpublished sauropod material allows for a revision of the present state of sauropod diversity of the Bombarral Sub-basin. These new specimens come from Pombal, Leiria, Batalha, Porto de Mós, Alcobaça and Caldas da Rainha, and include an almost complete posterior or middle dorsal neural spine and a partial caudal series. The systematic re-evaluation of the sauropod record of this sector indicates the presence of turiasaurs, diplodocines, titanosauriforms and an indeterminate eusauropod form. During the last part of the twentieth century, the discovery of fossil vertebrates has increased significantly in the sediments cropping out in the central and northern sectors of the Bombarral Sub-basin (Alcobaça and Bombarral Formations), improving our understanding of the Late Jurassic faunas of the Lusitanian Basin.

Upper Jurassic sauropod record in the Lusitanian Basin (Portugal): Geographical and lithostratigraphical distribution

Sauropod remains are relatively abundant in the Upper Jurassic sediments of the Lusitanian Basin. These dinosaurs are recorded in several sub-basins formed during the third rifting episode related to the evolution of the Lusitanian Basin. The Kimmeridgian-Tithonian sedimentary sequence is dominated by siliciclastic deposits, indicating a continental environment. Sauropods are present all along this mainly terrestrial sequence, being recorded in the Alcobaça, Praia da Amoreira-Porto Novo, Sobral, Freixial, and the Bombarral Formations, ranging from the early Kimmeridgian to the late Tithonian. Sauropoda is the most abundant dinosaur group in the Upper Jurassic fossil record of the Lusitanian Basin and is especially well-represented in the Bombarral and Turcifal Sub-basins. Several new specimens, so far unpublished, are reported here. The sauropod fauna identified mainly includes non-neosauropod eusauropods (including turiasaurs), diplodocoids (some specimens with diplodocine affinities), basal macronarians (non-camarasaurids and camarasaurids), and titanosauriforms (some specimens with brachiosaurid affinities). Macronarians, turiasaurs and diplodocoids are generally present along the entire Kimmeridgian-Tithonian continental to transitional deposits of the Lusitanian Basin, but the known fossil record for some more exclusive groups such as camarasaurids, brachiosaurids, and diplodocines, present a more restricted stratigraphic distribution. Pedro Mocho. The Dinosaur Institute, Natural History Museum of Los Angeles County, 900 Exposition Blvd., 90007 CA, Los Angeles, USA; Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, c/ Senda del Rey, 9, 28040 Madrid, Spain; Laboratório de Paleontologia e Paleoecologia, Sociedade de História Natural, Polígono Industrial do Alto do Ameal, Pav.H02 e H06, 2565-641, Torres Vedras, Portugal. pmocho@nhm.org Rafael Royo-Torres. Fundación Conjunto Paleontológico de Teruel-Dinópolis/Museo Aragonés de Paleontología, av. Sagunto s/n. E-44002 Teruel, Spain. royo@dinopolis.com Fernando Escaso. Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, c/ Senda del Rey, 9, 28040 Madrid, Spain; Laboratório de Paleontologia e Paleoecologia, Sociedade de História Natural, Polígono Mocho, Pedro, Royo-Torres, Rafael, Escaso, Fernando, Malafaia, Elisabete, de Miguel Chaves, Carlos, Narváez, Iván, Pérez-García, Adán, Pimentel, Nuno, Silva, Bruno C., and Ortega, Francisco. 2017. Upper Jurassic sauropod record in the Lusitanian Basin (Portugal): Geographical and lithostratigraphical distribution. Palaeontologia Electronica 20.2.27A: 1-50 palaeo-electronica.org/content/2017/1856-portuguese-sauropods Copyright:

Analysis of diversity, stratigraphic and geographical distribution of isolated theropod teeth from the Upper Jurassic of the Lusitanian Basin, Portugal

PurposeIsolated theropod teeth are abundant in the Upper Jurassic of the Lusitanian Basin and are an important source to reconstruct the diversity of this group as well as its geographic and stratigraphic distribution. However, reliably identification of isolated teeth is complex, especially for those morphotypes related to poorly represented groups. Herein a set of isolated theropod teeth collected in different sites from the Upper Jurassic of the Lusitanian Basin ranging from the late Kimmeridgian to late Tithonian in age are described and discussed.MethodsThese teeth were grouped in seventeen distinct morphotypes based first on morphology and comparative anatomy. Multivariate statistical analyses were performed in order to assign each morphotype to a certain taxon.ResultsThe current analysis shows the presence of several groups of theropods such as Ceratosaurus, Torvosaurus, and Allosaurus beside morphotypes identified as belonging to indeterminate Megalosauroidea and Allosauroidea and morphotypes tentatively assigned to Tyrannosauroidea, Dromaeosauridae, and Richardoestesia. This faunal composition, namely the presence of a non-megalosaurid megalosauroid possibly related to the piatnitzkysaurid Marshosaurus, indicates a higher diversity of theropods in the Late Jurassic of the Lusitanian Basin than previously known, based on more complete specimens. Results obtained from this analysis partially agree with previous studies of other collections with isolated theropod teeth from the Upper Jurassic of Portugal such as those of the Guimarota coal mine. However, the presence of velociraptorine dromaeosaurids, compsognathids, and troodontids reported from this site could not be confirmed in the sample herein analyzed. This analysis also indicates a great similarity of the theropod faunas from the Late Jurassic of the Lusitanian Basin and other European chronocorrelative localities such as those from Spain and Germany.ResumenObjetivoLos dientes aislados de dinosaurios terópodos son un registro abundante en el Jurásico Superior de la cuenca lusitánica pudiendo llegar ser una importante contribución para comprender la diversidad y la distribución geográfica y estratigráfica de estas faunas. Sin embargo, la identificación de dientes aislados y su asignación a un determinado taxón es compleja, especialmente en el caso de morfotipos relacionados con grupos poco conocidos en el mismo registro. En este estudio se presenta el resultado del análisis de un conjunto de dientes aislados de terópodos procedentes de diferentes localidades de la cuenca lusitánica datadas en el Jurásico Superior, concretamente entre el Kimmeridgiense superior y el Tithoniense superior.MétodosEstos dientes se han agrupado en diecisiete morfotipos a partir del estudio morfológicco y de la comparación anatómica. Se realizaron análisis estadístico multivariante para comprobar la identificación de cada morfotipo.ResultadosEl resultado de este análisis ha revelado una gran diversidad de grupos de terópodos que incluye Ceratosaurus, Torvosaurus y Allosaurus además de morfotipos identificados como pertenecientes a Megalosauroidea indet. y Allosauroidea indet. Además, se han reconocido también algunos morfotipos preliminarmente asignados a Tyrannosauroidea, Dromaeosauridae y Richardoestesia. Esta composición faunística, tal como la presencia de non-megalosauridos megalosauroides posiblemente relacionado al piatnitzkysaurido Marshosaurus, sugiere una mayor diversidad de terópodos de la que se conoce actualmente a partir de ejemplares más completos. Los resultados obtenidos soportan, en parte, algunos estudios previos de otras colecciones con dientes aislados del Jurásico Superior de Portugal, como por ejemplo los de la mina de Guimarota. No obstante, la presencia de terópodos velociraptorinos, compsognathidos y troodontidos, citados en Guimarota, no se ha podido confirmar en la muestra estudiada. Este análisis indica también una grande semejanza de las faunas de terópodos del Jurásico Superior de la cuenca lusitánica y de otras localidades sincrónicas europeas como por ejemplo de España y Alemania.

The westernmost records of extinct large European tortoises: the presence of Titanochelon (Testudinidae) in the Miocene of Portugal

Abstract The information about the European giant fossil tortoises has been greatly increased in recent years, based on the description of new material and the revision of previously published specimens. A genus with a wide paleobiogeographic and stratigraphic distribution, Titanochelon, has recently been described, containing all large testudinids from the European Neogene record. Its type species, Titanochelon bolivari, was described in the Spanish record. The presence of this species outside this country had not been justified. In this paper we present and describe fossil material of large testudinids from several sites in the Lisbon District (Portugal), from lower (Quinta da Barbacena; MN4), middle (Charneca do Lumiar and Quinta da Farinheira; MN5) and upper (Aveiras de Baixo; MN9) Miocene levels. This study allows us to confirm the presence of Titanochelon in the Portuguese record and justify, for the first time, the presence of Titanochelon bolivari outside Spain. Furthermore we are able to discuss the geographical and temporal distribution of the large tortoises in the middle Miocene of Europe, a relevant period for the diversity and evolution of this clade.

Carcharodontosaurian remains (Dinosauria, Theropoda) from the Upper Jurassic of Portugal

Abstract A new specimen of a theropod dinosaur found in the Upper Jurassic (Freixial Formation, late Tithonian) of the Lusitanian Basin is described. It corresponds to a single individual and includes a sequence of articulated caudal vertebrae, an almost complete right pes, and other fragments of the appendicular skeleton. The specimen includes the most complete pes of a theropod dinosaur currently known in the Lusitanian Basin and represents one of the youngest skeletal records of theropod dinosaurs currently known in the Portuguese Upper Jurassic. A systematic analysis of this specimen is performed and it shows a combination of characters that allows us to interpret it as belonging to an allosauroid taxon. Within this clade, the material from Cambelas shares a few features with some carcharodontosaurids, including the presence of a lateral lamina extending along the anterior end of the centrum in the caudal vertebra and of a low vertical crest on the lateral surface of the femoral lesser trochanter. The set of remains described here shares some unusual features with another specimen previously described in the Portuguese fossil record, which also presents some synapomorphies of Carcharodontosauria. However, no autapomorphy or exclusive character combination can be recognized in the specimen here described in order to describe it as a new form. The presence of this specimen suggests a greater diversity in the allosauroid theropod fauna from the Late Jurassic of the Lusitanian Basin than currently known and probably expands the temporal record of Carcharodontosauria up to upper Tithonian of southwestern Europe.