Search

The Upper Jurassic Lourinhã Formation is well known for its rich assemblage of fossil vertebrates. In this formation, ornithopod dinosaurs are represented by two iguanodontian species, Eousdryosaurus nanohallucis and Draconyx loureiroi. We recently became aware of unreported material belonging to the holotype of Draconyx loureiroi, consisting of partially articulated manual elements. We here re-describe the holotype specimen ML 357, including the newly discovered material. The specimen was subjected to CT-scanning and its surface data used to assess anatomical characters. Linear measurements of metatarsal III were used to estimate the body length of the specimen. The Draconyx loureiroi holotype was included in two datasets and analysed with maximum parsimony and Bayesian inference approaches to estimate evolutionary rates among Iguanodontia. We present evidence that Draconyx loureiroi is a valid taxon nested in Styracosterna and is clearly diagnosable by a unique combination of characters. Both maximum parsimony and Bayesian inference indicate high evolutionary rates across the Jurassic/Cretaceous transition for the base of Iguanodontia. Length estimation suggests that Draconyx loureiroi was a relatively small, bipedal and possibly cursorial animal. Given its basal phyletic position, we interpret this bauplan was the ancestral condition for Styracosterna, that only later in the Cretaceous evolved into giant quadrupedal forms.

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). Made available in DSpace on 2021-11-11T23:38:28Z (GMT). No. of bitstreams: 0 Previous issue date: 2020 publishersversion published

Thin fossil eggshell from Upper Cretaceous deposits of Europe, characterized by nodular ornamentation similar to modern gekkotan eggshell, has mostly been interpreted as gekkotan (=‘geckoid’) in origin. However, in some cases, as for the oogenus Pseudogeckoolithus, a theropod affinity has also been suggested. The true affinity of these fossil ‘geckoid’ eggshells has remained controversial due to the absence of analytical methods for identifying genuine gecko eggshell in the fossil record. In this study, we apply electron backscatter diffraction (EBSD) analysis to latest Cretaceous European ‘geckoid’ (including Pseudogeckoolithus) eggshell, in comparison with modern gekkotan and theropod (avian) eggshell. We found that Pseudogeckoolithus has a definite theropod eggshell-like crystallographic configuration, in clear contrast to that seen in modern ‘gecko eggshells’. Furthermore, the crystallography of the nodular ornamentation in Pseudogeckoolithus is similar to that seen in megapode eggshell, but different from that of gecko eggshell, despite superficial morphological similarity. The remarkable morphological similarities between Pseudogeckoolithus and modern gecko eggshells are thus convergent, and the ‘gekkotan affinity’ hypothesis can be dismissed for Pseudogeckoolithus. This study provides a template for differentiating true gekkotan from dinosaurian eggshells in the fossil record. The potential functional significance of eggshell ornamentation, lost in most modern birds, requires further study, and experimental zoological approach may shed light on this issue. Finally, the present results suggest caution about the dangers of using potentially homoplastic eggshell characters in eggshell parataxonomy.

Altres ajuts: CERCA Programme/Generalitat de Catalunya Altres ajuts: Fossil preparation was supported by the Servei de Museus-Departament de Cultura of the Generalitat de Catalunya (grants 2015/104328, CLT005/16/00008, CLT005/19/00045) and the Institut d'Estudis Ilerdencs (grant 201602412). The origin of the last sauropod dinosaur communities in Europe and their evolution during the final 15 million years of the Cretaceous have become a complex phylogenetic and palaeobiogeographic puzzle characterized by the controversy on the alleged coexistence of immigrant, Gondwana-related taxa alongside relictual and insular clades. In this context, we describe a new titanosaurian sauropod dinosaur, Abditosaurus kuehnei gen. et sp. nov., from the Late Cretaceous (Maastrichtian) Tremp Group of Catalonia (Spain). Phylogenetic analyses recover Abditosaurus separately from other European titanosaurs, within a clade of otherwise South American and African saltasaurines. The affinity of the new taxon with southern landmasses is reinforced by spatiotemporal co-occurrence with Gondwanan titanosaurian oospecies in southern Europe. The large size and the lack of osteohistological features potentially related to insular dwarfism or size reduction support the idea that Abditosaurus belongs to an immigrant lineage, unequivocally distinct from some of the island dwarfs of the European archipelago. The arrival of the Abditosaurus lineage to the Ibero-Armorican Island is hypothesized to have occurred during the earliest Maastrichtian (70.6 Ma), probably as a result of a global and regional sea-level drop that reactivated ancient dispersal routes between Africa and Europe. The arrival of large-bodied titanosaurs to the European archipelago produced dramatic changes in its insular ecosystems and important evolutionary changes in its dinosaur faunas, especially with respect to the 'island rule' effect.

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.

The authors have requested that this preprint be removed from Research Square.

Found in the Upper Jurassic outcrops of Lourinha, Portugal, and first published in 1997, the Paimogo dinosaur egg clutch is one of Portugal’s most remarkable fossils, with over one hundred eggs preserved in association with embryonic bones, of the allosauroid theropod Lourinhanosaurus. However, many questions about it have remained unanswered, even until the present day. After its discovery, this extraordinary fossil became the keystone of a small local museum, greatly kick-starting regional tourism, while also holding the fossils in trust for future generations to study. More than 20 years later, continually sustained paleontological interest from the public has even given rise to both a highly successful dinosaur theme park in the region and an aspiring UNESCO Geopark. Recently, a multidisciplinary team of preparators, paleontologists, sedimentologists, mineralogists, and geochemists revisited an unopened jacket from the original excavation using an array of techniques to address various questions. Studies are ongoing, but the corpus of information obtained and the methodologies utilized to gather data have offered an opportunity to design an exhibit around the history of the Paimogo clutch, highlighting the scientific methods involved, and asserting the importance of preserving geological heritage for the future, when new tools will doubtlessly become available to provide yet another new look at old fossils. Here, we describe our analytical procedures and present an innovative exhibit designed to introduce to the public the latest advances on the research behind an iconic piece of Portuguese geoheritage, increasing its value both as a research item and as an educational resource.

Crocodylomorphs were a diverse clade in the Late Jurassic of Portugal, with six taxa reported to date. Here we describe 126 isolated teeth recovered by screen-washing of sediments from Valmitão (Lourinhã, Portugal, late Kimmeridgian–Tithonian), a vertebrate microfossil assemblage in which at least five distinct crocodylomorph taxa are represented. Ten morphotypes are described and attributed to five clades (Lusitanisuchus, Atoposauridae, Goniopholididae, Bernissartiidae and an undetermined mesoeucrocodylian). Four different ecomorphotypes are here proposed according to ecological niches and feeding behaviours: these correspond to a diet based on arthropods and small vertebrates (Lusitanisuchus and Atoposauridae), a generalist diet (Goniopholididae), a durophagous diet (Bernissartiidae) and a carnivorous diet. Lusitanisuchus mitracostatus material from Guimarota is here redescribed to achieve a better illustration and comparison with the new material. This assemblage shares similar ecomorphotypes with other Mesozoic west-central European localities, where a diversity of crocodylomorphs lived together, avoiding direct ecological competition through niche partitioning. The absence of large marine crocodylomorphs, present in other contemporaneous assemblages, is here interpreted as evidence that the Valmitão assemblage was deposited in a freshwater environment, although sample bias cannot be completely ruled out. These affinities are further supported by the presence of lanceolate and leaf-shaped teeth associated with continental clades.

Throughout the evolutionary history of Avialae, several members of this clade have evolved into giant forms, in different time periods and ecological contexts. In Europe, the first birds that show ...

The South-Pyrenean Basin (northeastern Spain) has yielded a rich and diverse record of Upper Cretaceous (uppermost Campanian−uppermost Maastrichtian) vertebrate fossils, including the remains of some of the last European dinosaurs prior to the Cretaceous-Paleogene (K-Pg) extinction event. In this work, we update and characterize the vertebrate fossil record of the Arén Sandstone and Tremp formations in the Western Tremp Syncline, which is located in the Aragonese area of the Southern Pyrenees. The transitional and continental successions of these sedimentary units are dated to the late Maastrichtian, and exploration of their outcrops has led to the discovery of numerous fossil remains (bones, eggshells, and tracks) of dinosaurs, including hadrosauroids, sauropods, and theropods, along with other tetrapods such as crocodylomorphs, testudines, pterosaurs, squamates, and amphibians. In particular, this fossil record contains some of the youngest lambeosaurine hadrosaurids (Arenysaurus and Blasisaurus) and Mesozoic crocodylomorphs (Arenysuchus and Agaresuchus subjuniperus) in Europe, complementing the lower Maastrichtian fossil sites of the Eastern Tremp Syncline. In addition, faunal comparison with the fossil record of Hațeg island reveals the great change in the dinosaur assemblages resulting from the arrival of lambeosaurine hadrosaurids on the Ibero-Armorican island, whereas those on Haţeg remained stable. In the light of its paleontological richness, its stratigraphic continuity, and its calibration within the last few hundred thousand years of the Cretaceous, the Western Tremp Syncline is one of the best places in Europe to study the latest vertebrate assemblages of the European Archipelago before the end-Cretaceous mass extinction.

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.

The Sobrarbe-Pirineos UNESCO Global Geopark, located in the Central Pyrenees, is a region of remarkable geodiversity that includes extensive Eocene fossil-bearing sites and constitutes an important archive of paleobiodiversity. The Sobrarbe-Pirineos Geopark hosts outcrops of Eocene formations bearing an unusual abundance and diversity of fossils from marine and continental sedimentary environments, making the Sobrarbe-Pirineos Geopark a perfect window for learning about tropical ecosystems of the Eocene of southern Europe. These environments were in part tectonically controlled and offer a unique opportunity to understand how faunas changed in an active area. Here, we outline the main groups of fossils from the Sobrarbe-Pirineos Geopark, including popular examples such as the “Crocodile of Ordesa-Vio” and the sirenian Sobrarbesiren. The Geopark has been a major tool in the geoconservation of Eocene fossils.

A review of the onset of the synrift sedimentation and synsedimentary extensional tectonics of the Oliete sub-basin (northwestern Maestrazgo basin, East Spain) is presented here based on new data acquired after extensive sedimentological, structural and palaeontological analysis of the Barremian Blesa Fm. The lower boundary of the Blesa Fm is a prominent basal synrift unconformity overlying Jurassic units. This formation has been divided into three genetic stratigraphic sequences bounded by sub-basin-wide unconformities. The lower Blesa sequence (LBS) is characterized by distal alluvial to palustrine marls/clays grading upward to palustrine–lacustrine limestones. The LBS is bounded on top by a planar to irregular transgressive, hardened ferruginous surface, locally encrusted by oysters. Above this discontinuity, the middle Blesa sequence consists of oyster-rich limestones and marls deposited in a shallow restricted bay, which grade to distal alluvial and palustrine–lacustrine marls/clays and limestones towards the marginal areas of the basin. The boundary between the middle and upper Blesa sequence (UBS) is a regressive surface outlined by the local presence of an erosive conglomeratic bed. The UBS generally starts with red clays deposited in distal alluvial fan environments, which grade upwards to palustrine and lacustrine carbonates and marls/clays. The local presence of heterolithic alternations of clay with fine-to-medium sandstone and of cross-bedded sandstones indicates the local occurrence of siliciclastic coastal environments in the UBS. The upper boundary of the Blesa Fm is marked by widespread transgression, giving rise to the bioclastic limestones of the Alacon Fm. In the present paper, the stratigraphic position and palaeoenvironmental context of the abundant vertebrate remains found across the defined sequences within the Blesa Formation is reviewed. The results obtained are relevant for a further understanding of the tectosedimentary evolution of the studied basin. Successive stages of evolution are distinguished, including the initial uplift, breakup and erosion of the earlier Jurassic carbonate platform that took place during the Tithonian–Hauterivian; the onset of synrift sedimentation during the early Barremian, which was highly controlled by extensional faulting and differential block subsidence; the homogenization of the basin subsidence accompanied by the incursion of marine waters (sourced from southeastern areas) during the middle part of the Barremian; and the significant fall in base level, of possible climatic origin and also involving significant siliciclastic input in the northern areas of the Oliete sub-basin around the middle part of the late Barremian.

The Eocene record of turtle eggshells is scarce, with a single unconfirmed report from France. This scarcity contrasts with the great abundance of osteological remains, distributed over a wide palaeogeographical range as a result of climatic warmth. In this paper, we describe the first definitive Eocene Testudoolithidae eggshell fragments attributable to Testudines, most likely pleurodiran turtles, from the Eocene Sobrarbe Formation at the Castejon de Sobrarbe fossil site (CS-41) in northern Spain. The eggshells were found in association with osteological remains of at least four individuals of Eocenochelus eremberti (Pleurodira, Erymnochelyini) in an otherwise sirenian-dominated bonebed. Analysis of eggshell ultra- and microstructure allows comparison with eggshells from fossil and extant turtles. The eggshells are highly recrystallized but preserve relics of their original aragonitic radial ultrastructure as crystal phantoms. The barrel-shaped shell units, which are taller than wide, with compactituberculate ornamentation and funnel-shaped pore openings are similar to those of the Palaeocene ootaxon Haininchelys curiosa, whose holotype and paratype have been lost. The CS-41 eggshells are most similar to those of the pleurodiran Erymnochelys madagascariensis from Madagascar, the closest living relative to Eocenochelus, further supporting our attribution. Sedimentological and taphonomic analysis of the assemblage supports the hypothesis that the CS-41 fossil site was formed as the infilling of an abandoned tributary channel in the deltaic plain by an overbank or debris-flow deposit associated with a storm event. This storm resulted in the exhumation and subsequent re-sedimetation of the remains of a nesting ground of Eocenochelus eremberti, an otherwise coastal turtle, which would have entered inland streams to nest in the sand bars at the mouth of the Sobrarbe Deltaic Complex.

The Spheroolithidae oospecies Guegoolithus turolensis, putatively attributed to non-hadrosauroid styracosterns was first described in the Barremian of the Iberian Basin, and later reported in the Valanginian–Hauterivian of the Cameros Basin, with both occurrences separated by a few hundred kilometres but by over 10 million years. Here we apply electron backscatter diffraction to search for crystallographic and ultrastructural differences between both occurrences of this ootaxon. Not only did not we found significant differences between eggshells from the two basins, probably laid by different non-hadrosauroid styracostern species but also, we found significant similitudes with the Late Cretaceous Maiasaura eggshells from North America. All Spheroolithidae eggshells are characterized by a profusion of low angle misorientation boundaries within larger orientation domains, that fan out from the eisospherite. A very quick reorientation of the c-axis of the crystals is observed, and explained partially via competitive growth. This conservative eggshell architecture suggests that the physical properties of the eggshell of styracosterns remain unchanged for at least 80 million years, implying similar nesting and breeding behaviours.

espanolGalvesaurus herreroi es un sauropodo recuperado en la Formacion Villar del Arzobispo (Kimmeridgiense superior-Titoniense inferior), del termino municipal de Galve (Teruel). Sus relaciones filogeneticas han sido largamente discutidas, para dar solucion a este problema se ha llevado a cabo un analisis filogenetico, con nuevos datos y usando una moderna propuesta filogenetica (Carballido et al., 2017). Los caracteres de Galvesaurus se han codificado basandose en la redescripcion de los restos publicados y en la descripcion de dos fosiles ineditos: el coracoides derecho y un fragmento del pubis derecho. Los resultados del analisis inducen a proponer la inclusion de Galvesaurus en el clado Titanosauriformes, como taxon hermano de Lusotitan, y siendo incluidos ambos dentro de Brachiosauridae. Asi mismo, se ha realizado un estudio estratigrafico que situa el yacimiento de Galvesaurus en la parte baja de la Formacion Villar del Arzobispo, asignandole al sauropodo una edad Kimmeridgiense superior-Titoniense inferior. EnglishGalvesaurus herreroi is a sauropod from the Villar del Arzobispo Formation (late Kimmeridgian-early Tithonian), from the municipality of Galve (Teruel). Its phylogenetic relations have been long debated, so we carried out a phylogenetic analysis, using a new data matrix recently published by Carballido et al. (2017). The characters of Galvesaurus were coded on the basis of the redescription of the published remains and the description of two unpublished fossils: a right coracoid and a fragment of the right pubis. The results of the analysis suggest the inclusion of Galvesaurus in the clade Titanosauriformes, as a sister taxon to Lusotitan, these two taxa form part of the Brachiosauridae clade. Likewise, a stratigraphic study was undertaken, placing the Galvesaurus site in the lower part of the Villar del Arzobispo Formation, thus assigning the sauropod a late Kimmeridgian-early Tithonian age.

Fil: Moreno Azanza, Miguel. Universidade Nova de Lisboa; Portugal. Fil: Moreno Azanza, Miguel. Universidad de Zaragoza; España Fil: Gasca Pérez, José M. Universidad de Zaragoza; España Fil: Gasca Pérez, José M. Provincia de Neuquén. Ministerio de Energía, Ambiente y Servicios Públicos. Dirección Provincial de Minería. Museo Provincial de Ciencias Naturales Prof. "Dr. Juan A. Olsacher"; Argentina Fil: Díaz Martínez, Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigación en Paleobiología y Geología; Argentina Fil: Díaz Martínez, Ignacio. Universidad Nacional de Río Negro; Argentina Fil: Bauluz Lázaro, Blanca. Universidad de Zaragoza; España Fil: Canudo, José I. Universidad de Zaragoza; España Fil: Fernández, Arturo. Asociación de Amigos del Museo de Enciso; España Fil: Pérez Lorente, Félix. Universidad de La Rioja; España Here we describe the new fossil site of El Horcajo in the vicinity of Trevijano (La Rioja, Spain). It is located in the palustrine facies of the Enciso Group (Cameros Basin). This new locality has provided dozens of eggshell fragments, together with other vertebrate remains and charophyte fructifications, which allow dating of the locality as Valanginian-Hauterivian. Five ootaxa have being recognized: 1) the Spheroolithidae Guegoolithus turolensis; 2) a surprisingly thick Prismatoolithidae indet., with certain affi nities to the oogenus Sankofa, that may represent a new oogenus and oospecies for this oofamily; 3) the Krokolithidae Krokolithes sp.; 4) recrystallized ?Testudoolithidae eggshells; and 5) ?Geckoolithidae eggshells of uncertain affi nity. This oodiversity is similar to that of other eggshell microsites. The ootaxonomic list differs from the other Iberian locality of the same age, Pochancalo 1 (Villanueva de Huerva Formation, Zaragoza) in lacking sauropod and megalosauroid theropod eggshells, but shares the presence the ornithopod and coelurosaurian theropod eggshells with other younger microsites found in similar facies. The age of the new locality has immediate consequences for the tectono-sedimentary framework of the Cameros Basin, and the new data will need to be considered in future stratigraphical works. Furthermore, the reliability of eggshells as biostratigraphic markers is questioned, and the viability of Guegoolithus as a guide fossil for the lower Barremian is rejected. En este trabajo se describe el nuevo yacimiento fosilífero de El Horcajo en las cercanías de Trevijano (La Rioja, España). Se encuentra en facies palustres dentro del Grupo Enciso (Cuenca de Cameros). Este nuevo yacimiento ha proporcionado docenas de fragmentos de cáscara de huevo, junto con otros restos fósiles de vertebrados y fructifi caciones de carofi tas, que permiten datar el yacimiento como ValanginienseHauteriviense. Se han identificado cinco ootaxones: 1) Guegoolithus turolensis, de la oofamilia Spheroolithidae; 2) un Prismatoolithidae indet. sorprendentemente grueso y con afi nidades al oogénero Sankofa, que podría corresponder a un nuevo oogénero y ooespecie; 3) Krokolithes sp. de la oofamilia Krokolithidae; 4) cáscaras recristalizadas asignables con dudas a Testudoolithidae; y 5) cáscaras de afi nidad incierta, aquí asignadas con dudas a Geckoolithidae. Esta oodiversidad es similar a la de otros microyacimientos de cáscaras de huevo. La lista ootaxonómica difi ere de la obtenida en la otra única localidad de esta edad en Iberia, Pochancalo 1 (Formación Villanueva de Huerva, Zaragoza), por la ausencia de cáscaras de huevo asignables a saurópodos y terópodos megalosauroideos, pero comparte la presencia de cáscaras de huevo asignables a ornitópodos y terópodos celurosaurios con otros yacimientos más recientes que muestran facies similares. Este nuevo yacimiento tiene consecuencias inmediatas para el marco tectono-sedimentario de la cuenca de Cameros, siendo necesario considerar estos nuevos datos en futuros estudios estratigráfi cos. Además, se cuestiona la fi abilidad del uso de cáscaras de huevo como marcadores bioestratigráfi cos, ya que los nuevos datos ponen en duda la utilidad de Guegoolithus como fósil guía para el Barremiense inferior.

An enlarged theropod manual ungual (CSC1-4) from the Weald facies of Spain is described. The claw was found in the fossil locality of Cana Seca 1, Teruel province, within the El Castellar Formation of early Barremian (Early Cretaceous) in age. CSC1-4 is morphologically closer to megalosauroids than to any other theropod clade bearing enlarged manual claws and shows the greatest similarity to the manual ungual of digit I of Baryonyx walkeri. Both CS1-4 and this taxon share a particularly enlarged, elongated and transversely wide manual claw. CSC1-4, however, differs from Baryonyx's ungual in having less curvature, a straight dorsal edge in the proximal part, slightly more width above the grooves than below, and a certain asymmetry, with the lateral face more flattened. Taking into account the palaeogeographic and temporal context, these considerations suggest that they are closely related but distinct spinosaurid taxa. The presence of an enlarged manual claw in spinosaurids has been invoked as an anatomical feature typically associated with scavenging and hunting habits, as well as digging behaviour. The spinosaurid record from the Barremian of the Iberian Peninsula shows that members of this clade favored freshwater environments with some marine influence in this part of Europe.

Crocodiloid eggshell is considered to be one of the most conservative among amniotes. This contrasts with the high body diversity observed within the crocodylomorph lineage, which extebds from the Triassic to the present. This incongruence raises a fundamental question in palaeology: is the crocodylomorph eggshell structure that conservative, or are there variations in this structure that have been misidentified in the fossil record or remain undiscovered to taphonomic biases? In this paper we re-examine eggshells from the Barremian of northern Spain that were previously assigned to chelonians. We erect a new oogenus and oospecies, Mycomorphoolithus kohringi , characterized by thin eggshells with mushroom-shaped or inverted cone shell units with blocky extinction with smooth or slightly undulating outer surface, covered by a highly variable number of pores of irregular size and shape. These variations in the pore opening pattern are here interpreted as evidence of degradation of the eggshell during embryo development, a process that has only been described in modern alligatorids. After discarding its chelonian and dinosaurian affinities, we identify them as related to Krokolithidae, but with enough differences to justify exclusion from this oofamily. In addition, eggshells from the Berriasian of England previously reported as dinosaurian-spherulitic eggshells, are here assigned to undetermined oospecies of Mycomorphoolithus . Thus, the record of Mycomorphoolithus extends throughout most of the Lower Cretaceous. This long-surviving oogenus may represent eggshells of the non-eusuchian crocodylomorphs that are abundant in the microfossil sites where Mycomorphoolithus eggshells are found.

Sirenians are the only extant herbivorous mammals fully adapted to an aquatic lifestyle. They originated in Africa during the Paleocene from an undetermined clade of afrotherian mammals, and by the end of the Eocene they were widely distributed across the tropical latitudes. Here we introduce Sobrarbesiren cardieli gen. et sp. nov. It is the first adequately-known quadrupedal sirenian from Eurasia and the oldest record of this clade from western Europe. Fossils have been recovered from the middle Lutetian (SBZ15) site of Castejón de Sobrarbe-41 (Huesca, Spain), and comprise many cranial and postcranial remains, including pelvic girdle and hind limb bones, from at least six sirenian individuals of different ontogenetic stages. Sobrarbesiren shows a suite of characters previously considered synapomorphies of different clades of derived sirenians, such as the presence of the processus retroversus of the squamosal and the pterygoid fossa, combined with ancestral characters such as the presence of an alisphenoid canal, a permanent P5, at least two sacral vertebrae, a primitive pelvis and functional femora and fibulae. Sobrarbesiren is recovered as the sister taxon of Dugongidae and represents a transitional stage of adaptation to aquatic life between the amphibious quadrupedal prorastomids and the aquatic quadrupedal protosirenids.

The Barremian Mirambel Formation (Maestrazgo Basin, Iberian Chain, NE Spain) preserves different types of dinosaur and other vertebrate fossils (skeletal, eggshell and ichnological remains). A total of 31 vertebrate fossil sites and tracksites have been recognized within this unit in the Ladruñán area (Teruel province). Detailed stratigraphic, sedimentological and micropalaeontological analyses have also been performed in the unit. A vertical sedimentary trend from alluvial-dominated facies (meandering river and related overbank areas) to palustrine-lacustrine facies and back has been defined for the Mirambel Formation in this area. The depositional system was located close to the coastline, as indicated by sporadic marine input in the lower part of the unit. Most fossil remains were recovered by surface collection as well as by the usual techniques used for macrovertebrate excavations. The dinosaur record identified comprises ornithopods, theropods and sauropods. Four distinct track-bearing horizons have been identified. The heterolithic nature and aggradation characteristic of the Mirambel Formation are favourable factors for track formation and preservation. The dinosaur tracks consist of convex hyporeliefs or concave epireliefs that record the trackmakers as they frequented lakeshores, alluvial floodplains and fluvial courses. Macrovertebrate bonebeds occur in alluvial settings (poorly-drained floodplains and “ponds”). Microvertebrate concentrations are located in shallow lacustrine deposits. Isolated skeletal elements can be found in a great variety of deposits. Attritional accumulation in a low-energy depositional context is the general pattern of origin for the bone-bearing fossil sites of the Mirambel Formation. As regards the genetic framework, the resulting skeletal assemblages are predominantly the result of physical factors, with sedimentology as a key factor, rather than biological phenomena. Eggshell fragments are frequent throughout the unit but are clearly more common in palustrine-lacustrine deposits. These can be taken to be parautochthonous bioclasts from nearby areas and might be indicative of the preferential affinity of the egg-layers for wetlands and lakeshores. Fil: Gasca Pérez, José Manuel. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad de Zaragoza; España. Provincia de Neuquén. Ministerio de Energía, Ambiente y Servicios Públicos. Dirección Provincial de Minería. Museo Provincial de Ciencias Naturales Prof. "Dr. Juan A. Olsacher"; Argentina Fil: Moreno Azanza, Miguel. Universidad de Zaragoza; España. Universidade Nova de Lisboa; Portugal Fil: Bádenas, Beatriz. Universidad de Zaragoza; España Fil: Díaz Martínez, Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigación en Paleobiología y Geología; Argentina Fil: Castanera, Diego. Universidad de Zaragoza; España. Ludwig-Maximilians-Universität; Alemania Fil: Canudo, José I.. Universidad de Zaragoza; España Fil: Aurell, Marcos. Universidad de Zaragoza; España

We describe a new species of an ornithischian dinosaur, Isaberrysaura mollensis gen. et sp. nov. The specimen, consisting in an almost complete skull and incomplete postcranium was collected from the marine-deltaic deposits of the Los Molles Formation (Toarcian-Bajocian), being the first reported dinosaur for this unit, one of the oldest from Neuquén Basin, and the first neornithischian dinosaur known from the Jurassic of South America. Despite showing a general stegosaurian appearance, the extensive phylogenetic analysis carried out depicts Isaberrysaura mollensis gen. et sp. nov. as a basal ornithopod, suggesting that both Thyreophora and neornithischians could have achieved significant convergent features. The specimen was preserved articulated and with some of its gut content place in the middle-posterior part of the thoracic cavity. Such stomach content was identified as seeds, most of them belonging to the Cycadales group. This finding reveals a possible and unexpected role of this ornithischian species as seed-dispersal agent., Facultad de Ciencias Naturales y Museo

Altered iguanodontian dinosaur teeth have been studied to analyse the unusual replacement of biogenic apatite and to deduce how the alterations affect the preservation of these fossils. The fossils were recovered from alluvial lacustrine facies (Barremian, Early Cretaceous) in Teruel (Spain). Mandibular teeth and sediments were studied in hand sample, by X-ray diffraction (XRD) and by scanning electron microscopy (SEM). The combination of the different techniques shows that the teeth have undergone at least three pseudomorphic stages: (1) replacement of the organic component of the dentine and enamel by fluorapatite; (2) replacement of the fluorapatite by aluminium sulphate phosphate phases (crandallite–woodhouseite series); and (3) replacement of part of the tooth by gypsum and Al- and Fe-rich phosphates (vauxite) as a consequence of the superficial weathering of the fossil-rich outcrop. The presence of organic matter and sulphides in the host rocks was the main factor promoting the dissolution of the previous phases and crystallization of the new ones. This is the first description of the replacement of biogenic apatite in dinosaur teeth by aluminium and iron phosphates as a result of fossil diagenetic processes; these minerals are scarce in sedimentary environments and may have economic interest because they contain rare earth or radioactive elements. Additionally, our results indicate that, to ensure proper curating of the specimens from this fossil site, it is crucial to completely remove the host rock and strictly control storage humidity.

Spalacotheriids, basal trechnotheres with an acutely angled, reversed-triangle pattern of major molar cusps, are relatively rare pan-Laurasian mammals that ranged through much of the Cretaceous but are generally not well represented in the fossil record. Herein we describe newly collected specimens from the Molino Alto 1 locality, Teruel Province, Spain, and review previously described materials from the nearby locality of Galve; the sites lie in the same horizon in the upper El Castellar Formation and are of early Barremian age. On this basis, we recognize and name Aliaga molinensis, gen. et sp. nov., and provisionally place in the same genus a previously described species, cf. Aliaga henkeli, comb. nov., which appears to be the largest known spalacotheriid. The two species are characterized by a distinctive, multicuspate cingulid on lower cheek teeth. Based on acuteness of molar cusp triangulation and other features (e.g., planar molar shearing surfaces, elongation and cusp pattern of penultimat...

New fossils from the lower Barremian (Lower Cretaceous) of Spain increase the known occurrence of iguanodont dinosaurs (i.e. members of the clade Iguanodontia) in the Iberian Peninsula. A partial dentary and ischium from the fossil locality of La Cantalera 2 (Blesa Formation, municipality of Josa, Teruel province) testify to the presence of a new specimen in the early Barremian of Iberia, closely related to Delapparentia turolensis, which was previously only known in the type locality (La Maca-3, Camarillas Formation, municipality of Galve, Teruel province). These new fossils are assigned to Delapparentia turolensis (ischium) and cf. Delapparentia sp. (dentary). Moreover, the description and comparison of some of the characters of the ischium reveal that this bone is of previously unnoticed relevance for iguanodont systematics. The differences in overlapping postcranial material – observed by comparing ischial material from the provinces of Teruel and Burgos – lead us to conclude that late Hauterivian-early Barremian iguanodont diversity in Iberia consists of at least two different taxa: Delapparentia turolensis and another form closely related to Iguanodon. Additionally, dental morphotypes from La Cantalera bonebed evince the presence of at least three different iguanodonts in the lower Barremian of Iberia.

Sequence-stratigraphic, biostratigraphic and strontium-isotopic data have made it possible to update what is known of the stratigraphy and sedimentary evolution of the Kimmeridgian–Berriasian synrift successions recorded in four subbasins (Aguilon, Oliete, Morella, Galve) of the central Iberian Rift System (NE Spain). The studied successions are arranged in three unconformity-bounded synrift sequences. Synrift sequence 1A (Kimmeridgian–mid-early Tithonian) includes four transgressive-regressive sequences deposited on low-angle carbonate ramps, characterized by shallow-water grain-supported facies and coeval open-marine rhythmic successions of marls and lime mudstones (overall thickness ranging from 120 to 250 m). Ammonite biostratigraphy, combined with the last occurrence of Alveosepta jaccardi (Schrodt) and strontium-isotopic data constrains the Kimmeridgian-Tithonian boundary towards the uppermost part of synrift sequence 1A. Synrift sequence 1B (mid-early Tithonian to mid-Berriasian) consists of coastal to shallow-marine carbonate to mixed carbonate-siliciclastic successions, with a continuous record in the depocentral areas of the Morella and Galve subbasins (up to 365 m in thickness). Anchispirocyclina lusitanica (Egger) is common in the lower and middle part of the sequence, whereas charophytes of the lower Berriasian Globator maillardii maillardii Zone are recorded in its upper part. Strontium-isotopic data and ostracods whose first occurrence is Berriasian, indicate that the Tithonian-Berriasian boundary is located towards the mid-upper part of the synrift sequence 1B. Synrift sequence 1C (mid-Berriasian–mid-early Valanginian) is locally recorded in the Galve subbasin and consists of siliciclastic continental successions (up to 100 m in thickness). The available biostratigraphic data (charophytes, ostracods, sporomorphs) indicate that this sequence was deposited upwards from early Berriasian Globator maillardii incrassatus Zone. The presence of the lower Valanginian successions at the upper part of synrift sequence 1C in certain subsiding areas of the Galve subbasin cannot be ruled out. The sequence-stratigraphic and biostratigraphic data reported here indicate that the Galve and Morella subbasins started to develop during the Kimmeridgian. Successive stages of tectonic activity affected these subbasins during the mid-early Tithonian, the mid-Berriasian, and around the Berriasian-Valanginian transition. The stratigraphy and tectono-sedimentary evolution of the central Iberian Rift System indicate that the Berriasian successions are linked to the “Jurassic cycle”.

The Upper Cretaceous outcrops of the Pyrenees yield one of the most extensive and continuous records of paleoological remains anywhere in the world. Most of eggs and eggshells have been referred to...

This paper describes the skull of the eusuchian Allodaposuchus subjuniperus sp. nov. This new skull was recovered between the villages of Beranuy and Serraduy del Pon (Huesca, Spain). Stratigraphic...

Abnormalities in the histo- and ultrastructure of the amniote eggshell are often related to diverse factors, such as ambient stress during egg formation, pathologies altering the physiology of the egg-laying females, or evolutionarily selected modifications of the eggshell structure that vary the physical properties of the egg, for example increasing its strength so as to avoid fracture during incubation. When dealing with fossil materials, all the above hypotheses are plausible, but a detailed taphonomical study has to be performed to rule out the possibility that secondary processes of recrystallization have occurred during fossilization. Traditional analyses, such as optical microscopy inspection and cathodoluminescence, have proven not to be enough to understand the taphonomic story of some eggshells. Recently, electron backscatter diffraction has been used, in combination with other techniques, to better understand the alteration of fossil eggshells. Here we present a combined study using scanning electron microscopy, optical microscopy, cathodoluminescence and electron backscatter diffraction of eggshell fragments assigned to Megaloolithus cf. siruguei from the Upper Cretaceous outcrops of the Cameros Basin. We focus our study on the presence of secondary shell units that mimic most aspects of the ultrastructure of the eggshell mammillae, but grow far from the inner surface of the eggshell. We call these structures extra-spherulites, describe their crystal structure and demonstrate their secondary origin. Our study has important implications for the interpretation of secondary shell units as biological or pathological structures. Thus, electron backscatter diffraction complements other microscope techniques as a useful tool for understanding taphonomical alterations in fossil eggshells. Fil: Moreno Azanza, Miguel. Universidad de Zaragoza; España Fil: Bauluz, Blanca. Universidad de Zaragoza; España Fil: Canudo, José Ignacio. Universidad de Zaragoza; España Fil: Gasca Pérez, José Manuel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte; Argentina. Provincia de Neuquén. Ministerio de Energía, Ambiente y Servicios Públicos. Dirección Provincial de Minería. Museo Provincial de Ciencias Naturales Prof. ; Argentina. Universidad de Zaragoza; España Fil: Torcida Fernández Baldor, Fidel. Museo de Los Dinosaurios, Salas de Los Infantes, Burgos; España

A review of the stratigraphy of the Galve sub-basin (western Maestrazgo Basin, eastern Spain) around the Jurassic-Cretaceous transition is presented here, based on new data acquired after extensive geological mapping and logging complemented with facies analysis, new biostratigraphic data and a revision of the published information available. The results obtained are relevant for a more detailed understanding of the tecto-sedimentary evolution of the studied basin during the transition between two stages of rift evolution (i.e., syn-rift sequences 1 and 2). In addition, new information on the age and setting of numerous dinosaur fossil- and track-sites found across the Galve sub-basin and in the northern part of the nearby Penyagolosa sub-basin is provided here. Two new lithostratigraphic units are defined and characterized, the Aguilar del Alfambra and the Galve formations. The previous stratigraphic framework considered only two lithostratigraphic units (the Villar del Arzobispo and El Castellar formations) bounded by a single regional unconformity, and this resulted in significant misinterpretations. The whitish limestones, red lutites and cross-bedded sandstones of the Aguilar del Alfambra Formation were deposited in transitional environments, ranging from coastal lutitic plains to restricted lagoons. Of particular interest are the laminated micritic-peloidal limestones with abundant fenestral porosity (supratidal ponds to intertidal flats), which preserve common dinosaur footprints. This unit is bounded by widespread unconformities and is of very variable thickness (0-450 m), controlled by extensional tectonics operating at the climax of syn-rift sequence 1 during the latest Tithonian-middle Berriasian. The overlying Galve Formation is of variable thickness (from 0 to 100 m) and is also bounded by regional unconformities described in detail here. It consists of red lutites with cross-bedded and tabular-burrowed sandstones representing channel and overflow deposits in an alluvial floodplain. The sauropod dinosaur Aragosaurus ischiaticus found in this unit has a controversial age assignment. The age of the Galve Formation is poorly constrained from late Berriasian to Hauterivian, but new biostratigraphic data presented here, combined with the correlation with the nearby Penyagolosa and Salzedella sub-basins, suggest a possible equivalence to the upper Berriasian-lower Valanginian sequence deposited during the initial stage of syn-rift sequence 2. Fil: Aurell, Marcos. Universidad de Zaragoza; España Fil: Bádenas, Beatriz. Universidad de Zaragoza; España Fil: Gasca Pérez, José Manuel. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad de Zaragoza; España Fil: Canudo, José I.. Universidad de Zaragoza; España Fil: Liesa, Carlos L.. Universidad de Zaragoza; España Fil: Soria, Ana R.. Universidad de Zaragoza; España Fil: Moreno Azanza, Miguel. Universidad de Zaragoza; España Fil: Najes, Laura. Universidad de Zaragoza; España

A partially articulated postcranial skeleton of a small ornithischian dinosaur, Gideonmantellia amosanjuanae nov. gen. et sp., from the Early Cretaceous of Galve (Teruel province, Spain) is described. It was recovered in an outcrop of fluvial red clays from the Camarillas Formation, which is Barremian in age. This partial skeleton is recognised as a new ornithopod taxon by the following autapomorphies: (1) postacetabular process of the ilium with a brevis shelf that is noticeably medially expanded in its cranial part but narrow and horizontal in its caudal part; (2) rod-like prepubic process with its anterior end twisted and expanded; and (3) L-shaped first chevron. Our phylogenetic analysis indicates that Gideonmantellia represents a basal ornithopod taxon more derived than Orodromeus , the “Asian clade” (which includes Haya and others) and Hypsilophodon .

The sauropodAragosaurus ischiaticusSanz, Buscalioni, Casanovas & Santafé, 1987 was the first dinosaur to be described in Spain. The holotype was recovered from the site of Las Zabacheras (Galve, Teruel province). This site has traditionally been situated in the El Castellar Formation (in the lower part of the Wealden facies). Recently, it has been proposed that the remains ofAragosaurusstem from the Villar del Arzobispo Formation (late Tithonian–upper part of the early Berriasian), which would mean that the sauropod was almost 15 million years older than previously thought. Detailed field work has been carried out, making it possible to pinpoint the position of the low-angle unconformity between the Villar del Arzobispo Formation and the El Castellar Formation. This unconformity originated as a result of block tilting that occurred during the early stages of the Early Cretaceous rifting episode. The upper levels of the Jurassic sequence (i.e. the Villar del Arzobispo Formation) were exposed to erosion and karstification, leading to the formation of a discontinuous conglomeratic level. This level has been locally preserved at the bottom of the Wealden syn-rift sequence (i.e. the El Castellar Formation). The results of our detailed mapping demonstrate that theAragosaurusholotype was found in the lower part of the El Castellar Formation. Moreover, our revision of the existing datings suggests that the El Castellar Formation as a whole is Valanginian?–early Barremian in age. Given thatAragosauruswas located in its lower part, it is probably Valanginian?–Hauterivian in age.

Cuenca-Bescos, G., Badiola, A., Canudo, J.I., Gasca, J.M., and Moreno-Azanza, M. 2011. New dryolestidan mammal from the Hauterivian—Barremian transition of the Iberian Peninsula. Acta Palaeontologica Polonica 56 (2): 257–267.

Delapparentia turolensis Ruiz-Omenaca, 2011 is the only iguanodont taxon erected in the Barremian of Spain. It is described on the basis of a partial postcranial skeleton discovered in the 1950s near the village of Galve (Teruel Province), within the Camarillas Formation. Recently, new remains from the same individual have been recovered, and these are described here. Furthermore, after first-hand examinations of the holotype, the phylogenetic position of this taxon has been analysed for the first time, and its diagnosis is emended. Delapparentia turolensis is a large-sized, basal iguanodont which presents an autapomorphic, unusually high axial neural spine and a unique combination of postcranial characters. The ilium morphology differs from that of other basal iguanodonts and relates Delapparentia to the Valanginian Barilium dawsoni from England, with whom it shares two synapomorphies. In our phylogenetic analysis Delapparentia is recovered in a polytomy with Kukufeldia , Lanzhousaurus , Barilium and the clade equivalent to Iguanodontoidea.

Trigonoolithus amoae oogen. et oosp. nov. is described on the basis of abundant eggshell fragments from the La Cantalera 1 site in the Early Cretaceous (early Barremian) Blesa Formation, Teruel Province, northern Spain. The surface ornamentation, the most diagnostic feature of the new oogenus, consists of closely spaced sub-triangular or rounded protuberances that have not previously been reported in any other ootaxon. The eggshells present three distinct layers, with a gradual transition between prismatic and mammillary layers, and a poorly developed external layer with progressive prismatic to external layer transition, a combination of characters that allows them to be assigned to Prismatoolithidae. Phylogenetic analyses based on oological characters place Trigonoolithus at the base of Prismatoolithidae.

Fil: Cruzado Caballero, Penélope. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigación en Paleobiología y Geología; Argentina Fil: Canudo, José. Grupo Aragosaurus-IUCA, Área de Paleontología, Facultad de Ciencias , Universidad de Zaragoza; España. Fil: Moreno Azanza, Miguel. Grupo Aragosaurus-IUCA, Área de Paleontología, Facultad de Ciencias , Universidad de Zaragoza; España. Fil: Ruiz Omeñaca, José Ignacio. Grupo Aragosaurus-IUCA, Área de Paleontología, Facultad de Ciencias , Universidad de Zaragoza, Museo del Jurásico de Asturias (MUJA); España. Arenysaurus ardevoli is a lambeosaurine hadrosaurid from the late Maastrichtian of Aren (Huesca, northern ´ Spain) that has recently been described. The holotype is the first and the most complete lambeosaurine with a braincase from Europe. In this paper, we present a complete description of the postcranial skeleton, which was poorly described when the taxon was named because it was partially unprepared, and new information on several cranial bones (jugal, maxilla, and dentition). A new phylogenetic analysis of Arenysaurus and the closely related Blasisaurus canudoi, also from the late Maastrichtian of Aren, places them inside the Parasaurolophini in a dichotomy with ´ Parasaurolophus spp. Paleobiogeographically, the presence of Arenysaurus and its relationships with other lambeosaurines suggest at least one geodispersal event from Asia to Europe no later than the middle–late Campanian. .

La Cantalera is an accumulation site for fossil vertebrates consisting mainly of teeth and isolated postcranial remains. It has the greatest vertebrate biodiversity of any site from the Hauterivian-Barremian transition in the Iberian Peninsula. Up to now, 31 vertebrate taxa have been recognized: an osteichthyan (Teleostei indet.), two amphibians (Albanerpetonidae indet. and Discoglossidae indet.), a chelonian (Pleurosternidae? indet.), a lizard (Paramacellodidae? indet.), four crocodylomorphs (cf. Theriosuchus sp., Bernissartiidae indet., Goniopholididae indet., cf. Lisboasaurus sp.), two pterosaurs (Istiodactylidae? indet., Ornithocheiridae? indet.), four ornithopod dinosaurs (Iguanodontoidea indet., Hadrosauroidea? indet., “Hypsilophodontidae” indet., Rhabdodontidae? indet.), a thyreophoran (Polacanthidae indet.), a sauropod (Euhelopodidae indet.), eleven theropods (Carcharodontosauridae? indet., Baryonychinae indet., aff. Paronychodon sp., Maniraptora indet. 1-3, Dromaeosaurinae indet. 1-3, Velociraptorinae indet., Avialae? indet.) and three or four multituberculate mammals (Cantalera abadi, Eobaatar sp., Plagiaulacidae or Eobaataridae gen. et sp. indet., “Plagiaulacida” indet.). Nine ootaxa have also been distinguished at the site of La Cantalera. Oofamilies assigned to dinosaurs (Elongaloolithidae, Prismatoolithidae, cf. Spheroolithidae), to crocodiles (Krokolithidae) and eggshells of two oofamilies incertidae sedis are represented. This association is consistent with the record of bone and tooth remains from the site. La Cantalera was formed in a marshy environment where there was an intermittent body of water. The great majority of the vertebrate fossil remains lack evidence of transport, so the preserved association is a good representation of the vertebrate ecosystem in or around the marshy area of La Cantalera. The vertebrate association of La Cantalera displays certain differences with respect to those from lacustrine environments of the Hauterivian-Barremian transition of the Iberian Range. These differences include, for example, the absence of Chondrichthyes, the merely token presence of the osteichthyans, the scarcity of chelonians, the presence of exclusively multituberculate mammals, the lower diversity of sauropods and the greater diversity of theropods.