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Ceratosaurus tooth from the Lourinhã formation at Porto das Barcas (Portugal), showing vertical ridges on lingual surface (ML809).
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Late Jurassic theropod dinosaurs have been known in Portugal since 1863 but only now are they being fully understood, with the recognition of genera such as Allosaurus, Aviatyrannis, Ceratosaurus, Lourinhanosaurus, and Torvosaurus from the Lourinhã and Alcobaça Formations (Kimmeridgian/Tithonian). Ceratosaurus dentisulcatus can now be reported from...
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... material from Portugal was collected in Lourinhã Municipality (right femur and left tibia ML352 from Rodela do Valmitão, figure 2, tooth ML809 from Porto das Barcas, tooth ML737 from Peralta and tooth ML342 from Merendeiro; figure 1). The most identifiable mate- rial is from the Praia da Amoreira Member, in the lower part of the Lourinhã Formation (Kimmeridgian). ...
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Citations
... The principal objective of this study is to provide a comprehensive re-description and detailed anatomical analysis of Allosaurus europaeus ML415, including the partial skull, three cervical vertebrae, and several ribs, which represent the holotype specimen [10]. The secondary objective of this study is to discover, if possible, a new set of characters present in the specimen and compare them with those of other Allosaurus species using morphometric phylogenetic analysis. ...
... Dinosauria Owen, 1842 Saurischia Seeley, 1887 Theropoda Marsh, 1881 Neotheropoda Bakker, 1986 Tetanurae Gauthier, 1986 Avetheropoda Paul, 1988 Carnosauria Huene, 1920 Allosauroidea Currie and Zhao, 1994 Allosauridae Marsh, 1878 Allosaurus Marsh, 1877 Allosaurus europaeus Mateus et al., 2006 Holotype: ML415 is comprised of two blocks and several separated fragments. The first block contains a partially articulated skull, with most of the left posterior bones preserved and part of the atlanto-axis complex, including three teeth. ...
... Since it is poorly preserved, we cannot assess its entire shape; nevertheless, by combining both fossae and according to previous reconstructions [3,10], it may suggest that it does not look so different in shape compared to other Allosaurus species. However, the ridge that delineates this fossa, as previously mentioned, is far less pronounced in this specimen compared to other Allosaurus species, creating a shallower fossa in the contact between the lacrimal and jugal bones. ...
Allosaurus is one of the most famous theropod dinosaurs, but the validity and relationships between the different species have been confusing and often questioned. Portugal is relevant to the understanding of the genus in light of the discovery of A. europaeus ML415 from the Early Tithonian of Lourinhã and Allosaurus MNHNUL/AND.001 from Andrés. However, the exact classification and validity of these two specimens has always been controversial. The presence of Allosaurus in Portugal is strong evidence for a North America–Europe Late Jurassic dispersal, later supported by other taxa. A detailed cranial description and specimen-based phylogeny were performed and resolved many of the open questions: (1) The diversity of Allosaurus is limited to three named species: A. fragilis, A. europaeus, and A. jimmadseni. (2) Nine autapomorphies were found in A. europaeus, confirming the validity of the species. (3) Phylogenetic analyses place both Portuguese specimens in the genus Allosaurus, based on the following synapomorphies: jugal bone lateral view, relative heights of quadratojugal prongs, the dorsal prong is equal in height, the jugal bone in lateral view shows shallow accessory pneumatization of the antorbital fossa, the palatine pneumatic recess shape is small, and lacrimal horn morphology has a triangular horn. (4) The Andrés specimen is placed with the A. europaeus and they are considered here to be the same species, which is paleo-geographically and biochronologically congruent. (5) A. europaeus and A. jimmadseni are sister taxa and closer to each other than to A. fragilis. The genus is distributed in occurrences from the United States, Germany, and Portugal, and from the Late Kimmeridgian to the Late Tithonian, while the Cenomanian report from Japan is reidentified as Segnosaurus.
... Antunes and Mateus 2003;Escaso et al. 2007Escaso et al. , 2014Mateus et al. 2009Mateus et al. , 2014Mocho et al. 2017Mocho et al. , 2019Ortega et al. 2017). The fauna of theropod dinosaurs currently known in these levels is diverse and includes mostly medium-and large sized taxa, including Ceratosaurus (Mateus and Antunes 2000a;Malafaia et al. 2015), Torvosaurus (Mateus and Antunes 2000b;Araújo et al. 2013;Hendrickx et al. 2014;, Lourinhanosaurus (Mateus 1998 Mateus et al. 2006), and Lusovenator . However, this record also includes several specimens, mainly represented by isolated material that has been interpreted as belonging to small sized coelurosaurian theropods (e.g. ...
... On the other hand, most taxa of medium-or large-bodied sized theropods from the Upper Jurassic of Portugal (Ceratosaurus, Torvosaurus, and Allosaurus) may have been present since the early Late Jurassic, despite their osteological fossil record in the Lusitanian Basin is particularly well known for the late Kimmeridgian to the early Berriasian interval (e.g. Pérez-Moreno et al. 1999;Mateus et al. 2006;Hendrickx et al. 2014;Malafaia et al. 2015Malafaia et al. , 2017aMalafaia et al. , 2017b. This is also possibly the case for Lourinhanosaurus, which is represented by material restricted to Kimmeridgian to lower Tithonian sequences (Mateus 1998), but the fossil record of the clade to which it has been related, the Metriachantosauridae, is known in Laurasia at least since the early Middle Jurassic (Wu et al. 2009). ...
... More precise details of this classification have been suggested, with several works attributing the eggs and embryos of Paimogo to the allosauroid theropod Lourinhanosaurus antunesi, on the basis of the similar age and geological proximity, the relative proportions of the centra of the presacral vertebrae (Riqcles et al., 2001), and the presence of a large premaxilla showing four alveoli, which is unlike Allosaurus (Carrano et al., 2013). However, both studies predate the description of two new allosauroid theropod species from the Lourinhã Formation: Allosaurus europaeus and Lusovenator santosi (Mateus et al., 2006;Malafaia, E. et al., 2020). Further detailed taxonomic studies of the embryonic material are needed to improve the taxonomic attribution of the Paimogo egg assemblage. ...
The accurate identification of dinosaur egg accumulations as nests or clutches is crucial for understanding the reproductive behaviour of these extinct species. However, existing methods often rely on the presence of complete eggs and embryo remains, and sedimentological criteria that are only applicable to well-structured sediments. In this study, we introduce an innovative approach to characterize egg accumulations in structureless
sediments, where traditional nest structures may not be preserved. Our methodology employs a unique combination of sedimentological, taphonomic, geochemical, and geophysical proxies for the study of egg accumulations. We applied this approach to the egg accumulation from Paimogo (Jurassic, Portugal), traditionally interpreted as a nest. Our findings reveal that the Paimogo egg assemblage is a secondary deposit, resulting from a flooding event in a fluvial plain that dismantled several allosauroid and crocodylomorph clutches. The eggshell vapor conductance results, coupled with sedimentological evidence, suggest that allosauroid dinosaurs buried their eggs in the dry terrain of overbank areas close to a main channel during the breeding season, likely during the dry season to prevent the
embryos from drowning. This research underscores the necessity of multidisciplinary approaches in interpreting egg accumulations and offers a novel methodology for studying these accumulations in structureless sediments. Our findings provide new insights into the breeding behaviour and nesting preferences of these extinct organisms, contributing to our understanding of dinosaur ecology.
... Theropods are mainly represented by medium to large-sized forms belonging to Ceratosauria or tetanurans, including members of Megalosauridae and Allosauroidea (e.g. Mateus 1998Mateus , 2006Hendrickx and Mateus et al. 2014;Malafaia et al. 2010Malafaia et al. , 20152017a, b;, but some small theropods are also recorded. The analyses of the relationship between the Late Jurassic fauna of Portugal and North America have been a source of many studies. ...
Management of Natural History collections, particularly paleontological collections, is usually structured around specimens. However, it might not be feasible when the collections are incorporated from sources where the whole cycle of specimen collection is unknown, making it hard to determine what fossils belong to a single specimen and if these specimens belong to a single fossil-assemblage, taxon or individual. The Sociedade de História Natural from Torres Vedras (Portugal) manages such collections using a management strategy approach based on the spatial location from where the fossils were found. To achieve this, we have implemented a methodology for field collection and developed a spatial database in PostgreSQL/PostGIS to manage the entire stages of paleontological collections management. On top of the database, a second layer of web services, Web GIS (Geographical Information System) applications, and desktop GIS, all based on QGIS, are also under development to facilitate access to data stored in the database to internal and external researchers and partners, as well public institutions. This effort is made in close collaboration with in-house actors and stakeholders and has yielded promising results. The development of this system provides four advantages: (i) it played an essential role in supporting the transfer of the collection to new storage facilities; (ii) it provides overviews of the distribution of the characteristics of the collections; (iii) it enhances institutional partnerships by providing consistent and reliable data and(iv) it supports the establishment and development of research lines.
... The Lusitanian Basin has yielded one of the most extensive fossil collections of Late Jurassic dinosaurs and other vertebrates from Europe. The similarity with coeval faunas of the North American Morrison Formation has been well-documented, with several shared genera (e.g., Stegosaurus, Escaso et al. 2007; Ceratosaurus, Mateus and Antunes 2000a;Malafaia et al. 2015;Torvosaurus, Mateus and Antunes 2000b;Hendrickx and Mateus 2014;Malafaia et al. 2017c;Allosaurus, Pérez-Moreno et al. 1999;Mateus et al. 2006). Other European Upper Jurassic fossil records are scarce (particularly for theropod dinosaurs) and mostly consist of fragmentary material with uncertain phylogenetic interpretation (e.g., Allain and Pereda Suberbiola 2003;Canudo and Ruiz-Omeñaca 2003;Ruiz-Omeñaca et al. 2008;Lubbe et al. 2009;Cobos et al. 2014;Evers and Wings 2020). ...
Malafaia, E., Mocho, P., Escaso, F., Narvaéz, I., and Ortega, F. 2024. Taxonomic and stratigraphic update of the material
historically attributed to Megalosaurus from Portugal. Acta Palaeontologica Polonica 69 (2): 127–171. https://doi.org/10.4202/app.01113.2023
... There is a striking similarity between the Late Jurassic dinosaur faunas of North America and Europe. Numerous iconic dinosaur taxa, including †Allosaurus [35], †Torvosaurus [36], †Ceratosaurus [35], †Supersaurus [37] and †Stegosaurus [38] are present in penecontemporaneous deposits in both the North American Morrison and Portuguese Lourinhã Formations ( figure 5). The discovery of the North American species †Helioscopus dickersonae and its close relationship to †Ardeosaurus brevipes and †Limnoscansor digitatellus of the German Solnhofen Limestone establishes another broadly distributed Laurasian clade of Jurassic squamates, alongside the North American paramacellodids [24], the Morrison specimens referred to the (possible) pan-anguimorph †Dorsetisaurus [39], and to †Diablophis gilmorei [40] which was originally referred to the English taxon †Parviraptor [41]. ...
... There is a striking similarity between the Late Jurassic dinosaur faunas of North America and Europe. Numerous iconic dinosaur taxa, including †Allosaurus [35], †Torvosaurus [36], †Ceratosaurus [35], †Supersaurus [37] and †Stegosaurus [38] are present in penecontemporaneous deposits in both the North American Morrison and Portuguese Lourinhã Formations ( figure 5). The discovery of the North American species †Helioscopus dickersonae and its close relationship to †Ardeosaurus brevipes and †Limnoscansor digitatellus of the German Solnhofen Limestone establishes another broadly distributed Laurasian clade of Jurassic squamates, alongside the North American paramacellodids [24], the Morrison specimens referred to the (possible) pan-anguimorph †Dorsetisaurus [39], and to †Diablophis gilmorei [40] which was originally referred to the English taxon †Parviraptor [41]. ...
Geckos are a speciose and globally distributed clade of Squamata (lizards, including snakes and amphisbaenians) that are characterized by a host of modifications for nocturnal, scansorial and insectivorous ecologies. They are among the oldest divergences in the lizard crown, so understanding the origin of geckoes (Gekkota) is essential to understanding the origin of Squamata, the most species-rich extant tetrapod clade. However, the poor fossil record of gekkotans has obscured the sequence and timing of the assembly of their distinctive morphology. Here, we describe the first North American stem gekkotan based on a three-dimensionally preserved skull from the Morrison Formation of western North America. Despite its Late Jurassic age, the new species already possesses several key characteristics of the gekkotan skull along with retained ancestral features. We show that this new stem gekkotan, and several previously named species of uncertain phylogenetic relationships, comprise a widespread clade of early crown lizards, substantiating faunal homogeneity in Laurasia during the Late Jurassic that extended across disparate ecological, body-size and physiological classes.
... Despite being one of the most abundant and well-known theropod, Allosaurus still has a complex and not fully resolved taxonomic status. Currently, the record of Allosaurus in Portugal would include the set of cranial and postcranial elements so far undescribed from Andrés and the holotype of Allosaurus europaeus (Mateus et al. 2006), a partial skull associated with cervical vertebrae and ribs from the Upper Jurassic of Praia de Vale Frades (Lourinhã). Here we describe a set of axial elements that includes dorsal, sacral and caudal vertebrae collected in Andrés. ...
... Ceratosaurus (Infraorder Neotheropoda, Superfamily Ceratosauria, Family Ceratosauridae) was a large predator that lived during the Late Jurassic (Kimmeridgian and Tithonian) of North America and Europe (Madsen and Welles, 2000;Mateus et al., 2006;Foster, 2007). Ceratosaurus nasicornis Marsh 1884 is characterized by a large head with respect to the rest of its body, with a narrow and rectangular section (Fig. 5). ...
... Allosaurus (Infraorder Carnosauria, Superfamily Allosauroidea, Family Allosauridae) is a large theropod that habited in North America and Iberian Peninsula during the Late Jurassic (Oxfordian to Tithonian; Pérez-Moreno et al., 1999;Turner and Peterson, 1999;Mateus et al., 2006;Carpenter, 2010). Allosaurus fragilis Marsh 1877 is the bestknown specie. ...
The aim of this work is to obtain diverse morphometric data from digitized 3D models of scientifically accurate palaeoreconstructions of theropods from eight representative families. The analysed polyvinyl chloride (PVC) models belong to the genera Coelophysis , Dilophosaurus , Ceratosaurus , Allosaurus , Baryonyx , Carnotaurus , Giganotosaurus , and Tyrannosaurus. The scanned 3D models were scaled considering different body-size estimations of the literature. The 3D analysis of these genera provides information on the skull length and body length that allows for recognition of major evolutionary trends. The skull length/body length in the studied genera increases according with the size of the body from the smallest Coelophysis with a ratio of 0.093 to ratios of 0.119–0.120 for Tyrannosaurus and Giganotosaurus , the largest study theropods. The study of photogrammetric 3D models also provides morphometric information that cannot be obtained from the study of bones alone, but knowing that all reconstructions begin from the fossil bones, such as the surface/volume ratio (S/V). For the studied theropod genera surface/volume ratio ranges from 35.21 for Coelophysis to 5.55 for Tyrannosaurus . This parameter, closely related to the heat dissipation, help in the characterization of the metabolism of extinct taxa. Accordingly, slender primitive forms of the Early Jurassic (i.e. Coelophysis and Dilophosaurus ) had relatively smaller skulls and higher mass-specific metabolic rates than the robust large theropods of the Cretaceous (i.e. Giganotosaurus and Tyrannosaurus ) . This work presents a technique that, when applied to proper dinosaur models, provides extent and accurate data that may help in diverse study areas within the dinosaur palaeontology and palaeobiology.
... Although Foster (2007) noted that Torvosaurus is the second most abundant theropod taxon in the Morrison Formation, few specimens have been recorded so far, and the vast majority are fragmentary and occur in multi-taxon bonebeds in association with the much more abundant Allosaurus (Hanson & Makovicky 2014). Likewise, the European species of Torvosaurus is also the largest and most massive theropod dinosaur from the Late Jurassic of this continent, but, although megalosaurids seem to have been less rare in the European Late Jurassic than in North America (Rauhut et al., 2018), all specimens recorded so far are fragmentary (Antunes & Mateus 2003;Mateus et al. 2006;Malafaia et al. 2008Malafaia et al. , 2017aHendrickx & Mateus 2014). ...
A fragmentary maxilla from the middle Callovian Ornatenton Formation of the Wiehengebirge, north-western Germany, shows two autapomorphies of the theropod dinosaur genus Torvosaurus, a maxilla fenestra that is developed as a large and shallow but not sharply defined depression and an anteroposteriorly oriented ridge transversing the ventral part of the maxillary fenestra. This specimen represents the first occurrence of this genus from Germany and the oldest record of Torvosaurus, which is otherwise securely known from the Kimmeridgian-Tithonian of Portugal and the western USA. Given that the two closest relatives of Torvosaurus, Megalosaurus and Wiehenvenator, are known from the Bathonian of England and the Callovian of Germany, respectively, an evolutionary origin of derived megalosaurines in north-central Europe is indicated. The records of Torvosaurus in the Kimmeridgian-Tithonian of Portugal and the Morrison Formation of the western USA most probably represent dispersal of the genus from this area in the Late Jurassic.
... The Portuguese specimens represent some of the earliest evidence of these clades in Laurasia and have significant implications for understanding the paleobiogeographic context and dynamics of Late Jurassic theropod fauna in the peri-Atlantic realm. The Upper Jurassic theropod record in the Lusitanian Basin is mostly composed of medium-to large-sized taxa with a trans-Atlantic distribution, such as the genera Ceratosaurus (e.g., Malafaia et al., 2015), Torvosaurus (e.g., Hendrickx and Mateus, 2014b), and Allosaurus (e.g., Pérez-Moreno et al., 1999;Mateus et al., 2006). However, this record also includes some exclusive taxa interpreted as related to Jurassic forms from both Laurasian (e.g., Lourinhanosaurus antunesi; Mateus, 1998) and Gondwanan (e.g., Carcharodontosauria;Malafaia et al., 2019) landmasses. ...
Carcharodontosaurian allosauroids were temporally restricted to the Cretaceous, being known from all land masses with the exception of Antarctica. In addition to Veterupristisaurus from Tanzania, exceptions to this distribution have been reported recently, consisting on fragmentary materials from Upper Jurassic strata of China, Germany, and Portugal. Here, we propose a new Late Jurassic carcharodontosaurian taxon, Lusovenator santosi, gen. et sp. nov. based on the reevaluation of previously described specimens from the Lusitanian Basin, Portugal. The performed phylogenetic analysis recovered Lusovenator santosi as an early branching carcharodontosaurian allosauroid diagnosed by an exclusive combination of characters, including three autapomorphic features: (1) large recesses in neural arch of anterior dorsal vertebrae; (2) well-developed and continuous longitudinal laminae extending from the tip of the prezygapophyses to the distal end of the postzygapophyses in mid-caudal vertebrae; and (3) supraacetabular crest of ilium forming a prominent ventrolaterally projecting shelf. Lusovenator santosi is the oldest carcharodontosaurian allosauroid yet discovered from Laurasia and supports unequivocally the hypothesis of a pre-Cretaceous scenario for the radiation of the clade. The identification of this taxon highlights the high diversity of medium- to large-bodied theropods in the later part of the Late Jurassic of the Iberian Peninsula. Carcharodontosauria is not yet known in correlative levels of the North American Morrison Formation, and the existence of contacts after the late Tithonian between these landmasses could explain the distribution of this clade and other dinosaur groups present in the Iberian Jurassic and in the North American Lower Cretaceous.
