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Ceratosaurus tooth from the Lourinhã formation at Porto das Barcas (Portugal), showing vertical ridges on lingual surface (ML809).

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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... 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. ...
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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). ...
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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. ...
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... Besides, a study of the biomechanical significance of suture morphology of this taxon also used this configuration (Rayfield, 2005). Furthermore, the clear presence of an expression of the jugal on the rim of the antorbital fenestra was considered an important character to distinguish the European species of Allosaurus, A. europaeus, from its North American counterparts (Mateus, Walen & Antunes, 2006; see also Malafaia et al., 2007). ...
... Furthermore, the new observations have implications for the diagnosis of the European A. europaeus (Mateus, Walen & Antunes, 2006; Fig. 5D) as a distinct species, for which the jugal participation in the antorbital fenestra was listed as one of the few autapomorphic characters that differentiate it from the North American species. Besides, the authors listed the absence of a lacrimal-maxilla contact as a further apomorphy, which is related to the former character. ...
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... Some of the best and most complete theropod material in Europe comes from Late Kimmeridgian-Tithonian formations in the Lusitanian Basin of Portugal, including the Lourinhã and Alcobaça formations. The Portuguese theropod fauna includes the allosauroid Allosaurus europaeus (Mateus, Walen & Telles Antunes, 2006), the ceratosaur Ceratosaurus sp. (Mateus & Antunes, 2000), the megalosauroid Torvosaurus gurneyi (Hendrickx & Mateus, 2014;Malafaia et al., 2017), the allosauroid Lourinhanosaurus antuneso (Mateus, 1998;Benson, 2010), the tyrannosauroid Aviatyrannis jurassica (Rauhut, 2003b). ...
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Marine limestones and marls in the Langenberg Quarry provide unique insights into a Late Jurassic island ecosystem in central Europe. The beds yield a varied assemblage of terrestrial vertebrates including extremely rare bones of theropod from theropod dinosaurs, which we describe here for the first time. All of the theropod bones belong to relatively small individuals but represent a wide taxonomic range. The material comprises an allosauroid small pedal ungual and pedal phalanx, a ceratosaurian anterior chevron, a left fibula of a megalosauroid, and a distal caudal vertebra of a tetanuran. Additionally, a small pedal phalanx III-1 and the proximal part of a small right fibula can be assigned to indeterminate theropods. The ontogenetic stages of the material are currently unknown, although the assignment of some of the bones to juvenile individuals is plausible. The finds confirm the presence of several taxa of theropod dinosaurs in the archipelago and add to our growing understanding of theropod diversity and evolution during the Late Jurassic of Europe.
... Allosaurus has long played a crucial role in phylogenetic analyses of the Theropoda, either as a member of an ingroup or as an outgroup taxon in analyses of Coelurosauria (Rauhut, 2003;Benson, Carrano & Brusatte, 2010;Carrano, Benson & Sampson, 2012). Nineteen species of Allosaurus have been erected since 1877 (see Chure, 2000a;Mateus, Walen & Antunes, 2006;Dalman, 2014), although the holotype material for many has been neither fully illustrated nor described, and the validity of these species has not been critically evaluated. Many proposed synonymies are yet to be evaluated in detail, although we have a manuscript in preparation doing that. ...
... This crest is lowest anteriorly and highest (22 mm) slightly anterior to the cornual process or horn of the lacrimal. These crests are present in Allosaurus jimmadseni (DINO 11541, MOR 693, SMA 0005, and BYU 5253) as well as in Allosaurus europaeus (ML 415) (Mateus, Walen & Antunes, 2006). This dorsolateral nasal crest is not present in Allosaurus fragilis (USNM 4734, DINO 2560, BYU 9466, UMNH VP 7748, and UMNH VP 9149-see figure 11), andis not present in Sinraptor dongi, Neovenator, Acrocanthosaurus or Carcharodontosaurus. ...
... Instead, the jugal does contribute to the antorbital fossa. This is also the case in Allosaurus fragilis, but this is contrary to the published description of Allosaurus europaeus, in which the jugal intervenes between the lacrimal and maxilla (Mateus, Walen & Antunes, 2006). ...
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Allosaurus is one of the best known theropod dinosaurs from the Jurassic and a crucial taxon in phylogenetic analyses. On the basis of an in-depth, firsthand study of the bulk of Allosaurus specimens housed in North American institutions, we describe here a new theropod dinosaur from the Upper Jurassic Morrison Formation of Western North America, Allosaurus jimmadseni sp. nov., based upon a remarkably complete articulated skeleton and skull and a second specimen with an articulated skull and associated skeleton. The present study also assigns several other specimens to this new species, Allosaurus jimmadseni , which is characterized by a number of autapomorphies present on the dermal skull roof and additional characters present in the postcrania. In particular, whereas the ventral margin of the jugal of Allosaurus fragilis has pronounced sigmoidal convexity, the ventral margin is virtually straight in Allosaurus jimmadseni . The paired nasals of Allosaurus jimmadseni possess bilateral, blade-like crests along the lateral margin, forming a pronounced nasolacrimal crest that is absent in Allosaurus fragilis .
... The observations made from the Valmitão assemblage are consistent with this proposal and support the biogeographical and taxonomic variation seen in previous studies, suggesting that Atoposauridae were already diverse by the end of the Late Jurassic in Europe (Tennant & Mannion, 2014;Tennant et al., 2016;Schwarz et al., 2017). This high trophic diversity, in the context of freshwater crocodylomorphs (Wilberg et al., 2019) that were relatively small in size, represents an adaptation in an ecosystem otherwise dominated by dinosaurs, where predatory niches were already occupied by mediumand large-sized predators, such as the theropods Lourinhanosaurus, Allosaurus and Torvosaurus (Mateus et al., 2006). palaeoecoloGical differences between the valmitão vma and the Guimarota mine assemblaGe The apparent absence of Lisboasaurus and the scarcity of Lusitanisuchus in the Valmitão vma were unexpected given their relative abundance in the similarly aged Guimarota Mine, suggesting that they were relatively uncommon in the Lourinhã Formation. ...
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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.
... The observations made from the Valmitão assemblage are consistent with this proposal and support the biogeographical and taxonomic variation seen in previous studies, suggesting that Atoposauridae were already diverse by the end of the Late Jurassic in Europe (Tennant & Mannion, 2014;Tennant et al., 2016;Schwarz et al., 2017). This high trophic diversity, in the context of freshwater crocodylomorphs (Wilberg et al., 2019) that were relatively small in size, represents an adaptation in an ecosystem otherwise dominated by dinosaurs, where predatory niches were already occupied by mediumand large-sized predators, such as the theropods Lourinhanosaurus, Allosaurus and Torvosaurus (Mateus et al., 2006). palaeoecoloGical differences between the valmitão vma and the Guimarota mine assemblaGe The apparent absence of Lisboasaurus and the scarcity of Lusitanisuchus in the Valmitão vma were unexpected given their relative abundance in the similarly aged Guimarota Mine, suggesting that they were relatively uncommon in the Lourinhã Formation. ...
Article
Full-text available
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.
... The Lourinhã Formation is the Late Jurassic unit with the most abundant record of vertebrate fossils in Portugal and Europe, with a complex ecosystem as evidenced by the vast diversity of the fossil record [73]. The Lourinhã Formation has been repeatedly compared with the Morrison Formation in North America for their similarities in fauna, paleoenvironment, age and sedimentology (e.g., [21,73,[78][79][80][81]). Although the proto-North Atlantic started expanding during the Hettangian, and by the Kimmeridgian and Tithonian when much of these formations were deposited it already separated Eastern North America and Iberia by more than 300 km, a regression during the Callovian-Oxfordian with eustatic sea-level drop and uplift resulted in ephemeral land-bridges connecting the two landmasses [80]. ...
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The stegosaur species Miragaia longicollum was erected based on a partial anterior skeleton from the Upper Jurassic of Portugal. Until then, almost all stegosaur specimens in Portugal and Spain had been identified as Dacentrurus armatus, the sister taxon of M. longicollum and only other member of the clade Dacentrurinae. The holotypes of the two species have little overlap, since the holotype of D. armatus is mostly a posterior skeleton, so the classification of other specimens to either species is unclear and the validity of M. longicollum has been questioned and debated. Here we describe a largely complete specimen of M. longicollum discovered in 1959 in Atouguia da Baleia, Peniche, Portugal, consisting of both anterior and posterior portions of the skeleton. Comparisons to the holotypes of dacentrurines and other stegosaurs shed light on the convoluted relationships of this group. We conclude that M. longicollum is valid and rather different from D. armatus, and provide a revised diagnosis of M. longicollum, as well as revised diagnoses for D. armatus, Dacentrurinae, and the first diagnosis of the genus Miragaia, granting stability to these taxa and allowing new considerations to be given on the classification of other Iberian stegosaurs. This new specimen is, to date, the most complete dinosaur described from Portugal and the most complete stegosaur described from Europe. Miragaia shared anatomical features that show a close affinity to Alcovasaurus longispinus, confirming this to be the first known dacentrurine stegosaur in America, coherent with the hypothesis of an ephemeral land bridge between North America and Iberia that allowed faunal exchange.