ABSTRACTHerein we describe phytosaurs from thin fluvial overbank sandstones of the Upper Triassic Malmros Klint Formation of the Fleming Fjord Group (central East Greenland). The new sample includes more than 150 disarticulated bones and teeth from small to large specimens belonging to at least four individuals. The fossils mostly consist of teeth and postcranial elements and permit the recognition of a new species of Mystriosuchus, M. alleroq, diagnosed by an L-shaped quadratojugal whose anterior suture trends anterodorsally and a tripartite degree of heterodonty. Humeral diaphyseal histology of one specimen reveals a fairly compact cortex that surrounds a cancellous medullary region followed by a remodeling zone containing scattered secondary osteons. Primary bone tissue is parallel-fibred with a moderate to low vascular density. The cortex is cyclically interrupted by distinct growth marks indicating a seasonal environment. A change in growth rate from moderate to low is documented within the outer cortex, indicating that at least this individual was close to somatic maturity. Mystriosuchus has formerly been known as an exclusively European taxon. The new findings support the European faunal influence in East Greenland during the Late Triassic inferred from other taxa such as temnospondyls and archosaurs. The mid-late Norian age of European Mystriosuchus suggests an additional age constraint for the vertebrate-bearing portion of the Malmros Klint Formation.

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Abstract We present the first detailed braincase anatomical description and neuroanatomical study of Portugalosuchus azenhae, from the Cenomanian (Late Cretaceous) of Portugal. This eusuchian crocodylomorph was originally described as a putative Crocodylia and one of the oldest representatives of this clade; however, its phylogenetic position remains controversial. Based on new data obtained from high resolution Computed Tomography images (by micro-CT scan), this study aims to improve the original description of this taxon and also update the scarce neuroanatomical knowledge of Eusuchia and Crocodylia from this time interval, a key period to understand the origin and evolution of these clades. The resulting three-dimensional models from the CT data allowed a detailed description of its well-preserved neurocranium and internal cavities. Therefore, it was possible to reconstruct the cavities of the olfactory region, nasopharyngeal ducts, brain, nerves, carotid arteries, blood vessels, paratympanic sinus system and inner ear, which allowed to estimate some neurosensorial capabilities. By comparison with other crocodylomorphs, these analyses showed that Portugalosuchus, back in the Cenomanian, already displayed an olfactive acuity, sight, hearing and cognitive skills within the range of that observed in other basal eusuchians and crocodylians, including extant species. In addition, and in order to test its disputed phylogenetic position, these new anatomical data, which helped to correct and complete some of the original observations, were included in one of the most recent morphology-based phylogenies. The position of Portugalosuchus differs slightly from the original publication since it is now located as a “thoracosaurid” within Gavialoidea, but still as a crocodylian. Despite all this, to better contrast these results, additional phylogenetic analyses including this new morphological character coding together with DNA data should be performed.

Ostriches are known to be the fastest bipedal animal alive; to accomplish such an achievement, their anatomy evolved to sustain the stresses imposed by running at such velocities. Ostriches represent an excellent case study due to the fact that their locomotor kinematics have been extensively studied for their running capabilities. The shape and structure of ostrich bones are also known to be optimized to sustain the stresses imposed by the body mass and accelerations to which the bones are subjected during movements. This study focuses on the limb bones, investigating the structure of the bones as well as the material properties, and how both the structure and material evolved to maximise the performance while minimising the stresses applied to the bones themselves. The femoral shaft is hollowed and it presents an imbricate structure of fused bone ridges connected to the walls of the marrow cavity, while the tibial shaft is subdivided into regions having different mechanical characteristics. These adaptations indicate the optimization of both the structure and the material to bear the stresses. The regionalization of the material highlighted by the mechanical tests represents the capability of the bone to adapt to external stimuli during the life of an individual, optimizing not only the structure of the bone but the material itself.

Sauropod dinosaurs are well known for their massive sizes and long necks and tails. Among sauropods, flagellicaudatan dinosaurs are characterized by extreme tail elongation, which has led to hypotheses regarding tail function, often compared to a whip. Here, we analyse the dynamics of motion of a 3D model of an apatosaurine flagellicaudatan tail using multibody simulation and quantify the stress-bearing capabilities of the associated soft tissues. Such an elongated and slender structure would allow achieving tip velocities in the order of 30 m/s, or 100 km/h, far slower than the speed of sound, due to the combined effect of friction of the musculature and articulations, as well as aerodynamic drag. The material properties of the skin, tendons, and ligaments also support such evidence, proving that in life, the tail would not have withstood the stresses imposed by travelling at the speed of sound, irrespective of the conjectural ‘popper’, a hypothetical soft tissue structure analogue to the terminal portion of a bullwhip able to surpass the speed of sound.

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

Spinosaurids are some of the most enigmatic Mesozoic theropod dinosaurs due to their unique adaptations to aquatic environments and their relative scarcity. Their taxonomy has proven to be especially problematic. Recent discoveries from Western Europe in general, specifically Iberia, provide some of the best specimens for the understanding of their phylogeny, leading to the description of the spinosaurid Vallibonavenatrix cani and the recognition of the Iberian dinosaur Camarillasaurus cirugedae as one of them. Portuguese associated spinosaurid remains (ML1190) from the Papo Seco Formation (early Barremian) were previously assigned to Baryonyx walkeri but new material recovered in 2020 along with new phylogenetic analyses suggests a different phylogenetic placement, making their revision necessary. Here we show that these remains are not attributable to Baryonyx walkeri, but to a new genus and species, Iberospinus natarioi, gen. et sp. nov. The new taxon is characterized by the presence of a single Meckelian foramen in the Meckelian sulcus, a straight profile of the ventral surface of the dentary and a distal thickening of the acromion process of the pubis between other characters. Iberospinus natarioi is recovered as a sister taxon of the clade formed by Baryonyx and Suchomimus, and outside Spinosaurinae when Vallibonaventrix cani is excluded from the analysis. The description of this taxon reinforces Iberia as a hotspot for spinosaur biodiversity, with several endemic taxa for the region. As expected for the clade, the dentary displays a highly vascularized neurovascular network. The morphometric analysis of parts of the skeleton (pedal phalanx and caudal vertebrae, among others) shows an intermediate condition between basal tetanurans and spinosaurines.

{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.}

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The Algar do Vale da Pena is a cave that was discovered during limestone mining works around 1980, at the Natural Park of Serras de Aire e Candeeiros, Portugal. The entrance to the cave, through the collapsed roof, is a 30 meter near-vertical well, inaccessible without means of vertical progression and specialized equipment. Soon after the discovery of the cave, bone material belonging to bears and hundreds of bear claw marks were located inside it. Nevertheless, the technical descent and the extraordinary difficulties of retrieving the calcite flowstones that encased the bones prevented scientists from undertaking excavations for nearly 40 years. Since 2016 a series of expeditions by a team of paleontologists from NOVA University and Museum of Lourinhã recovered more than 20 disarticulated remains from five bone accumulations, separated more than five meters apart, which given the level of erosion and breakage, are para-autochthonous in origin. These are interpreted as belonging to different individuals. Morphological characters, like a relatively elongated third phalanx, the clear separation of paracone-metacone of the first upper molar and the ventral position of the mandibular condyle; as well as numerous morphometric characters, allow to ascribe the remains to Ursus arctos. Nevertheless, the main finding from the cave is a complete skull with an articulated mandible; which denotes absence of transport, encased within a 180 kg calcite flowstone, including other skeletal remains as well, which could not be extracted from the cave until 2021. This block was removed with the help of members of several speleological groups. It promises a wealth of information about fossil brown bears, being the most complete bear fossil ever recovered in Portugal. It is currently being dated trough magnetostratigraphy and prepared. We consider this intervention as an extraordinary example of citizen science applied to paleontology

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ABSTRACT Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

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ABSTRACTCrocodylomorph eggs and eggshells are known as old as the Late Jurassic and are frequent components of most multiootaxic eggshell assemblages. Classified within the oofamily Krokolithidae, thei histo- and ultrastructures are conservative throughout geological time, characterised by inverted-trapezoid-shaped shell units that grow from highly spaced basal knobs and present a diagnostic tabular ultrastructure. Here, we report 327 eggshell fragments from a new fossil site from the Maastrichtian of the Southern Pyrenees, Veracruz 1, and erect a new oogenus and oospecies, Pachykrokolithus excavatum oogen. et oosp. nov. characterised by crocodyloid morphotype and a prominent rugosocavate ornamentation. Eggshells from the slightly older locality of Blasi 2b, previously reported as aff. Krokolithidae, are also assigned to this new ootaxon. Different crocodylomorph taxa coexisted during the Late Cretaceous of the Tremp Basin, hindering the attribution of Pachykrokolithus excavatum oogen. et oosp. nov. to a single clade. Nevertheless, allodaposuchid eusuchians were dominant in this ecosystem, and are the most probable producers of Pachykrokolithus excavatum oogen. et oosp. nov. eggs.

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Here, we describe the first pterosaur remains from Angola, an assemblage of fourteen bones from the Lower Maastrichtian marine deposits of Bentiaba, Namibe Province. One new species is introduced, Epapatelo otyikokolo, gen. et sp. nov., which comprises an articulated partial left humerus and ulna as well as an articulated left ulna and radius (from a second individual). Phylogenetic analysis confirms a non-nyctosaurid pteranodontian attribution for this new taxon and supports a new apomorphy-based clade, Aponyctosauria, which is here defined. Late Cretaceous pteranodontians are rare in Sub-Saharan Africa and throughout the Southern Hemisphere. Preliminary histological analysis also reveals a likely sub-adult age for one of the specimens. This fossil assemblage provides a first glimpse of Angolan pterosaur paleobiodiversity providing further insight into the Gondwanan ecosystems of the Upper Cretaceous.

The record of Late Jurassic stegosaur tracks from the Lourinhã Formation (Kimmeridgian-Tithonian) is here revised. Thirty-eight dinosaur tracks, preserved as natural infill casts, are here reported, and thirty-two of them are attributed to the ichnogenus Deltapodus. Four of those present impressions of skin, with polygonal scales and random pattern. Deltapodus is the most common ichnogenus in the track record of the Lourinhã Formation. The sizes and shape suggest one single dacentrurine trackmaker, which could be Miragaia longicollum, also common in the same horizons.

The taxonomy, chronology and dispersal patterns of hippopotamuses in Western Europe are still a matter of debate, despite the long history of researchers on these topics. In this framework, the Portuguese material has never been investigated or recently revised, and it can contribute to increasing our comprehension about Quaternary hippopotamuses. The material considered in this work, collected from sites dated between 1 Ma and 0.13 Ma, is compared with fossil Pleistocene hippopotamuses from the Iberian Peninsula, Italy, Germany and Greece, in order to contribute into the discussion on evolutionary and dispersal patterns of these large semi-aquatic mammals in Europe during the Pleistocene. Portugal deposits recorded the presence of H. antiquus, starting from 1 Ma up to 0.4 Ma, and H. cf. amphibius during the late Middle Pleistocene. In particular, the occurrence of H. antiquus in Condeixa, dated around 0.4 Ma, can be considered as the LAD of this species in Europe. The paleobiogeography and dispersal patterns of European hippopotamuses are further discussed.

ABSTRACT: Type specimens (holotypes, neotypes, syntypes, etc.) are of crucial importance because they are the only tangible evidence of the nomenclatural act that supports the understanding of paleobiodiversity. The list of the vertebrate species whose type specimen is based on fossils from Portugal is presented here. We counted 206 species, of which there are 45 bony fishes, 39 Late Jurassic Mammaliamorpha, 33 Cenozoic mammals, 28 non-avian dinosaurs, 25 non-dinosaur reptiles, 11 Cenozoic Aves, 22 Chondrichthyes and three other vertebrates. Except for the Quaternary fossil record, the type specimens can be used as a shortcut for measuring the fossil record and paleobiodiversity through geological time and rock units because they correlate in 95% with the fossil record by epoch in the case-study of fossil vertebrates of Portugal.

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A remarkably well-preserved, almost complete and articulated new specimen (GP/2E 9266) of Tupandactylus navigans is here described for the Early Cretaceous Crato Formation of Brazil. The new specimen comprises an almost complete skeleton, preserving both the skull and post-cranium, associated with remarkable preservation of soft tissues, which makes it the most complete tapejarid known thus far. CT-Scanning was performed to allow the assessment of bones still covered by sediment. The specimen can be assigned to Tupa. navigans due to its vertical supra-premaxillary bony process and short and rounded parietal crest. It also bears the largest dentary crest among tapejarine pterosaurs and a notarium, which is absent in other representatives of the clade. The new specimen is here regarded as an adult individual. This is the first time that postcranial remains of Tupa. navigans are described, being also an unprecedented record of an articulated tapejarid skeleton from the Araripe Basin.

We report a new specimen of the plesiosaur Cardiocorax mukulu that includes the most complete plesiosaur skull from sub-Saharan Africa. The well-preserved three-dimensional nature of the skull offers rare insight into the cranial anatomy of elasmosaurid plesiosaurians. The new specimen of Cardiocorax mukulu was recovered from Bentiaba, Namibe Province in Angola, approximately three meters above the holotype. The new specimen also includes an atlas-axis complex, seventeen postaxial cervical vertebrae, partial ribs, a femur, and limb elements. It is identified as Cardiocorax mukulu based on an apomorphy shared with the holotype where the cervical neural spine is approximately as long anteroposteriorly as the centrum and exhibits a sinusoidal anterior margin. The new specimen is nearly identical to the holotype and previously referred material in all other aspects. Cardiocorax mukulu is returned in an early-branching or intermediate position in Elasmosauridae in four out of the six of our phylogenetic analyses. Cardiocorax mukulu lacks the elongated cervical vertebrae that is characteristic of the extremely long-necked elasmosaurines, and the broad skull with and a high number of maxillary teeth (28–40) which is characteristic of Aristonectinae. Currently, the most parsimonious explanation concerning elasmosaurid evolutionary relationships, is that Cardiocorax mukulu represents an older lineage of elasmosaurids in the Maastrichtian.

Found in the Upper Jurassic outcrops of Lourinhã, 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.

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The Late Triassic (Norian) outcrops of the Malmros Klint Formation, Jameson Land (Greenland) have yielded numerous specimens of non-sauropod sauropodomorphs. Relevant fossils were briefly reported in 1994 and were assigned to Plateosaurus trossingensis. However, continuous new findings of early non-sauropod sauropodomorphs around the globe facilitate comparisons and allow us to now revise this material. Here, the non-sauropod sauropodomorph Issi saaneq gen. et sp. nov. is described based on two almost complete and articulated skulls. The two skulls represent a middle-stage juvenile and a late-stage juvenile or subadult. Issi saaneq differs from all other sauropodomorphs by several unique traits: (1) a small foramen at the medial surface of the premaxilla; (2) an anteroposteriorly elongated dorsoposterior process of the squamosal; (3) a relatively high quadrate relative to rostrum height; (4) a well-developed posterodorsal process of the articular. These features cannot be explained by taphonomy, ontogeny, or intraspecific variation. Issi saaneq shows affinities to Brazilian plateosaurids and the European Plateosaurus, being recovered as the sister clade of the latter in our phylogenetic analysis. It is the northernmost record of a Late Triassic sauropodomorph, and a new dinosaur species erected for Greenland. Issi saaneq broadens our knowledge about the evolution of plateosaurid sauropodomorphs.

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A new plesiosaur partial skeleton, comprising most of the trunk and including axial, limb, and girdle bones, was collected in the lower Sinemurian (Coimbra Formation) of Praia da Concha, near São Pedro de Moel in central west Portugal. The specimen represents a new genus and species, Plesiopharos moelensis gen. et sp. nov. Phylogenetic analysis places this taxon at the base of Plesiosauroidea. Its position is based on this exclusive combination of characters: presence of a straight preaxial margin of the radius; transverse processes of mid-dorsal vertebrae horizontally oriented; ilium with sub-circular cross section of the shaft and subequal anteroposterior expansion of the dorsal blade; straight proximal end of the humerus; and ventral surface of the humerus with an anteroposteriorly long shallow groove between the epipodial facets. In addition, the new taxon has the following autapomorphies: iliac blade with less expanded, rounded and convex anterior flank; highly developed ischial facet of the ilium; apex of the neural spine of the first pectoral vertebra inclined posterodorsally with a small rounded tip. This taxon represents the most complete and the oldest plesiosaur species in the Iberian Peninsula. It is also the most complete, best preserved, and oldest marine vertebrate in the region and testifies to the incursion of marine reptiles in the newly formed proto-Atlantic sea, prior to the Atlantic Ocean floor spreading in the Early Cretaceous.

The Kap Stewart Group (Rhaetian-Sinemurian, Triassic–Early Jurassic) of the Jameson Land Basin in central East Greenland has traditionally been regarded as a strictly continental unit with delta and perennial lake sediments. New finds of plesiosaur bone remain in a thin storm deposited sandstone bed in the middle part of the Rhætelv Formation of the Kap Stewart Group, however, indicates a likely period of marine influence. At the study area at the eastern margin of the basin, the Rhætelv Formation is 300-m thick and overlies unconformably the Norian Fleming Fjord Group. The bone-bearing sandstone occurs 190 m above the base of the group and is closely associated with black laminated mudstones; palynological investigation of three samples from these mudstones indicates that they are of a younger Hettangian age. The Hettangian was a relatively short stage (201.3–199.5 Ma) and elsewhere characterized by two episodes of sea-level highstands. Assuming that the marine incursion in the Jameson land Basin evidenced by the plesiosaur fossil remains took place during the youngest of these sea-level highstands, the bone-bearing bed of the Rhætelv Formation can be dated to 200 Ma and thereby gives the first numerical age constraint of this hitherto poorly dated succession.

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A large, well-preserved vertebrate coprolite was found in a lacustrine sediment in the Malmros Klint Formation of the Late Triassic Fleming Fjord Group in the Jameson Land Basin, central East Greenland. The size and internal and external morphology of the coprolite is consistent with that of crocodilian coprolites and one end of the coprolite exhibits evidence of post-egestion trampling. As the associated vertebrate fauna of the Fleming Fjord Group contains abundant remains of pseudosuchian phytosaurs, the coprolite is interpreted as being from a large phytosaur.