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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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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.}

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.

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The geochemical signatures of dinosaur eggshells represent well-established proxies in paleoenvironmental and paleobiological research. The variable sampling procedures reported in the literature, however, deserve attention. In order to evaluate the impact of different sampling methodologies on carbon and oxygen isotope and elemental concentrations, grinding was contrasted with drilling to extract powder samples from eggshell fragments collected at several locations. Eggshell data were further contrasted with surface materials, encasing matrix and compared with independent proxies using petrographic and elemental techniques. Iron and manganese elemental concentrations revealed an enrichment sequence depending on the sampling strategy for the same eggshell fragment. This pattern can be mistaken for a variable state of preservation. In contrast, carbon and oxygen isotope values exhibited only subtle differences and lacked clear trends. This suggests that isotope data are less susceptible to different methodological approaches. It is shown that drilling offers a wider range of possibilities compared to grinding (e.g., faster and less destructive). Additionally, drilled powder samples can confidently be used for elemental and isotope analysis, excluding contamination, thus providing a more accurate set of proxy data from eggshell archives.

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ABSTRACTVirtual palaeontology is a growing field, leading palaeontologists to better understand the microanatomy of many extinct species. The application of techniques such as CT and μCT-scanning allows the researchers to study micro-anatomical features in a non-invasive way and make inferences on the palaeobiology of animals. Dinosaurs have been extensively studied using these techniques, with particular focus on the microanatomy of the cranium, whereas relatively little is known of other cranial elements, such as the lower jaw. Here, we aim to fill this gap, describing the microanatomy of the specimen ML 768, an isolated dentary belonging to a dryosaurid iguanodontian dinosaur from the Upper Jurassic of Lourinhã Fm. The dentary ML 768 was subjected to μCT-scanning, and subsequently the data were segmented in Avizo and rendered in Blender. We identified functional and replacement teeth, recognising remnants of old replacement cycles. Furthermore, we mapped a rich neurovascular network present in the dentary and compared it with reference literature. We found that the high vascularisation is shared with other cerapodan dinosaurs with high tooth replacement rates, although homoeostasis may have also played a role in the development of this condition. Further evidence is needed to appreciate the macroevolutionary significance of these findings.

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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.

An incomplete, yet remarkably-sized dentated rostrum and associated partial cervical vertebrae of a pterosaur (ML 2554) were recently discovered from the Late Jurassic (Late Kimmeridgian-Early Tithonian) Lourinhã Formation of Praia do Caniçal, of central west Portugal. This specimen exhibits features such as a spatulated anterior expansion of the rostrum, robust comb-like dentition, and pronounced rims of the tooth alveoli, indicating gnathosaurine affinities. Based on its further unique tooth and dentary morphology, a new genus and species, \textit{Lusognathus almadrava} gen. et spec. nov., is proposed, making this the first named pterosaur species found within Portugal. The presence of this taxon adds yet another element to the fluvio-deltaic lagoonal environment that has been suggested as representative of the Lourinhã Formation in the Late Jurassic, further contributing to the diversity and distribution of gnathosaurines worldwide.

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O sítio de Silveirinha é uma das localidades de mamíferos mais conhecidas da Paleontologia do Cenozoico de Portugal e da Europa em geral. Graças à sua rica e diversificada associação de mamíferos, com mais de 30 taxa, foi posicionado no Eocénico inferior (início do Ypresiano, MP7, ca. 55,8 M.a.), sendo o local mais antigo da Europa desta Época, devido à presença única de taxa típicos do Paleocénico superior, juntamente com outras espécies já características do Eocénico inferior. Este estudo irá rever o material de pequenos mamíferos deste sítio, conservado na coleção clássica da Universidade Nova de Lisboa, a fim de fazer uma actualização taxonómica à luz das publicações mais recentes.

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.

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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.

The jigsaw-puzzle fit of South America and Africa is an icon of plate tectonics and continental drift. Fieldwork in Angola since 2002 allows the correlation of onshore outcrops and offshore geophysical and well-core data in the context of rift, sag, salt, and post-salt drift phases of the opening of the central South Atlantic. These outcrops, ranging in age from >130 Ma to <71 Ma, record Early Cretaceous outpouring of the Etendeka-Paraná Large Igneous Province (Bero Volcanic Complex) and rifting, followed by continental carbonate and siliciclastic deposition (Tumbalunda Formation) during the sagging of the nascent central South Atlantic basin. By the Aptian, evaporation of sea water resulted in thick salt deposits (Bambata Formation), terminated by sea floor spreading. The Equatorial Atlantic Gateway began opening by the early Late Cretaceous (100 Ma) and allowed flow of currents between the North and South Atlantic, creating environmental conditions that heralded the introduction of marine reptiles. These dramatic outcrops are a unique element of geoheritage because they arguably comprise the most complete terrestrially exposed geological record of the puzzle-like icon of continental drift.

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Allosaurus Marsh, 1877 is one of the most iconic theropods of the Late Jurassic of USA and Portugal. However,
since the first description the genus was used as taxonomical wastebasket to which several species were
described. We revisit three type specimens, Allosaurus fragilis YPM 1930, A. atrox YPM 1890 and A. ferox USNM
2315. We interpret the vertebral material of A. atrox and support it as a junior synonym of A. fragilis, and A. ferox
as Allosaurus indet. rather than A. fragilis.

Turiasauria is a clade of basal sauropod dinosaurs hitherto only known from the Middle Jurassic (Bathonian) to the Lower Cretaceous (Valanginian). A new find of a shed tooth crown from the Lower Jurassic (Pliensbachian), Halse Formation of Bornholm, Denmark, is spoon-like, asymmetrical, and heart-shaped, which identifies the tooth as turiasaurian, pushing the origin of the Turiasauria some 17 My back into the Lower Jurassic. This suggests a North Pangean/Laurasian origin of the turiasaurian clade, which then, during the Middle to Late Jurassic, dispersed through Europe, India, and Africa, with their latest representatives found in the Early Cretaceous of England and North America. Furthermore, this is the first record of a sauropod from the Pliensbachian in Europe.

Iguanodontia is a diverse clade of herbivorous ornithischian dinosaurs that were speciose and abundant during the Jurassic and Cretaceous. Although the monophyly of Iguanodontia is well supported, their internal relationships have sparked heated debate due to several phylogenetic paradigm shifts. Late Jurassic basally branching iguanodontians in particular are not well understood in terms of their systematic affinities and evolutionary relevance. Their fossil record in Europe is meager compared with North America, with only a few species currently recognized. Two taxa are currently known from the Upper Jurassic of England, the basally branching styracosternan Cumnoria prestwichii and the putative dryosaurid Callovosaurus leedsi. In the Upper Jurassic of Portugal, the styracosternan Draconyx loureiroi and the dryosaurid Eousdryosaurus nanohallucis are presently the only described basally branching iguanodontians. Here we report a new species of early diverging iguanodontian from the Upper Jurassic Lourinhã Formation of western-central Portugal. The new species is clearly distinguished from all other coeval taxa by an exclusive combination of characters that include a tibia with a cnemial crest that is directed craniolaterally and a fibular condyle that is angled at 90° with respect to the proximal epiphysis, a fibula with symmetrical proximal margins, and a reduced metatarsal I. The phylogenetic relationships of the Lourinhã iguanodontian were explored using maximum parsimony and Bayesian inference. The two analyses recover the Lourinhã iguanodontian as an indeterminate dryomorphan, with more precise affinities precluded due to the current available material. Body size is estimated between 3 and 4 meters for the holotype specimen, adding to the diversity of small ornithopods already recognized in the paleoichnological record of the Lourinhã Formation. http://zoobank.org/urn:lsid:zoobank.org:pub:5F4D52D0-0F0B-4809-8561-BE58C7C97D45