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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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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.

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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 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.

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.

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.

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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.}

Three new additional skull specimens of Talarurus plicatospineus have been recovered from the Upper Cretaceous (Cenomanian–Santonian) Bayanshiree Formation, of Bayan Shiree cliffs, eastern Gobi Desert, Mongolia. The skulls feature unique characters such as an anteriorly protruded single internarial caputegulum, around 20 flat or concave nasal-area caputegulae surrounded by a wide sulcus, a vertically oriented elongate loreal caputegulum with a pitted surface, an elongate lacrimal caputegulum positioned above the posterodorsal border of the maxilla, two longitudinally arranged large frontoparietal caputegulae surrounded by smaller rhomboid caputegulae, small but elongate medial supraorbital caputegulae, a posterior supraorbital caputegulum that is four times larger than the anterior one, up to three transverse parallel grooves on the dorsal surface of the posterior supraorbital caputegulum, postocular caputegulae along the ventral to posterior rim of the orbit that extend almost to the anteroventral margin of the squamosal horn, a longitudinal furrow tapering towards the apex of the squamosal horn, a lateral nuchal caputegulum four to five times larger than other nuchal caputegulae, and a pterygovomerine keel with a ventral margin that is dorsally positioned to the alveolar ridge. The phylogenetic analysis result showed that Talarurus is sister to the clade that includes the derived Asian ankylosaurines (Saichania chulsanensis, Tarchia kielanae, and Zaraapelta nomadis). It also shows that there was dispersal of ankylosaurines from Asia into western North America before the Cenomanian. Moreover, the rostral differences between Talarurus and Tsagantegia, another ankylosaur from the same formation, suggest possible niche partitioning between these taxa.

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{We here describe an articulated partial skeleton of a small neosuchian crocodylomorph from the Lourinhã Formation (Late Jurassic, Portugal). The skeleton corresponds to the posterior region of the trunk and consists of dorsal, ventral and limb osteoderms, dorsal vertebrae, thoracic ribs and part of the left hindlimb. The paravertebral armour is composed of two rows of paired osteoderms with the lateral margins ventrally deflected and an anterior process for a ‘peg and groove’ articulation. We also compare its dermal armour with that of several Jurassic and Cretaceous neosuchian crocodylomorphs, establishing a detailed description of this type of osteoderms.These features are present in crocodylomorphs with a closed paravertebral armour bracing system. The exceptional 3D conservation of the specimen, and the performance of a micro-CT scan, allowed us to interpret the bracing system of this organism to assess if previous models were accurate. The characters observed in this specimen are congruent with Goniopholididae, a clade of large neosuchians abundant in most semi-aquatic ecosystems from the Jurassic and Early Cretaceous of Laurasia. However, its small size, contrasted with the sizes observed in goniopholidids, left indeterminate whether it could have been a dwarf or juvenile individual. Future histological analyses could shed light on this.}

This chapter provides an overview of the alpha paleobiodiversity of Angola based on the available fossil record that is limited to the sedimentary rocks, ranging in age from Precambrian to the present. The geological period with the highest paleobiodiversity in the Angolan fossil record is the Cretaceous, with more than 80{%} of the total known fossil taxa, especially marine molluscs, including ammonites as a majority among them. The vertebrates represent about 15{%} of the known fauna and about one tenth of them are species firstly described based on specimens from Angola.

Este capítulo apresenta uma visão geral da paleobiodiversidade alfa de Angola com base no registo fóssil disponível, o qual se limita às rochas sedimentares, a sua idade variando entre o Pré‑Câmbrico e o pre‑
sente. O período geológico com a maior paleobiodiversidade no registo fóssil angolano é o Cretácico, com mais de 80% do total dos táxones fósseis conhecidos, especialmente moluscos marinhos, sendo estes na sua maioria
amonites. Os vertebrados representam cerca de 15% da fauna conhecida e cerca de um décimo destes são espécies descritas pela primeira vez com base em espécimes de Angola.

This chapter provides an overview of the alpha paleobiodiversity of Angola based on the available fossil record that is limited to the sedimentary rocks, ranging in age from Precambrian to the present. The geological period with the highest paleobiodiversity in the Angolan fossil record is the Cretaceous, with more than 80{%} of the total known fossil taxa, especially marine molluscs, including ammonites as a majority among them. The vertebrates represent about 15{%} of the known fauna and about one tenth of them are species firstly described based on specimens from Angola.

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Abstract We report here new and the first mammaliamorph tracks from the Early Cretaceous of Africa. The tracksite, that also bears crocodylomorph and sauropod dinosaurian tracks, is in the Catoca diamond mine, Lunda Sul Province, Angola. The mammaliamorph tracks have a unique morphology, attributed to Catocapes angolanus ichnogen. et ichnosp. nov. and present an anterolateral projection of digit I and V. The tracks with an average length of 2.7 cm and width of 3.2 cm are the largest mammaliamorph tracks known from the Early Cretaceous unmatched in size in the skeletal fossil record. The crocodylomorph trackways and tracks are attributed to Angolaichnus adamanticus ichnogen. et ichnosp. nov. (‘ichnofamily’ Batrachopodidae) and present a functionally pentadactyl pes, an extremely outwardly rotated handprint, and an unusual tetradactyl and plantigrade manus. One medium-sized sauropod dinosaur trackway preserved skin impressions of a trackmaker with stride length of 1.6 m; a second is that of a small-sized sauropod trackmaker with a pace length of 75 cm.

A new stem cheloniid turtle, Cabindachelys landanensis, gen. et sp. nov., is represented by a nearly complete skull and partial hyoid collected in lower Paleocene shallow marine deposits, equivalent to the offshore Landana Formation, near the town of Landana in Cabinda, Angola. A partial chelonioid carapace previously reported from this locality is referred here to C. landanensis. Cabindachelys landanensis possesses clear synapomorphies of Pan-Cheloniidae, including a rod-like rostrum basisphenoidale, V-shaped basisphenoid crest, and secondary palate, but also retains a reduced foramen palatinum posterius, unlike most other pan-cheloniids. Phylogenetic analysis suggests that C. landanensis forms a weakly-supported clade with Erquelinnesia gosseleti, Euclastes acutirostris, Euclastes wielandi and Terlinguachelys fischbecki, although a close relationship between the protostegid T. fischbecki and these durophagous pan-cheloniids is unlikely. The Paleocene–Eocene strata near Landana have produced a number of turtle fossils, including the holotype specimen of the pleurodire Taphrosphys congolensis. A turtle humerus collected c. 1 m above the holotype skull of C. landanensis differs from humeri of chelonioids and Taphrosphys, indicating that a third turtle taxon is present at Landana. Cheloniid fossil material is rare in the Landana assemblage, in comparison with the abundant remains of Taphrosphys congolensis found throughout the stratigraphic section. This disparity implies that C. landanensis preferred open marine habitats, whereas Taphrosphys congolensis spent more time in nearshore environments. The appearance of new durophagous species such as C. landanensis in the early Paleocene reflects the rapid radiation of pan-cheloniids as they diversified into open niches following the K–Pg extinction.

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