NEW SPECIMENS OF ANGOLASAURUS BOCAGEI AND COMMENTS ON THE EARLY RADIATIONS OF PLIOPLATECARPINE MOSASAURS POLCYN, Michael, SMU, Dallas, TX, USA; JACOBS, Louis, SMU, Dallas, TX, USA; MATEUS, Octávio, Museu da Lourinhã, Lourinhã, Portugal; SCHULP, Anne, Natuurhistorisch Museum Maastricht, Maastricht, Netherlands New, well preserved material of the Turonian mosasaur Angolasaurus bocagei from the Tadi Beds of the Itombe Formation in northern Angola, allows detailed redescription of its morphology and reassessment of its phylogenetic relationships. Angolasaurus had been previously referred to the genus Platecarpus; however, phylogenetic analysis confirms the valid taxonomic status of A. bocagei, and reconstructs that taxon within a clade that also includes the genera Selmasaurus and Ectenosaurus. These forms are united by an elaborated infrastapedial process of the quadrate and a unique ridge-like descending process of the parietal forming the supraoccipital articulation, but also retain a relatively plesiomorphic configuration of the braincase. That clade is united with all other plioplatecarpines by a number of derived characters including the presence of a novel basicranial circulation pattern. In Africa, North and South America, early plioplatecarpines are known by the Middle Turonian and Angolasaurus and closely related forms appear by the Upper Turonian. Selmasaurus and Ectenosaurus are a rare faunal component of the Santonian and Campanian of North America. Platecarpus planifrons appears in the Coniacian of North America and represents the plesiomorphic condition of the clade containing the remaining species of Platecarpus and Plioplatecarpus, that appears in the Santonian and persist until the end of the Cretaceous, reaching global distribution. The temporal and geographic distribution of these radiations suggest influence of paleogeography and eustatic sea levels.

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Although the Late Jurassic of Portugal has provided abundant dinosaur fossils, material from the Early Cretaceous is scarce. This paper reports new cranial and postcranial material of the theropod dinosaur Baryonyx walkeri found in the Barremian (Papo Seco Formation) of Portugal. This specimen, found at Praia das Aguncheiras, Cabo Espichel, consists of a partial dentary, isolated teeth, pedal ungual, two calcanea, presacral and caudal vertebrae, fragmentary pubis, scapula, and rib fragments. It represents the most complete spinosaurid yet discovered in the Iberian Peninsula and the most complete dinosaur from the Early Cretaceous of Portugal. This specimen is confidently identified as a member of Baryonychinae due to the presence of conical teeth with flutes and denticles in a dentary rosette. The specimen ML1190 shares the following characteristics with Baryonyx walkeri: enamel surface with small (nearly vertical) wrinkles, variable denticle size along the carinae, 6–7 denticles per mm, wrinkles forming a 45 degree angle near the carinae, and tooth root longer than crown. In addition, dubious taxa based on teeth morphology such as Suchosaurus cultridens (Owen, 1840–1845), and Suchosaurus girardi (Sauvage 1897–98; Antunes & Mateus 2003) are discussed, based on comparisons with well-known material such as Baryonyx walkeri Charig & Milner, 1986. Suchosaurus cultridens and S. girardi are considered as nomina dubia due to the lack of diagnostic apomorphies, but both specimens are referred to Baryonychinae incertae sedis.

The end-Triassic extinction event (ETE), considered one of the ‘Big Five’ mass extinctions, marks a dividing line between early Mesozoic vertebrate assemblages, typically including abundant temnospondyls, basal synapsids and basal archosaurs, and ‘typical’ Mesozoic faunas dominated by dinosaurs, pterosaurs, crocodylomorphs, turtles and mammaliaforms. Recent geochemical work has provided strong evidence that the ETE is synchronous with, and likely caused by, the emplacement of the Central Atlantic magmatic province (CAMP). However, stratigraphic sections containing both terrestrial vertebrates and CAMP basalts are scarce, complicating attempts to examine terrestrial faunal changes during this extinction event. The Triassic–Jurassic Algarve Basin, southern Portugal, is an extensional rift basin to-marginal marine red beds (the ‘Grés de Silves’ Group) interbedded with CAMP basalts.... bonebed from the interval ‘AB1’ of the Grés de Silves. Preliminary excavations yielded at least nine well-preserved temnospondyl individuals represented by partial to nearly complete skulls and disarticulated postcranial elements of juvenile to adult ages. Nearly all material appears to represent a single species of metoposaurid referable to the genus Metoposaurus, well known from the late Carnian–early Norian of Germany and Poland. A number of char- acters of the occiput and mandible suggest that the Algarve material may represent a new species. This new material provides new data on the diversity and paleogeographical distri- bution of the metoposaurids, a highly autapomorphic and peculiar group composed of large aquatic carnivores with a unique elongated but brevirostral skull. This taxon also provides [...] Horizon may be within or close to the late Carnian–early Norian. Additional bone-bearing horizons within the ‘Grés de Silves’ provide a rare opportunity to examine terrestrial faunal change in the lead-up to the ETE.

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.

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Eleven new tracks from the Upper Jurassic of Portugal are described and attributed to the stegosaurian ichnogenus Deltapodus. One track exhibits exceptionally well−preserved impressions of skin on the plantar surface, showing the stegosaur foot to be covered by closely spaced skin tubercles of ca. 6 mm in size. The Deltapodus specimens from the Aalenian of England represent the oldest occurrence of stegosaurs and imply an earlier cladogenesis than is recognized in the body fossil record.

Stegosaurian dinosaurs have a quadrupedal stance, short forelimbs, short necks, and are generally considered to be low browsers. A new stegosaur, Miragaia longicollum gen. et sp. nov., from the Late Jurassic of Portugal, has a neck comprising at least 17 cervical vertebrae. This is eight additional cervical vertebrae when compared with the ancestral condition seen in basal ornithischians such as Scutellosaurus.
Miragaia has a higher cervical count than most of the iconically long-necked sauropod dinosaurs. Long neck length has been achieved by ‘cervicalization’ of anterior dorsal vertebrae and probable lengthening of centra. All these anatomical features are evolutionarily convergent with those exhibited in the necks of
sauropod dinosaurs. Miragaia longicollum is based upon a partial articulated skeleton, and includes the only known cranial remains from any European stegosaur. A well-resolved phylogeny supports a new clade that unites Miragaia and Dacentrurus as the sister group to Stegosaurus; this new topology challenges the common view of Dacentrurus as a basal stegosaur.

Portugal is rich on dinosaur remains (bones, eggs, and tracks) from Early Jurassic to Late
Cretaceous ages, but mainly from the Late Jurassic, in which dozen of tracksites have been reported.
Here are reported new or poorly known track localities:
1) Five tracksites share the preservation substrate (marine carbonated limestone), age (late Jurassic), geographic area (Leiria district of Portugal), kind of preservation (true tracks), and completeness (trackways of multiple individuals):
i) Praia dos Salgados includes eight trackways, mostly ornithopods and theropods, and one wide gauge sauropod, made in very soft sediment; some preserve the hallux impression.
ii) Serra de Mangues is mostly covered with vegetation but seems to include dozens of tracks comprising theropods, thyreophorans, ornithopods and sauropods.
iii) Sobral da Lagoa (Pedreira do Rio Real) include six trackways but poorly preserved;
and
iv) Serra de Bouro that preserves four sauropod trackways in one single layer.
v) Pedrógão, preserved, at least, one theropod trackway and several isolated tracks of
theropods and ornithopods were found in different layers in the Early Oxfordian.
2) The locality in Praia de Porto das Barcas yielded natural casts of stegosaur tracks
(including pes print with skin impression) and a very large sauropod pes print with about
1.2 m long pes.
3) A new pterosaur tracksite was found in the Late Jurassic of Peralta, Lourinhã (Sobral Member, Lourinhã Fm.; Late Kimmeridgian/Early Tithonian). More than 220 manus and pes tracks have been collected in about five square meters, all ascribed to pterosaurs. The tracks were produced in a thin mud layer that has been covered by sand which preserved them as sandstone mould infill (natural casts). The manus of the largest specimens is 13 cm wide and 5.5 cm long and the pes measures 14.5 cm in length and 9 cm in width. This shows the occurrence of very large pterosaurs in the Late Jurassic. Other pterosaur tracksites in the Late Jurassic of Portugal are: Porto das Barcas (Lourinhã Municipality), South of Consolação (Peniche Municipality), and Zambujal de Baixo (Sesimbra Municipality).

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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.

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.

We report a new chelonioid turtle on the basis of a nearly complete skull collected in lower Paleocene, shallow marine deposits, equivalent to the offshore Landana Formation, near the town of Landana in Cabinda Province, Angola. Chelonioid material previously reported from this locality is likely referable to this new taxon. The well-preserved skull is missing the left quadrate, squamosal, and prootic, both opisthotics, and the mandible. The skull possesses a rod-like basisphenoid rostrum, which is a synapomorphy of Chelonioidea, but it differs from other chelonioid skulls in that the contact between the parietal and squamosal is absent, and the posterior palatine foramen is present. Phylogenetic analysis recovers the new taxon as a basal chelonioid. The Paleocene– Eocene strata near Landana have produced a wealth of turtle fossils, including the holotype of the pleurodire Taphrosphys congolensis. A turtle humerus collected from the Landana locality differs morphologically from the humeri of chelonioids and Taphrosphys, indicating the presence of a third taxon. Chelonioid fossil material in the Landana assemblage is rare compared to the abundant fragmentary remains of Taphrosphys that are found throughout the stratigraphic section. This disparity in abundance suggests the new chelonioid taxon preferred open marine habitats, whereas Taphrosphys frequented nearshore environments.

We report a new chelonioid turtle on the basis of a nearly complete skull collected in lower Paleocene, shallow marine deposits, equivalent to the offshore Landana Formation, near the town of Landana in Cabinda Province, Angola. Chelonioid material previously reported from this locality is likely referable to this new taxon. The well-preserved skull is missing the left quadrate, squamosal, and prootic, both opisthotics, and the mandible. The skull possesses a rod-like basisphenoid rostrum, which is a synapomorphy of Chelonioidea, but it differs from other chelonioid skulls in that the contact between the parietal and squamosal is absent, and the posterior palatine foramen is present. Phylogenetic analysis recovers the new taxon as a basal chelonioid. The Paleocene– Eocene strata near Landana have produced a wealth of turtle fossils, including the holotype of the pleurodire Taphrosphys congolensis. A turtle humerus collected from the Landana locality differs morphologically from the humeri of chelonioids and Taphrosphys, indicating the presence of a third taxon. Chelonioid fossil material in the Landana assemblage is rare compared to the abundant fragmentary remains of Taphrosphys that are found throughout the stratigraphic section. This disparity in abundance suggests the new chelonioid taxon preferred open marine habitats, whereas Taphrosphys frequented nearshore environments.

We report a new chelonioid turtle on the basis of a nearly complete skull collected in lower Paleocene, shallow marine deposits, equivalent to the offshore Landana Formation, near the town of Landana in Cabinda Province, Angola. Chelonioid material previously reported from this locality is likely referable to this new taxon. The well-preserved skull is missing the left quadrate, squamosal, and prootic, both opisthotics, and the mandible. The skull possesses a rod-like basisphenoid rostrum, which is a synapomorphy of Chelonioidea, but it differs from other chelonioid skulls in that the contact between the parietal and squamosal is absent, and the posterior palatine foramen is present. Phylogenetic analysis recovers the new taxon as a basal chelonioid. The Paleocenetextendash Eocene strata near Landana have produced a wealth of turtle fossils, including the holotype of the pleurodire Taphrosphys congolensis. A turtle humerus collected from the Landana locality differs morphologically from the humeri of chelonioids and Taphrosphys, indicating the presence of a third taxon. Chelonioid fossil material in the Landana assemblage is rare compared to the abundant fragmentary remains of Taphrosphys that are found throughout the stratigraphic section. This disparity in abundance suggests the new chelonioid taxon preferred open marine habitats, whereas Taphrosphys frequented nearshore environments.

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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.

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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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Here we describe a new species of the mosasaurine genus Prognathodon from the Maastrichtian of Namibe, Angola, on the basis of five specimens which represent most of the cranial skeleton including the diagnostic quadrate. Phylogenetic analysis shows this new taxon, P. kianda nov. sp., to be the sister taxon to all other species of Prognathodon. It is unique amongst Prognathodon in possessing a high marginal tooth count and relatively small pterygoid teeth. The tooth morphology in the new taxon is reminiscent of some species of the genus Liodon, allowing association of Liodon-like dentition with otherwise Prognathodon-like crania, and thus resolves the long-standing question of the phylogenetic affinities of Liodon.

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A partial plesiosauroid skull from the São Gião Formation (Toarcian, Lower Jurassic) of Alhadas, Portugal is re−evaluated and described as a new taxon, Lusonectes sauvagei gen. et sp. nov. It has a single autapomorphy, a broad triangular parasphenoid cultriform process that is as long as the posterior interpterygoid vacuities, and also a unique character combination, including a jugal that contacts the orbital margin, a distinct parasphenoid–basisphenoid suture exposed between the posterior interpterygoid vacuities, lack of an anterior interpterygoid vacuity, and striations on the ventral surface of
the pterygoids. Phylogenetic analysis of Jurassic plesiosauroids places Lusonectes as outgroup to “microcleidid elasmosaurs”, equivalent to the clade Plesiosauridae. Lusonectes sauvagei is the only diagnostic plesiosaur from Portugal, and the westernmost occurrence of any plesiosaurian in Europe.

The end-Triassic extinction event (ETE), considered one of the ‘Big Five’ mass extinctions, marks a dividing line between early Mesozoic vertebrate assemblages, typically including abundant temnospondyls, basal synapsids and basal archosaurs, and ‘typical’ Mesozoic faunas dominated by dinosaurs, pterosaurs, crocodylomorphs, turtles and mammaliaforms.
Recent geochemical work has provided strong evidence that the ETE is synchronous with, and likely caused by, the emplacement of the Central Atlantic magmatic province (CAMP).
However, stratigraphic sections containing both terrestrial vertebrates and CAMP basalts are scarce, complicating attempts to examine terrestrial faunal changes during this extinction event. The Triassic–Jurassic Algarve Basin, southern Portugal, is an extensional rift basin

to-marginal marine red beds (the ‘Grés de Silves’ Group) interbedded with CAMP basalts.

bonebed from the interval ‘AB1’ of the Grés de Silves. Preliminary excavations yielded at least nine well-preserved temnospondyl individuals represented by partial to nearly complete skulls and disarticulated postcranial elements of juvenile to adult ages. Nearly all material appears to represent a single species of metoposaurid referable to the genus Metoposaurus, well known from the late Carnian–early Norian of Germany and Poland. A number of characters of the occiput and mandible suggest that the Algarve material may represent a new species. This new material provides new data on the diversity and paleogeographical distribution of the metoposaurids, a highly autapomorphic and peculiar group composed of large aquatic carnivores with a unique elongated but brevirostral skull. This taxon also provides

Horizon may be within or close to the late Carnian–early Norian. Additional bone-bearing horizons within the ‘Grés de Silves’ provide a rare opportunity to examine terrestrial faunal change in the lead-up to the ETE.