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.

Abstract: The paleontological collections of the Museum of Lourinhã, in Portugal, has a rich paleontological collection, particularly of Late Jurassic dinosaurs of the Lourinhã Formation (Kimmeridgian-Tithonian). Most salient highlights comprehend the following dinosaur holotype specimens: stegosaur Miragaia longicollum, theropod Lourinhanosaurus antunesi, sauropod Dinheirosaurus lourinhanensis, ornithopod Draconyx loureiroi, theropod Allosaurus europaeus, and, a mammal, Kuehneodon hahni. Other dinosaur specimens are referred including the nest and eggs and embryos of Lourinhanosaurus. Portugal is very productive in Late Jurassic vertebrates, being the seventh country bearing more dinosaur taxa.

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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.

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The Late Triassic (ca. 237-201.5 million years ago) was a transitional interval in the evolution of terrestrial ecosystems, during which ‘modern’ clades such as archosaurs and mammals were radiating while ‘archaic’ groups such as temnospondyl amphibians and basal synapsids remained abundant. Little is known about the Triassic terrestrial (nonmarine) vertebrates of the Iberian Peninsula. The Algarve Basin of southern Portugal is an extensional rift basin formed during the breakup of Pangaea, which is filled with terrestrial, lacustrine, and marginal marine siliciclastics of the Grés de Silves Formation, interbedded with CAMP basalts that mark the end-Triassic extinction (radioisotopically dated to ~198-201.5 Ma). Since 2009, our field project in the Algarve has discovered numerous vertebrate specimens within the Grés de Silves, including a monodominant bonebed containing hundreds of specimens of metoposaurids, a peculiar group of temnospondyls that filled crocodile-like predatory niches in lacustrine and fluvial environments. These specimens appear to belong to a new species of Metoposaurus, similar to M. diagnosticus and M. krasiejowensis from central Europe but possessing several putative autapomorphies of the braincase and lower jaw. We also discovered a mandible of a phytosaur, the first specimen of these long-snouted, semi-aquatic archosauriforms from the Iberian Peninsula. These discoveries of characteristic Carnian Norian taxa indicate that the fossil-bearing portion of the Grés de Silves is Late Triassic in age, and provide further evidence that metoposaurids and phytosaurs commonly occurred together in low palaeolatitudes during this time.

Portugal is ranked within the ten countries with more dinosaur taxa and the Lourinhã Formation is known
by the Late Jurassic findings of dinosaurs and other fossils. Often, studies of the external morphological
characteristics of the fossils are not sufficient and, observations of internal structures, using non-
destructive techniques, are required. The fossils here presented belong to the Museum da Lourinhã
(Portugal) and comprise a lower jaw of a basal crocodilian (possibly a Tomistomidae), eggshells and
several vertebrae from the exceptionally well preserved in ovo remains of Late Jurassic theropod dinosaur
Lourinhanosaurus. Neutron Tomography (NT) experiments with this material has been carried out at the
Geesthacht Neutron Facility in Germany. Additionally, eggshell fragments and several vertebrae have
been studied by Synchrotron-Radiation based Micro-Computed Tomography (SRμCT) at the HARWI II and
BW2 beamlines, respectively. These beamlines are operated by the Helmholtz-Zentrum Geesthacht at the
storage ring DORIS III at the Deutsches Elektronen–Synchrotron DESY in Hamburg, Germany. In both cases
(NT and SRμCT) complete 3D recordings have been obtained using a non-destructive procedure. The high-
quality tomographic datasets can be effectively studied through interactive digital visualization. Hence,
these visualization methods provide precious information about the 3D internal micro morphology of
fossils, like the network of the eggshell pores, often invisible in more traditional techniques, and provide a
direct window into the evolutionary history of organisms.

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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.

Forty partial fossil skulls belonging to beaked whales (Cetacea, Odontoceti, Ziphiidae) were collected by trawling and long-line fishing on Neogene (probably Late Early to Middle Miocene) layers of the Atlantic floor off the coasts of Portugal and Spain (Asturias and Galicia). e systematic study of the most diagnostic Iberian specimens, those preserving the rostrum and the dorsal part of the cranium, led to the recognition of two new genera (Globicetus n. gen. and Imocetus n. gen.) and four new species (Choneziphius leidyi n. sp., G. hiberus n. gen., n. sp., I. piscatus n. gen., n. sp., and Tusciziphius atlanticus n. sp.).
Based on the matrix of a previous work, the phylogenetic analysis places all the new taxa in the subfamily Ziphiinae Gray, 1850. More fragmentary specimens are tentatively referred to the genera Caviziphius Bianucci & Post, 2005 and Ziphirostrum du Bus, 1868. Among these new ziphiids, extremely bizarre skull morphologies are observed. In G. hiberus n. gen., n. sp. the proximal portion of the rostrum bears a voluminous premaxillary spheroid. In T. atlanticus n. sp. a medial premaxillary bulge is present on the rostrum; together with asymmetric
rostral maxillary eminences at the rostrum base, this bulge displays various degrees of elevation in different specimens, which may be interpreted as sexual dimorphism. Specimens of I. piscatus n. gen., n. sp. bear two sets of even crests: spur-like rostral maxillary crests and longitudinal maxillary crests laterally bordering a wide and long facial basin. A preliminary macroscopic observation of these elements indicates very dense bones, with a compactness comparable with that of cetacean ear bones. Questioning their function, the high medial rostral elements (the premaxillary spheroid of G. hiberus n. gen., n. sp. and the medial bulge of T. atlanticus n. sp.) remind the huge rostral maxillary crests of adult males of the extant Hyperoodon ampullatus (Forster, 1770). In the latter, the crests are very likely related to head-butting. However, they are made of much more spongy bone than in the fossil taxa studied here, and therefore possibly better mechanically suited for facing impacts. Other interpretations of these unusual bone specializations, related to deep-diving (ballast) and echolocation (sound reflection), fail to explain the diversity of shapes and the hypothetical sexual dimorphism observed in at least part of the taxa. e spur-like rostral maxillary crests and long maxillary crests limiting the large facial basin in I. piscatus n. gen., n. sp. and the excrescences on the maxilla at the rostrum base in Choneziphius spp. are instead interpreted as areas of origin for rostral and facial muscles, acting on the nasal passages, blowhole, and melon. From a palaeobiogeographic point of view, the newly described taxa further emphasize the differences in the North Atlantic (including Iberian Peninsula) and South African Neogene ziphiid faunal lists. Even if the stratigraphic context is poorly understood, leaving open the question of the geological age for most of the dredged specimens, these differences in the composition of cold to temperate northern and southern hemisphere fossil ziphiid faunas may be explained by a warm-water equatorial barrier.

The Spheroolithidae oospecies Guegoolithus turolensis, putatively attributed to non-hadrosauroid styracosterns was first described in the Barremian of the Iberian Basin, and later reported in the Valanginian–Hauterivian of the Cameros Basin, with both occurrences separated by a few hundred kilometres but by over 10 million years.

We here present fossil Crocodylomorpha eggshells from the Upper Jurassic Lourinhã Formation of Portugal, recovered from five sites: one nest from Cambelas with 13 eggs, and three partial eggs and various fragments from, Paimogo N (I), Paimogo S (II), Casal da Rola, and Peralta. All specimens but the nest were found in association with dinosaur egg material. Our research reveals that on a micro- and ultrastructural analysis, all samples present the typical characters consistent with crocodiloid eggshell morphotype, such as the shell unit shape, the organization of the eggshell layers, and the triangular blocky extinction observed with crossed nicols. We assign the material from the Lourinhã Formation to the oofamily Krokolithidae, making it the oldest crocodylomorph eggs known so far, as well as the best record for eggs of non- crocodylian crocodylomorphs. Furthermore, our study indicates that the basic structure of crocodiloid eggshells has remained stable since at least the Upper Jurassic.

We here present fossil Crocodylomorpha eggshells from the Upper Jurassic Lourinhã Formation of Portugal, recovered from five sites: one nest from Cambelas with 13 eggs, and three partial eggs and various fragments from, Paimogo N (I), Paimogo S (II), Casal da Rola, and Peralta. All specimens but the nest were found in association with dinosaur egg material. Our research reveals that on a micro- and ultrastructural analysis, all samples present the typical characters consistent with crocodiloid eggshell morphotype, such as the shell unit shape, the organization of the eggshell layers, and the triangular blocky extinction observed with crossed nicols. We assign the material from the Lourinhã Formation to the oofamily Krokolithidae, making it the oldest crocodylomorph eggs known so far, as well as the best record for eggs of non- crocodylian crocodylomorphs. Furthermore, our study indicates that the basic structure of crocodiloid eggshells has remained stable since at least the Upper Jurassic.

Thyreophoran dinosaurs have spike- and club-shaped osteoderms probably used for defensive weapons. The structural and histological variations have been little known. Here, we provide the comparisons of the internal structures in defensive weapons of ankylosaurs and stegosaurs, using spikes of a polacanthid (Gastonia) and a nodosaurid (Edmontonia), clubs of ankylosaurids (Saichania and Ankylosauridae indet. from Canada), and spikes of stegosaurids (Stegosaurus and Dacentrurus), which sheds light on understandings of evolutionary history and functional implications of defensive weapons in thyreophorans. In ankylosaurs, the structural and histological features of spikes and clubs are similar with those of small osteoderms in having thin compact bones, thick cancellous bones with large vascular canals, and abundant collagen fibers. A previous study demonstrated that each of three groups of ankylosaurs (polacanthid, nodosaurid, and ankylosaurid) has distinct characteristic arrangements of collagen fibers in small osteoderms. This study shows that spikes and clubs of ankylosaurs maintain the same characteristic features for each group despite of the differences in shapes and sizes. Conversely, the spike-shaped osteoderms in primitive (Dacentrurus) and derived (Stegosaurus) stegosaurids have similar structure to each other and are significantly different from the other types of stegosaur osteoderms (throat bony ossicles and plates) in having thick compact bones with a medullary cavity. These lack abundant collagen fibers unlike ankylosaur osteoderms. The spikes of ankylosaurs and stegosaurs are similar in shape, but their structural and histological features are different in having unique structures of collagen fibers for ankylosaurs and thick compact bones for stegosaurs, providing enough strength to have large spikes and to use them as defensive weapons. Although the shapes of ankylosaur clubs are different from spikes, the internal structures are similar, suggesting that ankylosaurs maintain similar structures despite of different shapes in osteoderms. These results indicate that ankylosaurs and stegosaurs used different strategies independently to evolve defensive weapons.

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The non-avian saurischians that have associated eggshells and embryos are represented only by the sauropodomorph Massospondylus and Coelurosauria (derived theropods), thus missing the basal theropod representatives. We report a dinosaur clutch containing several crushed eggs and embryonic material ascribed to the megalosaurid theropod Torvosaurus. It represents the first associated eggshells and embryos of megalosauroids, thus filling an important phylogenetic gap between two distantly related groups of saurischians. These fossils represent the only unequivocal basal theropod embryos found to date. The assemblage was found in early Tithonian fluvial overbank deposits of the Lourinhã Formation in West Portugal. The morphological, microstructural and chemical characterization results of the eggshell fragments indicate very mild diagenesis. Furthermore, these fossils allow unambiguous association of basal theropod osteology with a specific and unique new eggshell morphology.