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.

Theropod teeth are particularly abundant in the fossil record and frequently reported in the literature. Yet, the dentition of many theropods has not been described comprehensively, omitting details on the denticle shape, crown ornamentation and enamel texture. This paucity of information has been particularly striking in basal clades, thus making identification of isolated teeth difficult, and taxonomic assignments uncertain. We here provide a detailed description of the dentition of Megalosauridae, and a comparison to and distinction from superficially similar teeth of all major theropod clades. Megalosaurid dinosaurs are characterized by a mesial carina facing mesiolabially in most mesial teeth, centrally positioned carinae on both most mesial and lateral crowns, a mesial carina terminating above the cervix, and short to well-developed interdenticular sulci between distal denticles. A discriminant analysis performed on a dataset of numerical data collected on the teeth of 62 theropod taxa reveals that megalosaurid teeth are hardly distinguishable from other theropod clades with ziphodont dentition. This study highlights the importance of detailing anatomical descriptions and providing additional morphometric data on teeth with the purpose of helping to identify isolated theropod teeth in the future.

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

The skull-bone quadrate in nonavian theropods is very diverse morphologically alongside the disparity of the group as a whole. However this disparity has been underestimated for taxonomic purposes. In order to evaluate the phylogenetic potential and investigate the evolutionary transformations of the quadrate, we conducted a Catalano-Goloboff phylogenetic morphometric analysis as well as a cladistic analysis using 98 discrete quadrate related characters. The cladistic analysis provides a fully resolved tree mirroring to some degree the classification of nonavian theropods. The quadrate morphology by its own provides a wealth of data with strong phylogenetic signal and allows inference of major trends in the evolution of this bone. Important synapomorphies include: for Abelisauroidea, a lateral ramus extending to the ectocondyle; for Tetanurae, the absence of the lateral process; for Spinosauridae, a medial curvature of the ventral part of the pterygoid ramus occurring just above the mandibular articulation; for Avetheropoda, an anterior margin of the pterygoid flange formed by a roughly parabolic margin; and for Tyrannosauroidea, a semi-oval pterygoid flange shape in medial view. The Catalano-Goloboff phylogenetic morphometric analysis reveals two main morphotypes of the mandibular articulation of the quadrate linked to function. The first morphotype, characterized by an anteroposteriorly broad mandibular articulation with two ovoid/subcircular condyles roughly subequal in size, is found in Ceratosauria, Tyrannosauroidea and Oviraptorosauria. This morphotype allows a very weak displacement of the mandible laterally. The second morphotype is characterized by an elongate and anteroposteriorly narrow mandibular articulation and a long and parabolic/sigmoid ectocondyle. Present in Megalosauroidea, Carcharodontosauridae and Dromaeosauridae, this morphotype permits the lower jaw rami to be displaced laterally when the mouth opened.

Theropods form a taxonomically and morphologically diverse group of dinosaurs that include extant birds. Inferred relationships between theropod clades are complex and have changed dramatically over the past thirty years with the emergence of cladistic techniques. Here, we present a brief historical perspective of theropod discoveries and classification, as well as an overview on the current systematics of non-avian theropods. The first scientifically recorded theropod remains dating back to the 17th and 18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to the megalosaurid Megalosaurus. The latter was the first theropod genus to be named in 1824, and subsequent theropod material found before 1850 can all be referred to megalosauroids. In the fifty years from 1856 to 1906, theropod remains were reported from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organization of ‘family’ to ‘superfamily’ level clades. The earliest definitive theropods come from the Carnian of Argentina, and coelophysoids form the first significant theropod radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs, therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the end of the Cretaceous, though the megalosauroid clade did not extend into the Maastrichtian. Current debates are focused on the monophyly of deinonychosaurs, the position of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids. Some recent analyses have suggested a placement of dilophosaurids outside Coelophysoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria with troodontids placed more closely to avialans than dromaeosaurids.

The Lourinhã Formation (Kimmeridgian-Tithonian) of Central West Portugal is well known for its diversified dinosaur fauna similar to that of the Morrison Formation of North America; both areas share dinosaur taxa including the top predator Torvosaurus, reported in Portugal. The material assigned to the Portuguese T. tanneri, consisting of a right maxilla and an incomplete caudal centrum, was briefly described in the literature and a thorough description of these bones is here given for the first time. A comparison with material referred to Torvosaurus tanneri allows us to highlight some important differences justifying the creation of a distinct Eastern species. Torvosaurus gurneyi n. sp. displays two autapomorphies among Megalosauroidea, a maxilla possessing fewer than eleven teeth and an interdental wall nearly coincidental with the lateral wall of the maxillary body. In addition, it differs from T. tanneri by a reduced number of maxillary teeth, the absence of interdental plates terminating ventrally by broad V-shaped points and falling short relative to the lateral maxillary wall, and the absence of a protuberant ridge on the anterior part of the medial shelf, posterior to the anteromedial process. T. gurneyi is the largest theropod from the Lourinhã Formation of Portugal and the largest land predator discovered in Europe hitherto. This taxon supports the mechanism of vicariance that occurred in the Iberian Meseta during the Late Jurassic when the proto-Atlantic was already well formed. A fragment of maxilla from the Lourinhã Formation referred to Torvosaurus sp. is ascribed to this new species, and several other bones, including a femur, a tibia and embryonic material all from the Kimmeridgian-Tithonian of Portugal, are tentatively assigned to T. gurneyi. A standard terminology and notation of the theropod maxilla is also proposed and a record of the Torvosaurus material from Portugal is given.

Although nonavian theropod have received considerable interest in the last years, their ontogeny still remains poorly understood, especially the ontogenetical changes affecting their skull (Rauhut
and Fechner, 2005). The quadrate, for instance, is preserved in several embryos and juvenile specimens belonging to many clades of theropods such as the Tyrannosauridae (Carr, 1999), Compsognathidae (Dal Sasso and Maganuco, 2011), Therizinosauroidea (Kúndrat et al., 2007), Oviraptoridae (Norell et al., 1994; Norell et al., 2001; Weishampel et al., 2008) and Troodontidae (Varrichio et al., 2002) but very little is usually said about the anatomy of this bone and no one has ever investigated ontogenetical variation in the nonavian theropod quadrate. The discovery of two quadrates belonging to embryos of the sinraptorid Lourinhanosaurus antunesi from Portugal and five isolated quadrates pertaining to juvenile, subadult and adult specimens of Spinosauridae from Morocco fills this gap and allows some ontogenetic information to be drawn for this bone in these two specific clades of Theropoda.
I

Theropod dinosaurs form a highly diversified clade, and their teeth are some of the most common components of the Mesozoic dinosaur fossil record. This is the case in the Lourinhã Formation (Late Jurassic, Kimmeridgian-Tithonian) of Portugal, where theropod teeth are particularly abundant and diverse. Four isolated theropod teeth are here described and identified based on morphometric and anatomical data. They are included in a cladistic analysis performed on a data matrix of 141 dentition-based characters coded in 60 taxa, as well as a supermatrix combining our dataset with six recent datamatrices based on the whole theropod skeleton. The consensus tree resulting from the dentition-based data matrix reveals that theropod teeth provide reliable data for identification at approximately family level. Therefore, phylogenetic methods will help identifying theropod teeth with more confidence in the future. Although dental characters do not reliably indicate relationships among higher clades of theropods, they demonstrate interesting patterns of homoplasy suggesting dietary convergence in (1) alvarezsauroids, therizinosaurs and troodontids; (2) coelophysoids and spinosaurids; (3) compsognathids and dromaeosaurids; and (4) ceratosaurids, allosauroids and megalosaurids.

Based on morphometric and cladistic analyses, the biggest tooth from Lourinhã is referred to a mesial crown of the megalosaurid Torvosaurus tanneri, due to the elliptical cross section of the crown base, the large size and elongation of the crown, medially positioned mesial and distal carinae, and the coarse denticles. The smallest tooth is identified as Richardoestesia, and as a close relative of R. gilmorei based on the weak constriction between crown and root, the “eight-shaped” outline of the base crown and, on the distal carina, the average of ten symmetrically rounded denticles per mm, as well as a subequal number of denticles basally and at mid-crown. Finally, the two medium-sized teeth belong to the same taxon and exhibit pronounced interdenticular sulci between distal denticles, hooked distal denticles for one of them, an irregular enamel texture, and a straight distal margin, a combination of features only observed in abelisaurids. They provide the first record of Abelisauridae in the Jurassic of Laurasia and one of the oldest records of this clade in the world, suggesting a possible radiation of Abelisauridae in Europe well before the Upper Cretaceous.

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The quadrate in nonavian theropods is incredibly diverse morphologically; however this morphological disparity has been underestimated for taxonomic purposes. The quadrate topological homologies and anatomy, as well as the terminology, among nonavian theropod clades are reviewed. In order to evaluate the phylogenetic potential and investigate the evolutionary transformations of the quadrate, we conducted a Catalano-Goloboff phylogenetic morphometric analysis using 3 morphometric characters, a total of 28 landmarks coded for 23 taxa, as well as a cladistic analysis using 115 discrete quadrate-related characters coded for 43 taxa. The cladistic analysis provides a fully resolved tree mirroring the current classification of nonavian theropods. The quadrate morphology by its own provides a wealth of data with strong phylogenetic signal. Several unambiguous synapomorphies support nonavian theropod relationships and the resulting consensus tree allows inference of major trends in the evolution of this bone. Important synapomorphies include: for Abelisauridae, a lateral ramus extending to the ectocondyle; for Tetanurae, the absence of the lateral process; for Spinosauridae, a medial curvature of the ventral part of the pterygoid ramus occurring just above the mandibular articulation; for Neotetanurae, an anterior margin of the pterygoid flange formed by a roughly parabolic margin; and for Tyrannosauroidea, a semi-oval pterygoid flange shape in medial view. The Catalano-Goloboff phylogenetic morphometric analysis reveals two main morphotypes of the mandibular articulation of the quadrate linked to function. The first morphotype, characterized by an anteroposteriorly broad mandibular articulation with two ovoid/subcircular condyles roughly subequal in size, is found in Ceratosauria, Tyrannosauroidea and Oviraptorosauria. This morphotype allows a very weak displacement of the mandible laterally. The second morphotype is characterized by an elongate and anteroposteriorly narrow mandibular articulation and a long and parabolic/sigmoid ectocondyle. Present in Megalosauroidea, Allosauroidea and Dromaeosauridae, this morphotype permits the lower jaw rami to be displaced laterally when the mouth opened.

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Vertebrate-bearing fossiliferous outcrops of Cretaceous age in sub-Saharan Africa are rare because of younger superficial deposits, vegetation cover, and the widespread occurrence of Precambrian metamorphic plateau basement comprising much of the continent. However, one area of extensive marine and nonmarine
Cretaceous exposures is found between the plateau and the coast in Angola. The Angolan margin was formed in conjunction with the breakup of Gondwana and subsequent growth of the South Atlantic. Cretaceous deposits are constrained in age by the emplacement of oceanic crust, which began no later than magnetozone M3
(approximately 128 Ma, Barremian). Shallow marine facies are exposed in sea cliffs but equivalent facies become increasingly terrestrial inland. Few vertebrate fossils have been described from Angola aside from sharks.
Notable exceptions are the late Turonian mosasaurs Angolasaurus bocagei and Tylosaurus iembeensis from northern Angola. Those taxa are significant because they are among the earliest derived mosasaurs. Recent field work led to the discovery of a new skull of Angolasaursus as well as sharks, fish, plesiosaurs, the skull of a new taxon of turtle, additional mosasaurs, and the articulated forelimb of a sauropod dinosaur, the first reported dinosaur from Angola. In southern Angola, marine sediments spanning the Cretaceous-Paleogene boundary are found.

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Africa is the only continent that now straddles arid zones located beneath the descending limbs of both the northern and southern Hadley cells, and it has done so since it became a distinct continent in the Early Cretaceous. Since that time, Africa has drifted tectonically some 12 degrees north and rotated approximately 45 degrees counterclockwise. This changing latitudinal setting and position of the landmass under the relatively stable Hadley Cells is manifested as southward migration of climatic zones over the past 132 million years. Data from kerogen, X-ray diffraction analysis of sedimentary matrix, carbon isotopes from shell samples and tooth enamel,new 40Ar/39Ar radiometric dates, pollen and plant macrofossils, and fossil vertebrates indicate a productive upwelling system adjacent to a coastal desert since the opening of the South Atlantic Ocean; however, the position of the coastal desert has migrated southward as Africa drifted north, resulting in today's Skeleton Coast and Benguela Current. This migration has had a profound effect on the placement of the West African coast relative to areas of high marine productivity and resulting extensive hydrocarbon deposits, on the placement of arid zones relative to the continent especially the Skeleton Coast desert, on the climatic history of the Congo Basin (which shows a Late Cretaceous decrease in aridity based on the relative abundance of analcime in the Samba core), and in reducing the southern temperate region of Africa from 17% of continental area during the Cretaceous to 2% today. We show here that these related geographic and environmental changes drove ecological and evolutionary adjustments in southern African floras and faunas, specifically with respect to the distribution of anthropoid primates, the occurrence of modern relicts such as the gnetalean Welwitschia mirabilis, endemism as in the case of ice plants, and mammalian adaption to an open environment as in springhares. Africa's tectonic drift through climate zones has been a first-order environmental determinant since the Early Cretaceous.

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The Ørsted Dal Member of the Upper Triassic Fleming Fjord Formation in East Greenland is well known for its rich vertebrate fauna, represented by numerous specimens of both body and ichnofossils. In particular, the footprints of theropod dinosaurs have been described. Recently, an international expedition discovered several slabs with 100 small chirotheriid pes and manus imprints (pes length 4–4.5 cm) in siliciclastic deposits of this unit. They show strong similarities with Brachychirotherium, a characteristic Upper Triassic ichnogenus with a global distribution. A peculiar feature in the Fleming Fjord specimens is the lack of a fifth digit, even in more deeply impressed imprints. Therefore, the specimens are assigned here tentatively to cf. Brachychirotherium. Possibly, this characteristic is related to the extremely small size and early ontogenetic stage of the trackmaker. The record from Greenland is the first evidence of this morphotype from the Fleming Fjord Formation. Candidate trackmakers are crocodylian stem group archosaurs; however, a distinct correlation with known osteological taxa from this unit is not currently possible. While the occurrence of sauropodomorph plateosaurs in the bone record links the Greenland assemblage more closer to that from the Germanic Basin of central Europe, here the described footprints suggest a Pangaea-wide exchange.Supplementary material: Three-dimensional model of cf. Brachychirotherium pes–manus set (from MGUH 31233b) from the Upper Triassic Fleming Fjord Formation (Norian–Rhaetian) of East Greenland as pdf, ply and jpg files (3D model created by Oliver Wings; photographs taken by Jesper Milàn) is available at https://doi.org/10.6084/m9.figshare.c.2133546

The Late Triassic (Norian–early Rhaetian) Fleming Fjord Formation of central East Greenland preserves a diverse fossil fauna, including both body and trace fossils. Trackways of large quadrupedal archosaurs, although already reported in 1994 and mentioned in subsequent publications, are here described and figured in detail for the first time, based on photogrammetric data collected during fieldwork in 2012. Two trackways can be referred to Eosauropus, while a third, bipedal trackway may be referred to Evazoum, both of which have been considered to represent sauropodomorph dinosaur tracks. Both the Evazoum and the Eosauropus trackways are distinctly larger than other trackways referred to the respective ichnogenera. The trackmaker of the best preserved Eosauropus trackway is constrained using a synapomorphy-based approach. The quadrupedal posture, the entaxonic pes structure, and five weight-bearing digits indicate a derived sauropodiform trackmaker. Other features exhibited by the tracks, including the semi-digitigrade pes and the laterally deflected unguals, are commonly considered synapomorphies of more exclusive clades within Sauropoda. The present trackway documents an early acquisition of a eusauropod-like pes anatomy while retaining a well-developed claw on pedal digit IV, which is reduced in eusauropods. Although unequivocal evidence for sauropod dinosaurs is no older than the Early Jurassic, the present trackway provides evidence for a possible Triassic origin of the group.

Abstract
Late Jurassic dinosaur footprints were found on a coastline cliff in Lourinhã, Porto das Barcas, Lagido do Forno (coordinate 39°14.178’N, 9°20.397’W, Portugal) in June 2001. The locality is characterized by steep cliffs with high slopes that are composed of gray and red sandstones/ siltstones. The location belongs to the successions of Lusitanian Basin representing the Porto Novo Member of the Lourinhã Formation. Three natural infills of tridactyl tracks, possibly ascribed to ornithopod, a bipedal herbivore were found, representing a left foot movement, a right and a left one, respectively. Footprints are 300- 400mm wide and have a height of 330-360mm. The footprints are characterized by round fingers, which are elongated due to some degradation/ erosion. The footprints were collected from the field in 2001 and subsequently cleaned, consolidated and glued in the laboratory of the Museum of Lourinhã before being exhibited in a museum display. Stone matrix was removed and a consolidation product was applied, probably a polyvinyl acetate. The footprint with broken central digit was glued with an epoxy resin, Araldite. Both applied products were confirmed by analysis of μ- FTIR and both presented colour change and detachment surface problems. The footprints have been exposed in the palaeontology hall of the Museum of Lourinhã, Portugal from 2004 without climate controlling. These trace fossils form an important part of the palaeontological collection of Late Jurassic vertebrate fossils from Lourinhã Formation. Presently, it is considered a unique heritage in danger of disappearing due to high decay level of disaggregation of its geological structure. The footprints display several pathologies, such as “Blistering”, “Powdering”, “Exfoliation”’ as well as “Dirt”, “Fracture”’, “Inscriptions”, “Consolidants” and “Adhesives” and are now in very poor conditions. Laboratorial analysed were made to evaluate the presence of salts. Moreover a microclimatic study was conducted inside the museum to evaluate the influence of thermo-hygrometric parameters on the decay processes. The future interventions will depend on the results of consolidation trials that are currently under progress by using stone samples taken from the same layer and location from Porto das Barcas applying different commercial consolidation products.

This study focuses on the assessment of the degradation processes occurring in three sandstone infills of fossilized Late Jurassic ornithopod tridactyl footprints, found in 2001 in a coastline cliff in Porto das Barcas (Lourinhã, Portugal) and exhibited in a museum display since 2004. These dinosaur footprints present nowadays severe decay phenomena compromising their physical integrity and are leading gradually to their loss of value. The deterioration patterns were recorded, a map of their distribution was prepared and several samples were collected both in the dinosaur footprints and in the coastline cliff. Different analytical procedures were applied such as XRD, FTIR, FESEM and Ion Chromatography. A microclimatic survey was also performed and air temperature and relative humidity was measured during eight months both indoor and also outdoor. The decay patterns observed are a combination intrinsic and extrinsic factors the stone material, namely swelling of clay minerals in the rock matrix (smectite and chlorite-smectite mixed-layer), presence of salts (mainly chlorides), application of past conservation treatments (poly(vinyl) acetate and epoxy resins) and with the museum's indoor thermohygrometric conditions (mainly non-stable hygrometric conditions). This scientific knowledge is therefore essential to the sustainable preservation of this paleontological heritage.

Although diplodocoid sauropods from Africa and the Americas are well known, their European record remains largely neglected. Here we redescribe Dinheirosaurus lourinhanensis from the Late Jurassic of Portugal. The holotype comprises two posterior cervical vertebrae, the dorsal series and a caudal centrum. Redescription demonstrates its validity on the basis of three autapomorphies: (1) posteriorly restricted ventral keel on posterior cervical vertebrae; (2) three small subcircular fossae posterior to the lateral coel on posterior cervical neural spines; (3) accessory lamina linking the hyposphene with base of the posterior centrodiapophyseal lamina in middle-posterior dorsal vertebrae. Phylogenetic analysis places Dinheirosaurus as the sister taxon to Supersaurus, and this clade forms the sister taxon to other diplodocines. However, this position should be treated with caution as Dinheirosaurus displays several plesiomorphic features absent in other diplodocids (including unbifurcated presacral neural spines, and dorsolaterally projecting diapophyses on dorsal vertebrae) and only four additional steps are required to place Dinheirosaurus outside of Flagellicaudata. We identify Amazonsaurus as the basal-most rebbachisaurid and recover Zapalasaurus outside of the South American Limaysaurinae, suggesting the biogeographic history of rebbachisaurids is more complex than previously proposed. Review of the European diplodocoid record reveals evidence for the earliest known diplodocid, as well as additional diplodocid remains from the Late Jurassic of Spain. A Portuguese specimen, previously referred to Dinheirosaurus, displays strong similarities to Apatosaurus from the contemporaneous Morrison Formation of North America, indicating the presence of a second Late Jurassic Portuguese diplodocid taxon. Along with Dinheirosaurus, these Portuguese remains provide further evidence for a Late Jurassic palaeobiogeographic connection between Europe and North America. No dicraeosaurids are currently known from Europe, but rebbachisaurids are present in the Early Cretaceous, with weak evidence for the earliest known representative from the Late Jurassic of Spain; however, more complete material is required to recognize early members of this clade.

Titanosauriforms represent a diverse and globally distributed clade of neosauropod dinosaurs, but their inter-relationships remain poorly understood. Here we redescribe Lusotitan atalaiensis from the Late Jurassic Lourinhã Formation of Portugal, a taxon previously referred to Brachiosaurus. The lectotype includes cervical, dorsal, and caudal vertebrae, and elements from the forelimb, hindlimb, and pelvic girdle. Lusotitan is a valid taxon and can be diagnosed by six autapomorphies, including the presence of elongate postzygapophyses that project well beyond the posterior margin of the neural arch in anterior-to-middle caudal vertebrae. A new phylogenetic analysis, focused on elucidating the evolutionary relationships of basal titanosauriforms, is presented, comprising 63 taxa scored for 279 characters. Many of these characters are heavily revised or novel to our study, and a number of ingroup taxa have never previously been incorporated into a phylogenetic analysis. We treated quantitative characters as discrete and continuous data in two parallel analyses, and explored the effect of implied weighting. Although we recovered monophyletic brachiosaurid and somphospondylan sister clades within Titanosauriformes, their compositions were affected by alternative treatments of quantitative data and, especially, by the weighting of such data. This suggests that the treatment of quantitative data is important and the wrong decisions might lead to incorrect tree topologies. In particular, the diversity of Titanosauria was greatly increased by the use of implied weights. Our results support the generic separation of the contemporaneous taxa Brachiosaurus, Giraffatitan, and Lusotitan, with the latter recovered as either a brachiosaurid or the sister taxon to Titanosauriformes. Although Janenschia was recovered as a basal macronarian, outside Titanosauria, the sympatric Australodocus provides body fossil evidence for the pre-Cretaceous origin of titanosaurs. We recovered evidence for a sauropod with close affinities to the Chinese taxon Mamenchisaurus in the Late Jurassic Tendaguru beds of Africa, and present new information demonstrating the wider distribution of caudal pneumaticity within Titanosauria. The earliest known titanosauriform body fossils are from the late Oxfordian (Late Jurassic), although trackway evidence indicates a Middle Jurassic origin. Diversity increased throughout the Late Jurassic, and titanosauriforms did not undergo a severe extinction across the Jurassic/Cretaceous boundary, in contrast to diplodocids and non-neosauropods. Titanosauriform diversity increased in the Barremian and Aptian–Albian as a result of radiations of derived somphospondylans and lithostrotians, respectively, but there was a severe drop (up to 40%) in species numbers at, or near, the Albian/Cenomanian boundary, representing a faunal turnover whereby basal titanosauriforms were replaced by derived titanosaurs, although this transition occurred in a spatiotemporally staggered fashion.

A partial chelonian was collected from the Langenberg Formation quarry of Oker (near Goslar, Lower Saxony, Germany). The amniotes from this formation include other chelonians (possible Plesiochelyidae), the sauropod Europasaurus holgeri, theropods (Velociraptorinae), crocodylomorphs (Atoposauridae, Theriosuchus pusillus, Goniopholis simus, Machimosaurus hugii, Steneosaurus brevirostris) and pterosaurs (Dsungapteridae, ?Ornithocheiroidea, ?Ctenochasmatidae).

The reported specimen, DFMMh/FV 296, includes a skull part (articulated quadrate, squamosal, basisphenoid, and pterygoid), a disarticulated 40 cm long partial carapace, plastron, and one cervical vertebra.

The carapace bears fontanelles, trapezoidal suprapygal with straight edges, small last neurals

[....]
a plicated longitudinal pattern in the proximal end of the costals originating on the posterior side of the scute sulci and dissipating posteriorly, wide central opening in the plastron, xiphiplastra with little or no contact between each other, and both the hyo- and hypoplastron have digit-form buttress projections.

The specimen has a central plastra fontanelle, which is regarded as a feature of the clade including [....], Santanachelys, and Thalassemys. DFMMh/FV 296 differs from
[....]

edges and xiphiplastron participation in the central fontanelle is autapomorphic for the Thalassemys genus. A large central fontanelle with hyo and hypoplastron polygonal medial
[....]
contact between xiphiplastra and reduced size of the two last neural plates is shared between T. hugii and the Oker specimen. DFMMh/FV 296 differs from this species due to the presence of plastral projections. The different shape of the plastron (no polygonal-like margins or hyo- and hypoplastron projections) suggests that T. moseri [....]
ent genus. Therefore, we can assign this specimen to Thalassemys sp.

The paleontological richness of Morocco has been scientifically known since at least the early 20th century. The region of the Middle Atlas, more specifically the Boulemane area, has been however only sparsely studied since the 1960s when it provided vertebrate fossils from the Middle Jurassic. In September 2013, a Moroccan-Portuguese expedition to the village of Taghrout, Boulemane, made excavations in a Pleistocene fossil site that once was a small high-altitude sedimentary basin, uncharted in previous geological maps. The excavations yielded bone material from large mammals, the most common findings are elephants ascribed to Elephas, but artiodactyls, turtles, and in-situ Acheulean tools were also collected. This represents a new and important paleontological and archeological site. In addition to the discoveries of Taghrout, the expedition also retrieved Quaternary vertebrate material from a nearby cave and found new Jurassic localities, with arcosaur bones and dinosaur footprints, in El Mers.

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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.

This article presents a synthesis of Palaeozoic and Mesozoic fossil tetrapods from Greenland, including an updated review of the holotypes and a new photographic record of the main specimens. All fossil tetrapods found are from East Greenland, with at least 30 different known taxa: five stem tetrapods (Acanthostega gunnari, Ichthyostega eigili, I. stensioi, I. watsoni, and Ymeria denticulata) from the Late Devonian of the Aina Dal and Britta Dal Formations; four temnospondyl amphibians (Aquiloniferus kochi, Selenocara groenlandica, Stoschiosaurus nielseni, and Tupilakosaurus heilmani) from the Early Triassic of the Wordie Creek Group; two temnospondyls (Cyclotosaurus naraserluki and Gerrothorax cf. pulcherrimus), one testudinatan (cf. Proganochelys), two stagonolepids (Aetosaurus ferratus and Paratypothorax andressorum), the eudimorphodontid Arcticodactylus, undetermined archosaurs (phytosaurs and both sauropodomorph and theropod dinosaurs), the cynodont Mitredon cromptoni, and three mammals (Haramiyavia clemmenseni, Kuehneotherium, and cf. ?Brachyzostrodon), from the Late Triassic of the Fleming
Fjord Formation; one plesiosaur from the Early Jurassic of the Kap Stewart Formation; one plesiosaur and one ichthyosaur from the Late Jurassic of the Kap Leslie Formation, plus a previously unreported Late Jurassic plesiosaur from Kronprins Christian Land. Moreover, fossil tetrapod trackways are known from the Late Carboniferous (morphotype Limnopus) of the Mesters Vig Formation and at least four different morphologies (such as the crocodylomorph Brachychirotherium, the auropodomorph Eosauropus and Evazoum, and the theropodian Grallator) associated to archosaurian trackmakers are known from the Late Triassic of the Fleming Fjord Formation. The presence of rich fossiliferous tetrapod sites in East Greenland is linked to the presence of well-exposed continental and shallow marine deposits with most finds in terrestrial deposits from the Late Devonian and the Late Triassic.

Eggshells from the three extant crocodilian species Crocodylus mindorensis (Philippine Crocodile), Paleosuchus palpebrosus (Cuvier's Smooth-fronted Caiman or Musky Caiman) and Alligator mississippiensis (American Alligator or Common Alligator) were prepared for thin section and scanning electron microscope analyses and are described in order to improve the knowledge on crocodilian eggs anatomy and microstructure, and to find new apomorphies that can be used for identification. Both extant and fossil crocodilian eggs present an ornamentation that vary as anastomo-, ramo- or the here newly described rugosocavate type. The angusticaniculate pore system is a shared character for Crocodylomorpha eggshells and some dinosaurian and avian groups. Previously reported signs of incubated crocodilian eggs were found also on our only fertilised and hatched egg. Paleosuchus palpebrosus presents unique organization and morphology of the three eggshell layers, with a relatively thin middle layer characterised by dense and compact tabular microstructure.