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.

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 forelimb of a new sauropod dinosaur (Angolatitan adamastor n. gen. et sp.) from the Late Turonian of Iembe (Bengo Province) represents the first dinosaur discovery in Angola, and is one of the few occurrences of sauropod dinosaurs in sub-Saharan Africa collected with good chronological controls. The marginal marine sediments yielding the specimen are reported to be late Turonian in age and, thus it represents a non-titanosaurian sauropod in sub-Saharan Africa at a time taken to be dominated by titanosaurian forms. Moreover, Angolatitan adamastor is the only basal Somphospondyli known in the Late Cretaceous which implies in the existence of relict forms in Africa.

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.

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.

We present the δ13C and paleomagnetic stratigraphy for marine strata at the coast of southern Angola, anchored by an intercalated basalt with a whole rock 40Ar/39Ar radiometric age of 84.6 ± 1.5 Ma, being consistent with both invertebrate and vertebrate biostratigraphy. This is the first African stable carbon isotope record correlated to significant events in the global carbon cycle spanning the Late Cenomanian to Early Maastrichtian. A positive ∼ 3‰ excursion seen in bivalve shells below the basalt indicates the Cenomanian-Turonian Boundary Event at 93.9 Ma, during Oceanic Anoxic Event 2. Additional excursions above the basalt are correlated to patterns globally, including a negative ∼ 3‰ excursion near the top of the section interpreted as part of the Campanian-Maastrichtian Boundary Events. The age of the basalt ties the studied Bentiaba section to a pulse of Late Cretaceous magmatic activity around the South Atlantic and significant tectonic activity, including rotation, of the African continent.

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.

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The Bench 19 Bonebed at Bentiaba, Angola, is a unique concentration of marine vertebrates preserving six species of mosasaurs in sediments best correlated by magnetostratigraphy to chron C32n.1n between 71.4 and 71.64 Ma. The bonebed formed at a paleolatitude near 24°S, with an Atlantic width at that latitude approximating 2700 km, roughly half that of the current width. The locality lies on an uncharacteristically narrow continental shelf near transform faults that controlled the coastal outline of Africa in the formation of the South Atlantic Ocean. Biostratigraphic change through the Bentiaba section indicates that the accumulation occurred in an ecological time dimension within the 240 ky bin delimited by chron 32n.1n. The fauna occurs in a 10 m sand unit in the Mocuio Formation with bones and partial skeletons concentrated in, but not limited to, the basal 1–2 m. The sediment entombing the fossils is an immature feldspathic sand shown by detrital zircon ages to be derived from nearby granitic shield rocks. Specimens do not appear to have a strong preferred orientation and they are not concentrated in a strand line. Stable oxygen isotope analysis of associated bivalve shells indicates a water temperature of 18.5°C. The bonebed is clearly mixed with scattered dinosaur and pterosaur elements in a marine assemblage. Gut contents, scavenging marks and associated shed shark teeth in the Bench 19 Fauna indicate biological association and attrition due to feeding activities. The ecological diversity of mosasaur species is shown by tooth and body-size disparity and by δ13C analysis of tooth enamel, which indicate a variety of foraging areas and dietary niches. The Bench 19 Fauna was formed in arid latitudes along a coastal desert similar to that of modern Namibia on a narrow, tectonically controlled continental shelf, in shallow waters below wave base. The area was used as a foraging ground for diverse species, including molluscivorus Globidens phosphaticus, small species expected near the coast, abundant Prognathodon kianda, which fed on other mosasaurs at Bench 19, and species that may have been transient and opportunistic feeders in the area.

Recent fieldwork in the Namibe province in southern Angola yielded cranial and postcranial elements of at least two individuals of the rare and enigmatic mosasaur Halisaurus from a single small excavation. The genus Halisaurus is unique in retaining a primitive configuration of the temporal arcade, specifically a broad, vertically oriented contact between the parietal and the supratemporal. The supratemporal is broadly sutured to the opisthotic and prootic, unlike the condition in varanoids in which the simple lunate element lies between the parietal ramus and the squamosal and does not form a sutural contact with the opisthotic or prootic, but as in other halisaurines retains a plesiomorphic, vertically oriented contact with the parietal rami. The squamosal is lightly built and broadly arched as in Varanus. Comparison with known halisaurines indicates the new material is referable to the species Halisaurus arambourgi.
The locality that yielded the new specimens has also yielded a large number of isolated teeth, bones, articulated, and associated skeletons of Mosasaurus, Prognathodon, Globidens, and Plioplatecarpus, which with Halisaurus comprise a mosasaur assemblage most similar to that reported from the Maastrichtian of Morocco.

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Abstract Several analyses of diversity through geological time use global, synoptic databases, and this practice often makes it difficult to distinguish true signals in changing diversity from regional-scale sampling and/or geological artefacts. Here we investigate how echinoid diversity changed through the Mesozoic of the Lusitanian basin in Portugal based on a comprehensive, revised database, and seek to distinguish biological signal from geological or environmental constraints. The observed diversity pattern is far from having a defined trend, showing many fluctuations that appear to be linked with gaps in the geological record. This study revealed that, independently of the method used, whether correlation tests or model fitting, the diversity signal is not completely explained by the studied sampling proxies. Among the different proxies, marine facies variation in combination with outcrop area best explains the palaeodiversity curve.

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.

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.

ABSTRACT New elasmosaurid plesiosaur specimens are described from the Early Maastrichtian of Angola. Phylogenetic analyses reconstruct the Angolan taxon as an aristonectine elasmosaurid and the sister taxon of an unnamed form of similar age from New Zealand. Comparisons also indicate a close relationship with an unnamed form previously described from Patagonia. All of these specimens exhibit an ostensibly osteologically immature external morphology, but histological analysis of the Angolan material suggests an adult with paedomorphic traits. By extension, the similarity of the Angolan, New Zealand and Patagonian material indicates that these specimens represent a widespread paedomorphic yet unnamed taxon.

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.

ABSTRACT We report here a new elasmosaurid from the early Maastrichtian at Bentiaba, southern Angola. Phylogenetic analysis places the new taxon as the sister taxon to Styxosaurus snowii, and that clade as the sister of a clade composed of (Hydrotherosaurus alexandrae (Libonectes morgani + Elasmosaurus platyurus)). The new taxon has a reduced dorsal blade of the scapula, a feature unique amongst elasmosaurids, but convergent with cryptoclidid plesiosaurs, and indicates a longitudinal protraction-retraction limb cycle rowing style with simple pitch rotation at the glenohumeral articulation. Morphometric phylogenetic analysis of the coracoids of 40 eosauropterygian taxa suggests that there was a broad range of swimming styles within the clade.

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Adult large-bodied theropods are often found with numerous pathologies. A large, almost complete, probably adult Allosaurus specimen from the Howe Stephens Quarry, Morrison Formation (Late Kimmeridgian–Early Tithonian), Wyoming, shows multiple pathologies. Pathologic bones include the left dentary, two cervical vertebrae, one cervical and several dorsal ribs, the left scapula, the left humerus, right ischium, and two left pedal phalanges. These pathologies can be classified as follows: the fifth cervical vertebra, the scapula, several ribs and the ischium are traumatic, and a callus on the shaft of the left pedal phalanx II-2 is traumatic-infectious. Traumatically fractured elements exposed to frequent movement (e.g. the scapula and the ribs) show a tendency to develop pseudarthroses instead of callus healing. The pathologies in the lower jaw and a reduced flexor tubercle of the left pedal phalanx II-2 are most likely traumatic or developmental in origin. The pathologies on the fourth cervical are most likely developmental in origin or idiopathic, that on the left humerus is infectious or idiopathic, whereas left pedal phalanx IV-1 is classified as idiopathic. With exception of the ischium, all traumatic / traumatic-infectious pathologic elements show unambiguous evidences of healing, indicating that the respective pathologies did not cause the death of this individual. Alignment of the scapula and rib pathologies from the left side suggests that all may have been caused by a single traumatic event. The ischial fracture may have been fatal. The occurrence of multiple traumatic pathologies again underlines that large-bodied theropods experienced frequent injuries during life, indicating an active predatory lifestyle, and their survival perhaps supports a gregarious behavior for Allosaurus. Signs of infections are scarce and locally restricted, indicating a successful prevention of the spread of pathogens, as it is the case in extant reptiles (including birds).

Adult large-bodied theropods are often found with numerous pathologies. A large, almost complete, probably adult \\textitAllosaurus specimen from the Howe Stephens Quarry, Morrison Formation (Late Kimmeridgian–Early Tithonian), Wyoming, exhibits multiple pathologies. Pathologic bones include the left dentary, two cervical vertebrae, one cervical and several dorsal ribs, the left scapula, the left humerus, the right ischium, and two left pedal phalanges. These pathologies can be classified as follows: the fifth cervical vertebra, the scapula, several ribs and the ischium are probably traumatic, and a callus on the shaft of the left pedal phalanx II-2 is probably traumatic-infectious. Traumatically fractured elements exposed to frequent movement (e.g., the scapula and the ribs) show a tendency to develop pseudarthroses instead of a callus. The pathologies in the lower jaw and a reduced extensor tubercle of the left pedal phalanx II-2 are most likely traumatic or developmental in origin. The pathologies on the fourth cervical are most likely developmental in origin or idiopathic, that on the left humerus could be traumatic, developmental, infectious or idiopathic, whereas the left pedal phalanx IV-1 is classified as idiopathic. With exception of the ischium, all as traumatic/traumatic-infectious classified pathologic elements show unambiguous evidences of healing, indicating that the respective pathologies did not cause the death of this individual. Alignment of the scapula and rib pathologies from the left side suggests that all may have been caused by a single traumatic event. The ischial fracture may have been fatal. The occurrence of multiple lesions interpreted as traumatic pathologies again underlines that large-bodied theropods experienced frequent injuries during life, indicating an active predatory lifestyle, and their survival perhaps supports a gregarious behavior for \\textitAllosaurus. Alternatively, the frequent survival of traumatic events could be also related to the presence of non-endothermic metabolic rates that allow survival based on sporadic food consumption or scavenging behavior. Signs of pathologies consistent with infections are scarce and locally restricted, indicating a successful prevention of the spread of pathogens, as it is the case in extant reptiles (including birds).