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

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

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.

n/a

Although occurrences of marine reptiles have been previously reported from Angola, with the exception of two Turonian taxa, these reports were based largely on isolated teeth. Fieldwork in 2005 and 2006 yielded well-preserved remains of marine reptiles including plesiosaurs, turtles, and mosasaurs. The mosasaurs discussed here were recovered from two field areas: Turonian sediments at Iembe along the north coast and Maastrichtian sediments at Bentiaba on the south coast. The Turonian section near Iembe produced at least two new specimens of Angolasaurus bocagei and one fragmentary specimen of Tylosaurus iembeensis. One of the Angolasaurus specimens is represented by a well preserved, complete and articulated skull and partial postcrania, including portions of the forelimbs and pectoral girdle. The preservation of material from the Bentiaba locality is remarkable due to the grain support of the entombing sandstone, which preserves fine anatomical details with little apparent crushing, and in the number of articulated, semi-articulated, and associated skeletons. Identifications from the field and preliminary preparation show the Bentiaba mosasaur fauna is represented by at least five genera including Mosasaurus, Prognathodon, Globidens, Plioplatecarpus and Halisaurus. Collectively, these new specimens greatly expand our knowledge of the anatomy and systematics of Angolan mosasaurs.

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.

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.

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.

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.

n/a

n/a

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.

n/a

n/a

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.

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.