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2015
Araújo, R., Polcyn M. J., Lindgren J., Jacobs L. L., Schulp A. S., Mateus O., Gonçalves O. A., & Morais M. - L. (2015).  New aristonectine elasmosaurid plesiosaur specimens from the Early Maastrichtian of Angola and comments on paedomorphism in plesiosaurs. Netherlands Journal of Geosciences. FirstView, 1–16., 2 Abstractaraujo_et_al_2015_paedomorphism-libre.pdfWebsite

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.

Strganac, C., Jacobs L. L., Polcyn M. J., Ferguson K. M., Mateus O., Gonçalves O. A., Morais M. - L., & da Silva Tavares T. (2015).  Stable oxygen isotope chemostratigraphy and paleotemperature regime of mosasaurs at Bentiaba, Angola. Netherlands Journal of Geosciences. FirstView, 1–7., 2 Abstractstrganac_etal2015_stable_oxigen_isotopes.pdfWebsite

ABSTRACT Stable oxygen isotope values of inoceramid marine bivalve shells recovered from Bentiaba, Angola, are utilised as a proxy for paleotemperatures during the Late Cretaceous development of the African margin of the South Atlantic Ocean. The δ18O values derived from inoceramids show a long-term increase from –3.2‰ in the Late Turonian to values between –0.8 and –1.8‰ in the Late Campanian. Assuming a constant oceanic δ18O value, an ∼2‰ increase may reflect cooling of the shallow marine environment at Bentiaba by approximately 10°. Bentiaba values are offset by about +1‰ from published records for bathyal Inoceramus at Walvis Ridge. This offset in δ18O values suggests a temperature difference of ∼5° between coastal and deeper water offshore Angola. Cooler temperatures implied by the δ18O curve at Bentiaba coincide with the stratigraphic distribution of diverse marine amniotes, including mosasaurs, at Bentiaba.

Xing, L., Lockley M. G., Marty D., Zhang J., Wang Y., Klein H., McCrea R. T., Buckley L. G., Belvedere M., Mateus O., Gierliński G. D., Piñuela L., Persons, IV S. W., Wang F., Ran H., Dai H., & Xie X. (2015).  An Ornithopod-Dominated Tracksite from the Lower Cretaceous Jiaguan Formation (Barremian–Albian) of Qijiang, South-Central China: New Discoveries, Ichnotaxonomy, Preservation and Palaeoecology. PLoS ONE. 10, e0141059., 10, Number 10: Public Library of Science Abstractlida_et_al_2015_an_ornithopod-dominated_tracksite_from_the.pdfWebsite

The historically-famous Lotus Fortress site, a deep 1.5–3.0-meter-high, 200-meter-long horizonal notch high up in near-vertical sandstone cliffs comprising the Cretaceous Jiaguan Formation, has been known since the 13th Century as an impregnable defensive position. The site is also extraordinary for having multiple tetrapod track-bearing levels, of which the lower two form the floor of part of the notch, and yield very well preserved asseamblages of ornithopod, bird (avian theropod) and pterosaur tracks. Trackway counts indicate that ornithopods dominate (69%) accounting for at least 165 trackmakers, followed by bird (18%), sauropod (10%), and pterosaur (3%). Previous studies designated Lotus Fortress as the type locality of Caririchnium lotus and Wupus agilis both of which are recognized here as valid ichnotaxa. On the basis of multiple parallel trackways both are interpreted as representing the trackways of gregarious species. C. lotus is redescribed here in detail and interpreted to indicate two age cohorts representing subadults that were sometimes bipedal and larger quadrupedal adults. Two other previously described dinosaurian ichnospecies, are here reinterpreted as underprints and considered nomina dubia. Like a growing number of significant tetrapod tracksites in China the Lotus Fortress site reveals new information about the composition of tetrapod faunas from formations in which the skeletal record is sparse. In particular, the site shows the relatively high abundance of Caririchium in a region where saurischian ichnofaunas are often dominant. It is also the only site known to have yielded Wupus agilis. In combination with information from other tracksites from the Jiaguan formation and other Cretaceous formations in the region, the track record is proving increasingly impotant as a major source of information on the vertebrate faunas of the region. The Lotus Fortress site has been developed as a spectacular, geologically-, paleontologically- and a culturally-significant destination within Qijiang National Geological Park.

Araújo, R., Polcyn M. J., Schulp A. S., Mateus O., Jacobs L. L., Gonçalves O. A., & Morais M. - L. (2015).  A new elasmosaurid from the early Maastrichtian of Angola and the implications of girdle morphology on swimming style in plesiosaurs. Netherlands Journal of Geosciences. FirstView, 1–12., 1 Abstractaraujo_et_al_2015_a_new_elasmosaurid_from_the_early_maastrichtian_of_angola.pdfWebsite

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.

Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & de Andrade M. B. (2015).  Addendum to ‘Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)’. Royal Society Open Science. 2, , Number 2: The Royal Society Abstractyoung_et_al_2015_addendum_to_revision_of_the_late_jurassic_teleosaurid_genus_machimosaurus_crocodylomorpha_thalattosuchia.pdfWebsite

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Hendrickx, C., Mateus O., & Araújo R. (2015).  The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627–642. Abstracthendrickx_et_al_2015_theropod_teeth_app.pdfWebsite

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.

Mateus, O., Jacobs L. L., Polcyn M. J., Myers T. S., & Schulp A. S. (2015).  The fossil record of testudines from angola from the turonian to oligocene. Society of Vertebrate Paleontology Annual Meeting. 177., Dallasmateus_et_al_2015_testudines_angola_svp_abstract.pdf
Strganac, C., Jacobs L., Polcyn M., Mateus O., Myers T., Araújo R., Fergunson K. M., Gonçalves A. O., Morais M. L., Schulp A. S., da Tavares T. S., & Salminen J. (2015).  Geological Setting and Paleoecology of the Upper Cretaceous Bench 19 Marine Vertebrate Bonebed at Bentiaba, Angola. Netherlands Journal of Geosciences. 94(1), 121-136. Abstractstrganac_et_al_2014_geological_setting_bentiaba_angola.pdfWebsite

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.

Marzola, M., Russo J., & Mateus O. (2015).  Identification and comparison of modern and fossil crocodilian eggs and eggshell structures. Historical Biology. 27(1), 115-133. Abstractmarzola_et_al_2015_identification_and_comparison_of_modern_and_fossil_crocodilian_eggs_and_eggshell_structures.pdfWebsite

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.

Pereira, B. C., Benton M. J., Ruta M., & Mateus O. (2015).  Mesozoic echinoid diversity in Portugal: Investigating fossil record quality and environmental constraints on a regional scale. Palaeogeography, Palaeoclimatology, Palaeoecology. 424, 132 - 146. Abstractpereira_e_al_2015_mesozoic_echinoids_portugal.pdfWebsite

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.

Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch’s Journal of Vertebrate Palaeontology. 12(1), 1-73. Abstracthendrickx_etal_2015_non_avian_theropods_pjvp12_11.pdfWebsite

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.

Hendrickx, C., Mateus O., & Araújo R. (2015).  A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. e982797. Abstracthendrickx_et_al_2015_theropod_teeth_svp.pdfWebsite

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Polcyn, M. J., Jacobs L. L., Schulp A. S., Mateus O., & Araújo R. (2015).  Tethyan and Weddellian biogeographic mixing in the Maastrichtian of Angola. Annual Meeting of the Society of Vertebrate Paleontology. 196., Dallas, TXpolcyn_etal2015_mix_fauna_angola_svp_abstract.pdf
Polcyn, M. J., Jacobs L. L., Schulp A. S., & Mateus O. (2015).  Trolling the Cretaceous Seas: Marine Amniotes of Two West Coast Margins. Geological Society of America Annual Meeting. 55.: Geological Society of America Abstracts with Programs. Vol. 47, No. 4, p.55 Abstract

In this session we review the Upper Cretaceous marine amniote records from the west coasts of North America and Africa. Recent work by our group in Angola, on the west coast of Africa, has opened up new fossiliferous localities, producing well-preserved turtles, plesiosaurs, and mosasaurs, ranging in age from Late Turonian to Late Maastrichtian. These African localities were deposited in arid latitudes and highly productive upwelling zones along the passive margin of a growing South Atlantic Ocean. The fossil record of Cretaceous marine amniotes from the West Coast of North America is relatively meager when compared to the African record and the prolific fossil beds laid down in the epicontinental seas of the Western Interior Seaway and northern Europe. Nonetheless, these localities provide an important glimpse of a marine ecosystem that developed on the active margins of a deep ocean basin. Historically considered to be depauperate and endemic, the west coast fauna was characterized by unusual forms such as Plotosaurus, arguably one of the most derived mosasaurs; however, in recent years, additional taxa have been described, revealing species diversity and ecological partitioning within these communities and in some cases, faunal interchange with other regions. The large quantity of well-preserved fossils from the west coast of Africa is influenced in part by its paleogeographic position, deposited within highly productive areas of Hadley Cell controlled upwelling zones. By contrast, the North American west coast localities have been deposited in temperate and higher latitudes since the Late Cretaceous. Nonetheless, the North American and African faunas share some common characteristics in a possessing a mix of endemic and more cosmopolitan forms. Habitat partitioning reflected in tooth form and body size is comparable between the Angolan and the North American west coast, and there is remarkable convergence in taxa which appear to exploit certain like-niches.

Clemmensen, L. B., Milàn J., Adolfssen J. S., Estrup E. J., Frobøse N., Klein N., Mateus O., & Wings O. (2015).  The vertebrate-bearing Late Triassic Fleming Fjord Formation of central East Greenland revisited: stratigraphy, palaeoclimate and new palaeontological data. Geological Society, London, Special Publications. 434(1), 31-47. Abstractclemmensenetal2015greenland.pdfWebsite

In Late Triassic (Norian–Rhaetian) times, the Jameson Land Basin lay at 40° N on the northern part of the supercontinent Pangaea. This position placed the basin in a transition zone between the relatively dry interior of the supercontinent and its more humid periphery. Sedimentation in the Jameson Land Basin took place in a lake–mudflat system and was controlled by orbitally forced variations in precipitation. Vertebrate fossils have consistently been found in these lake deposits (Fleming Fjord Formation), and include fishes, dinosaurs, amphibians, turtles, aetosaurs and pterosaurs. Furthermore, the fauna includes mammaliaform teeth and skeletal material. New vertebrate fossils were found during a joint vertebrate palaeontological and sedimentological expedition to Jameson Land in 2012. These new finds include phytosaurs, a second stem testudinatan specimen and new material of sauropodomorph dinosaurs, including osteologically immature individuals. Phytosaurs are a group of predators common in the Late Triassic, but previously unreported from Greenland. The finding includes well-preserved partial skeletons that show the occurrence of four individuals of three size classes. The new finds support a late Norian–early Rhaetian age for the Fleming Fjord Formation, and add new information on the palaeogeographical and palaeolatitudinal distribution of Late Triassic faunal provinces.

Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & De Andrade M. B. (2015).  Addendum to ?Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)?. Royal Society Open Science. 2, , Number 2 Abstract
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Klein, H., Milàn J., Clemmensen L. B., Frobøse N., Mateus O., Klein N., Adolfssen J. S., Estrup E. J., & Wings O. (2015).  Archosaur footprints (cf. Brachychirotherium) with unusual morphology from the Upper Triassic Fleming Fjord Formation (Norian–Rhaetian) of East Greenland. Geological Society, London, Special Publications. 434, AbstractWebsite

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

Marzola, M., Mateus O., Schulp A. S., Jacobs L. L., Polcyn M. J., Pervov V., Goncalves A. O., & Morais M. L. (2015).  Comparative anatomy and systematics of Cretaceous mammal tracks of Angola. 13th Annual Meeting of the European Association of Vertebrate Palaeontologists - EAVP 2015. , Opole, Poland Abstract
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Marzola, M., Mateus O., Schulp {A. S. }, Jacobs {L. L. }, Polcyn {M. J. }, Pervov V., Goncalves {A. O. }, & Morais {M. L. } (2015).  Comparative anatomy and systematics of Cretaceous mammal tracks of Angola. 35. Abstract
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Hansen, B. B., Milàn J., Clemmensen L. B., Adolfssen J. S., Estrup E. J., Klein N., Mateus O., & Wings O. (2015).  Coprolites from the Late Triassic Kap Stewart Formation, Jameson Land, East Greenland: morphology, classification and prey inclusions. Geological Society, London, Special Publications. 434, AbstractWebsite

A large collection of vertebrate coprolites from black lacustrine shales in the Late Triassic (Rhaetian–Sinemurian) Kap Stewart Formation, East Greenland is examined with regard to internal and external morphology, prey inclusions, and possible relationships to the contemporary vertebrate fauna. A number of the coprolites were mineralogically examined by X-ray diffraction (XRD), showing the primary mineral composition to be apatite, clay minerals, carbonates and, occasionally, quartz in the form of secondary mineral grains. The coprolite assemblage shows multiple sizes and morphotypes of coprolites, and different types of prey inclusions, demonstrating that the coprolite assemblage originates from a variety of different producers.Supplementary material: A description of the size, shape, structure, texture, contents and preservation of the 328 specimens is available at https://doi.org/10.6084/m9.figshare.c.2134335

Mateus, O., Jacobs L. L., Polcyn {M. J. }, Myers T. S., & Schulp A. S. (2015).  The fossil record of testudines from Angola from the Turonian to Oligocene. : Journal of Vertebrate Paleontology Abstract
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Marzola, M., Russo J., & Mateus O. (2015).  Identification and comparison of modern and fossil crocodilian eggs and eggshell structures. Historical Biology. 27, 115-133., Number 1 Abstract
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Pereira, B. C., Benton M. J., Ruta M., & Mateus O. (2015).  Mesozoic echinoid diversity in Portugal: Investigating fossil record quality and environmental constraints on a regional scale. Palaeogeography, Palaeoclimatology, Palaeoecology. 424, 132-146. Abstract
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Brusatte, S. L., Butler R. J., Mateus O., & Steyer J. S. (2015).  A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls. Journal of Vertebrate Paleontology. 35, , Number 3 Abstract
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Hendrickx, C., Aráujo R., & Mateus O. (2015).  The non-avian theropod quadrate I: Standardized terminology with an overview of the anatomy and function. PeerJ. 2015, , Number 9 Abstract
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Xing, L., Lockley M. G., Marty D., Zhang J., Wang Y., Klein H., McCrea R. T., Buckley L. G., Belvedere M., Mateus O., Gierli?ski G. D., Piñuela L., Persons W. S., Wang F., Ran H., Dai H., & Xie X. (2015).  An ornithopod-dominated tracksite from the lower Cretaceous Jiaguan Formation (Barremian-Albian) of Qijiang, South-Central China: New discoveries, ichnotaxonomy, preservation and palaeoecology. PLoS ONE. 10, , Number 10 Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch’s Journal of Vertebrate Palaeontology. 12, 1-73. AbstractWebsite

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.

Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch{'}s Journal of Vertebrate Palaeontology. 12, 1-73., Number 1 Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. \á\}vio (2015).  An overview of non-avian theropod discoveries and classification. PalArch\’\}s Journal of Vertebrate Palaeontology. 12, 1-73. Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. \á\}vio (2015).  An overview of non-avian theropod discoveries and classification. PalArch\’\}s Journal of Vertebrate Palaeontology. 12, 1-73. Abstract
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Hendrickx, C., Mateus O., & Araujo R. (2015).  A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 35, , Number 5 Abstract
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Tschopp, E., Mateus O., & Benson R. B. J. (2015).  A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). {PeerJ}. 3, e857.: {PeerJ} AbstractWebsite
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Polcyn, {M. J. }, Jacobs L. L., Schulp A. S., & Mateus O. (2015).  Tethyan and Weddellian biogeographic mixing in the Maastrichtian of Angola. : Journal of Vertebrate Paleontology Abstract
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Polcyn, {M. J. }, Jacobs {L. L. }, Schulp {A. S. }, & Mateus O. (2015).  Trolling the Cretaceous Seas: Marine Amniotes of Two West Coast Margins. : Geological Society of America Abstracts with Programs. Vol. 47, No. 4, p.55 Abstract

In this session we review the Upper Cretaceous marine amniote records from the west coasts of North America and Africa. Recent work by our group in Angola, on the west coast of Africa, has opened up new fossiliferous localities, producing well-preserved turtles, plesiosaurs, and mosasaurs, ranging in age from Late Turonian to Late Maastrichtian. These African localities were deposited in arid latitudes and highly productive upwelling zones along the passive margin of a growing South Atlantic Ocean. The fossil record of Cretaceous marine amniotes from the West Coast of North America is relatively meager when compared to the African record and the prolific fossil beds laid down in the epicontinental seas of the Western Interior Seaway and northern Europe. Nonetheless, these localities provide an important glimpse of a marine ecosystem that developed on the active margins of a deep ocean basin. Historically considered to be depauperate and endemic, the west coast fauna was characterized by unusual forms such as Plotosaurus, arguably one of the most derived mosasaurs; however, in recent years, additional taxa have been described, revealing species diversity and ecological partitioning within these communities and in some cases, faunal interchange with other regions. The large quantity of well-preserved fossils from the west coast of Africa is influenced in part by its paleogeographic position, deposited within highly productive areas of Hadley Cell controlled upwelling zones. By contrast, the North American west coast localities have been deposited in temperate and higher latitudes since the Late Cretaceous. Nonetheless, the North American and African faunas share some common characteristics in a possessing a mix of endemic and more cosmopolitan forms. Habitat partitioning reflected in tooth form and body size is comparable between the Angolan and the North American west coast, and there is remarkable convergence in taxa which appear to exploit certain like-niches.

Clemmensen, L. B., Milàn J., Adolfssen J. S., Estrup E. J., Frobøse N., Klein N., Mateus O., & Wings O. (2015).  The vertebrate-bearing Late Triassic Fleming Fjord Formation of central East Greenland revisited: stratigraphy, palaeoclimate and new palaeontological data. Geological Society, London, Special Publications. 434, AbstractWebsite

In Late Triassic (Norian–Rhaetian) times, the Jameson Land Basin lay at 40° N on the northern part of the supercontinent Pangaea. This position placed the basin in a transition zone between the relatively dry interior of the supercontinent and its more humid periphery. Sedimentation in the Jameson Land Basin took place in a lake–mudflat system and was controlled by orbitally forced variations in precipitation. Vertebrate fossils have consistently been found in these lake deposits (Fleming Fjord Formation), and include fishes, dinosaurs, amphibians, turtles, aetosaurs and pterosaurs. Furthermore, the fauna includes mammaliaform teeth and skeletal material. New vertebrate fossils were found during a joint vertebrate palaeontological and sedimentological expedition to Jameson Land in 2012. These new finds include phytosaurs, a second stem testudinatan specimen and new material of sauropodomorph dinosaurs, including osteologically immature individuals. Phytosaurs are a group of predators common in the Late Triassic, but previously unreported from Greenland. The finding includes well-preserved partial skeletons that show the occurrence of four individuals of three size classes. The new finds support a late Norian–early Rhaetian age for the Fleming Fjord Formation, and add new information on the palaeogeographical and palaeolatitudinal distribution of Late Triassic faunal provinces.

2014
Marzola, M., Mateus O., Russo J., & Milàn J. (2014).  Comparison of modern and fossil Crocodylomorpha eggs and contribution to the oophylogeny of Amniota. XII Annual Meeting of the European Association of Vertebrate Palaeontologists. , p. 192, Regione Piemonte: European Association of Vertebrate Palaeontologists. Museo Regionale di Scienze Naturalimarzola_et_al._2014_comparison_of_modern_and_fossil_crocodylomorpha_eggs_and_contribution_to_the_oophylogeny_of_amniota-_eavp_2014.pdf
Hendrickx, C., Araújo R., & Mateus O. (2014).  The nonavian theropod quadrate II: systematic usefulness, major trends and cladistic and phylogenetic morphometrics analyses. PeerJ PrePrints. 2, e380v2., 2014 AbstractWebsite

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.

Polcyn, M. J., Jacobs L. L., Strganac C., Mateus O., Myers T. S., May S., Araújo R., Schulp A. S., & Morais M. L. (2014).  Geological and paleoecological setting of a marine vertebrate bonebed from the Lower Maastrichtian at Bentiaba, Angola. Secondary Adaptation of Tetrapods to Aquatic Life. , 2-4 Jun 2014, Washington DC, USA
Mateus, O. (2014).  Comparison of modern and fossil Crocodylomorpha eggs and contribution to the oophylogeny of Amniota. Annual Meeting of the European Association of Vertebrate Palaeontologists. XII Annual Meeting of the European Association of Vertebrate Palaeontologists, 192., 1 Abstract
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Mateus, O. (2014).  Cracking dinosaur endothermy: paleophysiology unscrambled. NA, , 1 Abstract

The amniote eggshell functions as a respiratory structure adapted for the optimal transmission of respiratory gasses to and from the embryo according to its physiological requirements. Therefore amniotes with higher oxygen requirements, such as those that sustain higher metabolic rates, can be expected to have eggshells that can maintain a greater gas flux to and from the egg. Studies of extant amniotes have found that eggshells of reduced porosity impose a limit on the metabolic rate of the offspring. Here we show a highly significant relationship between metabolic rates and eggshell porosity in extant amniotes that predicts highly endothermic metabolic rates in dinosaurs. This study finds the eggshell porosity of extant endotherms to be significantly higher than that of extant ectotherms. Eggshell porosity values of dinosaurs are found to be significantly higherthan that of extant ectotherms, but not extant endotherms. Dinosaur eggshells are commonly preserved in the fossil record, and porosity may be readily identified and measured. This provides a simple tool to identify metabolic rates in extinct egg-laying tetrapods whose eggs possessed a mineralized shell

Mateus, O. (2014).  Degradation processes and consolidation of Late Jurassic sandstone dinosaur tracks in museum environment (Museum of Lourinhã, Portugal). Geophysical Research Abstracts. Geophysical Research Abstracts, EGU2014–9026–1, 2014., 1 Abstract
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Mateus, O. (2014).  Elephas and other vertebrate fossils near Taghrout, Morocco. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 178., 1 Abstract
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Mateus, O. (2014).  Geological and paleoecological setting of a marine vertebrate bonebed from the Lower Maastrichtian at Bentiaba, Angola. Proceedings of the Secondary Adaptation of Tetrapods to Aquatic Life. NA., 1 Abstract

A single, geographically and temporally restricted horizon at Bentiaba, Angola (14.3° S), preserves a concentration of skeletons and isolated elements representing sharks, rays, bony fish, at least three species of turtles, two species of plesiosaurs, at least five species of mosasaurs, and rare volant and terrestrial forms. The concentration, referred to as the Bench 19 Fauna, formed on a narrow continental shelf at paleolatitude 24°S as predicted by paleomagnetic data and confirmed by plate motion models. The shelf evolved as a transform passive margin along faults associated with the opening of the South Atlantic. Latitude 24°S falls today along the coast of northern Namibia, an area of intense upwelling and hyperarid coastal desert. The Namibe Basin in southern Angola is separated from the Walvis Basin of Namibia by the Walvis Ridge, and the continental shelf in northern Namibia is eight times the width of that at Bentiaba. However, the sediment entombing the fossils at Bentiaba is an immature feldspathic sand, shown by detrital zircon ages to be derived from nearby exposed granitic shield rocks, suggesting similar climatic and drainage conditions between the two regions. Temporal control of the Bentiaba section is provided by magnetostratigraphy and stable carbon isotope chemostratigraphy anchored by an Ar40/Ar39radiometric date on basalt. The age of Bench 19 is constrained to chron C32n.1n and thus falls between 71.4 and 71.64 Ma. Massive bedding without hummocky cross-bedding or other sedimentary structures indicates deposition in shallow water below wave base. δ18O analysis of bivalve shells indicates a water temperature of 18° C immediately below Bench 19. Nearest neighbor distance peaks at 5 m (n=19

Hendrickx, C., & Mateus O. (2014).  Torvosaurus gurneyi n. sp., the largest terrestrial predator from Europe, and a proposed terminology of the maxilla anatomy in nonavian theropods. PLoS ONE. 9, e88905., 03, Number 3: Public Library of Science Abstracthendrickx_mateus_2014_torvosaurus_portugal.pdfWebsite

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.

Hendrickx, C., & Mateus O. (2014).  Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa. 3759, 1-74. Abstracthendrickx__mateus_2014._abelisauridae_dinosauria_theropoda_from_the_late_jurassic_of_portugal.pdf

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.

Strganac, C., Salminen J., Jacobs L. L., Ferguson K. M., Polcyn M. J., Mateus O., Schulp A. S., Morais M. L., Tavares T. S., & Gonçalves A. O. (2014).  Carbon isotope stratigraphy and 40Ar/39Ar age of the Cretaceous South Atlantic coast, Namibe Basin, Angola. Journal of African Earth Sciences. onine, 1-11. Abstractstrganac_et_al_2014_carbon_isotope_stratigraphy_magnetostratigraphy_and_40ar_39ar_age_of.pdfWebsite

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.