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Fernandes, A. E., Beccari V., Kellner A. W. A., & Mateus O. (2023).  A new gnathosaurine (Pterosauria, Archaeopterodactyloidea) from the Late Jurassic of Portugal. PeerJ. 11, e16048. Abstractfernandes_et_al_2023_lusognathus_peerj-16048.pdfWebsite

An incomplete, yet remarkably-sized dentated rostrum and associated partial cervical vertebrae of a pterosaur (ML 2554) were recently discovered from the Late Jurassic (Late Kimmeridgian-Early Tithonian) Lourinhã Formation of Praia do Caniçal, of central west Portugal. This specimen exhibits features such as a spatulated anterior expansion of the rostrum, robust comb-like dentition, and pronounced rims of the tooth alveoli, indicating gnathosaurine affinities. Based on its further unique tooth and dentary morphology, a new genus and species, \textit{Lusognathus almadrava} gen. et spec. nov., is proposed, making this the first named pterosaur species found within Portugal. The presence of this taxon adds yet another element to the fluvio-deltaic lagoonal environment that has been suggested as representative of the Lourinhã Formation in the Late Jurassic, further contributing to the diversity and distribution of gnathosaurines worldwide.

Ferrari, L., Rotatori F. M., Camilo B., Moreno-Azanza M., & Mateus O. (2022).  New specimen of dryomorphan (Ornithischia, Iguanodontia) remains from the Upper Jurassic of Portugal. Abstract book of the XIX Annual Conference of the European Association of Vertebrate Palaeontologists, Benevento/Pietraroja, Italy, 27th June-2nd July 2022.. 61-62.: PalaeoVertebrata, Special Volume 1- 2022, 224. Doi: 10.18563/pv.eavp Abstractferrarietal_2022.pdf

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Foth, C., Evers S., Pabst B., Mateus O., Flisch A., Patthey M., & Rauhut O. W. M. (2015).  New insights into the lifestyle of Allosaurus (Dinosauria: Theropoda) based on another specimen with multiple pathologies. PeerJ PrePrints. 3, e824v1., 2015 Abstractfoth_et_al_2015_peerj-preprints-824.pdfWebsite

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).

Foth, C., Evers S. W., Pabst B., Mateus O., Flisch A., Patthey M., & Rauhut O. W. M. (2015).  New insights into the lifestyle of \\textitAllosaurus (Dinosauria: Theropoda) based on another specimen with multiple pathologies. PeerJ. 3, e940., 5 AbstractWebsite

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).

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Gaspar, A., Avelar T., & Mateus O. (2007).  Criacionismo e Sociedade no Séc. XX.  Evolução e Criacionismo: Uma Relação Impossível. 133-160., Lisboa Abstract
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Gaspar, A., Avelar T., & Mateus O. (2007).  Criacionismo e Sociedade no Séc. XX. (Avelar, T., O. Mateus, Almada, F., Gaspar, A., Ed.).Evolução e Criacionismo: Uma Relação Impossível. 133-160., Lisboa: Quasi ed. gasparavelarmateus2007evoluoecriacio.pdf
Gaspar, A., Mateus O., & Almada F. (2007).  Os argumentos criacionistas em face da evidência científica. Evolução e Criacionismo: Uma Relação Impossível. 197-237., Lisboa Abstract
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Gaspar, A., Mateus O., & Almada F. (2007).  Os argumentos criacionistas em face da evidência científica. Evolução e Criacionismo: Uma Relação Impossível. 197-237., Lisboa: Quasi ed.gasparmateusealmada2007argumentosevo.pdf
Graf, J., Jacobs L. L., Polcyn M. J., Mateus O., & Schulp A. S. (2011).  New fossil whales from Angola. 71st Annual Meeting of the Society of Vertebrate Paleontology. 119., Jan: Abstracts of the 71st Annual Meeting of the Society of Vertebrate Paleontologygraf_et_al_mateus_2011_fossil_whales_from_angola_svp11abstracts.pdf
Guillaume, A. R. D., Moreno-Azanza M., & Mateus O. (2022).  New species of Albanerpetontidae (Lissamphibia) from the Lourinhã Formation. XXXVII Jornadas SEP y V Congreso Ibérico de Paleontología. 94., Cuenca: ISBN 13: 978-84-09-44478-6 Abstractguillaume_et_al_2022_albanerpetontidae_jornadas_sep_cuenca_abstract_2022.pdf

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Guillaume, A., Costa F., & Mateus O. (2017).  Skin impressions on stegosaur tracks from the Upper Jurassic of Portugal. 191–195. Abstract
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Guillaume, A. R., Moreno-Azanza M., & Mateus O. (2019).  New lissamphibian material from the Lourinhã Formation (Late Jurassic, Portugal). Journal of Vertebrate Paleontology, Program and Abstracts. 112.guillaume_et_al_2019_svp_abstract.pdf
Guillaume, A. R. D., Natário C., Mateus O., & Moreno-Azanza M. (2022).  Plasticity in the morphology of the fused frontals of Albanerpetontidae (Lissamphibia; Allocaudata). Historical Biology. 1 - 18., 2022: Taylor & Francis Abstractplasticity_in_the_morphology_of_the_fused_frontals_of_albanerpetontidae_lissamphibia_allocaudata.pdfWebsite

ABSTRACTAlbanerpetontidae form an enigmatic extinct group of lissamphibians, ranging from the early Bathonian to the early Pleistocene. The Upper Jurassic outcrops of Portugal yield a large collection of material, suitable for addressing the intraspecific variation in and diagnostic potential of the characteristic fused frontals. We revise 58 specimens from the Guimarota beds of the Kimmeridgian Alcobaça Formation and describe 62 new frontal bones from the Kimmeridgian ? Tithonian Lourinh? Formation. Smaller specimens exhibit a vermicular dorsal ornamentation, while it is polygonal in larger specimens and other albanerpetontids. Compared to small specimens, larger specimens display: (1) larger ventrolateral crests extending posteriorly after the parietal margin; (2) a relatively shorter internasal process; (3) a frontal width across posterior edges relatively smaller; and (4) a ventromedian crest less pronounced. Morphometric analyses suggest a single species with different ontogenetic stages. Specimens are attributed to aff. Celtedens sp., based on a bell-shaped outline with a curved orbital margin (although variable in Portuguese specimens), and a flabellate, bulbous-shaped internasal process. The species is more similar to C. megacephalus than C. ibericus, but its phylogenetic position comprises an unresolved trichotomy. Our results show that intraspecific variation and homoplasy render the fused frontal non-diagnostic below the generic level.

Guillaume, A. R. D., Moreno-Azanza M., Puértolas-Pascual E., & Mateus O. (2018).  Crocodylomorph teeth from the Lourinhã Formation, Portugal (Late Jurassic). XVI Annual Meeting of the European Association of Vertebrate Palaeontologists. 80., Caparica, Portugal June 26th-July 1st, 2018 Abstractguillaume_et_al_2018_eavp_abstract.pdf

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Guillaume, A. R. D., Ezquerro L., Moreno-Azanza M., & Mateus O. (2023).  Vertebrate microfossil assemblages from the Lourinhã Formation: a sneak peek on the paleoecology of the Late Jurassic in Portugal. The Anatomical Record. 306(#S1), 118-121.: Conference: 14th Symposium on Mesozoic Terrestrial Ecosystems and BiotaAt … Abstractguillaume_et_al_2023_vertebrate_microfossil.pdf

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Guillaume, A. R. D., Natário C., Moreno-Azanza M., & Mateus O. (2021).  New occurences of Celtedens (Lissamphibia: Albanerpetontidae) from the Lourinhã Formation. XVIII Conference of the European Association of Vertebrate Palaeontologists. 87. Abstractguillaume_et_al_2021_celtedens.pdf

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Guillaume, A., Costa F., & Mateus O. (2017).  Skin impressions on stegosaur tracks from the Upper Jurassic of Portugal. Abstract book of the XV Encuentro de Jóvenes Investigadores en Paleontología/XV Encontro de Jovens Investigadores em Paleontologia, Lisboa, 428 pp.. 191-195. Abstractguillaume_2017_skin.pdf

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Guillaume, A. R. D., Moreno-Azanza M., & Mateus O. (2018).  Microvertebrates from the Lourinhã Formation (Late Jurassic, Portugal). 1st Palaeontological Virtual Congress. online. Abstractguillaume-et-al_pvc2018_abstract.pdf

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Guillaume, A. R., Moreno-Azanza M., & Mateus O. (2019).  New lissamphibian material from the Lourinhã Formation (Late Jurassic, Portugal). Journal of Vertebrate Paleontology, Program and Abstracts. 112. Abstract
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Guillaume, A. R. D., Mateus O., & Moreno-Azanza M. (2023).  The Albanerpetontidae (Lissamphibia) from the Lourinhã Formation (Upper Jurassic, Portugal). The Anatomical Record. 306(#S1), 116-118. Abstractguillaume_et_al_2023_albanerpetontidae_lissamphibia.pdfWebsite

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Guillaume, A. R. D., Natário C., Moreno-Azanza M., & Mateus O. (2021).  New material ascribed to Albanerpetontidae (Lissamphibia; Allocaudata) from the Lourinhã Formation: evidence for intraspecific variation. XIX Encontro de Jovens Investigadores em Paleontologia. 50-51. Abstractguillaume_et_al_2021_albanerpetontidae.pdf

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Guillaume, A., Costa F., & Mateus O. (2017).  Skin impressions on stegosaur tracks from the Upper Jurassic of Portugal. Abstract book of the XV Encuentro de Jóvenes Investigadores en Paleontolog{\'ıa/XV Encontro de Jovens Investigadores em Paleontologia, Lisboa, 428 pp.. 191–195. Abstract
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Guillaume, A. R. D., Moreno-Azanza M., Puértolas-Pascual E., & Mateus O. (2018).  Crocodylomorph teeth from the Lourinhã Formation, Portugal (Late Jurassic). XVI Annual Meeting of the European Association of Vertebrate Palaeontologists. 80., Caparica, Portugal June 26th-July 1st, 2018 Abstract

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Guillaume, A. R. D., Costa F., & Mateus O. (2022).  Stegosaur tracks from the Upper Jurassic of Portugal: new occurrences and perspectives. Ciências da Terra / Earth Sciences Journal. 20(1), 37-60. Abstractguillaumeetal.pdf

The record of Late Jurassic stegosaur tracks from the Lourinhã Formation (Kimmeridgian-Tithonian) is here revised. Thirty-eight dinosaur tracks, preserved as natural infill casts, are here reported, and thirty-two of them are attributed to the ichnogenus Deltapodus. Four of those present impressions of skin, with polygonal scales and random pattern. Deltapodus is the most common ichnogenus in the track record of the Lourinhã Formation. The sizes and shape suggest one single dacentrurine trackmaker, which could be Miragaia longicollum, also common in the same horizons.

Guillaume, A. R. D., Moreno-Azanza M., Puértolas-Pascual E., & Mateus O. (2020).  Palaeobiodiversity of crocodylomorphs from the Lourinhã Formation based on the tooth record: insights into the palaeoecology of the Late Jurassic of Portugal. Zoological Journal of the Linnean Society. 189(2), 549–583., 11 Abstractguillaume_et_al_palaeobiodiversity_of_crocodylomorphs_from_the.pdfWebsite

{Crocodylomorphs were a diverse clade in the Late Jurassic of Portugal, with six taxa reported to date. Here we describe 126 isolated teeth recovered by screen-washing of sediments from Valmitão (Lourinhã, Portugal, late Kimmeridgian–Tithonian), a vertebrate microfossil assemblage in which at least five distinct crocodylomorph taxa are represented. Ten morphotypes are described and attributed to five clades (Lusitanisuchus, Atoposauridae, Goniopholididae, Bernissartiidae and an undetermined mesoeucrocodylian). Four different ecomorphotypes are here proposed according to ecological niches and feeding behaviours: these correspond to a diet based on arthropods and small vertebrates (Lusitanisuchus and Atoposauridae), a generalist diet (Goniopholididae), a durophagous diet (Bernissartiidae) and a carnivorous diet. Lusitanisuchus mitracostatus material from Guimarota is here redescribed to achieve a better illustration and comparison with the new material.This assemblage shares similar ecomorphotypes with other Mesozoic west-central European localities, where a diversity of crocodylomorphs lived together, avoiding direct ecological competition through niche partitioning. The absence of large marine crocodylomorphs, present in other contemporaneous assemblages, is here interpreted as evidence that the Valmitão assemblage was deposited in a freshwater environment, although sample bias cannot be completely ruled out. These affinities are further supported by the presence of lanceolate and leaf-shaped teeth associated with continental clades.}

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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

Hansen, B. B., Milàn J., Clemmensen L. B., Adolfssen J. S., Estrup E. J., Klein N., Mateus O., & Wings O. (2016).  Coprolites from the Late Triassic Kap Stewart Formation, Jameson Land, East Greenland: morphology, classification and prey inclusions. Geological Society, London, Special Publications. 434(1), 49-69. Abstracthansen_et_al_2015_coprolites_from_the_late_triassic_kap_stewart_formation_jameson_land_east_greenland.pdfWebsite

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

Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. 28 (3, Supplement), 89A-90A. Journal of Vertebrate Paleontology. 28, 89-90., Number Suppl. to 3 Abstract
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Hayashi, S., Redelstorff R., Mateus O., Watabe M., & Carpenter K. (2014).  Gigantism of stegosaurian osteoderms. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 145.hayashi_et_al_2014_gigantism_of_stegosaurian_osteoderms.pdf
Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. 28 (3, Supplement), 89A-90A. Journal of Vertebrate Paleontology. 28, 89–90., Number Suppl. to Abstract
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Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. Journal of Vertebrate Paleontology. 28(3, Supplement), 89A-90A., Number Suppl. to 3 Abstracthayashi_et_al_2008_histology_stegosaurs_defensive_weapons_of_thyreophoran_dinosaurs-_histological_comparisons_and_structural_differences_in_spikes_and_clubs_of_ankylosaurs_and_stegosaurs.pdfWebsite

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

Hendrickx, C., Araújo R., & Mateus O. (2012).  The nonavian theropod quadrate: systematics usefulness, major trends and phylogenetic morphometrics analysis. : Journal of Vertebrate Paleontology, Program and Abstracts, 2012, p.110. ISSN 1937-2809 Abstracthendrickx_araujo__mateus_2012_quadrate_theropods_svp_2012_abstract_book_meeting_abstracts.pdf.pdf

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

Hendrickx, C., Mateus O., Araújo R., & Choiniere J. (2019).  The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica. 22, , Number 3 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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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.

Hendrickx, C., Mateus O., & Buffetaut E. (2016).  Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.. PLoS ONE. 11, e0144695., 01, Number 1: Public Library of Science Abstracthendrickx_et_al_2016_morphofunctional_analysis_of_the_quadrate_of_spinosauridae_dinosauria.pdfWebsite

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

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., & Buffetaut E. (2016).  Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.. PLoS ONE. 11, e0144695., 01, Number 1: Public Library of Science AbstractWebsite

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

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.

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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Hendrickx, C., Bell P. R., Pittman M., Milner A. R. C., Cuesta E., O'Connor J., Loewen M., Currie P. J., Mateus O., Kaye T. G., & Delcourt R. (2022).  Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs. Biological Reviews. , Number n/a Abstracthendrickxetal.2021.morphologyanddistributionofscales.pdfWebsite

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.

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., Mateus O., Araújo R., & Choiniere J. (2019).  The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica. 22(3), 1-110. Abstractthe_distribution_of_dental_features_in_non-avian_t.pdfWebsite

Isolated theropod teeth are some of the most common fossils in the dinosaur fossil record and are continually reported in the literature. Recently developed quantitative methods have improved our ability to test the affinities of isolated teeth in a repeatable framework. But in most studies, teeth are diagnosed on qualitative characters. This can be problematic because the distribution of theropod dental characters is still poorly documented, and often restricted to one lineage. To help in the identification of isolated theropod teeth, and to more rigorously evaluate their taxonomic and phylogenetic potential, we evaluated dental features in two ways. We first analyzed the distribution of 34 qualitative dental characters in a broad sample of taxa. Functional properties for each dental feature were included to assess how functional similarity generates homoplasy. We then compiled a quantitative data matrix of 145 dental characters for 97 saurischian taxa. The latter was used to assess the degree of homoplasy of qualitative dental characters, address longstanding questions on the taxonomic and biostratigraphic value of theropod teeth, and explore the major evolutionary trends in the theropod dentition.

In smaller phylogenetic datasets for Theropoda, dental characters exhibit higher levels of homoplasy than non-dental characters, yet they still provide useful grouping information and optimize as local synapomorphies of smaller clades. In broader phylogenetic datasets, the degree of homoplasy displayed by dental and non-dental characters is not significantly different. Dental features on crown ornamentations, enamel texture and tooth microstructure have significantly less homoplasy than other dental features and can be used to identify many theropod taxa to ‘family’ or ‘sub-family’ level, and some taxa to genus or species. These features should, therefore, be a priority for investigations seeking to classify isolated teeth.

Our observations improve the taxonomic utility of theropod teeth and in some cases can help make isolated teeth useful as biostratigraphic markers. This proposed list of dental features in theropods should, therefore, facilitate future studies on the systematic paleontology of isolated teeth.