Publications in the Year: 2022

Conference Paper

Estraviz López, D, Mateus O.  2022.  Bearing the weight: Technical recovery of brown bear fossils (Ursus arctos) from Algar do Vale da Pena, Portugal. XX EJIP. , Cañaveral de León Abstractestraviz-lpezmateus2022bearingtheweighttechnicalrecoveryofbrownbearfossilsursusarctosfromalgardovaledapenaportugal.pdf

The Algar do Vale da Pena is a cave that was discovered during limestone mining works around 1980, at the Natural Park of Serras de Aire e Candeeiros, Portugal. The entrance to the cave, through the collapsed roof, is a 30 meter near-vertical well, inaccessible without means of vertical progression and specialized equipment. Soon after the discovery of the cave, bone material belonging to bears and hundreds of bear claw marks were located inside it. Nevertheless, the technical descent and the extraordinary difficulties of retrieving the calcite flowstones that encased the bones prevented scientists from undertaking excavations for nearly 40 years. Since 2016 a series of expeditions by a team of paleontologists from NOVA University and Museum of Lourinhã recovered more than 20 disarticulated remains from five bone accumulations, separated more than five meters apart, which given the level of erosion and breakage, are para-autochthonous in origin. These are interpreted as belonging to different individuals. Morphological characters, like a relatively elongated third phalanx, the clear separation of paracone-metacone of the first upper molar and the ventral position of the mandibular condyle; as well as numerous morphometric characters, allow to ascribe the remains to Ursus arctos. Nevertheless, the main finding from the cave is a complete skull with an articulated mandible; which denotes absence of transport, encased within a 180 kg calcite flowstone, including other skeletal remains as well, which could not be extracted from the cave until 2021. This block was removed with the help of members of several speleological groups. It promises a wealth of information about fossil brown bears, being the most complete bear fossil ever recovered in Portugal. It is currently being dated trough magnetostratigraphy and prepared. We consider this intervention as an extraordinary example of citizen science applied to paleontology

Conference Proceedings

Rotatori, FM, Camilo B, Bertozzo F, Mateus O, Moreno-Azanza M.  2022.  A basal ankylopollexian dinosaur from the Late Jurassic of Portugal and its implications for iguanodontian diversity. Abstract book of the XIX Annual Conference of the European Association of Vertebrate Palaeontologists, Benevento/Pietraroja, Italy, 27th June-2nd July 2022. :170-171.: PalaeoVertebrata, Special Volume 1- 2022, 224. Doi: 10.18563/pv.eavp Abstractrotatorietal_2022.pdf


Puértolas‐Pascual, E, Serrano‐Martínez A, Kuzmin IT, Mateus O.  2022.  Neuroanatomy of the Cenomanian crocodylomorph Portugalosuchus. XIX Annual Conference of the European Association of Vertebrate Paleontologists. :170-171. Abstractpuertolas-pascoal_et_al_2022_portugalosuchus_eavp_2022_abstract_volume.pdf


Moreno-Azanza, M, Pérez-Pueyo M, Puértolas-Pascual E, Núñez-Lahuerta C, Mateus O, Bauluz B, Bádenas B, Canudo JI.  2022.  Cáscaras de huevo de los últimos cocodrilomorfos del Cretácico (Huesca, España). XXXVII Jornadas de Paleontología SEP - V Congreso Ibérico de Paleontología. :p.119. Abstract2022_moreno-azanza_etal_sep.pdf


Saleiro, A, Guillaume ARD, Rotatori FM, Ríos-Ibañez M, López ED, Conti S, Martino R, Puértolas-Pascual E, Mateus O, Moreno-Azanza M.  2022.  A beta taxonomy approach to Late Jurassic and Early Cretaceous dinosaur assemblages. Abstract book of the XIX Annual Conference of the European Association of Vertebrate Palaeontologists, Benevento/Pietraroja, Italy, 27th June-2 nd July 2022.. :176-177.: PalaeoVertebrata, Special Volume 1- 2022, 224. Doi: 10.18563/pv.eavp2022 Abstractsaleiroetal_2022-eavp.pdf


Puértolas-Pascual, E, Serrano-Martínez A, Kuzmin IT, Mateus O.  2022.  Paleoneuroanatomía de Portugalosuchus, un cocodrilomorfo eusuquio del Cenomaniense de Portugal. XXXVII Jornadas SEP y V Congreso Ibérico de Paleontología. (Libro de resúmenes):153., Cuenca: ISBN 13: 978-84-09-44478-6 Abstractpuertolas-pascula_et_al_2022_portugalosuchus_jornadas_sep_cuenca_abstract_2022.pdf


López-Rojas, V, Puértolas-Pascual E, Marinheiro J, Mateus O, Mateus S.  2022.  A new goniopholidid skull from the Late Jurassic of Lourinhã, Portugal. XXVII Jornadas SEP e V Congreso Ibérico de Paleontologia. :101., Cuencalopez-rojas_et_al_2022_goniopholis_jornadas_sep_cuenca_abstract_2022.pdf
Guillaume, ARD, Moreno-Azanza M, Mateus O.  2022.  New species of Albanerpetontidae (Lissamphibia) from the Lourinhã Formation. XXXVII Jornadas SEP y V Congreso Ibérico de Paleontología. (Libro de resúmenes):94., Cuenca: ISBN 13: 978-84-09-44478-6 Abstractguillaume_et_al_2022_albanerpetontidae_jornadas_sep_cuenca_abstract_2022.pdf


Ríos, M, Estraviz-López D, Martino R, Lohmann P, Mateus O, Solounias N.  2022.  The first Sivatheriine (Artiodactyla, Giraffidae) from the Middle Miocene of Portugal. XIX Annual conference of the European Association of Vertebrate Palaeontologists (19th EAVP)At: Benevento, Italy. :167. Abstracteavp_2022_abstractvolume-176.pdf


Martino, R, Ríos M, Mateus O, Rook L, Pandolf L.  2022.  New insights into the hippopotamid (Mammalia, Hippopotamidae) from the Casino Basin (Tuscany, Italy). XIX Annual conference of the European Association of Vertebrate Palaeontologists (19th EAVP)At: Benevento, Italy. :119. Abstract
Fernandes, AF, Beccari V, Kellner AWA, Mateus O.  2022.  A new Gnathosaurine (Archaeopterodactyloidea, Pterosauria) from the Late Jurassic of Portugal. XIX Annual Conference of the European Association of Vertebrate Palaeontologists. :56-57. Abstractfernandes_et_al_2022_pterosaur_eavp_2022_abstract_volume.pdf


Ferrari, L, Rotatori FM, 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


Lategano, F, Conti S, Costa F, Mateus O, Lozar F.  2022.  Fighter or bluffer: comparison of bending and compression in models of the caudal spines of dacentrurine and stegosaurine stegosaurs. XIX Annual Conference of the European Association of Vertebrate Paleontologists. :113. Abstract

Journal Article

Reboleira, A, Mateus O.  2022.  Geologia, Grutas e Fauna Subterrânea do Planalto das Cesaredas, Portugal. Captar. 11:1-19.: DOI: Abstractreboleira_e_mateus_2022.pdf.pdf


Conti, S, Tschopp E, Mateus O, Zanoni A, Masarati P, Sala G.  2022.  Multibody analysis and soft tissue strength refute supersonic dinosaur tail, 2022. 12(1):19245. Abstractconti_et_al_2022._multibody_analysis_and_soft_tissue_strength_refute_supersonic_dinosaur_tail.pdfWebsite

Sauropod dinosaurs are well known for their massive sizes and long necks and tails. Among sauropods, flagellicaudatan dinosaurs are characterized by extreme tail elongation, which has led to hypotheses regarding tail function, often compared to a whip. Here, we analyse the dynamics of motion of a 3D model of an apatosaurine flagellicaudatan tail using multibody simulation and quantify the stress-bearing capabilities of the associated soft tissues. Such an elongated and slender structure would allow achieving tip velocities in the order of 30 m/s, or 100 km/h, far slower than the speed of sound, due to the combined effect of friction of the musculature and articulations, as well as aerodynamic drag. The material properties of the skin, tendons, and ligaments also support such evidence, proving that in life, the tail would not have withstood the stresses imposed by travelling at the speed of sound, irrespective of the conjectural ‘popper’, a hypothetical soft tissue structure analogue to the terminal portion of a bullwhip able to surpass the speed of sound.

Pereira, AM, Silva MM, Mateus O.  2022.  First record of Phymactis papillosa (Lesson, 1830), a Pacific south sea anemone in European shores. Journal of the Marine Biological Association of the United Kingdom. :1–4.: Cambridge University Press Abstractfirst-record-of-phymactis-papillosa-lesson-1830-a-pacific-south-sea-anemone-in-european-shores.pdfWebsite


Guillaume, ARD, 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.

Moreno-Azanza, M, Pérez-Pueyo M, Puértolas-Pascual E, Núñez-Lahuerta C, Mateus O, Bauluz B, Bádenas B, Canudo JI.  2022.  A new crocodylomorph related ootaxon from the late Maastrichtian of the Southern Pyrenees (Huesca, Spain). Historical Biology. :1-10.: Taylor & Francis Abstracta_new_crocodylomorph_related_ootaxon_from_the_late_maastrichtian_of_the_southern_pyrenees_huesca_spain.pdfWebsite

ABSTRACTCrocodylomorph eggs and eggshells are known as old as the Late Jurassic and are frequent components of most multiootaxic eggshell assemblages. Classified within the oofamily Krokolithidae, thei histo- and ultrastructures are conservative throughout geological time, characterised by inverted-trapezoid-shaped shell units that grow from highly spaced basal knobs and present a diagnostic tabular ultrastructure. Here, we report 327 eggshell fragments from a new fossil site from the Maastrichtian of the Southern Pyrenees, Veracruz 1, and erect a new oogenus and oospecies, Pachykrokolithus excavatum oogen. et oosp. nov. characterised by crocodyloid morphotype and a prominent rugosocavate ornamentation. Eggshells from the slightly older locality of Blasi 2b, previously reported as aff. Krokolithidae, are also assigned to this new ootaxon. Different crocodylomorph taxa coexisted during the Late Cretaceous of the Tremp Basin, hindering the attribution of Pachykrokolithus excavatum oogen. et oosp. nov. to a single clade. Nevertheless, allodaposuchid eusuchians were dominant in this ecosystem, and are the most probable producers of Pachykrokolithus excavatum oogen. et oosp. nov. eggs.

Hendrickx, C, Bell PR, Pittman M, Milner ARC, Cuesta E, O'Connor J, Loewen M, Currie PJ, Mateus O, Kaye TG, 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.

Guillaume, ARD, Natário C, Mateus O, Moreno-Azanza M.  2022.  Plasticity in the morphology of the fused frontals of Albanerpetontidae (Lissamphibia; Allocaudata), 2022. Historical Biology. :1-18.: 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.

Fernandes, AE, Mateus O, Andres B, Polcyn MJ, Schulp AS, Gonçalves AO, Jacobs LL.  2022.  Pterosaurs from the Late Cretaceous of Angola. Diversity. 14, Number 9 Abstractdiversity-14-00741.pdfWebsite

Here, we describe the first pterosaur remains from Angola, an assemblage of fourteen bones from the Lower Maastrichtian marine deposits of Bentiaba, Namibe Province. One new species is introduced, Epapatelo otyikokolo, gen. et sp. nov., which comprises an articulated partial left humerus and ulna as well as an articulated left ulna and radius (from a second individual). Phylogenetic analysis confirms a non-nyctosaurid pteranodontian attribution for this new taxon and supports a new apomorphy-based clade, Aponyctosauria, which is here defined. Late Cretaceous pteranodontians are rare in Sub-Saharan Africa and throughout the Southern Hemisphere. Preliminary histological analysis also reveals a likely sub-adult age for one of the specimens. This fossil assemblage provides a first glimpse of Angolan pterosaur paleobiodiversity providing further insight into the Gondwanan ecosystems of the Upper Cretaceous.

Rotatori, FM, Moreno-Azanza M, Mateus O.  2022.  Reappraisal and new material of the holotype of Draconyx loureiroi (Ornithischia: Iguanodontia) provide insights on the tempo and modo of evolution of thumb-spiked dinosaurs, 02. Zoological Journal of the Linnean Society. Abstractrotatori_etal2022.pdfWebsite

{The Upper Jurassic Lourinhã Formation is well known for its rich assemblage of fossil vertebrates. In this formation, ornithopod dinosaurs are represented by two iguanodontian species, Eousdryosaurus nanohallucis and Draconyx loureiroi. We recently became aware of unreported material belonging to the holotype of Draconyx loureiroi, consisting of partially articulated manual elements. We here re-describe the holotype specimen ML 357, including the newly discovered material. The specimen was subjected to CT-scanning and its surface data used to assess anatomical characters. Linear measurements of metatarsal III were used to estimate the body length of the specimen. The Draconyx loureiroi holotype was included in two datasets and analysed with maximum parsimony and Bayesian inference approaches to estimate evolutionary rates among Iguanodontia. We present evidence that Draconyx loureiroi is a valid taxon nested in Styracosterna and is clearly diagnosable by a unique combination of characters. Both maximum parsimony and Bayesian inference indicate high evolutionary rates across the Jurassic/Cretaceous transition for the base of Iguanodontia. Length estimation suggests that Draconyx loureiroi was a relatively small, bipedal and possibly cursorial animal. Given its basal phyletic position, we interpret this bauplan was the ancestral condition for Styracosterna, that only later in the Cretaceous evolved into giant quadrupedal forms.}

Martino, R, Ríos MI, Mateus O, Pandolfi L.  2022.  Taxonomy, chronology, and dispersal patterns of Western European Quaternary hippopotamuses: New insight from Portuguese fossil material. Quaternary International. Abstractmartino_et_al_2022_hippos_portugal.pdfWebsite

The taxonomy, chronology and dispersal patterns of hippopotamuses in Western Europe are still a matter of debate, despite the long history of researchers on these topics. In this framework, the Portuguese material has never been investigated or recently revised, and it can contribute to increasing our comprehension about Quaternary hippopotamuses. The material considered in this work, collected from sites dated between 1 Ma and 0.13 Ma, is compared with fossil Pleistocene hippopotamuses from the Iberian Peninsula, Italy, Germany and Greece, in order to contribute into the discussion on evolutionary and dispersal patterns of these large semi-aquatic mammals in Europe during the Pleistocene. Portugal deposits recorded the presence of H. antiquus, starting from 1 Ma up to 0.4 Ma, and H. cf. amphibius during the late Middle Pleistocene. In particular, the occurrence of H. antiquus in Condeixa, dated around 0.4 Ma, can be considered as the LAD of this species in Europe. The paleobiogeography and dispersal patterns of European hippopotamuses are further discussed.

Moreno-Azanza, M, Balauz B, Ezquerro L, Coimbra R, Mateus O, Nuñez-Lahuerta C.  2022.  Unbroken dinosaur eggs accumulated together do not guarantee autochthony of the fossil assemblage. Journal of Taphonomy. 16(1-4):129-130. Abstractmoreno-azanza_et_al_2020_unbroken_eggs.pdf


Mateus, O, Estraviz-López D.  2022.  A new theropod dinosaur from the early cretaceous (Barremian) of Cabo Espichel, Portugal: Implications for spinosaurid evolution, 02. PLOS ONE. 17:1-48., Number 2: Public Library of Science Abstractjournal.pone_.0262614.pdfWebsite

Spinosaurids are some of the most enigmatic Mesozoic theropod dinosaurs due to their unique adaptations to aquatic environments and their relative scarcity. Their taxonomy has proven to be especially problematic. Recent discoveries from Western Europe in general, specifically Iberia, provide some of the best specimens for the understanding of their phylogeny, leading to the description of the spinosaurid Vallibonavenatrix cani and the recognition of the Iberian dinosaur Camarillasaurus cirugedae as one of them. Portuguese associated spinosaurid remains (ML1190) from the Papo Seco Formation (early Barremian) were previously assigned to Baryonyx walkeri but new material recovered in 2020 along with new phylogenetic analyses suggests a different phylogenetic placement, making their revision necessary. Here we show that these remains are not attributable to Baryonyx walkeri, but to a new genus and species, Iberospinus natarioi, gen. et sp. nov. The new taxon is characterized by the presence of a single Meckelian foramen in the Meckelian sulcus, a straight profile of the ventral surface of the dentary and a distal thickening of the acromion process of the pubis between other characters. Iberospinus natarioi is recovered as a sister taxon of the clade formed by Baryonyx and Suchomimus, and outside Spinosaurinae when Vallibonaventrix cani is excluded from the analysis. The description of this taxon reinforces Iberia as a hotspot for spinosaur biodiversity, with several endemic taxa for the region. As expected for the clade, the dentary displays a highly vascularized neurovascular network. The morphometric analysis of parts of the skeleton (pedal phalanx and caudal vertebrae, among others) shows an intermediate condition between basal tetanurans and spinosaurines.