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2021
Milàn, J., Mateus O., Mau M., Rudra A., Sanei H., & Clemmensen L. B. (2021).  A possible phytosaurian (Archosauria, Pseudosuchia) coprolite from the Late Triassic Fleming Fjord Group of Jameson Land, central East Greenland. Bulletin of the Geological Society of Denmark. 69, 71-80. Abstractmilan_et_al_2021_coprolites_greenland_bull69-71-80.pdfWebsite

A large, well-preserved vertebrate coprolite was found in a lacustrine sediment in the Malmros Klint Formation of the Late Triassic Fleming Fjord Group in the Jameson Land Basin, central East Greenland. The size and internal and external morphology of the coprolite is consistent with that of crocodilian coprolites and one end of the coprolite exhibits evidence of post-egestion trampling. As the associated vertebrate fauna of the Fleming Fjord Group contains abundant remains of pseudosuchian phytosaurs, the coprolite is interpreted as being from a large phytosaur.

Conti, S., Mateus O., & Sala G. (2021).  Mechanical characterization of tibial bone material of an ostrich. Rossi V., Fanti F., Barbieri G., Cavalazzi B. & Scarponi D. (Editors) 2021. Paleodays 2021. Abstract Book del XXI Convegno della Società Paleontologica Italiana, live virtual edition: 127 pp.. , 15-17 June, Bologna (Italy): University of Bolognaconti_et_al_2021_ostrich_bone.pdf
Fernandes, A. E., Mateus O., Bauluz B., Coimbra R., Ezquerro L., Núñez-Lahuerta C., Suteu C., & Moreno-Azanza M. (2021).  The Paimogo Dinosaur Egg Clutch Revisited: Using One of Portugal’s Most Notable Fossils to Exhibit the Scientific Method. Geoheritage. 13(3), 66., 2021 Abstractfernandes_et_al-2021-geoheritage.pdfWebsite

Found in the Upper Jurassic outcrops of Lourinhã, Portugal, and first published in 1997, the Paimogo dinosaur egg clutch is one of Portugal’s most remarkable fossils, with over one hundred eggs preserved in association with embryonic bones, of the allosauroid theropod Lourinhanosaurus. However, many questions about it have remained unanswered, even until the present day. After its discovery, this extraordinary fossil became the keystone of a small local museum, greatly kick-starting regional tourism, while also holding the fossils in trust for future generations to study. More than 20 years later, continually sustained paleontological interest from the public has even given rise to both a highly successful dinosaur theme park in the region and an aspiring UNESCO Geopark. Recently, a multidisciplinary team of preparators, paleontologists, sedimentologists, mineralogists, and geochemists revisited an unopened jacket from the original excavation using an array of techniques to address various questions. Studies are ongoing, but the corpus of information obtained and the methodologies utilized to gather data have offered an opportunity to design an exhibit around the history of the Paimogo clutch, highlighting the scientific methods involved, and asserting the importance of preserving geological heritage for the future, when new tools will doubtlessly become available to provide yet another new look at old fossils. Here, we describe our analytical procedures and present an innovative exhibit designed to introduce to the public the latest advances on the research behind an iconic piece of Portuguese geoheritage, increasing its value both as a research item and as an educational resource.

Beccari, V., Pinheiro F. L., Nunes I., Anelli L. E., Mateus O., & Costa F. R. (2021).  Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade. PLOS ONE. 16(8), e0254789 - ., 2021/08/25: Public Library of Science Abstractbeccari_et_al_2021.pdfWebsite

A remarkably well-preserved, almost complete and articulated new specimen (GP/2E 9266) of Tupandactylus navigans is here described for the Early Cretaceous Crato Formation of Brazil. The new specimen comprises an almost complete skeleton, preserving both the skull and post-cranium, associated with remarkable preservation of soft tissues, which makes it the most complete tapejarid known thus far. CT-Scanning was performed to allow the assessment of bones still covered by sediment. The specimen can be assigned to Tupa. navigans due to its vertical supra-premaxillary bony process and short and rounded parietal crest. It also bears the largest dentary crest among tapejarine pterosaurs and a notarium, which is absent in other representatives of the clade. The new specimen is here regarded as an adult individual. This is the first time that postcranial remains of Tupa. navigans are described, being also an unprecedented record of an articulated tapejarid skeleton from the Araripe Basin.

Conti, S., Tschopp E., Sala G., & Mateus O. (2021).  Multibody simulations of diplodocid tail motion. Annual conference of the European Association of Vertebrate Palaeontologists. , 5th-9th July : European Association of Vertebrate Palaeontologistsconti_et_al_2021_diplodocid_tail._eavp_abstract.pdf
2022
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
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Rotatori, F. M., 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

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Saleiro, A., Guillaume A. R. D., Rotatori F. M., Ríos-Ibañez M., López E. D., 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

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Moreno-Azanza, M., Pérez-Pueyo M., Puértolas-Pascual E., Núñez-Lahuerta C., Mateus O., Bauluz B., Bádenas B., & Canudo J. I. (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

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Jésus, V. J. P., Mateus O., Milàn J., & Clemmensen L. B. (2022).  First occurrence of a frog-like batrachian (Amphibia) in the Late Triassic Fleming Fjord Group, central East Greenland. Bulletin of the Geological Society of Denmark. 70, 117–130. Abstractbull70-117-130.pdfWebsite

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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.

Moreno-Azanza, M., Pérez-Pueyo M., Puértolas-Pascual E., Núñez-Lahuerta C., Mateus O., Bauluz B., Bádenas B., & Canudo J. I. (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.

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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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.

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

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

2023
Rotatori, F. M., Chiarenza A. A., Moreno-Azanza M., & Mateus O. (2023).  Rise of a dinasty: macroevolutionary and biogeographic patterns of iguanodontian dinosaurs across the Jurassic–Cretaceous transition. EAVP Annual Conference of the European Association of Vertebrate Palaeontologists. 226.: Palaeovertebrata, Special Volume 1-23. DOI: 10.18563/pv.eavp2023 Abstract
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Coimbra, R., Moreno-Azanza M., Ezquerro L., Nuñez-Lahuerta C., Gasca J. M., Immenhauser A., Mateus O., & Rocha F. (2023).  Evaluating and comparing geochemical sampling protocols in dinosaur eggshells: refining Cretaceous ecosystem research. Cretaceous Research. 105632. Abstractsingle_file_coimbra_et_al._2023_cretresearch.pdfWebsite

The geochemical signatures of dinosaur eggshells represent well-established proxies in paleoenvironmental and paleobiological research. The variable sampling procedures reported in the literature, however, deserve attention. In order to evaluate the impact of different sampling methodologies on carbon and oxygen isotope and elemental concentrations, grinding was contrasted with drilling to extract powder samples from eggshell fragments collected at several locations. Eggshell data were further contrasted with surface materials, encasing matrix and compared with independent proxies using petrographic and elemental techniques. Iron and manganese elemental concentrations revealed an enrichment sequence depending on the sampling strategy for the same eggshell fragment. This pattern can be mistaken for a variable state of preservation. In contrast, carbon and oxygen isotope values exhibited only subtle differences and lacked clear trends. This suggests that isotope data are less susceptible to different methodological approaches. It is shown that drilling offers a wider range of possibilities compared to grinding (e.g., faster and less destructive). Additionally, drilled powder samples can confidently be used for elemental and isotope analysis, excluding contamination, thus providing a more accurate set of proxy data from eggshell archives.

Werneburg, I., Pommery Y., Ruciński M., Kästle B., Cohen G. J., Natchev N., Mateus O., & Ferreira G. D. (2023).  Functional morphology of the skull of Henodus chelyops (Placodontia). International Congress of Vertebrate Morphology Cairns - QLD - Australia 28 July - 1 August 2023. The Anatomical Record. 232-233. Abstractwerneburg_et_al_2023_henodus_icvm_2023_abstracts_updated_8_14-1693344432.pdf

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Rotatori, F. M., Quaranta M., Bertozzo F., Hübner T., Camilo B., Mateus O., & Moreno-Azanza M. (2023).  Hadrosaur-like vascularisation in the dentary of an early diverging iguanodontian dinosaur. Historical Biology. 1-6.: Taylor & Francis AbstractWebsite

ABSTRACTVirtual palaeontology is a growing field, leading palaeontologists to better understand the microanatomy of many extinct species. The application of techniques such as CT and μCT-scanning allows the researchers to study micro-anatomical features in a non-invasive way and make inferences on the palaeobiology of animals. Dinosaurs have been extensively studied using these techniques, with particular focus on the microanatomy of the cranium, whereas relatively little is known of other cranial elements, such as the lower jaw. Here, we aim to fill this gap, describing the microanatomy of the specimen ML 768, an isolated dentary belonging to a dryosaurid iguanodontian dinosaur from the Upper Jurassic of Lourinhã Fm. The dentary ML 768 was subjected to μCT-scanning, and subsequently the data were segmented in Avizo and rendered in Blender. We identified functional and replacement teeth, recognising remnants of old replacement cycles. Furthermore, we mapped a rich neurovascular network present in the dentary and compared it with reference literature. We found that the high vascularisation is shared with other cerapodan dinosaurs with high tooth replacement rates, although homoeostasis may have also played a role in the development of this condition. Further evidence is needed to appreciate the macroevolutionary significance of these findings.

Conti, S., Masarati P., Tschopp E., Zanoni A., Mateus O., & Sala G. (2023).  How to simulate soft tissues in extinct animals. Using sauropod dinosaurs as a case study. ECCOMAS Thematic Conference on Multibody Dynamics. Abstractconti_et_al_2023_id_218_424_eccomas_mbd_2023_congresso_lisbona.pdf

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Crespo, V. D., Sequero López C., Rios M., Estraviz López D., Gamonal A., Martino R., Riccetto M., Callapez P., Legoinha P., & Mateus O. (2023).  Revisiting the classical small mammal assemblage from the Eocene site of Silveirinha: a fresh look at old collections. XI Congresso Nacional de Geologia: Geociências e Desafios globaisAt: Universidade de CoimbraVolume: Livro de Resumos, págs 85-86. , Coimbra Abstractcrespov_etal_xicng2023_silveirinha.pdf

O sítio de Silveirinha é uma das localidades de mamíferos mais conhecidas da Paleontologia do Cenozoico de Portugal e da Europa em geral. Graças à sua rica e diversificada associação de mamíferos, com mais de 30 taxa, foi posicionado no Eocénico inferior (início do Ypresiano, MP7, ca. 55,8 M.a.), sendo o local mais antigo da Europa desta Época, devido à presença única de taxa típicos do Paleocénico superior, juntamente com outras espécies já características do Eocénico inferior. Este estudo irá rever o material de pequenos mamíferos deste sítio, conservado na coleção clássica da Universidade Nova de Lisboa, a fim de fazer uma actualização taxonómica à luz das publicações mais recentes.

Conti, S., Sala G., & Mateus O. (2023).  Smart Biomechanical Adaptation Revealed by the Structure of Ostrich Limb Bones. Biomimetics. 8(1), , Number 1 Abstractbiomimetics-08-00098.pdfWebsite

Ostriches are known to be the fastest bipedal animal alive; to accomplish such an achievement, their anatomy evolved to sustain the stresses imposed by running at such velocities. Ostriches represent an excellent case study due to the fact that their locomotor kinematics have been extensively studied for their running capabilities. The shape and structure of ostrich bones are also known to be optimized to sustain the stresses imposed by the body mass and accelerations to which the bones are subjected during movements. This study focuses on the limb bones, investigating the structure of the bones as well as the material properties, and how both the structure and material evolved to maximise the performance while minimising the stresses applied to the bones themselves. The femoral shaft is hollowed and it presents an imbricate structure of fused bone ridges connected to the walls of the marrow cavity, while the tibial shaft is subdivided into regions having different mechanical characteristics. These adaptations indicate the optimization of both the structure and the material to bear the stresses. The regionalization of the material highlighted by the mechanical tests represents the capability of the bone to adapt to external stimuli during the life of an individual, optimizing not only the structure of the bone but the material itself.

López-Rojas, V., Clemmensen L. B., Milàn J., Wings O., Klein N., & Mateus O. (2023).  A new phytosaur species (Archosauriformes) from the Upper Triassic of Jameson Land, central East Greenland. Journal of Vertebrate PaleontologyJournal of Vertebrate Paleontology. e2181086., 2023: Taylor & Francis Abstracta_new_phytosaur_species_archosauriformes_from_the_upper_triassic_of_jameson_land_central_east_greenland.pdfWebsite

ABSTRACTHerein we describe phytosaurs from thin fluvial overbank sandstones of the Upper Triassic Malmros Klint Formation of the Fleming Fjord Group (central East Greenland). The new sample includes more than 150 disarticulated bones and teeth from small to large specimens belonging to at least four individuals. The fossils mostly consist of teeth and postcranial elements and permit the recognition of a new species of Mystriosuchus, M. alleroq, diagnosed by an L-shaped quadratojugal whose anterior suture trends anterodorsally and a tripartite degree of heterodonty. Humeral diaphyseal histology of one specimen reveals a fairly compact cortex that surrounds a cancellous medullary region followed by a remodeling zone containing scattered secondary osteons. Primary bone tissue is parallel-fibred with a moderate to low vascular density. The cortex is cyclically interrupted by distinct growth marks indicating a seasonal environment. A change in growth rate from moderate to low is documented within the outer cortex, indicating that at least this individual was close to somatic maturity. Mystriosuchus has formerly been known as an exclusively European taxon. The new findings support the European faunal influence in East Greenland during the Late Triassic inferred from other taxa such as temnospondyls and archosaurs. The mid-late Norian age of European Mystriosuchus suggests an additional age constraint for the vertebrate-bearing portion of the Malmros Klint Formation.

2024
Rotatori, F. M., Ferrari L., Sequero C., Camilo B., Mateus O., & Moreno-Azanza M. (2024).  An unexpected early-diverging iguanodontian dinosaur (Ornithischia, Ornithopoda) from the Upper Jurassic of Portugal. Journal of Vertebrate Paleontology. e2310066.: Taylor & Francis AbstractWebsite

Iguanodontia is a diverse clade of herbivorous ornithischian dinosaurs that were speciose and abundant during the Jurassic and Cretaceous. Although the monophyly of Iguanodontia is well supported, their internal relationships have sparked heated debate due to several phylogenetic paradigm shifts. Late Jurassic basally branching iguanodontians in particular are not well understood in terms of their systematic affinities and evolutionary relevance. Their fossil record in Europe is meager compared with North America, with only a few species currently recognized. Two taxa are currently known from the Upper Jurassic of England, the basally branching styracosternan Cumnoria prestwichii and the putative dryosaurid Callovosaurus leedsi. In the Upper Jurassic of Portugal, the styracosternan Draconyx loureiroi and the dryosaurid Eousdryosaurus nanohallucis are presently the only described basally branching iguanodontians. Here we report a new species of early diverging iguanodontian from the Upper Jurassic Lourinhã Formation of western-central Portugal. The new species is clearly distinguished from all other coeval taxa by an exclusive combination of characters that include a tibia with a cnemial crest that is directed craniolaterally and a fibular condyle that is angled at 90° with respect to the proximal epiphysis, a fibula with symmetrical proximal margins, and a reduced metatarsal I. The phylogenetic relationships of the Lourinhã iguanodontian were explored using maximum parsimony and Bayesian inference. The two analyses recover the Lourinhã iguanodontian as an indeterminate dryomorphan, with more precise affinities precluded due to the current available material. Body size is estimated between 3 and 4 meters for the holotype specimen, adding to the diversity of small ornithopods already recognized in the paleoichnological record of the Lourinhã Formation. http://zoobank.org/urn:lsid:zoobank.org:pub:5F4D52D0-0F0B-4809-8561-BE58C7C97D45