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2013
da} Rocha, {R. E. B., Kullberg {J. C. R. }, & Mateus O. (2013).  A Bacia Lusitaniana: Estratigrafia, Paleogeografia e Tectónica. (Dias, Araújo, R., A, Terrinha, P., Kullberg, {J. C.}, Ed.).Geologia de Portugal no contexto da Ibéria. 195–347., 1: Escolar Editora Abstract
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Mateus, O. (2013).  Cabinda revisited: age and environment of new Cenozoic vertebrate fossils from northern Angola. 45, , 1 Abstract

In the early 20thcentury, Belgian naturalists reported Paleocene and Eocene sharks, the bothremydid pleurodiran turtleTaphosphrys(formerlyBantuchelys), and a neosuchian and the dyrosaurid crocodyliformCongosaurusfrom coastal outcrops near Landana in the northern province of Cabinda, Angola. In 1935, rare and fragmentary mammals were reported from strata at Malembo Point, south of Landana, and originally considered to be Miocene in age. Subsequent re-identification of the mammal fossils suggested that the Malembo deposits are Lower Oligocene based on the presence of an arsinoithere and hyracoids. An anthropoid canine, originally identified as a carnivore incisor, was also recognized. In four recent expeditions to Cabinda, Projecto PaleoAngola has traced and measured the stratigraphy between Landana and Malembo Point and sampled these deposits for pollen, stable isotopes, detrital zircons, paleomagnetic stratigraphy, and for U/Pb dating of bones, teeth, and coprolites. Although the faunas from all Cabinda localities are dominated by sharks and rays, new discoveries from Landana include a complete cheloniid cryptodire turtle skull, a small snake vertebra, and a bird bone. Discoveries from the Malembo level include a narrow-snouted crocodyliform similar toCongosaurusandEuthecodon, an arsinoithere anterior tooth, an upper molar similar to that of the ptolemaiidanKelba, an unidentified mammalian incisor, and a large primate-like premolar. Recent biostratigraphic advances in East Africa and the new fossil discoveries in Cabinda suggest similarity to late Oligocene faunas in Ethiopia and Kenya, although the large primate-like premolar is unique. To this day, the fossil localities of Malembo provide the only coastal, low latitude, low elevation record of West African Cenozoic terrestrial mammals. https://gsa.confex.com/gsa/2013AM/webprogram/Paper232115.html

Mateus, O. (2013).  Cathetosaurus as a valid sauropod genus and comparisons with Camarasaurus. Journal of Vertebrate Paleontology. 173., 1 Abstract
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Mateus, O. (2013).  Decapod crustacean body and ichnofossils from the Mesozoic of Portugal. NA, , 1 Abstract

Book of abstracts of the 5th Symposium on Mesozoic and Decapod Crustaceans

Mateus, O. (2013).  First definitive association between embryonic Allosaurus bones and prismatoolithus eggs in the Morrison Formation (Upper Jurassic, Wyoming, USA). Journal of Vertebrate Paleontology. 101., 1 Abstract

Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

Coelho,(ed)C., & et al (2013).  Arrábida - al-rábita. , Lisboa, 229 pp.: Associação de Município da Região de Setúbal
Kullberg, J. C., Rocha R. B., Soares A. F., Rey J., Terrinha P., Azerêdo A. C., Callapez P., Duarte, L.V., Kullberg M. C., Martins L., Miranda J. R., Alves C., Mata J., Madeira J., Mateus O., Moreira M., & Nogueira C. R. (2013).  A Bacia Lusitaniana: Estratigrafia, Paleogeografia e Tectónica. (Dias, R. Araújo, A, Terrinha, P. and Kullberg, J. C., Ed.).Geologia de Portugal no contexto da Ibéria. Volume II. 195-350., Lisboa: Escolar Editorakullberg_et_al_2013_a_bacia_lusitaniana.pdf
Bianucci, G., Miján I., Lambert O., Post K., & Mateus O. (2013).  Bizarre fossil beaked whales (Odontoceti, Ziphiidae) fished from the Atlantic Ocean floor off the Iberian Peninsula. Geodiversitas. 35(1), 105-153. Abstractbianucci_et_al_2013_fossil_beaked_whales_iberian_peninsula.pdf

Forty partial fossil skulls belonging to beaked whales (Cetacea, Odontoceti, Ziphiidae) were collected by trawling and long-line fishing on Neogene (probably Late Early to Middle Miocene) layers of the Atlantic floor off the coasts of Portugal and Spain (Asturias and Galicia). e systematic study of the most diagnostic Iberian specimens, those preserving the rostrum and the dorsal part of the cranium, led to the recognition of two new genera (Globicetus n. gen. and Imocetus n. gen.) and four new species (Choneziphius leidyi n. sp., G. hiberus n. gen., n. sp., I. piscatus n. gen., n. sp., and Tusciziphius atlanticus n. sp.).
Based on the matrix of a previous work, the phylogenetic analysis places all the new taxa in the subfamily Ziphiinae Gray, 1850. More fragmentary specimens are tentatively referred to the genera Caviziphius Bianucci & Post, 2005 and Ziphirostrum du Bus, 1868. Among these new ziphiids, extremely bizarre skull morphologies are observed. In G. hiberus n. gen., n. sp. the proximal portion of the rostrum bears a voluminous premaxillary spheroid. In T. atlanticus n. sp. a medial premaxillary bulge is present on the rostrum; together with asymmetric
rostral maxillary eminences at the rostrum base, this bulge displays various degrees of elevation in different specimens, which may be interpreted as sexual dimorphism. Specimens of I. piscatus n. gen., n. sp. bear two sets of even crests: spur-like rostral maxillary crests and longitudinal maxillary crests laterally bordering a wide and long facial basin. A preliminary macroscopic observation of these elements indicates very dense bones, with a compactness comparable with that of cetacean ear bones. Questioning their function, the high medial rostral elements (the premaxillary spheroid of G. hiberus n. gen., n. sp. and the medial bulge of T. atlanticus n. sp.) remind the huge rostral maxillary crests of adult males of the extant Hyperoodon ampullatus (Forster, 1770). In the latter, the crests are very likely related to head-butting. However, they are made of much more spongy bone than in the fossil taxa studied here, and therefore possibly better mechanically suited for facing impacts. Other interpretations of these unusual bone specializations, related to deep-diving (ballast) and echolocation (sound reflection), fail to explain the diversity of shapes and the hypothetical sexual dimorphism observed in at least part of the taxa. e spur-like rostral maxillary crests and long maxillary crests limiting the large facial basin in I. piscatus n. gen., n. sp. and the excrescences on the maxilla at the rostrum base in Choneziphius spp. are instead interpreted as areas of origin for rostral and facial muscles, acting on the nasal passages, blowhole, and melon. From a palaeobiogeographic point of view, the newly described taxa further emphasize the differences in the North Atlantic (including Iberian Peninsula) and South African Neogene ziphiid faunal lists. Even if the stratigraphic context is poorly understood, leaving open the question of the geological age for most of the dredged specimens, these differences in the composition of cold to temperate northern and southern hemisphere fossil ziphiid faunas may be explained by a warm-water equatorial barrier.

Jacobs, L. L., Myers T. S., Goncalves A. O., Graf J. F., Jacobs B. F., KAPPELMAN J. W., Mateus O., Polcyn M. J., RASBURY E. T., & Vineyard D. P. (2013).  Cabinda revisited: age and environment of new Cenozoic vertebrate fossils from northern Angola. Geological Society of America Abstracts with Programs. Vol. 45, No. 7, p.0.
Mateus, O., & Tschopp E. (2013).  Cathetosaurus as a valid sauropod genus and comparisons with Camarasaurus. Journal of Vertebrate Paleontology, Program and Abstracts, 2013. 173.mateus__tschopp_2013_cathetosaurs_camarasaurus__svp_meeting_abstracts_213.pdf
Tschopp, E., & Mateus O. (2013).  Clavicles, interclavicles, gastralia, and sternal ribs in sauropod dinosaurs: new reports from Diplodocidae and their morphological, functional and evolutionary implications. Journal of Anatomy. 222, 321-340. Abstracttschopp__mateus_2013_clavicles_interclavicles_gastralia_and_sternal_ribs_in_diplodocid.pdfWebsite

Ossified gastralia, clavicles and sternal ribs are known in a variety of reptilians, including dinosaurs. In sauropods, however, the identity of these bones is controversial. The peculiar shapes of these bones complicate their identification, which led to various differing interpretations in the past. Here we describe different elements from the chest region of diplodocids, found near Shell, Wyoming, USA. Five morphotypes are easily distinguishable: (A) elongated, relatively stout, curved elements with a spatulate and a bifurcate end resemble much the previously reported sauropod clavicles, but might actually represent interclavicles; (B) short, L-shaped elements, mostly preserved as a symmetrical pair, probably are the real clavicles, as indicated by new findings in diplodocids; (C) slender, rod-like bones with rugose ends are highly similar to elements identified as sauropod sternal ribs; (D) curved bones with wide, probably medial ends constitute the fourth morphotype, herein interpreted as gastralia; and (E) irregularly shaped elements, often with extended rugosities, are included into the fifth morphotype, tentatively identified as sternal ribs and/or intercostal elements. To our knowledge, the bones previously interpreted as sauropod clavicles were always found as single bones, which sheds doubt on the validity of their identification. Various lines of evidence presented herein suggest they might actually be interclavicles – which are single elements. This would be the first definitive evidence of interclavicles in dinosauromorphs. Previously supposed interclavicles in the early sauropodomorph Massospondylus or the theropods Oviraptor and Velociraptor were later reinterpreted as clavicles or furculae. Independent from their identification, the existence of the reported bones has both phylogenetic and functional significance. Their presence in non-neosauropod Eusauropoda and Flagellicaudata and probable absence in rebbachisaurs and Titanosauriformes shows a clear character polarity. This implicates that the ossification of these bones can be considered plesiomorphic for Sauropoda. The proposed presence of interclavicles in sauropods may give further support to a recent study, which finds a homology of the avian furcula with the interclavicle to be equally parsimonious to the traditional theory that furcula were formed by the fusion of the clavicles. Functional implications are the stabilizing of the chest region, which coincides with the development of elongated cervical and caudal vertebral columns or the use of the tail as defensive weapon. The loss of ossified chest bones coincides with more widely spaced limbs, and the evolution of a wide-gauge locomotor style.

Mateus, O. (2013).  Crocodylomorphs from the Mesozoic of Portugal and a new skull of eusuchian from the Late Cretaceous. 2013 Hwaseong International Dinosaurs Expedition Symposium, pp.66-67.. , Hwaseong, South Korea Abstractmateus_2013_crocodylomorphs_portugal_new_skull.pdf

The diversity of fossil crocodylomorphs in Portugal is high, with occurrence as old as Mystriosaurus (=Steneosaurus) bollensis from the Lower Jurassic. The Late Jurassic forms are the better documented, and include the following taxa: Machimosaurus hugii, Lisboasaurus estesi Seiffert, 1973, Lusitanisuchus mitrocostatus Seiffert, 1975; Schwarz & Fechner 2004, Theriosuchus guimarotae Schwarz and Salisbury 2005, Cf. Alligatorium, Goniopholis baryglyphaeus, and a crocodylomorph-like eggs in dinosaur nests (Mateus et al., 1998; Ricqlès et al., 2001). From the Lower Cretaceous were reported a few dinosaurs but its record is strangely scarce in crocodylomorphs (Mateus et al., 2011). The Upper Cretaceous crocodiles show a large diversity, but it is mostly based in fragmentary material that require revision, such as “Crocodylus” blavieri? Grey from the Upper Campanian - Maastrichtian of Viso, near Aveiro (initially reported by Sauvage 1897-98), Goniopholis cf. crassidens Owen 1841 and Oweniasuchus pulchelus Jonet 1981. Moreover there is a fascinating, but poorly understood, crocodylomorph diversity in the Cenomanian of Portugal, documented by fragmentary specimens that have been doubtfully assigned to Thoracosaurus Leidy 1852 of the Middle Cenomanian of Cacém, to the nomen dubium Oweniasuchus lusitanicus Sauvage 1897-98 (interpreted as a mesosuchian goniopholid) based in a fragmentary mandible from the Campanian-Maastrichtian, and also from the Middle Cenomanian of Portugal, Buffetaut and Lauverjat (1978) report an fragmentary unidentified possible dyrosaurid from Nazaré. All this specimens are too incomplete to be compared with the specimen here described. In contrast, Cenozoic crocodiles of Portugal are often known after complete skulls and several individuals. The taxa list include Iberosuchus macrodon (Lower to Middle Eocene), Tomistoma calaritanus (Early Miocene) and T. lusitanica (Burdigalian-Helvetian), and Diplocynodon sp. (Antunes, 1961, 1987, 1994).
At least, two different morphotypes of crocodylomorph eggs from the Late Jurassic of Lourinhã Formation are also known.
A new specimen here reported of crocodile based in a partial skull and mandible (ML1818) from the Uppermost Middle Cenomanian platform carbonates of Baixo Mondego, west central Portugal (Tentúgal Fm., Callapez, 2004). The taxon is phylogenetically positioned as a basal Eusuchia, due to the choanae enclosed by the pterygoid, and closely related with stem Crocodylia and Borealosuchus. This specimen represents the only well documented and valid eusuchian species in the Cenomanian of Europe and is the oldest representative of an eusuchian crocodylomorph, with the exception for the Barremian Hylaeochampsa vectiana.

Carrano, M., Mateus O., & Mitchell J. (2013).  First definitive association between embryonic Allosaurus bones and prismatoolithus eggs in the Morrison Formation (Upper Jurassic, Wyoming, USA). Annual Meeting of Vertebrate Paleontology. 101.: Journal of Vertebrate Paleontology, Program and Abstracts, 2013 Abstractcarrano_mateus_mitchell_2013_allosaurus_embryos_morrison_svp_abstract.pdf

Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we
report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery
also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

Strganac, C., Jacobs L. L., Ferguson K. M., Polcyn M. J., Mateus O., Schulp A. S., & Morais M. L. (2013).  Late Cretaceous marine reptiles and cooling at the South Atlantic coast inferred through stable oxygen isotopes of Inoceramus from the Namibe Basin, Angola. Geological Society of America Abstracts with Programs. Vol. 45, No. 7, p.0.
Polcyn, M. J., Jacobs L. L., Mateus O., Schulp A. S., Strganac C., Araújo R., Graf J. F., Vineyard D., & Myers T. S. (2013).  A marine vertebrate assemblage from the Campanian-Maastrichtian boundary at Bentiaba, Angola. Geological Society of America Abstracts with Programs. Vol. 45, No. 7, p.0. polcyn_et_al_2013_abstract__a_marine_vertebrate_assemblage_from_the_campanian-maastrichtian_boundary_at_bentiaba_angola_2013__gsa_27-30_october_2013.pdf
Mannion, P. D., Upchurch P., Barnes R. N., & Mateus O. (2013).  Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society. 168, 98-206. Abstractmannion_et_al_2013_sauropod_lusotitan_portugal.pdfWebsite

Titanosauriforms represent a diverse and globally distributed clade of neosauropod dinosaurs, but their inter-relationships remain poorly understood. Here we redescribe Lusotitan atalaiensis from the Late Jurassic Lourinhã Formation of Portugal, a taxon previously referred to Brachiosaurus. The lectotype includes cervical, dorsal, and caudal vertebrae, and elements from the forelimb, hindlimb, and pelvic girdle. Lusotitan is a valid taxon and can be diagnosed by six autapomorphies, including the presence of elongate postzygapophyses that project well beyond the posterior margin of the neural arch in anterior-to-middle caudal vertebrae. A new phylogenetic analysis, focused on elucidating the evolutionary relationships of basal titanosauriforms, is presented, comprising 63 taxa scored for 279 characters. Many of these characters are heavily revised or novel to our study, and a number of ingroup taxa have never previously been incorporated into a phylogenetic analysis. We treated quantitative characters as discrete and continuous data in two parallel analyses, and explored the effect of implied weighting. Although we recovered monophyletic brachiosaurid and somphospondylan sister clades within Titanosauriformes, their compositions were affected by alternative treatments of quantitative data and, especially, by the weighting of such data. This suggests that the treatment of quantitative data is important and the wrong decisions might lead to incorrect tree topologies. In particular, the diversity of Titanosauria was greatly increased by the use of implied weights. Our results support the generic separation of the contemporaneous taxa Brachiosaurus, Giraffatitan, and Lusotitan, with the latter recovered as either a brachiosaurid or the sister taxon to Titanosauriformes. Although Janenschia was recovered as a basal macronarian, outside Titanosauria, the sympatric Australodocus provides body fossil evidence for the pre-Cretaceous origin of titanosaurs. We recovered evidence for a sauropod with close affinities to the Chinese taxon Mamenchisaurus in the Late Jurassic Tendaguru beds of Africa, and present new information demonstrating the wider distribution of caudal pneumaticity within Titanosauria. The earliest known titanosauriform body fossils are from the late Oxfordian (Late Jurassic), although trackway evidence indicates a Middle Jurassic origin. Diversity increased throughout the Late Jurassic, and titanosauriforms did not undergo a severe extinction across the Jurassic/Cretaceous boundary, in contrast to diplodocids and non-neosauropods. Titanosauriform diversity increased in the Barremian and Aptian–Albian as a result of radiations of derived somphospondylans and lithostrotians, respectively, but there was a severe drop (up to 40%) in species numbers at, or near, the Albian/Cenomanian boundary, representing a faunal turnover whereby basal titanosauriforms were replaced by derived titanosaurs, although this transition occurred in a spatiotemporally staggered fashion.

Salminen, J., Dinis J., & Mateus O. (2013).  Preliminary magnetostratigraphy for Jurassic/Cretaceous transition in Porto da Calada, Portugal. In: Veikkolainen, T., Suhonen, K., Näränen, J., Kauristie, K., and Kaasalainen, S. (eds.). XXVI Geofysiikan päivät,. 119-122., May 21-22 2013 in Helsinkisalminen.johanna_gfp2013_portugal_preliminary_magnetostratigraphy_for_jurassic_cretaceous_transition_in.pdf
Tschopp, E., & Mateus O. (2013).  The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology. 11(7), 853–888. Abstracttschopp__mateus_2013_the_skull_and_neck_of_a_new_flagellicaudatan_sauropod_from_the_morrison.pdfWebsite

A new taxon of diplodocid sauropod, Kaatedocus siberi gen. et sp. nov., is recognized based on well-preserved cervical vertebrae and skull from the Morrison Formation (Kimmeridgian, Late Jurassic) of northern Wyoming, USA. A phylogenetic analysis places it inside Diplodocinae (Sauropoda: Flagellicaudata: Diplodocidae), as a sister taxon to a clade uniting Tornieria africana and the classical diplodocines Barosaurus lentus and Diplodocus. The taxon is diagnosed by a unique combination of plesiomorphic and derived traits, as well as the following unambiguous autapomorphies within Diplodocidae: frontals separated anteriorly by a U-shaped notch; squamosals restricted to the post-orbital region; presence of a postparietal foramen; a narrow, sharp and distinct sagittal nuchal crest; the paired basal tuber with a straight anterior edge in ventral view; anterior end of the prezygapophyses of mid- and posterior cervical vertebrae is often an anterior extension of the pre-epipophysis, which projects considerably anterior to the articular facet; anterodorsal corner of the lateral side of the posterior cervical vertebrae marked by a rugose tuberosity; posterior margin of the prezygapophyseal articular facet of posterior cervical vertebrae bordered posteriorly by conspicuous transverse sulcus; posterior cervical neural spines parallel to converging. The inclusion of K. siberi and several newly described characters into a previously published phylogenetic analysis recovers the new taxon as basal diplodocine, which concurs well with the low stratigraphical position of the holotype specimen. Dinheirosaurus and Supersaurus now represent the sister clade to Apatosaurus and Diplodocinae and therefore the most basal diplodocid genera. The geographical location in the less known northern parts of the Morrison Fm., where K. siberi was found, corroborates previous hypotheses on faunal provinces within the formation. The probable subadult ontogenetic stage of the holotype specimen allows analysis of ontogenetic changes and their influence on diplodocid phylogeny.

Brusatte, S. L., Butler R. J., Mateus O., Steyer J. S., & Whiteside J. H. (2013).  Terrestrial vertebrates from the Late Triassic of Portugal: new records of temnospondyls and archosauriforms from a Pangaean rift sequence. 61st Symposium on Vertebrate Palaeontology and Comparative Anatomy. 15-16., Edinburgh Abstractbrusatte_et_al_2013_triassic_algarve_portugal_temnospondyls_svpca.org_years_2013_edinburgh_abstracts.pdf

The Late Triassic (ca. 237-201.5 million years ago) was a transitional interval in the evolution of terrestrial ecosystems, during which ‘modern’ clades such as archosaurs and mammals were radiating while ‘archaic’ groups such as temnospondyl amphibians and basal synapsids remained abundant. Little is known about the Triassic terrestrial (nonmarine) vertebrates of the Iberian Peninsula. The Algarve Basin of southern Portugal is an extensional rift basin formed during the breakup of Pangaea, which is filled with terrestrial, lacustrine, and marginal marine siliciclastics of the Grés de Silves Formation, interbedded with CAMP basalts that mark the end-Triassic extinction (radioisotopically dated to ~198-201.5 Ma). Since 2009, our field project in the Algarve has discovered numerous vertebrate specimens within the Grés de Silves, including a monodominant bonebed containing hundreds of specimens of metoposaurids, a peculiar group of temnospondyls that filled crocodile-like predatory niches in lacustrine and fluvial environments. These specimens appear to belong to a new species of Metoposaurus, similar to M. diagnosticus and M. krasiejowensis from central Europe but possessing several putative autapomorphies of the braincase and lower jaw. We also discovered a mandible of a phytosaur, the first specimen of these long-snouted, semi-aquatic archosauriforms from the Iberian Peninsula. These discoveries of characteristic Carnian Norian taxa indicate that the fossil-bearing portion of the Grés de Silves is Late Triassic in age, and provide further evidence that metoposaurids and phytosaurs commonly occurred together in low palaeolatitudes during this time.

Ribeiro, V., Holwerda F., & Mateus O. (2013).  Theropod egg sites from the Lourinhã Formation, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2013. 198.ribeiro_et_al_2013_theropod_egg_sites_lourinha_svp_meeting_abstracts_213.pdf
Schulp, A. S., Polcyn M. J., Mateus O., & Jacobs L. L. (2013).  Two rare mosasaurs from the Maastrichtian of Angola and the Netherlands. Netherlands Journal of Geosciences — Geologie en Mijnbouw. 92(1), 3-10. Abstractschulp_et_al_2013_two_rare_mosasaurs_from_the_maastrichtian_of_angola__njg-92-1-schulp.pdf

We report here the addition of two rare mosasaur taxa to the Maastrichtian marine amniote fauna of Angola. The new specimens include a dentary fragment referable to the large carnivore Prognathodon saturator and an isolated tooth of the small durophage Carinodens belgicus. Both were recovered from Maastrichtian outcrops in southern Angola in 2011. Additionally, a complete posterior mandibular unit of a large mosasaur from the type Maastrichtian of The Netherlands, collected some time prior to 1879 and previously identified as ‘Mosasaurus giganteus’, is described and reassigned here to Prognathodon saturator; historical issues surrounding the taxonomic attribution of this specimen are clarified. The new material extends the known geographic distribution of Prognathodon saturator and Carinodens belgicus.

Mannion, P. D., Upchurch P., Barnes R. N., & Mateus O. (2013).  Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society. 168, 98-206., Number 1 Abstract
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Schulp, A. S., Polcyn M. J., Mateus O., & Jacobs L. L. (2013).  Two rare mosasaurs from the Maastrichtian of Angola and the Netherlands. Geologie en Mijnbouw/Netherlands Journal of Geosciences. 92, 3-10., Number 1 Abstract
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2012
Mannion, P. D., Upchurch P., Mateus O., Barnes R. N., & Jones M. E. H. (2012).  New information on the anatomy and systematic position of Dinheirosaurus lourinhanensis (Sauropoda: Diplodocoidea) from the Late Jurassic of Portugal, with a review of European diplodocoids. Journal of Systematic Palaeontology. 10(3), 521–551., Jan Abstractmannion_et_al_2012_new_information_on_the_anatomy_and_systematic_position_of_dinheirosaurus_lourinhanensis_sauropoda_-_diplodocoidea_from_the_late_jurassic_of_portugal_with_a_review_of_european_diplodocoids.pdf

Although diplodocoid sauropods from Africa and the Americas are well known, their European record remains largely neglected. Here we redescribe Dinheirosaurus lourinhanensis from the Late Jurassic of Portugal. The holotype comprises two posterior cervical vertebrae, the dorsal series and a caudal centrum. Redescription demonstrates its validity on the basis of three autapomorphies: (1) posteriorly restricted ventral keel on posterior cervical vertebrae; (2) three small subcircular fossae posterior to the lateral coel on posterior cervical neural spines; (3) accessory lamina linking the hyposphene with base of the posterior centrodiapophyseal lamina in middle-posterior dorsal vertebrae. Phylogenetic analysis places Dinheirosaurus as the sister taxon to Supersaurus, and this clade forms the sister taxon to other diplodocines. However, this position should be treated with caution as Dinheirosaurus displays several plesiomorphic features absent in other diplodocids (including unbifurcated presacral neural spines, and dorsolaterally projecting diapophyses on dorsal vertebrae) and only four additional steps are required to place Dinheirosaurus outside of Flagellicaudata. We identify Amazonsaurus as the basal-most rebbachisaurid and recover Zapalasaurus outside of the South American Limaysaurinae, suggesting the biogeographic history of rebbachisaurids is more complex than previously proposed. Review of the European diplodocoid record reveals evidence for the earliest known diplodocid, as well as additional diplodocid remains from the Late Jurassic of Spain. A Portuguese specimen, previously referred to Dinheirosaurus, displays strong similarities to Apatosaurus from the contemporaneous Morrison Formation of North America, indicating the presence of a second Late Jurassic Portuguese diplodocid taxon. Along with Dinheirosaurus, these Portuguese remains provide further evidence for a Late Jurassic palaeobiogeographic connection between Europe and North America. No dicraeosaurids are currently known from Europe, but rebbachisaurids are present in the Early Cretaceous, with weak evidence for the earliest known representative from the Late Jurassic of Spain; however, more complete material is required to recognize early members of this clade.

Smith, A. S., Araújo R., & Mateus O. (2012).  A new plesiosauroid from the Toarcian (Lower Jurassic) of Alhadas, Portugal. Acta Palaeontologica Polonica. 57(2), 257–266., Jan Abstractsmith__araujo__mateus_2012_a_new_plesiosauroid_from_the_toarcian_lower_jurassic_of_alhadas_portugal.pdf

A partial plesiosauroid skull from the São Gião Formation (Toarcian, Lower Jurassic) of Alhadas, Portugal is re−evaluated and described as a new taxon, Lusonectes sauvagei gen. et sp. nov. It has a single autapomorphy, a broad triangular parasphenoid cultriform process that is as long as the posterior interpterygoid vacuities, and also a unique character combination, including a jugal that contacts the orbital margin, a distinct parasphenoid–basisphenoid suture exposed between the posterior interpterygoid vacuities, lack of an anterior interpterygoid vacuity, and striations on the ventral surface of
the pterygoids. Phylogenetic analysis of Jurassic plesiosauroids places Lusonectes as outgroup to “microcleidid elasmosaurs”, equivalent to the clade Plesiosauridae. Lusonectes sauvagei is the only diagnostic plesiosaur from Portugal, and the westernmost occurrence of any plesiosaurian in Europe.

Myers, T. S., Tabor N. J., Jacobs L. L., & Mateus O. (2012).  Palaeoclimate of the Late Jurassic of Portugal: Comparison with the Western United States. Sedimentology. 59(6), 1695–1717., 2012//01/ Abstractmyers_et_al_2012_palaeoclimate_of_the_late_jurassic_of_portugal_comparison_with_the_western.pdfWebsite

Investigation of the palaeoclimatic conditions associated with Upper Jurassic strata in Portugal and comparison with published palaeoclimate reconstructions of the Upper Jurassic Morrison Formation in western North America provide important insights into the conditions that allowed two of the richest terrestrial faunas of this period to flourish. Geochemical analyses and observations of palaeosol morphology in the informally named Upper Jurassic Lourinhã formation of western Portugal indicate warm and wet palaeoclimatic conditions with strongly seasonal precipitation patterns. Palaeosol profiles are dominated by carbonate accumulations and abundant shrink-swell (vertic) features that are both indicative of seasonal variation in moisture availability. The δ18OSMOW and δDSMOW values of phyllosilicates sampled from palaeosol profiles range from +22·4‰ to +22·7‰ and −53·0‰ to −37·3‰, respectively. These isotope values correspond to temperatures of formation between 32°C and 39°C ± 3°, with an average of 36°C, which suggest surface temperatures between 27°C and 34°C (average 31°C). On average, these surface temperature estimates are 1°C higher than the highest summer temperatures modelled for Late Jurassic Iberia using general circulation models. Elemental analysis of matrix material from palaeosol B-horizons provides proxy (chemical index of alteration minus potassium) estimates of mean annual precipitation ranging from 766 to 1394 mm/year, with an average of approximately 1100 mm/year. Palaeoclimatic conditions during deposition of the Lourinhã formation are broadly similar to those inferred for the Morrison Formation, except somewhat wetter. Seasonal variation in moisture availability does not seem to have negatively impacted the ability of these environments to support rich and relatively abundant faunas. The similar climate between these two Late Jurassic terrestrial ecosystems is probably one of the factors which explains the similarity of their vertebrate faunas.

Mateus, O. (2012).  Age and paleoecology of mosasaurs and plesiosaurs from the Late Cretaceous South Atlantic margin at Bentiaba, Angola. Journal of Vertebrate Paleontology, Program and Abstracts, 2012. 180.. 180–181., 1 Abstract
n/a
Mateus, O. (2012).  Evidence for presence of clavicles and interclavicles in sauropod dinosaurs and its implications on the furcula-clavicle homology. Journal of Vertebrate Paleontology. 184–185., 1 Abstract

Clavicles and interclavicles are plesiomorphically present in Reptilia. However, several groups show reduction or even loss of these elements. Crocodylimorpha, e.g., lost the clavicles, whereas dinosaurs are generally interpreted to only preserve the clavicles, the theropod furcula representing an unique case of fused clavicles. In sauropods, reports of clavicles are relatively frequent in non-titanosauriforms. These elements are elongated, curved, and rather stout bones with a spatulate and a bifurcate end. However, they were always found as single bones, and differ from the relatively short and unbifurcated clavicles found articulated with the scapulae of basal sauropodomorphs. Elements from the Howe Quarry (Late Jurassic; Wyoming, USA) shed new light on these interpretations. Besides the elongated, curved bones (herein named morphotype A), also pairs of symmetric, L-shaped bones were recovered (morphotype B), associated with diplodocid dorsal and cervical vertebrae. Elements resembling morphotype B - articulated between the scapulae - have recently been reported from a diplodocid found near Tensleep, Wyoming. Taphonomic evidence, as well as the fact that they were preserved in symmetrical pairs, therefore implies that morphotype B represents the true sauropod clavicles. Contrary to earlier reports, morphotype A elements from the Howe Quarry, as well as of previously reported specimens show a symmetry plane following the long axis of the elements. It is thus possible that the morphotype A elements were single bones from the body midline. The only such element present in the pectoral girdle of tetrapods are the interclavicle and the furcula. Comparison with crocodilian and lacertiform interclavicles indicates that the bifurcate end of the sauropod elements might represent the reduced transverse processes of the anterior end, and the spatulate end would have covered the coracoids or sternal plates ventrally. The presence of both clavicles and interclavicles in the pectoral girdle stiffens the anterior trunk, and enhances considerably its stability. Such an enforcement might have been needed in diplodocids due to the strong lateral forces induced to the fore-limbs by the posteriorly placed center of mass (due to shorter fore- than hind-limbs), as well as lateral movements of the enormously elongated necks and tails. The absence of clavicles and interclavicles in titanosauriforms coincides with the development of wide-gauge locomotion style. The presence of interclavicles in sauropods supports the recently proposed homology of the furcula with the interclavicle, instead of representing fused clavicles. Interclavicles were thus not lost, but may have remained cartilaginous or have yet to be found in basal dinosauriforms.

Mateus, O. (2012).  Ontogenetical changes in the quadrate of basal tetanurans. Fundamental!. 101–104., 1 Abstract

Although nonavian theropod have received considerable interest in the last years, their ontogeny still remains poorly understood, especially the ontogenetical changes affecting their skull (Rauhutand Fechner, 2005). The quadrate, for instance, is preserved in several embryos and juvenile specimens belonging to many clades of theropods such as the Tyrannosauridae (Carr, 1999), Compsognathidae (Dal Sasso and Maganuco, 2011), Therizinosauroidea (Kúndrat et al., 2007), Oviraptoridae (Norell et al., 1994; Norell et al., 2001; Weishampel et al., 2008) and Troodontidae (Varrichio et al., 2002) but very little is usually said about the anatomy of this bone and no one has ever investigated ontogenetical variation in the nonavian theropod quadrate. The discovery of two quadrates belonging to embryos of the sinraptorid Lourinhanosaurus antunesi from Portugal and five isolated quadrates pertaining to juvenile, subadult and adult specimens of Spinosauridae from Morocco fills this gap and allows some ontogenetic information to be drawn for this bone in these two specific clades of Theropoda.

Mateus, O. (2012).  A preliminary report on coprolites from the Late Triassic part of the Kap Stewart Formation, Jameson Land, East Greenland. Bulletin of the New Mexico Museum of Natural History and Science. 57, 203–205., 1, Number NA Abstract

The basal part of the Triassic-Jurassic (Rhaetian-Sinemurian) Kap Stewart Formation, exposed at Jameson Land, East Greenland, yields an extensive coprolite collection from black, parallel-laminated mudstone (“paper shale”), representing an open lacustrine system. Preliminary investigations show three different types of coprolites: elongated cylindrical masses, composed of irregularly wrapped layers; elongated cylindrical masses with constriction marks; and spirally-coiled specimens.

Strganac, C., Ferguson, K.M, Jacobs J. L., Polcyn M. J., & Mateus O. (2012).  Age and paleoecology of mosasaurs and plesiosaurs from the Late Cretaceous South Atlantic margin at Bentiaba, Angola. Journal of Vertebrate Paleontology, Program and Abstracts, 2012. 180.strganac_et_al_mateus_2012_age_bentiaba_angola_2012_svp_abstract.pdf
Ribeiro, V., & Mateus O. (2012).  Chronology of the Late Jurassic dinosaur faunas, and other reptilian faunas, from Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, p. 161 ISSN 1937-2809. ribeiro__mateus_2012_chronology_dinosaurs_portugal_abstract_book_meeting_abstracts.pdf.pdf
Mateus, O., Polcyn M. J., Jacobs L. L., Araújo R., Schulp A. S., Marinheiro J., Pereira B., & Vineyard D. (2012).  Cretaceous amniotes from Angola: dinosaurs, pterosaurs, mosasaurs, plesiosaurs, and turtles. V Jornadas Internacionales sobre Paleontología de Dinosaurios y su Entorno. 71-105., Salas de los Infantes, Burgos Abstractmateus_et_al_2012_amniotes_from_angola_cretaceous_amniotes_from_angola_dinosaurs_pterosaurs_mosasaurs.pdf

Although rich in Cretaceous vertebrate fossils, prior to 2005 the amniote fossil record of Angola was poorly known. Two horizons and localities have yielded the majority of the vertebrate fossils collected thus far; the Turonian Itombe Formation of Iembe in Bengo Province and the Maastrichtian Mocuio Formation of Bentiaba in Namibe Province. Amniotes of the Mesozoic of Angola are currently restricted to the Cretaceous and include eucryptodire turtles, plesiosaurs, mosasaurs, pterosaurs, and dinosaurs. Recent collecting efforts have greatly expanded our knowledge of the amniote fauna of Angola and most of the taxa reported here were unknown prior to 2005.

Myers, T. S., Tabor N. J., Jacobs L. L., & Mateus O. (2012).  Estimating soil pCO2 using paleosol carbonates: implications for the relationship between primary productivity and faunal richness in ancient terrestrial ecosystems. Paleobiology. 38(4), 585–604. Abstractmyers_et_al_2012_estimating_soil_paleosols_portugal.pdf

In this paper we present a method for estimating soil pCO2 in ancient environments using the measured carbon-isotope values of pedogenic carbonates and plant-derived organic matter. The validity of soil pCO2 estimates proves to be highly dependent on the organic δ13C values used in the calculations. Organic matter should be sourced from the same paleosol profiles as sampled carbonates to yield the most reliable estimates of soil pCO2. In order to demonstrate the potential use of soil pCO2 estimates in paleoecological and paleoenvironmental studies, we compare samples from three Upper Jurassic localities. Soil pCO2 estimates, interpreted as a qualitative indicator of primary paleoproductivity, are used to rank the Late Jurassic terrestrial environments represented by the Morrison Formation in western North America, the informally named Lourinhã formation in Western Europe, and the Stanleyville Group in Central Africa. Because modern terrestrial environments show a positive correlation between primary productivity and faunal richness, a similar relationship is expected in ancient ecosystems. When the relative paleoproductivity levels inferred for each study area are compared with estimates of dinosaur generic richness, a positive correlation emerges. Both the Morrison and Lourinhã formations have high inferred productivity levels and high estimated faunal richness. In contrast, the Stanleyville Group appears to have had low primary productivity and low faunal richness. Paleoclimatic data available for each study area indicate that both productivity and faunal richness are positively linked to water availability, as observed in modern terrestrial ecosystems.

Tschopp, E., & Mateus O. (2012).  Evidence for presence of clavicles and interclavicles in sauropod dinosaurs and its implications on the furcula-clavicle homology. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, 184. ISSN 1937-2809 . 184. Abstracttschopp__mateus_2012_interclavicles_clavicles_svp_2012_abstract.pdf

Clavicles and interclavicles are plesiomorphically present in Reptilia. However, several groups show reduction or even loss of these elements. Crocodylimorpha, e.g., lost the clavicles, whereas dinosaurs are generally interpreted to only preserve the clavicles, the theropod furcula representing an unique case of fused clavicles. In sauropods, reports of clavicles are relatively frequent in non-titanosauriforms. These elements are elongated, curved, and rather stout bones with a spatulate and a bifurcate end. However, they were always found as single bones, and differ from the relatively short and unbifurcated clavicles found articulated with the scapulae of basal sauropodomorphs.
Elements from the Howe Quarry (Late Jurassic; Wyoming, USA) shed new light on these interpretations. Besides the elongated, curved bones (herein named morphotype A), also pairs of symmetric, L-shaped bones were recovered (morphotype B), associated with diplodocid dorsal and cervical vertebrae. Elements resembling morphotype B - articulated between the scapulae - have recently been reported from a diplodocid found near Tensleep, Wyoming. Taphonomic evidence, as well as the fact that they were preserved in symmetrical pairs, therefore implies that morphotype B represents the true sauropod clavicles.
Contrary to earlier reports, morphotype A elements from the Howe Quarry, as well as of previously reported specimens show a symmetry plane following the long axis of the elements. It is thus possible that the morphotype A elements were single bones from the body midline. The only such element present in the pectoral girdle of tetrapods are the interclavicle and the furcula. Comparison with crocodilian and lacertiform interclavicles indicates that the bifurcate end of the sauropod elements might represent the reduced transverse processes of the anterior end, and the spatulate end would have covered the coracoids or sternal plates ventrally.
The presence of both clavicles and interclavicles in the pectoral girdle stiffens the anterior trunk, and enhances considerably its stability. Such an enforcement might have been needed in diplodocids due to the strong lateral forces induced to the fore-limbs by the posteriorly placed center of mass (due to shorter fore- than hind-limbs), as well as lateral movements of the enormously elongated necks and tails. The absence of clavicles and interclavicles in titanosauriforms coincides with the development of wide-gauge locomotion style.
The presence of interclavicles in sauropods supports the recently proposed homology of the furcula with the interclavicle, instead of representing fused clavicles. Interclavicles were thus not lost, but may have remained cartilaginous or have yet to be found in basal dinosauriforms.

Araújo, R., Castanhinha R., Mateus O., & Martins R. (2012).  Late Jurassic theropod embryos from Porto das Barcas, Lourinhã formation, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, ISSN 1937-2809 . 57. Abstractaraujo_et_al_mateus_2012_dinosaur_eggs_portugal_svp_2012_abstract_book_meeting_abstracts.pdf.pdf

A clutch of several crushed eggs and embryos from the Late Jurassic (near the Kimmeridgian-Tithonian
boundary), Lourinhã Formation, Portugal contains a complete maxilla, erupted and scattered teeth,
and presacral vertebrae. The maxilla bears four teeth separated by individualized interdental plates,
the dorsal process of the maxilla is confluent with the maxillary body, the ventral rim of the antorbital
fossa is parallel to the tooth row, and the anterior border of the maxilla forms a right angle with the
ventral margin. The teeth are conical and recurved distally with carinae on mesial and distal sides. The
vertebrae are amphiplatyan, with a ventral pair of neurovascular foramina and heavily pitted articular
facets. These fossils allow unambiguous association of basal theropod osteology (Megalosauroidea) with
a new eggshell morphotype. Synchrotron micro-computed tomographic scanning (SRμCT), scanning
electron microscopy, and thin-sections under polarized and normal light revealed that the outer
ornamentation of the eggshell is composed of anastomosing ridges and islets, the pores communicate
near the outer region of the eggshells, and in radial section they are irregular canals that ramify towards
the surface. Micro-proto induced x-ray emission (micro-PIXE) analysis of the eggshell (excluding pores)
revealed the presence of Mg, Fe, Mn (0.33%, 0.27% and 0.18%, respectively) and several trace elements,
with a corresponding loss of Ca (39.4% detected but 40.0% expected for calcite), which suggests minimal
eggshell diagenesis. The eggshells do not luminesce, which could imply that no diagenetic alteration
took effect. However, the quenching effect of Fe2+ has to be taken into consideration. Conversely,
luminescence is observed in the pores since they are filled with sediment, composed of phyllosilicates,
as revealed by SRμCT, micro-PIXE and x-ray diffraction analyses.

Mateus, O. (2012).  New dinosaur and pterosaur tracksites from the Late Jurassic of Portugal. , Chongqing, China: 2012 Abstract Book of Qijiang International Dinosaur Tracks Symposium Abstractmateus_2012_dinosaur_tracks_portugal__abstract_book_qijiang_int_dinosaur_tracks_symposium.pdf

Portugal is rich on dinosaur remains (bones, eggs, and tracks) from Early Jurassic to Late
Cretaceous ages, but mainly from the Late Jurassic, in which dozen of tracksites have been reported.
Here are reported new or poorly known track localities:
1) Five tracksites share the preservation substrate (marine carbonated limestone), age (late Jurassic), geographic area (Leiria district of Portugal), kind of preservation (true tracks), and completeness (trackways of multiple individuals):
i) Praia dos Salgados includes eight trackways, mostly ornithopods and theropods, and one wide gauge sauropod, made in very soft sediment; some preserve the hallux impression.
ii) Serra de Mangues is mostly covered with vegetation but seems to include dozens of tracks comprising theropods, thyreophorans, ornithopods and sauropods.
iii) Sobral da Lagoa (Pedreira do Rio Real) include six trackways but poorly preserved;
and
iv) Serra de Bouro that preserves four sauropod trackways in one single layer.
v) Pedrógão, preserved, at least, one theropod trackway and several isolated tracks of
theropods and ornithopods were found in different layers in the Early Oxfordian.
2) The locality in Praia de Porto das Barcas yielded natural casts of stegosaur tracks
(including pes print with skin impression) and a very large sauropod pes print with about
1.2 m long pes.
3) A new pterosaur tracksite was found in the Late Jurassic of Peralta, Lourinhã (Sobral Member, Lourinhã Fm.; Late Kimmeridgian/Early Tithonian). More than 220 manus and pes tracks have been collected in about five square meters, all ascribed to pterosaurs. The tracks were produced in a thin mud layer that has been covered by sand which preserved them as sandstone mould infill (natural casts). The manus of the largest specimens is 13 cm wide and 5.5 cm long and the pes measures 14.5 cm in length and 9 cm in width. This shows the occurrence of very large pterosaurs in the Late Jurassic. Other pterosaur tracksites in the Late Jurassic of Portugal are: Porto das Barcas (Lourinhã Municipality), South of Consolação (Peniche Municipality), and Zambujal de Baixo (Sesimbra Municipality).

Vineyard, D., Mateus O., Jacobs L. L., Polcyn M. J., & Schulp A. (2012).  A new marine turtle from the Maastrichtian of Angola. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, 189. ISSN 1937-2809 . 189.vineyard_mateus_et_al_2012_euclastes_chelonia_turtle_angola_svp_2012_abstract.pdf
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. (2012).  Ontogenetical changes in the quadrate of basal tetanurans.. 10 th Annual Meeting of the European Association of Vertebrate Paleontologist ¡Fundamental! . 20, 101-104. Abstracthendrickx__mateus_2012_ontogenetical_changes_in_the_quadrate.pdf

Although nonavian theropod have received considerable interest in the last years, their ontogeny still remains poorly understood, especially the ontogenetical changes affecting their skull (Rauhut
and Fechner, 2005). The quadrate, for instance, is preserved in several embryos and juvenile specimens belonging to many clades of theropods such as the Tyrannosauridae (Carr, 1999), Compsognathidae (Dal Sasso and Maganuco, 2011), Therizinosauroidea (Kúndrat et al., 2007), Oviraptoridae (Norell et al., 1994; Norell et al., 2001; Weishampel et al., 2008) and Troodontidae (Varrichio et al., 2002) but very little is usually said about the anatomy of this bone and no one has ever investigated ontogenetical variation in the nonavian theropod quadrate. The discovery of two quadrates belonging to embryos of the sinraptorid Lourinhanosaurus antunesi from Portugal and five isolated quadrates pertaining to juvenile, subadult and adult specimens of Spinosauridae from Morocco fills this gap and allows some ontogenetic information to be drawn for this bone in these two specific clades of Theropoda.
I

Mateus, O., Carrano M. T., & Taquet P. (2012).  Osteology of the embryonic theropods from the Late Jurassic of Paimogo, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2012, p.137. ISSN 1937-2809. 137.mateus_et_al_2012_embryos_paimogo_portugal_svp_2012_abstract_book_meeting_abstracts.pdf.pdf