Publications in the Year: 2012

Conference Paper

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.

Mateus, O.  2012.  Ontogenetical changes in the quadrate of basal tetanurans, 1. Fundamental!. :101–104. 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.  Evidence for presence of clavicles and interclavicles in sauropod dinosaurs and its implications on the furcula-clavicle homology, 1. Journal of Vertebrate Paleontology. :184–185. 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.  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).

Mateus, O.  2012.  Age and paleoecology of mosasaurs and plesiosaurs from the Late Cretaceous South Atlantic margin at Bentiaba, Angola, 1. Journal of Vertebrate Paleontology, Program and Abstracts, 2012. 180.. :180–181. Abstract
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Martins, RMS, Araújo R, Mateus O, Castanhinha R, Pranzas PK, Beckmann F.  2012.  Tomography applied to the study of dinosaur fossils from the collection of the museum of Lourinhã (Portugal). I Congresso Internacional GeoCiencias na CPL. , Coimbra: Univ. Coimbramartins_et_al_2012_abstract.pdf

Conference Proceedings

Mateus, O, Polcyn MJ, Jacobs LL, Araújo R, Schulp AS, 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.

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, Carrano MT, 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
Martins, RMS, Beckmann F, Castanhinha R, Mateus O, Araújo R, Pranzas PK.  2012.  Synchrotron radiation-based micro-computed tomography applied to the characterization of dinosaur fossils from the Lourinhã Formation. 1 st . Meeting of Synchrotron Radiation Users from Portugal. :1-2., Caparica, Portugalmartins_et_al_2012_tomography_enurs_martins-rui_dinosaurfossils.pdf
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.

Journal Article

Smith, AS, Araújo R, Mateus O.  2012.  A new plesiosauroid from the Toarcian (Lower Jurassic) of Alhadas, Portugal. Acta Palaeontologica Polonica. 57:257-266., Number 2 Abstract
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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.

Martins, RMS, Castanhinha RA, Mateus O, Araújo R, Beckmann F, Schell N.  2012.  {Synchrotron radiation-based techniques applied to the study of dinosaur fossils from the collection of the museum of Lourinhã. Ciencia e Tecnologia dos Materiais. 24:153-156., Number 3-4 Abstract
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Smith, AS, Araújo R, Mateus O.  2012.  A new plesiosauroid from the Toarcian (Lower Jurassic) of Alhadas, Portugal, Jan. Acta Palaeontologica Polonica. 57(2):257–266. 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, TS, Tabor NJ, Jacobs LL, 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.

Strganac, C, Ferguson, K.M, Jacobs JL, Polcyn MJ, 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
Myers, TS, Tabor NJ, Jacobs LL, Mateus O.  2012.  Palaeoclimate of the late jurassic of portugal: Comparison with the western united states. Sedimentology. 59:1695-1717., Number 6 Abstract
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Mannion, PD, Upchurch P, Mateus O, Barnes RN, Jones MEH.  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, Jan. Journal of Systematic Palaeontology. 10(3):521–551. 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.

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

Mannion, PD, Upchurch P, Mateus O, Barnes RN, Jonese MEH.  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. 10:521-551., Number 3 Abstract
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Tschopp, E, Mateus O.  2012.  A sternal plate of a large-sized sauropod dinosaur from the Late Jurassic of Portugal. 10th Annual Meeting of the European Association of Vertebrate Paleontologists ¡Fundamental! . 20:263-266.: European Association of Vertebrate Paleontologist tschopp__mateus_2012_sternal_plate_sauropod_portugal.pdf