Article

A diverse Upper Jurassic dinosaur ichnofauna from central-west Portugal

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Mateus, O. & Milàn, J. 2009: A diverse Upper Jurassic dinosaur ichnofauna from central-west Portugal. Lethaia, Vol. 43, pp. 245–257.A newly discovered dinosaur track-assemblage from the Upper Jurassic Lourinhã Formation (Lusitanian Basin, central-west Portugal), comprises medium- to large-sized sauropod tracks with well-preserved impressions of soft tissue anatomy, stegosaur tracks and tracks from medium- to large-sized theropods. The 400-m-thick Lourinhã Formation consists of mostly aluvial sediments, deposited during the early rifting of the Atlantic Ocean in the Kimmeridgian and Tithonian. The stratigraphic succession shows several shifts between flood-plain mud and fluvial sands that favour preservation and fossilization of tracks. The studied track-assemblage is found preserved as natural casts on the underside of a thin bivalve-rich carbonate bed near the Tithonian–Kimmeridgian boundary. The diversity of the tracks from the new track assemblage is compared with similar faunas from the Upper Jurassic of Asturias, Spain and the Middle Jurassic Yorkshire Coast of England. The Portuguese record of Upper Jurassic dinosaur body fossils show close similarity to the track fauna from the Lourinhã Formation. □Dinosaur tracks, Lusitanian Basin, Portugal, skin impressions, Upper Jurassic.

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... Malafaia et al. [8] published a fragment of right maxilla (ALT-SHN. 116) whereas a mesialmost tooth (ML 962) was described by Hendrickx & Mateus [6]. Finally, embryonic remains (ML1188) discovered among a clutch of eggs have recently been reported by Araújo et al. [9]. ...
... In fact, the shallow maxillary fossa diagnostic of Torvosaurus tanneri [27] is not preserved in ALT-SHN. 116, and what has been interpreted as the anteriormost rim of the antorbital fenestra by Malafaia et al. ([8]: fig. 2B1) is, in fact, a diagnostic ridge located on the anteriormost corner of the lateral antorbital fossa (pers. ...
... However, the rim of the antorbital fenestra is not preserved in ALT-SHN. 116, and the position of the antorbital ridge relative to the antorbital fenestra cannot therefore be used as a diagnostic feature. Nonetheless, this fragment of maxilla includes several important features that support affinities with the genus Torvosaurus. ...
Article
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The Lourinhã Formation (Kimmeridgian-Tithonian) of Central West Portugal is well known for its diversified dinosaur fauna similar to that of the Morrison Formation of North America; both areas share dinosaur taxa including the top predator Torvosaurus, reported in Portugal. The material assigned to the Portuguese T. tanneri, consisting of a right maxilla and an incomplete caudal centrum, was briefly described in the literature and a thorough description of these bones is here given for the first time. A comparison with material referred to Torvosaurus tanneri allows us to highlight some important differences justifying the creation of a distinct Eastern species. Torvosaurus gurneyi n. sp. displays two autapomorphies among Megalosauroidea, a maxilla possessing fewer than eleven teeth and an interdental wall nearly coincidental with the lateral wall of the maxillary body. In addition, it differs from T. tanneri by a reduced number of maxillary teeth, the absence of interdental plates terminating ventrally by broad V-shaped points and falling short relative to the lateral maxillary wall, and the absence of a protuberant ridge on the anterior part of the medial shelf, posterior to the anteromedial process. T. gurneyi is the largest theropod from the Lourinhã Formation of Portugal and the largest land predator discovered in Europe hitherto. This taxon supports the mechanism of vicariance that occurred in the Iberian Meseta during the Late Jurassic when the proto-Atlantic was already well formed. A fragment of maxilla from the Lourinhã Formation referred to Torvosaurus sp. is ascribed to this new species, and several other bones, including a femur, a tibia and embryonic material all from the Kimmeridgian-Tithonian of Portugal, are tentatively assigned to T. gurneyi. A standard terminology and notation of the theropod maxilla is also proposed and a record of the Torvosaurus material from Portugal is given.
... Late Jurassic dinosaur tracksites are common and offer an abundance of fossils from localities all around the world (Fig. 1). In Europe, Late Jurassic dinosaur tracks are found in Switzerland (e.g., Meyer, 1993;Meyer and Lockley, 1996;Meyer and Thüring, 2003;Marty et al., 2003;Razzolini et al., 2017;Marty et al., 2018), France (Mazin et al., 1997(Mazin et al., , 2016Moreau et al., 2017), Spain (Canudo et al., 2005;Castanera et al., 2013;Alcalá et al., 2014;Cobos et al., 2014;Piñuela, 2015), Portugal (e.g., Lockley et al., 1994aLockley et al., , 1994bMeyer et al., 1994;Antunes and Mateus, 2003;Santos, 2008;Mateus and Milàn, 2010), Poland (Gierliński and Niedźwiedzki, 2002;Gierliński et al., 2009), Italy (Conti et al., 2005), Croatia (Mezga et al., 2007(Mezga et al., , 2017, whereas in North Africa the only occurrence of Late Jurassic dinosaur tracks is from Morocco (e.g., Dutuit and Ouazzou, 1980;Ishigaki, 1985bIshigaki, , 1985aBelvedere, 2008;Boutakiout et al., 2009;Marty et al., 2010;Nouri et al., 2011), probably due to sampling bias. ...
... In some cases, e.g., in Portugal (Mateus et al., 2006;Mateus and Milàn, 2010), Spain Rauhut et al., 2018), Germany (Lallensack et al., 2015) skeletal sites and tracksites are in close geological proximity allowing a more precise trackmaker identification. Dinosaur footprints have proved to be a valuable tool for palaeoecological analyses and reconstructions of faunal associations (e.g., Lockley, 1986;Belvedere et al., 2013). ...
... Portugal: The Late Jurassic deposits from the Lusitanian basin have yielded a large amount of tracksites located in different geological formations and localities (Lockley et al., 1992(Lockley et al., , 1994a(Lockley et al., , 1994b(Lockley et al., , 2000Lockley and Santos, 1993;Meyer et al., 1994;Antunes and Mateus, 2003;Santos, 2008;Mateus and Milàn, 2010;Castanera et al., 2016Castanera et al., , 2017. At Cabo Mondego (Figueira da Foz), tetradactyl footprints were identified in 1884 and this tracksite was the first to be described within the Lusitanian basin (Gomes, 1916). ...
Article
Late Jurassic theropod tracks are very common both in North Africa and Europe. Two recently described ichnotaxa Megalosauripus transjuranicus and Jurabrontes curtedulensis from the Kimmeridgian of Switzerland show the coexistence of two apex predators in the same palaeoenvironment. Similar tracks can be found in tracksites from the Iberian Peninsula and from Morocco. Here, we further explore the similarities among the Swiss ichnotaxa and the other tracks from Germany (Kimmeridgian), Spain (Tithonian-Berriasian), Portugal (Oxfordian-Tithonian) and Morocco (Kimmeridgian) through novel three-dimensional data comparisons. Specimens were grouped in two morphotypes: 1) large and gracile (30 < Foot Length<50cm) and 2) giant and robust (FL > 50cm). The analyses show a great morphological overlap among these two morphotypes and the Swiss ichnotaxa (Megalosauripus transjuranicus and Jurabrontes curtedulensis, respectively), even despite the differences in sedimentary environment and age. This suggests a widespread occurrence of similar ichnotaxa along the western margin of Tethys during the Late Jurassic. The new data support the hypothesis of a Gondwana-Laurasia faunal exchange during the Middle or early Late Jurassic, and the presence of migratory routes around the Tethys.
... The North American record yields the majority of the dinosaur skin reports (Bell, 2014), but some examples are described in Europe, Asia and South America, spanning a temporal range from the Jurassic to the Late Cretaceous. Regarding Europe, dinosaur sites containing skin impressions are reported from the Jurassic to the Late Cretaceous of Portugal and Spain (García-Ramos, Lires & Piñuela, 2002;Lockley et al. 2008;Mateus & Milàn, 2010;Vila et al. 2013;Navarrete et al. 2014). ...
... Such characteristics also appear in the Mirador de Vallcebre material. The shape and pattern of the skin impressions here described (polygonal, non-overlapping scales) are very similar to those reported by Currie, Badamgarav & Koppelhus (2003), Lockley et al. (2006Lockley et al. ( , 2008, Mateus & Milàn (2010) and Upchurch, Mannion & Taylor (2015) for various traces attributed to sauropods. They also resemble the integument traces preserved in a diplodocid sauropod track cast from the Morrison Formation (Platt & Hasiotis, 2006, fig. ...
... 7b, c). In terms of size, the scales from the Mirador de Vallcebre locality are larger than those of the Jurassic specimens from the United States (Platt & Hasiotis, 2006), of similar size to those from Korea and Mongolia (Currie, Badamgarav & Koppelhus, 2003;Lockley et al. 2006) and smaller than those from Portugal and Spain (Lockley et al. 2008;Mateus & Milàn, 2010). However, such texture and size variability might reflect different patterns from various parts of the animal's body and from different ages (Romano & Whyte, 2012). ...
Article
Southwestern Europe is one of the best regions for characterizing the dinosaur assemblages that prevailed just before the end-Cretaceous extinction. Aiming to better document this scenario, we provide the first evidence of dinosaur skin impressions in the red-beds of the Tremp Formation (southern Pyrenees). The impressions are assigned to sauropods (probably titanosaurians) on the basis of their scale morphology, arrangement and size. They represent a valuable tool for analysing the last occurrences of the sauropod clade before the K–Pg extinction, as they fall within chron C29r (latest Maastrichtian), thus representing some of the last in situ remains of this clade worldwide.
... In Europe, dinosaur tracks from the Late Jurassic and the Jurassic/Cretaceous boundary are reported from many countries such as England, France, Germany, Italy, Poland, Portugal, Spain and Switzerland (e.g. Dantas et al. 1994;Lockley and Meyer 2000 and references therein;Ensom 2002;Gierliński and Niedźwiedzki 2002;Gierliński and Sabath 2002;Marty et al. 2003;Garcίa-Hernández et al. 2003;Santisteban et al. 2003;Conti et al. 2005;Lockley et al. 2008;Marty 2008;Gierliński et al. 2009;Moratalla 2009;Mateus and Milàn 2010;Diedrich 2011;Avanzini et al. 2012;Lallensack et al. 2015). However, in France, with the exception of the Jura Mountains (eastern France; Fig. 1) where numerous Oxfordian-Tithonian trackways were recently discovered (Le Loeuff et al. 2006;Landry 2013;Cariou et al. 2014;Mazin et al. 2016), Late Jurassic dinosaur footprints were only reported from a few quarries of the Quercy area (Lot, southwestern France;Lange-Badré et al. 1996;Mazin et al. 1997Mazin et al. , 2000Gand et al. 2007;Mazin and Hantzpergue 2013;Fig. ...
... The morphology and dimension of the mark with digit impressions and the probable dinoturbation are very similar to small to medium-sized sauropod tracks (Platt and Hasiotis 2006;Lockley et al. 2008;Mateus and Milàn 2010;Thulborn 2012;Brusatte et al. 2015). Otherwise, this mark showing clear digit impressions but short heel imprint (Fig. 7b, c) is reminiscent of large thyreophoran tracks (e.g. ...
... Within the Late Jurassic dinosaur track record, the number of tracksites and trackways is clearly dominated by sauropods (D'Orazi Porchetti et al. 2016). In Europe, sauropod ichnites were reported from the Late Jurassic of many countries such as Germany (Lallensack et al. 2015), Poland (Gierliński and Niedźwiedzki 2002), Portugal Mateus and Milàn 2010), Spain (Lockley et al. 2007(Lockley et al. , 2008 and Switzerland (Meyer 1993;Ayer and Claude 2001;Marty 2008;Marty et al. 2003Marty et al. , 2010. In France, sauropod trackways were reported from several dozens of tracksite in the Oxfordian-Tithonian of the Jura Mountain (Le Loeuff et al. 2006;Landry 2013;Mazin et al. 2016). ...
Article
A new dinosaur tracksite was recently discovered in the Upper Jurassic deposits from the Oléron Island (Atlantic Ocean, western France). Tracks are located along the coastline in the tidal area of La Morelière near “Pointe de Chassiron”. They were observed in situ in an evaporitic interval from the “Purbeck beds” Unit. Sediments consist of calcite recrystallised after gypsum dissolution within limestone layers which are composed of thin laminites. The track-bearing surface shows abundant mud cracks and some ripple marks. Ichnofossils consist of medium-sized tridactyl footprints of theropods forming a narrow trackway. In order to obtain 3D reconstructions of the track-bearing surface, the trackway was scanned with a resolution of 0.5 mm using an Artec Eva 3D white-light scanner. As indicated by the track dimensions, the probable trackmaker is assigned to a medium-sized non-coelurosaur tetanuran, such as a megalosauroid or an allosauroid theropod. The palaeoenvironmental context is interpreted as a tidal flat in an evaporitic basin. In addition to the tridactyl tracks, we report sauropod- and thyreophoran-like tracks on two other surfaces from Chassiron–La Morelière.
... They characterised these tracks as nearly symmetrical, with absence of hallux, three relatively short digits emerging from a large basin-like depression representing a 'sole' or 'pad' to the foot, digits without evidence for phalangeal irregular, with an inward rotation of the distal end of digit III impression with respect to the trackway midline. Mateus and Milàn (2009, Fig. 9) have tentatively referred a giant theropod track with a PL of 79 cm from the Upper Jurassic Lourinhã Formation to Torvosaurus, which was, according to these authors, with a body length of 8-12 m, the largest theropod in the world during the Late Jurassic. They did not provide any ichnotaxonomical identification for this track, but this track differs from Jurabrontes because of the presence of a rounded heel area and notably because of a lateral swelling in the middle of dIII (Mateus & Milàn 2009;pers. ...
... Mateus and Milàn (2009, Fig. 9) have tentatively referred a giant theropod track with a PL of 79 cm from the Upper Jurassic Lourinhã Formation to Torvosaurus, which was, according to these authors, with a body length of 8-12 m, the largest theropod in the world during the Late Jurassic. They did not provide any ichnotaxonomical identification for this track, but this track differs from Jurabrontes because of the presence of a rounded heel area and notably because of a lateral swelling in the middle of dIII (Mateus & Milàn 2009;pers. obs. ...
... Megalosauridae or 'megalosaurs' are poorly understood, both in their anatomy and their phylogenetic affinities (Molnar et al. 1990;Holtz 2000;Benson et al. 2008), andThulborn (2001) stated, 'there exists no definite conception of megalosaurs or of their tracks' . However, Torvosaurus is a member of the Megalosauridae known from Colorado (Galton & Jensen 1979), and also known from Portugal Mateus & Milàn 2009, Hendrickx & Mateus 2014Malafaia et al. 2017). Mateus & Milàn (2009) have tentatively referred a giant-sized theropod track with a total length of 79 cm to Torvosaurus. ...
Article
Jurabrontes curtedulensis, a new ichnogenus and species of Late Jurassic giant theropod dinosaur track is described based on very well-preserved and morphologically-distinct tracks, all carefully excavated along federal highway A16 (Canton Jura, NW Switzerland). All trackways were systematically documented including parameter measurements, descriptions, outline drawings, orthophotos and laserscans. Jurabrontes is characterised by sub-equal track length and width, a small anterior triangle, weak mesaxony, three blunt digits (dII-III-IV) with pronounced (sub)triangular claw marks, a rounded heel, and clear phalangeal pad impressions. The combination of features of Jurabrontes is typical for a theropod (and not ornithopod) trackmaker. Jurabrontes is compared to other similar ichnotaxa and unnamed tracks of large theropods from the Early Jurassic to Late Cretaceous, from which it is clearly different. The sheer size of the largest tracks, that are amongst the largest worldwide and of similar size to Tyrannosauripus from the Late Cretaceous, suggests a ‘megalosaurid’ or large allosaurid theropod as a trackmaker. The presence of such large theropod tracks in tidal-flat deposits of the Jura carbonate platform and associated with small to large sauropod tracks has important palaeoecological implications for the dinosaur community and for palaeoenvironmental and palaeogeographical reconstructions. Jurabrontes - urn:lsid:zoobank.org:act:B482D2AF-637A-4B2D-8B0B-FEAD54CA2A26 J. curtedulensis- urn:lsid:zoobank.org:act:30D45944-5A2B-45E1-89B9-20298E475D51
... In other cases, the term axony was characterized in a purely morphological/dimensional fashion, without direct hints on the dynamic of locomotion and on different roles that pedal elements can play in body weight support and propulsive thrust (e.g., Schulp and Brokx 1999;Whyte and Romano 2001;Gierlinsky and Sabath 2002;Thulborn 2006;Mateus and Milàn 2010;Voigt et al. 2010;Klein et al. 2011;Meldrum et al. 2011;Costa Da Silva et al. 2012;Kim et al. 2012) (Figure 1). ...
... Baird 1957;Cobos et al. 2016) could instead offer the opportunity to define different axonies according to a specific phase of the locomotion cycle (see Padian and Olsen 1984), and then to the particular footprint portion involved in each locomotion phase. For example, in describing footprints attributed to sauropods, Mateus and Milàn (2010) correctly defined the pes as entaxonic, considering that the medial digits are the most deeply impressed. However, by analyzing the differential depth (as far as possible) from the figures presented in the paper (Mateus and Milàn 2010), the footprint proximal portion seems to be more deeply imprinted laterally, thus defining a condition of ectaxony during a specific phase of the locomotion cycle. ...
... For example, in describing footprints attributed to sauropods, Mateus and Milàn (2010) correctly defined the pes as entaxonic, considering that the medial digits are the most deeply impressed. However, by analyzing the differential depth (as far as possible) from the figures presented in the paper (Mateus and Milàn 2010), the footprint proximal portion seems to be more deeply imprinted laterally, thus defining a condition of ectaxony during a specific phase of the locomotion cycle. In the same paper, the authors defined the studied theropod footprints as mesaxonic; the material, however, should be regarded as ectaxonic (i.e. more imprinted laterally) at least in the initial touch down phase (Mateus and Milàn 2010, p. 250, fig. ...
Article
Two meanings of the term axony are found in the ichnological literature. Multiple meanings may prove to be a double-edge sword, complicating scientific communication. In vertebrate ichnology the first meaning of axony relies on aspects of locomotion related to the body weight support and propulsive thrust. A second one concerns axony as a purely geometric and dimensional descriptor. These approaches are based on a static view of the impression process, implying the loss of much important information. Here we report an analysis of shallowly impressed footprints referred to the ichnotaxa Ichniotherium sphaerodactylum and Dimetropus osageorum. The analysis was carried out by considering the track registration as a dynamic process and attempting to identify and describe axony conditions during movements. Variations in the axony conditions can be understood in the light of the producer's foot anatomy and the reciprocal relations between foot bone elements. The concept of axony can be a useful tool in ichnological practice only when it is related to the complex dynamic of locomotion and the resulting track registration. It can help in restoring the interconnections between track and track-maker, re-establishing the biological significance of tetrapod footprints.
... Furthermore, isolated sauropod casts in the Lourinhã area and the locality of Porto Dinheiro (Lourinhã Formation, Kimmeridgian-Tithonian) have been recovered as well. In these cases, the tracks have been preserved as sandstone casts in fluvial environments (Milàn, Christiansen, and Mateus, 2005;Mateus and Milàn, 2010). ...
... Nonetheless, some of them are actually right tracks, whereas others are preserved as a natural casts, so in these cases, they have been mirrored. Within the Lusitanian Basin tracksites, we have selected single tracks from trackways MNDPDSA-G1 (manus) and MNDPDSA-G5 (manus and pes) at Galinha tracksite; trackways PM5-1 (manus) and PM5-2 (pes) at Pedra da Mua level 5 tracksite (a gypsum plaster cast of the latter, MNHN-MG-P270, is housed in the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa); and isolated manus (ML 965) and pes casts (ML 1151) described by Milàn, Christiansen, and Mateus (2005) and Mateus and Milàn (2010) The specimens have been described morphologically and photographed. The values given in Tables 8.1 and 8.2 represent data taken from perpendicular photographs of the tracks using the software ImageJ, with measurements taken by the authors on the specimens and data from the original papers. ...
... MUJA-1896 shows a lateral notch behind digit V. Among the Portuguese localities, at Porto Dinheiro locality, pes casts showing four digit marks (track 8 in Fig. 8.3) have also been reported (Mateus and Milàn, 2010). ML 1151shows blunt claw marks and division into digital pads. ...
... Giant (PL>70 cm) theropod ichnotaxa such as Tyrannosauripus from the Late Cretaceous of New Mexico [93], Bellatoripes from the Late Cretaceous of Canada [94] and some other large to giant unnamed tracks from the Late Jurassic of Portugal [95] and Morocco [96,97] are not considered here, as they significantly differ from Megalosauripus and Megalosauripus transjuranicus. Giant (PL>50 cm) theropod tracks from Highway A16 that are significantly different from M. transjuranicus have recently been named Jurabrontes curtedulensis [98]. ...
... Megalosauridae or 'megalosaurs' are poorly understood, both in their anatomy and their phylogenetic affinities [149,150], and Thulborn [5] stated: "there exists no definite conception of megalosaurs or of their tracks". However, Torvosaurus is a member of the Megalosauridae known from Colorado [151] and Portugal [95,142]. Its body length ranges from 8-12 m, which is too large for the studied tracks. ...
Article
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A new ichnospecies of a large theropod dinosaur, Megalosauripus transjuranicus, is described from the Reuchenette Formation (Early–Late Kimmeridgian, Late Jurassic) of NW Switzerland. It is based on very well-preserved and morphologically-distinct tracks (impressions) and several trackways, including different preservational types from different tracksites and horizons. All trackways were excavated along federal Highway A16 near Courtedoux (Canton Jura) and systematically documented in the field including orthophotos and laserscans. The best-preserved tracks were recovered and additional tracks were casted. Megalosauripus transjuranicus is characterized by tridactyl tracks with clear claw and digital pad impressions, and notably an exceptionally large and round first phalangeal pad on the fourth digit (PIV1) that is connected to digit IV and forms the round heel area. Due to this combination of features, M. transjuranicus clearly is of theropod (and not ornitho-pod) origin. M. transjuranicus is compared to other Megalosauripus tracks and similar ichno-taxa and other unassigned tracks from the Early Jurassic to Early Cretaceous. It is clearly different from other ichnogenera assigned to large theropods such as Eubrontes–Grallator from the Late Triassic and Early Jurassic or Megalosauripus–Megalosauropus–Bueckebur-gichnus and Therangospodus tracks from the Late Jurassic and Early Cretaceous. A second tridactyl morphotype (called Morphotype II) is different from Megalosauripus trans-juranicus in being subsymmetric, longer than wide (sometimes almost as wide as long), with blunt toe impressions and no evidence for discrete phalangeal pad and claw marks. Some Morphotype II tracks are found in trackways that are assigned to M. transjuranicus, to M.? transjuranicus or M. cf. transjuranicus indicating that some Morphotype II tracks are intra-trackway preservational variants of a morphological continuum of Megalosauripus transjura-nicus. On the other hand, several up to 40 steps long trackways very consistently present Morphotype II features (notably blunt digits) and do not exhibit any of the features that are typical for Megalosauripus (notably phalangeal pads). Therefore, it is not very likely that these tracks are preservational variants of Megalosauripus transjuranicus or PLOS ONE | https://doi.
... Up until now it has not been possible to assess if this gap may be due to differential preservation linked to ecological segregation as seen for ornithischians and saurischians in the Morrison Formation (Foster 2013) or if it reflects inter-specific competition between different herbivorous taxa. Beside skeletal and dental ornithopod fossils, various tracks and track-sites have been reported from the Lourinhã Formation (Mateus and Milàn 2009) including a giant sized Iguanodontipus-like footprint which is 70 cm long, suggesting the presence of a large sized iguanodontian possibly larger than any previously known Late Jurassic species (Mateus and Milàn 2008), but also smaller sized camptosaurid tracks (as figured by Antunes and Mateus 2003: fig. 11) and Dinehichnus-like ones, probably made by dryosaurids or other small bipedal neornithischians (Mateus and Milàn 2009). ...
... Beside skeletal and dental ornithopod fossils, various tracks and track-sites have been reported from the Lourinhã Formation (Mateus and Milàn 2009) including a giant sized Iguanodontipus-like footprint which is 70 cm long, suggesting the presence of a large sized iguanodontian possibly larger than any previously known Late Jurassic species (Mateus and Milàn 2008), but also smaller sized camptosaurid tracks (as figured by Antunes and Mateus 2003: fig. 11) and Dinehichnus-like ones, probably made by dryosaurids or other small bipedal neornithischians (Mateus and Milàn 2009). ...
Article
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Ornithopods are one of the most speciose group of herbivorous dinosaurs, rising during the Jurassic and getting extinct at the Cretaceous–Paleogene boundary. However, most of the attention has been given to derived forms (hadrosaurids). Herein, cranial and post-cranial ornithopod material from the Upper Jurassic Lourinhã Formation and housed at Museu da Lourinhã is described and discussed. Comparison and phylogenetic analyses has allowed the attribution of the material either to Dryosauridae or to Ankylopollexia. The large-sized taxa conservatively ascribed to Ankylopollexia, resemble more closely Early Cretaceous styracosternans than Late Jurassic taxa. Due to the lack of autapomorphic characters, it was not possible to assign the material to any of the two valid Jurassic ornithopod Portuguese species, Draconyx loureiroi and Eousdryosaurus nanohallucis, although phylogenetic analyses hint a close relationship between the Lourinhã dryosaurid material and E. nanohallucis. Principal Component Analysis plotting limb bones proportions indicates a not fully mature ontogenetic stage for the Portuguese specimens. Comparing the Portuguese ornithopod fauna with the one in Morrison Formation and Kimmeridge Clay Formation, it is remarked the key-role of Portugal to understand biogeographic patterns in the distribution of iguanodontians.
... The lowest occurrence is at Vale de Pombas, Amoreira-Porto Novo Member, Late Kimmeridgian, while the uppermost occurrence is at Porto das Barcas, top of Praia Azul Member, base of Tithonian. In this study, 24 new occurrences of stegosaur tracks from the Lourinhã Formation are presented, as well as 13 tracks studied in previous papers (Mateus & Milàn, 2008;Mateus & Milàn, 2010;Mateus et al., 2011). The previous studies authors attributed the tracks to Deltapodus and described them as eleven pes and two manus, some of them being the biggest known of the ichnogenus. ...
... Small lizards, are also known from the Upper Jurassic. Cteniogenys reedi is a small lizard described for Guimarota and Lourinhã (Seiffert, 1973;Ribeiro & Mateus, 2012) There are nine known crocodylomorphs from the Portuguese Upper Jurassic (Mateus et Milàn, 2010). Some of them are small but others are the largest forms found, like the over nine meter long Machimosaurus hugii, whose skull, found in Guimarota mine, is quite complete (Krebs and Schwarz, 2000). ...
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No decurso da 4ª edição do Mestrado em Paleontologia (NOVA, UÉ), a Comissão de Curso, o Departamento de Ciências da Terra da FCT-NOVA e o Departamento de Geociências da ECT-Universidade de Évora, entenderam oportuno promover o Simpósio NOVAPALEO 2016, a realizar nos dias 13 e 14 de maio de 2016, com o intuito de reunir jovens investigadores em Paleontologia, partilhar experiências e colocar em destaque a investigação científica que tem vindo a ser desenvolvida em Portugal neste domínio do saber. O evento inclui duas Conferências por convite e um workshop.
... In well-registered sauropod manus or pes prints the sole shows impressions or wrinkles indicative of sole pads (cf. Farlow, Pittman, and Hawthorne 1989;Dalla Vecchia and Tarlao 2000;Mil an, Christiansen, and Mateus 2005;Platt and Hasiotis 2006;Mateus and Mil an 2010;Huh et al. 2003;Romano and Whyte 2012;Castanera et al. 2016b;Hall, Fragomeni, and Fowler 2016). Occasionally the skin texture of the sole is recorded as an abutting polygonal pattern (Lockley et al. 1998Currie, Badamgarav, and Koppelhus 2003;Platt and Hasiotis 2006;Kim et al. 2010;Mateus and Mil an 2010;Navarette et al. 2014;Piñuela Suarez 2015;Castanera et al. 2016a;Fondevilla et al. 2017;Paik et al. 2017). ...
... Farlow, Pittman, and Hawthorne 1989;Dalla Vecchia and Tarlao 2000;Mil an, Christiansen, and Mateus 2005;Platt and Hasiotis 2006;Mateus and Mil an 2010;Huh et al. 2003;Romano and Whyte 2012;Castanera et al. 2016b;Hall, Fragomeni, and Fowler 2016). Occasionally the skin texture of the sole is recorded as an abutting polygonal pattern (Lockley et al. 1998Currie, Badamgarav, and Koppelhus 2003;Platt and Hasiotis 2006;Kim et al. 2010;Mateus and Mil an 2010;Navarette et al. 2014;Piñuela Suarez 2015;Castanera et al. 2016a;Fondevilla et al. 2017;Paik et al. 2017). ...
Article
Three parallel, manus-only sauropod trackways from the Coffee Hollow A-Male tracksite (Glen Rose Formation, Kendall County, Texas) were studied separately by researchers from the Heritage Museum of the Texas Hill Country and the Houston Museum of Natural Sciences. Footprint and trackway measurements generally show good agreement between the two groups’ data sets. Footprints appear to be shallowly impressed true tracks rather than undertracks. One of the Coffee Hollow trackways shows marked asymmetry in the lengths of paces that begin with the left as opposed to the right forefoot, and two of the Coffee Hollow trackways are unusually broad. The Coffee Hollow trackways differ enough from the manus portions of other Glen Rose Formation sauropod trackways to suggest that they were made by a different kind of sauropod. Greater differential pressure exerted on the substrate by the forefeet than the hindfeet probably explains the Coffee Hollow trackways, like other manus-only sauropod trackways, but the possibility that they indicate unusual locomotion cannot at present be ruled out.
... Some authors have considered the relative width of a trackway as potentially providing insights into the identification of tridactyl trackmakers, with theropods considered more likely to display a narrow gauge trackway, and ornithopods a more wide gauge trackway (e.g. Lockley et al., 1998;Day et al., 2004;Kim et al., 2009;Mateus and Mil an, 2010). However, this is not diagnostic within either trackmaker grouping, as both can have a range of gauge widths (e.g. ...
... However, this is not diagnostic within either trackmaker grouping, as both can have a range of gauge widths (e.g. Day et al., 2002Day et al., , 2004Kim et al., 2009;Mateus and Mil an, 2010). Similar to theropodan trackmakers, ornithopodan trackmakers appear to have been capable of very narrow pedal gauges (e.g. ...
Article
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The largest dinosaurian tracks at Lark Quarry, central-western Queensland, Australia, were re-examined using revised analytical protocols that incorporate three-dimensional (3D) structure. Comparisons were made with archival photographs, replica specimens (c. 1977) and the in situ tracks (2013) to account for changes to the track surface. Damage caused both during and after the excavation of the tracks was evident, and in cases where the archival photographs and 1970's replicas strongly differ from the in situ tracks, it is apparent that restoration has modified the original track morphology.
... There are also reports of the ichnogenus from the Middle Jurassic Lealt Shale Formation in Scotland (dePolo et al., 2020), from the Middle Jurassic Lajas Formation in Argentina (Pazos et al., 2019), and from the Aptian Tirgan Formation in Iran (Abbassi et al., 2018). Since its first report from the Saltwick Formation, D. brodricki has been referred to tracks from the Lourinhã Formation, in Portugal (Mateus & Milàn, 2010;Mateus et al., 2011), the Morrison Formation, in Utah (Milàn & Chiappe, 2009), and the Villar del Arzobispo Formation, the Camarillas Formation, and the Artoles Formation, in Spain (Herrero Gascón & Pérez Lorente, 2017). There are two more Deltapodus ichnospecies currently described: D. ibericus Cobos et al., 2010 from the Villar del Arzobispo Formation (Cobos et al., 2010), and D. curriei Xing et al., 2013 from the Qingshuihee -Hutubihe Formations (Tugulu Group), in China (Xing et al., 2013). ...
... Sauropods seem also underrepresented in the track record of the Lourinhã Formation when comparing with the Morrison Formation (Foster et al., 2006;Lockley et al., 2015Lockley et al., , 2017. This underrepresentation could be due to a collecting bias, as sauropod tracks in the Lourinhã Formation can be big-sized and not easy to collect (Mateus & Milàn, 2010). On the other hand, the ankylosaur track record in both formations reflect the scarcity of their respective skeletal remain (Lockley et al., 2017), and it has been suggested that the underrepresentation of stegosaur tracks toward their skeletal fossil record could be due to a dichotomy between paleoenvironmental preference and preservational modes in the Morrison Formation (Lockley et al., 2017). ...
Article
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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.
... The Late Jurassic sauropod tracksites are the most significant in terms of number of footprints and were the subject of the first references in the late 1970s and the 1980s (Antunes 1976(Antunes , 1981 More recent research has shown considerable dinosaur ichnodiversity in the Upper Jurassic deposits of the Lusitanian Basin. For instance, various morphotypes have been described in the Lourinhã area, such as a giant ornithopod track (Mateus & Milàn 2008), plus tracks of medium-to large-sized sauropods, stegosaurs (Deltapodus) and medium-to large-sized theropods (Mateus & Milàn 2010a;Mateus et al. 2011). New research is providing data about the ichnotaxonomic relationships among some of the tracks described, as well as new findings Castanera et al. 2020). ...
... Nonetheless, the specimen has some features that are worthy of mention, namely the orientation of the digits and claw marks. Several examples of sauropod pes prints have been described in the Late Jurassic and Early Cretaceous of the Iberian Peninsula, including the Lusitanian Basin (Mateus & Milàn 2010a;Castanera et al. 2016b), being characterized by claw marks oriented anterolaterally for digits I-III. In the case of ICNO.35, the orientation of the three digits, including digit I, is more lateral than anterolateral. ...
Article
Jurassic units of the Lusitanian Basin, housed at the Sociedade de História Natural in Torres Vedras, are here described. They were collected from three different geological formations, the Praia da Amoreira‐Porto Novo (upper Kimmeridgian) and the Alcobaça (Kimmeridgian‐lower Tithonian) formations in the Consolação Sub‐basin and the Freixial Fm. (middle‐upper Tithonian) in the Turcifal Sub‐basin. Four different theropod morphotypes are identified as follows: cf. Jurabrontes isp., Megalosauripus cf. transjuranicus, Grallatoridae indet. and an indeterminate morphotype (Theropoda indet.) that have affinities with other Therangospodus‐like tracks described in Europe. An indeterminate sauropod track is also identified. These five morphotypes suggest high saurischian dinosaur ichnodiversity, similar to that seen in other European Late Jurassic areas (e.g. the Swiss Jura Mountains), but represent just a portion of the higher diversity exhibited by the osteological record in the Lusitanian Basin. Further, one crocodylomorph pes track identified as Crocodylopodus isp. and swim tracks assigned to Characichnos isp., possibly also produced by crocodylomorphs, are also identified. The newly identified ichnotaxa, together with the older and other recent identifications, indicate ichnodiversity comparable with the richest coeval Upper Jurassic units.
... Sítios com pegadas de dinossáurios terópodes são abundantes em Portugal e conhecidos desde o Jurássico Médio (Bathoniano) de Vale de Meios e Algar dos Potes (Santos et al., 2000a;Razzolini et al., 2016), Jurássico Superior (Oxfordiano) do Cabo Mondego (Gomes, 1916;Lockley et al., 1998), (Oxfordiano-Kimmeridgiano) da Praia do Cavalo e da Pedreira da Ribeira do Cavalo (entretanto desaparecida; Dantas et al., 1994), das Pedras Negras e da Praia dos Salgados (Santos, 1998), (Tithoniano) de Porto Dinheiro, Caniçal, Praia da Areia Branca, Escadinhas, Praia de Valmitão e Praia da Corva (Santos, 1998;Mateus & Milán, 2010), além de Lagosteiros (Lockley et al., 1994), bem como do Cretácico Inferior (Hauteriviano) de Lagosteiros (Antunes, 1976), Barremiano da Praia da Salema Santos et al., 2013) e de Areia do Mastro (Figueiredo et al., 2021), Aptiano de Praia Grande do Rodízio (Madeira & Dias, 1983), Aptiano-Albiano da Praia dos Olhos de Água (Mateus & Antunes, 2003), Albiano de Parede (Santos et al., 2015) e do Cretácico Superior (Cenomaniano) de Carenque (Santos et al., 1992) . ...
... As pegadas tridáctilas gigantes têm sido atribuídas em Portugal a Torvosaurus (Mateus & Milán, 2010;Belvedere et al., 2019). Atendendo ao registo osteológico encontrado até agora na Praia de Pedrógão, que apenas identifica a presença de terópodes indeterminados, e ao facto das pegadas poderem atingir 65 cm de comprimento, com proporções só comparáveis a Torvosaurus gurneyi (Hendrickx & Mateus, 2014) ou a terópodes carcharodontossaurídeos (Malafaia et al., 2018), permite-nos antecipar a identificação no futuro de táxones destes grandes terópodes no Oxfordiano médio da Praia de Pedrógão, como os mais prováveis produtores das pegadas gigantes que se encontram na Formação de Cabaços. ...
Article
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https://www.cpgp.pt/boletim.php?fbclid=IwAR1teB_2idgzSQ2c-C_BDXKM_bh69OPTt27UwB44VEb039yYmNHvOpE-Fw0
... The sum of the Las Sereas site data (Torcida Fernández-Baldor et al., 2001;2012; this study) suggest that its ichno-been described in the Lourinhã Formation and the Espichel section of Tithonian age (Lockley et al., 1994c;Santos, 2003;Mateus and Milàn, 2010). In both cases, well-preserved footprints are subtriangular in shape and have laterally curved digit impressions. ...
... Las Sereas 7 and La Pedraja (both part of Las Sereas site) are unique tracksites formed in a lacustrine/palustrine environment in the Tithonian-Berriasian interval of the Iberian Peninsula. In Portugal, some tracksites have been described in Tithonian distal alluvial fan deposits with some coastal intercalations (Mateus and Milàn, 2010) and in shallow marine environments (Lockley et al., 1994b;Meyer et al, 1994;Santos, 2003). In the Tithonian-Berriasian transition, several tracksites have been recorded in the Iberian Range (Spain). ...
Article
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The Las Sereas site includes at least 14 ichnological outcrops along 5.6 km, in the Lara area, southwest Burgos Province. 67 ichnites of dinosaurs are documented at Las Sereas 7, identified as theropod and sauropod trackways occurring in shallow carbonates of lacustrine environment. Sauropod trackways have intermediate-gauge and low heteropody, and show different anatomical features to other tracks found in the ichnological record, especially in the disposition and orientation of pes digits. They are similar to Polyonyx from the Middle Jurassic of Portugal. However, since they do not preserve reliable manus data they are classified as aff. Polyonyx. The three sauropod trackways are related to the same kind of trackmaker. They differ from each other only in size, and gregarious behavior has not been detected. Analysis of these trackways reveals changes in travel direction even when there are few tracks in each sequence. At the Las Sereas 7 tracksite, the pace length (PL), width of the angulation pattern (WAP) and the WAP/PL ratio and depth analysis via photogrammetry show a direction change in two sauropod trackways. This tracksite and that at La Pedraja are unique in the Tithonian-Berriasian interval of the Iberian Peninsula that occur in a lacustrine environment, and could be indicate of the relationship between the diversity of Iberian, Tithonian-Berriasian sauropod tracks and sedimentary environments.
... Tracks attributed to s -Late Jurassic of Spain, Portugal, Morocco and Utah (Alcala et al., 2012;Cobos et al., 2010;Mampel et al., 2010Mampel et al., -2011Lockley et al., 2008;Piñuela et al., 2007;Mateus et Milàn, 2009;Belvedere et Mietto, 2010;Milàn and Chiappe 2009;Lockley et al., 2017;Whyte and Romano, 1994). ...
Thesis
Les opportunités de reconstituer l’écologie et les comportements d’individus et d’espèces de vertébrés éteints sont rares. Plusieurs grands assemblages de vertébrés fossiles datant de la fin du Jurassique au début du Crétacé fournissent des informations sur la biodiversité et les communautés de vertébrés terrestres. Cependant, à l’exception du Lagerstätte de Las Hoyas (Crétacé inférieur, Espagne), les fossiles de ces assemblages ont été retrouvés sur un large éventail géographique, stratigraphique et/ou temporel. Cette thèse présente les résultats d’une étude taphonomique, couplée avec des données néotaphonomiques, paléontologiques, ichnologiques, sédimentologiques et géochimiques, de l’assemblage de macrorestes fossiles de vertébrés du site du Crétacé inférieur d’Angeac-Charente, situé dans l’Ouest de la France. Neuf campagnes de fouilles menées depuis 2010 ont permis de récolter plusieurs milliers de macrorestes de vertébrés appartenant à 16 taxons différents, auxquels viennent s’ajouter d’abondants microrestes appartenant à 27 taxons de vertébrés supplémentaires, plusieurs centaines de coprolithes de termites et de vertébrés riches en inclusions végétales et osseuses, des centaines de remplissages naturels d’empreintes de dinosaures, de nombreux restes de végétaux, des moules de mollusques bivalves et gastropodes, ainsi que des ostracodes. L’ensemble sédimentaire est dominé par des argiles de décantation interrompues par des dépôts localisés de plus haute énergie (cours d’eau, inondation) déposé dans un environnement de zone humide de type marécage. De nombreuses figures de déformations synsédimentaires formées en milieu liquide sont conservées sous forme de « scènes figées ». Des empreintes attribuées à l’ichnogenre Deltapodus et conservées sous forme de remplissages de dépôt direct ont été produites par un groupe multigénérationnel de stégosaures. De nombreuses empreintes de sauropodes sont conservées en 3-D ou en 4-D sous forme de remplissages de profondeurs variables. L’assemblage osseux est multitaxique, très diversifié et comprend des taxons terrestres, amphibies et aquatiques d’eau douce. Il contient une combinaison de macrofossiles et microfossiles et s’est formé de manière complexe, avec l’intervention de multiples processus d’origine biologique, écologique et physique. Il est dominé par les dinosaures, en particulier une nouvelle espèce d’ornithomimosaure non-ornithomimidé, dont de nombreux restent résultent d’une mort en masse d’un troupeau multigénérationnel dominé par des juvéniles et subadultes. D’abondants restes de sauropodes ont été transportés sur une plus ou moins grande distance avant d’être déposés sur le site. De nombreuses traces de morsure à la surface des restes de carapaces appartenant à plusieurs individus de tortues Pleurosternon bullockii résultent du comportement de prédation du crocodylomorphe Goniopholis, avec l’utilisation de la technique « casse-noix ». Les modifications osseuses post-dépositionnelles, comme la désarticulation, les déplacements, réorientations, fractures et traces de surface, résultent principalement de l’intense piétinement par les dinosaures. Le Lagerstätte d’Angeac-Charente ouvre une fenêtre d’une résolution spatiale et temporelle exceptionnellement fine sur un écosystème terrestre ouest-européen du tout début du Crétacé dont les organismes étaient en interaction directe entre eux et avec leur environnement.
... This is in accordance with the record from the Late Jurassic Alcobaça and Lourinhã formations (Kimmeridgian-Tithonian) of Portugal, where there have been described remains of both dryosaurids: Aff. Dryosaurus sp., and "camptosaurids": Draconyx loureroi and Uteodon aphanoecetes Malafaia et al., 2010;Mateus and Milàn, 2010). ...
... The recognition of gait abnormalities in dinosaur trackways has been reported in both isolated tracks (Ishigaki, 1986(Ishigaki, , 1989Mateus & Mil an, 2010) and trackways (Abel, 1935;Avanzini, Piñuela, & García Ramos, 2008;Dantas, Santos, Lockley, & Meyer, 1994;Lockley, Hunt, Moratalla, & Matsukawa, 1994;McCrea et al., 2014a,b;. Previously published examples usually show the lack of a digit or a different value of homologous interdigital divarication angles (Avanzini et al., 2008). ...
Article
Trackways can provide unique insight to animals locomotion through quantitative analysis of variation in track morphology. Long trackways additionally permit the study of trackmaker foot anatomy, providing more insight on limb kinematics. In this paper we have restudied the extensive tracksite at Barranco de La Canal-1 (Lower Cretaceous, La Rioja, NW Spain) focussing on a 25-m-long dinosaur (ornithopod) trackway that was noted by an earlier study (Casanovas et al., 1995; Pérez-Lorente, 2003) to display an irregular pace pattern. This asymmetric gait has been quantified and photogrammetric models undertaken for each track, thus revealing distinct differences between the right and the left tracks, particularly in the relative position of the lateral digits II–IV with respect to the central digit III. Given that the substrate at this site is homogenous, the consistent repetition of the collected morphological data suggests that differences recorded between the right and the left tracks can be linked to the foot anatomy, but more interestingly, to an injury or pathology on left digit II. We suggest that the abnormal condition registered in digit II impression of the left pes can be linked to the statistically significant limping behaviour of the trackmaker. Furthermore, the abnormal condition registered did not affect the dinosaur's speed.
... The recognition of gait abnormalities in dinosaur trackways has been reported in both isolated tracks (Ishigaki, 1986(Ishigaki, , 1989Mateus & Mil an, 2010) and trackways (Abel, 1935;Avanzini, Piñuela, & García Ramos, 2008;Dantas, Santos, Lockley, & Meyer, 1994;Lockley, Hunt, Moratalla, & Matsukawa, 1994;McCrea et al., 2014a,b;. Previously published examples usually show the lack of a digit or a different value of homologous interdigital divarication angles (Avanzini et al., 2008). ...
Article
Full-text available
Trackways can provide unique insight to animals locomotion through quantitative analysis of variation in track morphology. Long trackways additionally permit the study of trackmaker foot anatomy, providing more insight on limb kinematics. In this paper we have restudied the extensive tracksite at Barranco de La Canal-1 (Lower Cretaceous, La Rioja, NW Spain) focussing on a 25-m-long dinosaur (ornithopod) trackway that was noted by an earlier study (Casanovas et al., 1995; Pérez-Lorente, 2003) to display an irregular pace pattern. This asymmetric gait has been quantified and photogrammetric models undertaken for each track, thus revealing distinct differences between the right and the left tracks, particularly in the relative position of the lateral digits II–IV with respect to the central digit III. Given that the substrate at this site is homogenous, the consistent repetition of the collected morphological data suggests that differences recorded between the right and the left tracks can be linked to the foot anatomy, but more interestingly, to an injury or pathology on left digit II. We suggest that the abnormal condition registered in digit II impression of the left pes can be linked to the statistically significant limping behaviour of the trackmaker. Furthermore, the abnormal condition registered did not affect the dinosaur's speed.
... Another Upper Jurassic unit suitable for comparison with the Morrison Formation is the Kimmeridgian Vega Formation, which crops out in the coastal Asturias region of northern Spain. Although it lacks a prolifi c body-fossil record, abundant trace fossils reveal that the Asturias area supported a diverse terrestrial fauna in the Late Jurassic, including dinosaurs, pterosaurs, lizards, and turtles (Lockley et al., 2008;Mateus and Milán, 2010). Recent work by Gutierrez and Sheldon (2012) reconstructed a cool, semihumid to semiarid climate with MAP ranging from ~400 to 900 mm yr -1 , based on calculations using CIA -K and depth to Bk horizon. ...
Article
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Elemental analyses of paleosol B horizons in the Upper Jurassic Morrison Formation of the western United States provide estimates of mean annual precipitation (MAP) and allow determination of humidity regimes. Samples were collected from the lower Morrison Formation in New Mexico and the upper Morrison Formation in Wyoming and Montana. The chemical index of alteration minus potassium (CIA - K) and the calcium and magnesium weathering index (CALMAG) were used to estimate paleorainfall. CIA - K values calculated for paleosols without shrink-swell (vertic) features correspond to MAP estimates between 800 and 1100 mm yr(-1), with an average of 1000 mm yr(-1). CALMAG values, calculated for vertic paleosols, correspond to MAP estimates between 50 and 1200 mm yr(-1), with an average of 700 mm yr(-1). MAP estimates from the older New Mexico strata indicate that early Morrison environments were relatively arid. MAP estimates from the younger Wyoming and Montana deposits reflect wetter conditions in the northernmost part of the Morrison Formation, but the transition from arid interior environments was abrupt. Humidity provinces inferred from geochemical proxy-based estimates of evapotranspiration and energy influx from precipitation range from semiarid to superhumid, suggesting wetter conditions than the MAP estimates, but supporting the relative differences in moisture among the three study areas. Paleo precipitation patterns within the Morrison depositional basin do not match the modern latitudinal distribution of rainfall that arises from zonal atmospheric circulation. Comparison with the Upper Jurassic Lourinha Formation in Portugal and the Vega Formation in Spain reveals that MAP in Late Jurassic Portuguese environments was similar to that in the wet northern part of the Morrison Formation, although more arid conditions prevailed in some areas of Portugal. Inferred humidity regimes for the Lourinha Formation, which range from semiarid to superhumid, also indicate small-scale geographic variability in climate, although less pronounced than that observed in the Morrison Formation. Paleoenvironments in northern Spain were similar to the drier Morrison environments in the continental interior. Given the abrupt climatic transitions inferred here for the Morrison Formation, paleoprecipitation estimates derived from a geographically restricted sample may reflect only local conditions and should not necessarily be extrapolated to larger areas.
... Thus, less attention has had the infill of the tracks (casts and cross sections of the casts) usually preserved at the bottom of the layers, which in some cases have been misidentified and considered as non-biogenic sedimentary structures in the geological record (Loope, 1986;Nadon, 1993Nadon, , 2001Engelmann and Hasiotis, 1999). In the last decade, this has changed and several papers have described dinosaur tracks preserved as natural casts or in cross section (Difley and Ekdale, 2002;Currie et al., 2003;Romano and Whyte, 2003;Platt and Hasiotis, 2006;Castanera et al., 2010;Mateus and Milàn, 2010;Avanzini et al., 2012;Pascual et al., 2012) that generally are found isolated at the field, in reduced outcrops. Consequently, the outstanding of the described regional megatracksite comes from both the concentration and wide distribution of tracks on a kilometric-scale bed and their preservation as casts at the base of a tsunami deposit, consisting of a coarse-grained clastic sandstone complex. ...
Article
A thick multiple-bed tsunami deposits consisting of sandstones and conglomerates has been discovered and investigated in the Camarillas Formation (~ 130.6-128.4 Ma, Barremian age) in eastern Spain. The tsunami deposit is interbedded within red mudstones deposited in mud flats of a back-barrier system. It crops out along seven kilometres in length and at its base a great number of dinosaur tracks assigned to sauropods, ornithopods and theropods have been preserved as natural casts; then constituting an exceptional regional megatracksite associated with tsunami deposits. On the basis of sedimentological features and the lateral and vertical architecture of the involved lithofacies, up to five couplets of inflow-backflow deposits, formed by a tsunami wave train, have been recognized overlying the tracks. Although sedimentation mainly took place during backflow currents, inflows led to the removal of sand from a fronting barrier island and the rip-up of lagoonal carbonate and clay pebbles, depositing them in the protected back-barrier lagoon. Its unusually great thickness is interpreted, among others, as being the result of the filling of the previous low topography of the back-barrier lagoon.
... This way, climatic and perhaps geo− graphical constraints may have favoured interactions be− tween non−gregarious large theropods. Although many of the other tracksites displaying large theropod footprints were recorded in similar depositional environments worldwide, none shows a high density of large theropod trackways (e.g., Alonso and Marquillas 1986;Calvo 1991;Leonardi and Spezzamonte 1994;Lockley and Hunt 1994;Thulborn 2001;Moreno and Pino 2002;Moreno and Benton 2005;Mossman et al. 2003;Mateus and Milàn 2010). Interestingly, climatic conditions corresponding to these other tracksites range from subtropical to temperate, hence not as arid as the conditions at Querulpa Chico and Chacarilla. ...
Article
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In modern terrestrial ecosystems, the population size of large predators is low, and a similar pattern has usually been assumed for dinosaurs. However, fossil finds of monospecific, large theropod accumulations suggest that population dynamics were more complex. Here, we report two Early Cretaceous tracksites dominated by large theropod footprints, in Querulpa Chico (Peru) and Chacarilla (Chile). The two sites correspond to distinct depositional environments-tidal basin/delta (Querulpa Chico) and meandering river (Chacarilla)-with both subject to extensive arid or semiarid palaeoclimatic conditions. Although most trackways show no preferred orientation, a clear relationship between two trackmakers is observed in one instance. This observation, coupled with the high abundance of trackways belonging to distinct large theropods, and the exclusion of tracks of other animals, suggests some degree of grouping behaviour. The presence of freshwater sources in a dry climate and perhaps social behaviour such as pair bonding may have promoted interactions between large carnivores. Further, the occurrence of these two tracksites confirms that large theropod dinosaurs, possibly spinosaurids and/or carcharodontosaurids, existed on the western margin of Gondwana as early as the earliest Cretaceous.
... The Vale de Meios trackmakers are large theropods or megatheropods as their estimated hip heights overpass the threshold (250 cm) proposed by some authors 33,43 and the footprint length exceed 45 cm 20,43,44 . These theropod tracks are among the largest theropod tracks described worldwide 30,31,45,46 . Nevertheless, other very large tracks are known. ...
Article
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Abstract A new dinosaur tracksite in the Vale de Meios quarry (Serra de Aire Formation, Bathonian, Portugal)preserves more than 700 theropod tracks. They are organized in at least 80 unidirectional trackways arranged in a bimodal orientation pattern (W/NW and E/SE). Quantitative and qualitative comparisons reveal that the large tridactyl, elongated and asymmetric tracks resemble the typical Late Jurassic-Early Cretaceous Megalosauripus ichnogenus in all morphometric parameters. Few of the numerous tracks are preserved as elite tracks while the rest are preserved as different gradients of modified true tracks according to water content, erosive factors, radial fractures and internal overtrack formations. Taphonomical determinations are consistent with paleoenvironmental observations that indicate an inter-tidal flat located at the margin of a coastal barrier. The Megalosauripus tracks represent the oldest occurrence of this ichnotaxon and are attributed to large megalosaurid dinosaurs. Their occurrence in Vale de Meios tidal flat represents the unique paleoethological evidence of megalosaurids moving towards the lagoon, most likley during the low tide periods with feeding purposes.
... Los rastros de saurópodos conservados como contralmoldes (hiporrelieves convexos) son raros en el registro fósil (Meyer et al., 1994), encontrándose con relativa frecuencia icnitas aisladas con esta conservación (Milàn et al., 2005;Platt & Hasiotis, 2006;Mateus & Milàn, 2010). El objetivo principal de este trabajo es describir por primera vez el rastro del yacimiento José María Herrero (JMH) constituido por contramoldes de saurópodos. ...
Conference Paper
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The record of Jurassic chondrichthyans remains from the Iberian Peninsula is reviewed. This record is very poor on the entire peninsula, with the exception of the Guimarota site (Portugal), with several taxa described and properly illustrated. The present paper is the first bibliographical list of publications dealing with the sharks record from the Jurassic of the Iberian Peninsula.
... Los rastros de saurópodos conservados como contralmoldes (hiporrelieves convexos) son raros en el registro fósil (Meyer et al., 1994), encontrándose con relativa frecuencia icnitas aisladas con esta conservación (Milàn et al., 2005;Platt & Hasiotis, 2006;Mateus & Milàn, 2010). El objetivo principal de este trabajo es describir por primera vez el rastro del yacimiento José María Herrero (JMH) constituido por contramoldes de saurópodos. ...
Conference Paper
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The chondrichthyan faunas from the Late Jurassic of Asturias is reviewed in this work. All the material in this study (teeth, finspines and dorsal spines), currently under preparation, are part of the Museo del Jurásico de Asturias (Colunga) collections. This record is relatively scarce, and consists exclusively of hybodontiform sharks: Hybodontiformes indet., Asteracanthus, Hybodus, and Planohybodus. All specimens come from Tereñes and Lastres Formations (Kimmeridgian).
... The vertebrate faunal assemblage of the Amoreira-Porto Novo Member also includes the sauropod Dinheirosaurus lourinhanensis, the theropods Lourinhanosaurus antunesi, Torvosaurus gurneyi, and Ceratosaurus sp., as well as the ornithischians Miragaia longicollum, Hypsilophodon sp., 'Trimucrodon cuneatus,' and indeterminate remains attributable to basal Iguanodontia (Bonaparte and Mateus, 1999;Antunes and Mateus, 2003;Mateus, 2006;Mateus et al., 2006Mateus et al., , 2009. This geologic member has also produced dinosaur eggs and embryo remains (Mateus et al., 1998;Ricqlès et al., 2001), as well as several dinosaur tracks (Milàn et al., 2005;Mateus and Milàn, 2010). ...
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Here we describe a new partial sauropod skeleton from the late Kimmeridgian (Late Jurassic) of the Lourinhã Formation, central west Portugal. The closely associated specimen comprises a complete tooth (with root), a fragment of cervical neural arch, an anterior chevron, and an almost complete right pectoral girdle and forelimb. The new sauropod, Zby atlanticus, n. gen. et sp., can be diagnosed on the basis of four autapomorphies, including a prominent posteriorly projecting ridge on the humerus at the level of the deltopectoral crest. Nearly all anatomical features indicate that Zby is a non-neosauropod eusauropod. On the basis of several characters, including tooth morphology, extreme anteroposterior compression of the proximal end of the radius, and strong beveling of the lateral half of the distal end of the radius, Zby appears to be closely related to Turiasaurus riodevensis from approximately contemporaneous deposits in eastern Spain. However, these two genera can be distinguished from each other by a number of features pertaining to the forelimb. Whereas previously described Late Jurassic Portuguese sauropods show close relationships with taxa from the contemporaneous Morrison Formation of North America, it appears that turiasaurians were restricted to Europe. All adult sauropods recovered in the Late Jurassic of Portugal thus far are very large individuals: it is possible that the apparent absence of small- or medium-sized adult sauropods might be related to the occupation of lower-browsing niches by non-sauropods such as the long-necked stegosaur Miragaia longicollum.
... Paronychodon sp.; Zinke & Rauhut 1994;Zinke 1998;Mateus 2005). Moreover, theropod embryos and hatchlings, ascribed to Lourinhanosaurus (de Ricqlès et al. 2001;Mateus et al. 1998;, Allosaurus (Rauhut & Fechner 2005) and Torvosaurus (Araújo et al. 2013), were also collected in Portugal, and a diverse ichnological record is known (Mateus & Milàn 2010). ...
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Theropod dinosaurs form a highly diversified clade, and their teeth are some of the most common components of the Mesozoic dinosaur fossil record. This is the case in the Lourinhã Formation (Late Jurassic, Kimmeridgian-Tithonian) of Portugal, where theropod teeth are particularly abundant and diverse. Four isolated theropod teeth are here described and identified based on morphometric and anatomical data. They are included in a cladistic analysis performed on a data matrix of 141 dentition-based characters coded in 60 taxa, as well as a supermatrix combining our dataset with six recent datamatrices based on the whole theropod skeleton. The consensus tree resulting from the dentition-based data matrix reveals that theropod teeth provide reliable data for identification at approximately family level. Therefore, phylogenetic methods will help identifying theropod teeth with more confidence in the future. Although dental characters do not reliably indicate relationships among higher clades of theropods, they demonstrate interesting patterns of homoplasy suggesting dietary convergence in (1) alvarezsauroids, therizinosaurs and troodontids; (2) coelophysoids and spinosaurids; (3) compsognathids and dromaeosaurids; and (4) ceratosaurids, allosauroids and megalosaurids. Based on morphometric and cladistic analyses, the biggest tooth from Lourinhã is referred to a mesial crown of the megalosaurid Torvosaurus tanneri, due to the elliptical cross section of the crown base, the large size and elongation of the crown, medially positioned mesial and distal carinae, and the coarse denticles. The smallest tooth is identified as Richardoestesia, and as a close relative of R. gilmorei based on the weak constriction between crown and root, the "eight-shaped" outline of the base crown and, on the distal carina, the average of ten symmetrically rounded denticles per mm, as well as a subequal number of denticles basally and at mid-crown. Finally, the two medium-sized teeth belong to the same taxon and exhibit pronounced interdenticular sulci between distal denticles, hooked distal denticles for one of them, an irregular enamel texture, and a straight distal margin, a combination of features only observed in abelisaurids. They provide the first record of Abelisauridae in the Jurassic of Laurasia and one of the oldest records of this clade in the world, suggesting a possible radiation of Abelisauridae in Europe well before the Upper Cretaceous.
... have been described while the skeletal record is poor (Hernández Medrano et al., 2008;Moratalla and Hernàn, 2010;Pérez-Lorente, 2015). By contrast, the Mirambel Formation is one of those that preserves both kind of fossils (skeletal and ichnological remains), and thus the information taken from one can complement the other (e.g., Mateus and Milàn, 2010). In this regard, a combination of both sources of data can represent the census for a paleoecological reconstruction of the unit, thus indicating whether the ichnological record is consistent or not with the skeletal record (Lockley, 1991). ...
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Up to now, the ichnological vertebrate record from the Barremian Mirambel Formation (NE Spain) has remained completely unknown despite the fact that osteological findings have been reported in recent years. Here we provide an overview of 11 new dinosaur tracksites found during a fieldwork campaign in the year 2011. The majority of these tracksites (seven) preserve small- to medium-sized tridactyl tracks here assigned to indeterminate theropods. Only one footprint presents enough characters to classify it as Megalosauripus isp. Ornithopod tracks identified as Caririchnium isp. and Iguanodontipodidae indet. and sauropod tracks are recorded at two tracksites. The footprints are preserved in a variety of paleoenvironmental conditions and thus display different kinds of preservation (true tracks, shallow undertracks, natural casts and undertrack casts). The ichnological record from the Mirambel Formation seems to be theropod dominated. This is a clear discrepancy with the osteological record identified in this formation, which shows a predominance of ornithopod dinosaurs.
... L = track length; LI = length of digit I; LII = length of digit II; LIII = length of digit III; LIV = length of digit IV; LV = length of digit V; I-II = angle between digits I and II; II-III = angle between digits II and III; III-IV = angle between digits III and IV; IV-V = angle between digits IV and V; I-V = angle between digits I and V; W = track width. L W L/W LI LII LIII LIV LV I-II II-III III-IV IV- palmar impression (i.e., Mateus and Milàn, 2010) or the striation in natural infill casts resulting of the skin dragging in the mud (Czerkas, 1994;Milàn et al., 2005;Foster and Hunt-Foster, 2011). The near-absence of preserved skin impressions in non-avian dinosaurs associated to the conservative anatomy of little variation has been the main reasons for neglecting skin for taxonomical identification within Dinosauria. ...
... Stratigraphic section of the Guxian tracksite, Shanxi Province, China. Belvedere and Mietto, 2010), Parabrontopodus and Brontopodus from Jura, France(Loeuff et al., 2006;Mazin et al., 2016), Parabrontopodus from Istria, Croatia(Mezga et al., 2007), narrow-gauge trackways from Cameros Basin in Spain(Moratalla, 2009), wide-and narrow-gauge trackways from central-west Portugal(Mateus and Milàn, 2010), first giant sauropod footprints similar to Brachiosaurus from Zimbabwe ...
Article
This paper presents the first report of sauropod tracks from the Upper Jurassic of Shanxi Province, China. Dinosaur tracks appear concentrated in five trackways, in different stratigraphic levels of the Late Jurassic Tianchihe Formation. Tracks are dominantly small and medium-size sauropod tracks and are tentatively assigned to Brontopodus based on preserved track morphology, trackway pattern and statistical analysis. The Tianchihe Formation in which the tracks appear shows a gradual change from meandering fluvial to sandy braided fluvial depositional systems developed in a seasonally arid environment. Comparisons of the evaluated speed of bipedal to quadruped trackways indicate that the slower walk more easily produces pes-dominated overprints. Trackways in the Guxian tracksite appear following different orientations, suggesting that these trackways were produced by different sauropods at different times. An unusual trackway following a curved pattern has been identified in the site and could represent a special locomotion character or a social behavior. The presence of eolian deposits in central Shanxi Province could have acted as a paleogeographic and paleoenvironmental barrier for the dispersion of the Yanliao Biota that survived in northern Hebei-western Liaoning and northestern Shanxi Province to the Ordos Basin during the Late Jurassic.
... Small to medium-sized tridactyl tracks assigned to ornithopods are scarce in the Late Jurassic European record compared with other types of tridactyl dinosaur footprints such as those of theropods, which are quite abundant in areas such as the Jura platform Mazin et al. 2017;Castanera et al. 2018), the Lusitanian Basin (Lockley and Santos 1993;Lockley et al. 1994;Mateus and Milàn 2010) or the Asturian Basin (Lockley et al. 2008;Piñuela Suarez 2015). This scarcity of ornithopod tracks is a consequence of an abundance of sauropod-and theropod-dominated ichnofaunas preserved in carbonate facies in the socalled Brontopodus ichnofacies, which are characterized by a rather low ichnodiversity (Hunt and Lucas 2007). ...
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The Sociedade de História Natural in Torres Vedras, Portugal houses an extensive collection of as yet undescribed dinosaur tracks with ornithopod affinities. They have been collected from different Late Jurassic (Kimmeridgian–Tithonian) geological formations (Praia de Amoreira-Porto Novo, Alcobaça, Sobral, and Freixial) that outcrop along the Portuguese coast, and belong to two different sub-basins of the Lusitanian Basin (the Consolação and Turcifal sub-basins). Three main morphotypes can be distinguished on the basis of size, mesaxony and the morphology of the metatarsophalangeal pad impression. The minute to small-sized morphotype is similar to the Anomoepus-like tracks identified in other Late Jurassic areas. The small to medium-sized morphotype resembles the Late Jurassic–Early Cretaceous ichnotaxon Dinehichnus, already known in the Lusitanian Basin. Interestingly, these two morphotypes can be distinguished qualitatively (slightly different size, metatarsophalangeal pad impression and digit morphology) but are nevertheless difficult to discriminate by quantitatively analysing their length-width ratio and mesaxony. The third morphotype is considered a large ornithopod footprint belonging to the ichnofamily Iguanodontipodidae. This ichnofamily is typical for Cretaceous tracksites but the new material suggests that it might also be present in the Late Jurassic. The three morphotypes show a negative correlation between size and mesaxony, so the smaller tracks show the stronger mesaxony, and the larger ones weaker mesaxony. The Upper Jurassic ornithopod record from the Lusitanian Basin has yielded both small and medium-sized ornithopod remains, mainly iguanodontians such as dryosaurids and ankylopollexians, which are the main candidates to be the trackmakers.
... Accordingly, the palaeoenvironmental information from one type can complement that from another (e.g. Mateus and Milàn, 2010). The unit crops out in the Ladruñán anticline (Teruel province), providing a remarkable number of vertebrate fossil localities bearing bones, ichnites and/or eggshells in a limited area. ...
... The deltoid shape with a thick tetradactyl pes and thick pentadactyl crescent shape of manus imprints of quadrupedal trackways has been reported from the Early Cretaceous of south England and attributed to nodosaurids (Wright et al., 1998). Although Deltapodus of the Tirgan Formation is much smaller than the holotype of Deltapodus from the Middle Jurassic of Yorkshire of England (Whyte and Romano, 1994;Whyte et al., 2007) or elsewhere Milàn and Chiappe, 2009;Mateus and Milàn, 2010;Belvedere and Mietto, 2010), they have, however, a configuration and digit number similar to Deltapodus and attributable to stegosaurs. Among abundant and diverse dinosaur footprints of the Late Jurassic Morrison Formation (Marsh, 1899;Lockley et al., 1986;Foster and Lockley, 2006), relatively small Deltapodus has been considered as a probable ankylosaurian pedal imprint 15 cm long and 12 cm wide (Gierlinski et al., 2010), and Kimmeridgian Deltapodus of Morocco have 16.6 cm and 25.7 cm length and 105 cm and 17.4 cm width (Belvedere and Mietto, 2010). ...
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Numerous dinosaur footprints are reported from the Cretaceous of Iran for the first time. These footprints are small and they have been discovered in the one of uppermost layers of the Tirgan Formation (Neocomian-Aptian), Gelian section, south Shirvan, Northeast Iran. The Tirgan Formation comprises Orbitolina-bearing bioclastic and oolitic limestones (200 m), which were deposited in the tidal flat to shallow marine environments of a carbonate ramp. Footprints were impressed in the subaerial conditions of an intertidal environment. Based on microfossils and stratigraphic position, they are Aptian in age. Most footprints are weathered and remained as indistinct holes, but five distinctive trackways are distinguishable. One of the trackways belongs to small sauropod with a narrow gauge. The means of pes footprint length and footprint width are 27.8 cm and 37.7 cm, respectively. Means of footprint length and footprint width of the manus in this trackway are 14.1 cm and 19.3 cm. Well preserved plantigrade stegosaur footprints are circular to ellipsoidal in outline and include tridactyl pes and tetradactyl manus imprints. The means of footprint length and footprint width equal 15.5 cm and 12.8 cm in the pes and 6.4 cm and 10.3 in the manus. These stegosaur footprints are assigned to Deltapodus isp. Abundant, small, quadrupedal ornithopod footprints (footprint length of pes and manus equals 13.0 cm and 9.7 cm; footprint width of pes and manus equals 18.4 and 15.0 cm) are attributable to Caririchnium and Ornithopodichnus, though they have a lower footprint length/footprint width ratio than the well-known ichnospecies of these ichnotaxa. It seems that dwarfism of the Tirgan Formation's dinosaur track-makers was due to the position of the Iranian microplate in northern part of Tethys, which is explainable by the island rule.
... Stegosaur tracks, firstly identified in 1990s (White & Romano, 1994Lockley & Hunt, 1998) have since become relatively common in the Lourinhã Formation (Mateus & Milàn, 2010;Mateus et al., 2011), with the first report of the ichnogenus Deltapodus Whyte & Romano, 1994in 2008 (Mateus & Milàn, 2008). However, impressions of skin scales had not been investigated in detail yet, and only recently had the presence of a hoof (Herrero Gascón & Pérez-Lorente, 2016), despite its suggestion in the morphology of Deltapodus (Whyte & Romano, 2001). ...
... The final print morphotype in Group A (Av) was assigned to a new ichnotaxon, Deltapodus brodricki [61], after the distinctive 'delta-shaped' pes imprint and the name of the person who first described dinosaur prints from the area. This print type has now been recognized across the Jurassic world and has been recorded from Spain, Portugal, Morocco and Utah (USA) [74], and most recently in the Cretaceous of China [75]. Group B contained 17 tridactyl prints that had been made by bipeds ( Figure 4). ...
Article
Despite the Hebrides and Cleveland basins being geographically close, research has not previously been carried out to determine faunal similarities and assess the possibility of links between the dinosaur populations. The palaeogeography of both areas during the Middle Jurassic shows that there were no elevated landmasses being eroded to produce conglomeratic material in the basins at that time. The low-lying landscape and connected shorelines may have provided connectivity between the two dinosaur populations. The dinosaur fauna of the Hebrides and Cleveland basins has been assessed based primarily on the abundant ichnites found in both areas as well as their skeletal remains. In the two basins, the dinosaur faunas are very similar, consisting of non-neosauropod eusauropods, a possible basal titanosauriform, large and small theropods and ornithopods and europodan thyreophorans. The main difference in the faunas is in the sizes. In the Cleveland Basin, the ichnites suggest that there were medium and large theropods alongside small to medium sized ornithopods, whereas, in the Hebrides Basin, the theropods were from small to large and the ornithopods were medium to large. It is suggested that migrations could have taken place between the two areas during the Middle Jurassic. A tentative food chain from the herbivorous dinosaurs to the top predators can be inferred from the footprints.
... Lourinhã is a city (and municipality) in the District of Lisbon in the Oeste subregion of Portugal with a population slightly exceeding 9 thousand citizens (https://censos.ine.pt). Geologically Lourinhã is Upper Jurassic Formation exposing in several localities in the Lusitanian Basin (central-west Portugal) (Mateus and Milàn, 2010) in close proximity to the Lourinhã city. Within the Formation, consisting of mostly alluvial sediments, deposited during the early rifting of the Atlantic Ocean (Mateus and Milan, 2010) skeletal elements of e.g. ...
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Fossils of dinosaurs and other tetrapods have long aroused interest of scientists and the public opinion alike. Every finding of a new (especially large) species receives coverage in national and international media, and thus, local fossil discoveries might constitute a good basis for local tourism development. The paper aims to examine whether fos-siliferous sites on their own may be enough for the development of palaeontological tourism to occur, or do they require the support of additional amusement infrastructure. For this purpose, the interest in chosen localities was analysed using Google and Wikipedia searches, and was further discussed against a survey on dinoparks and their elements. The above-mentioned data reveal that local tourism can be indeed predicated on local paleontological findings, however, it is deemed considerably more efficient if such attractions are backed with an extensive infrastructure of amusement theme parks.
... The first was a cast reported from the Tidwell Member , below the Salt Wash Member in Garfield County, Utah, the second was a cast reported from and Brushy Basin Member above the Saltwash Member, in San Juan County (Milan and Chiappe, 2009) and the third, an impression from near the Moab Giants site, is a small, isolated track (only 15 cm long) originally described as a "small Deltapodus print of possible ankylosaurian affinity" (Gierlinski et al., 2010). Given the increased number of reports of well-preserved Deltapodus in trackways (notably Cobos et al., 2010;Mateus and Milàn, 2010;Mateus et al., 2011;Xing et al., 2013), interpreted as stegosaurian, the interpretation of this small, isolated impression of Deltapodus as ankylosaurian is open to question. ...
Article
Recent construction at the Moab Giants dinosaur museum property ~ 10 miles (~ 16 km) north of Moab, in Grand County, Utah in 2014–2015 revealed a number of moderately well preserved dinosaur tracks from the Upper Jurassic, Salt Wash Member of the Morrison Formation. The best specimens were preserved as natural casts on the underside of massive sandstone beds, many containing dense assemblages of invertebrate traces. Here we describe several tridactyl theropod tracks, an ornithischian manus attributable to ichnogenus Stegopodus and an ornithischian pes track attributed to ichnogenus Deltapodus. Variably preserved Deltapodus have now been reported from the Tidwell, Salt Wash and Brushy Basin members of the Morrison Formation from Garfield, Grand and San Juan counties respectively, suggesting a wide distribution in space and time. The Salt Wash specimen is considered to be the most representative of Deltapodus morphologies reported from large samples in other regions, notably in Spain, Portugal western China and North Africa. Differences between Stegopodus and Deltapodus are reviewed. Deltapodus is reported from Europe and Asia as well as North America, in the Middle Jurassic through Late Cretaceous, and is often represented by abundant trackways. Stegopodus is presently reported from the Jurassic where it occurs in the Late Jurassic of North America as isolated tracks, and, with some debate, as trackways and isolated tracks from Europe. A middle Jurassic occurrence from Morocco is also reported.
... The final print morphotype in Group A (Av) was assigned to a new ichnotaxon, Deltapodus brodricki [61], after the distinctive 'delta-shaped' pes imprint and the name of the person who first described dinosaur prints from the area. This print type has now been recognized across the Jurassic world and has been recorded from Spain, Portugal, Morocco and Utah (USA) [74], and most recently in the Cretaceous of China [75]. Group B contained 17 tridactyl prints that had been made by bipeds ( Figure 4). ...
Preprint
Despite the Hebrides and Cleveland basins being geographically close, research has not previously been carried out to determine faunal similarities and assess the possibility of links between the dinosaur populations. The palaeogeography of both areas during the Middle Jurassic shows that there were no elevated landmasses being eroded to produce conglomeratic material in the basins at that time. The low-lying landscape and connected shorelines may have provided connectivity between the two dinosaur populations. The dinosaur fauna of the Hebrides and Cleveland basins has been assessed based primarily on the abundant ichnites found in both areas as well as their skeletal remains. In the two basins the dinosaur faunas are very similar, consisting of non-neosauropod eusauropods, a possible basal titanosauriform, large and small theropods and ornithopods and europodan thyreophorans. The main difference in the faunas is in the sizes. In the Cleveland Basin the ichnites suggest that there were medium and large theropods alongside small to medium sized ornithopods whereas in the Hebrides Basin the theropods were from small to large and the ornithopods were medium to large. It is suggested that migrations could have taken place between the two areas during the Middle Jurassic. A tentative food chain from the herbivorous dinosaurs to the top predators can be inferred from the footprints.
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The Kimmeridgian Vega, Tereñes and Lastres formations of Asturias have yielded a rich vertebrate fauna, represented by both abundant tracks and osteological remains. However, skeletal remains of theropod dinosaurs are rare, and the diversity of theropod tracks has only partially been documented in the literature. Here we describe the only non-dental osteological theropod remain recovered so far, an isolated anterior caudal vertebra, as well as the largest theropod tracks found. The caudal vertebra can be shown to represent a megalosaurine megalosaurid and represents the largest theropod skeletal remain described from Europe so far. The tracks are also amongst the largest theropod footprints reported from any setting and can be assigned to two different morphotypes, one being characterized by its robustness and a weak mesaxony, and the other characterized by a strong mesaxony, representing a more gracile trackmaker. We discuss the recently proposed distinction between robust and gracile large to giant theropod tracks and their possible trackmakers during the Late Jurassic-Berriasian. In the absence of complete pedal skeletons of most basal tetanurans, the identity of the maker of Jurassic giant theropod tracks is difficult to establish. However, the notable robustness of megalosaurine megalosaurids fits well with the described robust morphotypes, whereas more slender large theropod tracks might have been made by a variety of basal tetanurans, including allosaurids, metriocanthosaurids or afrovenatorine megalosaurids, or even exceptionally large ceratosaurs. Concerning osteological remains of large theropods from the Late Jurassic of Europe, megalosaurids seem to be more abundant than previously recognized and occur in basically all Jurassic deposits where theropod remains have been found, whereas allosauroids seem to be represented by allosaurids in Western Europe and metriacanthosaurids in more eastern areas. Short-term fluctuations in sea level might have allowed exchange of large theropods between the islands that constituted Europe during the Late Jurassic.
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The stegosaur species Miragaia longicollum was erected based on a partial anterior skeleton from the Upper Jurassic of Portugal. Until then, almost all stegosaur specimens in Portugal and Spain had been identified as Dacentrurus armatus, the sister taxon of M. longicollum and only other member of the clade Dacentrurinae. The holotypes of the two species have little overlap, since the holotype of D. armatus is mostly a posterior skeleton, so the classification of other specimens to either species is unclear and the validity of M. longicollum has been questioned and debated. Here we describe a largely complete specimen of M. longicollum discovered in 1959 in Atouguia da Baleia, Peniche, Portugal, consisting of both anterior and posterior portions of the skeleton. Comparisons to the holotypes of dacentrurines and other stegosaurs shed light on the convoluted relationships of this group. We conclude that M. longicollum is valid and rather different from D. armatus, and provide a revised diagnosis of M. longicollum, as well as revised diagnoses for D. armatus, Dacentrurinae, and the first diagnosis of the genus Miragaia, granting stability to these taxa and allowing new considerations to be given on the classification of other Iberian stegosaurs. This new specimen is, to date, the most complete dinosaur described from Portugal and the most complete stegosaur described from Europe. Miragaia shared anatomical features that show a close affinity to Alcovasaurus longispinus, confirming this to be the first known dacentrurine stegosaur in America, coherent with the hypothesis of an ephemeral land bridge between North America and Iberia that allowed faunal exchange.
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The Upper Jurassic of Portugal has been globally known for its microfossil vertebrate fauna thanks to the Konzentrat-Lagerstätte of the Guimarota mine, which provided thousands of bone fragments, isolated teeth, and even complete specimens. Other vertebrate microfossil assemblages have been studied around the world. Besides Guimarota, no other Portuguese Jurassic assemblage has been extensively studied. Hereby is presented a revision of the state of the art on Portuguese microvertebrate record, and the first microvertebrate studies on three localities from the Lourinhã Formation (Late Jurassic) hosted by a Portuguese institution; Porto das Barcas, Zimbral, and Valmitão has provided 2,497 microvertebrates skeletal remains and teeth, from which 824 specimens have been identified, described and assessed to the conservative-most taxa. The stratigraphy and sedimentology of the localities suggest that Porto das Barcas and Zimbral were floodplain mud deposits, and Valmitão was an oxbow lake mud deposit, with a slow rate of sedimentation. The remains have been attributed to fishes, amphibians, squamates, crocodylomorphs, and dinosaurs; but unfortunately, no mammaliaform material has been collected. Paleoecological analyses suggest Zimbral and Valmitão were dominated by a terrestrial fauna and more diverse than Porto das Barcas, dominated by an amphibious fauna. The Lourinhã Formation appears to have been closer to the shoreline than American localities in the Morrison and Cloverly Formations were, but more continental than Buenache and Las Hoyas localities (Spain) with swamp to lacustrine paleoenvironments. A detailed study on 125 crocodylomorph teeth from Valmitão support the presence of Goniopholididae, at least two Atoposauridae taxa, and Bernissartiidae in the Late Jurassic of Portugal, with a fauna either dominated by relative small individuals, either juveniles or adults or small taxa.
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At Cabo Mondego (western central Portugal), the Upper Jurassic marine to coastal succession contains several stratigraphic levels preserving dinosaur footprints on the surface bedding plane, as well as convolute bedding and soft sediment injection structures interpreted as dinoturbation structures. At least nineteen new three-dimensional structures observed in cross-sections are interpreted as produced by dinosaur trampling. The identification of three-dimensional structures of dinosaur footprints provides an important complement to the information obtained from footprints preserved on single bedding surfaces, such as the substrate consistency, potential trackmaker identification, and the possibility to enhance the distinction of sauropods and tridactyl dinosaurs, and paleoenvironmental interpretations. In the lower part of the Arenitos da Boa Viagem Formation, eight levels of probable lowermost Kimmeridgian age (ca. 157–156 Ma), displaying the above-mentioned deformational structures, were analyzed in detail. They support interpretations concerning the relationship between the footprints and the substrate consistency at the time of their formation. Three distinct cohesiveness patterns, defined by the penetration of the feet from the paleosurface, are the result of different degrees of substrate cohesiveness. Identifying the trackmakers of levels belonging to the middle Oxfordian–lower Kimmeridgian has important implications for Late Jurassic ecosystem reconstructions, as the footprints observed in Cabo Mondego indicate a change in the morphotypes throughout the Upper Jurassic succession.
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The occurrence and features of skin impressions in a sauropod footprint, the largest (>50 cm in diameter) reported to date for this taxon, from the Lower Cretaceous Haman Formation (Albian) in Korea are described, and its preservation and paleoenvironmental implications are interpreted. The skin impression-bearing deposits are floodplain sediments formed by sheetflood processes. The large impression is preserved in silty mudstone with microbial lenses and wisps overlying a planar- to cross-laminated and fine-grained sandstone to siltstone bed. The paleoenvironment of the skin impression-bearing deposits is interpreted as a saline sandflat to mudflat where microbial mats can form around lakes or ponds under semi-arid paleoclimatic conditions with alternating wetting and drying intervals. These paleoenvironmental conditions would have permitted the distinct preservation of skin impressions in a dinosaur footprint. The observations here suggest that some sauropod dinosaurs in the Cretaceous had a well-developed polygonal skin texture covering nearly the whole of their foot pads, as seen in modern elephants, which would increase stability when walking on muddy and wet ground.
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A recently discovered Early Cretaceous (early late Albian) dinosaur tracksite at Parede beach (Cascais, Portugal) reveals evidence of dinoturbation and at least two sauropod trackways. One of these trackways can be classified as narrow-gauge, which represents unique evidence in the Albian of the Iberian Peninsula and provides for the improvement of knowledge of this kind of trackway and its probable trackmaker, in an age when the sauropod record is scarce. These dinosaur tracks are preserved on the upper surface of a marly limestone bed that belongs to the Galé Formation (Água Doce Member, middle to lower upper Albian). The study of thin-sections of the beds C22/24 and C26 in the Parede section has revealed a microfacies composed of foraminifers, radiolarians, ostracods, corals, bivalves, gastropods, and echinoids in a mainly wackestone texture with biomicritic matrix. These assemblages match with the lithofacies, marine molluscs, echinids, and ichnofossils sampled from the section and indicate a shallow marine, inner shelf palaeoenvironment with a shallowing-upward trend. The biofacies and the sequence analysis are compatible with the early late Albian age attributed to the tracksite. These tracks and the moderate dinoturbation index indicate sauropod activity in this palaeoenvironment. Titanosaurs can be dismissed as possible trackmakers on the basis of the narrow-gauge trackway, and probably by the kidney-shaped manus morphology and the pes-dominated configuration of the trackway. Narrow-gauge sauropod trackways have been positively associated with coastal palaeoenvironments, and the Parede tracksite supports this interpretation. In addition, this tracksite adds new data about the presence of sauropod pes-dominated trackways in cohesive substrates. As the Portuguese Cretaceous sauropod osteological remains are very scarce, the Parede tracksite yields new and relevant evidence of these dinosaurs. Furthermore, the Parede tracksite is the youngest evidence of sauropods in the Portuguese record and some of the rare evidence of sauropods in Europe during the Albian. This discovery enhances the palaeobiological data for the Early Cretaceous Sauropoda of the Iberian Peninsula, where the osteological remains of these dinosaurs are relatively scarce in this region of southwestern Europe. Therefore, this occurrence is also of overall interest due to its impact on Cretaceous Sauropoda palaeobiogeography.
Article
Ichnological evidence from the Ravenscar Group (Middle Jurassic) of Yorkshire reveals a distinctive type of swimming track associated with walking tracks of the ichnogenus Deltapodus. The morphology of the prints suggests that both track types were made by the same type of animal. Since Deltapodus is interpreted as having been made by a stegosaur, the associated footprints suggest that stegosaurs could swim. This has significant implications for their palaeobiology and dispersal. The swimming prints are assigned to the ichnotaxon Characichnos isp. © 2015 The Author(s). Published by The Geological Society of London for the Yorkshire Geological Society. All rights reserved.
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Extensive and well-preserved tracksites in the coastally exposed Lower Cretaceous (Valanginian–Barremian) Broome Sandstone of the Dampier Peninsula provide almost the entire fossil record of dinosaurs from the western half of the Australian continent. Tracks near the town of Broome were described in the late 1960s as Megalosauropus broomensis and attributed to a medium-sized theropod trackmaker. Brief reports in the early 1990s suggested the occurrence of at least another nine types of tracks, referable to theropod, sauropod, ornithopod, and thyreophoran trackmakers, at scattered tracksites spread over more than 80 km of coastline north of Broome, potentially representing one of the world's most diverse dinosaurian ichnofaunas. More recently, it has been proposed that this number could be as high as 16 and that the sites are spread over more than 200 km. However, the only substantial research that has been published on these more recent discoveries is a preliminary study of the sauropod tracks and an account of the ways in which the heavy passage of sauropod trackmakers may have shaped the Dampier Peninsula's Early Cretaceous landscape. With the other types of dinosaurian tracks in the Broome Sandstone remaining undescribed, and the full extent and nature of the Dampier Peninsula's dinosaurian tracksites yet to be adequately addressed, the overall scientific significance of the ichnofauna has remained enigmatic. At the request of the area's Goolarabooloo Traditional Custodians, 400+ hours of ichnological survey work was undertaken from 2011 to 2016 on the 25 km stretch of coastline in the Yanijarri–Lurujarri section of the Dampier Peninsula, inclusive of the coastline at Walmadany (James Price Point). Forty-eight discrete dinosaurian tracksites were identified in this area, and thousands of tracks were examined and measured in situ and using three-dimensional photogrammetry. Tracksites were concentrated in three main areas along the coast: Yanijarri in the north, Walmadany in the middle, and Kardilakan–Jajal Buru in the south. Lithofacies analysis revealed 16 repeated facies types that occurred in three distinctive lithofacies associations, indicative of an environmental transgression between the distal fluvial to deltaic portions of a large braid plain, with migrating sand bodies and periodic sheet floods. The main dinosaurian track-bearing horizons seem to have been generated between periodic sheet floods that blanketed the preexisting sand bodies within the braid plain portion of a tidally influenced delta, with much of the original, gently undulating topography now preserved over large expanses of the present day intertidal reef system. Of the tracks examined, 150 could be identified and are assignable to a least eleven and possibly as many as 21 different track types: five different types of theropod tracks, at least six types of sauropod tracks, four types of ornithopod tracks, and six types of thyreophoran tracks. Eleven of these track types can formally be assigned or compared to existing or new ichnotaxa, whereas the remaining ten represent morphotypes that, although distinct, are currently too poorly represented to confidently assign to existing or new ichnotaxa. Among the ichnotaxa that we have recognized, only two (Megalosauropus broomensis and Wintonopus latomorum) belong to existing ichnotaxa, and two compare to existing ichnotaxa but display a suite of morphological features suggesting that they may be distinct in their own right and are therefore placed in open nomenclature. Six of the ichnotaxa that we have identified are new: one theropod ichnotaxon, Yangtzepus clarkei, ichnosp. nov.; one sauropod ichnotaxon, Oobardjidama foulkesi, ichnogen. et ichnosp. nov.; two ornithopod ichnotaxa, Wintonopus middletonae, ichnosp. nov., and Walmadanyichnus hunteri, ichnogen. et ichnosp. nov.; and two thyreophoran ichnotaxa, Garbina roeorum, ichnogen. et ichnosp. nov., and Luluichnus mueckei, ichnogen. et ichnosp. nov. The level of diversity of the main track types is comparable across areas where tracksites are concentrated: Kardilakan–Jajal Buru (12), Walmadany (11), and Yanijarri (10). The overall diversity of the dinosaurian ichnofauna of the Broome Sandstone in the Yanijarri–Lurujarri section of the Dampier Peninsula is unparalleled in Australia, and even globally. In addition to being the primary record of non-avian dinosaurs in the western half of Australia, this ichnofauna provides our only detailed glimpse of Australia's dinosaurian fauna during the first half of the Early Cretaceous. It indicates that the general composition of Australia's mid-Cretaceous dinosaurian fauna was already in place by the Valanginian–Barremian. Both sauropods and ornithopods were diverse and abundant, and thyreophorans were the only type of quadrupedal ornithischians. Important aspects of the fauna that are not seen in the Australian mid-Cretaceous body fossil record are the presence of stegosaurians, an overall higher diversity of thyreophorans and theropods, and the presence of large-bodied hadrosauroid-like ornithopods and very large-bodied sauropods. In many respects, these differences suggest a holdover from the Late Jurassic, when the majority of dinosaurian clades had a more cosmopolitan distribution prior to the fragmentation of Pangea. Although the record for the Lower Cretaceous of Gondwana is sparse, a similar mix of taxa occurs in the Barremian–lower Aptian La Amarga Formation of Argentina and the Berriasian–Hauterivian Kirkwood Formation of South Africa. The persistence of this fauna across the Jurassic-Cretaceous boundary in South America, Africa, and Australia might be characteristic of Gondwanan dinosaurian faunas more broadly. It suggests that the extinction event that affected Laurasian dinosaurian faunas across the Jurassic-Cretaceous boundary may not have been as extreme in Gondwana, and this difference may have foreshadowed the onset of Laurasian-Eurogondwanan provincialism. The disappearance of stegosaurians and the apparent drop in diversity of theropods by the mid-Cretaceous suggests that, similar to South America, Australia passed through a period of faunal turnover between the Valanginian and Aptian. -------- In: Society of Vertebrate Paleontology Memoir (Journal of Vertebrate Paleontology Vol. 36, supplement to 6, November 2016).
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Martin, A.J. 2016. A close look at Victoria's first known dinosaur tracks. Memoirs of Museum Victoria 74: 63–71. Lower Cretaceous (Aptian-Albian) rocks of Victoria, Australia are well known for their dinosaur body fossils, but not so much for their trace fossils. For example, the first known dinosaur track from the Eumeralla Formation (Albian) of Knowledge Creek, Victoria, was not discovered until 1980. This specimen, along with two more Eumeralla tracks found at Skenes Creek in 1989, constituted all of the dinosaur tracks recognised in Lower Cretaceous strata of southern Australia until the late 2000s. Unfortunately, none of these first-known dinosaur tracks of Victoria were properly described and diagnosed. Hence, the main purpose of this study is to document these trace fossils more thoroughly. Remarkably, the Knowledge Creek and one of the Skenes Creek tracks are nearly identical in size and form; both tracks are attributed to small ornithopods. Although poorly expressed, the second probable track from Skenes Creek provides a search image for less obvious dinosaur tracks in Lower Cretaceous strata of Victoria. The Skenes Creek tracks were also likely from the same trackway, and thus may represent the first discovered dinosaur trackway from Victoria. These tracks are the first confirmed ornithopod tracks for Victoria, augmenting abundant body fossil evidence of small ornithopods ('hypsilophodontids') in formerly polar environments during the Early Cretaceous.
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Although tracks of dinosaurs are well known from Upper Jurassic sediments, tracks of non-dinosaurian vertebrates are fairly rare. The Upper Jurassic Lastres Formations of Asturias in northern Spain contain many vertebrate tracksites that include footprints and trackways of non-dinosaurian tetrapods. Several of these tracks are natural casts of pentadactyl to tridactyl footprints with digits connected by arched structures. The digits are short with deep scratch marks oriented anteriorly. The Asturian tracks show a high degree of morphological similarity to other specimens previously described as possible turtle tracks. Observations from extant turtle trackways show some surprising similarities with the fossil material. The tracks are here interpreted as having been made by turtles partially buoyed by water or by turtles walking in a slightly wet subaerial environment.
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Fossil eggs from the theropod nesting site found at Paimogo, Lourinhã (Upper Kimmeridgian -Tithonian), were identified with dinosauroid obliquiprismatic morphotype and crocodiloid basic type. Most eggshells have been affected by dissolution and recrystallization. The study of obliquiprismatic eggshell morphology indicates a high humidity nesting environment, about the same as required by crocodiles. Palavras-chave: Ninho; terópode; morfotipo obliquiprismático; tipo básico crocodiloide; diagénese; ambiente de nidificação. Resumo: Ovos fósseis do local de nidificação de terópodes encontrado em Paimogo, Lourinhã (Kimmeridgiano superior -Titoniano), foram identificados como pertencentes ao morfotipo dinossauroide obliquiprismático e ao tipo básico crocodiloide. A maior dinosaur and crocodile eggs from Paimogo nesting site. Memórias da Academia de Ciências de Lisboa, 37: 83–99. 2
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A synthesis on the state of art on dinosaur knowledge in Portugal is presented. The following genera have been recognized: Ceratosaurus, Torvosaurus, Lourinhanosaurus, Allosaurus, cf. Compsognathus, Stokesosaurus, cf. Richardoestesia, cf. Archaeopteryx, Euronychodon, cf. Paronychodon, Dinheirosaurus, Lourinhasaurus, Lusotitan, cf. Pleurocoelus, Lusitanosaurus, Dacentrurus, Dracopelta, Phyllodon, Hypsilophodon, Alocodon, Trimucrodon, Draconyx, Iguanodon, and Taveirosaurus. Most are from Late Jurassic localities at the Lourinhã area and Guimarota. A new genus, Lusotitan, is here raised to include the Late Jurassic ‘Brachiosaurus’ atalaiensis. Lower Cretaceous until Cenomanian material is scarce, except for dinosaur footprints. An interesting Late-Cretaceous, mostly small dinosaur association has been collected between Aveiro and Taveiro. To cite this article: M.T. Antunes, O. Mateus, C. R. Palevol 2 (2003) 77–95.
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Some small and medium-sized crocodylomorph footprints are described from the Late Jurassic (Kimmeridgian) coastal and deltaic units of the northern Spain (Asturias). There are at least four footprint morphotypes. Three of them, with well preserved trackways, are included in the ichnogenus Crocodylopodus (Crocodylopodus isp. and Crocodylopodus meijidei); the fourth one, documented by some isolated large footprints, is referable to the ichnogenus Hatcherichnus. This ichnoassociation confirms the presence of small crocodilians in palaeoenvironments apparently dominated by dinosaurs. The presence of Hatcherichnus seems to confirm the affinity between the Iberian and North American ichnofaunas.
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A new Allosauroid dinosaur (Saurischia: Theropoda) was found at Peralta near Lourinhã, Portugal (Upper Jurassic, Lower Tithonian). It is described under the name Lourinhanosaurus antunesi. It's diagnosable by the all vertebral centra longer than tall, neural spines of the anterior caudal vertebrae with a well-developed spike- like anterior process, the pubic blade is perforated by a large vertical ellipsoidal foramen and the lesser trochanter is well separate from the main body of the femur in lateral view. In the rib cage were found 32 gastroliths. This is the first non-avian theropod found with gastroliths.
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AbstrAct -The Upper Jurassic Lourinhã Formation (Lusitanian Basin, Portugal) contains a diverse dinosaur fauna comprising theropods, sauropods, stegosaurs, ankylosaurs and several genera of ornithopods. The sedimentology in the area favours preservation of tracksways, and tracks from most of the dinosaurs are also represented by skeletal remains. During fieldwork in the summer of 2003 a new, large, tridactyl track was found at the beach of Vale Frades, approximately 6 km north of Lourinhã (central west Portugal). The track was found together with a stegosaur track on a clay bed exposed within the intertidal zone. Due to the immediate danger of erosion, the track was collected and is now on display at Museu da Lou-rinhã. The track is 70 cm long and 69 cm wide, the toes are short and broad, with indications of short blunt claws, and there is a high angle of divarication between the outer digits. The shape and dimensions of the track identifies it as deriving from an ornithopod dinosaur with an estimated hip height around three metres. Although very large ornithopods are known from the Cretaceous, the largest known Jurassic ornithopod is Camptosaurus from North America, and the largest known from Portugal is the camptosaurid Draconyx loureiroi. Neither of these reached the body size suggested by the new track. So far the track described herein is the only evidence for a Jurassic ornithopod of that size.
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Isolated teeth of small theropod dinosaurs from the Upper Jurassic lignite coal mine of Guimarota (near Leiria, Portugal) are described and illustrated. The well known Upper Jurassic theropods from Europe,Archaeopteryx andCompsognathus, are the most common taxa in the Guimarota assemblage. One morphotype is closely related to an allosaurid theropod. Six further morphotypes of theropod teeth are also described, which are closely related to Cretaceous theropods such as dromaeosaurids, troodontids, tyrannosaurids,Richardoestesia andParonychodon. A Late Jurassic origin of these groups of theropods, which is very often postulated, is discussed. Isolierte Zähne theropoder Dinosaurier aus dem Oberen Jura der Kohlengrube Guimarota (bei Leiria, Portugal) werden beschrieben und abgebildet. Die häufigsten Zahnfunde ähneln morphologisch den Bezahnungen der oberjurassischen TheropodenArchaeopteryx undCompsognathus. Ein Morphotyp deutet auf das Vorhandensein eines Allosauriden hin. Sechs weitere Morphotypen von Theropoden-Zähnen zeigen große Ähnlichkeiten mit kreidezeitlichen Formen, wie Dromaeosauriden, Troodontiden, Tyrannosauriden,Richardoestesia undParonychodon. Für diese Gruppen wird ein spätjurassischer Ursprung diskutiert, wie er von einigen Autoren bereits vermutet wurde.
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We report new sauropod material from the upper sandstones of the Bombarral Formation of the Lourinhã Group (Upper Jurassic, Tithonian) at the Praia da Areia Branca cliffs (Lourinhä region, west coast of Portugal). The new specimen is represented by an articulated, well-preserved, partial tail of an animal of an estimated length of 22 m. The specimen was recovered from beds related to a fluvial/upper delta plain environment of a deltaic system. Currently, nine caudal vertebrae and four chevrons have been prepared. The specimen bears some tooth marks that are attributed to a large theropod. All the recovered bones are articulated, suggesting that an almost complete individual is most probably preserved in the lower layer of the bottom of the cliff. A geophysical test was performed to test this hypothesis. These analyses are not completely conclusive but the presence of resistive anomalies is discussed.
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We describe the first American stegosaur track of the ichnospecies Deltapodus brodricki, collected in the Upper Jurassic Morrison Formation of San Juan County, southeastern Utah, United States. The track is preserved as a natural cast on the underside of a slab of fluvial sandstone and consists of a well-preserved pes track and the eroded remains of a manus track. Previously, Deltapodus was known only from the Middle Jurassic Yorkshire coast of England and the Upper Jurassic of Portugal and Spain. The new discovery thus substantially extends the geographic record of this ichnospecies and highlights the similarities between the Late Jurassic dinosaur faunas of North America and those of Western Europe.
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Dinosaur tracks are abundant in the Middle Jurassic rocks of Yorkshire and indeed characterize the non-marine sequences developed within the Cleveland Basin. These tracks and associated trackways provide valuable evidence of the possible diversity of the dinosaur communities, their potential makers and behaviour and useful insights into the habitats and palaeo-environment during the time of deposition. The uneven historical development of research into Yorkshire dinosaur tracks is reviewed and the Middle Jurassic lithostratigraphy, biostratigraphy and chronostratigraphy of the region is outlined. Next, the probable palaeoenvironment of the Middle Jurassic Cleveland Basin, generally regarded as a coastal plain and fluvial complex, is briefly summarized. The terminology used to describe the dominant preservational types of dinosaur tracks, such as surface, transmitted and underprints, is clearly defined, with examples from the Yorkshire sequences. The Yorkshire tracks show considerable morphological diversity and at present 29 different morphotypes have been recognized, which possibly represent at least 15 ich-notaxa. These morphotypes include both quadrupedal and bipedal forms, as well as a distinctive suite of raking prints resulting from swimming activity. The distribution and abundance of the known dinosaur tracks within the Middle Jurassic rocks of Yorkshire is described. For the first time, a range chart of dinosaur tracks is presented that illustrates the persistence of some morphotypes throughout the Ravenscar Group (Middle Jurassic) of the Cleveland Basin. Track distribution and diversity data allow reconstruction of the Yorkshire dinosaur communities that were made up of between 7-10 common types, belonging to sauropods, stegosaurids, ornithopods and theropods. The area is a 'megatracksite' of global importance.
Autopodial bones from the holotype of the ankylosaur Dracopelta zbyszewskii, from the Upper Jurassic of Portugal, are described for the first time. This material was found in association with a partial rib cage from the Tithonian of Praia do Sul (Assenta), near Torres Vedras and Ribamar da Ericeira (not Ribamar of the Lourinhã area as previously stated), on the west coast of Portugal. The remains are preserved articulated in a block of sandstone and consist of three digits (II, III and IV) and partial metapodials from a right manus (?) of an adult individual. The specimen is one of the few articulated autopodia known for an ankylosaur. Dracopelta can be diagnosed by the presence of proximal phalanges II and III as long as wide.
Chapter
The carnosaur manus (hand) was usually tridactyl, with three fingers conventionally identified as I, II and III. In early or persistently primitive-looking carnosaurs there was sometimes a remnant of digit IV, and in advanced forms such as Tyrannosaurus the outermost of the three fingers was sometimes so reduced that the manus was functionally didactyl. Overall, the hand was much smaller than the foot and seems to have had no major supportive role in locomotion. Its slender fingers terminated in strong curved claws, so large that they have sometimes been mistaken for those of the hindfoot (Russell 1970: 11). The fingers were not very widely divergent, in some instances almost parallel. In the standard tridactyl manus the phalangeal formula was 2:3:4:0:0; in the didactyl manus of tyrannosaurs the formula was 2:3:0:0:0.
Article
Fragmentary theropod remains from the Upper Jurassic (Kimmeridgian) of Guimarota, Portugal, represent a new taxon of theropod dinosaurs, Aviatyrannis jurassica gen. et sp. nov. Together with Stokesosaurus from the Morrison Formation of North America, Aviatyrannis represents the oldest known tyrannosauroid, indicating that tyrannosauroid origins may be found in the Middle-Late Jurassic of Europe/North America. Furthermore, current evidence suggests that early tyrannosaurs were rather small animals, which is in general accordance with their origin amongst the generally rather small coelurosaurs.
Article
A recently discovered dinosaur tracksite from the Upper Jurassic Morrison Formation, Bighorn Basin, Wyoming, contains abundant sauropod tracks that exhibit varying degrees of preservation. Most of these tracks appear as indistinct bulges on the bottoms of sandstone beds, but several are well preserved and show foot-pad and skin impressions. Three track morphotypes are recognized: a sauropod pes print, a Brontopodus-like manus print, and a diplodocid manus print. The Brontopodus-like manus print most likely represents the footprint of a brachiosaur. This morphotype also contains evidence of phalangeal nodes—the first reported for a sauropod manus. The diplodocid manus print is unique because it contains impressions of a substantial ungual on digit I and a heel pad. A partial sauropod track cast also contains an impression of interlocking, polygonal scales. This is only the second known North American sauropod footprint that contains skin impressions. The spectrum of preservational quality of the tracks and associated trace fossils is used to infer the relative moisture content of the original substrate. Moisture content of the original substrate is estimated to have been moist to borderline saturated. Observations of the tracks at the study areas also are used to establish a list of features that can be used to distinguish deep vertebrate tracks from load casts resulting from gravity-induced soft-sediment deformation.
Article
Dinosaur tracks are abundant in the Middle Jurassic rocks of Yorkshire and indeed characterize the non-marine sequences developed within the Cleveland Basin. These tracks and associated trackways provide valuable evidence of the possible diversity of the dinosaur communities, their potential makers and behaviour and useful insights into the habitats and palaeoenvironment during the time of deposition. The uneven historical development of research into Yorkshire dinosaur tracks is reviewed and the Middle Jurassic lithostratigraphy, biostratigraphy and chronostratigraphy of the region is outlined. Next, the probable palaeoenvironment of the Middle Jurassic Cleveland Basin, generally regarded as a coastal plain and fluvial complex, is briefly summarized. The terminology used to describe the dominant preservational types of dinosaur tracks, such as surface, transmitted and underprints, is clearly defined, with examples from the Yorkshire sequences. The Yorkshire tracks show considerable morphological diversity and at present 29 different morphotypes have been recognized, which possibly represent at least 15 ichnotaxa. These morphotypes include both quadrupedal and bipedal forms, as well as a distinctive suite of raking prints resulting from swimming activity. The distribution and abundance of the known dinosaur tracks within the Middle Jurassic rocks of Yorkshire is described. For the first time, a range chart of dinosaur tracks is presented that illustrates the persistence of some morphotypes throughout the Ravenscar Group (Middle Jurassic) of the Cleveland Basin. Track distribution and diversity data allow reconstruction of the Yorkshire dinosaur communities that were made up of between 7-10 common types, belonging to sauropods, stegosaurids, ornithopods and theropods. The area is a 'megatracksite' of global importance.
Article
The Lusitanian Basin of west-central Portugai is an Iberian Atlantic margin basin. During a late Jurassic rifting event, a series of alluvial fans distributed coarse, basement-derived clastics, southeastward from the western margin of the basin. The distal part of this sequence, which is about 140 m thick, consists of heterolithic, massive sandstone andm assive mudrock facies, arranged in crude fining-upwards cycles 1-15 m thick. These form discrete packets of sediment in a strongly aggradational sequence and are interpreted as the products of sheetflooding as the active area of sedimentation on the fan surface switched. Channelized flow was not necessarily an early feature of cycle development. Considerable fluctuations in discharge occurred and were probably related to climatic factors. Large quantities of tractional and suspended sediment were transported at high stages, and were rapidly deposited as flows waned. Caliche soil profiles indicate a seasonal semi-and climate. However, other features such as widespread soft-sediment deformation, the lack of desiccation structures and the absence of evaporites suggests that areas of active sedimentation often remained wet. Three fining-upward megacycles are distinguished. These indicate rapid advance of fan toes triggered by tectonic activity followed by gradual retreat.