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A new species of Struthiosaurus (Dinosauria: Ankylosauria) from the Upper Cretaceous of Villeveyrac (Southern France)
Abstract
A new species of the ankylosaur Struthiosaurus from the Upper Cretaceous (lower Campanian) of the Villeveyrac-Mèze Basin, southern France, is described from a partial skeleton that includes distal dorsal vertebrae, synsacrum and pelvic girdle. Struthiosaurus languedocensis, sp. nov. is a small-sized nodosaurid (less than 3 m length) characterised by distal dorsal centra that are very compressed laterally and hourglass in shape in ventral view; ischium directed immediately caudal from the acetabulum, with a robust shaft that does not taper distally and that is weakly curved in a caudoventral orientation. The synsacrum of S. languedocensis consists of ten co-ossified vertebrae, including five dorsals, four sacrals and a caudal. Among ankylosaurs, only Polacanthus foxii has a similar synsacral count, but there are significant differences in the pelvic structure between Struthiosaurus and Polacanthus. The presence of an ischium lacking a distinct nodosaurid-like ventral flexion appears to be diagnostic for the genus Struthiosaurus, as suggested by additional remains recovered from the upper Campanian of Laño (Iberian Peninsula). Current data suggests that Struthiosaurus is represented by different species in southwestern and central Europe.
... Ventrally, anterior dorsal centra possess a broad, gentle ridge that becomes less distinct extending posteriorly along the column (Figs 6B, 7B). This differs from the condition in Hylaeosaurus and Struthiosaurus languedocensis (Garcia & Pereda Suberbiola, 2003), which both possess a keel ventrally on dorsal centra . ...
... Transverse processes are only preserved on two of the anterior dorsals; these project dorsolaterally, and are triangular in cross-section with a flat dorsal surface and a ventral lamina that extends to the top of the parapophysis (Fig. 6, tp), as in Pinacosaurus (Burns et al., 2015) and Sauropelta (YPM 5154, 5495). A single prezygapophysis is preserved on a more posterior free dorsal; it is elliptical, faces ventromedially and would have been joined to its counterpart ventrally (Fig. 7A, D, przyg), as in Ankylosaurus (Brown, 1908), Pinacosaurus (Burns et al., 2015), Euoplocephalus (AMNH 5337), Struthiosaurus languedocensis (Garcia & Pereda Suberbiola, 2003), Polacanthus and Sauropelta (YPM 5154, 5495). In posterior view, a sharp, very prominent lamina arises dorsal to the neural canal and extends to the base of the postzygapophyses (Figs 6F, 7F, lam). ...
... This is similar to the condition in Gastonia burgei (BYU 11640) and Hylaeosaurus , but a sharp ridge and fossae are not present in Sauropelta (YPM 5154, 5495), Polacanthus or Euoplocephalus (AMNH 5337), which have a ridge but no fossae. Postzygapophyses are only preserved on the anterior-most free dorsal (Fig. 6F, pozyg); they are rounded in outline, face ventrally and slightly laterally, and are separated at their bases by a deep slit-like fossa, similar to Sauropelta (YPM 5154, 5495), Struthiosaurus languedocensis (Garcia & Pereda Suberbiola, 2003), Mymooropelta (Kirkland & Carpenter, 1994), Europelta (Kirkland et al., 2013) and Gastonia burgei (BYU 11640). This contrasts with the condition in Ankylosaurus (Brown, 1908) and Euoplocephalus (AMNH 5337), in which the postzygapophyses are joined ventrally. ...
... al., 2004). The small-bodied Struthiosaurus languedocensis has a maximum width of the preacetabular process reaching 12.5 cm and an estimated body length ranging between 2.5 and 3 m (Garcia and Suberbiola, 2003). Based on these comparisons, we estimate the body length of IVPP V26052 to be around 5e5.5 m, longer than Jinyunpelta but shorter than Polacanthus, Euoplocephalus, or Edmontonia. ...
... (Arbour and Mallon, 2017), Borealopelta, Sauropelta (Brown et al., 2017), and Scolosaurus (Penkalski and Blows, 2013). Small-to medium-sized ankylosaurians with a body length around 3e5 m include Gargoyleosaurus (3e3.5 m) (Kilbourne and Carpenter, 2005) (Kirkland and Carpenter, 1994;Garcia and Suberbiola, 2003;Leahey et al., 2015). Thus, given the lack of information indicating the precise ontogenetic stage of the specimen, the ankylosaurian specimen from the Longjing Formation (IVPP V26052) at least represents a moderate body-sized ankylosaurian dinosaur. ...
... Ankylosaurian remains are very abundant in China and have been reported from the Jurassic of the Junggar Basin in Xinjiang Uyghur Autonomous Region (Augustin et al., 2020;Dong, 1993); the Lower Cretaceous of Yixian (Xu et al., 2001), the Lower to Upper Cretaceous of Beipiao (Dong, 2002;Lü et al., 2007a;Yang et al., 2017), the Lower Cretaceous of Fuxin (Amiot et al., 2010), the Lower Cretaceous of Lingyuan (Han et al., 2014), and the Lower Cretaceous of Kazuo (Ji et al., 2014) in Liaoning Province; the Lower Cretaceous of Yongjing in Gansu Province (Yang et al., 2013), the Lower Cretaceous of Urad Houqi (Arbour and Currie, 2016; Bohlin, 1953), the Lower and Upper Cretaceous of Alashan (Vickaryous et al., 2001;Young, 1935), and the Upper Cretaceous of Bayan Mandahu (Burns et al., 2011;Godefroit et al., 1999) in Inner Mongolia Autonomous Region; the Upper Cretaceous of Tianzhen (Barrett et al., 1998;Pang and Cheng, 1998), and Zuoyun (Jia, 2018) in Shanxi Province; the Upper Cretaceous of Laiyang in Shandong Province (Buffetaut, 1995); the Cretaceous of Ruyang in Henan Province (Xu et al., 2007); the Cretaceous of Lishui (Lü et al., 2007b), Dongyang (Chen et al., 2013) and Jinyun in Zhejiang Province (Fig. 4). Among them, the localities of Beipiao in Liaoning, Zuoyun in Shanxi, and Lishui, Dongyang, and Jinyun in Zhejiang have similar geological ages to the Longjing Formation (Chen et al., 2013;Yang et al., 2017;Jia, 2018;Zheng et al., 2018). ...
We here report an ankylosaurian ilium from the Albian–Cenomanian Longjing Formation of Yanji City, Jilin Province, northeastern China. The diagnostic features allowing referral of this specimen to Ankylosauria include: an ilium rotated horizontally dorsal to the acetabulum with the primitive lateral surface facing ventrally; long and laterally divergent preacetabular process; and a shallow, cup-like acetabulum. This new specimen represents the first definitive ankylosaurian dinosaur from Jilin Province, and the easternmost ankylosaurian occurrence in China. The present discovery demonstrates the potential for future finds in the Longjing Formation, especially in the Longshan fossil beds of Yanji City, such as additional ankylosaurian or other dinosaur remains that might provide significant data on dinosaur evolution during the middle Cretaceous of eastern Asia.
... comm.), amphibians (Albanerpetondidae and the oldest representative of the Palaeobatrachidae), squamates (the mosasauroid specimens described here), turtles (Solemys and Polysternon provinciale), crocodylians (the eusuchians Allodaposuchus and Acynodon), a possible pterosaur, and dinosaurs. The latter includes two ornithopods (the endemic European Rhabdodon and an indeterminate small form), one ankylosaur (the nodosaurid Struthiosaurus languedocensis Garcia and Pereda-Suberbiola, 2003), and at least one theropod (referred to as Dromaeosauridae indet.). ...
... List of the vertebrate fossils from the lignite bearing bed of the lower part of the series (Early Campanian) in the L'Olivet mine from Villeveyrac (Hérault Department). Established on the basis of newly collected material and after Buffetaut et al. (1996), de Lapparent de Broin and Murelaga (1999), Tong and Gaffney (2000) and Garcia and Pereda-Suberbiola (2003). In bold, the taxon whose fossils are described in this paper. ...
... Tableau 1 Liste des fossiles de vertébrés provenant du niveau ligniteux à la base de la série dans la carrière (Campanien inférieur) de la mine de l'Olivet à Villeveyrac (Hérault). Établie sur la base de nouvelles récoltes de matériel et d'après Buffetaut et al. (1996), de Lapparent de Broin et Murelaga (1999), Tong et Gaffney (2000) et Garcia et Pereda-Suberbiola (2003). En gras, le taxon dont les fossiles sont décrits dans ce travail. ...
... comm, 2014) and the Santonian Hungarosaurus based on six associated skeletons (Ősi 2005;Ősi and Makádi 2009;Ősi 2015, in the paper). The Santonian to Maastrichtian ranged Struthiosaurus is known by ten different associated skeletons (Nopcsa 1929;Pereda-Suberbiola and Galton 2001;Garcia and Pereda-Suberbiola 2003;Ősi and Pereda-Suberbiola 2017), but they came from different tectonical units (Romania, Austria, Hungary, Spain) and the remains are poorly overlapping with each other to prevent comparative work. ...
... Dental wear analysis fulfilled on the in situ teeth of this specimen shows that a palinal jaw movement completed the powerstroke of Hungarosaurus (Ősi et al. 2014b). Although the teeth of S. languedocensis from France (Garcia and Pereda-Suberbiola 2003;Ősi et al. 2016a, b) and those referred to S. austriacus (Nopcsa 1929) also bear dental wear due to tooth-tooth contact (precise shearing bite), scratch orientation is still unknown; thus, the direction of jaw movement in Struthiosaurus is ambiguous. Stratigraphic position: MTM 2007.25 was discovered from the strongly cemented grey sandstone bed (unit 2), which is the covering bed of the most important fossiliferous layer (unit 1). ...
... True ankylosaurian diversity in Europe is, however, greatly masked by the lack of overlapping diagnostic elements in many assemblages (Ősi 2015). Thus, except for a few elements (Pereda-Suberbiola 1993a), all the Campanian-Maastrichtian ankylosaur material have been referred to as Struthiosaurus (Bunzel 1871;Seeley 1881;Nopcsa 1915Nopcsa , 1929Pereda-Suberbiola 1992, 1993b, 1999Galton 1994, 2001;Garcia and Pereda-Suberbiola 2003;Company 2004;Ősi et al. 2014c;Ősi and Pereda-Suberbiola 2017). Study and reconstruction of armour composition and cervical and pelvic armour morphology might help to challenge this view in the future (Burns 2008;Arbour et al. 2014;Burns and Currie 2014;Ősi and Pereda-Suberbiola 2017;Ősi and Rudolf 2017). ...
Ankylosaurian fossils are usually standard elements of Cretaceous continental vertebrate localities; however, bone-yielding horizons including more than one individual are extremely rare. Here, we present a unique assemblage of 12 partial, articulated or associated ankylosaurian skeletons and thousands of isolated bones and teeth discovered from the Santonian Iharkút vertebrate locality, western Hungary. Collected from an area of 600 m2 and from a single bone bed, this material is one of the richest ankylosaurian accumulation worldwide. The 12 skeletons are not monospecific but mostly based on the pelvic armour composition: six of them are from Hungarosaurus, two are referred to Struthiosaurus and four can be assigned to Nodosauridae indet. Sedimentological and taphonomical examinations revealed a single mass mortality event as the cause of the death and accumulation of these quadruped animals that are described here. The ankylosaur assemblage from Iharkút suggests at least a temporarily gregarious behaviour of these animals and also shows that Hungarosaurus and Struthiosaurus might live in the same moist habitat or at least preferred relatively close environments.
... comm.), amphibians (Albanerpetondidae and the oldest representative of the Palaeobatrachidae), squamates (the mosasauroid specimens described here), turtles (Solemys and Polysternon provinciale), crocodylians (the eusuchians Allodaposuchus and Acynodon), a possible pterosaur, and dinosaurs. The latter includes two ornithopods (the endemic European Rhabdodon and an indeterminate small form), one ankylosaur (the nodosaurid Struthiosaurus languedocensis Garcia and Pereda-Suberbiola, 2003), and at least one theropod (referred to as Dromaeosauridae indet.). ...
... List of the vertebrate fossils from the lignite bearing bed of the lower part of the series (Early Campanian) in the L'Olivet mine from Villeveyrac (Hérault Department). Established on the basis of newly collected material and after Buffetaut et al. (1996), de Lapparent de Broin and Murelaga (1999), Tong and Gaffney (2000) and Garcia and Pereda-Suberbiola (2003). In bold, the taxon whose fossils are described in this paper. ...
... Tableau 1 Liste des fossiles de vertébrés provenant du niveau ligniteux à la base de la série dans la carrière (Campanien inférieur) de la mine de l'Olivet à Villeveyrac (Hérault). Établie sur la base de nouvelles récoltes de matériel et d'après Buffetaut et al. (1996), de Lapparent de Broin et Murelaga (1999), Tong et Gaffney (2000) et Garcia et Pereda-Suberbiola (2003). En gras, le taxon dont les fossiles sont décrits dans ce travail. ...
New mosasauroid material from the Villeveyrac-L’Olivet mine (Hérault, southern France)is described. This historical and fossiliferous bauxite mine has yielded from the samestratigraphical level (Early Campanian) diverse continental vertebrates, including fishes(Lepisosteidae and Phyllodontidae), amphibians (Albanerpetondidae and the oldest repre-sentative of the Palaeobatrachidae), turtles (Solemydidae and Bothremydidae), squamates(Mosasauroidea), crocodilians (Eusuchia), dinosaurs (Rhabdodontidae, Nodosauridae andDromaeosauridae), and a possible pterosaur. The mosasauroid specimens represent differ-ent individuals of a single taxon, belonging to a large plesiopedal tethysaurine-like form(about 3 meters long). This discovery adds to previous reports of plesiomorphic freshwatermosasauroids in the Late Cretaceous dinosaur-bearing sites of the European archipelago.
... comm.), amphibians (Albanerpetondidae and the oldest representative of the Palaeobatrachidae), squamates (the mosasauroid specimens described here), turtles (Solemys and Polysternon provinciale), crocodylians (the eusuchians Allodaposuchus and Acynodon), a possible pterosaur, and dinosaurs. The latter includes two ornithopods (the endemic European Rhabdodon and an indeterminate small form), one ankylosaur (the nodosaurid Struthiosaurus languedocensis Garcia and Pereda-Suberbiola, 2003), and at least one theropod (referred to as Dromaeosauridae indet.). ...
... List of the vertebrate fossils from the lignite bearing bed of the lower part of the series (Early Campanian) in the L'Olivet mine from Villeveyrac (Hérault Department). Established on the basis of newly collected material and after Buffetaut et al. (1996), de Lapparent de Broin and Murelaga (1999), Tong and Gaffney (2000) and Garcia and Pereda-Suberbiola (2003). In bold, the taxon whose fossils are described in this paper. ...
... Tableau 1 Liste des fossiles de vertébrés provenant du niveau ligniteux à la base de la série dans la carrière (Campanien inférieur) de la mine de l'Olivet à Villeveyrac (Hérault). Établie sur la base de nouvelles récoltes de matériel et d'après Buffetaut et al. (1996), de Lapparent de Broin et Murelaga (1999), Tong et Gaffney (2000) et Garcia et Pereda-Suberbiola (2003). En gras, le taxon dont les fossiles sont décrits dans ce travail. ...
New mosasauroid material from the Villeveyrac-L’Olivet mine (Hérault, southern France)is described. This historical and fossiliferous bauxite mine has yielded from the samestratigraphical level (Early Campanian) diverse continental vertebrates, including fishes(Lepisosteidae and Phyllodontidae), amphibians (Albanerpetondidae and the oldest repre-sentative of the Palaeobatrachidae), turtles (Solemydidae and Bothremydidae), squamates(Mosasauroidea), crocodilians (Eusuchia), dinosaurs (Rhabdodontidae, Nodosauridae andDromaeosauridae), and a possible pterosaur. The mosasauroid specimens represent differ-ent individuals of a single taxon, belonging to a large plesiopedal tethysaurine-like form(about 3 meters long). This discovery adds to previous reports of plesiomorphic freshwatermosasauroids in the Late Cretaceous dinosaur-bearing sites of the European archipelago.
... Galobart cally and temporally match with the studied egg type. This premise restricts our search to late Campanian-early Maastrichtian titanosaurs, rhabdodontids, nodosaurid ankylosaurs, and both large and small theropods (Le Loeuff 1991;Le Loeuff et al. 1994;Laurent et al. 2002;Riera et al. 2009 (Garcia and Pereda-Suberbiola 2003). This feature has been related to limb biomechanical, but it could have also played a relevant reproductive role. ...
... Like in extant birds, the backward orientation of ischia in Struthiosaurus produces an opened ventral space that significantly increases the pelvic cavity (Figure 7(B)). The pelvic aperture of Struthiosaurus languedocencis measures about 29 cm in wide (Garcia and Pereda-Suberbiola 2003), whereas that of Struthiosaurus sp. from Laño locality (N Spain; Pereda-Suberbiola 1999) is about 26 cm wide. In both cases, a spherical egg of 15-19 cm in diameter could have easily passed away through the pelvic passage ( Figure 6(B)), even if the abdominal cavity is filled with inner tissues. ...
... Thus, albeit for exclusion, the only remaining candidates are the nodosaurid ankylosaurs. In SW Europe this group of armoured dinosaurs is represented by a single genus, Struthiosaurus (Pereda-Suberbiola 1999;Garcia and Pereda-Suberbiola 2003;Ö si and Prondvai 2013). Despite that Struthiosaurus could be small in size(Ö si and Prondvai 2013), it shows a unique pelvic configuration among ankylosaurians in having the ischia horizontally projected backward without caudal closing ...
The fossil record of dinosaur eggs and eggshells from the uppermost Cretaceous strata of south-western Europe is composed of both worldwide-distributed and endemic egg types. In this study, we are reviewing the enigmatic European oogenus Cairanoolithus, which after analysing abundant material from classic and new localities it is reassigned to a new oofamily (Cairanoolithidae oofam. nov.) in the light of the unique combination of structural characters. The new oofamily includes one oogenus and two oospecies. Cairanoolithid eggs share several features with other ornithopod egg types indeed.
Furthermore, our phylogenetic analysis places Cairanoolithus as the sister ootaxon of the ornithopod ootaxa, being considered the most basal ornithischian egg type known so far. Although neither embryonic nor bones remains are known in association with cairanoolithid eggs so far, several taxonomic attributions have been proposed for this egg type over time. On the basis of microstructural features, phylogenetic results and anatomical constrains, we discuss in this paper previous taxonomic attributions and provide new evidence for suggesting a plausible nodosaurid affinity.
... The preacetabular process is very long and transversely wide, and diverges laterally from the vertebral column at an angle of approximately 45u. The lateral margin of the preacetabular process is straight in ventral view, as in ankylosaurids, but unlike the condition in most nodosaurids, such as Sauropelta [25], Struthiosaurus [48], and Zhejiangosaurus [49], in which it is laterally curved. The postacetabular process is subtriangular in outline. ...
... The shaft of the ischium is unique in being narrower in its mid-shaft region and widening towards to the distal end, prior to tapering again further distally, whereas in other ankylosaurians the shaft either tapers distally along the whole shaft (e.g. Ankylosaurus [28], Sauropelta, Edmontonia [48]) or remains sub-equal in size along the whole shaft (e. g., Euoplocephalus: [31]), or is just slightly expanded at the distal end (e.g., Cedarpelta: [44]) (Fig. 9). The proximal end of the ischium is straight in lateral view. ...
... The proximal end of the ischium is straight in lateral view. This is unlike the convex and fan-like ischium in the ankylosaurids Ankylosaurus [50] and Euoplocephalus [51], and also unlike the concave proximal ischia of the nodosaurids Struthiosaurus and Edmontonia [48]. The proximal end of the ischium lacks the medial wall present in the basal ankylosaurid Cedarpelta [20,44]. ...
A new ankylosaurid, Chuanqilong chaoyangensis gen. et sp. nov., is described here based on a nearly complete skeleton from the Lower Cretaceous Jiufotang Formation of Baishizui Village, Lingyuan City, Liaoning Province, China. Chuanqilong chaoyangensis can be diagnosed on the basis of two autapomorphies (glenoid fossa for quadrate at same level as the dentary tooth row; distally tapering ischium with constricted midshaft) and also a unique combination of character states (slender, wedge-like lacrimal; long retroarticular process; humerus with strongly expanded proximal end; ratio of humerus to femur length = 0.88). Although a phylogenetic analysis places Chuanqilong chaoyangensis as the sister taxon of the sympatric Liaoningosaurus near the base of the Ankylosauridae, the two taxa can be distinguished on the basis of many features, such as tooth morphology and ischial shape, which are not ontogeny-related. Chuanqilong chaoyangensis represents the fourth ankylosaurid species reported from the Cretaceous of Liaoning, China, suggesting a relatively high diversity in Cretaceous Liaoning.
... During the following 60 years no significant ankylosaurian material has been unearthed from the Upper Cretaceous of Europe (for an overview of this period see Pereda-Suberbiola, 1992, 1993a, preventing the recognition of the true diversity and distribution of this group. Recent discoveries of new ankylosaurian material in northern Spain (Astibia et al., 1990;Pereda-Suberbiola, 1999;Pereda-Suberbiola et al., 1995) and southern France (Buffetaut and Le Loeuff, 1991;Garcia and Pereda-Suberbiola, 2003;Pereda-Suberbiola, 1993a) supported the presence of Struthiosaurus in western Europe; a third species of the genus, Struthiosaurus languedocensis, has been also described (Garcia and Pereda-Suberbiola, 2003) from the Campanian of Villeveyrac (southern France). Furthermore, associated and isolated postcranial elements referred to as Struthiosaurus sp. have been recently reported from eastern Spain (Chera) by Company (2004) ( Table 1). ...
... During the following 60 years no significant ankylosaurian material has been unearthed from the Upper Cretaceous of Europe (for an overview of this period see Pereda-Suberbiola, 1992, 1993a, preventing the recognition of the true diversity and distribution of this group. Recent discoveries of new ankylosaurian material in northern Spain (Astibia et al., 1990;Pereda-Suberbiola, 1999;Pereda-Suberbiola et al., 1995) and southern France (Buffetaut and Le Loeuff, 1991;Garcia and Pereda-Suberbiola, 2003;Pereda-Suberbiola, 1993a) supported the presence of Struthiosaurus in western Europe; a third species of the genus, Struthiosaurus languedocensis, has been also described (Garcia and Pereda-Suberbiola, 2003) from the Campanian of Villeveyrac (southern France). Furthermore, associated and isolated postcranial elements referred to as Struthiosaurus sp. have been recently reported from eastern Spain (Chera) by Company (2004) ( Table 1). ...
... The tooth (LPB (FGGUB) R.2182) markedly differs from those of other ankylosaurs (e.g., S. austriacus [PIUW 2349/105b, Pereda-Suberbiola andGalton, 2001: fig. 10.2N], S. languedocensis [Garcia and Pereda-Suberbiola, 2003], Edmontonia spp, Sauropelta edwardsorum and Euoplocephalus tutus [Coombs, 1990;Ő.A. personal observation], Ankylosaurus magniventris [Carpenter, 2004], Gargoyleosaurus parkpini [DMNH 27726], H. tormai [MTM 2007.25.2], Peloroplites [Carpenter et al., 2008], Pinacosaurus grangeri [ZPAL MGD-II/1], Tarchia gigantea [PIN N3142 250], or Saichania chulsanensis [Carpenter et al., 2011]) in the lack of a central cusp surrounded mesiodistally by several minor cusps, and in having only six, more or less equally sized, apically pointed cusps separated by deeply incised grooves. ...
Ankylosaurian remains from the Transylvanian Basin, Romania, are extremely rare. More than 100 years after the discovery of the first and only better-known assemblage, namely the type material of Struthiosaurus transylvanicus, new ankylosaurian material has been discovered in the Maastrichtian of the Hateg Basin, as well as at another locality (Vurpar), in the Transylvanian Basin, that is described here. The material consists of one tooth in a small jaw fragment (from the Hateg Basin) and at least two accummulations of associated, as well as several isolated, postcranial elements (from Vurpar). No diagnostic elements are preserved that would overlap with the type of S. transylvanicus, so we cannot assign any of the new specimens to this species. The tooth shows marked differences compared to those of other anklyosaurs including S. austriacus and Hungarosaurus in having only six, more or less equally sized, apically pointed cusps separated by deep grooves. The postcranial material from Vurpar represents at least three different individuals. The humerus is the most diagnostic element among the postcranial remains being most similar both in size and morphology to humeri referred to as Struthiosaurus from different European localities, thus here we refer the humerus and probably associated elements preserved in one assemblage to as cf. Struthiosaurus sp.; the remaining specimens from Vurpar are retained as Nodosauridae indet Histological studies have confirmed the adult nature of all sampled bones in the Vurpar ankylosaur material suggesting that these fully grown animals were of similar size to Struthiosaurus, a small-bodied nodosaurid the ontogenetic status of which, however, has never been investigated histologically. The obviously diminished body size of the Transylvanian ankylosaurs compared to other members of the clade could be explained by insular dwarfism using the same histology-based argument as presented for Magyarosaurus.
... Only nodosaurids have been described from the Upper Cretaceous of Europe with Struthiosaurus austriacus described from the Campanian of Austria in 1871 [19][20][21][22][23][24] followed by Struthiosaurus transylvanicus [25,26,27] from the uppermost Cretaceous (upper Maastrichtian) strata of Romania. Until recently, all Late Cretaceous ankylosaur fossils in Europe have been assigned to Struthiosaurus [28][29][30] including Struthiosaurus languedocensis from the Campanian of southern France [31]. The primitive nodosaurid Hungarosaurus tormai [32,33] from the mid-Late Cretaceous (Santonian) is now known from multiple specimens and has become the best documented ankylosaur in Europe. ...
... Likewise, lower Cenomanian teeth assigned to ''Acanthopholis'' have more deeply divided denticles in what is a proportionally taller tooth [17]. The teeth of Struthiosaurus languedocensis [31] from the lower Campanian of France also differ in size and in having longer, lower tooth crowns. ...
... The sacrum of Struthiosaurus languedocensis [31] is similar overall, but based on the description is not so strongly anteroposteriorly arched as in Europelta. Similarly, the sacrum of Hungarosaurus, as exhibited at the Hungarian Natural History Museum, appears to be moderately arched. ...
Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids- the Struthiosaurinae.
... Since Lapparent [105], many discoveries have been made [13,23,45,87,110,113]. The most productive dinosaur sites are Fox Amphoux [15,105] and La Boucharde [3,8] in the Aix-en-Provence Basin; Champ-Garimond [152] in Gard; Villeveyrac [44,82] and Cruzy [46] in Hérault, the Upper Aude Valley [30,31,36,113,114] and Montplaisir [118,119] in Aude, and the Plantaurel [112] in Ariège. ...
... Ankylosaur remains are not abundant, and most of the known material is indeterminate [3, 110,136,137]. A new species of the nodosaurid Struthiosaurus has recently been described from a partial skeleton that includes dorsal vertebrae, synsacrum and pelvic girdle from the Early Campanian of Villeveyrac (Hérault) (Fig. 4) [13,82]. The species Rhodanosaurus ludgunensis Nopcsa, 1929 is based on non-diagnostic material from Hérault and is regarded here as a nomen dubium [136]. ...
... The species Rhodanosaurus ludgunensis Nopcsa, 1929 is based on non-diagnostic material from Hérault and is regarded here as a nomen dubium [136]. All ankylosaur remains appear to belong to nodosaurids; there is no current evidence of ankylosaurids in the Latest Cretaceous of southern France [82]. ...
The French dinosaur record is one of the most extensive in Europe; it ranges stratigraphically from the Late Triassic to the Latest Cretaceous. All major clades of dinosaurs but marginocephalians are known. About 20 species are based on significant material; the theropods are the best represented. Most of these taxa have been described or revised in recent years. Important specimens have been discovered in the Late Triassic of eastern France, the Middle Jurassic of Normandy, and the Late Cretaceous of Provence and Languedoc. The ichnological record is good for the Late Triassic-Early Jurassic, and the Late Cretaceous egg sites are among the richest in the world. To cite this article: R. Allain, X.P. Suberbiola, Palevol 2 (2003) 27–44.
... During the Late Cretaceous, Europe was covered by a shallow epicontinental sea with numerous islands that were home to a wide variety of terrestrial vertebrates (Csiki-Sava et al., 2015). The ankylosaurian dinosaur Struthiosaurus, known from cranial and postcranial remains from Austria, Romania, France and Spain (Nopcsa, 1929;Pereda-Suberbiola et al., 1995;Pereda-Suberbiola & Galton, 2001;Garcia & Pereda-Suberbiola, 2003) ranked amongst these Late Cretaceous island-dwellers. Traditionally included in the family Nodosauridae, Struthiosaurus is widely acknowledged to belong to the smallest known ankylosaurs, with an estimated body length of two to three metres (Pereda-Suberbiola, 1992;Pereda-Suberbiola & Galton, 2001;Riguetti et al., 2022) and a mass of ~130 kg (Benson et al., 2018). ...
... Traditionally included in the family Nodosauridae, Struthiosaurus is widely acknowledged to belong to the smallest known ankylosaurs, with an estimated body length of two to three metres (Pereda-Suberbiola, 1992;Pereda-Suberbiola & Galton, 2001;Riguetti et al., 2022) and a mass of ~130 kg (Benson et al., 2018). As currently accepted, the genus Struthiosaurus includes three species: S. austriacus Bunzel, 1871 from the lower Campanian of Austria, S. languedocensis Garcia & Pereda-Suberbiola, 2003 from the Campanian of France and S. transylvanicus Nopcsa, 1915 from the Maastrichtian of Romania. In addition, ankylosaurian remains tentatively referred to as cf. ...
... It was in a close association with a left dorsal rib, the latter having a Tshaped cross-section in its anterior half ( Fig. 2A). The vertebral centrum is hourglass-shaped, but not as concave ventrally and laterally as the last dorsal of Struthiosaurus languedocensis (UM2 OLV-D50; Garcia and Pereda-Suberbiola, 2003). The ventral and lateral margins of the anterior, slightly concave articular surface are slightly eroded. ...
... Among the non-osteodermal skeletal elements, the sigmoidal lateral edge of the ilium in dorsal view and the relatively short postacetabular part of the ilium (Fig. 2) is more reminiscent to that of Struthiosaurus than of Hungarosaurus, further supporting the Struthiosaurus affinity of this specimen. The synsacra of Struthiosaurus languedocensis from Villeveyrac and Struthiosaurus sp. from Laño are composed of ten fused vertebrae (Garcia and Pereda-Suberbiola, 2003); nine vertebrae are known in the cf. Struthiosaurus synsacrum from Iharkút and eight or nine elements in the synsacral material of Hungarosaurus ( } Osi, 2005; } Osi and Mak adi, 2009). ...
The pelvic armor elements in the ankylosaurian material from the Upper Cretaceous of Iharkút, Hungary are described here. Among these, a new articulated hip region of a small bodied ankylosaur is referred here to cf. Struthiosaurus sp. It preserves, uniquely among Late Cretaceous European ankylosaurs, an in situ pelvic armor composed of among others four, keeled, oval to circular osteoderms lying centrally and arranged longitudinally above the synsacral neural spines. This is the first indication of this type of pelvic osteoderm arrangement in an ankylosaur, increasing our knowledge on this poorly known part of the ankylosaur skeleton. Some additional pelvic osteoderms are also described that help to reconstruct and distinguish the pelvic armor of the two Late Cretaceous European ankylosaurs Struthiosaurus and Hungarosaurus. Both taxa have some fused parts in the pelvic armor but most probably neither of them had a single, fused pelvic shield as that of the Early Cretaceous Polacanthus. Interwoven texture on the ventral surface of the osteoderms, observed in both European taxa and known in other ankylosaurs (e.g. Polacanthus, Nodosaurus), is suggested here to be a characteristic feature of the non-keeled, fused pelvic armor elements of Ankylosauria.
... En Europe, le Cénomanien n'avait jusqu'à présent livré que quelques restes (ostéodermes et dent) d'Angleterre décrits sous le nom d'Acanthopholis horridus (Huxley, 1867), auxquels s'ajoute la dent isolée provenant de Touraine mentionnée plus haut (Buffetaut & Brignon, 1999). Au sein des nodosauridés, la forme européenne la mieux connue correspond au genre Struthiosaurus, du Campanien-Maastricthien de France, d'Espagne, d'Autriche et de Roumanie (Astibia et al., 1990 ;Pereda-Suberbiola, 1992 ;Buffetaut et al., 1997 ;Garcia & Pereda-Suberbiola, 2003 ;Vickaryous et al., 2004). Les dents de Struthiosaurus sont d'ailleurs assez proches de nos spécimens (cf. ...
... Les dents de Struthiosaurus sont d'ailleurs assez proches de nos spécimens (cf. Garcia & Pereda-Suberbiola, 2003 ; fig. 4a Néraudeau et al. (2003). ...
M. François Guillocheau, Univ. Rennes (Président)
Mme Angela D. Buscalioni, Univ. Madrid (Rapporteur)
M. Jean-Claude Rage, MNHN, Paris (Rapporteur)
M. Jean-Michel Mazin, Univ. Lyon 1 (Examinateur)
M. Henri Cappetta, Univ. Montpellier 2 (Co-directeur de thèse)
M. Didier Néraudeau, Univ. Rennes 1 (Directeur de thèse)
... Ankylosaur remains from the Upper Cretaceous of Europe are relatively poorly known and up to now only two valid genera have been identified. Struthiosaurus Bunzel, 1871 is widespread in Europe, having been reported from several important continental vertebrate localities of early CampanianeMaastrichtian age (e.g., Seeley, 1881;Nopcsa, 1929;Pereda-Suberbiola, 1992, 1999Buffetaut et al., 1996;Garcia and Pereda-Suberbiola, 2003;Codrea et al., 2010). Three currently valid species of this genus are known. ...
... In stratigraphical order, S. austriacus Bunzel, 1871 was described from the lower Campanian Gosau beds of Austria based on some cranial and mainly postcranial material of at least three individuals of different ontogenetic stages (Pereda-Suberbiola and Galton, 2001). Similarly aged, early Campanian Struthiosaurus languedocensis Garcia and Pereda-Suberbiola, 2003 from Villeveyrac, southern France was described on the basis of an articulated skeleton consisting of the pelvic girdle, synsacrum and posterior dorsal vertebrae. The youngest species is S. transylvanicus Nopcsa, 1915 from the Maastrichtian deposits of the Hat¸eg Basin, Romania that is known on the basis of most probably associated cranial and postcranial material (Nopcsa, 1929). ...
... (MCNA 7420.4) (Garcia and Pereda-Suberbiola 2003). The distal shaft is expanded with a distinct triangular bulge anteriorly. ...
Huaxiazhoulong shouwen gen. et sp. nov. is a new ankylosaurid recovered from the Upper Cretaceous Tangbian Formation of Jiangxi Province, southern China. Huaxiazhoulong shouwen can be diagnosed on the basis of three autapomorphies (the middle shaft and distal end of the ischium are expanded; the ratio of width of distal end to minimum shaft width is greater than 3 in humerus, the maximum length of femur to humerus length ratio is about 1.45) and a unique combination of characters (the centra of anterior caudal vertebrae in anterior view is heart-shaped; the dorsal surface of scapula is straight; the scapulocoracoid has a large medial brace; the humeral head and deltopectoral crest are separated by a distinct notch anteriorly). The phylogenetic analysis shows that Huaxiazhoulong shouwen is an early member of Ankylosauridae. ARTICLE HISTORY
... Similarly, diversity of the European Lambeosaurinae may be high, with a wide range of jaw morphologies and species occurring, because hadrosaurids in Europe had limited competition, with relatively few other ornithischian lineages present. These included rhabdodontid iguanodontians 94 , basal hadrosauroids [95][96][97] , and struthiosaurine ankylosaurs 98,99 . North American lambeosaurines competed with multiple lineages of hadrosaurids as well as other ornithischians such as thescelosaurids, pachycephalosaurids, leptoceratopsids, nodosaurids and ankylosaurids, as well as herbivorous coelurosaurs such as ornithomimids, deinocheirids, caenagnathids, and possibly troodontids 100 . ...
In the Late Cretaceous, northern and southern hemispheres evolved distinct dinosaurian faunas. Titanosaurians and abelisaurids dominated the Gondwanan continents; hadrosaurids, ceratopsians and tyrannosaurs dominated North America and Asia. Recently, a lambeosaurine hadrosaurid, Ajnabia odysseus, was reported from the late Maastrichtian phosphates of the Oulad Abdoun Basin Morocco, suggesting dispersal between Laurasia and Gondwana. Here we report new fossils from the phosphates of Morocco showing lambeosaurines achieved high diversity in the late Maastrichtian of North Africa. A skull represents a new dwarf lambeosaurine, Minqaria bata. Minqaria resembles Ajnabia odysseus in size, but differs in the ventrally positioned jugal facet and sinusoidal toothrow. The animal is small, ~ 3.5 m long, but the fused braincase shows it was mature. A humerus and a femur belong to larger hadrosaurids, ~ 6 m long, implying at least three species coexisted. The diversity of hadrosaurids in Europe and Africa suggests a dispersal-driven radiation, with lambeosaurines diversifying to take advantage of low ornithischian diversity. African lambeosaurines are small compared to North American and Asia hadrosaurids however, perhaps due to competition with titanosaurians. Hadrosaurids are unknown from eastern Africa, suggesting Moroccan hadrosaurids may be part of a distinct insular fauna, and represent an island radiation.
... In southern France, several dinosaur-bearing Campanian localities are known, mainly in Bouches-du-Rhône, Var and Hérault. These levels have yielded a diverse dinosaur fauna that comprises ankylosaurs, ornithopods, sauropods, non-avian theropods, and birds (e.g., [5,[55][56][57]). The fossil record of non-avian theropod dinosaurs is represented by several cranial and postcranial materials mostly assigned to abelisaurids (e.g., [5]). ...
The Lo Hueco fossil site (Cuenca, Spain) is one of the most relevant localities for the study of Late Cretaceous continental vertebrate faunas from Europe. The fossil record of theropod dinosaurs from this locality is represented by scarce isolated postcranial materials that were preliminarily attributed to abelisaurids and to a possible giant bird, in addition to a large assemblage of isolated teeth that were related to different maniraptoran clades. Here, we describe an isolated partial left tibia articulated with the proximal tarsals and discuss their taxonomic affinities. A review of the European fossil record of Late Cretaceous theropods was performed to analyze possible changes in the faunistical composition during this period. The specimen from Lo Hueco exhibits some characters that have been interpreted as apomorphies for maniraptoran coelurosaurs and a combination of features compatible with deinonychosaurians. Within this clade, the specimen is more favorably comparable with velociraptorine dromaeosaurids and is tentatively interpreted as a member of this group. This specimen is one of the few non-dental specimens of dromaeosaurids described thus far from the Upper Cretaceous of the Iberian Peninsula and contributes to a better understanding of the composition and evolutionary history of the European theropod fauna during the last stages of the Mesozoic.
... The three species of the nodosaurid Struthiosaurus (S. austriacus from the lower Campanian of Austria, Pereda-Suberbiola and Galton, 2001; S. languedocensis from the Campanian-Maastrichtian of France, García and Pereda-Suberbiola, 2003; and S. transylvanicus from the Maastrichtian of Romania, Nopcsa, 1915) display relatively small body sizes. However, these smaller body sizes may be explained by the basal position of Struthiosaurus spp. ...
Pararhabdodon isonensis was the first species of lambeosaurine hadrosaurid described in Europe and is one of the last non-avian dinosaurs that lived before the K-Pg extinction. Yet, its relationship with other Ibero-Armorican lambeosaurines has remained controversial due to the lack of overlapping diagnostic material among taxa. Newfound hindlimb materials reveal a unique character for the species that reinforces its distinction from other European lambeosaurines and its postulated close relationship with Tsintaosaurus spinorhinus. P. isonensis becomes restricted to the upper Maastrichtian Talarn Formation.
Our osteohistological analysis indicates that Pararhabdodon isonensis probably reached adult body sizes comparable to those of other Ibero-Armorican lambeosaurines and nearing the body sizes of North American and Asian taxa. Its histomorphology indicates a relatively low growth rate, suggesting the achievement of larger body sizes over longer time periods, perhaps facilitated by a relatively low predation pressure. Unlike coeval dinosaurian clades of the Late Cretaceous European Archipelago, P. isonensis and at least some of the other Ibero-Armorican lambeosaurines that reached similarly large body sizes, like Adynomosaurus arcanus, represent exceptions to the ‘island rule’.
... Coombs (1971) reported that in Euoplocephalus rib fusion to dorsal vertebrae occurs as early as the second dorsal in the series. It is also the case that in ankylosaurs the last three (Sauropelta; Ostrom, 1970) to six (Struthiosaurus; Garcia & Pereda Suberbiola, 2003) dorsal vertebrae become fused together to form a presacral rod. Centra and zygapophyses are fused in the presacral rod (and, of course, the accompanying ribs), but the neural spines tend to vary. ...
Scelidosaurus fossils were first discovered during the commercial quarrying of the Liassic sea-cliffs between Charmouth and Lyme Regis in Dorset during the late 1850s. The original specimens included a well-preserved skull embedded in a block of argillaceous limestone (marlstone). Shortly after this skull was retrieved, a series of more-or-less contiguous marlstone slabs were recovered, containing most of the skeleton of the same animal (NHMUK R1111). After rudimentary (hammer and chisel) mechanical preparation, Owen published descriptions of this material (Owen, 1861, 1863). These two monographs have been the sole references pertaining to the anatomy of Scelidosaurus for >150 years. The skeleton of the lectotype of Scelidosaurus harrisonii (NHMUK R1111) has since been extracted from the surrounding matrix using an acid-immersion technique. Some additional specimens held in the collections of the Natural History Museum London, the Bristol City Museum and the Sedgwick Museum in Cambridge provide anatomical material that allows detailed description of this taxon, for which we have had, until now, a surprisingly poor understanding.
Axial skeleton: The axial skeleton of Scelidosaurus comprises eight cervical, 16 dorsal, four sacral and > 40 caudal vertebrae. During ontogeny, the posterior centrum articular surface of the 16th dorsal vertebra develops a firm, ligament-bonded junction with the succeeding sacral centrum. Apart from the atlas rib, which is single headed, double-headed ribs are present throughout the presacral vertebral series, and none shows any indication of fusion to its associated vertebra. However, those ribs attached to cervical vertebrae 2–4 were evidently bound firmly by connective tissue to rugose diapophyses. The last two (presacral) dorsal ribs show merger of the capitulum and tuberculum, meaning that they are separated by only a step. The angulation and arching of the dorsal ribs suggest that these animals had a broad (barrel-like) torso. Intercostal uncinate plates were present, attached to the posterior margins of some of the largest dorsal ribs. Their attachment sites are clearly marked, and these plates might have been composed of calcified cartilage in larger individuals. The sacral vertebrae fuse progressively during ontogeny, in an anterior-to-posterior sequence. The sacral ribs are long and robust, and tilt the iliac blade outward dorsally. A sacricostal ‘yoke’ (created by the fusion of the distal ends of adjacent sacral ribs) never forms. The base of the tail has a unique ball-and-socket-style joint between the centra of caudal vertebrae 1 and 2 in only one skeleton. This might have permitted powerful, but controlled, movements of the tail as a defensive weapon (or increased flexibility at the base of the tail, which might have been necessary for reproduction). Caudal ribs are initially long, blade-shaped projections that gradually decrease in size and become stub-like remnants that persist as far back as the midtail (approximately caudal vertebra 25). Haemal arches (chevrons) disappear nearer to the distal end of the tail (approximately caudal vertebra 35). Ossified tendons are preserved as epaxial bundles that are clustered in the ‘axillary’ trough (between the neural spine and transverse processes on either side of the midline). Ossified tendons are restricted to the dorsal and sacral region. Flattened ossified tendons are fused to the sides of sacral neural spines. In life, the ossified tendons might have formed a low-angled trellis-like arrangement.
Appendicular skeleton: The pectoral girdle comprises a long scapula, with a distally expanded blade. The proximal portion is expanded and supports an oblique promontory, forming an acromial process anteriorly and a thick, collar-like structure posteriorly above the glenoid. Between these two features is a shallow basin, bordered ventrally by a sutural edge for the coracoid. The scapula–coracoid suture remains unfused in large (5-m-long) individuals. The coracoid bears a discrete foramen and forms a subcircular dished plate, with the shallowest of embayments along its posterior edge. Clavicles are present as small fusiform bones attached to the acromial process of the scapulae and leading edge of each coracoid. A sternum was reported as ‘some partially ossified element of the endoskeleton’ Owen (1863: 13), but subsequent preparation of the skeleton has removed all trace of this material. The humerus is relatively long and has a prominent rectangular and proximally positioned deltopectoral crest. The ulna is robust and tapers distally, but there is no evidence of an olecranon process. The radius is more rod-like and terminates distally in an enlarged, subcircular and convex articular surface for the carpus. The carpus is represented by an array of five discoid carpals. The manus is pentadactyl and asymmetrical, with short, divergent metacarpals and digits that terminate in small, arched and pointed unguals on digits 1–3 (only). The phalangeal formula of the manus is 2-3-4-3-2. The pelvis is dominated by a long ilium; the preacetabular process is arched, transversely broad, and curves laterally. In juveniles, this process is short and horizontal, but during ontogeny it increases considerably in length and becomes arched. The iliac blade is tilted laterally, meaning that its dorsal blade partly overhangs the femur. The acetabulum forms a partial cupola, and there is a curtain-like medial wall that reduces the acetabular fenestra to a comparatively low, triangular opening between the pubis and ischium. The postacetabular portion of the ilium is long and supports a brevis shelf. The ischium has a long, laterally compressed shaft that hangs almost vertically beneath the ilium, and there is no obturator process. The pubis has a long, narrow shaft and a relatively short, deep, laterally compressed prepubic process that twists laterally (its distal end lies almost perpendicular to the long axis of the ilium). The articular pad on the pubis for the femoral head faces posteriorly. The obturator foramen is not fully enclosed within the pubis, but its foramen is closed off posteriorly by the pubic peduncle of the ischium. The femur is stout and has a slightly medially offset femoral head, and the greater trochanter forms a sloping shoulder continuous with, and lateral to, the femoral head. The anterior (lesser) trochanter is prominent and forms a thick, thumb-shaped projection on the anterolateral corner of the femoral shaft. The fourth trochanter is pendent and positioned at midshaft. In larger individuals, it appears to become thickened and reinforced by becoming coated with metaplastic bone derived from the tendons attached to its surface. The distal end of the femoral shaft is slightly curved and expands to form condyles. There is a deep and broad posterior intercondylar groove, but the anterior intercondylar groove is barely discernible in juveniles and not much better developed in subadults. The tibia and fibula are shorter than the femur. The tibia is structurally dominant, and the shorter fibula is comparatively slender and bowed. The proximal tarsals are firmly bound by connective tissue to the distal ends of the tibia and fibula. The distal end of the tibia is stepped, which aids the firm interlock between the crus and proximal tarsals. There appear to be two roughly discoid tarsals (distal tarsals 3 and 4), and a rudiment of distal tarsal 5 appears to be sutured to the lateral margin of distal tarsal 4. Five metatarsals are preserved, but the fifth is a splint of bone attached to the proximal end of metatarsal 4. Metatarsals 2–4 are dominant, long and are syndesmotically interlocked proximally, but their shafts splay apart distally. Metatarsal 1 is much shorter than the other three, but it retains two functional phalanges (including a short, pointed ungual). The foot is anatomically tetradactyl but functionally tridactyl. The pedal digit formula is 2-3-4-5-0. The digits diverge, but each appears to curve medially along its length, creating the impression of asymmetry. This asymmetry is emphasized, because the three principal unguals are also twisted medially. The ungual of digit 2 is the largest and most robust of the three, whereas that of digit 4 is the smallest and least robust.
The general girth of the torso and the displacement of the abdomen posteriorly (a consequence of the opisthopubic pelvic construction in this dinosaur) constrained the excursion of the hindlimb during the protraction phase of the locomotor cycle. The anterolateral displacement of the hindlimb during protraction is in accord with the freedom of motion that is evident at the acetabulum, the susceptibility of the hindlimb to torsion between and within its component parts, and the asymmetry of the foot. It is probable that thyreophorans (notably, ankylosaurs) used a similar oblique-parasagittal hindlimb excursion to accommodate their equally large and wide abdomens. This surmise accords with the structure of the pelves and hindlimbs of ankylosaurs. Derived stegosaurs might have obviated this ‘problem’, in part, because their hindlimbs were longer and their torsos and abdomens narrower and capable of being ‘stretched’ vertically to a greater extent. Nevertheless, the structure of their acetabula and hindlimbs indicates that the oblique-parasagittal style of hindlimb excursion remained a possibility and might be an evolutionary remnant of the locomotor style of basal, shorter-limbed stegosaurs.
A reconstruction of the endoskeleton of Scelidosaurus is presented on the basis of this updated description. Although quadrupedal, this animal was only facultatively so, judged by its forelimb-to-hindlimb proportions and structure; it therefore betrays bipedality in its ancestry.
... Nodosaurids are the ankylosaurians commonly known in several fossil sites from Campanian-Maastrichtian formations of south-western Europe (Csiki-Sava et al., 2015) (Fig. 11). Garcia and Pereda-Suberbiola (2003) defined the species Struthiosaurus languedocensis in the lower Campanian of the Villeveyrac-Meze Basin (Hérault, France). Nevertheless, due to the lack of diagnostic features most of the nodosaurid remains cannot be identified beyond higher taxonomic levels (Pereda-Suberbiola, 1992;Laurent et al., 2001;Laurent, 2003;Buffetaut, 2005). ...
In south-western Europe, the uppermost Cretaceous continental deposits in the different sedimentary basins of Iberia (Portugal and north and central Spain), the Pyrenees (Spain and France) as well as Languedoc and Provence (southern France) provide one of the few terrestrial records that allow a comprehensive study of the Campanian- Maastrichtian dinosaur assemblages. For the last years the southern Pyrenees has been the target of intense geological, palaeontological and geochronological research. Hundreds of fossil localities are now framed in high-resolution lithological sections. The succession of these sites, most of them located in the Tremp Syncline, is based on the physical correlation of rock bodies, as permitted by the general outcropping conditions. Outside this syncline, correlation is supported by geochronologic and biostratigraphic data (mainly magnetostratigraphy and planktic foraminifera biostratigraphy). The integration of the entire dataset sheds new light on the Maastrichtian dinosaur turnover, characterized by a shift from a sauropod-dominated to a hadrosauroid-dominated faunal assemblage. This turnover was progressive and involved immigrants from North America, Eurasia and Gondwana, which probably reached the study area after a sea level drop. This faunal change was mainly triggered by the arrival of lambeosaurine hadrosauroids, a group that rapidly displaced the rest of the herbivorous clades of the region. Some of the extinction events suffered by the “preturnover” faunas during the Maastrichtian coincide with marine isotopic and sea-level drop events, suggesting that faunal competition was not the only cause of the observed changes in dinosaur composition. Despite this faunal replacement, the resulting ecosystem after the turnover shows no major loss of biodiversity before the Cretaceous- Paleogene extinction event.
... El registre de dinosaures del Campanià i del Maastrichtià trobat a França, Espanya i Portugal és un dels més rics d'Europa, i es troba entre els més importants del món pel que fa al Cretaci terminal. A partir dels anys 90 un gran nombre de treballs científics van permetre avançar en el camp de la taxonomia i diversitat, la paleobiogeografia i la successió faunística en aquesta regió (per exemple, entre molts altres, Le Loeuff , 1995Le Loeuff et al., 1994;Antunes i SigogneauRussell, 1992Casanovas et al., 1999a i b;; López-Martínez et al., 1999; LópezMartínez, 2003;Pereda-Suberbiola, 1999;Garcia et al., 1999Garcia et al., , 2010Garcia i Pereda-Suberbiola, 2003;Allain i Pereda-Suberbiola, 2003;Company et al., 2009;Valentin et al., 2012;Vila et al., 2012Vila et al., , 2016 Prieto- Csiki-Sava et al., 2015;Pérez-Garcia et al., 2016;Díez Díaz et al., 2016). Gràcies a tota aquesta recerca, s'han pogut identificar dues associacions de dinosaures que es van succeir durant el Cretaci terminal a l'illa Ibero-Armoricana: ...
http://hdl.handle.net/10803/402396
The terrestrial Campanian/Maastrichtian geological and paleontological record from southwestern Europe is one of the best outside North America to study the last 15 million years previous to the Cretaceous-Paleogene mass extinction. The narrative of the last dinosaur communities from the Ibero-Armorican Island (European Archipelago) is addressed by studying their occurrences, associated paleoenvironments and magnetostratigraphic dating.
Such integrated works in the Mesozoic portion of the Tremp Group (South-Pyrenean Basin) includes the study of footprints of the ichnogenus Hadrosauropodus, linked to hadrosaur dinosaurs. These tracks are abundant in the fluvial coastal plains of the upper Maastrichtian. In addition to the fluvial ecosystem, hadrosaurs also colonized the coastal environment (e.g. lagoons), as reveals the L’Espinau site (amongst other localities). In contrast, sauropods were dominant in the Campanian and the lower Maastrichtian coastal and inland environments of the region, but were still present in those settings until the uppermost Maastrichtian, as evidenced by bones, eggshell and tracks, and skin impressions. However, their remains were much scarcer than those of hadrosaurs at this age.
The dinosaur faunal succession of southwestern Europe has been improved by means of: 1) habitat understanding, 2) dating and integrating the Aude record (northern Pyrenees), 3) improving of the age calibration of the Isona sector (southern Pyrenees), and 4) integrating of the Campanian- Maastrichtian dinosaur fossil record from the rest of France, Spain and Portugal (Provence and Iberian areas). Dating refinements permit a new model for the Maastrichtian dinosaur faunal succession. Instead of a rapid faunal shift from titanosaurian-dominated herbivorous assemblages to hadrosaur-dominated communities around the early-late Maastrichtian boundary, the achieved data show that the extinction of major clades and the apparition of new ones took place diachronously and was not time coincident. On the contrary, a coexistence period of about two milion years between older and newer Ibero-Armorican dinosaur inhabitants have been identified.
... Finally, a mixed fauna emerged in Europe (Ezcurra and Agnolín, 2012) combining lineages with Laurasian affinities, including lambeosaurine hadrosaurids, with Gondwanan lineages such as Titanosauridae (Sanz et al., 1999;Díaz et al., 2012;Díaz et al., 2016), and endemics such as Rhabdodontidae (Allain and Pereda-Suberbiola, 2003;Weishampel et al., 2003;Osi et al., 2012) and struthiosaurine nodosaurids (Pereda- Suberbiola and Galton, 2001;Garcia and Pereda-Suberbiola, 2003;Ősi, 2005). As in the Gondwanan continents, abelisaurids were the apex predators in these faunas (Ősi and Buffetaut, 2011;Tortosa et al., 2014). ...
During the latest Cretaceous, distinct dinosaur faunas were found in Laurasia and Gondwana. Tyrannosaurids, hadrosaurids, and ceratopsians dominated in North America and Asia, while abelisaurids and titanosaurids dominated in South America, India, and Madagascar. Little is known about dinosaur faunas from the latest Cretaceous of Africa, however. Here, a new abelisaurid theropod, Chenanisaurus barbaricus, is described from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa on the basis of a partial dentary and isolated teeth. Chenanisaurus is both one of the largest abelisaurids, and one of the youngest known African dinosaurs. Along with previously reported titanosaurid remains, Chenanisaurus documents the persistence of a classic Gondwanan abelisaurid-titanosaurid fauna in mainland Africa until just prior to the end-Cretaceous mass extinction. The animal is unusual both in terms of its large size and the unusually short and robust jaw. Although it resembles South American carnotaurines in having a deep, bowed mandible, phylogenetic analysis suggests that Chenanisaurus may represent a lineage of abelisaurids that is distinct from those previously described from the latest Cretaceous of South America, Indo-Madagascar, and Europe, consistent with the hypothesis that the fragmentation of Gondwana led to the evolution of endemic dinosaur faunas during the Late Cretaceous.
... Nodosaurid teeth are more blade-like and usually more complex than the cusp-like teeth of ankylosaurids (Mallon & Anderson 2014a) in having a larger crown, a rough enamel surface, crenelated cingulum and fluting that is confluent with the grooves of the marginal cusps (Coombs 1990; Figure 6(C)). to S. languedocensis (Garcia & Suberbiola 2003) bear informative wear facets. The wear facet on UM2 OLD-19 CV, covering approximately the 30% of the crown, is steeply inclined (ca. ...
Ankylosaurian dinosaurs were low-browsing quadrupeds that were traditionally thought of as simple orthal pulpers exhibiting minimal tooth occlusion during feeding, as in many extant lizards. Recent studies, however, have demonstrated that effective chewing with tooth occlusion and palinal jaw movement was present in some members of this group. Qualitative and quantitative analysis of feeding characters (i.e. craniodental features, tooth wear patterns, origin and insertion of jaw adductors) reveal at least three different jaw mechanisms during the evolution of Ankylosauria. Whereas, in basal members, food processing was restricted to simple orthal pulping, in late Early and Late Cretaceous North American and European forms a precise tooth occlusion evolved convergently in many lineages (including nodosaurids and ankylosaurids) complemented by palinal power stroke. In contrast, Asian forms retained the primitive mode of feeding without any biphasal chewing, a phenomenon that might relate to the different types of vegetation consumed by these low-level feeders in different habitats on different landmasses. Further, a progressive widening of the muzzle is demonstrated both in Late Cretaceous North American and Asian ankylosaurs, and the width and general shape of the muzzle probably correlates with foraging time and food type, as in herbivorous mammals.
... However, this morphology is also found stegosaurs. Indeed, the ilium of Scelidosaurus has a larger posterior process and supraacetabular process than that of ankylosaurs (BMNH R1111; AMNH 5409; Coombs, 1978;Garcia and Pereda Suberbiola, 2003); both of these features are, in fact, very similar in Scelidosaurus and Huayangosaurus (ZDM T7001). Furthermore, the ilium of Scutellosaurus, the most basal thyreophoran , bears an incipient lateral expansion (MNA Pl.175; Colbert, 1981). ...
Previous descriptions of the postcranial skeleton of the primitive stegosaur Huayangosaurus taibaii (Middle Jurassic: People's Republic of China) are insufficient for character determinations in cladistic analysis. Reexamination of the postcranium has revealed several important characters, most of which are retained plesiomorphies that have not been identified previously. For example, Huayangosaurus retains ossified tendons, the loss of which has been used as a synapomorphy of Stegosauria. These purported stegosaurian synapomorphies now serve to unite the monophyletic clade Stegosauridae, defined as all stegosaurs more closely related to Stegosaurus than to Huayangosaurus. In addition, some anatomical characters previously used as ‘ankylosauromorph’ synapomorphies have now been identified in Huayangosaurus: this suggests that these characters occur more basally within Thyreophora. The plesiomorphic status of Huayangosaurus suggests that substantial convergence between ankylosaurs and stegosaurs occurred during their respective evolutionary histories, reflecting coincident functional morphological changes associated with the adoption of obligate quadrupedality.
... Los números entre corchetes hacen referencia a los mapas de la Figura 2. Se indica con un asterisco el material cuyas afinidades son controvertidas. Abreviaturas: Gr, Grupo; Fm., Formación.Struthiosaurus en el Santoniense-Maastrichtiense(Garcia y Pereda-Suberbiola, 2003;Ősi, 2005;Ősi et al., 2014).En el hemisferio sur, se han descrito restos esqueléticos de anquilosaurios en el Cretácico Superior de la Antártida, Argentina, Nueva Zelanda, Madagascar y la India ...
Thyreophora es un clado de dinosaurios ornitisquios que reúne a estegosaurios, anquilosaurios y formas basales como Scelidosaurus. Su registro fósil se extiende desde el Jurásico Inferior hasta el Cretácico Superior. Muchos de los fósiles de tireóforos descubiertos hasta la fecha provienen de yacimientos situados en el hemisferio norte. No obstante, el registro gondwánico comprende relevantes restos esqueléti- cos y/o icnitas en Sudamérica, África, Madagascar, Australia, Nueva Zelanda y la Antártida. Los tireóforos podrían estar representados en África desde el Jurásico Inferior–Medio. Se ha documentado su presencia en el Jurásico Superior de Tanzania (estegosaurio Kentrosaurus) y en el lí- mite Jurásico–Cretácico de Bolivia (huellas). Durante el Cretácico Temprano, los estegosaurios estuvieron presentes en Sudáfrica (Paranthodon) y la Argentina (forma indeterminada), y los anquilosaurios en Australia (Minmi). Los anquilosaurios también tienen registro en el Cretácico Superior de Sudamérica (restos esqueléticos en la Argentina y huellas en Bolivia), la Antártida (Antarctopelta), Nueva Zelanda y posiblemente Madagascar. La presencia de anquilosaurios y estegosaurios posibles en el Cretácico Superior de la India está sin confirmar. Desde un punto de vista paleobiogeográfico, los tireóforos gondwánicos parecen provenir de diferentes dispersiones desde Laurasia. Los estegosaurios afri- canos serían el testimonio de dos eventos de dispersión ocurridos durante el Jurásico Medio–Tardío. Los anquilosaurios gondwánicos tampoco resultan de una radiación única: Minmi podría representar un linaje relictual establecido en Australia durante el Jurásico antes de la dicotomía Nodosauridae-Ankylosauridae, mientras que los nodosáuridos de la Argentina y la Antártida serían el resultado de una o varias dispersiones desde América del Norte durante el Cretácico Tardío.
... Until the discovery of the fossil assemblage assigned to the species Hungarosurus tormai Ősi, 2005 from the Santonian of Iharkút, western Hun-gary, which includes disarticulated cranial bones (Ősi, 2005;Ősi and Makádi, 2009), mainly the type material of Struthiosaurus, Struthiosaurus austriacus Bunzel, 1870 (Gosau beds, Lower Campanian, Austria) and a second species, Struthiosaurus transylvanicus Nopcsa, 1915 (Sinpetru beds, Maastrichtian, Romania), were available to provide ŐSI, PEREDA SUBERBIOLA, & FÖLDES: SKULL OF HUNGAROSAURUS 2 information on the cranial morphology of European Late Cretaceous ankylosaurs. The ankylosaur material published in the last two decades from Campano-Maastrichtian sediments of southern France and northern Spain, including Struthiosaurus languedocensis Garcia and Pereda Suberbiola, 2003 ('Fuvelian' beds, Lower Campanian, France), is composed mainly of postcranial material (e.g., Pereda Suberbiola, 1993, 1999Garcia and Pereda Suberbiola, 2003), and the cranial elements are represented only by edentulous maxillary and dentary fragments and isolated teeth (Pereda Suberbiola, , 1999. A new partial skull from the Chera locality, in Valencia, eastern Spain is now under description (Company et al., 2009;work currently in progress). ...
A partial skull of ankylosaur from the Upper Cretaceous (Santonian) Csehbánya Formation in Iharkút and the endocranial cast taken from it are described. The morphology of the exoccipital, the elongated 'neck' region of the basioccipital, the shape of the occipital condyle, and the different flexure of the medulla relative to the forebrain unambiguously differentiate this specimen from the basicranium of Struthiosaurus, so it is assigned to Hungarosaurus sp. Whereas the endocranial cast reflects a brain generally similar to those of other ankylosaurs, the dorsally hypertrophied cerebellum (also present is Struthiosaurus transylvanicus) is quite unusual within the group suggesting a more sophisticated cerebral coordination of posture and movement, and perhaps a more cursorial locomotary habit than predicted for other ankylosaurs.
... Rhabdodontids are also present, to the exclusion of titanosaurians or hadrosaurians, in slightly older European faunas inhabiting low-lying alluvial plains and marshes of Hungary (Iharkút; } Osi et al., 2012b), Austria (Muthmannsdorf; Bunzel, 1871;} Osi et al., 2012a), and southern France (Villeveyrac; Buffetaut et al., 1996;Garcia and Pereda Suberbiola, 2003). However, at these sites rhabdodontids are always associated with, and even outnumbered by, basal nodosaurids, which is also true in the basal part of the Ş ard Formation, at Vurp ar Vremir, 2010;Brusatte et al., 2013b). ...
... The ilium in the majority of ankylosaurs has a straight or gently convex lateral edge in dorsal view. In contrast, the lateral edge of the holotype of Taohelong is strongly sigmoidal, a condition only present in a few other ankylosaurs, including Sauropelta edwardsorum (Coombs 1978), Struthiosaurus languedocensis and Struthiosaurus sp. from the Iberian peninsula (Garcia & Pereda Suberbiola 2003). Besides Taohelong, ankylosaurs with a sigmoidal lateral edge to the ilium are present only in the Late Cretaceous. ...
The Ankylosauria is a group of herbivorous, quadrupedal, armoured dinosaurs subdivided into at least two major clades, the Ankylosauridae and the Nodosauridae. The most derived members of Ankylosauridae had a unique tail club formed from modified, tightly interlocking distal caudal vertebrae and enlarged osteoderms that envelop the terminus of the tail. We review all known ankylosaurid species, as well as ankylosaurs of uncertain affinities, in order to conduct a revised phylogenetic analysis of the clade. The revised phylogenetic analysis resulted in a monophyletic Ankylosauridae consisting of Ahshislepelta, Aletopelta, Gastonia, Gobisaurus, Liaoningosaurus, Shamosaurus and a suite of derived ankylosaurids (Ankylosaurinae). There is convincing evidence for the presence of nodosaurids in Asia during the Early Cretaceous. In the mid Cretaceous, Asian nodosaurids were replaced by ankylosaurine ankylosaurids. Ankylosaurines migrated into North America from Asia between the Albian and Campanian, where they diversified into a clade of ankylosaurines, here named Ankylosaurini, characterized by arched snouts and numerous flat cranial caputegulae. There is no evidence for any ankylosaurids in Gondwana; Ankylosauridae appears to be completely restricted to Asia and North America. The genus Crichtonpelta gen. nov. is created, type species Crichtonsaurus benxiensis Lü et al.
... Among sauropods, titanosaurians are represented by at least three lithostrotian genera: Ampelosaurus, Lirainosaurus and Magyarosaurus [Huene, 1932;Le Loeuff, 1993, 2005aSanz et al., 1999;Upchurch et al., 2004]. "Hypselosaurus" is a nomen dubium [Le Loeuff, 1993] Hungarosaurus [Pereda Suberbiola and Galton, 2001;Garcia and Pereda Suberbiola, 2003;Ösi, 2005]. Ornithopods include both basal and derived iguanodontians. ...
The continental tetrapod assemblages from the Santonian-Maastrichtian of Europe consist of dinosaurs (theropods: Abelisauroidea, Alvarezsauridae, Dromaeosauridae, ?Oviraptorosauria, ?Troodontidae, and birds: Enantiornithes, basal Ornithurae; sauropods: Titanosauria; ankylosaurs: Nodosauridae; ornithopods: Hadrosauridae, Rhabdodontidae; and neoceratopsians), pterosaurs (Azhdarchidae), crocodyliforms (eusuchians: Alligatoroidea, Gavialoidea, ?Hylaeochampsidae; sebecosuchian-like ziphosuchians; and, probably, basal neosuchians), choristoderes (?Champsosauridae), squamates (lacertilians: Iguanidae s.l., Paramacellodidae, Polyglyphanodontinae, Varanoidea; snakes: Madtsoiidae; possible amphisbaenians), turtles (cryptodires: Chelydroidea, Kallokibotionidae, Solemydidae; pleurodires: Bothremydidae, Dortokidae), lissamphibians (Albanerpetontidae; anurans: Discoglossidae, Palaeobatrachidae; caudates: Batrachosauroididae, Salamandridae), and mammals (multituberculates: Kogaionidae, ?"Paracimexomys group"; marsupials: Herpetotheriidae; eutherians: "Zhelestidae"). The palaeobiogeographical affinities of the Late Cretaceous continental tetrapods of Europe are complex. The faunas are commonly considered to show a mixed pattern resulting from the addition of "Asiamerican" and Gondwanan forms to European taxa. Albanerpetontids, both paramacellodid and polyglyphonodontine lizards, and hadrosaurid dinosaurs are taxa with Palaeolaurasian (or, in some case, even Neopangean) affinities. Other forms, such as paleobatrachid and batrachosauroidid lissamphibians, solemydid turtles, alligatoroid crocodyliforms, and nodosaurid dinosaurs can be considered as Euramerican taxa. Kallokibotionid and dortokid turtles, rhabdodontid dinosaurs and kogaionid mammals are endemic to Europe. The Gondwanan taxa have been regarded as African immigrants that reached southern Europe via the Mediterranean Tethyan sill. Abelisaurid and titanosaurid dinosaurs, trematochampsid crocodyliforms, podocnemidid and bothremydid turtles, and boid and madtsoiid snakes constitute the basic pattern of the so-called "Eurogondwanan fauna". However, the validity of some of these taxa is a disputed matter (Titanosauridae, Trematochampsidae), and the presence of other taxa in the Late Cretaceous of Europe is based on controversial data (Boidae, Podocnemididae). Only Abelisauroidea, Madtsoiidae and Bothremydidae (and, yet for confirming, Sebecosuchia) provide evidence of interchanges between Africa and Europe. At least abelisauroids might have reached Europe before the Late Cretaceous. In conclusion, most of the continental tetrapod taxa from the latest Cretaceous of Europe show affinities with those of northern Hemisphere landmasses. Latest Cretaceous trans-Tethyan dispersal events between Africa and Europe remain poorly documented.
... tered elements, including maxilla and lower jaw remains, isolated teeth, vertebrae and ribs, limb bones, and osteoderms (Pereda Suberbiola, 1993a. These remains have been assigned to the ankylosaur Struthiosaurus on the basis of features observed in the lower jaw, synsacrum, pelvis, and dermal armour (Pereda Suberbiola et al., 1995;Garcia and Pereda Suberbiola, 2003). Due to minor differences relative to known species of Struthiosaurus from the Campanian-Maastrichtian of Europe, the Laño material is tentatively referred to as Struthiosaurus sp. ...
The vertebrate-bearing beds of the Laño quarry (Condado de Treviño) are among the most relevant sites from the Late Cretaceous of Europe. Geologically, Laño and the adjacent region are set on the southern limb of the South-Cantabrian Synclinorium (SE Basque-Cantabrian Region, northern Iberian Peninsula). The Laño sites were discovered in 1984; thousands of bones and teeth, including microfossils, have been collected during the prospection in the field and excavation campaigns. The vertebrate remains occur at two different stratigraphic horizons within a continental to shallow marine succession of Late Campanian-Maastrichtian age. The lower horizon contains the Laño 1 and Laño 2 sites, whereas the upper horizon contains the Albaina site. In the Laño sites, three fossiliferous beds (called L1A, L1B and L2) are known within an alluvial system composed mainly of fluvial sands and silts. The sedimentary structures are consistent with channel areas within an extensive braided river system. Based mainly on stratigraphic correlations, the fluvial beds of Laño are regarded as Late Campanian to Early Maastrichtian in age. These deposits have yielded a very diverse vertebrate assemblage, which consists of nearly
40 species, including actinopterygians, lissamphibians, lepidosaurs, turtles, crocodyliforms, dinosaurs, pterosaurs, and mammals. Seven genera and ten species have been erected to date in Laño. With reference to the marine vertebrate association of Albaina, it consists of at
least 37 species, including sharks and rays, actinopterygians, mosasaurids, and plesiosaurs. Two genera and species of rhinobatoids (family
indet.) and two new species of rhinobatids have been erected in Albaina. The fossil association indicates a Late (but not latest) Maastrichtian age. Recently, isolated turtle and dinosaur fossils have been discovered in the sublittoral beds of Albaina. The Laño quarry is one of the most noteworthy Campanian-Maastrichtian vertebrate localities of Europe by its taxonomic diversity, and provides useful information about the composition and affinities of both continental and marine vertebrate faunas from the latest Cretaceous of southwestern Europe.
... However, this taxon is now considered to be a nomen dubium (Pereda-Suberbiola and Barrett 1999) and therefore this material, along with the teeth, is referred to as Nodosauridae indeterminate. From various Late Cretaceous localities in Europe some teeth have been referred to the small nodosaurid dinosaur Struthiosaurus Bunzel, 1870, e.g., three isolated teeth (UM2 OLV-D18-20 CV) from the lower Campanian of Southern France (Garcia and Pereda-Suberbiola 2003). Finally, the holotype of the nodosaurid Hungarosaurus tormai Ősi, 2005 from the upper Santonian western Hungary comprises 21 teeth, with cranial and postcranial remains. ...
We present new nodosaurid teeth from the Valanginian of Bexhill, Sussex and the Barremian of the Isle of Wight, the first from the Lower Cretaceous of the United Kingdom. Teeth found during the mid-1800s from the Valanginian and ascribed to the nodosaurid Hylaeosaurus are probably from sauropod dinosaurs. The Isle of Wight tooth could possibly be referred to Polacanthus foxii, the teeth of which are unknown. These new English nodosaurid teeth are similar to those of North American and European Jurassic to Late Cretaceous nodosaurids, especially the American Gastonia, Texasetes, Mymoorapelta, Gargoyleosaurus, and the European Hungarosaurus.
... Ankylosaur remains are also known from several localities of late Campanian to early Maastrichtian age in Provence and Languedoc. The material from Quarante (Hérault) described by Nopcsa (1929) as Rhodanosaurus ludgunensis (usually considered a nomen dubium: Garcia and Pereda-Suberbiola 2003) belongs to that faunal complex. Newly discovered material from Cruzy (Hérault; Fig. 7F) may provide a better understanding of the ankylosaurs from that time interval. ...
The Late Cretaceous was a time of tremendous global change, as the final stages of the Age of Dinosaurs were shaped by climate and sea level fluctuations and witness to marked paleogeographic and faunal changes, before the end-Cretaceous bolide impact. The terrestrial fossil record of Late Cretaceous Europe is becoming increasingly better understood, based largely on intensive fieldwork over the past two decades, promising new insights into latest Cretaceous faunal evolution. We review the terrestrial Late Cretaceous record from Europe and discuss its importance for understanding the paleogeography, ecology, evolution, and extinction of land-dwelling vertebrates. We review the major Late Cretaceous faunas from Austria, Hungary, France, Spain, Portugal, and Romania, as well as more fragmentary records from elsewhere in Europe. We discuss the paleogeographic background and history of assembly of these faunas, and argue that they are comprised of an endemic ‘core’ supplemented with various immigration waves. These faunas lived on an island archipelago, and we describe how this insular setting led to ecological peculiarities such as low diversity, a preponderance of primitive taxa, and marked changes in morphology (particularly body size dwarfing). We conclude by discussing the importance of the European record in understanding the end-Cretaceous extinction and show that there is no clear evidence that dinosaurs or other groups were undergoing long-term declines in Europe prior to the bolide impact.
... Vertebrae and ribs are quite similar to other nodosaurid ankylosaurs and they strongly resemble those of Struthiosaurus spp. (Pereda- Suberbiola and Galton, 2001;Garcia and Pereda-Suberbiola, 2003). Except for Minmi paravertebra (Molnar and Frey, 1987), Hungarosaurus is the only ankylosaur known to have paravertebral elements. ...
The composition of the Late Cretaceous (Santonian) continental vertebrate
fauna of Iharkút (Csehbánya Formation, Bakony, western Hungary, Central
Europe) is reviewed here. In the last decade, fieldwork has produced almost
5,000 associated and isolated bones and teeth belonging to at least 24 different
genera, represented by pycnodontiform and lepisosteid fishes, albanerpetontid
and anuran amphibians, dortokid, bothremydid and cryptodiran
turtles, scincomorphan and mosasauroid lizards, mesoeucrocodylian and
eusuchian crocodilians, nodosaurid ankylosaurs, rhabdodontid ornithopods,
basal tetanuran, abelisaurid, paravian, and enantionthine theropods, and
azhdarchid pterosaurs. Remains of mammals are still unknown from the
locality. Because of its Santonian age, the discovered fauna fills an important
and still underrepresented temporal gap in the Cretaceous vertebrate record
of Europe. The fauna is a mixture of Euramerican and Gondwanan forms.
The first group consists of numerous taxa (e.g., hylaeochampsid crocodilians,
nodosaurid ankylosaurs, rhabdodontid ornithopods, basal tetanurans),
the closest relatives of which are stratigraphically much older (Late Jurassic–
late Early Cretaceous) forms. These members of the fauna are suggested
to be relict forms surviving in an insular habitat of the Iharkút area
within the western Tethyan archipelago. At least bothremydid turtles further
strengthen the immigration of Gondwanan forms into Europe during the
Late Cretaceous. The supposed insular habitat of the Iharkút fauna is also
supported by the presence of a peculiar small-bodied heterodont crocodilian
with specialized feeding preference, and of pycondontiform fishes and
mosasaurs that colonized freshwater environments.
... JOURNAL OF PALEONTOLOGY, V. 88, NO. 3, 2014al., 2008aMartin, 2010;Martin and Delfino, 2010;Puértolas-Pascual et al., 2013) and Acynodon (Buscalioni et al., 1997(Buscalioni et al., , 1999Martin et al., 2006;Martin, 2007;Delfino et al., 2008b;Martin and Delfino, 2010), or the basal nodosaurid dinosaur Struthiosaurus (Nopcsa, 1929;Pereda-Suberbiola, 1999;Garcia and Pereda-Suberbiola, 2003;} Osi and Prondvai, 2013). Such coherent distributional patterns suggest a similar paleobiogeographic history of these taxa. ...
We present a detailed morphological description of the type-locality cranial material of Theriosuchus sympiestodon Martin, Rabi, and Csiki, 2010 from the Maastrichtian Densus -Ciula Formation of the Hat xeg Basin, Romania together with new material of isolated cranial elements and teeth from various sites of the same general area. The recognition of several individuals of distinct sizes allows for an assessment of ontogenetic variation in this taxon. New material, consisting of isolated teeth and an incomplete maxilla with in situ teeth, coming from various late Campanian/ early Maastrichtian sites in southern France is referable to ?Theriosuchus sp. and hints to a rare but widespread distribution of Theriosuchus in the Late Cretaceous European archipelago. INTRODUCTION
... In the paleogeographic region of southwestern Europe, the so-called Ibero-Armorican domain, of particular interest are the works assessing the diversity of the most abundant latest Cretaceous dinosaur taxa: the hadrosauroids (Prieto-Marquez et al., 2006;Pereda-Suberbiola et al., 2009a,b;Cruzado-Caballero et al., 2010;Prieto-Márquez et al., 2013) and sauropods (Vila et al., 2012;Díez Díaz et al., 2013). Conversely, other contemporaneous groups of dinosaurs reported from the uppermost Cretaceous (Campanian and Maastrichtian) of this region such as rhabdodontid ornithopods, nodosaurid ankylosaurs, and theropods (Allain and Pereda-Suberbiola, 2003;Garcia and Pereda-Suberbiola, 2003;Riera et al., 2009;Torices et al., 2013) have received less attention. ...
The scarcity of diagnostic skeletal elements in the latest Cretaceous theropod record of the Ibero-
Armorican domain (southwestern Europe) prevents to perform accurate phylogenetic, paleobiogeographic,
and diversity studies. In contrast, eggs and eggshells of theropod dinosaurs are relatively
abundant and well known in this region from which several ootaxa have been described. Here, we
describe the first Late Maastrichtian theropod ootaxon (Prismatoolithus trempii oosp. nov.) from SW
Europe and demonstrate that oological record can be used as a proxy for assessing diversity of eggproducers
and may help to complement their scarce bone record. The performed analyses indicate
that the theropod taxa and ootaxa reach their diversity maxima during the Late Campanian and start to
decrease near the CampanianeMaastrichtian boundary at both global and regional scales. The oological
diversity of theropods in the Ibero-Armorican domain is consistent with the theropod diversity identified
at high taxonomic level. Two distinct assemblages of theropod ootaxa can be recognized in the latest
Cretaceous of the Ibero-Armorican domain. Their temporal transition can be correlated with other
dinosaur faunal changes recorded in the region. This faunal turnover took place around the EarlyeLate
Maastrichtian boundary, involving ornithopods, sauropods, ankylosaurs and, according to the present
results, theropods as well.
... Rhabdodontids are also present, to the exclusion of titanosaurians or hadrosaurians, in slightly older European faunas inhabiting low-lying alluvial plains and marshes of Hungary (Iharkút; } Osi et al., 2012b), Austria (Muthmannsdorf; Bunzel, 1871;} Osi et al., 2012a), and southern France (Villeveyrac; Buffetaut et al., 1996;Garcia and Pereda Suberbiola, 2003). However, at these sites rhabdodontids are always associated with, and even outnumbered by, basal nodosaurids, which is also true in the basal part of the Ş ard Formation, at Vurp ar Vremir, 2010;Brusatte et al., 2013b). ...
The Transylvanian region of Romania preserves some of the most unusual and iconic dinosaurs in the
global fossil record, including dwarfed herbivores and aberrant carnivores that lived during the very
latest Cretaceous (Maastrichtian) in an ancient island ecosystem (the Hat¸ eg Island). A series of artificial
outcrops recently exposed during a hydroelectric project, the Petres¸ ti-Arini section near Sebes¸ in the
Transylvanian Basin, records a 400þ meter sequence documenting the transition from fully marine to
terrestrial environments during the CampanianeMaastrichtian. Calcareous nannofossil biostratigraphy
indicates that the lower marine beds in this section, part of the uppermost Bozes¸ Formation, can be
assigned to the CC22 biozone, corresponding to the loweremid upper Campanian. These beds smoothly
transition, via a brackish-water unit, into the fully continental Maastrichtian Sebes¸ Formation. Dinosaur
and pterosaur fossils from the uppermost Bozes¸ Formation can be assigned a late Campanian age making
them the oldest well-dated terrestrial fossils from the Hat¸ eg Island, and indicating that the classic Hat¸ eg
dinosaur fauna was becoming established by this time, coincident with the first emergence of widespread
land areas. Vertebrate fossils occur throughout the overlying Sebes¸ Formation at the site and are
dominated by the small-bodied herbivorous dinosaur Zalmoxes. The dominance of Zalmoxes, and the
absence of many taxa commonly seen elsewhere in Maastrichtian sites in Romania, suggests the possibility
that either the Petres¸ ti-Arini section preserves a somewhat unusual near-shore environment, or
the earliest Hat¸ eg Island dinosaur communities were structured differently from the more diverse
communities later in the Maastrichtian. Alternatively, due to the limited sample size available from the
studied succession, it is also conceivable that sampling biases give an incomplete portrayal of the Petres
¸ ti-Arini local fauna. Support for any one of these alternative hypotheses requires further data from
Petres¸ ti-Arini as well as from the larger Transylvania area.
... The first specimen (MA BZN 5) is very similar to the broad diamond-shaped tooth referred to an indeterminate nodosaurid by Buffetaut and Brignon (1999) from the Cenomanian of Touraine, western France. It is also similar to a tooth from the Campanian of southern France referred to Struthiosaurus by Garcia and Pereda Suberbiola (2003), a nodosaurid that is widely distributed in the Campanian-Maastrichtian deposits of southern Europe (Pereda Suberbiola, 1992;Vickaryous et al., 2004). The second specimen (MA BZN 6), which is taller, recurved and bears a stronger labial cingulum, is more reminiscent of some asymmetrical teeth of Edmontonia (Campanian-Maastrichtian: North America) figured by Carpenter (1990) and Coombs (1990Coombs ( , 1995. ...
The teeth of six dinosaur taxa (Carcharodontosauridae indet., Dromaeosauridae indet., Troodontidae indet., Brachiosauridae indet., Iguanodontoidea indet., and Nodosauridae indet.) are identified and described from the early Cenomanian of the Charentes region, western France. The composition and paleoecology of this coastal, insular fauna is discussed. The Charentes dinosaur assemblage shares affinities with Asiamerican and Gondwanan faunas. This clarifies and highlights the role of the European islands in the paleobiogeography of Cretaceous dinosaurs.
... The discovery of nodosaurid remains in one site dated as Late Maastrichtian extends their range (Laurent et al., 1999). As with sauropods, the restriction of nodosaurid remains to the grey unit may be a palaeoenvironmental bias caused by habitat preference, i.e., they lived mainly in coastal settings (Pereda-Suberbiola, 1992;Coombs and Deméré, 1996;Garcia and Pereda-Suberbiola, 2003; see also Butler and Barrett, 2008). ...
The succession of Maastrichtian deposits in the Tremp basin (South Pyrenean area, Spain) has provided long sections, with numerous dinosaur sites found in the Arén and Tremp Formations. The dinosaur record consists on titanosaur sauropods, dromaeosaurid theropods, hadrosaur ornithopods (including lambeosaurines) and nodosaurid ankylosaurians. Correlation of the sections enabled a clear succession of dinosaur localities to be established and thereby the succession of dinosaur faunas. Titanosaur remains, are present throughout the series, but are particularly abundant in the basal strata. Theropods and hadrosaur ornithopods are found all through the series. Lambeosaurine hadrosaur remains are restricted to sandstone channel beds close to the K/T boundary. The palaeoenvironmental conditions in each site were deduced from the sedimentological study, which allows establishing a relationship between environments and dinosaur taxa, and thus the over- or under-representation of taxa in the vertical succession to be unravelled. Titanosaurs and ornithopods are common in (but not exclusive to) marine-to-continental transitional and continental environments, respectively. Ankylosaurians are so far restricted to marine-to-continental transitional environments, but the sample is scarce. The fossil richness and the possibility of correlations among the different local sections make the Tremp Basin one of the best places in Europe to understand the final history on non-avian dinosaurs. The fossil record from the Tremp Basin suggests that they were well-diversified during the whole Maastrichtian until their extinction (as observed in North America and Asia). Sauropods were still present at the end of the Maastrichtian and lambeosaurines occurred at least in the upper part of the Maastrichtian.
... During the Late Campanian and Early Maastrichtian, rather few non-sauropod large herbivores are known however: the ornithopods Rhabdodon priscus (Matheron, 1869) and Rhabdodon septimanicus Le Loeuff, 1991b (cf. Chanthasit, 2010) as well as the ankylosaur Struthiosaurus languedocensis Garcia and Pereda-Suberbiola, 2003. Hadrosaurs, which became extremely abundant in the Late Maastrichtian (see Le Loeuff et al., 1994), may have been present too although they were apparently extremely rare before the Late Maastrichtian when up to five species of hadrosaurs may have coexisted on the Ibero-Armorican landmass according to Pereda-Suberbiola et al. (2009), together with at least two sauropods (Vila et al., 2009) 1 and an ankylosaur. ...
Recent discoveries in southern France and northern Spain suggest that the morphology of titanosaurian teeth shows much greater variations that previously thought. It is suggested that the different morphotypes are informative at specific or generic level and that titanosaurian genera may indeed be recognized by their isolated teeth. It is also confirmed that juvenile titanosaurian teeth have a rather uniform, cylindrical morphology. Four different morphotypes are described for the Ibero-Armorican Island in the Late Cretaceous.
The lower Albian track-bearing surface of the San Leonardo quarry (Molfetta, Apulia) is characterised by more than 800 footprints, produced by both quadrupedal and bipedal dinosaurs. Six well-preserved bipedal trackways, composed of tridactyl footprints are attributed to medium-to large-sized theropod dinosaurs. Only one clear but poorly preserved trackway and numerous isolated manus-pes couples have been attributed to quadrupedal dinosaurs. The tridactyl ichnoassemblage, analysed using both traditional methods and close-range photogrammetry, is represented by weakly mesaxonic and robust specimens. Morphological comparison with Upper Jurassic and Lower Cretaceous theropod tracks from surrounding areas, supported by morphometric analyses, points out a highest affinity with the specimens from Switzerland and North Africa. Nevertheless, a set of unique characters appears to justify the establishment of a new ichnospecies, Jurabrontes melphicticus. Additionally, the photogrammetric models of the quadrupedal trackway and four isolated manus-pes sets suggest they belong to the same morphotype, represented by asymmetrical tetradactyl pes and highly digitigrade tetra- or pentadactyl manus. These tracks share numerous morphological characters with both the ichnogenera Tetrapodosaurus and Metatetrapodus and thus can be attributed to a medium-sized ankylosaurian trackmaker.
Nodosauridae is a group of thyreophoran dinosaurs characterized by a collar of prominent osteoderms. In comparison to its sister group, the often club-tailed ankylosaurids, a different lifestyle of nodosaurids could be assumed based on their neuroanatomy and weaponry, e.g., regarding applied defensive strategies. The holotype of the nodosaurid Struthiosaurus austriacus consists of a single partial braincase from the Late Cretaceous of Austria. Since neuroanatomy is considered to be associated with ecological tendencies, we created digital models of the braincase based on micro-CT data. The cranial endocast of S. austriacus generally resembles those of its relatives. A network of vascular canals surrounding the brain cavity further supports special thermoregulatory adaptations within Ankylosauria. The horizontal orientation of the lateral semicircular canal independently confirms previous appraisals of head posture for S. austriacus and, hence, strengthens the usage of the LSC as proxy for habitual head posture in fossil tetrapods. The short anterior and angular lateral semicircular canals, combined with the relatively shortest dinosaurian cochlear duct known so far and the lack of a floccular recess suggest a rather inert lifestyle without the necessity of sophisticated senses for equilibrium and hearing in S. austriacus. These observations agree with an animal that adapted to a comparatively inactive lifestyle with limited social interactions.
The TuronianeConiacian continental fossil record in Europe is scarce. Here we present a new fossil assemblage of early Coniacian age that was systematically collected from the coal-bearing Gosau Group
of the Tiefengraben locality near St. Wolfgang, Austria. The diverse assemblage is composed of at least 60 taxa including sporomorphs and Normapolles-related pollen, seeds and leaves of angiosperms and
gymnosperms, charophytes, gastropods, bivalves, ostracods, termites, fishes, crocodiles and dinosaurs.
Concerning charophytes, ostracods, gastropods, crocodiles and dinosaurs, the discovered specimens either extend the temporal and spatial range of specific groups (in some cases as possible relict forms) or suggest the occurrence of new taxa. The discovered remains of algae, molluscs, ostracods, calcareous nannofossils and lepisosteid fish represent a mixed faunal assemblage from different palaeohabitats, from marginal marine to low salinity and freshwater or terrestrial environments. As Normapolles-related angiosperm plants dominate the flora with a relatively high number of dentate leaves, a slightly cooler microenvironment compared to other Turonian-Coniacian Central European localities is indicated. The characteristically grooved crocodylian teeth of Tiefengraben differ from the previously known Upper Cretaceous European crocodyliform teeth and suggest a more diverse crocodyliform fauna in the region.
Dinosaurs are represented by teeth of at least three different theropods, the largest of which is referred here to as basal tetanurans. The fossil assemblage of this early Gosau Group occurrence is of great importance for our understanding of the continental floristic and faunistic composition of the western Tethyan archipelago during the CenomanianeCampanian gap.
The Turonian–Coniacian continental fossil record in Europe is scarce. Here we present a new fossil assemblage of early Coniacian age that was systematically collected from the coal-bearing Gosau Group of the Tiefengraben locality near St. Wolfgang, Austria. The diverse assemblage is composed of at least 60 taxa including sporomorphs and Normapolles-related pollen, seeds and leaves of angiosperms and gymnosperms, charophytes, gastropods, bivalves, ostracods, termites, fishes, crocodiles and dinosaurs. Concerning charophytes, ostracods, gastropods, crocodiles and dinosaurs, the discovered specimens either extend the temporal and spatial range of specific groups (in some cases as possible relict forms) or suggest the occurrence of new taxa. The discovered remains of algae, molluscs, ostracods, calcareous nannofossils and lepisosteid fish represent a mixed faunal assemblage from different palaeohabitats, from marginal marine to low salinity and freshwater or terrestrial environments. As Normapolles-related angiosperm plants dominate the flora with a relatively high number of dentate leaves, a slightly cooler microenvironment compared to other Turonian–Coniacian Central European localities is indicated. The characteristically grooved crocodylian teeth of Tiefengraben differ from the previously known Upper Cretaceous European crocodyliform teeth and suggest a more diverse crocodyliform fauna in the region. Dinosaurs are represented by teeth of at least three different theropods, the largest of which is referred here to as basal tetanurans. The fossil assemblage of this early Gosau Group occurrence is of great importance for our understanding of the continental floristic and faunistic composition of the western Tethyan archipelago during the Cenomanian–Campanian gap.
A new, systematically collected vertebrate assemblage from the upper Turonian Schönleiten Formation of Gams bei Hieflau, Austria (Northern Calcareous Alps) is described here. The material consists of teeth and bones of chondrichthyan (Selachimorpha indet., Ptychotrygon gueveli) and osteichthyan fishes (cf. Lepisosteus sp., Vidalamiinae indet., Pycnodontidae indet.), amphibians (cf. Thaumastosaurus sp.), lizards (’Lacertilia’ indet.), mosasauroids (Tethysaurinae indet.), Theriosuchus-like and other, peculiar crocodyliforms and theropod dinosaurs (cf. Paronychodon sp.). The faunal composition is most similar to the Santonian fauna of Iharkút, western Hungary, a phenomenon that can be explained by the spatial and temporal closeness of the two landmasses. The Gams assemblage is of great importance since continental vertebrates from the Cenomanian to Santonian interval of Europe are extremely scarce, and it provides a small but significant link between older and younger vertebrate faunas in the region.
The polacanthid ankylosaur Gastonia burgei, from the Lower Cretaceous Cedar Mountain Formation, Utah, USA, is described in detail for the first time and compared with material from a stratigraphically higher monospecific Gastonia bone bed. Taphonomy and sedimentology of the bone bed place the site within the lower part of the Ruby Ranch Member and suggest mass mortality, either due to drought or drowning while crossing a swollen river. Burial of the scavenged and disarticulated bones was by crevasse splay. Gastonia is characterized by a skull that is subtriangular in dorsal view, being nearly as wide as long, a cranial surface with a pustulate texture, and anteroventrally projecting basipterygoid processes. In the postcrania, the scapula has a well-developed, arcuate acromion flange that attaches to the midshaft. The body armor includes laterally projecting tetrahedron osteoderms that are grooved along the posterior sides, dorsoventrally compressed triangular plates along the sides of the body, and coossified pelvic shield of large, raised osteoderms surrounded by rosettes of smaller osteoderms. A new species of Gastonia, G. lorriemcwhinneyae n. sp., is identified from the bone bed material. It differs from Gastonia burgei in having a flat skull roof, short paroccipital processes that is proportionally less expanded distally, short postacetabular process of ilium that is only 36% of the length of the preacetabular process as compared to 56% in G. burgei, and an ischium that is smoothly curved ventromedially without kink at its midpoint. Its discovery adds a new species of ankylosaur to the Polacanthidae family found in the Cedar Mountain Formation.
Here we present the magnetostratigraphic dating of the Laño locality (Condado de Treviño, northern Iberian Peninsula), one of the most noteworthy Campanian-Maastrichtian vertebrate sites of Europe. A composite section of 75 m thickness (Laño quarry) constructed from multiple, overlapping profiles and a continuous one (Faido) have been sampled for magnetostratigraphy. Thermal demagnetization techniques were systematically applied to 161 standard specimens and allowed characterizing the characteristic remanent magnetism, mostly carried out by magnetite. The palaeomagnetic signal is slightly scattered due to variety of lithologies, but the primary character can be guaranteed, since the normal and reverse directions are pseudo antiparallel; 346, 28 (α 95: 11.9°, k: 5.3) and 175, -35 (α 95: 16.4°, k: 4.6). Reliable samples allowed us to build the local polarity sequence made of eight magnetozones that has been used to correlate to the Global Polarity Time Scale. The age of the lower part of the Laño-village succession is basal late Campanian (Hoplitoplacenticeras marroti ammonite zone) and fits with the long reversed zone that must correlate to Chron C33r. The pattern of magnetozones allows tracking the section up to C30r at the upper part of the profile. In this correlation, the Laño vertebrate site is regarded as latest Campanian in age as it falls within the C32n (≈ 72–73.5 Ma). The combined lithostratigraphic and magnetostratigraphic analyses have yielded additional conclusions regarding the vertebrate assemblages that are representative of the Late Campanian of the Iberian Peninsula, in addition to highlight an older occurrence in Europe of some vertebrate groups such as salamandrid lissamphians and anguid lizards (or amphisbaenians).
The Zorralbo locality of the eastern Cameros Basin, near Soria, Spain, has produced a diverse dinosaur assemblage from the Lower Cretaceous Golmayo Formation. Ankylosaurs are represented by dorsal vertebrae and ribs, a fragmentary sacrum and ilium, and several types of dermal armour. Most, if not all, of the material probably belongs to a single medium to large-sized adult individual. The Soria remains are referred to Polacanthus on the basis of the presence of conical, ungrooved presacral spines, a sacropelvic shield composed of irregularly arranged bosses and small tubercles, large spined plates, and hollow-based triangular caudal plates with an extended posterior basal edge and a pointed spine. Polacanthus is well known from the Wealden Group (Barremian-Aptian) of the Isle of Wight and from the Weald Clay Group of West Sussex (England). In addition, isolated remains have been reported from the penicontemporaneous formations of the Iberian Peninsula. The Soria outcrop is currently the most productive Polacanthus site outside England. Moreover, it has yielded the oldest record (late Hauterivian to basal Barremian according to charophyte association) of this ankylosaur known to date in Europe. Minor anatomic differences between the Soria material and the taxa P. foxii (type-species) and P. rudgwickensis suggest the presence of a third species of Polacanthus in the Iberian Peninsula, but additional material is needed to confirm this interpretation.
A new titanosaur, Atsinganosaurus velauciensis, gen. and sp. nov. is described from well-preserved remains from the new Upper Cretaceous locality of Velaux-La Bastide Neuve (Aix-en-Provence Basin, France). This taxon is mainly diagnosed by a combination of characters, which differentiates it without ambiguity from other European Late Cretaceous taxa (Lirainosaurus, Ampelosaurus and Magyarosaurus). Atsinganosaurus confirms the presence in western Europe during the latest Cretaceous of a third titanosaurian species, slender and less derived which allows us to better understand the evolutionary and paleobiogeographical history of this group during the Cretaceous.
A well-preserved articulated skeleton of an ankylosaurian dinosaur from the Late Cretaceous of Bayan Mandahu (Inner Mongolia) is described. The specimen consists of the skull, lower jaws and a great part of the postcranial skeleton, including the cervical armour and tail-club in place. This material is referred to the ankylosaurid genus Pinacosaurus Gilmore, 1933. A new species, P. mephistocephalus, is erected on the basis of significant differences mainly in the nostril region relative to P. grangeri Gilmore, 1933. The new specimen, probably a subadult individual, provides further data about the composition and arrangement of the dermal armour of Pinacosaurus. Moreover, the comparisons with both young and adult specimens of Pinacosaurus allow a better understanding of ontogenetic changes in this dinosaur.
Le Campanien et le Maastrichtien du Languedoc ont été caractérisés paléontologiquement par des Mollusques, très progressivement, de 1830 à 1970. De nombreux gisements de Charophytes, découverts et étudiés depuis, permettent de préciser un certain nombre de points, comme l'apparition des matériaux pyrénéens à Villeveyrac au Rognacien inférieur, seulement, alors qu'ils sont présents dans les Corbières dès la base du Bégudien. Trois Charophytes nouvelles sont également décrites.
The turtle fauna of Laño includes four taxa which are known only from a restricted area in the late Campanian-
Maastrichtian of southern France and northern Iberian Peninsula. They are compared to other turtles of the world
and their relationships are studied by means of a short cladistic analysis. One is a cryptodire, Solemys, the latest
representative of a Laurasian Jurassic and Cretaceous family (Solemydidae), only known in western Europe at
Cretaceous times. European solemydids are revised and a new species of the genus Helochelydra, H. nopcsai,
is erected for the material from the Wealden (Lower Cretaceous) of the Isle of Wight, England. The other Laño
turtles are pleurodires: Dortoka is a representative of a primitive group (Dortokidae) considered as European in
origin; Polysternon and ?Elochelys are bothremydids and they probably come from the northern part of Africa,
where the family originates during the Early Cretaceous. All the genera are not known after Maastrichtian but
the Bothremydidae and Dortokidae persist in the Tertiary.
Key-words: Chelonians, Late Cretaceous, Iberian Peninsula, Endemism, New taxon, Ecology, Phylogenetic
relationships, Palaeobiogeography.
The dinosaurian fauna of the Niobrara Chalk Formation constitutes the best known assemblage from the middle Santonian-early Campanian interval of the Late Cretaceous of North America. The fauna consists of both hadrosaurid ornithopods and nodosaurid ankylosaurs.The nodosaurid specimens from the Smoky Hill Chalk Member of the Niobrara Chalk Formation include the holotype of Hierosaurus sternbergii (here regarded as nomen dubium), material of “Hierosaurus” coleii (herein designated the type species of the new genus Niobrarasaurus) and two specimens that may belong to N. coleii, but are better interpreted as Nodosauridae incertae sedis.Claosaurus agilis, the sole hadrosaurid from the Niobrara Chalk Formation, constitutes a poorly known, but distinct species having a basal relationship among remaining hadrosaurids.
Two partial skeletons of juvenile individuals of Stegosaurus with estimated body lengths of about 1.5 m (5 ft) and 2.6 m (8.5 ft) are described from the Morrison Formation (Upper Jurassic) of Utah and Wyoming. These juveniles differ from adults in the absence of fusion in composite bones (sacrum, scapulocoracoid, tibia–fibula–astragalus–calcaneum), the smoothness of the surface of the bones, the smaller size of ridges for muscular attachment, a proportionally more slender and elongate scapula, the relatively small size of the olecranon process of the ulna and the head of the femur, and the possible absence of dermal plates (but tail spines were present). Apart from the ratios of the length of the femur to those of the humerus and ilium, and the form of the scapula, ilium and fibula, the bones of juvenile individuals of Stegosaurus are very similar to those of juvenile individuals of the stegosaur Kentrosaurus (Upper Jurassic, East Africa). A probable sexual dimorphism in the sacrum with either four (?female) or five (?male) sacral ribs occurs in Kentrosaurus. The additional sacral rib attaches anterior to the other four; a rib in the Utah juvenile has the same distinctive shape, and it thus may have been a male (sacra of Stegosaurus with four sacral ribs have been previously illustrated).
A skeletal reconstruction and life restoration are presented for the Early Cretaceous nodosaurid ankylosaur Sauropelta edwardsi. This composite is based on several partial skeletons having armor preserved in situ from the Lower Cretaceous Cloverly Formation of Wyoming and Montana. Comparison of foot morphology with that of the ichnofossil Tetrapodosaurus borealis, from the Lower Cretaceous Gething Formation of British Columbia, suggests that the footprint is that of Sauropelta. A hypothesis is presented suggesting that most ankylosaurs are found on their backs because of bloating.- Author
Previously life reconstructions of ankylosaurid dinosaurs are incorrect in that they show uniformly shaped keeled plates arranged in neat longitudinal and transverse rows, and large spines projecting laterally from the flanks of the body. Articulated armor of Euoplocephalus tutus (previously described as the types "Scolosaurus cutleri" Nopsca and "Dyoplosaurus acutosquameus" Parks) from the Late Cretaceous Oldman Formation shows that armor shape and size varied considerably along neck, back, and tail. An attempt is made to reconstruct the skeleton and arrangement of the dermal armor of Euoplocephalus.
New remains of armoured dinosaurs from the late Cretaceous Lano locality in the north of the Iberian Peninsula are described and referred to the nodosaurid Struthiosaurus Bunzel. The more interesting material includes lower jaws and cervical dermal plates. The significance of the osteological variability in Struthiosaurus is discussed. It may reflect either one or several sources of intraspecific variation, or the occurrence of several species within Struthiosaurus. -Authors
The magnetostratigraphy of various sections in the Cretaceous-Tertiary basin of Aix-en-Provence allows us to correlate the local lithostratigraphy with marine stages. The Valdonnian corresponds to the Santonian, the Fuvelian and the Begudian with the Campanian, and the Gornacian with the Maastrichtian. -English summary
The skeleton of a small ankylosaur, referred to the genus Minmi (Molnar 1980), was recovered from the Albian Allaru Mudstone of north-central Queensland. The specimen shows most of the contacts between the cranial bones and preserves most of the dorsal armor in place. Unusual features include an apparently very large inferior process of the premaxilla, thin ventral sheets of the nasals, an apparently rod-like, vertical lachrymal, and a pronounced coronoid process on the mandible. The ilium is joined to two sacrais by a broad 'bridge' of bone. The skeleton is thought to derive from a mature or almost mature individual, and may have been mummified before burial.
Approximately half of existing dinosaur species belonging to the Order Ornithischia, or the 'bird-hipped' dinosaurs, which include such familiar forms as the stegosaurs, ankylosaurs, hadrosaurs, and ceratopsids. Although ornithischians are generally conceded to have descended from a common ancestor, little is known about the pattern of descent. Comparison of more recently discovered ornithischian fossils from China and Mongolia to better-known North American forms has shed light on the pattern of evolutionary diversification among ornithischians, a pattern that began approximately 200 My ago and ended abruptly nearly 140 My later at the end of the Cretaceous.-from Author
A partial nodosaurid ankylosaur skeleton, consisting primarily of the ilia, hindlimbs, posterior dorsal armor, plus partial forelimb elements and additional armor, was recovered from the marine Point Loma Formation, late Campanian age, north of San Diego, California. The specimen is similar to contemporaneous species of Panoplosaurus and Edmontonia from terrestrial sediments of the western interior, but there are also similarities to the armor of Stegopelta landerensis from marine sediments of earliest Cenomanian age from Wyoming. Skeletal elements critical for generic determination are not preserved, and the specimen is identified as Nodosauridae, incertae sedis. An associated marine invertebrate fauna and nannoplankton flora have revealed some discrepencies in the correlation systems used for Upper Cretaceous marine rocks of coastal California. Hollow limb bones of the specimen are interpreted as a preservational artifact. Nodosaurids had broad ecological tolerances, and visited riparian and coastal environments more frequently than other dinosaurs. However, a review of morphologic and distributional evidence fails to support a theory of amphibious or aquatic habits for nodosaurids.
During the last few years, systematic prospections and excavations in the non-marine Campanian andMaastrichtian of southern France, from Provence in the East to the valley of the Garonne in the West, have considerably increased our knowledge of the continental vertebrates (fishes, amphibians, turtles, squamates, crocodilians, pterosaurs, dinosaurs, birds and mammals) from that time interval. A succession of faunal assemblages, corresponding to the Early Campanian, the Late Campanian/Early Maastrichtian and the Late Maastrichtian, can now be recognised, with a marked change in the dinosaur fauna during the Maastrichtian, but no clear evidence of decline during the last million years of the Cretaceous. The biogeographical complexity of the Late Cretaceous vertebrate assemblages from southwestern Europe is underlined.RésuméCes dernières années, des prospections et des fouilles systématiques dans le Campanien et leMaastrichtien continentaux du Sud de la France, de la Provence à l'Est à la vallée de la Garonne à l'Ouest, ont considérablement augmenté notre connaissance des vertébrés continentaux (poissons, amphibiens, tortues, squamates, crocodiliens, ptérosaures, dinosaures, oiseaux et mammifères) de cette période. Une succession d'assemblages fauniques, correspondant au Campanien inférieur, au Campanien supérieur/Maastrichtien inférieur et au Maastrichtien supérieur, peut désormais être reconnue, avec un changement marqué dans la faune dinosaurienne au cours du Maastrichtien, mais pas de preuve claire d'un déclin pendant les derniers millions d'années du Crétacé. La complexité biogéographique des assemblages de vertébrés du Crétacé supérieur du Sud-Ouest de l'Europe est soulignée.
A partial nodosaurid ankylosaur skeleton from Lower Cretaceous littoral deposits of Texas represents a new genus and species, Texasetes pleurohalio. It is distinguished by a prong-like scapular spine that is directed toward the innermost point of the glenoid, development of a small prespinous fossa, and retention of a splint-like fourth trochanter on the femur. Preservation in marginal marine deposits is not indicative of normal habitat preferences. T. pleurohalio is more advanced than Hoplitosaurus marshi, and the latter may be the most primitive nodosaurid for which a considerable part of the specimen is known. The proposed synonymy of Hoplitosaurus and Polacanthus is rejected.
Pawpawsaurus campbelli gen. et sp. nov. from the Paw Paw Formation (late Albian), Tarrant County, Texas, appears more primitive than other pre-Campanian nodosaurids, Silvisaurus condrayi and Sauropelta edwardsi. New cranial synapomorphies for the Nodosauridae are prominent W-shaped basioccipital tubera, anteriorly concave and anteroposteriorly flattened quadrate, and transversely continuous and straight posterior margin of the pterygoid aligned with the quadrate shaft. These synapomorphies are closely related to the downward orientation of the nodosaurid head in life. A pair of bony eyelids recovered with the skull are the first discovered for the Nodosauridae. Other Paw Paw Formation nodosaurid remains, including new postcranial elements and a baby nodosaurid, are taxonomically indeterminate.
The Gosau formation, nearly corresponding in age to the Upper Greensand of this country, is represented at Neue Welt, near Wiener Noustadt, by freshwater deposits full of such freshwater shells as Melania and Unio, and land-plants such as Banksia and Pecopteris. The formation and its fauna have been described by Profs. Suess, Zittel, and many others; but, although the late Dr. Stoliczka detected a tooth imbedded in the coal of the formation, no important knowledge was obtained of the vertebrate fauna of the Gosau beds until Prof. Suess was so fortunate as to obtain the assistance of Bergverwalter Pawlowitsch in conducting excavations. These were carried on with admirable skill; timber drift-ways were driven into the rocks, with the result that they penetrated into a perfect cemetery of the remains of Cretaceous reptiles. The remarkable collection thus obtained was intrusted for description to Dr. Emanuel Bünzel, whose memoir upon it was published in 1871 in the ‘Transactions of the Imperial-Royal Geological Institution.’ Subsequently more specimens were discovered; and in Easter 1879 my honoured friend, Prof. Suess, invited me to visit Vienna to examine these specimens, with the object of making them available for the advancement of knowledge by publication. With the assistance of the Royal Society I gladly undertook this work, and spent a month in Vienna studying the thousands of fragments which had been obtained. The great, mass of these, mere comminuted bones, proved of but little value; or, rather, the time that I could give to their study enabled
The bone described by the author was found by Prof. Suess in a coal-mine belonging to the Gosau formation at Grünbach, near Wiener Neustadt, from which Prof. Suess obtained numerous other bones also belonging to terrestrial reptiles.
The author stated that the reptilian character of the bone appeared at the first glance, as it shows a single condyle and a temporal fossa resembling that of a Crocodile; but it has some peculiarities, such as the convexity of the occiput and its gentle passage into the roof of the skull, without forming an angle as in other reptiles, the transverse ridge in the occipital region, the want of sutures between the bones, the globular form of the condyle, the horizontality of the base, the ascending direction of the clivus, and the large brain-cavity, which render it impossible to refer the animal to which this bone belonged to any recognized order of Reptiles.
The known skulls of Dinosaurians have a steep occiput, and exhibit more of the Lacertilian type; the fragment described by the author rather resembled that of a bird. The author consequently suggested the formation of a new order of fossil Reptiles, Ornithocephala, nearly allied to the Ornithoscelida of Prof. Huxley.
For the animal of which this fragment is the only known relic, the author proposed the generic name of Struthiosaurus.
The author describes the large dorsal shield, which has been recently restored and now exhibits the grouping of the keeled and tuberculated fragments, which in their disconnected and scattered condition had formerly been regarded as portions of separate scutes. This unique specimen shows Polacanthus to have possessed a more complete dermal armature than any other Dinosaur yet described.
The origin and early evolution of many major dinosaur groups are poorly known because specimens are rare. One of these groups, the Ankylosauria, or armour-plated dinosaurs, is best known from well-preserved specimens from the Upper Cretaceous period of Asia and North America. Here we describe a well-preserved skull of an earlier, Late Jurassic ankylosaur, which will be important in clarifying the early history of this group. The specimen, Gargoyleosaurus parkpini gen. et sp. nov., was collected from the Upper Jurassic Morrison Formation of Wyoming, USA. Despite its geological age, the skull shows features seen in Late Cretaceous ankylosaurs, including fusion of bone armour to the surface of the skull and mandible and closure of two skull openings, the antorbital and upper temporal fenestrae. The new taxon also has characters common to the two ankylosaur families, the Ankylosauridae and Nodosauridae, supporting the proposal that the Ankylosauria originated from a single ancestor. Nevertheless, specialized characters place Gargoyleosaurus as the most primitive, or basal, member of the Ankylosauridae.
A specimen of a juvenile ankylosaur from the Upper Cretaceous Horseshoe Canyon Formation (Red Deer River, Alberta, Canada) includes a nearly complete pes, the first reported for the genus Euoplocephalus. Differences between juvenile and adult ankylosaurs are qualitatively similar to differences reported for juvenile and adult stegosaurs.
In 1923, H. C. T’an and O. Zdansky collected remains of an ankylosaurid dinosaur in the Late Cretaceous Wangshi Group of the Laiyang region, in eastern Shandong (China). Apart from a few caudal vertebrae, this material, which is kept at the Palaeontological Institution of the University of Uppsala (Sweden), was never described or figured. It includes a well-preserved sacrum with the attached right ilium and part of the presacral rod, caudal vertebrae, a left femur and a dermal scute. This material is referred to an ankylosaurid of the genus Pinacosaurus Gilmore, 1933, on the basis of the widely divergent ilium bearing a strong ventral ridge and of the slenderness of the femur. In the absence of cranial material, a specific attribution is difficult and the Uppsala material is referred to as Pinacosaurus cf. grangeri (P. grangeri being the only generally accepted species of Pinacosaurus). This is the first record of Pinacosaurus outside the Gobi Basin of Mongolia and northwestern China. In the Gobi Basin, Pinacosaurus has been reported only from the Djadokhta Formation or its equivalents, of supposed Campanian age, and it is suggested that at least the part of the Wangshi Group which yielded the Shandong Pinacosaurus may be of roughly the same age as the Djadokhta Formation.