Lower Cretaceous Of Europe Research Articles

Ornithopod tracks and bones: Paleoecology and an unusual evidence of quadrupedal locomotion in the Lower Cretaceous of eastern Iberia (Teruel, Spain)

Dinosaur tracks and osteological remains in the El Castellar Formation (upper Hauterivian–lower Barremian) in the Peñagolosa subbasin, southwest sector of the Maestrazgo Basin (Teruel Province, Spain), are scarce and fragmentary, respectively. Here, we report several unexpected, well-defined natural casts of large ornithopods from several sites of the El Castellar Formation. Moreover, an unusual manus-pes track set preserved as convex hyporelief has been found in the lowermost sandstone level of the Camarillas Formation, the first evidence of dinosaur for the base of this unit at the Peñagolosa subbasin. The ichnological comparison allows us to ascribe the studied tracks to Caririchnium. Their trackmakers were large styracosternans related to Iguanodon, and the presence of a manus-pes track set verifies that some of them may have a quadrupedal locomotion. This associated manus-pes track set constitutes one of the oldest ichnological pieces of evidence of quadrupedalism in large ornithopods from the Lower Cretaceous of Iberia and increases the limited record of this kind of track in the Lower Cretaceous of Europe. In addition, we describe some osteological fossils of these dinosaurs from the El Castellar Formation. These skeletal remains are related to medium to large-sized styracosternans, some of which closely resemble those of other European styracosternans such as Iguanodon. The evidence provided here verifies that these dinosaurs were frequent components in the coastal wetland, and fluvial to deltaic systems developed, during the late Hauterivian–early Barremian, in the Peñagolosa subbasin, possibly because these may have had a source of fresh vegetation and may have provided protection.

A Lower Cretaceous Lagerstätte from France: a taphonomic overview of the Angeac‐Charente vertebrate assemblage

Terrestrial ecosystems from the Lower Cretaceous of Europe and bonebeds formed in swampy environments are poorly known. The Berriasian-early Valanginian Angeac-Charente site in France represents an example of both. Nine field campaigns have yielded thousands of fossils of over a hundred taxa, including 16 taxa from vertebrate macroremains with numerous trample and crocodile bite marks; 22 taxa from the abundant vertebrate microremains; >10 vertebrate coprolite morphotypes with plant and vertebrate inclusions; abundant sauropod and stegosaur tracks including some preserved in ‘4-D’; termite coprolites; mollusc moulds; ostracods and plants, including coniferous wood, cones, leaves and cuticle fragments, charophytes and pollen. The richness, diversity and preservation of the fossils qualify the site as a fossil-Lagerstätte. The site represents a ‘snapshot’ into a Lower Cretaceous ecosystem. This is supported by REE analyses of biogenic apatite and sediment samples, the fossils being found in a single stratigraphical interval and the record of sedimentological and taphonomic ‘frozen scenes’. The Angeac-Charente bonebed is highly diverse, dominated by an ornithomimosaur taxon, and contains both macro- and microfossils. This indicates a complex formation, likely primarily influenced by ecological and biologic processes, but also significant physical processes. These include crocodyliform predation and/or scavenging on turtles, ornithomimosaurs and fishes; probable mass mortality occurrence of an ornithomimosaur herd; possible social behaviour of stegosaurs; limited hydraulic transport of most sauropod bones and intense dinoturbation.

Avian and crocodilian eggshells from the upper Barremian site of Vadillos-1 (Lower Cretaceous, Cuenca province, Spain)

The new upper Barremian microfossil site of Vadillos-1 (Lower Cretaceous, Beteta Gorges, Cuenca province, Spain) has yielded a rich fossil assemblage consisting of numerous eggshell fragments along with diverse skeletal remains of fishes, amphibians, turtles, crocodyliforms and dinosaurs. Crocodilian eggshells constitute the main fraction of the eggshell assemblage which includes abundant material belonging to the oofamily Krokolithidae, as well as a new type of crocodilian eggshells (Neokrokolithes trigonalis oogen. et oosp. nov.) characterised by a type of ornamentation and microstructure of the basal knobs not described before. Few fragments of the oospecies cf. Mycomorphoolithus kohringi, attributed to non-eusuchian crocodylomorphs, have been also found in this site thus expanding the record of this oospecies to the upper Barremian. The eggshell assemblage has also yielded the first record of avian eggshells (Tristratioolithus minuta oogen. et oosp. nov.) from the Lower Cretaceous of Europe. Comparisons with Lower Cretaceous avian eggshells suggest its possible attribution to Enantiornithes. The tiny eggshell fragments assemblage found at Vadillos-1 site is representative of a small sized fauna. Besides, the diversity appears strongly biased with >90% of the eggshells belonging to crocodilians and approximately 9% to Aves. As established for similar outcrops, this bias may reflect differences in preservation of eggshell types or represent, at least in part, a wetland ground nesting site.

A new Brachiosauridae Sauropod dinosaur from the lower Cretaceous of Europe (Soria Province, Spain)

A new dinosaur sauropod from the Golmayo Formation (upper Hauterivian-lower Barremian) in Soria province (Spain) is described as Soriatitan golmayensis gen. et sp. nov. The new material consists of one tooth, three dorsal vertebrae with ribs, a partial sacrum, five caudal centra, two caudal vertebral spines, one chevron, a humerus, an ulna, a radius, two partial ilia, two ischia, a fragment of pubis, and a partial femur. Cladistic analysis identified the material as belonging to Brachiosaruidae within Titanosauriformes. Sauropod material from the Golmayo Formation shares a diagnostic feature with Abydosaurus, Cedarosaurus, Tastavinsaurus and Venenosaurus including anteriorly deflected anterior–middle caudal neural spines and with Cedarosaurus a proximodistally straight lateral margin between the proximal head and the shaft of the humerus. Eight characters are potential autapomorphies in the sauropod from Spain, including the presence of a large rectangular ventral ridge below the preacetabular process of the ilium, the same length of the pubic and ischiadic blades in the ischium, and two lateral ridges in the lateral area of the deltopectoral crest of the humerus. The presence of Early Cretaceous brachiosarids in both, North America and Europe, give support to the hypothesis of a connection between the tectonic plates of these continents at some point during the Early Cretaceous.

A new basal elasmosaurid (Sauropterygia: Plesiosauria) from the Lower Cretaceous of Germany

ABSTRACTHere we report on a new basal elasmosaurid plesiosaurian, Lagenanectes richterae, gen. et sp. nov., from the Lower Cretaceous (probably Upper Hauterivian) of Germany. The material includes a partial skull (cranium and mandible), the atlas-axis complex, additional cervical vertebrae, caudal vertebrae, an ilium, and limb elements. The basioccipital and atlas intercentrum are pathologically deformed, probably due to an osteomyelitic infection. Two potential autapomorphies were found in the mandible: (1) the alveolar margin at the symphysis is laterally expanded with the rostral-most alveoli being markedly procumbent and situated along the lateral margins of the dentaries; and (2) the ventral midline at the symphysis is produced into a prominent wedge-shaped platform indented by numerous irregular pits. Lagenanectes richterae, gen. et sp. nov., also shows a number of typical elasmosaurid traits, including a longitudinal lateral ridge on the cervical vertebral centra (although a ventral notch is absent) and teeth with oval cross-sections. Lagenanectes richterae, gen. et sp. nov., is one of the best-preserved plesiosaurians from the Lower Cretaceous of Europe.

Phylogeny of the Ichthyopterygia incorporating recent discoveries from South China

ABSTRACTDuring the last decade, abundant ichthyopterygian material has been found from the Triassic of South China as well as the Upper Jurassic and Lower Cretaceous of Europe and South America, significantly expanding our knowledge of ichthyopterygian diversity through the Mesozoic. Previous phylogenetic hypotheses of the group no longer account for these extensive additions, necessitating a new phylogenetic framework for the entire Ichthyopterygia to enable evolutionary studies of the group. We present here a comprehensive phylogenetic hypothesis for Ichthyopterygia based on cladistic analysis of 163 characters coded for 59 ingroup and five outgroup taxa. The monophyly of Ichthyopterygia is strongly supported by a Bremer index value of 7. Five major groups of Ichthyopterygia during the Triassic, viz., Grippioidea, Cymbospondylidae, Mixosauridae, Shastasauridae, and Toretocnemidae, are well supported by Bremer index values between 3 and 5. Major clades that evolved in the Triassic, including Merriamosauria, Euichthyosauria, and Parvipelvia, are also robustly supported, whereas most post-Triassic clades are very weakly supported with a Bremer index value of 1, with a few exceptions, such as Thunnosauria and Ophthalmosauridae. The traditional Shastasauridae is expanded to comprise six genera but excludes Callawayia, which is more closely related to Parvipelvia than to Shastasauridae. ‘C.’ wolonggangensis is a shastasaurid but does not form a monophyletic clade with Callawayia neoscapularis or Guizhouichthyosaurus tangae as previously asserted. The new phylogenetic hypothesis is generally consistent with the stratigraphic occurrences of each taxon especially for the Triassic taxa.http://zoobank.org/urn:lsid:zoobank.org:pub:901B6FB6-2D80-4AE9-B017-A0226175AFDASUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Ji, C., D.-Y. Jiang, R. Motani, O. Rieppel, W.-C. Hao, and Z.-Y. Sun. 2015. Phylogeny of the Ichthyopterygia incorporating the recent discoveries from South China. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1025956.

A review of large Cretaceous ornithopod tracks, with special reference to their ichnotaxonomy

Trackways of ornithopods are well-known from the Lower Cretaceous of Europe, North America, and East Asia. For historical reasons, most large ornithopod footprints are associated with the genus Iguanodon or, more generally, with the family Iguanodontidae. Moreover, this general category of footprints is considered to be sufficiently dominant at this time as to characterize a global Early Cretaceous biochron. However, six valid ornithopod ichnogenera have been named from the Cretaceous, including several that are represented by multiple ichnospecies: these are Amblydactylus (two ichnospecies); Caririchnium (four ichnospecies); Iguanodontipus, Ornithopodichnus originally named from Lower Cretaceous deposits and Hadrosauropodus (two ichnospecies); and Jiayinosauropus based on Upper Cretaceous tracks. It has recently been suggested that ornithopod ichnotaxonomy is oversplit and that Caririchnium is a senior subjective synonym of Hadrosauropodus and Amblydactylus is a senior subjective synonym of Iguanodontipus. Although it is agreed that many ornithopod tracks are difficult to differentiate, this proposed synonymy is questionable because it was not based on a detailed study of the holotypes, and did not consider all valid ornithopod ichnotaxa or the variation reported within the six named ichnogenera and 11 named ichnospecies reviewed here. We therefore emphasize the need to base comparisons between ichnotaxa on type material, and not on selected referred material. It is concluded that there is considerable variation in the morphology of the holotypes, as well as variation in size and quality of the samples and the mode of preservation. Conversely, there is considerable overlap in morphology among other tracks that have been informally attributed to these ichnotaxa. These factors make it difficult to synonymize any of the existing ichnotaxa without detailed revision of the samples from which the type material originates. Nevertheless, a review of the type material of all ichnotaxa is presented as a basis for further discussion and, as a first step, the ichnofamily Iguanodontipodidae is proposed to accommodate Amblydactylus, Caririchnium and Iguanodontipus.

Spheroolithid eggshells in the Lower Cretaceous of Europe. Implications for eggshell evolution in ornithischian dinosaurs

Early Cretaceous spheroolithid eggshells have always been extremely scarce, unlike their putative egg layers, ornithopod dinosaurs, which are ubiquitous in Early Cretaceous ecosystems. In this paper, we re-evaluate eggshells previously assigned to the oofamily Elongatoolithidae, an oofamily related to derived theropod dinosaurs (including maniraptorans and oviraptorids), and reassign them to the oofamily Spheroolithidae, on the basis of the lack of a squamatic ultrastructure, and the presence of a prolatocanaliculate pore system and undulating extinction, typical of spheroolithid eggshells. We erect a new oogenus, Guegoolithus to include material previously reported as Macroolithus turolensis Amo-Sanjuan, Canudo and Cuenca-Bescós 2000, and new material from several lower Barremian localities of the Maestrazgo Basin in Spain. Guegoolithus eggshells exhibit an acicular radial ultrastructure in the lower part of the shell units, which develops into a tabular radial ultrastructure in the upper part, giving an appearance of a two-layered eggshell in thin sections that is not seen in SEM pictures. These features fill the gap between the two-layered prismatic eggshell of ceratopsians and the prolatospherulitic ornithopod eggshell of derived hadrosaurs.

The Basal Nodosaurid Ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (Lower Albian) Escucha Formation of Northeastern Spain

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.

The European Early Cretaceous Cryptodiran TurtleChitracephalus dumoniiand the Diversity of a Poorly Known Lineage of Turtles

Chitracephalus dumonii was named based on some of the most complete turtle remains from the Lower Cretaceous of Europe, and yet the taxon has barely been mentioned since. Indeed, new specimens were erroneously attributed to a new taxon, ldquo;Salasemys pulcherrima”. The synonymy is recognized here, and this extends the geographical range of this turtle and provides examples of individuals at different stages of ontogenetic development. The peculiar structure of its shell, and its ontogenetic development, are unique to this taxon. The systematic position of C. dumonii was previously unclear, usually being referred to Testudinata incertae sedis. Here, it is placed in a cladistic analysis, which shows that C. dumonii, and the recently described Hoyasemys jimenezi form part of a Lower Cretaceous European clade of Cryptodira that includes “macrobaenid”, “sinemydid”, and panchelonioidean turtles.

Galvechelone lopezmartinezae gen. et sp. nov., a new cryptodiran turtle in the Lower Cretaceous of Europe

Abstract: The Spanish town of Galve (Teruel) is notable because of the abundance of Upper Jurassic and, especially, Lower Cretaceous vertebrates recorded there. Although most groups have been studied in detail, information on turtles is very limited even though the material is relatively abundant. So far, no turtle taxa have been identified at the generic level. The only Lower Cretaceous articulated specimen from Galve is analysed here. It is identified as a representative of Cryptodira, Galvechelone lopezmartinezae gen. et sp. nov. Galvechelone lopezmartinezae is determined as a taxon belonging to the node that groups the turtles traditionally assigned to ‘Macrobaenidae’ and ‘Sinemydidae’, and other taxa such as the members of Panchelonioidea. This node, very abundant in the Lower Cretaceous of Asia, and with a broad subsequent distribution, has recently been recognized in the Lower Cretaceous of Europe. The diversity of basal members of Eucryptodira in the European Late Jurassic (represented by Thalassemydidae, Plesiochelyidae and Eurysternidae) was high. Owing to a relative scarcity of well‐preserved early Cretaceous turtles from Europe, the knowledge of this group of reptiles is limited. The study of the new turtle from Galve, together with the recently described Hoyasemys jimenezi, and the recently completed review of the enigmatic Chitracephalus dumonii demonstrate that members of the cryptodiran node grouping ‘Macrobaenidae’, ‘Sinemydidae’ and Panchelonioidea were also very diverse in this period.

High diversity of pancryptodiran turtles in the Lower Cretaceous of Europe

Turtles traditionally identified as basal Eucryptodira members were very abundant and diverse in the European Upper Jurassic. However, the European Cretaceous record of these turtles is poorly known. Brodiechelys brodiei, from the Wealden of the UK, is a turtle traditionally assigned to Eucryptodira. This taxon, recognized on the basis of very little material, has been poorly described and figured. More than a dozen shells are here attributed to B. brodiei. This allows a new diagnosis to be proposed for the taxon, and recognizes that their shells are subject to great intraspecific variability. This conclusion affects many characters used in the diagnosis of taxa defined by scarce material. The analysis of Hylaeochelys, a European Lower Cretaceous taxon traditionally identified as another basal Eucryptodira member, also allows high variability to be observed and several synonyms to be recognized. Hylaeochelys belli is identified here as the only valid species of this genus. Brodiechelys brodiei, H. belli and other European and Asian Jurassic and Lower Cretaceous taxa, previously identified as basal members of Eucryptodira, are incorporated into two hypotheses of phylogenetic relationships. A basal dichotomy in Pancryptodira is obtained. In one of the nodes, taxa traditionally assigned to Paracryptodira, Plesiochelyidae and H. belli are included. The other node groups B. brodiei, taxa traditionally assigned to “Xinjiangchelyidae”, and the Cryptodira members. This study shows a high diversity of Pancryptodira members in the Lower Cretaceous of Europe, assigned to several nodes. Representatives of Paracryptodira, other members of the node grouping the traditional members of Plesiochelyidae and Paracryptodira, and several members of the node grouping Xinjiangchelys latimarginalis and Cryptodira are identified.

A new Berriasian species ofGoniopholis(Mesoeucrocodylia, Neosuchia) from England, and a review of the genus

A new species, Goniopholis kiplingi sp. nov., based on an exceptionally preserved skull from the Lower Cretaceous of England is described in detail. It shows great similarity with Goniopholis simus and Goniopholis baryglyphaeus, but can be distinguished by the presence of longer lachrymals, smooth (not edged) dorsal surface of the quadrate, and proportionally longer rostrum. A comprehensive phylogenetic analysis of Mesoeucrocodylia (104 taxa; 486 characters) focused on goniopholidids (14 species) places G. kiplingi as sister-group of G. simus, and as part of a monophyletic group also containing G. baryglyphaeus. The relationships of Nannosuchus gracilidens and three undescribed European taxa are explored, and preliminary analyses of Denazinosuchus kirtlandicus (Upper Cretaceous, USA) and ‘Goniopholis’ phuwiangensis (Lower Cretaceous, Thailand) are presented. The assignment of taxa to the genus Goniopholis is discussed. Goniopholis, in its traditional sense, is considered paraphyletic and a restricted updated definition is proposed, with comments on the evolution of other goniopholidids. Morphological characteristics of fragmentary material attributed to Goniopholis are not considered sufficient to secure their generic/specific assignment, and provide no support for the presence of Goniopholis in Gondwanan and/or Upper Cretaceous sedimentary units. Currently Goniopholis is restricted to the Upper Jurassic−Lower Cretaceous of Europe. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, S66–S108.

A SPALACOLESTINE SPALACOTHERIID (MAMMALIA, TRECHNOTHERIA) FROM THE EARLY CRETACEOUS (BARREMIAN) OF SOUTHERN ENGLAND AND ITS BEARING ON SPALACOTHERIID EVOLUTION

Abstract: Bulk screening of Early Cretaceous (Barremian) Wealden Group strata of the Wessex Formation exposed on the south‐west and south‐east coasts of the Isle of Wight, southern England, has resulted in the recovery of fragmentary remains pertaining to a new spalacolestine spalacotheriid mammal, Yaverlestes gassoni gen. et sp. nov. These represent the first European record of the Spalacolestinae. The remains comprise a dentulous incomplete dentary and isolated upper and lower molariforms, the former representing the most substantial mammal remains yet recovered from the Wealden Group. Hitherto, six species of spalacotheriid mammal were known from the Lower Cretaceous of Europe. All are referred to the genus Spalacotherium but in the case of taxa diagnosed on the basis of isolated lower teeth and other specimens where the post‐canine dentition is incompletely known, it is now evident that these referrals should be treated with caution. Furthermore, the new Wessex Formation spalacotheriid and recently described spalacotheriids from the ?Barremian of Japan, and the Barremian and Aptian of China exhibit combinations of characters that suggest that spalacotheriids were more diverse and that their evolution was more complex than previously recognized. The systematic position of an isolated tooth from the basal Cretaceous Lourinhã Formation of Portugal is discussed and the tooth reassigned to the Spalacotheriidae. Together with the new Wessex Formation taxon, eight species are now known from the Lower Cretaceous of Europe. The discovery of a spalacolestine in the Barremian Wessex Formation supports the concept of faunal interchange between Europe, Asia and North America during the Early Cretaceous. It also supports derivation of North American spalacotheriids from a European or Eurasian ancestor.

An atoposaurid neosuchian (Archosauria: Crocodyliformes) from the Lower Cretaceous of Inner Mongolia (People's Republic of China)

Cranial and postcranial remains from the Laolonghuoze locality in the Ordos Basin, Inner Mongolia (China), represent a previously unrecorded taxon of crocodyliform archosaurs from the Lower Cretaceous Luohandong Formation (Zhidan Group). Several cranial features indicate that this form is referable to the Atoposauridae, which were previously definitely known only from the Upper Jurassic and Lower Cretaceous of Europe. Certain derived characters further indicate that the Ordos material is closely related to, if not actually referable to, Theriosuchus. The Ordos atoposaurid differs from the species of Theriosuchus in only a few features. In view of its fragmentary nature, the new material is identified as cf. Theriosuchus sp. at present.

New and little-known genera of cheilostome Bryozoa from the New Zealand region

Six new or little-known genera of cheilostome Bryozoa are recorded from the New Zealand region. Villicharixa n. gen. (Electridae), established for a species from South Island and from southern Chile, closely resembles forms from the lower Cretaceous of Europe. Daisyella n. gen. (Calloporidae) includes three uniserial species (two New Zealand and one Antarctic). Rhabdozoum Hincks (Rhabdozoidae), a little-known southern Australian genus, is newly recorded from the Otago Shelf, South Island. Plagiopora MacGillivray (Catenicellidae), a hitherto monotypic? Miocene Australian genus of the little-known subfamily Ditaxiporinae, is represented in Recent seas by a new species from the Norfolk Ridge. Yrbozoon n. gen. (Celleporidae) and Harpago n. gen. (Lekythoporidae), both monotypic, are based on new species from submarine plateaux southeast of New Zealand.

Goniopholis phuwiangensis nov. sp., a newmesosuchian crocodile from the Mesozoic of north-eastern Thailand

A new species of Goniopholis (Order Crocodylia, Suborder Mesosuchia, Family Goniopholididae) is described as G. phuwiangensis on the basis of a dentary from the Sao Khua Formation at Phu Wiang, in north-eastern Thailand. It is closely related to forms from the Upper Jurassic of North America and Europe and the Lower Cretaceous of Europe, and suggests a Late Jurassic to Early Cretaceous age for the Sao Khua Formation. The occurrence of Goniopholis in Thailand is indicative of Laurasian faunal affinities.

Valdosaurus, a hypsilophodontid dinosaur from the Lower Cretaceous of Europe and Africa

Material of the hypsilophodontid dinosaur Valdosauruscanaliculatus (Ornithischia: Ornithopoda) is described from the Lower Cretaceous (Barremian) of southern England and a new species is recognized from the Lower Cretaceous (Aptian) of Niger, West Africa. This occurrence of Valdosaurus in Europe and Africa provides evidence of a land connection between these continents across Tethys sometime in the early Cretaceous.