New Trigonioidoidea (Bivalvia; Unionoida) from the Early Cretaceous of Spain

Abstract: Two taxa belonging to the Trigonioidoidea (Order Unionoida) are described from the Early Cretaceous of Spain. Nippononaia (Paranippononaia) camerana subgen. et sp. nov. is described from the Aptian of the Cameros Basin of the north‐west Iberian Range. Subnippononaia fordi Barker et al., 1997 is described from new material from the Calizas de la Huérguina Formation (Late Barremian) from Las Hoyas and Buenache de la Sierra, Cuenca Province, and Subnippononaia is raised to generic status. The stratigraphy and palaeoenvironment of the Cornago G exposure of the Enciso Group where Nippononaia (Paranippononaia) camerana originates is interpreted as a fresh to brackish water system.     

Plio-Pleistocene evolution of the genus Castor (Rodentia, Mammalia) in Europe: C. fiber plicidens of Pietrafitta (Perugia, Central Italy)

In the lignite sediments of Pietrafitta (Tiberino Basin, Umbria, Central Italy), a rich fossil assemblage of vertebrate, invertebrate and plant remains belonging to the Farneta Faunal Unit (Late Villafranchian, Early Pleistocene) was collected. Among them fossil beaver cranial remains are relatively abundant. Western-Central Europe Villafranchian beaver remains were assigned to C. plicidens by some authors because molar occlusal surface patterns show complex enamel crenulations. Several recent authors have classified them as C. fiber while analysing other morphological patterns. Our samples have been compared to Plio-Pleistocene fossil remains and to living European populations of the genus Castor. New morphometric parameters on molar occlusal surfaces have been defined and statistical analyses (One-Way MANOVA, Principal Component Analysis and Canonical Discriminant Analysis) have been performed on them. The results point out a subspecific separation between the Late Villafranchian beaver of Pietrafitta, Quarata and San Giovanni in Valdarno (Umbria and Tuscany) and C. fiber populations. St. Vallier (France) Late Pliocene and Mosbach 2 (Germany) Middle Pleistocene beavers classified respectively by Viret and Friant as C. plicidens, show a C. f. fiber molar teeth pattern. Therefore, C. fiber plicidens did not occur in Central-Western Europe and this subspecific name may be used only for the local populations of Valdarno and Tiberino Basin (San Giovanni in Valdarno, Quarata, Pietrafitta and a few localities of the same area), at that time peripheral populations, probably semi-isolated during the Late Villafranchian, and at the southern limit of the geographic range of C. fiber.     

A male cone of Pseudofrenelopsis dalatzensis with in situ pollen grains from the Lower Cretaceous of Northeast China

A Classopollis-containing male cone found in close association with Pseudofrenelopsis dalatzensis from the Lower Cretaceous Dalazi Formation of Yanji Basin, eastern Jilin, China was studied using scanning electron microscopy and described as Classostrobus dalatzensis sp. nov. The small oval cone, borne on the top of a stalk (fertile short shoot), consists of helically arranged microsporophylls, each with a rhomboid and expanded distal head. The short shoot is divided into indistinct nodes and internodes with fine longitudinal striations. Cuticles of the microsporophyll and the internode of the short shoot are generally similar to Pseudofrenelopsis dalatzensis internode cuticles, all of them having papillate outer surfaces. Stomata are similar in structure and arrangement, with papillae of subsidiary cells overhanging the stomatal pit. Epidermal cells are rectangular or isodiametrical, each with a robust papilla on the periclinal wall. Hypodermal cells are present between the stomatal files. The internal structure of the in situ Classopollis pollen grains was studied in detail. Pollen grain is small, with thin sexine, poorly developed nexine and may be immature. This is the first record of in situ pollen grains Classopollis in China of which the internal structure is known in detail.     

A New Sail-Backed Styracosternan (Dinosauria: Ornithopoda) from the Early Cretaceous of Morella,Spain

A new styracosternan ornithopod genus and species is here described based on a partial postcranial skeleton and an associated dentary tooth of a single specimen from the Arcillas de Morella Formation (Early Cretaceous, late Barremian) at the Morella locality, (Castellón, Spain). Morelladon beltrani gen. et sp. nov. is diagnosed by eight autapomorphic features. The set of autapomorphies includes: very elongated and vertical neural spines of the dorsal vertebrae, midline keel on ventral surface of the second to fourth sacral vertebrae restricted to the anterior half of the centrum, a posterodorsally inclined medial ridge on the postacetabular process of the ilium that meets its dorsal margin and distal end of the straight ischial shaft laterally expanded, among others. Phylogenetic analyses reveal that the new Iberian form is more closely related to its synchronic and sympatric contemporary European taxa Iguanodon bernissartensis and Mantellisaurus atherfieldensis, known from Western Europe, than to other Early Cretaceous Iberian styracosternans (Delapparentia turolensis and Proa valdearinnoensis). The recognition of Morelladon beltrani gen. et sp. nov. indicates that the Iberian Peninsula was home to a highly diverse medium to large bodied styracosternan assemblage during the Early Cretaceous.     

Evidence of predation in Early Cretaceous unionoid bivalves from freshwater sediments in the Cameros Basin,Spain

Here, we present evidence of possible vertebrate predation on freshwater bivalves from the Lower Cretaceous strata of the Cameros Basin (Spain). The described collection contains the largest number of vertebrate‐inflicted shell injuries in freshwater bivalve shells yet reported in the Mesozoic continental record. Several types of shell damage on fossil shells of Protopleurobema numantina (Bivalvia: Unionoida) are described and their respective modes of formation interpreted in the context of morphological attributes of the shell injuries and the inferred tooth morphology of predators that could have inflicted such injuries. Detailed study of these bite marks shows similarities with the well‐documented injuries in the shells of marine molluscs, namely ammonoids, that have likewise been attributed to reptilian predators. The most parsimonious interpretation suggests crocodiles as the vertebrates interacting with the bivalves in the Cameros Basin. □Barremian–Aptian; bite marks; freshwater bivalves; predation; reptile; Unionoida.     

The affinities of the ostracod genus Cypridea Bosquet, 1852, and its allies, with consideration of implications for the phylogeny of nonmarine cypridoidean ostracods

The nonmarine ostracod genus Cypridea s.l., characterized by an antero-ventral “beak” (rostrum and alveolus) in both valves, achieved high diversity and global distribution in the Early Cretaceous but declined in the Late Cretaceous and became extinct during the Paleogene. Although it clearly belongs to the Superfamily Cypridoidea (Order Podocopida, Suborder Cypridocopina), the precise affinities of Cypridea s.l. have been controversial, different authors variously suggesting it to be most closely related to the cypridoidean families Ilyocyprididae, Cyprididae or Notodromadidae. Since Cypridea s.l. was responsible for much of the explosive radiation of nonmarine cypridoidean taxa during the Mesozoic, a clear understanding of its affinities is crucial to the elucidation of nonmarine ostracod phylogeny. We evaluate some of the key morphological features of cypridoidean carapaces as indicators of phylogenetic affinity, paying special attention to adductor muscle scar patterns and the structure of the anterior marginal zone. The morphology of Cypridea s.l. is compared with certain cypridoideans that bear similar beak-like or lip-like antero-ventral marginal structures, notably genera of the Family Cyprididae such as Bennelongia, Chlamydotheca, Cypris and Talicypridea, and consider whether these similarities represent close phylogenetic relationships or homeomorphy.     

The Genera Haplocytheridea Stephenson, 1936 and Cytheridea Bosquet, 1852 (Subphylum Crustacea, Class Ostracoda) from the Early Miocene Pirabas Formation, Brazil

The genera Haplocytheridea and Cytheridea have Tethyan origins and a wide paleobiogeographic and stratigraphic distribution, ranging from the Cretaceous to Recent. Both of them have been recorded in northern South America, but only Cytheridea has been found in the northern Cretaceous basins of Brazil. The genus Haplocytheridea is recorded, for the first time, in the Early Miocene deposits of the Pirabas Formation of northern Brazil, from a 20 m-thick succession of carbonate and argillaceous rocks deposited in lagoonal and restricted platform environments. Among the ten Haplocytheridea species identified, four are new: H. variopunctata n. sp., H. sandbergi n. sp., H. pirabasensis n. sp. and H. sinuosa n. sp. In addition, three new species of Cytheridea are also described: C. coimbrai n. sp., C. pirabasensis n. sp. and C. purperae n. sp. The highest frequencies and abundances of both genera, in the studied section, are thought to be associated with nearshore to brackish water settings.     

Endosymbiotic bacteria nodulating a new endemic lupine Lupinus mariae-josephi from alkaline soils in Eastern Spain represent a new lineage within the Bradyrhizobium genus

Lupinus mariae-josephi is a recently described endemic Lupinus species from a small area in Eastern Spain where it thrives in soils with active lime and high pH. The L. mariae-josephi root symbionts were shown to be very slow-growing bacteria with different phenotypic and symbiotic characteristics from those of Bradyrhizobium strains nodulating other Lupinus. Their phylogenetic status was examined by multilocus sequence analyses of four housekeeping genes (16S rRNA, glnII, recA, and atpD) and showed the existence of a distinct evolutionary lineage for L. mariae-josephi that also included Bradyrhizobium jicamae. Within this lineage, the tested isolates clustered in three different sub-groups that might correspond to novel sister Bradyrhizobium species. These core gene analyses consistently showed that all the endosymbiotic bacteria isolated from other Lupinus species of the Iberian Peninsula were related to strains of the B. canariense or B. japonicum lineages and were separate from the L. mariae-josephi isolates. Phylogenetic analysis based on nodC symbiotic gene sequences showed that L. mariae-josephi bacteria also constituted a new symbiotic lineage distant from those previously defined in the genus Bradyrhizobium. In contrast, the nodC genes of isolates from other Lupinus spp. from the Iberian Peninsula were again clearly related to the B. canariense and B. japonicum bv. genistearum lineages. Speciation of L. mariae-josephi bradyrhizobia may result from the colonization of a singular habitat by their unique legume host.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

Ethological variations in gauge in sauropod trackways from the Berriasian of Spain

Castanera, D., Pascual, C., Canudo, J.I., Hernández, N. & Barco, J., 2012: Ethological variations in gauge in sauropod trackways from the Berriasian of Spain. Lethaia, Vol. 45, pp. 476–489. Two sauropod trackways from the Early Cretaceous (Berriasian) of the Cameros Basin (Spain) show important variations in trackway gauge, along the same trackway. These variations seem to be associated with different behaviours; on the one hand with a significant variation in the direction of travel (turning phenomena) and on the other hand with a decrease in speed, probably associated with the properties of the substrate. These variations in sauropod trackways provide the trackway gauge debate with new data, supporting the hypothesis that the walking style and substrate properties may in some cases determine this sauropod character. The study of turning sauropod trackways improves our knowledge of sauropod locomotion. □Ethology, intermediate‐gauge, sauropod trackways, substrate properties, wide‐gauge.     

The origin and early evolution of metatherian mammals: the Cretaceous record

Metatherians, which comprise marsupials and their closest fossil relatives, were one of the most dominant clades of mammals during the Cretaceous and are the most diverse clade of living mammals after Placentalia. Our understanding of this group has increased greatly over the past 20 years, with the discovery of new specimens and the application of new analytical tools. Here we provide a review of the phylogenetic relationships of metatherians with respect to other mammals, discuss the taxonomic definition and diagnosis of Metatheria, outline the Cretaceous history of major metatherian clades, describe the paleobiology, biogeography, and macroevolution of Cretaceous metatherians, and provide a physical and climatic background of Cretaceous metatherian faunas. Metatherians are a clade of boreosphendian mammals that must have originated by the Late Jurassic, but the first unequivocal metatherian fossil is from the Early Cretaceous of Asia. Metatherians have the distinctive tightly interlocking occlusal molar pattern of tribosphenic mammals, but differ from Eutheria in their dental formula and tooth replacement pattern, which may be related to the metatherian reproductive process which includes an extended period of lactation followed by birth of extremely altricial young. Metatherians were widespread over Laurasia during the Cretaceous, with members present in Asia, Europe, and North America by the early Late Cretaceous. In particular, they were taxonomically and morphologically diverse and relatively abundant in the Late Cretaceous of western North America, where they have been used to examine patterns of biogeography, macroevolution, diversification, and extinction through the Late Cretaceous and across the Cretaceous-Paleogene (K-Pg) boundary. Metatherian diversification patterns suggest that they were not strongly affected by a Cretaceous Terrestrial Revolution, but they clearly underwent a severe extinction across the K-Pg boundary.     

First report of a green toad (Bufo viridis sensu lato) in the Early Pleistocene of Spain: Palaeobiogeographical and palaeoecological implications

For the first time unequivocal fossil remains of a green toad (Bufo viridis s.l.) are described in the Iberian Peninsula. The fossils come from the Cueva Victoria site, a late Early Pleistocene (ca. 1.1–1.2 Ma) karstic filling in semi-arid southeastern Spain (Murcia region). By extension, other remains from two other Early Pleistocene Spanish localities, Barranco León D (ca. 1.3 Ma) and Almenara-Casablanca 3 (ca. 1.1 Ma), are cautiously attributed to the group B. viridis. The B. viridis group was previously reported with some uncertainty to the west of its current distribution area in Western Europe (Spain and France) in the Pliocene (Bufo cf. viridis) and less probably in the Early Miocene (Bufo aff. viridis). Since no osteological differences have been established between the recently described extant species of B. viridis s.l. (e.g. Bufo balearicus, Bufo siculus, Bufo boulengeri, B. viridis sensu stricto and Bufo variabilis) no precise palaeobiogeographical relationships can be drawn for the Spanish fossils. However, the occurrence of a third species of bufonid toad during the Pleistocene in the South of the Iberian Peninsula raises some interesting ecological questions in relation to the local disappearance of the green toad, which can be hypothetically linked to the intensification of the Pleistocene glacial/interglacial climate dynamic or to probable competition with another toad, Bufo calamita.     

On the spinosaurid theropod teeth from the early Barremian (Early Cretaceous) Blesa Formation (Spain)

Spinosaurids are one of the most abundant theropods from the Early Cretaceous of the Iberian Peninsula. The published fossil record consists mainly of teeth. A good example is La Cantalera-1 site in the Blesa Formation (Spain). This site represents a marshy environment with periodic droughts, resulting in a non-permanent body of water. Theropod dinosaur teeth are frequent. Seventeen spinosaurid teeth have been studied using qualitative and quantitative features, including a statistical analysis, in order to compare them with the known spinosaurid record from Spain. In La Cantalera-1 we recognise two different morphotypes: Baryonychinae indet. and Spinosaurinae? indet. Baryonychinae indet. was produced by baryonychines close to Baryonyx; teeth with similar morphology have been described in other sites from the Early Cretaceous of the Iberian Peninsula. Teeth tentatively attributed to Spinosaurinae? indet. were produced by spinosaurines close to Spinosaurus. This morphotype could provide new evidence of the presence of spinosaurines in the Early Cretaceous of the Iberian Peninsula. The small size of the teeth from La Cantalera-1 when compared with those from other sites is interpreted as the result of the conditions characteristic of the deposit area, which represents an ecosystem unable to sustain large-sized organisms.     

A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical implications

The Abelisauridae are a family of mainly Cretaceous theropod dinosaurs with a wide distribution across the Gondwanan land masses. Although their presence in Europe was reported twenty-five years ago, it has often been considered as controversial largely because of the incompleteness of the available specimens. We report here the discovery of well-preserved abelisaurid material, including a highly diagnostic braincase, at a Late Cretaceous (late Campanian) locality in the Aix-en-Provence Basin, near the eponym city in south-eastern France. A new abelisaurid taxon is erected, Arcovenator escotae gen. nov., sp. nov., on the basis of cranial and postcranial material. A phylogenetic analysis reveals that the new Abelisauridae from Provence is more closely related to taxa from India and Madagascar than to South American forms. Moreover, Genusaurus, Tarascosaurus and the previous Late Cretaceous discoveries are identified as basal abelisaurids. Contrary to previously proposed palaeobiogeographical models of abelisaurid evolution, the presence of the new taxon in Europe suggests that Europe and Africa may have played a major role in abelisaurid dispersal, which apparently involved crossing marine barriers.     

Tentaculitoid affinities of the tubeworm-like fossil Tymbochoos sinclairi (Okulitch, 1937) from the Ordovician of North America

Tymbochoos sinclairi (Okulitch, 1937) has a laminar tube structure and pseudopuncta similar to the tentaculitoids. This suggests that Tymbochoos belongs to the Tentaculitoidea Bou?ek, 1964.     

Evidence for the longest stratigraphic range of a post-Triassic Ichthyosaur: a Leptonectes tenuirostris from the Pliensbachian (Lower Jurassic) of Switzerland

A three-dimensional well-preserved ichthyosaur skull and parts of the postcranial skeleton are attributed to the species Leptonectes tenuirostris (Conybeare, 1822). It was found vertically embedded in Pliensbachian deposits representing three successive biozones (ibex to margaritatus Zone). The find is dated as early Late Pliensbachian (margaritatus Zone) by a rich ammonite and ostracod fauna. It is the first record of the genus Leptonectes from Switzerland and from the Late Pliensbachian. It is so far the best preserved and most complete ichthyosaur from this time interval worldwide. With diagnostic specimens known from the Rhaetian (Late Keuper) up to the early Late Pliensbachian (Middle Liassic), L. tenuirostris (Conybeare, 1822) has the most extensive stratigraphic range documented for any post-Triassic ichthyosaur so far.     

The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula

Despite the postulation of the “herpetofaunal stability hypothesis”, which suggests that the herpetofauna underwent relatively few changes during the Pleistocene, the amphibian and squamate faunas of western Europe are known to have progressively diminished during the Pliocene and the beginning of the Pleistocene. Iberian Early Pleistocene sites continue to document the presence of “exotic” herpetofaunal elements that are supposed to have disappeared from the mainland, such as Oriental vipers, agamid lizards, the anguids Pseudopus and Dopasia, and a representative of the green toad group (Bufo viridis sensu lato), as well as possibly gekkonids, a scincid lizard (Lygosominae), the anguid Ragesaurus and blind snake (Scolecophidia). The geographical and temporal pattern of these progressive southward withdrawals and extirpations on the Iberian Peninsula shows that extirpation events occurred in northern Spain at roughly the Olduvai paleomagnetic event and in southern Spain just before the Jaramillo paleomagnetic reversal, thus permitting us to establish a framework that can be used to complement the biochronological zonation of the Iberian Early Pleistocene.