Biostratigraphic and biogeographic implications of a hadrosaurid (Ornithopoda: Dinosauria) from the Upper Cretaceous Almond Formation of Wyoming,USA

The results of Barnum Brown's 1937 expedition to the Almond Formation of Wyoming consisted of two unidentified ceratopsian skulls and a partial hadrosaurid specimen (AMNH 3651). The hadrosaurid is here attributed to the Maastrichtian genus Saurolophus, verifying previous biostratigraphic correlations of this formation using ammonite zones. Fossiliferous lower Maastrichtian formations occurring latitudinally between those of Alberta, Canada, and southwestern Texas, USA, such as the Almond Formation, are essential for testing the effects and duration of apparent hadrosaurid faunal segregation earlier in the Campanian, and indirectly aiding in the placement of faunal boundaries that are currently unknown for the late Campanian. The discovery of Saurolophus in Wyoming, a close relative of the Campanian genus Prosaurolophus, affirms that the segregation of hadrosaurid faunas established in the late Campanian (～75 Ma) continued for at least 3 million years. Combining occurrences of Saurolophus from Mongolia and the Moreno Formation of California with those of Alberta, Canada, this genus appears to have had one of the largest geographic ranges of any equivalent clade of hadrosaurid dinosaur, although species level distributions are still uncertain.     

Latest Campanian and Maastrichtian Siderolitidae (larger benthic foraminifera) from the Pyrenees (S France and NE Spain)

The siderolitids from the uppermost Campanian and Maastrichtian deposits of the Pyrenees have been re-studied. This has revealed a high diversity and rapid replacement of taxa, confirming the group as a good tool for high resolution biostratigraphy. Two genera have been found in the uppermost Campanian–Maastrichtian interval in the Pyrenean deposits: Siderolites Lamarck, and Wannierina Robles-Salcedo. Siderolites, with canaliferous spines or denticulate periphery, is represented by four species replacing each other from the latest Campanian to Maastrichtian: Siderolites praecalcitrapoides (latest Campanian), S. pyrenaicus sp. nov. (early Maastrichtian), Siderolites calcitrapoides (late Maastrichtian) and Siderolites denticulatus (late Maastrichtian). Wannierina is characterised by well-developed keels and ramified marginal canals. Two species of Wannierina have been identified and they succeeded one another from latest Campanian to early Maastrichtian: Wannierina vilavellensis sp. nov. (latest Campanian) and Wannierina cataluniensis (early Maastrichtian). The species of the genus Siderolites inhabited shallow waters of tropical to subtropical platforms with moderate-to-high water-energy conditions and those of the genus Wannierina are typical of deep–water low-energy environments but still in the eutrophic zone.     

Conifer fossil woods from the Santa Marta Formation (Upper Cretaceous), Brandy Bay,James Ross Island,Antarctica

Conifer fossil woods from the Santonian to lower Campanian Santa Marta Formation are anatomically studied in detail. The wood flora is dominated by conifers, mainly Araucariaceae. Conifer fossil woods represent 84% of the samples (26 of 31) of the assemblage. The remaining samples are dicotyledon woods which are not included in this study. Woods are mostly calcified and preservation is often excellent. Fossil woods include three species of Agathoxylon, Phyllocladoxylon antarcticum, Cupressinoxylon hallei and a new species of Cupressinoxylon, C. rotundum Pujana sp. nov. The conifer-dominated forests are consistent with the pollen and macrofloras previously described from sediments of the same stratigraphic unit. This fossil wood assemblage contributes to understand the floristic changes that took place in the Cretaceous forests.     

Calcareous nannofossils biostratigraphy of the Late Campanian–Early Maastrichtian Age, Block 16, Jiza’-Qamar Basin, Eastern Yemen

One hundred and thirty nine samples have been studied from the Late Campanian–Early Maastrichtian of three deep wells drilled in Jiza’-Qamar Basin, Eastern Yemen to determine the calcareous nannofossil zones and the age of the sediments. Forty-seven calcareous nannofossil species were identified and four biozones were determined in the present study (CC21–CC24). These biozones are assigned to the Late Campanian–Early Maastrichtian ages. Most of the studied species in this work refer to tropical–subtropical environment. The Campanian–Maastrichtian Boundary was determined in Al-Fatk well based on the last occurrence of Eiffelithus eximus and the last occurrences of Uniplanarius sissinghii and Uniplanarius trifidus.     

Pararhabdodon isonensis and Tsintaosaurus spinorhinus: a new clade of lambeosaurine hadrosaurids from Eurasia

We present new anatomical information showing that Koutalisaurus kohlerorum, from the Maastrichtian of Lleida Province, northeastern Spain, is most probably the junior synonym of Pararhabdodon isonensis from the same region. Dentary and maxillary characters previously considered as autapomorphies of K. kohlerorum and P. isonensis, respectively, are shown to be synapomorphies uniting the latter with Tsintaosaurus spinorhinus from the Campanian of the Wangshi Group, Shandong Province, China.This study provided conclusive evidence of the presence of the Lambeosaurinae in Europe. Tsintaosaurus spinorhinus and Pararhabdodon isonensis were inferred to form a clade of basal lambeosaurines characterized by a maxilla with an elevated articular facet for the jugal (continuous with the ectopterygoid ridge) and an extremely medially projected symphyseal region of the dentary. This clade originated in Asia during the middle or late Campanian. Pararhabdodon isonensis or its ancestors migrated from Asia to the Iberian island of the European archipelago. Reconstruction of ancestral areas by Fitch parsimony attributes the European occurrence of P. isonensis to a single dispersal event from Asia no later than middle to late Campanian.     

Petreşti-Arini – An important but ephemeral Upper Cretaceous continental vertebrate site in the southwestern Transylvanian Basin,Romania

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 Haţeg Island). A series of artificial outcrops recently exposed during a hydroelectric project, the Petreşti-Arini section near Sebeş in the Transylvanian Basin, records a 400+ meter sequence documenting the transition from fully marine to terrestrial environments during the Campanian–Maastrichtian. Calcareous nannofossil biostratigraphy indicates that the lower marine beds in this section, part of the uppermost Bozeş Formation, can be assigned to the CC22 biozone, corresponding to the lower–mid upper Campanian. These beds smoothly transition, via a brackish-water unit, into the fully continental Maastrichtian Sebeş Formation. Dinosaur and pterosaur fossils from the uppermost Bozeş Formation can be assigned a late Campanian age making them the oldest well-dated terrestrial fossils from the Haţeg Island, and indicating that the classic Haţeg dinosaur fauna was becoming established by this time, coincident with the first emergence of widespread land areas. Vertebrate fossils occur throughout the overlying Sebeş 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 Petreşti-Arini section preserves a somewhat unusual near-shore environment, or the earliest Haţeg 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 Petreşti-Arini local fauna. Support for any one of these alternative hypotheses requires further data from Petreşti-Arini as well as from the larger Transylvania area.     

Upper Maastrichtian shallow marine environments and neoselachian assemblages in North Iberian palaeomargin (Castilian Ramp,Spain)

Upper Maastrichtian deposits formed in a nearshore subtidal environment within the Valdenoceda Formation (Castilian Ramp, North Iberian margin) are described together with two recently found selachian assemblages. Rare earth element concentrations (REE) have been used to assess the degree of taphonomic mixing and reworking, observing that it is minor or non-existent, and differences in degree of preservation and ecologic mixing can be explained by biostratinomic processes. The patterns of REE also helped to obtain a better understanding of the depositional environment, including the diagenetic history from burial to final degree of bone preservation.The fossil assemblages here described are close to that of the late Maastrichtian of Albaina (in the enclave of Condado de Treviño, Burgos), both in the Basque-Cantabrian Region, but their age may be slightly older (early late Maastrichtian). In total, the new assemblages consist of 17 taxa, assigned to 11 genera of shallow-water dwellers combined with individuals from the outer shelf. They represent cosmopolitan taxa (Squalicorax pristodontus, Serratolamna serrata and Rhombodus binkhorsti) together with local species (Rhinobatos echavei, Rhinobatos ibericus). Although there are not significant differences between Albaina and Quintanilla la Ojada faunas, the new assemblages add interesting taphonomic and geochemical information to the few existing uppermost Cretaceous deposits with fossil sharks in southwestern Europe.     

Constraining the age of the last marine sediments in the late Cretaceous of central south Pyrenees (NE Spain): Insights from larger benthic foraminifera and strontium isotope stratigraphy

The uppermost Cretaceous (upper Campanian–Maastrichtian) marine deposits of the central south Pyrenees host a rich larger benthic foraminiferal fauna and several rudist-rich levels. These marine deposits are directly overlain by the continental facies of the Arén and Tremp Formations, which are famous for their fossil dinosaur remains. Larger benthic foraminiferal distribution documents an important faunal turnover in all the carbonate platform environments within the photic zone, from open marine to littoral areas. Biostratigraphy indicates that this turnover occurred close to the Campanian-Maastrichtian boundary. This is also confirmed by strontium isotope stratigraphy which indicates an earliest Maastrichtian age for the appearance of the larger benthic foraminiferal assemblage constituted by Lepidorbitoides socialis, Clypeorbis mammillata, Wannierina cataluniensis, Orbitoides gruenbachensis, Siderolites aff. calcitrapoides, Fascispira colomi, Omphalocyclus macroporus and Laffiteina mengaudi. In particular, a numerical age of 71 Ma is obtained for the Hippurites radiosus level, just a few meters below the first continental deposits of the Arén sensu stricto Formation. The youngest marine sediments of the central south Pyrenees are early Maastrichtian in age. This is also an important constraint for the age of the end-Cretaceous dinosaur fossil localities of the Tremp basin.     

Dentaries of a caenagnathid (Dinosauria: Theropoda) from the Nemegt Formation of the Gobi Desert in Mongolia

A symphyseal region of the fused dentaries of a caenagnathid theropod is described from the Upper Cretaceous Nemegt Formation at the Bugin Tsav locality in the Mongolian Gobi Desert. In contrast to the high diversity of Caenagnathidae in the upper Campanian to Maastrichtian in North America, only specimens of a single caenagnathid, Elmisaurus raurus, have been reported in the coeval strata in Asia. Although dentaries are commonly-found bones in the fossil record of Caenagnathidae, the present specimen is the first discovery of caenagnathid dentaries from the upper Campanian to Maastrichtian in Asia. The Nemegt Formation is unique for its diverse oviraptorosaurian fauna that includes both Caenagnathidae and Oviraptoridae as well as the non-caenagnathoid Avimimus portentosus. Hypothesized coexistence of eolian and fluvial environments in the Gobi Basin during the deposition of the Nemegt Formation might explain such co-occurrence of Caenagnathidae and Oviraptoridae.     

Dinosaur eggshells from the Santonian Milk River Formation of Alberta,Canada

The North American fossil record of dinosaur eggshells for the Cretaceous is primarily restricted to formations of the middle (Albian–Cenomanian) and uppermost (Campanian–Maastrichtian) stages, with a large gap in the record for intermediate stages. Here we describe a dinosaur eggshell assemblage from a formation that represents an intermediate and poorly fossiliferous stage of the Upper Cretaceous, the Santonian Milk River Formation of southern Alberta, Canada. The Milk River eggshell assemblage contains five eggshell taxa: Continuoolithus, Porituberoolithus, Prismatoolithus, Spheroolithus, and Triprismatoolithus. These ootaxa are most similar to those reported from younger Campanian–Maastrichtian formations of the northern Western Interior than they are to ootaxa reported from older middle Cretaceous formations (i.e., predominantly Macroelongatoolithus). Characteristics of the Milk River ootaxa indicate that they are ascribable to at least one ornithopod and four small theropod species. The taxonomic affinity of the eggshell assemblage is consistent with the dinosaur fauna known based on isolated teeth and fragmentary skeletal remains from the formation, although most ornithischians and large theropods are not represented by eggshell. Relative to the Milk River Formation eggshell, similar oospecies occurring in younger Cretaceous deposits tend to be somewhat thicker, which may reflect an increase in body size of various dinosaur lineages during the Late Cretaceous.     

A neoselachian shark fauna from the Late Cretaceous of Senegal

A selachian fauna is described for the first time from the Late Cretaceous (Campanian–Maastrichtian) of Senegal. So far, the Campanian Paki Formation has only yielded a single tooth of Rhombodus sp. whereas the Cap de Naze Formation has yielded a more diverse fauna including juvenile Cretalamna cf. Cretalamna biauriculata, Serratolamna serrata, Carcharias cf. Carcharias heathi, ?Carcharias sp., Squalicorax pristodontus, Schizorhiza stromeri, Parapaleobates sp., Rhombodus binkhorsti and Rhombodus andriesi. Teeth of juvenile Cretalamna largely dominate the assemblage. Such an assemblage confirms a Late Maastrichtian age for the unit 3 in the Cap de Naze Formation. The assemblage, although composed of cosmopolitan taxa, is similar to the contemporaneous selachian assemblage from the phosphates of Morocco.     

Pelvic elements of the giant bird Gargantuavis from the Upper Cretaceous of Cruzy (southern France), with remarks on pneumatisation

A synsacrum fragment and an incomplete ilium of the giant bird Gargantuavis philoinos are described from the upper Campanian/lower Maastrichtian Montplo-Nord locality at Cruzy (Hérault, southern France). The ilium provides new information about the pelvic morphology of Gargantuavis. Both the synsacrum and ilium are extensively pneumatised. This extensive pneumatisation may be a plesiomorphic feature, but may also have had a weight-reduction function in a large flightless bird.     

An Ichthyornis-like bird from the earliest Late Cretaceous (Cenomanian) of European Russia

Ornithuromorph birds (the clade which includes modern avian radiation) first appeared in the Early Cretaceous in Asia and achieved a great diversity during the latest ages of the Late Cretaceous (Campanian and Maastrichtian). The evolutionary history of orithuromorphs during the first 17 MYAs of the Late Cretaceous (Cenomanian to Santonian ages) remains very poorly known, as the fossil record for this time interval is largely restricted to several isolated finds of the classic avian genus Ichthyornis in North America. Here we describe an isolated distal tibiotarsus of an evolutionary advanced bird, morphologically similar to Ichthyornis, from the middle Cenomanian of Saratov Province, European Russia. This is the first documentation of an Ichthyornis-like bird in the Old World. The find further constitutes only the second pre-Campanian record of the Late Cretaceous Ornithuromorpha in Eurasia, the second record of Cenomanian birds in Russia. This discovery shows that Ichthyornis-like birds enjoyed a wide geographical distribution as early as the beginning of the Late Cretaceous. Given that the earliest and the most primitive ornithuromorph birds are known from Asia, the new find supports a Eurasian origin for Ichthyornithidae.     

Titanoceratops ouranos, a giant horned dinosaur from the late Campanian of New Mexico

At the end of the Cretaceous, 65.5 million years ago, the giant ceratopsids Triceratops and Torosaurus dominated North America’s dinosaur fauna. The origins of these giant ceratopsids, the Triceratopsini, are poorly understood. This paper describes Titanoceratops ouranos, a giant ceratopsid from the late Campanian (73-74 Ma) of New Mexico, and the earliest known triceratopsin. The holotype was previously interpreted as an aberrant and exceptionally large specimen of Pentaceratops sternbergi, but the animal does not show the diagnostic features of Pentaceratops. Instead, cladistic analysis shows that Titanoceratops is the sister taxon of a clade formed by Eotriceratops, Triceratops, and Torosaurus. With an estimated mass of 6.5 tons, Titanoceratops is among the largest dinosaurs known from the Campanian of North America, and rivaled Triceratops in size. The recognition of Titanoceratops suggests that giant chasmosaurines evolved once, among the Triceratopsini, and that the group evolved large size five million years earlier than previously thought. The giant horned dinosaurs probably originated in the southern part of the North American continent during the Campanian but only became widespread during the Maastrichtian.     

A new species of Pluridens (Mosasauridae: Halisaurinae) from the upper Campanian of Southern Nigeria

The Upper Cretaceous of Africa has produced a diverse fauna of mosasaurs, including the highly specialized, long-jawed Pluridens. The type of Pluridens walkeri comes from the Maastrichtian Farin-Doutchi Formation of Niger, with a second, referred specimen coming from the Campanian section of the Campanian-Maastrichtian Nkporo Shale near Calabar, in southern Nigeria. Comparisons of this referred specimen with the holotype suggest that it represents a distinct and more primitive species. The Calabar jaw resembles P. walkeri in being long and narrow anteriorly with a shallow subdental shelf, and in having small, numerous, recurved teeth with medially positioned replacement pits. However, it lacks many of the derived features that characterize Pluridens walkeri, such as the extremely long and straight jaw, the extreme lateral protrusion and subcircular section of the dentary, strong transverse expansion of the dental thecae, and extreme reduction and increase in number of the teeth. The Calabar Pluridens is therefore referred to a new species, Pluridens calabaria. Following recent studies, Pluridens is considered to represent a highly derived member of the Halisaurinae. The marked differences between the Campanian and Maastrichtian species of the genus underscore the rapid pace of mosasaur evolution during the Cretaceous.     

A new titanosaur (Dinosauria,Sauropoda) from the Upper Cretaceous of Lo Hueco (Cuenca,Spain)

The upper Campanian-lower Maastrichtian site of Lo Hueco (Cuenca, Spain) has provided a set of well-preserved partial skeletons in anatomical connection or with a low dispersion of their skeletal elements. One partial skeleton is herein described and a new titanosaurian sauropod is established, Lohuecotitan pandafilandi. This titanosaur is diagnosed by eight autapomorphic features: dorsally and ventrally widened or bifurcated posterior centrodiapophyseal lamina in anterior and middle dorsal vertebrae; short postspinal lamina with a transversely expanded distal end represented by smooth scars in the dorsal vertebrae; anteriormost caudals with the medial spinoprezygapophyseal and medial spinopostzygapophyseal laminae ventrally connected with the prespinal and postspinal laminae, respectively; anterior caudal neural spines with a dorsal projection of the prespinal and postspinal laminae; anterior caudal neural spines bears a “greek-cross”-like cross-section; middle caudal centra having two round and rough structures in the dorsal edge of the posterior articulation, which extends to the dorsal surface of the centrum; the articular ends of the rami of the haemal arches are divided in two articular surfaces; and tuberosity between the anterior and the lateral trochanter of the fibula. The herein performed phylogenetic analysis considered L. pandafilandi as a member of Lithostrotia more derived than Malawisaurus. The known palaeodiversity of the Late Cretaceous Ibero-Armorican titanosaurs is increasing, and further analyses focused on this group will be necessary to better understand the evolutionary history of European titanosaurs and to clarify their relationships within Titanosauria.     

Morphometric analysis of the genus Omphalocyclus from the Late Cretaceous of Turkey: new data on its stratigraphic distribution in Mediterranean Tethys and description of two new taxa

Omphalocyclus is an orbitoidal taxon poorly known from the Late Cretaceous benthonic foraminiferal inventory of the Tethyan realm. Apart from its debatable diagnosis only in the late Maastrichtian of western Tethys, the genus has also been discovered in much older beds in association with Orbitoides and Lepidorbitoides having rather primitive developmental stages. The morphometric analysis of A-forms in successive assemblages, ranging in age from (late) Campanian to terminal Maastrichtian, enables the documentation of phylogenetic changes for the first time. The most conspicuous change is found to be the general increase in the size of the embryon, which on average doubles by the end of the Maastrichtian. This trend is followed by the increase in the number of epi-embryonic chamberlets, which is however, not as significant as the former parameter. Omphalocyclus in the stratigraphically lowermost populations has mainly three to four primary epi-embryonic and no accessory epi-embryonic chamberlets. With the introduction of radial stolons, which seems to have taken place in horizons referable to the Gansserina gansseri Zone, only several accessory epi-embryonic chamberlets arise from the tritoconch. Instead, epi-embryonic chamberlets become rather larger in size and they also cover a wider portion of the embryon along its thick outer wall. Considering the suitable change in embryon size, and also some other morphologic features in successive populations, two new species, O. anatoliensis sp. nov. and O. cideensis sp. nov. are erected in late Campanian and late Campanian-early Maastrichtian populations respectively. The most advanced specimens in the late Maastrichtian are attributed to the long-known species O. macroporus [Lamarck, J.B., 1816. Histoire naturelle des animaux sans vertèbres, vol. 2. Verdière, Paris, pp. 1–568]. Thus, a tentative subdivision scheme comprising three successive species, O. anatoliensis – O. cideensis – O. macroporus, is proposed.