Opportunistic exploitation of dinosaur dung: fossil snails in coprolites from the Upper Cretaceous Two Medicine Formation of Montana

Multiple associations of fossil snails with dinosaur coprolites demonstrate that snails and dinosaurs not only shared ancient habitats but were trophically linked via dinosaur dung. Over 130 fossil snails representing at least seven different taxa have been found on or within herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana. The terrestrial snail Megomphix is the most common taxon, but three other terrestrial taxa (Prograngerella, Hendersonia and Polygyrella) and three aquatic snails (Lioplacodes, ?Viviparus and a physid) also occur in coprolites. At least 46% of the shells in the faeces are whole or nearly so, indicating that most (if not all) of the snails were not ingested by dinosaurs, but were post‐depositional visitors to the dung pats. The sizeable, moist and microbially enriched dinosaur faeces would have provided both food and roosting sites for the ancient snails, and the large number of snail–coprolite associations reflect recurring, opportunistic exploitation of dung. The terrestrial taxa in the coprolites suggest that this Late Cretaceous locality included sufficiently moist detrital or vegetative cover for snails when dinosaur dung was not present. The aquatic snails probably entered the faeces during flood events. Dinosaur dung would have provided an abundant but patchy influx of resources that was probably seasonally available in the ancient environment.     

Early Cretaceous (Valanginian - Hauterivian) calcareous nannofossils and isotopes of the northern hemisphere: proxies for the understanding of Cretaceous climate

From three boreholes (DSDP Site 535; ODP Site 638; BGS borehole 81/43) of the Central Atlantic and the North Sea Basin 379 samples of early Cretaceous age (Valanginian-Hauterivian) were examined. The localities cover a S-N transect of approximately 3000 km stretching from 17°N to 40°N palaeolatitude. The distribution of calcareous nannofossils and fluctuations of the stable isotopes (δ¹³C, δ¹⁸O) have been recorded and were compared with results of recent studies. We differentiate between high nutrient indicators and oligotrophic taxa and propose a four step scheme to characterize the trophic level of the surface water. (1) High abundances of the fertility group (Biscutum constans/Zeugrhabdotus spp.) combined with a high dominance of B. constans and low abundances of Watznaueria barnesae/W. fossacincta represent a high nutrient environment (eutrophic setting). (2) High abundances of the fertility group combined with a high dominance of Zeugrhabdotus spp. and low abundances of W. barnesae/W. fossacincta reflect enhanced nutrient contents of the surface water (mesotrophic setting). (3) Enhanced abundances of the fertility group combined with high abundances of W. barnesae/W. fossacincta indicate slightly increased nutrient contents of the surface water (meso- to oligotrophic setting). (4) Low abundances of the fertility group and high abundances of W. barnesae/W. fossacincta are of low nutrient affinities (oligotrophic setting). Our estimations of seawater palaeotemperatures in combination with literature data show a distinctive trend for the Valanginian to Hauterivian interval. A general decrease of water temperature from the Valanginian to the early Hauterivian is obvious. This decrease of temperature coincides with the southward migration of the high latitudinal cold water species Crucibiscutum salebrosum to lower latitudes. Our findings shed new light on the evolution of the earliest Cretaceous climate, which may be characterized as a warm greenhouse world with interludes of short cooling.     

Oichnus Bromley Borings in the Irregular Echinoid Hemipneustes Agassiz from the Type Maastrichtian (Upper Cretaceous,The Netherlands and Belgium)

Three ichnospecies of Oichnus Bromley occur in tests of the large holasteroid echinoid Hemipneustes striatoradiatus (Leske) in the type area of the Maastrichtian (Upper Cretaceous) in The Netherlands and Belgium; Oichnus simplex Bromley (penetrative), Oichnus paraboloides Bromley (nonpenetrative and showing two distinct morphologies), and Oichnus excavatus isp. nov. (nonpenetrative). The two distinct morphologies of O. paraboloides (both shallow, one with a central boss) are gregarious, but do not occur together on the same specimens, suggesting they were generated by different taxa. Oichnus paraboloides with a central boss occurs on H. striatoradiatus from the upper Nekum Member, Maastricht Formation (Maastrichtian, Upper Cretaceous). Tests of the host echinoid are smaller in the overlying Meerssen Member, Maastricht Formation, where they are infested by O. excavatus, the largest borings considered herein, which have concave walls and a large central boss. Blisters inside tests from the Meerssen Member show that this infestation occurred when the echinoid was alive. It is postulated that producers of these borings in H. striatoradiatus may have been genetically related and increased in size during the Maastrichtian even as the host echinoids showed a size decrease. This size increase in H. striatoradiatus was genetic and cannot be related to increase in size of borings.     

Representatives of the family Bochianitidae (Ammonoidea) from the Lower Cretaceous of the Crimean Mountains

The heteromorph ammonites of the family Bochianitidae from the Lower Cretaceous of the Crimean Mountains are revised. The validity of the genus Janenschites, separated from the genus Bochianites is confirmed. The species Bochianites neocomiensis (d’Orbigny), B. goubechensis Mandov, B. levis sp. nov. and B. crymensis sp. nov. are described from the Berriasian and the species Janenschites oosteri (Sarasin et Schöndelmayer) and J. incisus sp. nov. are described from the Lower Barremian. The family Bochianitidae first appeared at the beginning of the Berriasian in the southern regions (Africa and the Crimea), and spread to the northern regions of western Europe in the Valanginian-Hauterivian.     

Pithonelloid wall-type of the Late Cretaceous calcareous dinoflagellate cyst genus Tetratropis

The late Cretaceous calcareous dinoflagellate genus Tetratropis features both a pithonelloid wall-type (evenly inclined wall-components, proven here by a polarisation optical revision) and a peridinialean paratabulation strongly suggesting a dinoflagellate origin of at least part of the Pithonelloideae. This affinity with dinoflagellates sheds more light on the palaeoecology of the Pithonelloideae (commonly termed “calcispheres”), which are characteristic of the middle to Late Cretaceous. The very short-term stratigraphic occurrence of all Tetratropis species is comparable to the distribution pattern of other calcareous dinoflagellate cyst species with a distinctive paratabulation and is thought to reflect a narrow palaeoecological niche. Tetratropis species can be interpreted either as paratabulated morphotypes of otherwise atabulate Pithonelloideae formed under exceptional palaeoenvironmental conditions or as invaders from a highly specific palaeoecological niche during short-term palaeoceanographic events probably related to the initiation of the Late Cretaceous global cooling.     

Reconstruction of Late Cretaceous coastal paleotemperature from halite deposits of the Late Cretaceous Nongbok Formation (Khorat Plateau,Laos)

Cretaceous evaporites of the Maha Sarakhan Formation in Thailand (e.g., the Nongbok Formation, Laos) have been studied for almost a century as the huge potash deposits in the world. The consistently high local paleotemperatures should lead to huge salt deposits during the evaporation process. Primary fluid inclusions in halite can provide surface brine water temperatures directly and quantitatively. Until now, there have been no data published from paleotemperature of primary fluid inclusions of Cretaceous halite. The non-marine halite from the Cretaceous Nongbok Formation (Laos) precipitated from shallow brine waters with temperatures of 17.7–42.3 °C.     

Fossil evidence for a herbaceous diversification of early eudicot angiosperms during the Early Cretaceous

Eudicot flowering plants comprise roughly 70% of land plant species diversity today, but their early evolution is not well understood. Fossil evidence has been largely restricted to their distinctive tricolpate pollen grains and this has limited our understanding of the ecological strategies that characterized their primary radiation. I describe megafossils of an Early Cretaceous eudicot from the Potomac Group in Maryland and Virginia, USA that are complete enough to allow reconstruction of important life-history traits. I draw on quantitative and qualitative analysis of functional traits, phylogenetic analysis and sedimentological evidence to reconstruct the biology of this extinct species. These plants were small and locally rare but widespread, fast-growing herbs. They had complex leaves and they were colonizers of bright, wet, disturbance-prone habitats. Other early eudicot megafossils appear to be herbaceous rather than woody, suggesting that this habit was characteristic of their primary radiation. A mostly herbaceous initial diversification of eudicots could simultaneously explain the heretofore sparse megafossil record as well as their rapid diversification during the Early Cretaceous because the angiosperm capacity for fast reproduction and fast evolution is best expressed in herbs.     

Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation on Bornholm, Denmark

The dinosaur fauna of the palynologically dated lower Berriasian Skyttegård Member of the Rabekke Formation on the Baltic island of Bornholm, Denmark, is represented by isolated tooth crowns. The assemblage is restricted to small maniraptoran theropods, assigned to the Dromaeosauridae incertae sedis and Maniraptora incertae sedis. The dromaeosaurid teeth are characterized by their labiolingually compressed and distally curved crowns that are each equipped with a lingually flexed mesial carina and a distinctly denticulated distal cutting edge. A morphologically aberrant tooth crown (referred to as Maniraptora incertae sedis) has triangular denticles of uneven width, a feature occasionally found in Upper Cretaceous hesperornithiform toothed diving birds, but also in premaxillary teeth of the velociraptorine Nuthetes from the Lower Cretaceous of England.     

Mesofossils with platanaceous affinity from the Dakota Formation (Cretaceous) in Kansas, USA

The Platanaceae holds a basal position in the phylogeny of eudicots and therefore is of great interest to angiosperm systematists. The fossil record of the family is found in strata ranging from the Cretaceous to Recent in America, Europe and Asia. The research on the Platanaceae in the Dakota Formation can be traced back to 19th century; however, mesofossils of reproductive organs of the Platanaceae were never reported in the Midwest of North America before. This paper reports several specimens of Friisicarpus (Platanaceae) from the Dakota Formation in Kansas, USA. It complements the existing fossil records, and provides more information on reproductive biology of the family. The comparison with similar fossils from eastern North America and Europe provides some hints on biostratigraphy of the Cretaceous.     

New radiolarians from the earliest Cretaceous of the Sultanate of Oman (Wahrah Formation,Jebel Buwaydah)

In a section from the central Oman Mountains (Wahrah Formation, Jebel Buwaydah) well preserved radiolarians were extracted. Stratigraphically they represent typical earliest Cretaceous species (Bernasian). Besides well known Tethyan species allowing an exact age determination, unknown morphotypes are common in some samples. Some of these represent new species, which are described herein. The composition and diversity of the faunas hints to a depositional environment in the lower bathyal.     

New material of Xyelydidae (Hymenoptera) from the Lower Cretaceous Yixian Formation of China

Chifengilyda robusta, a new extinct sawfly genus and new species assigned to Xyelidae, is described and figured here based on one fossil specimen from the Lower Cretaceous Yixian Formation of Ningcheng in Inner Mongolia, China. Chifengilyda robusta n. gen. n. sp. differs from other xyelydids in having a forewing with pterostigma completely sclerotized, Sc closer to C, Sc1 about 3 times as long as Sc2, Sc1 intersecting C far beyond Rs base, 2r-m postfurcal and cell 1mcu relatively large. This new taxon further enriches the diversity of Xyelydidae in the Early Cretaceous, and even enhances our understanding of the evolution and inter-genera and interspecies relationships of this family.     

Specialization of bone structure in Pachyvaranus crassispondylus Arambourg, 1952, an aquatic squamate from the Late Cretaceous of the southern Tethyan margin

The histological organization of the vertebrae of the Maastrichtian squamate Pachyvaranus crassispondylus Arambourg, 1952 , was compared to that of various extant squamates, in order to further document the causes and functional consequences of the so-called 'pachyostosis', frequently observed in Late Cretaceous squamates. The vertebrae of Pachyvaranus are composed of the same basic bone tissue types as those of extant lizards and snakes. In particular, periosteal cortices are made of a pseudolamellar (or 'parallel-fibred') tissue, with radial vascular canals, Sharpey's fibres and conspicuous cyclic growth marks that are strictly identical to that found in extant varanids. Conversely, the vertebrae of Pachyvaranus are extremely compact, whereas those of extant squamates are very cancellous and lightly built. This difference is due to the absence in Pachyvaranus of a broad internal resorption field that, in extant lizards and snakes, transforms compact cortices into loose spongy formations. This absence of inner bone resorption typically corresponds to an osteosclerotic process. In Pachyvaranus , cortical hyperplasy, or pachyostosis stricto sensu , was restricted to the walls of the neural spine. Extreme vertebral porosity is likely to be a primitive condition in squamates, because all lizards and snakes examined in this study display this feature. Therefore, the high vertebral compactness observed in Pachyvaranus would be a derived condition arising from the loss (or de-differentiation) of a morphogenetic process: the broad internal resorption of the vertebrae. Possible palaeoecological bearings of these results are discussed.     

A surprisingly derived hydrometrid (Hemiptera,Gerromorpha) from the Crato Formation,Early Cretaceous of Brazil

In the present paper, we review the fossil record of the Hydrometridae (Hemiptera, Gerromorpha) and present a new species from the Early Cretaceous Crato Formation of Northeastern Brazil, Christometra paradoxa gen. et sp. nov. This species is based on a new specimen (a female), as well as a previously figured one (a male), providing a rare case of preservation of sexually dimorphic features in the fossil record. This is the third species coming from this deposit, which is Aptian-Albian in age and the oldest deposit to have yielded hydrometrids so far. Only five other Mesozoic species are known, being slightly younger in age (Cenomanian). So far, phylogenetic analyses have recovered Cretaceous hydrometrids as basal relative to Cenozoic genera but, Christometra paradoxa exhibits several advanced characteristics that unite it in a clade together with the extant genera Hydrometra and Bacillometroides, in a more derived position than any previously known fossil hydrometrid.

The present publication is registered in the Official Register of Zoological Nomenclature (Zoobank), under the registration number http://zoobank.org/urn:lsid:zoobank.org:pub:3CFA88AB-3CBC-4CCC-8196-698ECC863947. The registration number for the nomenclatural act of the genus is http://zoobank.org/urn:lsid:zoobank.org:act:84744426-1259-4864-8E3F-E43E0DAB2021, and that of the species is http://zoobank.org/urn:lsid:zoobank.org:act:23700AB2-F7AD-4F50-A5E7-CB28868079B2.     

Hadrosauroid Dinosaurs from the Late Cretaceous of the Sultanate of Oman

Fragmentary post-cranial remains (femora, tibia, vertebrae) of ornithischian dinosaurs from the Late Cretaceous of the Sultanate of Oman are described and referred to hadrosauroids. The specimens come from the Al-Khod Conglomerate, of latest Campanian to Maastrichtian age, in the north-eastern part of the country. Although the fragmentary condition of the fossils precludes a precise identification, various characters, including the shape of the fourth trochanter of the femur and the morphology of its distal end, support an attribution to hadrosauroids. With the possible exception of a possible phalanx from Angola, this group of ornithopod dinosaurs, which apparently originated in Laurasia, was hitherto unreported from the Afro-Arabian plate. From a paleobiogeographical point of view, the presence of hadrosauroids in Oman in all likelihood is a result of trans-Tethys dispersal from Asia or Europe, probably by way of islands in the Tethys shown on all recent paleogeographical maps of that area. Whether hadrosauroids were widespread on the Afro-Arabian landmass in the latest Cretaceous, or where restricted to the « Oman island » shown on some paleogeographical maps, remains to be determined.     

Basal titanosauriform (Dinosauria, Sauropoda) teeth from the Lower Cretaceous Yixian Formation of Liaoning Province, China

The Yixian Formation of Liaoning Province, People's Republic of China has yielded a diverse fauna of non-avian dinosaurs, but is dominated by small-bodied taxa. Here, we describe a series of isolated teeth from the Lujiatun Beds of the formation that are referable to a basal titanosauriform sauropod. Some of the teeth possess a distinctive circular boss on the lingual surface, which suggests that they are referable to cf. Euhelopus sp. This identification provides some additional support for biostratigraphical correlations between the Jehol Group and the Mengyin Formation of Shandong Province that suggest an Early Cretaceous age for the latter unit. Moreover, the titanosauriform affinities of the teeth provide further evidence for the dominance of this sauropod clade in eastern Asia during the Cretaceous.     

Phaeodarian Radiolaria from the Upper Cretaceous beds of central Japan

Upper Cretaceous Phaeodarea (Radiolaria) were recovered from the Shoya Formation, which crops out 100 km northwest of Tokyo, central Japan. The Shoya Formation consists of about 600 m-thick marine sedimentary rocks, represented by alternating beds of sandstone and mudstone, which are overlain by about 10 m of Phaeodarian-bearing siliceous mudstone. The latter is assigned to the Upper Cretaceous (late Campanian to early Maastrichtian) based on the associated Polycystine Radiolarian fossils. In spite of the poor general preservation of nearly all Phaeodarian specimens as recrystallized quartz infillings, three new Phaeodarian species, Challengeranium cretaceum, Challengeron paleotriangulum, and Medusetta fossilis, were identified on the basis of their shape and ornamentation. Our finding, together with two other very recent reports of fossil Phaeodarians clearly document that the origin of Phaeodarian Radiolarians can be extended back to at least the Upper Cretaceous.     

New mammal remains from the Late Cretaceous Bostobe Formation (Northeast Aral Sea Region,Kazakhstan)

Four recently collected mammal specimens from the Upper Cretaceous (Santonian–?Campanian) Bostobe Formation in the northeastern Aral Sea Region, Kazakhstan are attributed to Asioryctitheria indet. (an edentulous dentary fragment) and the zhelestid Parazhelestes sp. cf. P. mynbulakensis (a maxillary fragment with a double-rooted canine, an M1, and a dentary fragment including m3). These new records double the known mammal fauna from this formation, which previously included the zhelestid Zhalmouzia bazhanovi and Zhelestidae indet. The taxonomic and ecological structure of the mammal assemblage from the Bostobe Formation can, on present evidence, be considered close to the other eutherian dominated Late Cretaceous mammal assemblages of Central Asia. This region is important in particular in the search for Late Cretaceous ancestors of crown-group eutherian mammal clades (Placentalia).     

An enigmatic Ephedra-like fossil lacking micropylar tube from the Lower Cretaceous Yixian Formation of Liaoning,China

Angiosperms and gymnosperms are two well-separated groups in seed plants according to the current understanding. The huge gap between these two groups constitutes a serious threat against the Darwinism, which expects a continuous transitional series between them. The Lower Cretaceous Yixian Formation of Liaoning, China is famous for its megafossil angiosperms, including some early angiosperms and putative gnetalean plants. Here we document another Ephedra-like fossil plant, Pseudoephedra n. gen. n. sp., from the Yixian Formation on the basis of light microscopic (LM) and scanning electron microscopic (SEM) observations. Although its general morphology demonstrates a great resemblance to Ephedra, the expected micropylar tube characteristic of Ephedra is missing in Pseudoephedra. Instead a solid projection is seen on the top of the female parts. Such a puzzling character combination makes Pseudoephedra perplexing in seed plant phylogeny. If put in Ephedraceae (Gnetales), Pseudoephedra would destroy the only synapomorphy (micropylar tube) of the BEG clade. If put in angiosperms, Pseudoephedra would bridge the formerly huge gap between gymnosperms and angiosperms. Apparently, further investigation is needed to clarify the uncertain position of Pseudoephedra.     

On ichthyosaur remains from the Cretaceous of the Voronezh Region

A new ichthyosaur, Platypterygius ochevi sp. nov., from the Albian-Cenomanian of the Voronezh Region is described based on a partial forefin. It differs from congeners in the very large facet for the lageniformis on the humerus.     

Extramorphological Features of Sauropod Dinosaur Tracks in the Uhangri Formation (Cretaceous), Korea

Sauropod footprints in the Uhangri Formation (Cretaceous) of Korea exhibit an unusual pattern of morphology, with the interior of each print partitioned into a series of pockets by conspicuous radial crests. The crests are evidently extramorphological features and have been interpreted as upwellings of sediment extruded through the floor of the footprint following its fracture by impact of the trackmaker's foot. That explanation entails some inconsistencies, and an alternative explanation is proposed here. The alternative explanation envisages delamination of a superficial sheet of sediment that was lifted into a canopy, which subsequently collapsed in radiating folds. The superficial sheet of sediment might have been lifted by either or both of two mechanisms—by adhering to the underside of the trackmaker's foot or by being forced upward into a blister-like dome by the backflow of water previously displaced by impact of the trackmaker's foot. These alternative explanations draw attention to minor morphological features that were previously unexplained.