Structure of fossil dinosaur eggshell from the Aix Basin,France

At least four types of dinosaur eggshell are distinguished among samples of fossil eggshell from Late Cretaceous deposits of southern France. Recognition is based on shell microstructure, porosity, and shell thickness. Estimated values of water vapour conductance for these dinosaur eggshells are much greather than predicted for avian eggs, suggesting that the eggs were incubated under conditions of high humidity, such as would occur underground.     

Sediment and eggshell interactions: using abrasion to assess transport in fossil eggshell accumulations

Recent excavation at the Late Cretaceous Egg Mountain locality in Montana revealed abundant Spheroolithus eggshell fragments. The depositional history of the fragments is problematic due to the vertical dispersal of eggshell in homogeneous mudstone. The results of previous studies assessing eggshell transport are based on thin, modern eggshell occurring on well-defined horizons. Therefore, the assessment of depositional history required a different method. Chicken (Gallus gallus domesticus) and ostrich (Struthio camelus) eggshells were placed in a tumbler with water and quartz sand for 168–504 h to simulate transport. The resulting wear on these fragments, revealed under scanning electron microscope, was compared to unabraded eggshell. In addition, the modern eggshell was compared to fossil eggshells from a nesting site, crevasse splay and channel deposits, and Egg Mountain. Fossil eggshells from high-energy paleoenvironments display similar edge rounding to modern eggshell placed in the tumbler. In contrast, eggshell from Egg Mountain lacked edge rounding and resembled fossil eggshell from a nesting site and unabraded modern eggshell, suggesting that the Egg Mountain locality represents a parautochthonous assemblage. This study indicates that sediment interaction leaves distinct patterns of abrasion on eggshell, which may be useful for assessing transport of eggshell at fossil localities.     

Parataxonomy and palaeobiogeographic significance of dinosaur eggshell fragments from the Upper Cretaceous strata of the Cauvery Basin,South India

Abstract

In the past, fossilised dinosaur eggshells have been extensively documented from the Upper Cretaceous Lameta Formation of Central India and as many as nine oospecies are known at present from this formation. Compared to this, only one dinosaur oospecies has been described from the Cretaceous succession of the Cauvery Basin. However, the first fossil egg from India, identified as a chelonian egg, was documented from the Aptian – Albian Karai Formation of the Cauvery Basin in 1957. Following this, a solitary titanosaurid dinosaur egg was described from the Upper Cretaceous (Lower Maastrichtian) Kallankuruchhi Formation, Cauvery Basin in 1996. More recently, we have recovered isolated eggshell fragments from the marine part of the Upper Cretaceous (Late Maastrichtian) Kallamedu Formation. Based on eggshell morphology, microstructure and ultrastructure, these eggshell fragments are assigned to the oospecies Fusioolithus baghensis. The new find from the Cauvery Basin is important from palaeobiogeographic point of view as the oofamily Fusioolithidae is found in the Upper Cretaceous strata of India, France, Argentina and Morocco. Based on the common occurrence of similar oospecies in South America, Africa, Europe and India, a Late Cretaceous palaeobiogeographic connection between India and South America as well as Europe via Africa is suggested.     

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.     

Titanosaur eggshells from the Tremp Formation (Upper Cretaceous,Southern Pyrenees,Spain)

Hundreds of megaloolithid eggshells have been found at the Costa de la Coma site (Tremp Fm, South-Central Pyrenees, Spain), the microstructural features (spherulitic eggshell units, thickness, ornamentation and pore system) of which agree with the Megaloolithus oogenus diagnosis. Comparisons with other megaloolithids from the Tremp Basin, southern France and South America, however, showed that these eggshells share many features with the South American oospecies Megaloolithus patagonicus. A cladistic analysis of the qualitative eggshell features of the main dinosaur taxa was performed. The results grouped the Costa de la Coma eggshells with Megaloolithus patagonicus, forming a clade separated from other megaloolithids, and revealed a polytomy of megaloolithids (associated with sauropods) and spheroolithids (associated with hadrosaurs). The finding of Megaloolithus cf. patagonicus in the Tremp Fm agrees with skeletal evidence provided by four indeterminate titanosaur forms from the Formation's late Cretaceous deposits. Other Megaloolithus oospecies of the Tremp Fm may belong to hadrosaurs, the skeletal remains of which are also abundant in the Tremp Basin.     

Electron backscatter diffraction (EBSD) analysis of maniraptoran eggshells with important implications for microstructural and taphonomic interpretations

Electron backscatter diffraction (EBSD) is a useful tool for gathering crystallographic information from carbonate biominerals because it maps out the orientation of crystal grains very precisely. EBSD has become popular in invertebrate palaeontology but its application in vertebrate palaeontology remains limited. However, the study of fossil eggshells is a field where EBSD has wide potential applicability and provides a quantitative approach to fossil eggshell research as well as new qualitative data. Here we analyse fossil and extant maniraptoran dinosaur eggshells using EBSD analysis emphasizing four different aspects. The mapping imaging clarifies previously ambiguous characters such as squamatic ultrastructure and allows a more objective evaluation of avian and non‐avian maniraptoran eggshell. In particular, our results imply that the presence of an external zone in the manirpatoran eggshell is not diagnostic of avian eggshell. EBSD analysis can be also used for differentiating true pore canals from cracks in the eggshell radial section, thereby determining the biological genuineness or otherwise of a pore‐like structure. Finally, the misorientation angle distribution of the material shows a clear dichotomy that may reflect reproductive brooding strategy, although further studies on contact incubation of palaeognaths and neognaths are needed to confirm this.     

Hoof-Like Unguals,Skin, and Foot Movements Deduced from Deltapodus Casts of the Galve Basin (Upper Jurassic-Lower Cretaceous,Teruel, Spain)

Galve (Teruel, Spain) is a town in the interior of a synclinal fold with Upper Jurassic marine limestones along its flanks, and, in its core, Upper Jurassic–Lower Cretaceous continental and shoreline sediments crop out. The core sediments cover an area about 8 km², and contain a concentration of sites with footprints, bones, and eggshells of dinosaurs. The footprints are both shafts and natural casts. Some casts are attributed to stegosaurs (Deltapodus). The Deltapodus casts are characterized by features that allow us to make direct observations on the skin formed by polygonal scales, and ellipsoidal “hooves,” as well as deductions on the movement of the limbs during walking. According to the opinion of some authors, dinosaur footprints are indicators of the motion of their limbs and sometimes of the whole body. So far, results have been deduced from theropod, ornithopod, and sauropod footprints. This article shows the results obtained from analis of the aforementioned Deltapodus casts, i.e., forelimb movement similar to that of the forelimbs of sauropods, and the rigid structure of the autopodial part of the hind limb.     

The family Phryganeidae (Insecta,Trichoptera) from the Mesozoic and Cenozoic of Asia (with a brief worldwide overview of the fossil caddisfly fauna)

The following new taxa of the family Phryganeidae are described: Proagrypnia sinitsae gen. et sp. nov. from the Upper Jurassic or Lower Cretaceous of Russia (Chernovskie Kopi locality) and Baissophryganoides popovi sp. nov. from the Lower Cretaceous of Mongolia (Anda-Khuduk locality). A larval case of Folindusia (Acrindusia) phryganoides sp. nov. from the Paleocene of Russia (Arkhara–Boguchan brown coal field) is described. A worldwide overview of the records of fossil adult insects and larval cases of Phryganeidae is presented.     

白垩纪双壳类Isodomella在藏北的发现及其意义

产于西藏北部改则县川巴和麻米附近的1个双壳类动物群,包括 Isodomella choinbaensis sp. nov. 和 Caestocorbula (Parmicorbula) perlonga sp. nov. 等9种,时代为早白垩世 Valanginian 至早 Aptian 期.从 Isodomella 揭示出的铰齿看,它应归入 Corbiculidae 科;所共生的种类反映 Isodomella 为与正常海有密切联系的半咸水生活的种类.Isodomella 被认为是日本特有的属,在日本以外尚属首次发现.这个属的动物从日本到藏北的迁移路线,很可能是经我国台湾、南海,绕中南半岛折向西北,又经我国滇西、藏东到达藏北的.这个动物群与东特提斯北部边缘的动物群联系密切,说明川巴和麻米所在的拉萨地块当时已是欧亚大陆的一部分.     

Nest taphonomy of common terns (Sterna hirundo) on Poplar Island,Chesapeake Bay,Maryland

Mesozoic non-avian theropod dinosaurs displayed a diverse range of egg types and clutch forms, suggesting a variety of nesting behaviours, some of which may be shared with birds. More accurate inferences of these behaviours require taphonomic studies of modern nesting sites. Here, we document common tern (Sterna hirundo) nesting sites on Poplar Island in Chesapeake Bay, Maryland. Nests were surveyed on multiple occasions, documenting nest composition, density and distribution, as well as eggshell concentration and orientation. Three colonies yielded 79 tern nests with 193 eggs. Twelve nests hatched, 7 were predated, 30 failed and the fates of 30 remain unknown. Abundant eggshells occurred at the centre of the nests. Concave-up eggshell in or on the nest surface characterised hatched and predated nests, whereas eggshell forced into the subsurface seemed to favour concave-down orientation and may reflect post-hatching extended use of the nest. Eggs buried deep within the nest materials and/or substrate suggest adult abandonment. Quantitative data on eggshell orientation and observations regarding controls on nest distribution, egg predation and incorporation of intact eggs into the substrate at modern nesting sites provide physical guidelines for the improved interpretation of fossil nesting localities.     

Morphometric analysis of dinosaur eggshells: constraints of size on shape

Studies on fossil eggs and eggshells have been traditionally focused on analysis of microstructure and systematic classification, but few attempts have been made to examine the variability of eggshell characters. This work examines the variability of one diagnostic character, shape of the calcitic units, of some dinosaur eggshells by using outline shape analysis (elliptic Fourier analysis). The studied material belongs to five different oospecies of the parafamily Megaloolithidae Zhao, 1979. To test shape variation within and between oospecies, an analysis on 129 outlines was carried out and their variability was summarised by principal component analysis. We then tested if the shape of megaloolithid units changes according to size (i.e. allometry). A regression analysis between eggshell unit shape and two different size measurements (centroid size and eggshell thickness) showed that the thinnest eggshells (thickness ≤ 1 mm) have different shapes than thicker ones ( ≥ 2.5 mm). Our results stress that the variability in the shape of eggshell units in Megaloolithus oospecies is correlated to size (i.e. eggshell thickness), a phenomenon that should be then considered carefully when determining oospecies, since a single oospecies can show both a wide range of eggshell thickness and combined with different eggshell unit shapes.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

A re-evaluation of aff. Megaloolithidae eggshell fragments from the uppermost Cretaceous of the Pyrenees and implications for crocodylomorph eggshell structure

The Upper Cretaceous outcrops of the Pyrenees yield one of the most extensive and continuous records of paleoological remains anywhere in the world. Most of eggs and eggshells have been referred to the oofamily Megaloolithidae. In this study, we present a revision of eggshell fragments from the Blasi 2 locality, lattermost Maastrichtian in age, previously assigned to aff. Megaloolithidae. The presence of a blocky extinction pattern and basal knobs supports a crocodilian affinity of these materials. We classify them as Krokolithidae indet. Three structural layers can be recognised in the Blasi 2 eggshells, a feature that is shared with other recent eggshells (e.g. Crocodylus porosus and Crocodylus niloticus) and fossil crocodylomorph eggshells (Krokolitheswilsoni), which were previously described as single layered. The new proposed affinity of the Blasi 2 eggshells reduces the Megaloolithidae oodiversity of the last few million years of the Cretaceous in the Pyrenees to only two valid ootaxa, Megaloolithusmamillare and Megaloolithusbaghensis. The lack of more complete material precludes the erection of new ootaxa based on the Blasi 2 material.     

The earliest fossil schizopterid bug (Insecta: Heteroptera) in the Lower Cretaceous amber of Lebanon

The schizopterid bug Libanohypselosoma popovi n. gen., n. sp. belonging to the subfamily Hypselosomatinae is described from the Lower Cretaceous amber of Lebanon. This fossil is the earliest record of the Schizopteridae. The species is distinguished from its related taxa, a discussion is given.     

A new peculiar dinosaur egg,Sankofa pyrenaica oogen. nov. oosp. nov. from the Upper Cretaceous coastal deposits of the Aren Formation,south‐central Pyrenees,Lleida, Catalonia,Spain

Abstract: A new type of small, ovoid dinosaur egg, Sankofa pyrenaica oogen. nov. oosp. nov., with a prismatic type eggshell is described from upper Cretaceous (upper Campanian–Maastrichtian) deposits of the Montsec area, South Pyrenean Central Unit, Lleida, Catalonia, Spain. This egg type was sub‐vertically laid in only two rich monospecific sites of a single stratigraphic layer from coastal deposits of the Aren Formation, interpreted as an emerged beach ridge of a barrier island – lagoon depositional system. The size and shape of these eggs with their asymmetric poles are roughly similar to modern hen eggs, which is unusual in the Cretaceous fossil egg record. Its phylogenetic position clusters with bird and Troodontid eggs. A morphospace analysis of egg shapes shows the similarity of the new egg to a Campanian fossil bird egg from Argentina, both being intermediate between modern‐bird eggs and extinct nonavian theropod eggs. However, the eggshell microstructure of Sankofa pyrenaica differs from that of bird eggs in its incipient squamatic texture. It has a peculiar pattern of interlocking small crystals in the middle of the palisade layer, instead of the thick squamatic structure commonly present in modern avian eggshells. This new egg type is attributed to a small theropod, probably with a single oviduct like birds and whose mosaic distribution of features is a combination between that of birds and nonavian theropods. This enhances the arguments supporting the close phylogenetic relationships between both groups.     

An unusual lizard (Reptilia: Squamata) from the Early Cretaceous of Las Hoyas, Spain

The Early Cretaceous (Barremian) lepidosaurian assemblage of Las Hoyas (Cuenca Province, Castilla-La Mancha), Spain, resembles that of roughly contemporaneous localities at Montsec (Catalonia), Una (Castilla-La Mancha) and Galve (Aragon) in being dominated by the scincomorph lizard Meyasaums ( Ilerdaesaurus ). It differs in the presence of two previously unknown taxa, one of which, described here, shows a strikingly unusual morphology in its ribs and distal limb skeleton suggestive of climbing specialization. This form is sufficiently distinct from other known taxa to warrant the erection of a new genus and species, Scandensia ciervensis gen. et sp. nov. Cladistic analysis of Scandensia suggests that it forms a sister taxon to living squamates (Iguania + Scleroglossa and all descendants of their most recent common ancestor).     

Anatomical differentiation of isolated scales of amiiform fishes (Amiiformes,Actinopterygii) from the Early Cretaceous of Las Hoyas (Cuenca,Spain)

The scales of amiiform fishes are more different from each other than previously stated. The anatomy of the scales of the three amiiform taxa from the Early Cretaceous (Barremian) of Las Hoyas (Cuenca, Spain) is described in detail. The differences between them has allowed the segregation of isolated scales form the fossil record of this site into the three taxa, providing relatively large population samples that can be studied from a palaeobiological and palaeoecological point of view.     

Palaeobotanical remains associated with dinosaur fossils from the Camarillas Formation (Barremian) of Galve (Teruel,Spain)

A well-preserved macroflora and rich palynological assemblages corresponding to the Camarillas Formation (early-middle Barremian) in the San Cristóbal and Galve Mine sites from the Galve sub-basin in northeastern Spain are presented here. These remains represent the first fossil plant evidence from these deposits. Within plant macroremains, the cheirolepidiacean Pseudofrenelopsis aff. varians (Fontaine) Watson has been found. The palynological assemblage yielded well-preserved spores and pollen grains, mainly dominated by the genus Classopollis. Spores are also abundantly represented by schizaeacean spores (Cicatricosisporites and Plicatella). This spore assemblage supports an early-middle Barremian age for these localities. It is noteworthy that small basal angiosperm pollen grains of the genera Crassipollis and Retimonocolpites, together with other indeterminate grains, have been reported here. This flora constitutes the primary food producer for dinosaur at that time, and thus the information of the flora is important for the understanding of the ecological background for the dinosaur evolution during the Early Cretaceous.     

浙江天台晚白垩世伤齿龙(troodontids)蛋化石的新发现

浙江天台上白垩统赤城山组中发掘出属伤齿龙类的棱柱形蛋化石。研究认为,棱柱形蛋从伤齿龙类恐龙中产出就具有较高刚度的蛋壳,可直接插入松软的泥沙里,明显区别于其他大多数恐龙及现代爬行类所产的韧性蛋壳蛋,韧性蛋壳的恐龙蛋往往因重力作用产生变形而呈扁球形和扁椭球形。伤齿龙类的棱柱形蛋的刚性蛋壳很接近现代鸟蛋的蛋壳,暗示伤齿龙类恐龙生殖系统在蛋壳形成方面与现代鸟类有一定相似性。同时,伤齿龙类的棱柱形蛋或多或少地以向外倾斜的形式排布,反映出伤齿龙类恐龙产卵时可能面向巢穴外,是一种对生存环境的适应行为。     

Implications de l’analyse microstructurale et géochimique des œufs de dinosaures de la cairanne (Bassin d’Aix en Provence,France, Rognacien inférieur)

Microstructural and geochemical analyses of the dinosaur eggshells at La Cairanne (Aix en Provence Basin, Lower Rognacien) show the presence of diagenesis. Diagenetic processes have altered the thickness of the shells and some structural data. The chemical composition has been also modified. Thus it is difficult to use these parameters to reconstruct the paleoenvironment. It is possible, indeed probable, that the problems of the previous authors about the definitions of eggshell morphotypes in the Aix Basin are due to diagenetic variations.