Parataxonomic review of the Upper Cretaceous dinosaur eggshells belonging to the oofamily Megaloolithidae from India and Argentina

The eggshell oospecies from India and Argentina are compared and reviewed in detail. These eggshells resemble each other in having a nodular outer surface ornamentation and clearly arched growth lines of the shell units. Microstructurally, the eggshell oospecies belonging to the oofamily Megaloolithidae shows fan-like shell units, which are sharply separated from each other throughout the thickness of the eggshell and can be traced up to the surface of the eggshell. Comparisons between four oospecies from India and Argentina reveal three groupings, which show similarities between megaloolithids of both countries: (1) Megaloolithus jabalpurensis, M. matleyi and M. patagonicus; (2) M. cylindricus, M. rahioliensis and Tipo 1d; and (3) M.megadermus and Tipo 1e. The other two types of eggshell oospecies from India and Argentina show partially fused external nodes and shell units. As a result, growth lines enter into the adjacent shell units with a marked concavity. A new oogenus Fusioolithus have been erected due to fusion between shell units and tubospherulitic morphotype, which include two new oospecies F. baghensis and F. berthei. Till date, morphostructurally, a total of 15 eggshell oospecies belonging to different oofamilies have been recorded from India and seven oospecies from Argentina.     

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.     

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.     

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.     

A new oospecies of Similifaveoloolithidae from the Xiuning Basin,Late Cretaceous of Anhui,China

Well-preserved dinosaur eggs from the Cretaceous Huizhou Formation in the Xiuning Basin, Anhui Province, China, are analysed in this paper. We describe a new oospecies, Similifaveoloolithus qiyunshanensis, based on several distinct characters of external morphology, size, eggshell thickness, and internal microstructure. Radial sections of this new oospecies show branched eggshell units with a fused layer near the outer surface, while numerous irregular pores and cones constitute a honeycomb pattern in tangential sections. The discovery of S. qiyunshanensis expands the distribution of Similifaveoloolithidae dinosaur eggs in China and provides new fossils for researching dinosaur eggshell formation mechanisms which are different from those of the currently known oofamilies. The dinosaur-egg-bearing strata in the Huizhou Formation have been dated to the early Late Cretaceous (Cenomanian-Turonian) on the basis of a similar dinosaur egg assemblage in the Tiantai Basin in Zhejiang Province. The eggs described in this paper are thought to have been laid in a buried nest while enrichment of trace elements in eggshells may have been caused by their ingestion into the body of the dinosaur producer. We suggest that the paleoclimate of this habitat was semi-arid to arid and that this environment was favourable for the preservation of eggs.http://zoobank.org/LSIDurn:lsid:zoobank.org:pub:9D63A51F-5A4B-4249-901D-42A33E9FA933     

Unusually thick dinosaur eggshell fragments from the Spanish Late Cretaceous

Fieldwork carried out recently in the southeastern branch of the Iberian Range (Valencia Province, Spain) has led to the collection of a large volume of dinosaur eggshell fragments of unusual thickness. These specimens, up to 4.9 mm thick, were recovered from palustrine grey marls of the upper Campanian-lower Maastrichtian Sierra Perenchiza Formation, which comprises a wetland paleoenvironment deposit. These eggshell fragments have a characteristic compactituberculate ornamentation, dinosauroid-spherulitic organisation, and exhibit a complex canaliculate respiratory system. The external tuberculate surface of the shell as well as the internal microstructure enable referral to Megaloolithus aff. siruguei, the most common megaloolithid oospecies known from the Iberian Peninsula and southern France. The biostratigraphic range of M. siruguei matches the temporal distribution of titanosaurid dinosaurs across the Iberian Range, tentatively considered to be potential producers.     

A new egg with avian egg shape from the Upper Cretaceous of Zhejiang Province,China

A new egg is described from the Upper Cretaceous lacustrine deposit of the Chichengshan Formation in the Tiantai Basin, Zhejiang Province, southeast China. The new specimen shares eggshell micro-features with members of the oofamily Stalicoolithidae, Paraspheroolithus of the oofamily Spheroolithidae and Mosaicoolithus (oofamily indet.), with barrel-shaped cones, prolatocanaliculate pore system, horizontal accretion lines and light stripes throughout the eggshell. However, the new egg differs from the aforementioned ootaxa by its small size and asymmetrical shape, revealing new morphological variation among eggs with microstructure similar to that of Paraspheroolithus, Mosaicoolithus and Stalicoolithidae. We refer the new egg to a new oogenus and oospecies, thus increasing the diversity of the Tiantai Basin oofauna. The Tiantai Basin has yielded a variety of dinosaur eggs and one turtle clutch. Comparatively, the new egg is surprisingly small and ovoid, a morphology usually associated with avian eggs, although the absence of a squamatic layer excludes the egg from being referred to this group.     

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.     

Great spotted cuckoo eggshell microstructure characteristics can make eggs stronger

Obligate avian brood parasites lay stronger eggs than their hosts or non‐parasitic relatives because they are rounder and have a thicker eggshell. Additionally, some other characteristics of the brood parasitic eggshells related to their microstructure such as size and orientation of calcite crystal units could also contribute to generating even stronger shells. An eggshell microstructure formed by small randomly oriented calcite crystal units can increase the robustness of the eggshells of birds. Here, the eggshell microstructure of avian brood parasites as well as their hosts have been characterized in detail, using X‐ray diffraction analyses to estimate the size and degree of orientation of calcite crystal units making the eggshell. Specifically, the brood parasitic great spotted cuckoo Clamator glandarius and two hosts (jackdaws Corvus monedula and magpie Pica pica) and one non‐host species (the pigeon, Columba livia domestica) were considered. Calcite crystal of the eggshell of the brood parasitic species was smaller and more randomly oriented than those of the eggshells of non‐parasitic species, which suggest that eggshell microstructure would contribute to explain why parasitic eggs are more resistant to breakage than those of their hosts.     

浙江天台盆地晚白垩世网形蛋类新类型及网形蛋类的分类订正(英文)

记述了产自浙江省天台盆地的网形蛋类新材料,建立一新蛋属——拟网形蛋属(Paradictyoo lithusoogen.nov.),两个新蛋种:庒前拟网形蛋(Paradictyoolithus zhuangqianensis oogen.etoosp.nov.)和下西山拟网形蛋(P.xiaxishanensis oogen.et oosp.nov.)。在此基础上,重新系统描述其他已知4种网形蛋属成员的结构特征,并对网形蛋类的分类进行了订正:网形蛋属(Dictyoolithus)仅保留红坡网形蛋(D.hongpoensis Zhao,1994)一个蛋种;建立一新蛋属:原网形蛋属(Protodictyoolithus oogen.nov.),将发现于河南西峡盆地的内乡网形蛋(D.neixiangensis Zhao,1994)和发现于山东莱阳盆地的蒋氏网形蛋(D.jiangi Liu&Zhao,2004)分别修订为:内乡原网形蛋(修订种)(Protodictyoolithus neixiangensis(Zhao,1994)comb.nov.)和蒋氏原网形蛋(修订种)(P.jiangi(Liu&Zhao,2004)comb.nov.);此外,发现于吉林公主岭早白垩世的公主岭网形蛋(D.gongzhulingensis Wang et al.,2006)并不属于网形蛋类,而应归入似蜂窝蛋类(Similifaveoolithus),将其修订为公主岭似蜂窝蛋(修订种)(Similifaveoolithus gongzhulingensis(Wangetal.2006)comb.nov.)。     

The oldest known avian eggshell,Plagioolithus fukuiensis,from the Lower Cretaceous (upper Barremian) Kitadani Formation,Fukui, Japan

Fossil record of Early Cretaceous birds may be geographically biased, and eggs and eggshells predating the Late Cretaceous were unknown. Here, we report the oldest known bird eggshell (FPDM-V-0009175) collected from the upper Barremian Kitadani Formation in Katsuyama City, Fukui, Japan. The Kitadani Formation likely represents fluvial environments. Thin-section and scanning electron microscope analyses revealed diagnostic characters of FPDM-V-0009175, including thin (0.44 mm) shell, smooth external surface, non-branching and narrow pore canals with relatively constant width, three structural layers, oblique crystal orientation from vertical in the external layer, and mammillary to continuous to external layer thickness ratio of 1:1:0.44. These characters allow assignment of FPDM-V-0009175 to a new oogenus and oospecies, Plagioolithus fukuiensis, and suggest it belonging to a bird. The three-layered eggshell structure is seen in extant and extinct birds, Plagioolithus fukuiensis, and non-avian theropods. Therefore, such structure may be plesiomorphic among theropods, appearing in the late Barremian or earlier. As the first bird body fossil from the Mesozoic of Japan, Plagioohlithus fukuiensis suggests extensive distribution and abundance of birds in the Barremian East Asia. Plagioolithus fukuiensis indicates that the late-Barremian birds inhabited and reproduced in the fluvial environments within a basin located along the eastern margin of the Asian continent.http://www.zoobank.org/urn:lsid:zoobank.org:pub:90A8CCE8-57F7-473C-855C-B2CCE2BBD1A2     

Upper Cretaceous paleogeography of the Central Southern Pyrenean Basins (Catalonia, Spain) from microfacies analysis and charophyte biostratigraphy

The most extensive latest Cretaceous deposits of the Pyrenees are non-marine facies traditionally called “Garumnian”, which represent the first continentalization of the Pyrenean Foreland Basins. The age and paleogeography of the basal Garumnian facies in the different parts of the Pyrenean Basin has been a matter of debate. Charophyte biostratigraphy and microfacies analysis suggest that the continentalization was diachronous in the Central Southern Pyrenean basins, i.e., between the Àger and Tremp basins, and the emersion progressed from south to north. In the Àger Basin, to the south, the first Upper Cretaceous non-marine rocks are represented by the La Maçana Formation. This unit is mainly formed by freshwater lacustrine limestones organized into hundreds of shallowing-upwards sequences ranging from deeper lacustrine facies, through marginal, well-illuminated environments dominated by characean meadows, and finishing with lakeshores dominated by clavatoracean meadows. The charophyte assemblage from the La Maçana Fm is mainly formed by Peckichara cancellata, P. sertulata, Microchara cristata, M. parazensis, Platychara caudata, and Clavator brachycerus, which belong to the Peckichara cancellata charophyte biozone (Late Campanian). In contrast, in the Tremp Basin, located to the north, the beginning of the non-marine sedimentation is organized into a few cycles of the well-known La Posa Fm. These cycles begin with brackish deposits formed by thick marls with euryhaline molluscs at the base, followed by lignite and brackish to freshwater limestones. The brackish facies are dominated by porocharaceans. The charophyte assemblage of the La Posa Formation is formed by Feistiella malladae, Peckichara sertulata, Microchara cristata, and Clavator brachycerus, which belong to the Septorella ultima charophyte biozone (Early Maastrichtian).     

Two new theropod egg sites from the Late Jurassic Lourinhã Formation,Portugal

Two new Late Jurassic (uppermost Late Kimmeridgian) dinosaur eggshell sites are described, Casal da Rola and Porto das Barcas, both near Lourinhã, central-west Portugal. Casal da Rola yields eggshells with an obliquiprismatic morphotype comparable to those from a nest with the associated fossil embryos from Paimogo, tentatively assigned to the theropod Lourinhanosaurus antunesi. The Porto das Barcas eggshells have a dendrospherulitic morphotype with a prolatocanaliculate pore system. This morphotype was also recognised in eggshells from a clutch with associated Torvosaurus embryos at the Porto das Barcas locality. A preliminary cladistic analysis of eggshell morphology suggests theropod affinities for the Casal da Rola eggs, but is unable to resolve the phylogenetic position of the Porto das Barcas eggs. The eggshells at both sites are preserved in distal flood plain mudstones and siltstones. Carbonate concretions within the deposits indicate paleosol development.     

Egg colouration predicts brood size,telomere length and body condition of spotless starling fledglings

Understanding the impressive interspecific variation in avian eggshell colouration has attracted the attention of evolutionary ecologists for more than a century. Several functional explanations predict positive covariation between eggshell pigmentation and phenotypic quality of nestlings. We test this prediction in spotless starlings Sturnus unicolor by using biometric measurements and telomere length of hatchlings and fledglings as proxies of phenotypic quality. Female spotless starlings lay immaculate blue‐green eggs, a sexually selected signal directed to males. Pigmentation predicts positive associations with concentration of antioxidants and testosterone in the yolk and with paternal provisioning effort during nestling growth. Eggshell pigmentation (blue‐green chroma) is not associated with telomere length of hatchlings, which suggests weak maternal effects on this trait. However, we find negative associations of eggshell colouration with both body condition and telomere length of fledglings. Moreover, we find positive associations between eggshell colouration and clutch size, which suggests that sibling competition is higher in nests with more coloured eggshells. Previous works demonstrated that level of sibling competition is positively related to telomere erosion and, thus, the detected negative associations between eggshell colouration, body condition and telomere length of fledglings would reflect higher level of competition in nests with more coloured eggshells. We therefore speculate with the possibility that females that lay larger clutches also lay more coloured eggshells that elicit increased paternal provisioning effort and, thus, raise larger broods at the expense of telomere erosion of their offspring.     

Fine structure of the subitaneous eggshell of the sessile rotifer Stephanoceros millsii (Monogononta) with observations on vesicle trafficking in the integument during ontogeny

Rotifers that engage in cyclical parthenogenesis produce two types of eggs: subitaneous eggs that hatch as clonal females and meiotic eggs that hatch as haploid males, or if fertilized, as females after a period of diapause (resting eggs). The ultrastructure of resting eggshells is known for some motile species, but there are limited data on subitaneous eggshells, and no data on any eggshells of sessile rotifers. Here, we investigated the ultrastructure of the subitaneous eggshell of the sessile rotifer Stephanoceros millsii and its potential origins of secretion, the maternal vitellarium and embryonic integument. We also explored secretory activity in the larval and adult integuments to determine whether activity changes during ontogeny. The eggshell consists of a single layer with two sublayers: an external granular sublayer apparently derived from the maternal vitellarium, and an internal flocculent sublayer secreted by the embryonic integument that may form a hatching membrane or glycocalyx. Secretory activity remains high in both the larva and adult and appears to be the source of the thickening glycocalyx. Altogether, the subitaneous eggshell of S. millsii is the thinnest among monogonont rotifers. Thin eggshells may have evolved in response to the added protection provided by the mother’s extracorporeal tube.     

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.     

A distinctive new species of vireo (Passeriformes: Vireonidae) from the Western Andes of Colombia

On the basis of field observations and two specimens from two localities in the Chocó biogeographic region of western Colombia, we here describe Vireo masteri, sp. nov. This species appears to be most closely related to Vireo carmioli of Costa Rica and west Panama but is much smaller and has a distinctive long superciliary stripe, a feature otherwise unknown in the subgenus Vireo, of which it is the southernmost known taxon. Vireo masteri appears to be restricted to a narrow belt of premontane pluvial forests from 1200 to 1600 m on the Pacific slope of the Western Andes. We present notes on the behaviour, ecology and conservation of this new species. Con base en observaciones en el campo y dos especímenes de dos localidades en la región biogeográfica del Chocó colombiano, describimos una nueva y distintiva especie de vireo o chiví, Vireo masteri, sp. nov. Esta especie parece mas afin a Vireo carmiloi de Costa Rica y Panamá que a cualquier otra especie, pero es de menor tamaño y tiene una lista superciliar larga y llamativa, caracteristica única en la subgénero Vireo, del cual es la forma más sureña. Vireo masteri aparentemente se restringe a una faja altitudinal angosta (1200–1600 msnm) de bosque pluvial premontano en la vertiente del Pacífico de la Cordillera Occental de los Andes colombianos. Presentamos anotaciones sobre la ecología, el comportamiento, y la conservación de esta neueva especie.     

A rare mineral,vaterite, acts as a shock absorber in the eggshell of a communally nesting bird

Birds’ eggshells are primarily composed of calcite, an abundant polymorph of calcium carbonate (CaCO₃). However, the eggshells of some species are coated with spherules of vaterite, a rare and thermodynamically unstable polymorph of CaCO₃, the function of which remains unknown. We experimentally tested the mechanical and physiological effects of the vaterite coating on eggshells of the Greater Ani Crotophaga major, a tropical cuckoo. Vaterite removal did not affect vapour conductance rates across the eggshell, indicating that the vaterite coating does not influence gas exchange during embryonic development. However, nanoindentation revealed that the hardness and elasticity of vaterite is similar to that of calcite, and white light interferometry showed that the vaterite layer increased the total thickness of the shell cuticle by up to 10%. Furthermore, calculations of contact mechanics found that when two eggs come into contact, the depth of the surface deformation caused by the contact is far less than the thickness of the vaterite coating. These results suggest that the layer of vaterite spherules may act as a shock absorber for the underlying calcite shell, protecting it from mechanical damage caused by collision with other eggs in the nest and reducing the risk of eggshell fracture during incubation.