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.     

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.     

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     

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.)。     

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.     

Eggshell Types and Their Evolutionary Correlation with Life-History Strategies in Squamates

The eggshell is an important physiological structure for the embryo. It enables gas exchange, physical protection and is a calcium reserve. Most squamates (lizards, snakes, worm lizards) lay parchment-shelled eggs, whereas only some gekkotan species, a subgroup of lizards, have strongly calcified eggshells. In viviparous (live-bearing) squamates the eggshell is reduced or completely missing (hereafter “shell-less”). Recent studies showed that life-history strategies of gekkotan species differ between species with parchment- and rigid-shelled eggshells. Here we test if the three different eggshell types found in the squamates are also associated with different life-history strategies. We first investigated the influence of the phylogeny on the trait “eggshell type” and on six life-history traits of 32 squamate species. Phylogenetic principal component analysis (pPCA) was then conducted to identify an association between life-history strategies and eggshell types. Finally, we also considered adult weight in the pPCA to examine its potential effect on this association. Eggshell types in squamates show a strong phylogenetic signal at a low taxonomical level. Four out of the six life-history traits showed also a phylogenetic signal (birth size, clutch size, clutches per year and age at female maturity), while two had none (incubation time, maximum longevity). The pPCA suggested an association of life-history strategies and eggshell types, which disappeared when adult weight was included in the analysis. We conclude that the variability seen in eggshell types of squamates is weakly influenced by phylogeny. Eggshell types correlate with different life-history strategies, and mainly reflect differences in adult weights of species.     

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.     

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.     

First report of fossil turtle eggshell west of the Colorado Plateau

The first fossil turtle eggshell from North America west of the Colorado Plateau (UCMP 218504) is described, adding to the growing global record of chelonian eggshell. Ten blocks of mudstone were collected from the Upper Miocene (Clarendonian) Orinda Formation, Contra Costa County, California, during the excavation of the Caldecott Tunnel Fourth Bore. They contain rigid eggshell averaging 500 μm in thickness. Closely packed shell units with a height/width ratio of 2:1 are composed of radiating, acicular crystals originating from a basal core, a set of characters that support the identification of the eggshell as chelonian, and further as testudinoid. Egg diameter is estimated at between 34.2 and 42.4 mm based on shell curvature. Adult carapace length is estimated to be 70.7 cm, a size similar to fossils of Hesperotestudo known from California and present in the Orinda Formation. UCMP 218504 greatly extends the geographic record of turtle fossil eggshell and adds to the limited number of mid-Neogene eggs. It also provides evidence for a warm climate in Miocene California and demonstrates the south-eastern contraction of conditions appropriate for tortoise reproduction during the last 10 million years.     

Magellanic Penguin eggshell pores: does number matter?

Most penguin species lay two eggs 3–4 days apart that hatch 1–2 days apart. Hypotheses to explain the shorter incubation period for the second egg include differences in porosity of the eggshells. Eggs with more or larger pores and/or thinner shells have higher gas exchange rates and faster embryonic development. We used eggshells from hatchling Magellanic Penguins Spheniscus magellanicus to test whether eggshell thickness and pore density affect incubation period. We expected: (i) second eggs to have thinner shells and/or more pores than first eggs; (ii) eggs to have thinner shells and/or more pores if they were laid by older females, later in the season, or in burrow nests (more humid than open nests under bushes). We found no support for these hypotheses. Egg pores and shell thickness were not related to incubation period. Eggs from the same clutch were similar in eggshell thickness and pore density. Female age, laying date and nest type were poor predictors of eggshell thickness and pore density. Egg pores, we conclude, have little explanatory power for differences in incubation time of eggs in Magellanic Penguins, and suggest that synchronous onset of incubation matters.     

Structure of shells from eggs of kinosternid turtles

Shells from eggs of five species of kinosternid turtle (Sternotherus minor, Kinosternon flavescens, K. baurii, K. Hirtipes, and K. alamosae) were examined with light and scanning electron microscopy. Except for possible differences among species in thickness of eggshells, structure of shells from all eggs was similiar. In general, kinosternid turtles lay eggs having a rigid calcareous layer composed of calcium carbonate in the form of aragonite. The calcareous layer is organized into individual shell units with needlelike crystallites radiating from a common center. Most of the thickness of the eggshell is attributable to the calcareous layer, with the fibrous shell membrane comprising only a small fraction of shell thickness. Pores are found in the calcareous layer, but they are not numereous. The outer surface of the eggshells is sculptured and may have a thick, organic layer in places. The outer surface of the shell membrane of decalcified eggshells is studded with spherical cores which presumably nucleate growth of shell units during shell formation. The shell membrane detaches from eggs incubated to hatching, carrying with it remnants of the calcareous layer. Such changes in shell structure presumably reflect withdrawal of calcium from the eggshell by developing embryos.     

Effects of nanosized zinc oxide and γ-polyglutamic acid on eggshell quality and serum parameters of aged laying hens

In order to increase zinc (Zn) absorption and improve eggshell quality, diets for aged laying hens were supplemented with different Zn sources and the effects on egg production, eggshell quality and serum parameters were recorded. Seventy-five 64-week old brown layers were individually caged and randomly allotted to five treatment groups: an un-supplemented Control and four groups, where the following Zn sources were added to the diets, ZnO (group ZnO), Zn-methionine (group Zn-Met), nanosized ZnO (group nanoZnO) and γ-PGA-nano ZnO (group γ-PGA-nanoZnO). The Zn level was maintained at 80 mg/kg in the supplemented diet, while the Control group received Zn at 40 mg/kg diet. The results indicated that the average daily feed intake was highest in group nanoZnO (p < 0.05), whereas further performance parameters were not influenced by treatments. Eggshell thickness was increased in group γ-PGA-nanoZnO (p < 0.05). Zn content in eggshells and Zn concentration in serum were increased in groups Zn-Met, nanoZnO and γ-PGA-nanoZnO (p < 0.05). Serum ghrelin concentration was significantly elevated in all Zn-supplemented groups, but further increased in groups nanoZnO and γ-PGA-nanoZnO (p < 0.05). Carbonic anhydrase activity was highest in group γ-PGA-nanoZnO and lowest in group Zn-Met (p < 0.05). Compared to the Control and group ZnO, supplementation of γ-PGA-nanoZnO and nanoZnO increased serum IgG levels (p < 0.001). In conclusion, dietary supplementation of nanoZnO and γ-PGA-nanoZnO increased Zn content in eggshells, serum Zn concentration, ghrelin and IgG levels of aged layers when compared to regular ZnO. Compared to Zn-Met, the serum carbonic anhydrase activity and ghrelin levels were also increased. Thus, nanosized ZnO alone or mixed with γ-PGA has positive effects on the Zn status of aged layers.     

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.     

江西萍乡地区晚白垩世副蜂窝蛋类一新蛋种

依据采自江西省萍乡地区上白垩统周田组的恐龙蛋化石,记述了蜂窝蛋科(Faveoloolithidae)副蜂窝蛋属(Parafaveoloolithus)一新蛋种。材料包括一枚较完整的蛋和若干破碎蛋壳。根据蛋壳弦切面呈蜂窝状结构,径切面蛋壳中、上部蛋壳局部呈现由6~10个以上壳单元成群聚集一起等特征,将其订为一新蛋种——萍乡副蜂窝蛋(Parafaveoloolithus pingxiangensis oosp.nov.)。     

卵壳的超微结构特征

本文运用扫描电镜研究了美洲鸵鸟,鹂鹋,非洲鸵鸟,普通家鸡,环颈雉,绿头鸭,王企鹅等七种现生鸟蛋壳和更新世安氏鸵鸟蛋壳以及六种白垩纪恐龙蛋壳（长形长形蛋Ｅｌｏｎｇａｔｏｏｌｉｔｈｕｓｅｌｏｎｇａｔｕｓ,安氏长形蛋Ｅｌｏｎｇａｔｏｏｌｉｔｈｕｓａｎｄｒｅｗｓｉ,瑶屯巨形蛋Ｍａｃｒｏｏｌｉｔｈｕｓｙａｏｔｕｎｅｎｓｉｓ粗皮巨形蛋Ｍａｃｒｏｏｌｉｔｈｕｓｒｕｇｕｓｔｕｓ,将军顶圆形蛋Ｓｐｈｅｒｏｏｌｉｔｈ     

Is greater eggshell density an alternative mechanism by which parasitic cuckoos increase the strength of their eggs?

Brood parasites lay unusually strong eggs, presumably to prevent puncture ejection of the eggs by hosts. However, it has been suggested that eggs of some parasitic cuckoos have normal strength. This suggestion was based on the eggshell thickness and shape of these eggs. Here, we propose that there may be other structural adaptations (such as increased eggshell density) that make the cuckoo eggs unusually strong. In this study, we compared the eggshell density of parasitic cuckoos to the eggshell density of two control groups, non-parasitic cuckoos and a sample of non-passerine species. The comparisons to both control groups demonstrated that the parasitic cuckoos have eggshells of significantly higher density than would be expected for their size. Our results support the hypothesis that the higher eggshell density is an alternative mechanism by which some cuckoos increase the strength of their eggs.     

Genetic variation in eggshell crystal size and orientation is large and these traits are correlated with shell thickness and are associated with eggshell matrix protein markers

The size and orientation of calcium carbonate crystals influence the structure and strength of the eggshells of chickens. In this study, estimates of heritability were found to be high (0.6) for crystal size and moderate (0.3) for crystal orientation. There was a strong positive correlation (0.65) for crystal size and orientation with the thickness of the shell and, in particular, with the thickness of the mammillary layer. Correlations with shell breaking strength were positive but with a high standard error. This was contrary to expectations, as in man-made materials smaller crystals would be stronger. We believe the results of this study support the hypothesis that the structural organization of shell, and in particular the mammillary layer, is influenced by crystal size and orientation, especially during the initial phase of calcification. Genetic associations for crystal measurements were observed between haplotype blocks or individual markers for a number of eggshell matrix proteins. Ovalbumin and ovotransferrin (LTF) markers for example were associated with crystal size, while ovocleidin-116 and ovocalyxin-32 (RARRES1) markers were associated with crystal orientation. The location of these proteins in the eggshell is consistent with different phases of the shell-formation process. In conclusion, the variability of crystal size, and to a lesser extent orientation, appears to have a large genetic component, and the formation of calcite crystals are intimately related to the ultrastructure of the eggshell. Moreover, this study also provides evidence that proteins in the shell influence the variability of crystal traits and, in turn, the shell's thickness profile. The crystal measurements and/or the associated genetic markers may therefore prove to be useful in selection programs to improve eggshell quality.     

The effect of developmental stage on eggshell thickness variation in endangered falcons

We compared eggshell thickness of hatched eggs with that of non-developed eggs in endangered falcon taxa to explore the effect of embryo development on eggshell thinning. To our knowledge, this has never been examined before in falcons, despite the fact that eggshell thinning due to pollutants and environmental contamination is often considered the most common cause of egg failure in falcons. Because of the endangered nature of these birds, and the difficulty in gaining access to the nests and their eggs, there is a large gap in our knowledge regarding eggshell thickness variation and the factors affecting it. We used a linear mixed-effects (LME) model to explore the variation in eggshell thickness (n = 335 eggs) in relation to the developmental stage of the eggs, but also in relation to the falcon taxa, the laying sequence and the study zone. Female identity (n = 69) and clutch identity (n = 98) were also included in the LME model. Our results are consistent with the prediction that eggshell thickness decreases during incubation because of the important effect of calcium uptake by the embryo during development. Our results also show that eggs laid later in the sequence had significantly thinner eggshells. In this study, we provide the first quantitative data on eggshell thickness variation of hatched eggs in different falcon taxa that were not subjected to contamination or food limitation (i.e., bred under captive conditions). Because eggshell thickness strongly influences survival and because the species examined in this study are endangered, our data represent a valuable control for future studies on the effects of pollution on eggshells from wild populations and thus are an important contribution to the conservation of falcons.     

Global diversity and adaptations of avian eggshell thickness indices

The amniote eggshell is a fundamental aspect of the embryo life-support system, protecting it from UV light, microbes and mechanical damage, while regulating gas exchange and providing calcium for growth. The thickness of eggshells is highly diverse across modern birds and influences multiple eggshell functions, yet the selective pressures driving eggshell thickness have not been clearly identified. Here, we use a global dataset of avian eggshell thickness indices for 4260 (> 41%) avian species to assess trends in eggshell thickness across the phylogeny, as these indices are strongly correlated with direct measures of eggshell thickness and are non-destructive to the sample. We analysed the dataset within a phylogenetic framework to assess the relative importance of climatic and ecological explanations for variation in eggshell thickness indices. The distribution of avian eggshell thickness indices across species was found to be primarily driven by phylogenetic relatedness, in addition to evolutionary processes that do not match a Brownian model of evolution. Across modern birds, thicker eggshells were more prevalent in species (1) with precocial young, (2) which exhibit a scavenger-based diet, (3) which primarily feed on vertebrates or plants (excluding nectivores, seed and fruit specialists) and (4) which breed in open habitats. Thicker eggshells found in species with precocial young probably enable higher rates of calcium removal for the more advanced embryo development. Excessive light transmission through the shell damages developing embryos, while too little light transmission can impede development. Eggs in shaded habitats experience low light exposure, and thus thinner shells are more prevalent in these environments potentially to increase light transmission through the shell. Overall, variation in eggshell thickness indices appears to be driven largely by phylogeny, with certain life-history traits linked to embryo growth rate, calcium content of their food, and the need to mitigate excessive light transmission through the shell.     

Avian eggshell pigments are not consistently correlated with colour measurements or egg constituents in two Turdus thrushes

The colourful appearance of avian eggshells is a prominent aspect of maternal reproductive effort in birds. Some differences in eggshell coloration have been reported to co‐vary with various measures of maternal condition and these patterns support the hypothesis that, in some bird species, several aspects of eggshell colour (i.e. primary chroma and brightness) function as a signal of maternal and offspring quality to induce greater paternal investment. We directly quantified eggshell pigment concentrations of blackbird Turdus merula and song thrush T. philomelos eggshells and tested how the two key pigments (protoporphyrin IX and biliverdin) co‐varied with other eggshell traits and egg constituents as measures of maternal reproductive investment, including total yolk carotenoid concentration, total lipid concentration, yolk mass, and shell thickness. Contrary to predictions, we detected few statistical patterns overall. We found that protoporphyrin IX concentration was negatively associated with blue‐green chroma in blackbirds but not in song thrush. The concentration of protoporphyrin IX was significantly greater in blackbirds and also showed different patterns of association with total yolk lipids and yolk carotenoid concentrations between these two species (significant species interaction terms). Our results reveal that it is not appropriate to simply assume in these two avian species that reflectance‐based eggshell colour measures are a suitable proxy for eggshell pigment concentrations or can be used as consistent predictors of maternal reproductive investment. These results highlight the need to assess and validate the strength and direction of the statistical relationships between eggshell colour measures, pigment concentrations, and maternal resource deposition in the egg for other species of birds.