The age of the geese from the parent flock and the laying period affect the features of the eggs.

The study aimed to assess the goose hatching egg features in four reproductive seasons from 3 stages of laying. Three hundred sixty eggs were used in the study from geese in the first, second, third, and fourth laying season. From each group, 90 eggs were analyzed (30 eggs from the beginning, the peak, and the end of the laying cycle). The structure of the egg and morphological and physical features of the yolk, albumen, and eggshell were analyzed. It was shown that the weight and structure of eggs increased, but the shape index was lower in 2-yr-old geese, as well Haugh's units decreased. The yolk share was lower in the first year, but albumen and eggshell were higher than in other groups. The eggshell whiteness was higher in the first year than in the second, and third. The pores' quantity was higher in the first year in the blunt and equatorial parts, but the total number in the egg was the highest in the fourth year. The yolk, albumen, and eggshell' density increased with the age. Changes in laying periods were inversely proportional to the changes shown depending on the layers' age. Geese's age and laying period impact the eggs' features. Based on the egg quality features, the incubation conditions could be adapted, as well as it can be treated as an indicator of the effectiveness of hatching and goslings quality. Research has shown that the biological value of hatching eggs changes with the age of the geese and the laying period.

First ovum-in-ovo pathological titanosaurid egg throws light on the reproductive biology of sauropod dinosaurs.

Pathologic eggs have been documented in the amniote eggs of birds, turtles, and dinosaurs. These eggs occur either in the form of one egg within another egg, a condition known as ovum-in-ovo or multi-shelled eggs showing additional pathological eggshell layer/s besides the primary shell layer. Though multi-shelled eggs and eggshells were previously recorded only  in reptiles and ovum-in-ovo eggs in birds, now it has been shown that multi-shelled egg pathology occurs in birds as well. However, no ovum-in-ovo egg has been reported  in dinosaurs or for that matter  in other reptiles. Here we describe an ovum-in-ovo pathological egg from a titanosaurid dinosaur nest from the Upper Cretaceous Lameta Formation of western Central India which makes it the first report of this pathology in dinosaurs. Birds possess a specialized uterus while other amniotes have a generalized uterus. However, alligators and crocodiles retain a specialized uterus like birds along with a reptilian mode of egg-laying. The discovery of ovum-in-ovo egg from a titanosaurid dinosaur nest suggests that their oviduct morphology was similar to that of birds opening up the possibility for sequential laying of eggs in this group of sauropod dinosaurs. This new find underscores that the ovum-in-ovo pathology is not unique to birds and sauropods share a reproductive behavior very similar to that of other archosaurs.

First titanosaur dinosaur nesting site from the Late Cretaceous of Brazil.

Titanosaurs were successful herbivorous dinosaurs widely distributed in all continents during the Cretaceous, with the major diversity in South America. The success of titanosaurs was probably due to several physiological and ecological factors, in addition to a series of morphological traits they achieved during their evolutionary history. However, the generalist nesting behaviour using different palaeoenvironments and strategies was key to accomplish that success. Titanosaur nesting sites have been found extensively around the world, with notable records in Spain, France, Romania, India, and, especially, Argentina. Here, we describe the first titanosaur nesting site from the Late Cretaceous of Brazil that represents the most boreal nesting site for South America. Several egg-clutches, partially preserved, isolated eggs and many eggshell fragments were discovered in an Inceptisol palaeosol profile of the mining Lafarge Quarry, at the Ponte Alta District (Uberaba Municipality, Minas Gerais State), corresponding to the Serra da Galga Formation (Bauru Group, Bauru Basin). Although classical mechanical preparation and CT scans have not revealed embryonic remains in ovo, the eggs and eggshell features match those eggs containing titanosaurian embryos found worldwide. The morphology of the egg-clutches and observations of the sedimentary characteristics bolster the hypothesis that these sauropods were burrow-nester dinosaurs, as was already suggested for the group based on other nesting sites. The egg-clutches distributed in two levels along the Lafarge outcrops, together with the geopalaeontological data collected, provide clear evidence for the first colonial nesting and breeding area of titanosaur dinosaurs in Brazil.

Significance of eggshell morphology as an additional tool to distinguish species of sand flies (Diptera: Psychodidae: Phlebotominae).

Morphological characteristics of eggshells are important in sand fly ootaxonomy. In this study, eggshells from Phlebotomus stantoni Newstead, Sergentomyia khawi (Raynal), and Grassomyia indica (Theodor) sand flies collected in Chiang Mai province, Thailand were examined and characterized using light microscopy (LM) and scanning electron microscopy (SEM). Then, eggshell morphology of these three species was described for the first time. Each gravid female was forced to lay eggs by decapitation and the eggs were collected for SEM analysis. Egg laying females were identified by morphological examination and molecular typing using cytochrome b (Cytb) as a molecular marker. The chorionic sculpturing of Ph. stantoni eggs combines two patterns on the same egg: unconnected parallel ridges and reticular patterns. Sergentomyia khawi and Gr. indica have similar chorionic polygonal patterns, but their exochorionic morphology and aeropylar area are different. Results indicate that eggshell morphological characteristics such as chorionic pattern, exochorionic morphology, inter-ridge/boundary area, aeropylar area (including the number of aeropyles) and basal layer, can be useful to develop morphological identification keys of eggs. These can serve as an additional tool to distinguish species of sand flies. In addition, the chorionic sculpturing of the eggs of the three species of sand flies observed by LM is useful for species identification in gravid females with spermathecae obscured by eggs.

Shifts in eggshell thickness are related to changes in locomotor ecology in dinosaurs.

Birds share an array of unique characteristics among extant land vertebrates. Among these, external and microstructural characteristics of extant bird eggs have been linked to changes in reproductive strategy that arose among non-avian theropod dinosaurs. More recently, differences in egg proportions recovered in crown birds relative to other dinosaurs were suggested as possibly linked to avian flight, but dense sampling close to its proposed origin was lacking. Here we assess the evolution of eggshell thickness in a targeted sample of 114 dinosaurs including birds, and test the relationship of eggshell thickness with potential life history correlates and locomotor mode using phylogenetic comparative methods. Only egg mass and flight are identified as significant predictors of eggshell thickness. While a high correlation between egg mass and eggshell thickness is expected, that relationship is much stronger in flying taxa, which show a significantly higher slope and lower residual variance than flightless species. This suggests stabilizing selection of eggshell thickness among theropods, as recovered for other traits in extant birds (e.g. genome size, metabolic rate). Within living birds, Eufalconimorphae present an apomorphic increase in relative eggshell thickness which remains unexplained, as few morphological synapomorphies of this clade have been identified.

Geometrical characteristics of eggs from 3 poultry species.

We studied the correlations between egg geometrical parameters (i.e., egg shape index, sphericity, geometric mean diameter, surface area, and volume) and eggshell qualities, or the organic matrix in eggshell. Eggs were collected from 5 poultry breeds belonging to 3 species (commercial Hy-line Brown Chicken, Shaoxing Duck, Jinding Duck, Taihu Goose, and Zhedong White Goose). The geometrical parameters showed high variation among 3 species of poultry, and even between breeds in the same species. The five geometrical parameters were grouped into 2 sets, one contained shape index and sphericity, the other comprised geometric mean diameter, surface area, and volume. The parameters in the same set can be perfectly fitted to one another. Egg weight, shell membrane weight, and calcified shell weight were significantly correlated with geometric mean diameter, surface area, and volume. In accordance with false discovery rate-adjusted P value, both shell membrane relative weight and calcified shell thickness showed no significant correlations with any of the geometrical parameters. However, the correlations between geometrical parameters and other shell variables (calcified shell weight, shell relative weight, calcified shell thickness uniformity, and eggshell breaking strength) depend on breed. Both constitutive proportions and percentage contents of 3 eggshell matrix components (acid-insoluble, water-insoluble, and both acid and water facultative-soluble matrix) had no effects on egg shape and size. The correlations between the amounts of various shell matrix, egg shape and size depend on breed or species. This study provides a methodology and the correlation between geometrical parameters and eggshell qualities, and between geometrical parameters and organic matrix components in calcified shells.

Integrative description of a new Dactylobiotus (Eutardigrada: Parachela) from Antarctica that reveals an intraspecific variation in tardigrade egg morphology.

Tardigrades constitute one of the most important group in the challenging Antarctic terrestrial ecosystem. Living in various habitats, tardigrades play major roles as consumers and decomposers in the trophic networks of Antarctic terrestrial and freshwater environments; yet we still know little about their biodiversity. The Eutardigrada is a species rich class, for which the eggshell morphology is one of the key morphological characters. Tardigrade egg morphology shows a diverse appearance, and it is known that, despite rare, intraspecific variation is caused by seasonality, epigenetics, and external environmental conditions. Here we report Dactylobiotus ovimutans sp. nov. from King George Island, Antarctica. Interestingly, we observed a range of eggshell morphologies from the new species, although the population was cultured under controlled laboratory condition. Thus, seasonality, environmental conditions, and food source are eliminated, leaving an epigenetic factor as a main cause for variability in this case.

A giant soft-shelled egg from the Late Cretaceous of Antarctica.

Egg size and structure reflect important constraints on the reproductive and life-history characteristics of vertebrates1. More than two-thirds of all extant amniotes lay eggs2. During the Mesozoic era (around 250 million to 65 million years ago), body sizes reached extremes; nevertheless, the largest known egg belongs to the only recently extinct elephant bird3, which was roughly 66 million years younger than the last nonavian dinosaurs and giant marine reptiles. Here we report a new type of egg discovered in nearshore marine deposits from the Late Cretaceous period (roughly 68 million years ago) of Antarctica. It exceeds all nonavian dinosaur eggs in volume and differs from them in structure. Although the elephant bird egg is slightly larger, its eggshell is roughly five times thicker and shows a substantial prismatic layer and complex pore structure4. By contrast, the new fossil, visibly collapsed and folded, presents a thin eggshell with a layered structure that lacks a prismatic layer and distinct pores, and is similar to that of most extant lizards and snakes (Lepidosauria)5. The identity of the animal that laid the egg is unknown, but these preserved morphologies are consistent with the skeletal remains of mosasaurs (large marine lepidosaurs) found nearby. They are not consistent with described morphologies of dinosaur eggs of a similar size class. Phylogenetic analyses of traits for 259 lepidosaur species plus outgroups suggest that the egg belonged to an individual that was at least 7 metres long, hypothesized to be a giant marine reptile, all clades of which have previously been proposed to show live birth6. Such a large egg with a relatively thin eggshell may reflect derived constraints associated with body shape, reproductive investment linked with gigantism, and lepidosaurian viviparity, in which a 'vestigial' egg is laid and hatches immediately7.

The first dinosaur egg was soft.

Calcified eggshells protect developing embryos against environmental stress and contribute to reproductive success1. As modern crocodilians and birds lay hard-shelled eggs, this eggshell type has been inferred for non-avian dinosaurs. Known dinosaur eggshells are characterized by an innermost membrane, an overlying protein matrix containing calcite, and an outermost waxy cuticle2-7. The calcitic eggshell consists of one or more ultrastructural layers that differ markedly among the three major dinosaur clades, as do the configurations of respiratory pores. So far, only hadrosaurid, a few sauropodomorph and tetanuran eggshells have been discovered; the paucity of the fossil record and the lack of intermediate eggshell types challenge efforts to homologize eggshell structures across all dinosaurs8-18. Here we present mineralogical, organochemical and ultrastructural evidence for an originally non-biomineralized, soft-shelled nature of exceptionally preserved ornithischian Protoceratops and basal sauropodomorph Mussaurus eggs. Statistical evaluation of in situ Raman spectra obtained for a representative set of hard- and soft-shelled, fossil and extant diapsid eggshells clusters the originally organic but secondarily phosphatized Protoceratops and the organic Mussaurus eggshells with soft, non-biomineralized eggshells. Histology corroborates the organic composition of these soft-shelled dinosaur eggs, revealing a stratified arrangement resembling turtle soft eggshell. Through an ancestral-state reconstruction of composition and ultrastructure, we compare eggshells from Protoceratops and Mussaurus with those from other diapsids, revealing that the first dinosaur egg was soft-shelled. The calcified, hard-shelled dinosaur egg evolved independently at least three times throughout the Mesozoic era, explaining the bias towards eggshells of derived dinosaurs in the fossil record.

The shell gland in laying and natural moulting commercial egg-type chickens: A histomorphological and ultrastructural study.

The purpose of the present study was to determine the histological and ultrastructural changes in the luminal epithelium of the shell gland associated with natural moulting. Samples of the shell gland from laying (32 weeks old) and moulting (75 weeks old) hens were studied using histological, histochemical and electron microscopic techniques. In addition, TUNEL was used to demonstrate the distribution of apoptotic cells in the luminal epithelium of the shell gland. Autophagy, characterized by the presence of autophagosomes and autolysosomes, was evident in the early stages of degeneration in non-ciliated, ciliated and mitochondrial cells. The intermediate and advanced stages of regression in non-ciliated as well as mitochondrial cells occurred via apoptosis, while both apoptotic and necrotic ciliated cells were observed during the later stages of degeneration. The results of the present study suggest that a synergy of autophagy, apoptosis and necrosis is involved in the involution of the shell gland during natural moulting.

Ostrich eggshell bead diameter in the Holocene: Regional variation with the spread of herding in eastern and southern Africa.

Despite their ubiquity in Holocene African archaeological assemblages, ostrich eggshell (OES) beads are rarely studied in detail. An exception is in southern Africa, where there is a proposed relationship between OES bead diameter and the arrival of herding ~2000 years before present. In 1987, Leon Jacobson first observed that beads from forager sites in Namibia tended to be smaller than those associated with herder sites. Studies examining bead size around the Western Cape have generally confirmed Jacobson's findings, though the driving forces of the diameter change remain unknown. Since this time, diameter has become an informal way of distinguishing forager and herder assemblages in southern Africa, but no large-scale studies of OES bead variation have been undertaken. Here we present an expanded analysis of Holocene OES bead diameters from southern, and for the first time, eastern Africa. Results reveal distinct patterns in OES bead size over time, reflecting different local dynamics associated with the spread of herding. In southern Africa, OES diameters display low variability and smaller absolute size through time. While larger beads begin to appear <2000 years ago, most beads in our study remained smaller. In contrast, eastern African OES bead diameters are consistently larger over the last 10,000 years and show no appreciable size change with the introduction of herding. Notably, larger beads thought to be associated with herders in southern Africa fall within the range of eastern African beads, indicating a potential connection between these regions in the Late Holocene consistent with genetic findings. Regional differences in bead size are subtle, on the order of millimeters, yet offer a potentially important line of evidence for investigating the spread of herding in sub-Saharan Africa. In order to understand the meaning of these changes, we encourage additional studies of OES bead assemblages and urge researchers to report individual bead diameters, rather than averages by level.

An Early Cretaceous enantiornithine (Aves) preserving an unlaid egg and probable medullary bone.

Understanding non-crown dinosaur reproduction is hindered by a paucity of directly associated adults with reproductive traces. Here we describe a new enantiornithine, Avimaia schweitzerae gen. et sp. nov., from the Lower Cretaceous Xiagou Formation with an unlaid egg two-dimensionally preserved within the abdominothoracic cavity. Ground-sections reveal abnormal eggshell proportions, and multiple eggshell layers best interpreted as a multi-layered egg resulting from prolonged oviductal retention. Fragments of the shell membrane and cuticle are both preserved. SEM reveals that the cuticle consists of nanostructures resembling those found in neornithine eggs adapted for infection-prone environments, which are hypothesized to represent the ancestral avian condition. The femur preserves small amounts of probable medullary bone, a tissue found today only in reproductively active female birds. To our knowledge, no other occurrence of Mesozoic medullary bone is associated with indications of reproductive activity, such as a preserved egg, making our identification unique, and strongly supported.

Structure and evolutionary implications of the earliest (Sinemurian, Early Jurassic) dinosaur eggs and eggshells.

One of the fossil record's most puzzling features is the absence of preserved eggs or eggshell for the first third of the known 315 million year history of amniote evolution. Our meagre understanding of the origin and evolution of calcareous eggshell and amniotic eggs in general, is largely based on Middle Jurassic to Late Cretaceous fossils. For dinosaurs, the most parsimonious inference yields a thick, hard shelled egg, so richly represented in the Late Cretaceous fossil record. Here, we show that a thin calcareous layer (≤100 µm) with interlocking units of radiating crystals (mammillae) and a thick shell membrane already characterize the oldest known amniote eggs, belonging to three coeval, but widely distributed Early Jurassic basal sauropodomorph dinosaurs. This thin shell layer strongly contrasts with the considerably thicker calcareous shells of Late Jurassic dinosaurs. Phylogenetic analyses and their Sinemurian age indicate that the thin eggshell of basal sauropodomorphs represents a major evolutionary innovation at the base of Dinosauria and that the much thicker eggshell of sauropods, theropods, and ornithischian dinosaurs evolved independently. Advanced mineralization of amniote eggshell (≥150 µm in thickness) in general occurred not earlier than Middle Jurassic and may correspond with a global trend of increase in atmospheric oxygen.

Previtellogenic oocytes of South African largemouth bass Micropterus salmoides Lacépède 1802 (Actinopterygii, Perciformes) - the Balbiani body, cortical alveoli and developing eggshell.

The ovaries of the largemouth bass Micropterus salmoides, an alien and invasive species in South Africa, contain a germinal epithelium which consists of germline and somatic cells, as well as previtellogenic and late vitellogenic ovarian follicles. The ovarian follicle consists of an oocyte surrounded by follicular cells and a basal lamina; thecal cells adjacent to this lamina are covered by an extracellular matrix. In this article, we describe the Balbiani body and the polarization and ultrastructure of the cytoplasm (ooplasm) in previtellogenic oocytes. The nucleoplasm in all examined oocytes contains lampbrush chromosomes, nuclear bodies and several nucleoli near the nuclear envelope. The ultrastructure of the nucleoli is described. Numerous nuage aggregations are present in the perinuclear cytoplasm in germline cells as well as in the ooplasm. Possible roles of these aggregations are discussed. The ooplasm contains the Balbiani body, which defines the future vegetal region in early previtellogenic oocytes. It is comprised of nuage aggregations, rough endoplasmic reticulum, Golgi apparatus, mitochondria, complexes of mitochondria with nuage-like material, and lysosome-like organelles. In mid-previtellogenic oocytes, the Balbiani body surrounds the nucleus and later disperses in the ooplasm. The lysosome-like organelles fuse and transform into vesicles containing material which is highly electron dense. As a result of the fusion of the vesicles of Golgi and rough endoplasmic reticulum, the cortical alveoli arise and distribute uniformly throughout the ooplasm of late previtellogenic oocytes. During this stage, the deposition of the eggshell (zona radiata) begins. The eggshell is penetrated by canals containing microvilli and consists of the following: the internal and the external egg envelope. In the external envelope three sublayers can be distinguished.

A mixed vertebrate eggshell assemblage from the Transylvanian Late Cretaceous.

A Late Cretaceous-aged multi-taxon nesting site from Romania preserved in three dimensions reveals the earliest example of nest site sharing yet known from the vertebrate fossil record. Eggshell and osteological evidence combined in this single accumulation demonstrate that at least four vertebrate taxa including enantiornithine birds and another avian of indeterminate affinities as well as crocodylomorphs and gekkotan squamates nested together in the same place. Colonial nesting in enantiornithines was previously described from this site; here, we present the first fossil evidence that other vertebrates also nested in the same place, perhaps exploiting the presence of the large bird colony. We describe four distinct eggshell morphotypes that have been collected from this site and draw palaeoecological inferences based on this unique multi-taxon nesting association.

Experimental increase in temperature affects eggshell thickness, and not egg mass, eggshell spottiness or egg composition in the great tit (Parus major).

Phenotypic effects of global warming have been documented in many different taxa. However, the importance of transgenerational phenotypic plasticity in these adaptations are seldom studied. In birds, temperature could affect egg characteristics. Higher temperatures during egg-laying may reduce maintenance costs for females and allow a higher investment in reproduction. Yet, females may also use temperatures as a cue for the risk of mismatch latter in the season. Thus, higher temperatures may be correlated to an acceleration of embryonic development (e.g. via hormonal manipulation). We performed an experiment in which night-time temperature was increased in the nestbox by approximately 1 °C throughout the entire laying period in great tits (Parus major). We collected one pre-treatment egg (beginning of the laying sequence) and one post-treatment egg (end of the laying sequence). Egg content (yolk androgens and lysozymes in the albumen), eggshell coloration, eggshell mass, egg mass, and shape were not affected by the treatment. However, last-laid eggs in clutches from control nestboxes had a thicker eggshell than last-laid eggs from heated nestboxes, suggesting a putative slight decrease of maternal investment with the experimental increase of temperature. We also observed effects of the laying sequence on egg characteristics. Eggs that were laid late in the laying sequence were heavier, larger, had larger spots and higher yolk androgens than eggs laid earlier. Lysozyme concentration decrease with the laying sequence in late clutches only. Thus, effects of temperature may also change with the laying sequence and it would be interesting in the future to tests the effects on first-laid eggs.

Effect of direct fed microbial in layer diets, on egg laying performance and health response of indigenous Ghanaian guinea fowls (Numida meleagris).

The aim of the present experiment was to examine the effect of direct fed microbial (DFM) on egg laying performance and health response of indigenous Ghanaian guinea fowls (Numida meleagris). A total of 216, 9-wk-old guinea keets (already sexed) were randomly assigned to 4 DFM administering treatments. These included control, daily, 3 consecutive days per week, and 7 d repeated every other week at 1.5 mL/L through water for 30 wk. Feed intake, body weight gain, feed conversion ratio, egg weight, yolk weight, egg albumen weight, shell weight, shell thickness, mortality, hematological and serum biochemical parameters were recorded. Birds on daily DFM treatment consumed the least feed (P = 0.007) with an increased weight gain (P = 0.009). Hen-day egg production, eggshell thickness and eggshell weight, albumen weight, albumen height, yolk weight, and yolk height were not different between treatments (P > 0.05). Egg weight was higher in daily supplementation of DFM in water than other treatments (P < 0.0001). Serum albumin was higher (P = 0.024) in daily DFM treatment, whereas serum low-density lipoprotein (LDL) cholesterol concentration was reduced (P = 0.017). The results showed that supplementing DFM daily at 1.5 mL/L of water resulted in increased total proteins and albumin as well as reduced LDL cholesterol. It could be concluded that these health indices increased the quality of guinea fowls eggs and may have improved the hatchability of eggs in the 1.5 mL/L DFM supplementation and there the supplementing DFM daily at 1.5 mL/L is recommended in Guinea fowls to increase hatchling numbers.

Dinosaur egg colour had a single evolutionary origin.

Birds are the only living amniotes with coloured eggs1-4, which have long been considered to be an avian innovation1,3. A recent study has demonstrated the presence of both red-brown protoporphyrin IX and blue-green biliverdin5-the pigments responsible for all the variation in avian egg colour-in fossilized eggshell of a nonavian dinosaur6. This raises the fundamental question of whether modern birds inherited egg colour from their nonavian dinosaur ancestors, or whether egg colour evolved independently multiple times. Here we present a phylogenetic assessment of egg colour in nonavian dinosaurs. We applied high-resolution Raman microspectroscopy to eggshells that represent all of the major clades of dinosaurs, and found that egg colour pigments were preserved in all eumaniraptorans: egg colour had a single evolutionary origin in nonavian theropod dinosaurs. The absence of colour in ornithischian and sauropod eggs represents a true signal rather than a taphonomic artefact. Pigment surface maps revealed that nonavian eumaniraptoran eggs were spotted and speckled, and colour pattern diversity in these eggs approaches that in extant birds, which indicates that reproductive behaviours in nonavian dinosaurs were far more complex than previously known3. Depth profiles demonstrated identical mechanisms of pigment deposition in nonavian and avian dinosaur eggs. Birds were not the first amniotes to produce coloured eggs: as with many other characteristics7,8 this is an attribute that evolved deep within the dinosaur tree and long before the spectacular radiation of modern birds.

Common guillemot (Uria aalge) eggs are not self-cleaning.

Birds are arguably the most evolutionarily successful extant vertebrate taxon, in part because of their ability to reproduce in virtually all terrestrial habitats. Common guillemots, Uria aalge, incubate their single egg in an unusual and harsh environment; on exposed cliff ledges, without a nest, and in close proximity to conspecifics. As a consequence, the surface of guillemot eggshells is frequently contaminated with faeces, dirt, water and other detritus, which may impede gas exchange or facilitate microbial infection of the developing embryo. Despite this, guillemot chicks survive incubation and hatch from eggs heavily covered with debris. To establish how guillemot eggs cope with external debris, we tested three hypotheses: (1) contamination by debris does not reduce gas exchange efficacy of the eggshell to a degree that may impede normal embryo development; (2) the guillemot eggshell surface is self-cleaning; (3) shell accessory material (SAM) prevents debris from blocking pores, allowing relatively unrestricted gas diffusion across the eggshell. We showed that natural debris reduces the conductance of gases across the guillemot eggshell by blocking gas exchange pores. Despite this problem, we found no evidence that guillemot eggshells are self-cleaning, but instead showed that the presence of SAM on the eggshell surface largely prevents pore blockages from occurring. Our results demonstrate that SAM is a crucial feature of the eggshell surface in a species with eggs that are frequently in contact with debris, acting to minimise pore blockages and thus ensure a sufficient rate of gas diffusion for embryo development.

An Intermediate Incubation Period and Primitive Brooding in a Theropod Dinosaur.

Non-avian dinosaurs such as oviraptorosaurs and troodontids share several important reproductive characters with modern birds, including eggshell microstructure and iterative egg production. Nevertheless, debate exists concerning their incubation strategies. Here we estimate incubation period for the troodontid, Troodon formosus, by examining a near-term embryonic tooth. Synchrotron scanning and histologic thin sections allowed counting of daily (von Ebner) growth lines. The tooth preserves 31 intact lines with an average spacing of 3.3 ± 0.96 µm. Adding 8 more for the missing crown tip gives a total age of 39 days. Modern crocodilians begin to establish their functional dentition at approximately 47% through incubation. Thus, this tooth age suggests a Troodon incubation period of 74 days, falling midway between avian (44.4 days) and reptilian (107.3 days) values predicted by the Troodon egg mass (314 g). An accelerated incubation relative to modern reptiles supports brooding and concurs with a suite of features in oviraptorosaurs and troodontids (sequential laying, large complex clutches, and precocial young) that appear dependent upon both adult body and incubation temperatures elevated over ambient conditions. However, the largely buried condition of Troodon clutches may have prohibited efficient brooding, necessitating longer incubation than that of modern birds with fully exposed eggs.