Out of Africa: Juvenile Dispersal of Black-Shouldered Kites in the Emerging European Population.

Knowledge of animal dispersal patterns is of great importance for the conservation and maintenance of natural populations. We here analyze juvenile dispersal of the poorly studied Black-shouldered Kite (Elanus caeruleus) monitored in southwestern Spain in an ongoing long-term study initiated in 2003. The European population of Black-shouldered kites is thought to be a recent one funded by colonizing African birds, as no kites have been found in the European fossil record, and the breeding population has progressively expanded to the North in the late 20th and 21st centuries. We obtained information on movements behavior during dispersal from 47 juveniles Kites after marking 384 nestlings with wing tags and three nestlings with radio transmitter. We have tested two competing hypotheses (i.e., the Resources Competition Hypothesis and the Wandering Hypothesis (WH)) that may explain the leptokurtic distribution of the natal dispersal distance in Elanus. After independence, juvenile females dispersed farther from the natal areas than males, as is common in birds. On average, males and females dispersed from their natal areas over 9 (i.e., 26.15 km) and 15 (i.e., 43.79 km) breeding territories, respectively. A male and two females dispersed further than 100 km from their natal nest. Our results indicated some evidence supporting the competition-for-resources hypotheses since nestlings hatched from high quality territories stayed closer from natal areas than nestlings hatched from low quality territories and also nestlings hatched first within the brood also tend to recruit closer to their natal area than later hatched nestlings which tend to disperse further away from their natal area. The information provided by these crucial demographic parameters will be used for the elaboration of future conservation plans for the management of this colonizing species in Europe.

Leucistic plumage as a result of progressive greying in a cryptic nocturnal bird.

Leucism, broadly defined as the lack of melanin pigmentation, occurs in many animal species. Most studies on leucism and other colour aberrations are based on opportunistic observations or small cross-sectional samples, thus limiting our ability to produce reliable results and test theoretical predictions. This study combines cross-sectional and longitudinal data collected in 2016-2020 from a population of red-necked nightjars (Caprimulgus ruficollis). The goals of the study are (i) to investigate sex and age effects on partial leucism, (ii) to separate within-subject effects (progressive greying) from between-subject effects (selective disappearance), and (iii) to examine differences in body mass, structural size, and life span between leucistic and non-leucistic individuals. The probability of leucism in nightjars increased from juveniles to adults at similar rates in males and females. Our longitudinal analysis and life-span comparisons indicated a minor contribution of selective disappearance to age-related changes in leucism, but rather suggested that the loss of melanin from feathers can be attributed to progressive greying in ageing adults. Body mass and size were consistently smaller (5% and 1.5%, respectively) in leucistic than in non-leucistic nightjars, although the reason for this difference remains unclear. Our study sheds light on the sources and mechanisms of variation in leucism in natural populations and its relationship with important life-history traits, such as life span.

Taphonomic information from the modern vertebrate death assemblage of Doñana National Park, Spain.

Modern death assemblages provide insights about the early stages of fossilization and useful ecological information about the species inhabiting the ecosystem. We present the results of taphonomic monitoring of modern vertebrate carcasses and bones from Doñana National Park, a Mediterranean coastal ecosystem in Andalusia, Spain. Ten different habitats were surveyed. Half of them occur in active depositional environments (marshland, lake margin, river margin, beach and dunes). Most of the skeletal remains belong to land mammals larger than 5 kg in body weight (mainly wild and feral ungulates). Overall, the Doñana bone assemblage shows good preservation with little damage to the bones, partly as a consequence of the low predator pressure on large vertebrates. Assemblages from active depositional habitats differ significantly from other habitats in terms of the higher incidence of breakage and chewing marks on bones in the latter, which result from scavenging, mainly by wild boar and red fox. The lake-margin and river-margin death assemblages have high concentrations of well preserved bones that are undergoing burial and offer the greatest potential to produce fossil assemblages. The spatial distribution of species in the Doñana death assemblage generally reflects the preferred habitats of the species in life. Meadows seem to be a preferred winter habitat for male deer, given the high number of shed antlers recorded there. This study is further proof that taphonomy can provide powerful insights to better understand the ecology of modern species and to infer past and future scenarios for the fossil record.

Long-Term Determinants of Tuberculosis in the Ungulate Host Community of Doñana National Park.

Animal tuberculosis (TB) is endemic in wild boar (Sus scrofa), red deer (Cervus elaphus), fallow deer (Dama dama) and cattle in south and central Spain. In order to clarify the processes that operate in the medium and long-term, we studied TB at the wildlife-livestock interface in Doñana National Park for 14 years (2006-2018) in relation to host density, stochastic factors (rainfall) and environmental features (e.g., aggregation points such as waterholes). Wild boar showed the highest prevalence of TB (76.7%), followed by red deer (42.5%), fallow deer (14.4%) and cattle (10.7%). We found evidence of relevant epidemiological processes which operate over the long-term and interact with host and community ecology. Interestingly, the effect of high wild boar population density on increased TB rates was mediated by sows, which could determine high incidence in young individuals already in maternal groups. Rainfall significantly determined a higher risk of TB in male red deer, probably mediated by sex-related differences in life history traits that determined more susceptibility and/or exposure in comparison to females. The positive association between the prevalence of TB in fallow deer and cattle may indicate significant interspecies transmission (in either direction) and/or similar exposure to risk factors mediated by ecological overlapping of grazing species. The identification of long-term drivers of TB provided evidence that its control in extensive pastoral systems can only be achieved by targeting all relevant hosts and integrating measures related to all the factors involved, such as: population abundance and the aggregation of wild and domestic ungulates, environmental exposure to mycobacteria, cattle testing and culling campaigns and adjustments of appropriate densities.

Correlates of individual variation in the porphyrin-based fluorescence of red-necked nightjars (Caprimulgus ruficollis).

Many nocturnal animals, including invertebrates such as scorpions and a variety of vertebrate species, including toadlets, flying squirrels, owls, and nightjars, emit bright fluorescence under ultraviolet light. However, the ecological significance of this unique coloration so attached to nocturnality remains obscure. Here, we used an intensively studied population of migratory red-necked nightjars (Caprimulgus ruficollis) to investigate inter-individual variation in porphyrin-based pink fluorescence according to sex, age, body condition, time of the year, and the extent of white plumage patches known to be involved in sexual communication. Males and females exhibited a similar extent of pink fluorescence on the under-side of the wings in both juvenile and adult birds, but males had larger white patches than females. Body condition predicted the extent of pink fluorescence in juvenile birds, but not in adults. On average, the extent of pink fluorescence in juveniles increased by ca. 20% for every 10-g increase in body mass. For both age classes, there was a slight seasonal increase (1-4% per week) in the amount of fluorescence. Our results suggest that the porphyrin-based coloration of nightjars might signal individual quality, at least in their first potential breeding season, although the ability of these and other nocturnal birds to perceive fluorescence remains to be unequivocally proven.

Cosmetic colouring by Bearded Vultures Gypaetus barbatus: still no evidence for an antibacterial function.

Bearded Vultures regularly visit ferruginous springs for cosmetic purposes to obtain their reddish plumage colouration. Different hypotheses have been proposed to explain this deliberate application of adventitious colouration: (1) to signal individual dominance status; (2) to exploit an anti-bacterial effect of iron oxides or ochre to reduce feather degradation by bacteria and, in parallel (3) to enable incubating birds to transfer this protection to their developing embryos to increase hatching success. Here, we re-evaluate the antibacterial hypothesis using three experimental approaches: (a) by applying feather-degrading bacteria to stained and unstained bearded vulture feathers; (b) by assessing the antibacterial activity of ochre; and (c) by comparing the breeding success of orange individuals with pale ones. Our findings suggest that the in vitro addition of feather degrading Bacillus licheniformis to naturally stained Bearded Vulture feathers did not retard feather degradation compared to controls. Iron particles from red soil (ochre) or iron salts had no antibacterial effect on the growth of three species of bacteria (Escherichia coli, Kocuria rhizophila and Bacillus licheniformis), incubated either in the dark or under visible light. Finally, breeding success did not differ between territories occupied by pale individuals versus orange ones. These results run counter to the hypothesis that iron oxides have an antibacterial role in Bearded Vultures. The use of red soils by Bearded Vultures may function as a territorial status signal, but may also be involved in other processes, such as pair formation and the long-term maintenance of the pair bond, as suggested for the closely related Egyptian vulture Neophron percnopterus.

Unmanned aircraft systems as a new source of disturbance for wildlife: A systematic review.

The use of small Unmanned Aircraft Systems (UAS; also known as "drones") for professional and personal-leisure use is increasing enormously. UAS operate at low altitudes (<500 m) and in any terrain, thus they are susceptible to interact with local fauna, generating a new type of anthropogenic disturbance that has not been systematically evaluated. To address this gap, we performed a review of the existent literature about animals' responses to UAS flights and conducted a pooled analysis of the data to determine the probability and intensity of the disturbance, and to identify the factors influencing animals' reactions towards the small aircraft. We found that wildlife reactions depended on both the UAS attributes (flight pattern, engine type and size of aircraft) and the characteristics of animals themselves (type of animal, life-history stage and level of aggregation). Target-oriented flight patterns, larger UAS sizes, and fuel-powered (noisier) engines evoked the strongest reactions in wildlife. Animals during the non-breeding period and in large groups were more likely to show behavioral reactions to UAS, and birds are more prone to react than other taxa. We discuss the implications of these results in the context of wildlife disturbance and suggest guidelines for conservationists, users and manufacturers to minimize the impact of UAS. In addition, we propose that the legal framework needs to be adapted so that appropriate actions can be undertaken when wildlife is negatively affected by these emergent practices.

The bare head of the Northern bald ibis (Geronticus eremita) fulfills a thermoregulatory function.

BACKGROUND: Dark pigments provide animals with several adaptive benefits such as protection against ultraviolet (UV) radiation and mechanical abrasion, but may also impose several constraints like a high absorbance of solar radiation. Endotherms, with relatively constant and high body temperatures, may be especially prone to thermoregulatory limitations if dark coloured and inhabiting hot environments. It is therefore expected that adaptations have specifically evolved because of these limitations. Bare, highly vascularised head skin may have evolved in birds with dark plumage from hot geographical regions because of favouring heat dissipation. Using the Northern bald ibis (Geronticus eremita) as a model species, we measured the surface temperature (Tsurf) of the head, the bill and the black feathered body of 11 birds along ambient temperatures (Ta) ranging from 21 to 42.5 °C employing thermal imaging. RESULTS: While Tsurf of the bill and the feathered body was only slightly above Ta, head Tsurf was considerably higher, by up to 12 °C. Estimated values of heat loss followed similar variations. We also found that the red colour intensity of the head of ibises increased with head Tsurf, suggesting that birds are capable of controlling blood flow and the thermoregulatory function of the head. CONCLUSIONS: These findings are consistent with the hypothesis that bare skin has evolved in dark pigmented birds inhabiting hot environments because of their ability to dissipate heat.

Tropical bat as mammalian model for skin carotenoid metabolism.

Animals cannot synthesize carotenoid pigments de novo, and must consume them in their diet. Most mammals, including humans, are indiscriminate accumulators of carotenoids but inefficiently distribute them to some tissues and organs, such as skin. This limits the potential capacity of these organisms to benefit from the antioxidant and immunostimulatory functions that carotenoids fulfill. Indeed, to date, no mammal has been known to have evolved physiological mechanisms to incorporate and deposit carotenoids in the skin or hair, and mammals have therefore been assumed to rely entirely on other pigments such as melanins to color their integument. Here we use high-performance liquid chromatography (HPLC) in combination with time-of-flight mass spectrometry (HPLC-TOF/MS) to show that the frugivorous Honduran white bat Ectophylla alba colors its skin bright yellow with the deposition of the xanthophyll lutein. The Honduran white bat is thus a mammalian model that may help developing strategies to improve the assimilation of lutein in humans to avoid macular degeneration. This represents a change of paradigm in animal physiology showing that some mammals actually have the capacity to accumulate dietary carotenoids in the integument. In addition, we have also discovered that the majority of the lutein in the skin of Honduran white bats is present in esterified form with fatty acids, thereby permitting longer-lasting coloration and suggesting bright color traits may have an overlooked role in the visual communication of bats.

Decoration Increases the Conspicuousness of Raptor Nests.

Avian nests are frequently concealed or camouflaged, but a number of species builds noticeable nests or use conspicuous materials for nest decoration. In most cases, nest decoration has a role in mate choice or provides thermoregulatory or antiparasitic benefits. In territorial species however, decorations may serve additional or complementary functions, such as extended phenotypic signaling of nest-site occupancy and social status to potential intruders. The latter may benefit both signaler and receiver by minimizing the risk of aggressive interactions, especially in organisms with dangerous weaponry. Support for this hypothesis was recently found in a population of black kites (Milvus migrans), a territorial raptor that decorates its nest with white artificial materials. However, the crucial assumption that nest decorations increased nest-site visibility to conspecifics was not assessed, a key aspect given that black kite nests may be well concealed within the canopy. Here, we used an unmanned aircraft system to take pictures of black kite nests, with and without an experimentally placed decoration, from different altitudes and distances simulating the perspective of a flying and approaching, prospecting intruder. The pictures were shown to human volunteers through a standardized routine to determine whether detection rates varied according the nest decoration status and distance. Decorated nests consistently showed a higher detection frequency and a lower detection-latency, compared to undecorated versions of the same nests. Our results confirm that nest decoration in this species may act as a signaling medium that enhances nest visibility for aerial receivers, even at large distances. This finding complements previous work on this communication system, which showed that nest decoration was a threat informing trespassing conspecifics on the social dominance, territory quality and fighting capabilities of the signaler.

Unmanned Aircraft Systems complement biologging in spatial ecology studies.

The knowledge about the spatial ecology and distribution of organisms is important for both basic and applied science. Biologging is one of the most popular methods for obtaining information about spatial distribution of animals, but requires capturing the animals and is often limited by costs and data retrieval. Unmanned Aircraft Systems (UAS) have proven their efficacy for wildlife surveillance and habitat monitoring, but their potential contribution to the prediction of animal distribution patterns and abundance has not been thoroughly evaluated. In this study, we assess the usefulness of UAS overflights to (1) get data to model the distribution of free-ranging cattle for a comparison with results obtained from biologged (GPS-GSM collared) cattle and (2) predict species densities for a comparison with actual density in a protected area. UAS and biologging derived data models provided similar distribution patterns. Predictions from the UAS model overestimated cattle densities, which may be associated with higher aggregated distributions of this species. Overall, while the particular researcher interests and species characteristics will influence the method of choice for each study, we demonstrate here that UAS constitute a noninvasive methodology able to provide accurate spatial data useful for ecological research, wildlife management and rangeland planning.

A rock engraving made by Neanderthals in Gibraltar.

The production of purposely made painted or engraved designs on cave walls--a means of recording and transmitting symbolic codes in a durable manner--is recognized as a major cognitive step in human evolution. Considered exclusive to modern humans, this behavior has been used to argue in favor of significant cognitive differences between our direct ancestors and contemporary archaic hominins, including the Neanderthals. Here we present the first known example of an abstract pattern engraved by Neanderthals, from Gorham's Cave in Gibraltar. It consists of a deeply impressed cross-hatching carved into the bedrock of the cave that has remained covered by an undisturbed archaeological level containing Mousterian artifacts made by Neanderthals and is older than 39 cal kyr BP. Geochemical analysis of the epigenetic coating over the engravings and experimental replication show that the engraving was made before accumulation of the archaeological layers, and that most of the lines composing the design were made by repeatedly and carefully passing a pointed lithic tool into the grooves, excluding the possibility of an unintentional or utilitarian origin (e.g., food or fur processing). This discovery demonstrates the capacity of the Neanderthals for abstract thought and expression through the use of geometric forms.

The earliest pigeon fanciers.

Feral Pigeons have colonised all corners of the Earth, having developed a close association with humans and their activities. The wild ancestor of the Feral Pigeon, the Rock Dove, is a species of rocky habitats, nesting typically on cliff ledges and at the entrance to large caves. This habit would have brought them into close contact with cave-dwelling humans, a relationship usually linked to the development of dwellings in the Neolithic. We show that the association between humans and Rock Doves is an ancient one with its roots in the Palaeolithic and predates the arrival of modern humans into Europe. At Gorham's Cave, Gibraltar, the Neanderthals exploited Rock Doves for food for a period of over 40 thousand years, the earliest evidence dating to at least 67 thousand years ago. We show that the exploitation was not casual or sporadic, having found repeated evidence of the practice in different, widely spaced, temporal contexts within the cave. Our results point to hitherto unappreciated capacities of the Neanderthals to exploit birds as food resources on a regular basis. More so, they were practising it long before the arrival of modern humans and had therefore invented it independently.

Determinants and short-term physiological consequences of PHA immune response in lesser kestrel nestlings.

Individual immune responses are likely affected by genetic, physiological, and environmental determinants. We studied the determinants and short-term consequences of Phytohaemagglutinin (PHA) induced immune response, a commonly used immune challenge eliciting both innate and acquired immunity, on lesser kestrel (Falco naumanni) nestlings in semi-captivity conditions and with a homogeneous diet composition. We conducted a repeated measures analyses of a set of blood parameters (carotenoids, triglycerides, ß-hydroxybutyrate, cholesterol, uric acid, urea, total proteins, and total antioxidant capacity), metabolic (resting metabolic rate), genotypic (MHC class II B heterozygosity), and biometric (body mass) variables. PHA challenge did not affect the studied physiological parameters on a short-term basis (<12 hr), except plasma concentrations of triglycerides and carotenoids, which decreased and increased, respectively. Uric acid was the only physiological parameter correlated with the PHA induced immune response (skin swelling), but the change of body mass, cholesterol, total antioxidant capacity, and triglycerides between sessions (i.e., post-pre treatment) were also positively correlated to PHA response. No relationships were detected between MHC gene heterozygosity or resting metabolic rate and PHA response. Our results indicate that PHA response in lesser kestrel nestlings growing in optimal conditions does not imply a severe energetic cost 12 hr after challenge, but is condition-dependent as a rapid mobilization of carotenoids and decrease of triglycerides is elicited on a short-term basis.

Birds of a feather: Neanderthal exploitation of raptors and corvids.

The hypothesis that Neanderthals exploited birds for the use of their feathers or claws as personal ornaments in symbolic behaviour is revolutionary as it assigns unprecedented cognitive abilities to these hominins. This inference, however, is based on modest faunal samples and thus may not represent a regular or systematic behaviour. Here we address this issue by looking for evidence of such behaviour across a large temporal and geographical framework. Our analyses try to answer four main questions: 1) does a Neanderthal to raptor-corvid connection exist at a large scale, thus avoiding associations that might be regarded as local in space or time?; 2) did Middle (associated with Neanderthals) and Upper Palaeolithic (associated with modern humans) sites contain a greater range of these species than Late Pleistocene paleontological sites?; 3) is there a taphonomic association between Neanderthals and corvids-raptors at Middle Palaeolithic sites on Gibraltar, specifically Gorham's, Vanguard and Ibex Caves? and; 4) was the extraction of wing feathers a local phenomenon exclusive to the Neanderthals at these sites or was it a geographically wider phenomenon?. We compiled a database of 1699 Pleistocene Palearctic sites based on fossil bird sites. We also compiled a taphonomical database from the Middle Palaeolithic assemblages of Gibraltar. We establish a clear, previously unknown and widespread, association between Neanderthals, raptors and corvids. We show that the association involved the direct intervention of Neanderthals on the bones of these birds, which we interpret as evidence of extraction of large flight feathers. The large number of bones, the variety of species processed and the different temporal periods when the behaviour is observed, indicate that this was a systematic, geographically and temporally broad, activity that the Neanderthals undertook. Our results, providing clear evidence that Neanderthal cognitive capacities were comparable to those of Modern Humans, constitute a major advance in the study of human evolution.

Specific appetite for carotenoids in a colorful bird.

BACKGROUND: Since carotenoids have physiological functions necessary for maintaining health, individuals should be selected to actively seek and develop a specific appetite for these compounds. METHODOLOGY/PRINCIPAL FINDINGS: Great tits Parus major in a diet choice experiment, both in captivity and the field, preferred carotenoid-enriched diets to control diets. The food items did not differ in any other aspects measured besides carotenoid content. CONCLUSIONS/SIGNIFICANCE: Specific appetite for carotenoids is here demonstrated for the first time, placing these compounds on a par with essential nutrients as sodium or calcium.

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.

Increase in membrane thickness during development compensates for eggshell thinning due to calcium uptake by the embryo in falcons.

We compared membrane thickness of fully developed eggs with those of non-developed eggs in different endangered falcon taxa. To our knowledge, membrane thickness variation during development has never been examined before in falcons or any other wild bird. Yet, the egg membrane constitutes an important protective barrier for the developing embryo. Because eggshell thinning is a general process that occurs during bird development, caused by calcium uptake by the embryo, eggs are expected to be less protected and vulnerable to breakage near the end of development. Thus, egg membranes could play an important protective role in the later stages of development by getting relatively thicker. We used linear mixed models to explore the variation in membrane thickness (n = 378 eggs) in relation to developmental stage, taxon, female age, mass and identity (73 females), egg-laying sequence (105 clutches) and the study zone. Our results are consistent with the prediction that egg membranes are thicker in fully developed eggs than in non-developed eggs, suggesting that the increase in membrane thickness during development may compensate for eggshell thinning. In addition, our data shown that thicker membranes are associated with larger, heavier and relatively wider eggs, as well as with eggs that had thinner eggshells. Egg-laying sequence, female age and the study zone did not explain the observed variation of membrane thickness in the falcon taxa studied. As we provide quantitative data on membrane thickness variation during development in falcons not subjected to contamination or food limitation (i.e. bred under captive conditions), our data may be used as a reference for studies on eggs from natural populations. Considering the large variation in membrane thickness and the multiple factors affecting on it and its importance in the protection of the embryo, we encourage other researchers to include measurements on membranes in studies exploring eggshell thickness variation.

The liver but not the skin is the site for conversion of a red carotenoid in a passerine bird.

Carotenoids may provide numerous health benefits and are also responsible for the integumentary coloration of many bird species. Despite their importance, many aspects of their metabolism are still poorly known, and even basic issues such as the anatomical sites of conversion remain controversial. Recent studies suggest that the transformation of carotenoid pigments takes place directly in the follicles during feather growth, even though the liver has been previously recognised as a storing organ for these pigments with a certain potential for conversion. In this context, we analysed the carotenoid profile of plasma, liver, skin and feathers of male Common Crossbills (Loxia curvirostra). Interestingly, the derivative feather pigment 3-hydroxy-echinenone was detected in the liver and in the bloodstream (i.e. the necessary vehicle to transport metabolites to colorful peripheral tissues). Our results demonstrate for the first time with empirical data that the liver may act as the main site for the synthesis of integumentary carotenoids. This finding contradicts previous assumptions and raises the question of possible inter-specific differences in the site of carotenoid conversion in birds.

Determinants of eggshell strength in endangered raptors.

We compared eggshell strength in a group of falcon taxa including the peregrine falcon (Falco peregrinus peregrinus), the red shaheen falcon (F. peregrinus babylonicus), the saker falcon (F. cherrug), the gyr falcon (F. rusticolus) and some interspecific and intraspecific hybrids. Our results showed that smaller falcons (<1,000 g) of the peregrine group have eggshells that are significantly softer (x=13.3 N) and thinner (x=0.26 mm) (n=107 eggs) than larger falcons (>1,000 g) of the gyr-saker group (x=20.8 N and 0.39 mm, respectively, n=81 eggs). We found a significant positive correlation between egg hardness and eggshell thickness. Linear mixed models showed that clutches from heavier females consisted of larger and harder eggs with thicker shells and thicker egg membranes. Eggs produced by older females and eggs laid later in the laying sequence were relatively smaller and softer and had relatively thin egg membranes and eggshells. Individual females, irrespective of their age, contributed significantly to the observed variation in egg strength. Egg size and hardness of hybrid eggs were similar to that of the pure species suggesting that hybridization does not affect eggshell hardness or thickness. Our study provides quantitative evidence of several factors, other than levels of contamination, which may affect eggshell thickness and hardness in falcons.