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Mutualistic symbioses between animals and bacteria depend on acquisition of appropriate symbionts while avoiding exploitation by non-beneficial microbes. The mode of acquisition of symbionts would determine, not only the probability of encountering but also evolutionary outcomes of mutualistic counterparts. The microbiome inhabiting the uropygial gland of the European hoopoe (Upupa epops) includes a variety of bacterial strains, some of them providing antimicrobial benefits. Here, the mode of acquisition and stability of this microbiome is analyzed by means of Automated rRNA Intergenic Spacer Analysis and two different experiments. The first experiment impeded mothers' access to their glands, thus avoiding direct transmission of microorganisms from female to offspring secretions. The second experiment explored the stability of the microbiomes by inoculating glands with secretions from alien nests. The first experiment provoked a reduction in similarity of microbiomes of mother and nestlings. Interestingly, some bacterial strains were more often detected when females had not access to their glands, suggesting antagonistic effects among bacteria from different sources. The second experiment caused an increase in richness of the microbiome of receivers in terms of prevalence of Operational Taxonomic Units (OTUs) that reduced differences in microbiomes of donors and receivers. That occurred because OTUs that were present in donors but not in receivers incorporated to the microbiome of the latter, which provoked that cross-inoculated nestlings got similar final microbiomes that included the most prevalent OTUs. The results are therefore consistent with a central role of vertical transmission in bacterial acquisition by nestling hoopoes and support the idea that the typical composition of the hoopoe gland microbiome is reached by the incorporation of some bacteria during the nestling period. This scenario suggests the existence of a coevolved core microbiome composed by a mix of specialized vertically transmitted strains and facultative symbionts able to coexist with them. The implications of this mixed mode of transmission for the evolution of the mutualism are discussed.

In this work we investigated the between-colony spatial, behavioural and trophic segregation of two sub-populations of the elusive Macaronesian shearwaters Puffinus baroli breeding only ~340 km apart in Cima Islet (Porto Santo Island) and Selvagem Grande Island. Global location sensing (gls) loggers were used in combination with the trophic ecology of tracked individuals, inferred from the isotopic signatures of wing feathers. Results suggest that these two Macaronesian shearwater sub-populations do segregate during the non-breeding period in some 'sub-population-specific' regions, by responding to different oceanographic characteristics (habitat modelling). Within these disparate areas, both sub-populations behave differently (at-sea activity) and prey on disparate trophic niches (stable isotope analysis). One hypothesis would be that each sub-population have evolved and adapted to feed on particular and 'sub-population-specific' resources, and the segregation observed at the three different levels (spatial, behavioural and trophic) might be in fact a result of such adaptation, from the emergence of 'cultural foraging patterns'. Finally, when comparing to the results of former studies reporting on the spatial, behavioural and trophic choices of Macaronesian shearwater populations breeding on Azores and Canary Islands, we realized the high ecological plasticity of this species inhabiting and foraging over the North-East Atlantic Ocean.

Diet studies present numerous methodological challenges. We evaluated the usefulness of commercially available trail-cameras for analyzing the diet of Northern Goshawks (Accipiter gentilis) as a model for nesting raptors during the period 2007-2011. We compared diet estimates obtained by direct camera monitoring of 80 nests with four indirect analyses of prey remains collected from the nests and surroundings (pellets, bones, feather-and-hair remains, and feather-hair-and-bone remains combined). In addition, we evaluated the performance of the trail-cameras and whether camera monitoring affected Goshawk behavior. The sensitivity of each diet-analysis method depended on prey size and taxonomic group, with no method providing unbiased estimates for all prey sizes and types. The cameras registered the greatest number of prey items and were probably the least biased method for estimating diet composition. Nevertheless this direct method yielded the largest proportion of prey unidentified to species level, and it underestimated small prey. Our trail-camera system was able to operate without maintenance for longer periods than what has been reported in previous studies with other types of cameras. Initially Goshawks showed distrust toward the cameras but they usually became habituated to its presence within 1-2 days. The habituation period was shorter for breeding pairs that had previous experience with cameras. Using trail-cameras to monitor prey provisioning to nests is an effective tool for studying the diet of nesting raptors. However, the technique is limited by technical failures and difficulties in identifying certain prey types. Our study also shows that cameras can alter adult Goshawk behavior, an aspect that must be controlled to minimize potential negative impacts.

Predation is one of the main causes of adult mortality and breeding failure for ground-nesting birds. Micro-habitat structure around nests plays a critical role in minimizing predation risk. Plovers nest in sites with little vegetation cover to maximize the incubating adult visibility, but many studies suggest a trade-off between nest-crypsis and predator detection strategies. However, this trade-off has not been explored in detail because methods used so far do not allow estimating the visibility with regards to critical factors such as slope or plant permeability to vision. Here, we tested the hypothesis that Kentish plovers select exposed sites according to a predator detection strategy, and the hypothesis that more concealed nests survive longer according to a crypsis strategy. To this end, we obtained an accurate estimation of the incubating adult's field of vision through a custom built inverted periscope. Our results showed that plovers selected nest sites with higher visibility than control points randomly selected with regards to humans and dogs, although nests located in sites with higher vegetation cover survived longer. In addition, the flushing distance (i.e., the distance at which incubating adults leave the nest when they detect a potential predator) decreased with vegetation cover. Consequently, the advantages of concealing the nest were limited by the ability to detect predators, thus indirectly supporting the existence of the trade-off between crypsis and predator detection. Finally, human disturbance also constrained nest choice, forcing plovers to move to inland sites that were less suitable because of higher vegetation cover, and modulated flushing behavior, since plovers that were habituated to humans left their nests closer to potential predators. This constraint on the width of suitable breeding habitat is particularly relevant for the conservation of Kentish Plover in sand beaches, especially under the current context of coastal regression and increase of recreational activities.

Understanding biodiversity distribution is a primary goal of community ecology. At a landscape scale, bee communities are affected by habitat composition, anthropogenic land use, and fragmentation. However, little information is available on local-scale spatial distribution of bee communities within habitats that are uniform at the landscape scale. We studied a bee community along with floral and nesting resources over a 32 km2 area of uninterrupted Mediterranean scrubland. Our objectives were (i) to analyze floral and nesting resource composition at the habitat scale. We ask whether these resources follow a geographical pattern across the scrubland at bee-foraging relevant distances; (ii) to analyze the distribution of bee composition across the scrubland. Bees being highly mobile organisms, we ask whether bee composition shows a homogeneous distribution or else varies spatially. If so, we ask whether this variation is irregular or follows a geographical pattern and whether bees respond primarily to flower or to nesting resources; and (iii) to establish whether body size influences the response to local resource availability and ultimately spatial distribution. We obtained 6580 specimens belonging to 98 species. Despite bee mobility and the absence of environmental barriers, our bee community shows a clear geographical pattern. This pattern is mostly attributable to heterogeneous distribution of small (<55 mg) species (with presumed smaller foraging ranges), and is mostly explained by flower resources rather than nesting substrates. Even then, a large proportion (54.8%) of spatial variability remains unexplained by flower or nesting resources. We conclude that bee communities are strongly conditioned by local effects and may exhibit spatial heterogeneity patterns at a scale as low as 500-1000 m in patches of homogeneous habitat. These results have important implications for local pollination dynamics and spatial variation of plant-pollinator networks.

Background: We describe the first occurrence in the fossil record of an aquatic avian twig-nest with five eggs in situ (Early Miocene Tudela Formation, Ebro Basin, Spain). Extensive outcrops of this formation reveal autochthonous avian osteological and oological fossils that represent a single taxon identified as a basal phoenicopterid. Although the eggshell structure is definitively phoenicopterid, the characteristics of both the nest and the eggs are similar to those of modern grebes. These observations allow us to address the origin of the disparities between the sister taxa Podicipedidae and Phoenicopteridae crown clades, and traces the evolution of the nesting and reproductive environments for phoenicopteriforms., Methodology/principal Findings: Multi-disciplinary analyses performed on fossilized vegetation and eggshells from the eggs in the nest and its embedding sediments indicate that this new phoenicopterid thrived under a semi-arid climate in an oligohaline (seasonally mesohaline) shallow endorheic lacustine environment. High-end microcharacterizations including SEM, TEM, and EBSD techniques were pivotal to identifying these phoenicopterid eggshells. Anatomical comparisons of the fossil bones with those of Phoenicopteriformes and Podicipediformes crown clades and extinct palaelodids confirm that this avian fossil assemblage belongs to a new and basal phoenicopterid., Conclusions/significance: Although the Podicipediformes-Phoenicopteriformes sister group relationship is now well supported, flamingos and grebes exhibit feeding, reproductive, and nesting strategies that diverge significantly. Our multi-disciplinary study is the first to reveal that the phoenicopteriform reproductive behaviour, nesting ecology and nest characteristics derived from grebe-like type strategies to reach the extremely specialized conditions observed in modern flamingo crown groups. Furthermore, our study enables us to map ecological and reproductive characters on the Phoenicopteriformes evolutionary lineage. Our results demonstrate that the nesting paleoenvironments of flamingos were closely linked to the unique ecology of this locality, which is a direct result of special climatic (high evaporitic regime) and geological (fault system) conditions.

Theoretical models aimed at explaining the evolution of honest, informative begging signals employed by nestling birds to solicit food from their parents, require that dishonest signalers incur a net viability cost in order to prevent runaway escalation of signal intensity over evolutionary time. Previous attempts to determine such a cost empirically have identified two candidate physiological costs associated with exaggerated begging: a growth and an immunological cost. However, they failed to take into account the fact that those costs are potentially offset by the fact that nestlings that invest more in begging are also likely to obtain more food. In this study, we test experimentally whether a 25% increase in ingested food compensates for growth and immunological costs of extra begging in southern shrike (Lanius meridionalis) nestlings. Three nestmates matched by size were given three treatments: low begging, high begging-same food intake, and high begging-extra food intake. We found that, while a higher food intake did effectively compensate for the growth cost, it failed to compensate for the immunological cost, measured as T-cell mediated immune response against an innocuous mitogen. Thus, we show for the first time that escalated begging has an associated physiological net cost likely to affect nestling survival negatively.

The presence of conspecifics can strongly modulate the quality of a breeding site. Both positive and negative effects of conspecifics can act on the same individuals, with the final balance between its costs and benefits depending on individual characteristics. A particular case of inter-individual variation found in many avian species is chromatic variability. Among birds, plumage coloration can co-vary with morphology, physiology and behavior as well as with age. These relationships suggest that cost-benefit balances of conspecific presence may be different for individuals with different colorations. We investigated whether inter-individual variability affects population regulation and expansion processes by analyzing potential differences in density-dependent productivity and settlement patterns in relation to plumage coloration in a population of a long-lived avian species recently undergoing a notable increase in numbers and distribution range. Our results show strong variation in the effect of density on productivity of breeding pairs depending on plumage coloration of their members. Productivity of dark birds decreased along the breeding density gradient while that of lighter breeders remained unchanged with conspecific density. In a similar way, our results showed an uneven occupation of localities by individuals with different plumage coloration in relation to local densities, with the breeding of lighter harriers more aggregated than that of dark-brown ones. At a population scale, darker birds had higher probability of colonization of the most isolated, empty sites. Explanations for species range expansion and population regulation usually make the inferred assumption that species traits are similar among individuals. However, in most species, there could be individual variation in niche requirements or dispersal propensities among individuals with different traits. Our results contribute to the growing appreciation that the individual traits, but not the average trait at the level of species, are important during population regulation and expansion processes.

Background: Megaloolithid eggs have long been associated with sauropod dinosaurs. Despite their extensive and worldwide fossil record, interpretations of egg size and shape, clutch morphology, and incubation strategy vary. The Pinyes locality in the Upper Cretaceous Tremp Formation in the southern Pyrenees, Catalonia provides new information for addressing these issues. Nine horizons containing Megaloolithus siruguei clutches are exposed near the village of Coll de Nargó. Tectonic deformation in the study area strongly influenced egg size and shape, which could potentially lead to misinterpretation of reproductive biology if 2D and 3D maps are not corrected for bed dip that results from tectonism., Methodology/findings: Detailed taphonomic study and three-dimensional modelling of fossil eggs show that intact M. siruguei clutches contained 20-28 eggs, which is substantially larger than commonly reported from Europe and India. Linear and grouped eggs occur in three superimposed levels and form an asymmetric, elongate, bowl-shaped profile in lateral view. Computed tomography data support previous interpretations that the eggs hatched within the substrate. Megaloolithid clutch sizes reported from other European and Indian localities are typically less than 15 eggs; however, these clutches often include linear or grouped eggs that resemble those of the larger Pinyes clutches and may reflect preservation of incomplete clutches., Conclusions/significance: We propose that 25 eggs represent a typical megaloolithid clutch size and smaller egg clusters that display linear or grouped egg arrangements reported at Pinyes and other localities may represent eroded remnants of larger clutches. The similarity of megaloolithid clutch morphology from localities worldwide strongly suggests common reproductive behaviour. The distinct clutch geometry at Pinyes and other localities likely resulted from the asymmetrical, inclined, and laterally compressed titanosaur pes unguals of the female, using the hind foot for scratch-digging during nest excavation.

Visual signals are crucial for parent-offspring communication, although their functioning has been neglected for nocturnal birds. Here, we investigated parental preference for nestling coloration in nocturnal conditions--a question hitherto unexplored--in a nocturnal raptor, the scops owl (Otus scops). We assessed how parents allocated food during the night in relation to a manipulation of ultraviolet (UV) reflectance of the cere (skin above the beak) of their offspring. Reflectance of the cere shows a marked peak in the UV part of the spectrum, and location of the UV peak is related to nestling body mass (i.e. heavier nestlings have a UV peak at lower wavelengths). We found evidence of parental bias in favour of lighter offspring: UV-reduced nestlings gained more weight during the night than their control siblings. This study provides the first experimental evidence of the use of visual cues for parent-offspring communication in a nocturnal bird.

In many litter-bearing mammals and in a few viviparous reptiles the sex ratio of the entire brood or the sex of the adjacent fetuses induces sex-specific differences in the hatchling's phenotype. This study examines whether the sex of incubation neighbours affects hatchling characteristics in oviparous common lizards (Lacerta vivipara). Oviparous common lizards lay eggs with thin eggshells and, therefore, are an optimal model organism for studying the effects of hormone leakage among developing embryos since the strongest evidence for prenatal sex ratio effects on offspring development comes from viviparous populations of the same species. Groups of three eggs were incubated together and were categorised according to the sex of the resulting hatchlings as either homosex (three hatchlings of the same sex) or heterosex (one male or one female hatchling plus two siblings of the opposite sex). Hatchlings incubated adjacent to siblings of the same sex had larger body mass and body condition. Males tended to have lower ventral scale counts when incubated with other males. Conversely, females tended to have more ventral scales when incubated with other females, indicative of a more feminised phenotype. There was also a significant interaction between hatchling sex and incubation environment with respect to the length of the fourth digit of the hindlimb, likely indicative of masculinisation in heterosex females. This study suggests steroid diffusion between adjacent eggs in a minimally manipulative experiment and provides the first evidence for developmental effects of the exogenous hormonal environment in near natural conditions in an oviparous amniote. Implications of these results for the evolution of within-clutch sex ratio are discussed.

Compared to non-flying nest-dwelling ectoparasites, the biology of most species of flying ectoparasites and its potential impact on avian hosts is poorly known and rarely, if ever, reported. In this study we explore for the first time the factors that may affect biting midge (Diptera: Ceratopogonidae) and black fly (Diptera: Simuliidae) abundances in the nest cavity of a bird, the hole-nesting blue tit Cyanistes caeruleus, and report their effects on adults and nestlings during reproduction. The abundance of biting midges was positively associated with nest mass, parental provisioning effort and abundance of blowflies and black flies, while negatively associated with nestling condition. Furthermore, a medication treatment to reduce blood parasitaemias in adult birds revealed that biting midges were more abundant in nests of females whose blood parasitaemias were experimentally reduced. This finding would be in accordance with these insect vectors attacking preferentially uninfected or less infected hosts to increase their own survival. The abundance of black flies in the population was lower than that of biting midges and increased in nests with later hatching dates. No significant effect of black fly abundance on adult or nestling condition was detected. Blood-sucking flying insects may impose specific, particular selection pressures on their hosts and more research is needed to better understand these host-parasite associations.

Background: Departures from power law group size frequency distributions have been proposed as a useful tool to link individual behavior with population patterns and dynamics, although examples are scarce for wild animal populations., Methodology/principal Findings: We studied a population of Lesser kestrels (Falco naumanni) breeding in groups (colonies) from one to ca. 40 breeding pairs in 10,000 km(2) in NE Spain. A 3.5 fold steady population increase occurred during the eight-year study period, accompanied by a geographical expansion from an initial subpopulation which in turn remained stable in numbers. This population instability was mainly driven by first-breeders, which are less competitive at breeding sites, being relegated to breed solitarily or in small colony sizes, and disperse farther than adults. Colony size frequency distributions shifted from an initial power law to a truncated power law mirroring population increase. Thus, we hypothesized that population instability was behind the truncation of the power law. Accordingly, we found a power law distribution through years in the initial subpopulation, and a match between the power law breakpoint (at ca. ten pairs) and those colony sizes from which the despotic behavior of colony owners started to impair the settlement of newcomers. Moreover, the instability hypothesis was further supported by snapshot data from another population of Lesser kestrels in SW Spain suffering a population decline., Conclusions/significance: Appropriate analysis of the scaling properties of grouping patterns has unraveled the link between local agonistic processes and large-scale (population) grouping patterns in a wild bird population.

We investigated how the expression of carotenoid-based plumage coloration (lightness and chroma) in nestling blue tits Parus caeruleus is associated with forest structure in oak forests of central Spain. We found evidence of a reduced expression of carotenoid-based coloration in nestlings growing up in successionally young and structurally simple forest territories. Our results suggest that breast feather coloration can be used as an indicator of nestling quality because nestlings with more intense yellow plumage coloration had larger body size and stronger immune responses to the injection of phytohemagglutinin (PHA). Given the association of forest structural complexity with carotenoid-based plumage coloration, our findings suggest that variation in habitat structure may have a significant impact on forest birds in their first stages of life which has implications for forest management practices.

Adjustment of offspring sex ratios should be favoured by natural selection when parents are capable of facultatively altering brood sex ratios and of recognizing the circumstances that predict the probable fitness benefit of producing sons and daughters. Although experimental studies have shown that female birds may adjust offspring sex ratios in response to changes in their own condition and in the external appearance of their mate, and male attributes other than his external morphology are also thought to act as signals of male quality, it is not known whether females will respond to changes in such signals, in the absence of any change in the appearance of the male himself. Here, we experimentally manipulated a male courtship display, the green plants carried to the nest by male spotless starlings (Sturnus unicolor), without changing any physical attributes of the male himself, and examined whether this influenced female decisions on offspring sex ratio. We found that in an environment in which female starlings were producing more daughters than sons, experimental enhancement of the green nesting material caused females to significantly increase the number of male eggs produced and thereby removed the female bias. This effect was consistent in 2 years and at two localities. This demonstrates that the green material, whose function has long puzzled biologists, conveys important information to the female and that she facultatively adjusts offspring production accordingly.