Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

Age and breeding success related to nest position in a White stork Ciconia ciconia colony

Coloniality is a breeding system that may produce benefits in terms of breeding success, although these advantages could vary according to factors such as colony size or nest position. We studied breeder's age in relation to nest position (peripheral or central) within the colony. In addition, we studied the relationship between breeding success and nest position, controlling for breeder's age, a highly correlated factor, in a White Stork Ciconia ciconia colony over a 7-year period. Our results show that central nests are mainly occupied by adult birds and had lower failure rates. However, controlling for breeder's age, nest position per se did not explain breeding success. The scarce predation and the lack of human disturbance in the study colony could explain the absence of differences in breeding success between different nest positions within the colony.     

Factors associated with nest survival of Black‐throated Sparrows,desert‐breeding nest‐site generalists

Black‐throated Sparrows (Amphispiza bilineata) are common breeding birds throughout the desert regions of North America and can be considered nest‐site generalists. Information about how spatial (e.g., vegetation) and temporal factors influence nest survival of these sparrows is lacking throughout their range. Our objective was to examine the spatial and temporal factors associated with nest survival of Black‐throated Sparrows at the nest and nest‐patch scales in the predator‐rich environment of the northern Chihuahuan Desert of New Mexico. We used a logistic‐exposure model fit within a Bayesian framework to model the daily survival probability of Black‐throated Sparrow nests. Predation was the leading cause of nest failure, accounting for 86% of failed nests. We found evidence of negative associations between nest survival and both vegetative cover above nests and shrub density within 5 m of nests. We found no support for other habitat covariates, but did find strong evidence that daily survival rate was higher earlier in the breeding season and during the egg‐laying stage. A decline in nest survival later in the breeding period may be due to increased predator activity due to warmer ambient temperatures, whereas lower survival during the incubation and nestling stages could be a result of increased activity at nests. A generalist approach to nest‐site selection may be an adaptive response to the presence of a diverse assemblage of nest predators that results in the reduced influence of spatial factors on nest survival for Black‐throated Sparrows.     

Effects of temperature and nest heat exposure on nestling growth,dehydration and survival in a Mediterranean hole‐nesting passerine

Variable environments impose constraints on adaptation by modifying selection gradients unpredictably. Optimal bird development requires an adequate thermal range, outside which temperatures can alter nestling physiology, condition and survival. We studied the effect of temperature and nest heat exposure on the reproductive success of a population of double‐brooded Spotless Starlings Sturnus unicolor breeding in a nestbox colony in central Spain with a marked intra‐seasonal variation in temperature. We assessed whether the effect of temperature differed between first and second broods, thus constraining optimal nest‐site choice. Ambient temperature changed greatly during the chick‐rearing period and had a strong influence on nestling mass and all body size measures we recorded, although patterns of clutch size or nestling mortality were not influenced. This effect differed between first and second broods: nestlings were found to have longer wings and bills with increasing temperature in first broods, whereas the effect was the opposite in second broods. Ambient temperature was not related to nestling body mass or tarsus‐length in first broods, but in second broods, nestlings were lighter and had smaller tarsi with higher ambient temperatures. The exposure of nestboxes to heat influenced nestling morphology: heat exposure index was negatively related to nestling body mass and wing‐length in second broods, but not in first broods. Furthermore, there was a positive relationship between nest heat exposure and nestling dehydration. Our results suggest that optimal nest choice is constrained by varying environmental conditions in birds breeding over prolonged periods, and that there should be selection for parents to switch from sun‐exposed to sun‐protected nest‐sites as the season progresses. However, nest‐site availability and competition for sites are likely to impose constraints on this choice.     

Breeding habitat and nest‐site selection by an obligatory “nest‐cleptoparasite”, the Amur Falcon Falco amurensis

The selection of a nest site is crucial for successful reproduction of birds. Animals which re‐use or occupy nest sites constructed by other species often have limited choice. Little is known about the criteria of nest‐stealing species to choose suitable nesting sites and habitats. Here, we analyze breeding‐site selection of an obligatory “nest‐cleptoparasite”, the Amur Falcon Falco amurensis. We collected data on nest sites at Muraviovka Park in the Russian Far East, where the species breeds exclusively in nests of the Eurasian Magpie Pica pica. We sampled 117 Eurasian Magpie nests, 38 of which were occupied by Amur Falcons. Nest‐specific variables were assessed, and a recently developed habitat classification map was used to derive landscape metrics. We found that Amur Falcons chose a wide range of nesting sites, but significantly preferred nests with a domed roof. Breeding pairs of Eurasian Hobby Falco subbuteo and Eurasian Magpie were often found to breed near the nest in about the same distance as neighboring Amur Falcon pairs. Additionally, the occurrence of the species was positively associated with bare soil cover, forest cover, and shrub patches within their home range and negatively with the distance to wetlands. Areas of wetlands and fallow land might be used for foraging since Amur Falcons mostly depend on an insect diet. Additionally, we found that rarely burned habitats were preferred. Overall, the effect of landscape variables on the choice of actual nest sites appeared to be rather small. We used different classification methods to predict the probability of occurrence, of which the Random forest method showed the highest accuracy. The areas determined as suitable habitat showed a high concordance with the actual nest locations. We conclude that Amur Falcons prefer to occupy newly built (domed) nests to ensure high nest quality, as well as nests surrounded by available feeding habitats.     

Equal reproductive success of phenotypes in the Larus glaucescens–occidentalis complex

Glaucous‐winged gulls Larus glaucescens and western gulls L. occidentalis hybridize extensively where their ranges overlap along the coasts of Washington and Oregon, producing a continuum of phenotypic intergrades between the two parental species. This zone often is considered an example of geographically bounded hybrid superiority, but studies of relative success among parental types and hybrids have not provided consistent support for this model. We tested the predictions of the dynamic‐equilibrium and geographically bounded hybrid superiority hypotheses by studying mate choice and reproductive success among gulls on Protection Island, Washington, the largest breeding colony of glaucous‐winged/western gulls within the hybrid zone. The dynamic‐equilibrium hypothesis posits that hybridization due to dispersal balances selection against less fit hybrids and assortative mating is adaptive. Geographically bounded hybrid superiority posits that hybrids are better fit than parental types within an ecotone between the environments to which the parental species are adapted, and a preference for hybrid mates is adaptive. Additionally, we investigated whether hatching success and nest site choice are correlated for Protection Island gulls. We assigned a hybrid index to each sample bird by examining plumage melanism and bare part coloration in the field. Sheltered nests contained larger clutches and exhibited increased hatching success, but choice of nest habitat was not associated with hybrid index. Western gull‐like pairs produced smaller third eggs; however, hybrid index was not correlated with clutch size or hatching success. Protection Island gulls did exhibit assortative mating. In short, we did not find strong support for either geographically bounded hybrid superiority or the dynamic‐equilibrium hypothesis.     

Mechanisms of colony selection by first‐year Lesser Kestrels Falco naumanni

In colonial species, first‐time breeders may use the number of settled conspecifics in colony selection, but such a relationship is confused by the correlation between colony size and nest‐site availability. To distinguish conspecific attraction from neutral colony selection, we experimentally increased nest availability for first‐year Lesser Kestrels Falco naumanni, allowing us to dissociate the number of vacant nest‐sites from colony size at the arrival time of first‐year birds. Under natural conditions, the number of first‐year birds settling was positively correlated with both the number of philopatric and the total number of breeding pairs (colony size) already settled. However, the probability of occupation of experimentally manipulated nests by first‐year birds was independent of colony size. In experimental colonies, the number of first‐year birds settling was positively correlated with the number of manipulated nest‐sites but not with the number of conspecifics. Overall, these results support a neutral colony selection by first‐year Lesser Kestrels based on nest‐site availability.     

Model of burrow selection predicts pattern of burrow switching by Leach's Storm‐Petrels

Patterns of nest site selection exhibited at the scale of a population should result from initial preferences of individuals occupying nest sites as well as preferences exhibited by individuals moving between nest sites. We tested whether nest‐site preferences measured at the population scale were predictive of patterns of burrow switching by Leach's Storm‐Petrels (Oceanodroma leucorhoa), a long‐lived seabird that nests in underground burrows. Breeding pairs generally choose from the pool of available existing burrows rather than constructing new burrows, and a portion of the burrows in a colony remains unused in any breeding season. We quantified burrow preference at a colony on Kent Island, New Brunswick, over four breeding seasons. We used a classification and regression tree analysis to build a predictive model of nest‐site selection. Preferentially occupied burrows were drier, longer, had larger nest chambers, and were in areas of higher burrow density. To measure preferences during burrow switching, we tracked individuals that switched burrows, comparing characteristics of the burrows in which these birds were originally found to those they inhabited at the end of the study period. Characteristics preferred by switching individuals were a subset of those observed at the scale of the population; individuals moved to burrows that were drier, longer, and had larger nest chambers. Our results show how preferences of individuals that move between nest sites contribute to nest site preferences exhibited at the population scales commonly tested.     

Safety first: terrestrial predators drive selection of highly specific nesting sites in colonial‐breeding birds

Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.     

Delayed egg‐laying and shortened incubation duration of Arctic‐breeding shorebirds coincide with climate cooling

Biological impacts of climate change are exemplified by shifts in phenology. As the timing of breeding advances, the within‐season relationships between timing of breeding and reproductive traits may change and cause long‐term changes in the population mean value of reproductive traits. We investigated long‐term changes in the timing of breeding and within‐season patterns of clutch size, egg volume, incubation duration, and daily nest survival of three shorebird species between two decades. Based on previously known within‐season patterns and assuming a warming trend, we hypothesized that the timing of clutch initiation would advance between decades and would be coupled with increases in mean clutch size, egg volume, and daily nest survival rate. We monitored 1,378 nests of western sandpipers, semipalmated sandpipers, and red‐necked phalaropes at a subarctic site during 1993–1996 and 2010–2014. Sandpipers have biparental incubation, whereas phalaropes have uniparental incubation. We found an unexpected long‐term cooling trend during the early part of the breeding season. Three species delayed clutch initiation by 5 days in the 2010s relative to the 1990s. Clutch size and daily nest survival showed strong within‐season declines in sandpipers, but not in phalaropes. Egg volume showed strong within‐season declines in one species of sandpiper, but increased in phalaropes. Despite the within‐season patterns in traits and shifts in phenology, clutch size, egg volume, and daily nest survival were similar between decades. In contrast, incubation duration did not show within‐season variation, but decreased by 2 days in sandpipers and increased by 2 days in phalaropes. Shorebirds demonstrated variable breeding phenology and incubation duration in relation to climate cooling, but little change in nonphenological components of traits. Our results indicate that the breeding phenology of shorebirds is closely associated with the temperature conditions on breeding ground, the effects of which can vary among reproductive traits and among sympatric species.     

Use of novel nest boxes by carmine bee‐eaters (Merops nubicus) in captivity

Carmine bee‐eaters make attractive additions to zoo aviaries but breeding programs have had challenges and limited success. The objectives of this study were to document nesting behavior of Carmine bee‐eaters in a captive setting and compare reproductive success between a novel nest box (plastic, 17 × 30 × 22 cm) and a PVC pipe model used previously (30 cm long, 8 cm in diameter). Three bee‐eater pairs were given access to seven nest chambers (six novel boxes, one PVC model). Behavioral observations occurred during a 15‐min period in the morning or afternoon before egg production and continued until chicks fledged for a total of 87 observation periods (21.75 hr). All occurrences by an individual bird entering or exiting a nest tunnel, food provision, and the time (min) spent inside a nest cavity were documented. Additionally, daily temperature within each nest chamber was recorded. Before eggs were produced the average daily temperature (23.02°C) within the nest chambers did not differ, suggesting that nest cavity choice was not influenced by temperature. No differences were detected among pairs in percent of observed time spent inside their nest cavities or number of times a nest tunnel was entered during the incubation or fledging periods. During incubation females spent a greater percent of observed time inside the nest cavity than males (P=0.02). During the fledging period food provision did not differ between the pairs, however males entered their nest tunnels more often per hour than females (P=0.03), and males tended to provide food more often than females (P=0.053). Two pairs nested in novel nest boxes and successfully fledged one chick each. The pair that nested in the PVC model did not fledge a chick. A nest box that aids in keeping eggs intact is essential for breeding bee‐eaters in captivity, and maintaining captive populations will provide opportunities for zoo visitors to enjoy these birds and will reduce the need to remove birds from the wild. Zoo Biol 0:1–13, 2007. © 2007 Wiley‐Liss, Inc.     

Within‐season variation in sexual selection in a fish with dynamic sex roles

The strength of sexual selection may vary between species, among populations and within populations over time. While there is growing evidence that sexual selection may vary between years, less is known about variation in sexual selection within a season. Here, we investigate within‐season variation in sexual selection in male two‐spotted gobies (Gobiusculus flavescens). This marine fish experiences a seasonal change in the operational sex ratio from male‐ to female‐biased, resulting in a dramatic decrease in male mating competition over the breeding season. We therefore expected stronger sexual selection on males early in the season. We sampled nests and nest‐holding males early and late in the breeding season and used microsatellite markers to determine male mating and reproductive success. We first analysed sexual selection associated with the acquisition of nests by comparing nest‐holding males to population samples. Among nest‐holders, we calculated the potential strength of sexual selection and selection on phenotypic traits. We found remarkable within‐season variation in sexual selection. Selection on male body size related to nest acquisition changed from positive to negative over the season. The opportunity for sexual selection among nest‐holders was significantly greater early in the season rather than late in the season, partly due to more unmated males. Overall, our study documents a within‐season change in sexual selection that corresponds with a predictable change in the operational sex ratio. We suggest that many species may experience within‐season changes in sexual selection and that such dynamics are important for understanding how sexual selection operates in the wild.     

Honey Buzzard Pernis apivorus nest‐site selection in relation to habitat and the distribution of Goshawks Accipiter gentilis

The selection of a suitable nest‐site is critical for successful reproduction. Species' preferences for nest‐sites have presumably evolved in relation to local habitat resources and/or interactions with other species. The importance of these two components in the nest‐site selection of the Eurasian Honey Buzzard Pernis apivorus was assessed in two study areas in eastern Austria. There was almost no difference in macro‐ and micro‐habitat features between nest‐sites and random plots, suggesting that Honey Buzzards did not base their choice of nest‐site on habitat characteristics. However, nests were placed significantly further from nests of Northern Goshawk Accipiter gentilis than would be expected if nest‐sites had been chosen at random. Furthermore, in one study area Honey Buzzards appeared to favour areas close to human settlements, perhaps indicating a mechanism to avoid Goshawks, which tend to avoid the proximity of humans. No habitat variable was significantly associated with the loss of Honey Buzzard young, but predation was higher in territories closer to breeding pairs of Goshawks at both study sites. Although Honey Buzzards are restricted to nesting in forests, their choice of nest‐site therefore appears to be largely dictated by the distribution of predators. Studies of habitat association may yield misleading results if the effects of predation risk on distribution are not considered.     

Nest survival is influenced by parental behaviour and heterospecifics in a mixed‐species colony

Studies of avian nest success often focus on examining influences of variation in environmental and seasonal factors. However, in‐depth evaluations can also incorporate variation in individual incubation behaviour to further advance our understanding of avian reproductive ecology. We examined these relationships in colonially nesting Black‐crowned Night‐Herons Nycticorax nycticorax using intensive video‐monitoring methods to quantify incubation behaviours. We modelled nest survival as a function of both extrinsic factors and incubation behaviours over a 3‐year period (2010–12) on Alcatraz Island, USA. Model‐averaged parameter estimates indicated that nest survival increased as a function of greater incubation constancy (% of time spent incubating eggs within a 24‐h period), and average daily precipitation throughout the nesting stage. Common Ravens Corvus corax are the only known nest predator of Night‐Herons on Alcatraz Island, as on many other coastal Pacific islands. We also investigated the effects of heterospecific nesting of California Gulls Larus californicus and Western Gulls Larus occidentalis in a mixed‐species colony with Night‐Herons, based on nesting proximity data collected over a 2‐year period (2011–12). This second analysis indicated that, in addition to incubation behaviours, nesting heterospecifics are an important factor for explaining variation in Night‐Heron nest survival. However, contrary to our original expectation, we found that Night‐Herons experienced increased nest survival with increasing distance from gull colony boundaries. These results may apply to other areas with multiple colonial nesting species and similar predator communities and climatic patterns.     

Fine‐scale bird monitoring from light unmanned aircraft systems

Unmanned aircraft systems (UAS) are remote‐controlled devices capable of collecting information from difficult‐to‐access places while minimizing disturbance. Although UAS are increasingly used in many research disciplines, their application to wildlife research remains to be explored in depth. Here, we report on the use of a small UAS to monitor temporal changes in breeding population size in a Black‐headed Gull Chroicocephalus ridibundus colony. This method makes it possible to obtain georeferenced data on nest locations without causing colony disturbance, which would not otherwise be possible via direct ground observations.     

Nest‐site selection by Slender‐billed Parakeets in a Chilean agricultural‐forest mosaic

ABSTRACT Species in the family Psittacidae may be particularly vulnerable to anthropogenic habitat transformations that reduce availability of suitable breeding sites at different spatial scales. In southern Chile, loss of native forest cover due to agricultural conversion may impact populations of Slender‐billed Parakeets (Enicognathus leptorhynchus), endemic secondary cavity‐nesting psittacids. Our objective was to assess nest‐site selection by Slender‐billed Parakeets in an agricultural‐forest mosaic of southern Chile at two spatial scales: nest trees and the habitat surrounding those trees. During the 2008–2009 breeding seasons, we identified nest sites (N= 31) by observing parakeet behavior and using information provided by local residents. Most (29/31) nests were in mature Nothofagus obliqua trees. By comparing trees used for nesting with randomly selected, unused trees, we found that the probability of a tree being selected as a nest site was positively related to the number of cavity entrances, less dead crown, and more basal injuries (e.g., fire scars). At the nesting‐habitat scale, nest site selection was positively associated with the extent of basal injuries and number of cavity entrances in trees within 50 m of nest trees. These variables are likely important because they allow nesting parakeets to minimize cavity search times in potential nesting areas, thereby reducing energetic demands and potential exposure to predators. Slender‐billed Parakeets may thus use a hierarchical process to select nest sites; after a habitat patch is chosen, parakeets may then inspect individual trees in search of a suitable nest site. Effective strategies to ensure persistence of Slender‐billed Parakeets in agricultural‐forest mosaics should include preservation of both individual and groups of scattered mature trees.     

Causes of variable reproductive performance by Southern Ground‐hornbill Bucorvus leadbeateri and implications for management

Range occupancy of the cooperatively breeding Southern Ground‐Hornbill Bucorvus leadbeateri in South Africa has decreased by 65% in the last three generations and the effective management of the remaining populations is hampered by a lack of fundamental understanding of the factors determining reproductive performance. We examined the influence of social and environmental factors on the species' reproductive success in South Africa using data gathered from 23 groups over eight breeding seasons. Some groups had access to artificial nest‐sites, others did not. High rainfall (> 500 mm) over the breeding season led to a decrease in reproductive success, with groups being most successful in years when rain in the vicinity of the nest ranged from 300 to 500 mm. Groups breeding in natural nests were successful only when the proportion of open woodland surrounding the nest‐site was high. Those that bred in artificial nests, where overall breeding success was more than twice as high as those in natural nests, were less dependent on the availability of open woodland. Large groups (more than three birds) bred more successfully than groups comprising only two to three individuals. Group size, helper effects and rainfall cannot be managed to increase the productivity of Ground‐Hornbills but the fact that the availability of artificial nest‐sites and the amount of open woodland around the nest‐site both contribute positively to breeding performance identifies practical and simple management options for increasing the reproductive output of Southern Ground‐Hornbill populations.     

The effects of heterospecifics and climatic conditions on incubation behavior within a mixed‐species colony

Parental incubation behavior largely influences nest survival, a critical demographic process in avian population dynamics, and behaviors vary across species with different life history breeding strategies. Although research has identified nest survival advantages of mixing colonies, behavioral mechanisms that might explain these effects is largely lacking. We examined parental incubation behavior using video‐monitoring techniques on Alcatraz Island, California, of black‐crowned night‐heron Nycticorax nycticorax (hereinafter, night‐heron) in a mixed‐species colony with California gulls Larus californicus and western gulls L. occidentalis. We first quantified general nesting behaviors (i.e. incubation constancy, and nest attendance), and a suite of specific nesting behaviors (i.e. inactivity, vigilance, preening, and nest maintenance) with respect to six different daily time periods. We employed linear mixed effects models to investigate environmental and temporal factors as sources of variation in incubation constancy and nest attendance using 211 nest days across three nesting seasons (2010–2012). We found incubation constancy (percent of time on the eggs) and nest attendance (percent of time at the nest) were lower for nests that were located < 3 m from one or more gull nest, which indirectly supports the predator protection hypothesis, whereby heterospecifics provide protection allowing more time for foraging and other self‐maintenance activities. To our knowledge, this is the first empirical evidence of the influence of one nesting species on the incubation behavior of another. We also identified distinct differences between incubation constancy and nest attentiveness, indicating that these biparental incubating species do not share similar energetic constraints as those that are observed for uniparental species. Additionally, we found that variation in incubation behavior was a function of temperature and precipitation, where the strength of these effects was dependent on the time of day. Overall, these findings strengthen our understanding of incubation behavior and nest ecology of a colonial‐nesting species.     

The role of nest‐site selection and cereal production in differential nest predation in Common Quail Coturnix coturnix and hybrid quail C. coturnix × C. japonica

In Europe, farm‐reared quail used for restocking purposes are often hybrids between the Common Quail Coturnix coturnix and the Japanese Quail Coturnix japonica. These hybrids interbreed with wild Common Quail populations and suffer higher rates of nest predation, which would suggest that the two quail types have different nest‐site selection patterns. We monitored 103 radiotagged nesting females (77 wild Common Quail over 18 breeding seasons and 26 hybrid quail over six breeding seasons) and analysed their nest placement behaviour. Our results did not provide any evidence that the quail types showed different nest‐site selection trends, as they all preferred to nest near cereal field margins. The higher nest predation rate experienced by hybrids might therefore be explained by less effective anti‐predator behaviour among hybrid females during incubation. As no edge effect on nest predation was observed, the preference for nesting close to cereal margins could be due to the greater food resources of field margins. Moreover, Common Quail nest predation decreased with a rise in barley straw production, supporting evidence that nest cover plays an important role in the prevention of nest predation in this species. No such relationship was found for hybrid quail, but this result is not conclusive, as the data on barley straw production for hybrid quail covered a much smaller range of interannual variability. Barley straw production in Catalonia decreased significantly between 1992 and 2012, possibly as a result of a change in the barley varieties used. This practice could entail a conservation threat for ground‐nesting bird species.     

Age‐dependence and individual heterogeneity in reproductive success of greater sage‐grouse

Research on iteroparous species has shown that reproductive success may increase with age until the onset of senescence. However, from the population perspective, increased reproductive success with age could be a consequence of within‐individual variation (e.g. ageing, breeding experience, foraging ability hypotheses), between‐individual variation (e.g. individual heterogeneity, frailty, selection, delayed breeding hypotheses), or a combination thereof. We evaluated within‐ and between‐individual variation in reproductive success of greater sage‐grouse (Centrocercus urophasianus; sage‐grouse), a galliforme of conservation concern throughout western North America. We monitored female reproductive activity from 1998–2010 and used generalized linear mixed models incorporating within‐subject centering to evaluate and separate within‐ and between‐individual effects. We detected positive effects of within‐individual variation on nest initiation and success where ageing increased the likelihood of both parameters, which appears to support the breeding experience and/or foraging ability hypotheses. However, nest initiation was also affected by between‐individual variation whereby the likelihood of initiation was higher for individuals with higher mean age (i.e. survived longer), as is predicted by the frailty and selection hypotheses. Our results indicate both within‐ and between‐individual variation affect reproductive output of sage‐grouse, but the effects of each varied by measure of reproductive output. Our results corroborate previous findings that suggest population age parameters (i.e. cross‐sectional) should be interpreted with caution due to potential entanglement of within‐ and between‐individual processes. Moreover, the relative role and strength of within‐ and between‐individual processes appeared to vary by measure of reproductive output in our results, which further emphasizes the need for longitudinal analysis of age effects, even in relatively short‐lived iteroparous animals, to adequately interpret biological processes.