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The phenology, distribution, and size composition of plankton communities are changing rapidly in response to warming. This may lead to shifts in the prey fields of planktivorous fish, which play a key role in transferring energy up marine food chains. Here, we use 60 + years of Continuous Plankton Recorder data to explore temporal trends in key taxa and community traits in the prey field of planktivorous lesser sandeels (Ammodytes marinus) in the North Sea, the Faroes and southern Iceland. We found marked spatial variation in the prey field, with Calanus copepods generally being much more common in the northern part of the study area. In the western North Sea, the estimated amount of available energy in the prey field has decreased by more than 50% since the 1960s. This decrease was accompanied by declining abundances of small copepods, and shifts in the timing of peak annual prey abundances. Further, the estimated average prey community body size has increased in several of the locations considered. Overall, our results point to the importance of regional studies of prey fields, and caution against inferring ecological consequences based only on large-scale trends in key taxa or mean community traits.

Synchrony in demographic rates between spatially disjunct populations is a widespread phenomenon, although the underlying mechanisms are often not known. This synchrony and its spatial patterns can have important consequences for the long-term persistence of metapopulations and can also be used to infer drivers of population dynamics. Here, we examined spatial patterns of synchrony in the breeding success of black-legged kittiwakes Rissa tridactyla in the UK, using an extensive dataset on kittiwake breeding success and 2 different ways of measuring synchrony: one reflecting synchrony in inter-annual fluctuations only (rdiff) and one reflecting synchrony in both inter-annual fluctuations and long-term trends (r). We found that between-colony synchrony in breeding success decreased with distance up to just over 200 km but that some colony pairs showed stronger or weaker synchrony than expected based on distance. This was also reflected in the configuration of spatially coherent clusters of kittiwake colonies with synchronous breeding success. Further, we compared the support for different drivers of these spatial patterns, including trophic interactions and weather conditions. We found that the spatial dynamics of the kittiwakes’ main prey in this region, the lesser sandeel Ammodytes marinus, appeared to play some role in generating synchrony in long-term patterns, but their role in generating synchrony in inter-annual fluctuations was less clear. The study shows that examining spatial patterns in synchrony can provide useful information for inferring potential drivers and the spatial scale over which they are acting.

Funding: The infrared thermography equipment and software were funded by the Biotechnology and Biological Sciences Research Council, UK, grant number BB/K002775/1. The corticosterone hormonal work was supported by British Society for Neuroendocrinology Project Support Grant awarded to NE and CW. Understanding how biological markers of stress relate to stressor magnitude is much needed and can be used in welfare assessment. Changes in body surface temperature can be measured using infrared thermography (IRT) as a marker of a physiological response to acute stress. While an avian study has shown that changes in body surface temperature can reflect the intensity of acute stress, little is known about surface temperature responses to stressors of different magnitudes and its sex-specificity in mammals, and how they correlate with hormonal and behavioural responses. We used IRT to collect continuous surface temperature measurements of tail and eye of adult male and female rats (Rattus norvegicus), for 30 minutes after exposure to one of three stressors (small cage, encircling handling or rodent restraint cone) for one minute, and cross-validated the thermal response with plasma corticosterone (CORT) and behavioural assessment. To obtain individual baseline temperatures and thermal responses to stress, rats were imaged in a test arena (to which they were habituated) for 30 seconds before and 30 minutes after being exposed to the stressor. In response to the three stressors, tail temperature initially decreased and then recovered to, or overshot the baseline temperature. Tail temperature dynamics differed between stressors; being restrained in the small cage was associated with the smallest drop in temperature, in male rats, and the fastest thermal recovery, in both sexes. Increases in eye temperature only distinguished between stressors early in the response and only in females. The post stressor increase in eye temperature was greater in the right eye of males and the left eye of females. In both sexes encircling may have been associated with the fastest increase in CORT. These results were in line with observed behavioural changes, with greater movement in rats exposed to the small cage and higher immobility after encircling. The female tail and eye temperature, as well as the CORT concentrations did not return to pre-stressor levels in the observation period, in conjunction with the greater occurrence of escape-related behaviours in female rats. These results suggest that female rats are more vulnerable to acute restraint stress compared to male rats and emphasise the importance of using both sexes in future investigations of stressor magnitude. This study demonstrates that acute stress induced changes in mammalian surface temperature measured with IRT relate to the magnitude of restraint stress, indicate sex differences and correlate with hormonal and behavioural responses. Thus, IRT has the potential to become a non-invasive method of continuous welfare assessment in unrestrained mammals. Publisher PDF

Human impacts alter landscapes with consequences for the distribution and availability of high‐quality food resources to populations inhabiting those landscapes, which may impact on the reproductive output of individuals in those populations. The sensitivity of wild populations to changes in food resources may vary among stages of the annual cycle. For example, in birds, effects are likely to be greater during costly stages such as egg production. Here we compare assimilated diet (from stable isotope analysis of chick feathers) and egg traits (egg size, shape, eggshell colour and maculation, using pattern‐analysis software) in Herring Gulls Larus argentatus, across seven colonies in southwest Scotland and Northern Ireland. The Herring Gull is an opportunistic, generalist forager on both marine and terrestrial resources which frequently exploits anthropogenic food sources such as fishery discards and human refuse. We found that larger eggs were laid in colonies where females consumed either a higher proportion of marine resources or terrestrial resources; smaller eggs were laid in colonies where females had an intermediate diet. In colonies where females consumed more marine items, they also laid eggs with higher maculation (intensity and size of spots) compared with colonies where females mainly consumed terrestrial food. We also found smaller and more pointed eggs, suggestive of resource shortages, in larger colonies. Generalist foragers are often thought to have the capacity to buffer themselves against changes in the food web, provided that enough alternative food is available. However, this study highlights that specializing on the most profitable or available resources has consequences for egg traits even in an opportunistic generalist forager exploiting a large range of habitats. If variation in egg traits is related to reproductive output, then understanding the impact of assimilated diet on reproduction early in the breeding season can provide important insights into how populations will respond to landscapes altered by human impact.

Capsule: Herring Gull Larus argentatus colonies located along coastlines that are relatively sheltered from wave impact, associated with high intertidal prey availability, and with extensive farmland habitat in the vicinity, have increased in size over the last three decades.Aims: To investigate potential habitat drivers, associated with local food availability, of variation in Herring Gull colony sizes.Methods: Colony sizes were compared for 68 Herring Gull colonies across southwest Scotland and Northern Ireland to relate to the availability of potential foraging habitats surrounding each colony.Results: Changes in Herring Gull colony sizes were spatially clustered, with between-colony differences in colony size related to coastline exposure and amount of farmland within the foraging range. At the end of the census period, colonies located in areas with the greatest availability of intertidal prey and farmland habitat were larger compared to those that were not.Conclusion: Spatially clustered var...

Determining how resource use impacts on a species’ demography is important, especially in habitats which are being altered by anthropogenic land-use change. If changes result in species consuming resources of reduced quality their demographic traits may be adversely affected. Generalist species are useful when investigating changes in resource availability as they can switch to alternatives if their preferred food becomes unavailable. For species that can forage on marine and terrestrial resources, it is often not known whether a switch from marine to terrestrial resources will have negative consequences. The herring gull, Larus argentatus, is one widespread generalist that opportunistically forages within marine and terrestrial habitats that are increasingly altered by humans. We determined marine and terrestrial resource use of gulls from eight colonies over two years across south-west Scotland and Northern Ireland using pellets and stable isotope analysis of chick feathers, which gave comparable results. Herring gulls in the study region used very little marine offshore resources but birds from colonies located in areas with sheltered coastlines, which provide abundant and diverse marine food from the intertidal zone, foraged more on marine intertidal resources. In contrast, colonies closer to built-up areas used more terrestrial resources. Herring gulls raised larger broods in colonies where they consumed a higher proportion of marine resource. Therefore, where generalist species, such as gulls, switch to an alternative resources that is available to them within their foraging range this may come at a cost of lower breeding success.

Pellets of indigestible material regurgitated by Herring Gulls Larus argentatus breeding on Lady Isle, Firth of Clyde, Scotland, were collected in 2018 and 2019 and examined for earthworm chaetae. Nearly two-thirds (65.6%) of the 314 pellets came from gulls that had consumed one or more earthworms. Significantly fewer pellets (57.6%) contained chaetae in 2018, a relatively dry May to July, than in 2019 (72.0%) when rainfall was close to the long-term average for May to July. There were significant associations between the presence of large quantities of terrestrial vegetation in the pellet and the detection of large numbers of earthworm chaetae and/or fragments of terrestrial arthropods, suggesting that recent consumption of these food items is likely when pellets contain large quantities of vegetation.

Reactions to acute stressors are critical for survival. Yet, the challenges of assessing underlying physiological processes in the field limit our understanding of how variation in the acute stress response relates to fitness in free-living animals. Glucocorticoid secretion during acute stress can be measured from blood plasma concentrations, but each blood sample can only provide information for one point in time. Also, the number of samples that can be extracted from an individual in the field is usually limited to avoid compromising welfare. This restricts capacity for repeated assessment, and therefore temporal resolution of findings within- and between-acute stress responses - both of which are important for determining links between acute stress and fitness. Acute stress induces additional body surface temperature changes that can be measured non-invasively, and at high frequencies using thermal imaging, offering opportunities to overcome these limitations. But, this method's usefulness in the field depends on the extent that environmental conditions affect the body surface temperature response, which remains poorly understood. We assessed the relative importance of individual physiology (baseline glucocorticoid concentrations) and environmental conditions (air temperature and relative humidity) in determining the eye region surface temperature (Teye) response to acute stress, in wild blue tits (Cyanistes caeruleus) during trapping, handling and blood sampling. When controlling for between-individual baseline variation, Teye initially dropped rapidly below, and then recovered above baseline, before declining more slowly until the end of the test, 160 s after trap closure. One measure of the amplitude of this response – the size of the initial drop in Teye – was dependent on environmental conditions, but not baseline corticosterone. Whereas, two properties defining response dynamics – the timing of the initial drop, and the slope of the subsequent recovery – were related to baseline corticosterone concentrations, independently of environmental conditions. This suggests inferring the acute stress response using thermal imaging of Teye will be practical under fluctuating environmental conditions in the field., Highlights • We used thermal imaging (TI) to infer acute physiological stress in wild blue tits. • Body surface temperature fluctuated during trapping, handling and blood sampling. • A rapid initial drop was followed by recovery above baseline, then a slower decline. • Baseline corticosterone, not environmental conditions predicted two response dynamics. • Using TI to infer acute stress should be practical in varying environmental conditions.

Methods to obtain moult data from wild birds have not changed much over the last century and most studies still depend on checking museum specimens or capturing birds. Here we assess the applicability of systematic field photography for detecting and scoring moult in adult Black Skimmers Rynchops niger from southern Brazil. Moult data extracted from photographs have a high within- (RGLMM = 0.98) and between- (RGLMM = 0.97) observer repeatability and show very good fit to current Underhill-Zucchini moult models (R² = 0.75). Photography offers the advantages of being less invasive, requiring less equipment and human effort, being feasible in areas where captures may not be possible, and causing less disturbance so enhancing the number of sampled individuals. This article is protected by copyright. All rights reserved.

Between-population variation of changes in numbers can provide insights into factors influencing variation in demography and how population size or density is regulated. Here, spatio-temporal patterns of population change of Herring Gull (Larus argentatus), Lesser Black-backed Gull (L. fuscus) and Great Black-backed Gull (L. marinus) in the British Isles are described. The aim of this study was to test for density-dependence and spatial variation in population trends as two possible, but not mutually exclusive, explanations of population changes with important implications for the understanding of these changes. Between 1969 and 2013, the three species showed different population trends with Herring Gulls showing a strong decline, Lesser Black-backed Gulls an increase until 2000 but then a decline since, and Great Black-backed Gulls showing no clear pattern. Population changes in Herring Gulls varied between different regions of the British Isles with decreases in the northern and western parts ...

The differential allocation hypothesis (DAH) predicts that an individual should vary its reproductive\ud investment according to the attractiveness of its mate. A recently revised version of the DAH makes\ud explicit that investment can be positive, i.e. higher for the offspring of attractive males which should be\ud of higher quality, or negative, i.e. higher for offspring of unattractive males, for example compensating for\ud inheriting poor paternal genes. Moreover, investment can be made by the father and the mother. Here,\ud we tested whether experimental manipulation of male attractiveness affected parental investment at\ud different reproductive stages and thus influenced fitness-related traits in offspring. In two aviaries, all\ud male zebra finches, Taeniopygia guttata, were given red leg rings to increase attractiveness and in two\ud aviaries all males received green leg rings to decrease attractiveness. This controlled for assortative\ud mating between treatments. Ring colour was merely an experimental manipulation of male attractiveness,\ud not paternal quality, so we might expect additional investment to elevate offspring quality. Eggs\ud were cross-fostered between and within treatments to allow differentiation of effects of investment in\ud eggs and nestlings. Clutch and brood sizes were standardized. Both positive and negative investment\ud were observed: Eggs from red-ringed fathers had higher yolk to albumen ratios than eggs from greenringed\ud fathers. Nestlings from eggs laid and incubated by parents in the red-ringed group had higher\ud hatching masses than those in the green-ringed group. Both parents in the green-ringed group fed\ud nestlings more frequently than red-ringed parents. Offspring performance was influenced by the\ud treatment of both foster and biological parents, but combined effects of these different investment\ud patterns on fitness-related traits were ambiguous. Male attractiveness appeared to affect patterns of\ud reproductive investment but not consistently across all forms of reproductive investment suggesting that\ud the costs and benefits of differential allocation vary among individuals and across contexts.

A major aim of evolutionary biology is to understand why patterns of genomic variation vary among populations and species. Large-scale genomic studies of widespread species are useful for studying how the environment and demographic history shape patterns of genomic divergence, and with the continually decreasing cost of sequencing, such studies are now becoming feasible. Here, we carry out one of the most comprehensive surveys of genomic variation in a wild vertebrate to date; the great tit ( Parus major ) HapMap project. We screened ca 500,000 SNP markers across 647 individuals from 29 populations, spanning almost the entire geographic range of the European great tit subspecies. We found that genome-wide variation was consistent with a recent colonisation across Europe from a single refugium in the Balkans and/or Turkey, with bottlenecks and reduced genetic diversity in island populations. Differentiation across the genome was highly heterogeneous, with clear “islands of differentiation” even among populations which are ostensibly panmictic. These islands of differentiation were consistently found in regions of low recombination, suggesting that background selection can rapidly promote population differentiation among even the most recently colonised populations. We also detected genomic outlier regions that were unique to peripheral great tit populations, most likely as a result of recent directional selection at the range edges of this species. These “unique” outlier regions contained candidate genes for morphology, thermal adaptation and colouration, supporting previous research in this species, and providing avenues for future investigation. Our study suggests that comprehensive screens of genomic variation in wild organisms can provide unique insights into evolution. Author summary Studying patterns of genetic variation is a useful way of determining why populations and species differ in nature. Genetic variation is shaped by natural selection, but also by the present and past size of populations, the amount of migration, and by features of the genome, such as variation in recombination rate, of the organism being studied. Teasing apart the effects of these different processes on genomic diversity is difficult, but one way that this can be achieved is by studying genomic variation across the entire range of a species. We performed a continental-scale analysis of genetic variation in the great tit - a widespread songbird that has been the focus of extensive ecological research. We first used genomic data to reconstruct the historical colonisation of great tits across Europe, and showed that during the last ice age, this species was likely restricted to a single region in Eastern Europe, from which they spread across the continent. We then studied how patterns of variation differ along the genome, and show that recombination rate is a key driver of variation among all populations. Importantly, by comparing many populations we were able to identify genes that have been subject to natural selection in specific geographical regions. We found that natural selection appeared to be strongest in populations on the edges of the great tit’s range acting on traits such as morphology, stress response and colouration. Large-scale genetic analyses such as ours are therefore useful approaches for understanding how evolution operates in the wild.

There is little evidence documenting the prevalence of plastic nest incorporation for different seabird species and populations, and even less detailing the source of such debris as nesting material. This study presents a baseline dataset on the presence of plastic in the nests of five seabird species on Lady Isle, Scotland using a novel and repeatable methodology for quantifying plastic incorporated into nests. Plastic was found in 24.5% to 80% of nests of all species. We analysed pellets of regurgitated material and the spatial distribution of herring gull nests containing plastic in the context of the tide and nesting habitat. Differences in the types of plastic found in pellets and nests suggests that plastic incorporated into herring gull nests was not derived at foraging sites and likely collected from the local environment. Targeted beach cleans before the breeding season could help minimise the quantity of plastic available to herring gulls.

Co-existence between great tits Parus major and blue tits Cyanistes caeruleus, \ud but also other hole nesting taxa, constitutes a classic example of species\ud co-occurrence resulting in potential interference and exploitation competition\ud for food and for breeding and roosting sites. However, the spatial and temporal\ud variation in co-existence and its consequences for competition remain poorly\ud understood. 2.We used an extensive database on reproduction in nest boxes by\ud great and blue tits based on 87 study plots across Europe and Northern Africa\ud during 1957-2012 for a total of 19,075 great tit and 16,729 blue tit clutches to \ud assess correlative evidence for a relationship between laying date and clutch\ud size, respectively, and density consistent with effects of intraspecific and\ud interspecific competition. 3.In an initial set of analyses, we statistically\ud controlled for a suite of site specific variables. We found evidence for an\ud effect of intraspecific competition on blue tit laying date (later laying at\ud higher density) and clutch size (smaller clutch size at higher density), but no\ud evidence of significant effects of intraspecific competition in great tits, nor\ud effects of interspecific competition for either species. 4.To further control for\ud site-specific variation caused by a range of potentially confounding variables,\ud we compared means and variances in laying date and clutch size of great and blue \ud tits among three categories of difference in density between the two species. We \ud exploited the fact that means and variances are generally positively correlated. \ud If interspecific competition occurs, we predicted a reduction in mean and an\ud increase in variance in clutch size in great tit and blue tit when density of\ud heterospecifics is higher than the density of conspecifics and for intraspecific \ud competition this reduction would occur when density of conspecifics is higher\ud than the density of heterospecifics. Such comparisons of temporal patterns of\ud means and variances revealed evidence, for both species, consistent with\ud intraspecific competition and to a smaller extent with interspecific competition.\ud 5.These findings suggest that competition associated with reproductive behaviour \ud between blue and great tits is widespread, but also varies across large spatial\ud and temporal scales. This article is protected by copyright.

Body temperature (Tb) is a valuable parameter when assessing the physiological state of animals, but its widespread measurement is often constrained by methods that are invasive or require frequent recapture of animals. Alternatives based on automated remote sensing of peripheral Tb show promise, but little is known about their strengths and limitations. We measured peripheral Tb in great tits Parus major with subcutaneously implanted passive integrated transponders (PIT tags) and externally attached radio transmitters to determine repeatability of measurements, sensitivity of each method to variation in ambient temperature (Ta) and wind speed, the relationship between methods, and their ability to capture circadian variation in Tb. Repeatability of measurements by radio transmitters was high (> 80%) when readings were taken within 20 min, but reduced to 16% when measures were spaced 3.5 h apart. PIT tag data for the 3.5 h interval were more repeatable (33%) and less variable (cv). Data were affected by Ta with a stronger effect on the externally attached transmitters, but the influence of wind speed was small for both methods. There was a significant positive relationship between transmitter- and PIT tag temperature during both days and nights. Both methods were equally suited to detect diel changes in peripheral Tb. However, transmitters offered longer detection distance and better temporal resolution. These qualities should be considered when deciding how to collect Tb data remotely. If properly deployed, both methods allow measurement of peripheral Tb over a wide range of natural systems and conditions in small, free-ranging, birds.

© 2014 The Authors. Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Our understanding of fundamental organismal biology has been disproportionately influenced by studies of a relatively small number of ‘model’ species extensively studied in captivity. Laboratory populations of model species are commonly subject to a number of forms of past and current selection that may affect experimental outcomes. Here, we examine these processes and their outcomes in one of the most widely used vertebrate species in the laboratory – the zebra finch (Taeniopygia guttata). This important model species is used for research across a broad range of fields, partly due to the ease with which it can be bred in captivity. However despite this perceived amenability, we demonstrate extensive variation in the success with which different laboratories and studies bred their subjects, and overall only 64% of all females that were given the opportunity, bred successfully in the laboratory. We identify and review several environmental, husbandry, life-history and behavioural factors that potentially contribute to this variation. The variation in reproductive success across individuals could lead to biases in experimental outcomes and drive some of the heterogeneity in research outcomes across studies. The zebra finch remains an excellent captive animal system and our aim is to sharpen the insight that future studies of this species can provide, both to our understanding of this species and also with respect to the reproduction of captive animals more widely. We hope to improve systematic reporting methods and that further investigation of the issues we raise will lead both to advances in our fundamental understanding of avian reproduction as well as to improvements in future welfare and experimental efficiency.

Marie Vaugoyeau [et al.], The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole-nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large-scale study showing a species-specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size., E. Barba acknowledges funding of project CGL2013-48001-C2-1-P (Spanish Ministry of Science and Innovation).

Increasing temperatures associated with climate change may generate phenological mismatches that disrupt previously synchronous trophic interactions. Most work on mismatch has focused on temporal trends, whereas spatial variation in the degree of trophic synchrony has largely been neglected, even though the degree to which mismatch varies in space has implications for meso-scale population dynamics and evolution. Here we quantify latitudinal trends in phenological mismatch, using phenological data on an oak-caterpillar-bird system from across the UK. Increasing latitude delays phenology of all species, but more so for oak, resulting in a shorter interval between leaf emergence and peak caterpillar biomass at northern locations. Asynchrony found between peak caterpillar biomass and peak nestling demand of blue tits, great tits and pied flycatchers increases in earlier (warm) springs. There is no evidence of spatial variation in the timing of peak nestling demand relative to peak caterpillar biomass for any species. Phenological mismatch alone is thus unlikely to explain spatial variation in population trends. Given projections of continued spring warming, we predict that temperate forest birds will become increasingly mismatched with peak caterpillar timing. Latitudinal invariance in the direction of mismatch may act as a double-edged sword that presents no opportunities for spatial buffering from the effects of mismatch on population size, but generates spatially consistent directional selection on timing, which could facilitate rapid evolutionary change.

Parent–offspring conflict theory posits that parents and offspring differ in the optimal duration of parental care. Accordingly, the timing of the termination of parental care could be contested between parents and offspring. The outcome of this conflict might be related to the degree of parental sensitivity to offspring demand. By cross-fostering zebra finch, Taeniopygia guttata, chicks of different ages among breeding pairs, we examined whether parents continue parental behaviour for a fixed time period, or are sensitive to offspring age. Parents provisioned chicks in the nest for longer when they received younger foster chicks and for less time when they received older foster chicks. All chicks fledged and became independent at the same age. Our results show that zebra finch parents extend the period of parental care in response to protracted demands from their brood.

Whether parental effort can be negotiated between partners over ecological time and adjusted across different contexts is not well understood. We manipulated male extra-pair copulation (EPC) opportunity in captive zebra finches, Taeniopygia guttata, to test whether males adjust incubation effort to the mating context and to examine how females respond to their partner’s effort. Birds without previous breeding experience were paired randomly and bred with the same partner twice. In the first breeding attempt, half the males received EPC opportunities with ‘extra-pair females’ during incubation, while the other half did not. Males that received EPC opportunities in the first breeding attempt did not in the second breeding attempt and vice versa. We recorded incubation effort on days when EPC opportunities were not presented. In their first breeding attempt, males with EPC opportunities incubated less than those without. Females compensated fully for the deficit in male care so that a pair’s combined incubation effort was unchanged. In the second attempt, when a male’s opportunity for EPCs was switched, individuals showed the same level of incubation effort that they had previously, irrespective of the current availability of extra-pair females. This suggests that division of effort was negotiated in the first breeding attempt and maintained without significant adjustments in the second attempt. The effects of male EPC opportunity in the first breeding attempt on subsequent incubation effort suggests that individual parental decisions can be shaped by previous experience and this may partly explain conflicting results in studies where individuals’ histories were not known.

The Black Skimmer Rynchops niger is a New World's coastal migratory bird with three subspecies based on distinctive morphometrics, plumage, and breeding distribution. Its population is decreasing in North America while numbers are poorly known in South America. Here we conducted an extensive review of the literature on the Black Skimmer to fill gaps in the knowledge of each subspecies' natural history and suggest where future research should focus. We used 41,247 records collected from databases and literature to establish the overall occurrence and location of breeding colonies for each subspecies. We also collected information on the phenology of life-history events described in 150 papers and books to compare the annual cycle of the three subspecies. The North American subspecies R. n. niger breeds, migrates and stages along the coast with few records inland. The South American R. n. intercedens and Amazonian R. n. cinerascens subspecies breed inland and migrate mostly to coastal staging sites. The pre...

Models of climate change predict that its effects on animal populations will not always be negative, but most studies indicate negative associations between changes in climate and the phenology of animal migration and reproduction. For some populations, however, climate change may render particular environments more favourable, with positive effects on population growth. We used a 30-year population dataset on over 2000 Common Eiders Somateria mollissima at a colony in southwest Iceland to examine the response of this species to climate fluctuations. Eiders are strongly dependent on suitable climatic conditions for successful reproduction and survival. Temperatures in southwest Iceland, in both winter and summer, have generally increased over the past 30 years but have shown considerable fluctuation. We show that females laid earlier following mild winters and that year-to-year variation in the number of nests was related to the temperature during the breeding season 2 years previously. Milder summers could have positive effects on breeding success and offspring survival, producing an increase in nest numbers 2 years later when most Eiders recruit into the breeding population. In this part of their range, Eiders could benefit from a general warming of the climate.

Very few studies have examined, at the field scale, the potential for faecal residues in the dung of avermectin-treated cattle to affect dung-breeding insects. The current study examined populations of dung beetles (Scarabaeidae: Aphodius) using pitfall traps baited with dung from untreated cattle on 26 fields across eight farms in southwest Scotland. The fields were grazed either by untreated cattle or by cattle treated with an avermectin product, i.e. doramectin or ivermectin. During the two-year study, significantly more beetles were trapped in fields grazed by treated cattle (n=9377 beetles) than in fields where cattle remained untreated (n=2483 beetles). Additional trials showed that beetles preferentially colonised dung of untreated versus doramectin-treated cattle. This may explain the higher captures of beetles in traps baited with dung of untreated cattle, which were located in fields of treated cattle. Given that Aphodius beetles avoided dung of treated cattle in the current study, the potential harmful effects of avermectin residues in cattle dung could be reduced through livestock management practices that maximise the availability of dung from untreated livestock in areas where avermectins are being used.

1. In species that actively maintain optimal conditions for offspring development, the quality of the breeding site may have a substantial effect on reproductive effort and breeding success. Here, we studied the effect of nest shelter on reproductive costs for incubating female common eiders (Somateria mollissima) nesting in an arctic environment using both a correlative and an experimental approach. 2. The correlative data showed that females that nested in naturally well-sheltered nests experienced milder temperatures, laid larger clutches, provided a more favourable thermal environment for their eggs and had a higher hatching success than those at exposed nest-sites. 3. We added artificial shelters to exposed nest-sites and compared them to unmanipulated exposed nest-sites to examine the effect of nest shelter in females of similar quality. Costs of nesting in exposed sites were greater as evidenced by the reduced female mass loss and more favourable thermal conditions for the eggs at artificially sheltered nest-sites relative to exposed nest-sites. However, there was no difference in hatchings success between artificially sheltered and exposed nest-sites. 4. This study shows even small changes in the climatic conditions at the nest-site can have substantial consequences for reproductive effort, but the association between nest-site quality and breeding success in eiders is due to better quality individuals occupying better nest-sites.

An organism’s pattern of development can have important long-term fitness effects. In species where the sexes differ in size or other phenotypic traits, they may also have different optimal developmental rates. This influences both parental sex allocation strategies and susceptibility of the sexes to early developmental conditions. However, sex differences in developmental rate and vulnerability to environment during the embryonic period are not well understood. In birds, sibling competition and hatching asynchrony may select for accelerated embryonic development of the last offspring in order to reduce their competitive disadvantage after hatching. They may advance their hatching in response to vocal stimuli by the older siblings. It is, however, unclear whether this flexibility in developmental rates is sex specific. In this study, we experimentally manipulated between-embryo contact and tested whether this affected the pre-natal developmental rate and post-hatching performance of male and female offspring from last-laid eggs in the herring gull. Post-hatching performance was measured both in competitive and non-competitive situations. Among young incubated in isolation, males hatched faster than females, but both sexes fledged in similar, relatively good condition. Among young incubated with normal between-embryo contact, hatching time did not differ between sexes, but males fledged in poorer condition than females, regardless of whether they were reared singly or in a brood. These results suggest that male and female offspring differ in their ability to mitigate the costs of hatching asynchrony.

Many populations have recovered from severe bottlenecks either naturally or through intensive conservation management. In the past, however, few conservation programs have monitored the genetic health of recovering populations. We conducted a conservation genetic assessment of a small, reintroduced population of Mauritius Kestrel (Falco punctatus) to determine whether genetic deterioration has occurred since its reintroduction. We used pedigree analysis that partially accounted for individuals of unknown origin to document that (1) inbreeding occurred frequently (2.6% increase per generation; N(eI)= 18.9), (2) 25% of breeding pairs were composed of either closely or moderately related individuals, (3) genetic diversity has been lost from the population (1.6% loss per generation; N(eV)= 32.1) less rapidly than the corresponding increase in inbreeding, and (4) ignoring the contribution of unknown individuals to a pedigree will bias the metrics derived from that pedigree, ultimately obscuring the prevailing genetic dynamics. The rates of inbreeding and loss of genetic variation in the subpopulation of Mauritius Kestrel we examined were extreme and among the highest yet documented in a wild vertebrate population. Thus, genetic deterioration may affect this population's long-term viability. Remedial conservation strategies are needed to reduce the impact of inbreeding and loss of genetic variation in this species. We suggest that schemes to monitor genetic variation after reintroduction should be an integral component of endangered species recovery programs.

Oxidative stress (OS) as a proximate mechanism for life-history trade-offs is widespread in the literature. One such resource allocation trade-off involves growth rate, and theory suggests that OS might act as both a constraint on and a cost of growth, yet studies investigating this have produced conflicting results. Here, we use meta-analysis to investigate whether increased OS levels impact on growth (OS as a constraint on growth) and whether greater growth rates can increase OS (OS as a cost of growth). The role of OS as a constraint on growth was supported by the meta-analysis. Greater OS, in terms of either increased damage or reduced levels of antioxidants, was associated with reduced growth although the effect depended on the experimental manipulation used. Our results also support an oxidative cost of growth, at least in terms of increased oxidative damage, although faster growth was not associated with a change in antioxidant levels. These findings that OS can act as a constraint on growth support theoretical links between OS and animal life histories and provide evidence for a growth-self-maintenance trade-off. Furthermore, the apparent oxidative costs of growth imply individuals cannot alter this trade-off when faced with enhanced growth. We offer a starting platform for future research and recommend the use of oxidative damage biomarkers in nonlethal tissue to investigate the growth-OS relationship further.

The effects of avermectin exposure on natural populations of the yellow dung fly,Scathophaga stercorariaLinnaeus, were investigated at the field scale on farms in south-west Scotland. Pastures forming the focus of the study were grazed with either untreated cattle or cattle receiving standard, manufacturer-recommended treatment regimes of an avermectin product. Flies were sampled between April and July in 2002 and 2003 using dung-baited pitfall traps. Abundance and wing asymmetry inS. stercorariapopulations were examined in relation to a range of environmental and management variables (including avermectin exposure, pasture management intensity, weather and season). Data used for abundance analyses were collected in fields where treated cattle had been dosed with either doramectin or ivermectin, while the data for the asymmetry analyses were from a subset of fields where treated cattle had been dosed with doramectin only. While abundance ofS. stercorariavaried significantly between years and with season, there was no difference in their abundance between fields grazed by avermectin-treated or untreated cattle. Asymmetry was significantly higher in fly populations in fields grazed by doramectin-treated cattle, suggesting that exposure to doramectin during development could have imposed some degree of environmental stress. While these results suggest that exposure to doramectin residues in dung on grazed pastures may have sublethal effects on the insects developing in that dung, there was no evident avermectin effect on the abundance of adultS. stercorariaoccurring in the pastures.

The quality of conditions provided by avian parents will have consequences for both parental and offspring fitness. While many components of avian reproduction appear to vary with parental age, the effect of age on incubation has largely been ignored so far. In this study, we tested whether young herring gulls provide a different incubation environment from mature ones and whether this has consequences for offspring performance. Laying and rearing conditions were standardised using a cross-fostering protocol. Egg predation rates tended to be higher in the nests of young parents. However, nest site, nest construction and egg temperature during incubation did not vary with parental age. Overall, the duration of incubation was shorter in young compared to mature birds and this reflected the later laying date of the former, since incubation duration generally decreased across the season. However, male eggs incubated by young parents had longer incubation periods than predicted for their laying dates. In contrast, incubation length of female eggs incubated by young pairs, and of male and female eggs incubated by mature birds did not deviate from the expected for any given laying date. Offspring that had been incubated by young parents had considerably poorer survival than those incubated by mature pairs, despite being reared under standardized, favourable conditions (singly, by mature parents). This was due to increased mortality among female chicks that had been incubated by young parents. The chicks incubated as eggs by young and mature birds, which survived until fledging, did not differ in body mass and size growth, or body condition. The results of this study demonstrate that parental age can influence offspring performance via variation in incubation environment, and that females are more susceptible than males to conditions experienced during embryonic development.

Asynchronous hatching is believed to result from incubation starting before laying is completed and leads to a hierarchy in chick size, with detrimental effects on the small, last-hatching chick. If chicks have some control over the timing of hatching, however, they might be able to compensate at least partly for the hatching spread. In this study on lesser black-backed gulls, Larus fuscus, we examined whether the timing of hatching varied with laying order. To separate their effects, we manipulated the hatching order relative to the laying order within broods of three chicks. We exchanged eggs between nests, so that chicks from first-laid eggs hatched last, and chicks from last-laid eggs hatched first. When hatching last, chicks from last-laid eggs took significantly less time to hatch than chicks from first-laid eggs. However, hatching duration did not differ between chicks from first- and last-laid eggs when hatching first. Hence, only offspring from last-laid eggs had the ability to accelerate hatching, and they did this only when their siblings had already hatched. Accelerated hatching might benefit the last-laid offspring by reducing hatching intervals, but it also resulted in low hatchling mass. As this ability to accelerate hatching was not found in first-laid offspring, differences in intrinsic egg quality may be responsible for hatching duration. Differences in egg quality, however, arise from differential allocation of resources across the laying sequence by the parent and may hence suggest a parental strategy to alleviate the detrimental effects of within-brood hierarchies on the last-hatching chick.

Summary 1Poorer performance in young compared with older age classes is common among iteroparous animals, and may result from age effects acting on different phases of reproduction. However, such effects have generally not been teased apart, and as a result their separate consequences for offspring fitness have remained unclear. 2We tested whether young and mature Herring Gulls produce offspring of different quality due to differences in their egg-production capacity. An experimental protocol was used to standardize incubation and rearing conditions. 3Eggs laid by young females were smaller than eggs of mature birds. They had a reduced absolute amount of albumen, which reflected mainly lower water content. 4Within the young age class, female eggs were smaller than male eggs and had significantly paler yolk. This was due to young birds having reduced water content of the albumen, as well as carotenoid content in eggs containing a female embryo. In the mature class there were no such differences in resource allocation in relation to embryo sex. The sex ratio of clutches did not vary with parental age. 5Hatchlings from eggs produced by young pairs were smaller skeletally but had similar mass to hatchlings from eggs laid by mature birds. When reared by mature foster parents, they did not differ in structural growth or survival. However, they showed a different pattern of growth in body mass. 6Our results suggest that poorer reproductive performance of young breeders is not simply a consequence of poorer egg quality, as this was largely compensated for when the eggs of young pairs were incubated and reared by mature foster parents. However, it is also possible that adverse effects of lower egg quality are not manifest in the offspring until later in life.

Suboptimal conditions during embryonic development can affect offspring fitness. Both egg quality and incubation behavior can affect hatching success, hatching mass, and subsequent offspring performance. These effects may differ between male and female offspring. We manipulated the prebreeding body condition of Zebra Finches (Taeniopygia guttata), using diets of different protein content. To separate possible effects on egg quality of parental body condition and incubation conditions, we did a cross-fostering experiment. We analyzed embryo survival and hatching mass with respect to body condition of the egg-laying parent, body condition of the incubating foster parent, and offspring sex. Embryos were not affected by the condition of the egg-laying parent. Eggs incubated by parents in better condition suffered less embryo mortality than those incubated by parents in poorer condition, but only when overall embryo mortality was low. Hatching mass was also affected by the incubating foster parent’s body condition. And hatchlings incubated by parents in good condition were heavier than those incubated by parents in poor condition. Female hatchlings from late-laid eggs were heavier, in comparison with the size of the egg from which they hatched, than female hatchlings from earlier-laid eggs. No such effect was found for males. Therefore, male and female embryos may differ in their sensitivity to suboptimal conditions during embryonic development. These results suggest that parental body condition during incubation can affect offspring fitness.Efectos de las Condiciones de Incubación y el Sexo de las Crías sobre el Desarrollo Embrional y la Supervivencia en Taeniopygia guttata

Gorman, H. E., Arnold, K. E. and Nager R. G. 2005. Incubation effort in relation to male attractiveness in zebra finches Taeniopygia guttata . / J. Avian Biol. 36: 413/420. The division of labour in parental care between the two sexes varies between and within species. In birds, parents have been shown to invest more into egg production and nestling care when paired with an attractive rather than an unattractive mate, as predicted by the differential allocation hypothesis. Here we investigate variation in the female’s and male’s share of incubation behaviour, a vital, and costly, period of parental care during which the embryo is vulnerable to perturbations in developmental conditions. We manipulated the attractiveness of male zebra finches Taeniopygia guttata , using red or green leg-rings. To simulate their natural social environment we allowed them to breed in outdoor aviaries. All males within an aviary were given the same coloured ring to avoid ring-colour related assortative mating. Males within a colony, however, were still expected to show some variation in attractiveness with the earliest laying females possibly pairing with the most attractive males. Indeed we found that both factors played a role in explaining female incubation effort. Among females mated to red ringed males, earlier laying females contributed significantly more to incubation than late laying females, but no such pattern was found in females mated to green ringed males. Overall, there were no differences in the level of incubation provided by both parents between treatment groups, suggesting some compensation within the pair. Hatching success was correlated with a pair’s total incubation effort. These results suggest that variation in the division of parental care between the sexes is in agreement with both increased effort of females mated with attractive males, and females compensating for the reduced effort of attractive males seeking further mating opportunities. These two factors can act at the same time in natural populations and both should be considered when explaining variation in division of labour between the sexes.

Females mated to attractive males are predicted to produce male-biased broods. Previous studies on zebra finches, Taeniopygia guttata, in which colored leg rings were used to alter male attractiveness, support this hypothesis. However, because molecular sexing techniques were not available, it was not known when during development this bias arose. Also, because both attractive (red-ringed) and unattractive (green-ringed) males were within the same aviary, assortative mating between treatments may have confounded the results. Using two different experimental designs, we tested whether the sex ratio of zebra finch eggs and chicks differed in response to paternal ring color whilst controlling for assortative mating between treatments. In the aviary experiment, birds could interact socially, but all males in an aviary had the same leg ring color. In the cage experiment, each female was randomly assigned a red- or green-ringed mate, thus also eliminating assortative mating within treatments. Offspring were sexed based on plumage or using a molecular method. The sex ratio at laying did not differ between treatments in either the aviary (n = 313 eggs) or cage (n = 151 eggs) experiments, suggesting that female zebra finches do not manipulate the primary sex ratio in response to their mate's ring color. However, in the cage experiment we found greater male embryonic mortality in the attractive group, which resulted in a female-biased sex ratio at sexual maturity, that is, in the opposite direction to that found in previous studies. Possible explanations for the disparity between our results and those of previous studies are considered. Copyright 2005.

Egg composition, which is under maternal control, can have a profound effect on offspring fitness. The presence of maternal testosterone and carotenoids in avian egg yolk, for example, is thought to enhance the development and competitive ability of the offspring and protect the hatching and growing chick against oxidative stress. Egg quality often differs between females and such variation can be due to differences in maternal social environment, e.g. breeding density. However, this is confounded by the possibility that the quality of individuals breeding in high- or low-density areas may vary. We tested if maternal social environment influences egg composition in a colonial seabird, the lesser black-backed gull (Larus fuscus). To control for confounding effects of female quality, we experimentally manipulated maternal social environment during egg formation. We increased the frequency of intra-specific interactions (i.e. aggressive encounters with conspecifics other than nest mates) in which the females were involved, by placing an elevated platform in their territory. Females that took part in more intra-specific interactions produced a heavier last egg, but the yolk testosterone concentration in eggs laid by control and experimental females did not differ. Differences in yolk testosterone concentration in relation to embryo sex were found neither in the control nor in the experimental group. In contrast, within the control group, eggs with a male embryo contained more carotenoids than eggs with a female embryo. Moreover, experimental females that had been involved in more intra-specific interactions produced female eggs with higher carotenoid levels compared to female eggs of control birds. An experimental increase in carotenoid levels was not observed in eggs containing a male embryo. Our results suggest that intra-specific interactions experienced by female birds during egg formation can influence conditions for embryonic development.

State dependence is thought to be an important factor in resource allocation decisions, particularly in those decisions relating to parental care. Incubation behaviour is a costly part of avian parental care, and therefore likely to depend on the parent's body condition. We manipulated the body condition of zebra finches, Taeniopygia guttata, during incubation by using prelaying diets of different protein content, and induced birds to lay a similar number of eggs. We tested the hypothesis that birds in better body condition would invest more in incubation than birds in poorer condition when incubating on the same diet. Females that had received a high-protein prelaying diet lost more body mass than those that had received a low-protein diet. They also increased the length of their incubation bouts between early and middle incubation, whereas females that had received a low-protein prelaying diet increased bout length only between middle and late incubation. There were no differences between males from the two diet groups, and males were responsible for a lesser proportion of incubation than females. These results indicate that incubation behaviour is state dependent in female zebra finches, and that birds of different body condition adopt different incubation strategies. We found no differences in incubation duration and hatching success between the two incubation strategies in captivity, but the potential risk of nest predation in the wild may differ. We suggest that only females in good condition can afford to adopt a strategy of increasing bout length early in incubation; females in poorer condition first have to recover their body condition after having produced a clutch. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour

Egg quality is a phenotype of, and can profoundly influence fitness in, both mother and offspring. However, the physiological mechanisms that underlie this maternal effect are poorly understood. Carotenoids are hypothesized to enhance antioxidant activity and immune function, and are responsible for the pigmentation of egg yolk. The proximate basis and consequences of this maternal investment, however, have not previously been studied in wild birds. In this supplemental feeding study of lesser black-backed gulls, Larus fuscus, carotenoid-fed females are shown to have increased integument pigmentation, higher plasma concentrations of carotenoids and antioxidant activity, and lower plasma concentrations of immunoglobulins (Igs) in comparison with controls. In turn, carotenoid-fed females produced eggs containing high carotenoid but low Ig concentrations (i.e. passive immunity), whereas control females produced eggs containing low carotenoid but high Ig concentrations. Within-clutch patterns of these resources varied over the laying sequence in a similar manner in both carotenoid-fed and control nests. Our results suggest that carotenoids could be one resource responsible for egg quality maternal effects in birds. We discuss the possible implications of carotenoid-mediated effects on phenotype for fitness in mothers and their offspring.

Summary In many incubating birds heat transfer from parent to egg is facilitated by the brood patch, an area of ventral abdominal skin that becomes highly vascularised, swells and loses its down feathers around the time of laying. Only the female develops a brood patch in most passerine species, but some males incubate and maintain the eggs at similar temperatures to females without a brood patch. Here we used a novel application of infra-red thermography (IRT) to examine sex differences in parental care from a physiological perspective. Using incubating male and female zebra finches (Taeniopygia guttata), a species in which the male lacks a brood patch, we measured the surface temperature of the ventral plumage overlying the abdomen and a reference area that does not contact the eggs (thorax) twice per pair. In half of the pairs clutch size was experimentally enlarged between the two sets of measurements to increase incubation demand. We found that the temperature differential between abdomen and thorax plumage was greater in females than in males, and that abdomen plumage was warmer after clutch enlargement than before it in females but not in males. These findings are consistent with morphological sex differences in brood patch development and suggest that male and female zebra finches differ in the way they regulate abdomen versus general body surface temperature in response to variation in incubation demand.

Møller, A.P. [et al.], Secondary hole-nesting birds that do not construct nest holes themselves and hence regularly breed in nest boxes constitute important model systems for field studies in many biological disciplines with hundreds of scientists and amateurs involved. Those research groups are spread over wide geographic areas that experience considerable variation in environmental conditions, and researchers provide nest boxes of varying designs that may inadvertently introduce spatial and temporal variation in reproductive parameters. We quantified the relationship between mean clutch size and nest box size and material after controlling for a range of environmental variables in four of the most widely used model species in the Western Palaearctic: great tit Parus major, blue tit Cyanistes caeruleus, pied flycatcher Ficedula hypoleuca and collared flycatcher F. albicollis from 365 populations and 79 610 clutches. Nest floor area and nest box material varied non-randomly across latitudes and longitudes, showing that scientists did not adopt a random box design. Clutch size increased with nest floor area in great tits, but not in blue tits and flycatchers. Clutch size of blue tits was larger in wooden than in concrete nest boxes. These findings demonstrate that the size of nest boxes and material used to construct nest boxes can differentially affect clutch size in different species. The findings also suggest that the nest box design may affect not only focal species, but also indirectly other species through the effects of nest box design on productivity and therefore potentially population density and hence interspecific competition. © 2014 The Authors. Methods in Ecology and Evolution © 2014 British Ecological Society., J. Morales by the Spanish Research Council-CSIC and the European Social Fund (programme Junta para la Ampliación de Estudios).

Studies of sexual selection have mostly focused on female choice and male–male competition. However, when a male contributes substantially to a breeding attempt, he would benefit from choosing a female whose reproductive potential is high in order to maximize his reproductive success. We examined male mate choice in zebra finches, Taeniopygia guttata, with respect to experimental manipulation of female fecundity. This was enhanced by a dietary supplement prior to contact with the males. When housed with females of differing fecundity, male zebra finches preferentially direct their courtship towards more fecund females. This outcome could be a consequence of female–female interactions or direct male preference. In this experiment, we gave males a choice between standard and supplemented females in an experimental arena where the females were spatially segregated, and could not interact with each other, thereby excluding the possibility of female–female competition. Males spent significantly more time with the more fecund female, they hopped more times on the perch opposite that female and were twice as likely to choose her than the standard female. The standard females were not, however, unreceptive to males, nor did they appear to differ in their activity patterns in response to the male. These results show that males are able to distinguish females that have higher fecundity, and that this is not the result of relative female competitiveness or dominance status.  2001 The Association for the Study of Animal Behaviour

Measurements of costs of reproduction are essential for our understanding of the evolution of reproductive effort. While in birds the effects of increased chick-rearing effort on subsequent survival and fecundity have been relatively well studied experimentally, costs associated with increased egg-production effort have received relatively little attention. We experimentally increased the egg-production effort of individually marked Lesser Black-backed Gulls Larus fuscus and followed their breeding performance in the next year. In the season following increased egg production, females, but not males, were less likely to be resighted in the study plot and those that did return were less likely to produce a clutch compared to control birds. It is unclear whether the observed effect on local return rate represents differential survival, differences in breeding propensity or differences in dispersal between experimental and control females. In any event, all of these would adversely affect the fitness of experimental females. In addition, those experimental females that did breed invested less in egg production the following season, which again is likely to affect breeding performance. Thus, this study provides evidence that there is an inter-brood trade-off between current egg-production effort and future fitness in birds.

Empirical evidence is growing that the offspring sex ratio in birds can be biased in relation to the body condition of parents during breeding. The sex ratio bias may come about because (1) the actual production of the two sexes may be skewed and/or (2) there may be a sex bias in early nestling mortality contingent on parental condition. By manipulating parental condition and giving them a control brood to rear, thereby eliminating effects operating via the eggs, we examined the extent to which parental condition influences the post-hatching survival of male and female lesser black-backed gulls, Larus fuscus. We found that the pre-fledging survival of male chicks was strongly reduced in all-male broods reared by parents in poor condition. Pre-fledging survival of female chicks was, however, unaffected by parental condition or brood sex composition. Thus, independently of any production biases, sex differences in nestling mortality alone can bias the offspring sex ratio at fledging in relation to the prevailing rearing conditions. In other studies on gulls we have, however, also shown that females in poor condition at laying preferentially produce female eggs. Clearly a bias in fledging sex ratio can occur within the same species due to a combination of differential production and differential post-laying mortality; the latter can involve a differential effect of poor egg quality on male and female offspring, differential effects of brood sex composition on their survival and a difference in the capacity of parents to rear males and females. All of these processes need to be taken into account in attempting to understand offspring sex ratios.