Female and male Leach's Storm Petrels () pursue different foraging strategies during the incubation period
Reproduction in procellariiform birds is characterized by a single egg clutch, slow development, a long breeding season and obligate biparental care. Female Leach's Storm Petrels , nearly monomorphic members of this order, produce eggs that are between 20 and 25% of adult body weight. We tested whether female foraging behaviour differs from male foraging behaviour during the ~ 44-day incubation period across seven breeding colonies in the Northwest Atlantic. Over six breeding seasons, we used a combination of Global Positioning System and Global Location Sensor devices to measure characteristics of individual foraging trips during the incubation period. Females travelled significantly greater distances and went farther from the breeding colony than did males on individual foraging trips. For both sexes, the longer the foraging trip, the greater the distance. Independent of trip duration, females travelled farther, and spent a greater proportion of their foraging trips prospecting widely as defined by behavioural categories derived from a Hidden Markov Model. For both sexes, trip duration decreased with date. Sex differences in these foraging metrics were apparently not a consequence of morphological differences or spatial segregation. Our data are consistent with the idea that female foraging strategies differed from male foraging strategies during incubation in ways that would be expected if females were still compensating for egg formation.
Sex ratio of Western Bluebirds is mediated by phenology and clutch size
Mothers may produce more of one sex to maximize their fitness if there are differences in the cost of producing each sex or there are differences in their relative reproductive value. Breeding date and clutch size are known to influence offspring sex ratios in birds through sex differences in dispersal, social behaviours, differential mortality, and available food resources. We tested if breeding date, clutch size and drought conditions influenced offspring sex ratios in a sexually size-monomorphic species, the Western Bluebird, by interrogating a 21-year dataset. After controlling for differential mortality, we found that hatch dates late in the breeding season were associated with the production of more females, suggesting that the value of producing males declines as the breeding season progresses. When clutch size was taken into account, small clutches yielded significantly more females late in the breeding season compared to the early and middle parts of the breeding season that produced significantly more males. Large clutches early in the season tended to produce more females, although this was not significant. Drought severity was not correlated with sex ratio adjustment. We propose and discuss several explanations for these patterns, including male offspring, but not female offspring, acting as helpers, increased female nestling provisioning late in the breeding season, differences in food abundance, and egg-laying order. Future work will help to uncover the mechanisms leading to these patterns. Identifying patterns and mechanisms of sex ratio skew from long-term datasets is important for informing predictions regarding life-history trade-offs in wildlife populations.
Urban gulls adapt foraging schedule to human-activity patterns
Numerous animals are able to adapt to temporal patterns in natural food availability, but whether species living in relatively novel environments such as cities can adapt to anthropogenic activity cycles is less well understood. We aimed to assess the extent to which urban gulls have adapted their foraging schedule to anthropogenic food source fluctuations related to human activity by combining field observations at three distinct urban feeding grounds (park, school and waste centre) with global positioning system (GPS) tracking data of gulls visiting similar types of feeding grounds throughout the same city. We found that the birds' foraging patterns closely matched the timing of school breaks and the opening and closing times of the waste centre, but gull activity in the park appeared to correspond to the availability of natural food sources. Overall, this suggests that gulls may have the behavioural flexibility to adapt their foraging behaviour to human time schedules when beneficial and that this trait could potentially enable them to thrive in cities.
Parental care in the Small Tree Finch in relation to parasitism and environmental factors
The parental food compensation hypothesis suggests that parents may compensate for the negative effects of parasites on chicks by increased food provisioning. However, this ability differs widely among host species and may also depend on ecological factors such as adverse weather conditions and habitat quality. Although weed management can improve habitat quality, management measures can bring about a temporary decrease in food availability and thus may reduce parents' ability to provide their nestlings with enough energy. In our study we investigated the interaction of parasitism and weed management, and the influence of climate on feeding rates in a Darwin's tree finch species, which is negatively impacted by two invasive species. The larvae of the invasive parasitic fly ingest the blood and body tissues of tree finch nestlings, and the invasive Blackberry affects one of the main habitats of Darwin's tree finches. We compared parental food provisioning of the Small Tree Finch in parasitized and parasite-free nests in three different areas, which differed in invasive weed management (no management, short-term and long-term management). In a parasite reduction experiment, we investigated whether the Small Tree Finch increases food provisioning rates to nestlings when parasitized and whether this ability depends on weed management conditions and precipitation. Our results provide no evidence that Small Tree Finches can compensate with additional food provisioning when parasitized with . However, we found an increase in male effort in the short-term management area, which might indicate that males compensate for lower food quality with increased provisioning effort. Furthermore, parental food provisioning was lower during rainfall, which provides an explanation for the negative influence of rain on breeding success found in earlier studies. Like other Darwin's finches, the Small Tree Finch seems to lack the ability to compensate for the negative effects of parasitism, which is one explanation for why this invasive parasite has such a devastating effect on this host species.
Genetic isolation in an endemic African habitat specialist
The Chestnut-banded Plover is a Near-Threatened shorebird species endemic to mainland Africa. We examined levels of genetic differentiation between its two morphologically and geographically distinct subspecies, in southern Africa (population size 11 000-16 000) and in eastern Africa (population size 6500). In contrast to other plover species that maintain genetic connectivity over thousands of kilometres across continental Africa, we found profound genetic differences between remote sampling sites. Phylogenetic network analysis based on four nuclear and two mitochondrial gene regions, and population genetic structure analyses based on 11 microsatellite loci, indicated strong genetic divergence, with 2.36% mitochondrial sequence divergence between individuals sampled in Namibia (southern Africa) and those of Kenya and Tanzania (eastern Africa). This distinction between southern and eastern African populations was also supported by highly distinct genetic clusters based on microsatellite markers (global = 0.309, GST' = 0.510, = 0.182). Behavioural factors that may promote genetic differentiation in this species include habitat specialization, monogamous mating behaviour and sedentariness. Reliance on an extremely small number of saline lakes for breeding and limited dispersal between populations are likely to promote reproductive and genetic isolation between eastern and southern Africa. We suggest that the two Chestnut-banded Plover subspecies may warrant elevation to full species status. To assess this distinction fully, additional sample collection will be needed, with analysis of genetic and phenotypic traits from across the species' entire breeding range.
Co-occurrence of birds and bats in natural nest-holes
Assessing the effects of repeated handling on the physiology and condition of semi-precocial nestlings
Repeated exposure to elevated levels of glucocorticoids during development can have long-term detrimental effects on survival and fitness, potentially associated with increased telomere attrition. Nestling birds are regularly handled for ecological research, yet few authors have considered the potential for handling-induced stress to influence hormonally mediated phenotypic development or bias interpretations of subsequent focal measurements. We experimentally manipulated the handling experience of the semi-precocial nestlings of European Storm Petrel to simulate handling in a typical field study and examined cumulative effects on physiology and condition in late postnatal development. Neither baseline corticosterone (the primary glucocorticoid in birds), telomere length nor body condition varied with the number of handling episodes. The absence of a response could be explained if Storm Petrels did not perceive handling to be stressful or if there is dissociation of the hypothalamic-pituitary-adrenal axis from stressful stimuli in early life. Eliciting a response to a stressor may be maladaptive for cavity-dwelling young that are unable to escape or defend themselves. Furthermore, avoiding elevated overall glucocorticoid exposure may be particularly important in a long-lived species, in which accelerated early-life telomere erosion could impact negatively upon longevity. We propose that the level of colony-wide disturbance induced by investigator handling of young could be important in underlining species-specific responses. Storm Petrel nestlings appear unresponsive to investigator handling within the limits of handling in a typical field study and handling at this level should not bias physiological and morphological measurements.
Maximizing dietary information retrievable from carcasses of Great Cormorants using a combined morphological and molecular analytical approach
Avian carcasses can provide important information on the trophic ecology of birds. Usually, the number of carcasses available for examination is limited and therefore it is important to gain as much dietary information per specimen as possible. In piscivorous birds and raptors, the stomach has been the primary source of dietary information, whereas the gut (intestine) has so far been neglected as it usually contains only a few morphologically identifiable hard parts of prey. Molecular approaches have the potential to retrieve dietary information from the gut, although this has not yet been verified. As well as identifying the prey, it is important to estimate any secondary predation to avoid food web errors in dietary analyses. The assignment of accidentally consumed prey is notoriously difficult regardless of the prey identification approach used. In the present study, morphological and molecular analyses were, for the first time, combined to maximize the dietary information retrievable from the complete digestive tract of Great Cormorants . Moreover, a novel approach based on predator-prey size ratios was applied to these piscivorous birds to minimize the number of samples that might contain secondarily predated prey. The stomach contents of the examined birds were found to provide the most dietary information when morphological and molecular analyses were used in combination. However, compared with the morphological approach, the molecular analysis increased the number of fish species detected by 39%. The molecular approach also permitted the identification of fish DNA in the Cormorant guts. Predator-prey size ratios derived from morphological analysis of fish hard parts can reduce the incidence of potential confounding influence of secondarily predated prey by 80%. Our findings demonstrate that a combination of morphological and molecular approaches maximizes the trophic information retrievable from bird carcasses.
The sight of an adult brood parasite near the nest is an insufficient cue for a honeyguide host to reject foreign eggs
Hosts of brood-parasitic birds typically evolve anti-parasitism defences, including mobbing of parasitic intruders at the nest and the ability to recognize and reject foreign eggs from their clutches. The Greater Honeyguide is a virulent brood parasite that punctures host eggs and kills host young, and accordingly, a common host, the Little Bee-eater frequently rejects entire clutches that have been parasitized. We predicted that given the high costs of accidentally rejecting an entire clutch, and that the experimental addition of a foreign egg is insufficient to induce this defence, Bee-eaters require the sight of an adult parasite near the nest as an additional cue for parasitism before they reject a clutch. We found that many Little Bee-eater parents mobbed Greater Honeyguide dummies while ignoring barbet control dummies, showing that they recognized them as a threat. Surprisingly, however, neither a dummy Honeyguide nor the presence of a foreign egg, either separately or in combination, was sufficient to stimulate egg rejection.
Ontogenic differences in sexual size dimorphism across four plover populations
Sexual size dimorphism (SSD) among adults is commonly observed in animals and is considered to be adaptive. However, the ontogenic emergence of SSD, i.e. the timing of divergence in body size between males and females, has only recently received attention. It is widely acknowledged that the ontogeny of SSD may differ between species, but it remains unclear how variable the ontogeny of SSD is within species. Kentish Plovers and Snowy Plovers are closely related wader species that exhibit similar, moderate (. 4%), male-biased adult SSD. To assess when SSD emerges we recorded tarsus length variation among 759 offspring in four populations of these species. Tarsus length of chicks was measured on the day of hatching and up to three times on recapture before fledging. In one population (Mexico, Snowy Plovers), males and females differed in size from the day of hatching, whereas growth rates differed between the sexes in two populations (Turkey and United Arab Emirates, both Kentish Plovers). In contrast, a fourth population (Cape Verde, Kentish Plovers) showed no significant SSD in juveniles. Our results suggest that adult SSD can emerge at different stages of development (prenatal, postnatal and post-juvenile) in different populations of the same species. We discuss the proximate mechanisms that may underlie these developmental differences.
Widespread supplementary feeding in domestic gardens explains the return of reintroduced Red Kites to an urban area
Reintroductions are commonly used to mitigate biodiversity loss. One prominent example is that of the Red Kite , a charismatic raptor of conservation concern. This species has been reintroduced across the UK over the last 25 years following its near extinction after centuries of persecution. The species was not expected to recolonize urban areas; its historical association with human settlements is attributed to scavenging on human waste and refuse, a resource now greatly reduced on the streets of modern European cities. However, the species has become a common daytime visitor to a large conurbation centred on the town of Reading, southern England, approximately 20 km from the first English reintroduction site. Given a near-absence of breeding and roost sites, we investigated foraging opportunities and habitat associations that might explain use by Red Kites of this urban area. Surveys of discarded human foods and road-kill suggested that these could support at most 13-29 Kites per day. Face-to-face surveys of a cross-section of residents revealed that 4.5% (equivalent to 4349 households) provided supplementary food for Red Kites in their gardens. Using estimates of per-household resource provision from another study, we calculated that this is potentially sufficient to feed 142-320 Kites, a substantial proportion of the total estimated to visit the conurbation each day (between 140 and 440). Road transects found positive associations between Red Kites and residential areas. We suggest that the decision made by thousands of householders to provide supplementary food for Red Kites in their gardens is the primary factor explaining their daytime abundance in this urban area.
Isotopic analysis of island House Martins indicates marine provenance of nutrients
The presence of one of the largest colonies of House Martins in Europe on the small island of Stora Karlsö, Sweden, led us to investigate the source of their food by analysis of stable isotopes of carbon and nitrogen. Carbon isotopic values of House Martin nestlings were the same as those of Common Guillemot nestlings fed on marine fish, but differed from local Collared Flycatcher nestlings fed on woodland insects. We infer that these House Martins fed their chicks almost exclusively on insects that had used nutrients derived from seabirds, indicating a dependence on the presence of a large seabird colony. We suggest by extension that some populations of island passerines of high conservation importance may also be dependent on nutrient subsidies from seabird colonies.
Sex-specific costs of reproduction in Eastern Bluebirds Sialia sialis
In species with bi-parental care, individuals must partition energy between parental effort and mating effort. Typically, female songbirds invest more than males in reproductive activities such as egg-laying and incubation, but males invest more in secondary sexual traits used in attracting mates. Animals that breed more than once within a season must also allocate time and energy between first and subsequent breeding attempts and between current and future breeding seasons. To investigate strategies of reproductive investment by males and females and the consequences of such strategies, we manipulated the size of broods of Eastern Bluebirds Sialia sialis. Pairs with enlarged first broods were less likely to produce a second clutch or took longer to initiate one than pairs with reduced broods. After rearing enlarged broods, females were less likely than males to survive to the following year. Although plumage coloration is a sexually selected trait in Eastern Bluebirds that is influenced by nutritional stress, we did not detect an effect of brood-size manipulation on female coloration. Past research, however, demonstrates that, in males, plumage colour is negatively affected by increasing brood size. We suggest that there are sex-specific strategies of reproductive investment in Eastern Bluebirds, and that researchers should incorporate measures of residual reproductive value in studies of life-history evolution.
Declining winter survival and fitness implications associated with latitudinal distribution in Norwegian Greylag Geese
The Norwegian Greylag Goose population has been increasing steadily over the past few decades, causing increasing nuisance in terms of agricultural crop damage. This, in combination with the importance of Greylags as a hunting target, has called for demographic estimates for the population to assist in management decisions. To this end, we analysed long-term mark-recapture data using Cormack-Jolly-Seber models embedded in program MARK to obtain survival estimates for the population. No sex-specific difference, or age effect on survival after juveniles had completed their first migration (3 months of age), was evident. Mean first-year survival was reported as 0.485 and annual survival of older birds as 0.700. On a monthly basis, survival in Greylags during summer and winter was very similar over the study period. A significant linear decline in winter survival from 0.909 to 0.807 was, however, apparent during the study period. Over the second half of the study (1994-2002), summer survival was about 3% lower than in the first half (1986-94) but no linear relationship was evident. We found a significant inverse relationship between Greylag survival during summer and latitudinal distribution in Norway. A similar relationship was evident between survival and annual bag numbers. The changes in adult survival observed in this study are likely to have had a substantial impact on the growth rate of the Norwegian Greylag population.