Family-based territoriality vs flocking in wintering common cranes Grus grus

We describe winter territoriality in common cranes, Grus grus , a long-lived migrant species with long-lasting pair bonds and parental care extending throughout the winter. Cranes are territorial in the breeding season, and usually gregarious during migration and wintering. Only 2% of the families present in our study area were territorial, all other families foraged in flocks with immatures and adult pairs. Territorial pairs defended the same winter territory year after year, but only when they had offspring. They were gregarious otherwise. The average breeding success measured throughout several years was higher in territorial pairs. Winter territories were small (0.7 km² on average), but included a higher diversity of habitats than the areas visited by gregarious birds (11.7 km² on average). Adults of territorial families showed longer vigilance times, and lower food intake rates than did adults in flocks, which were compensated with a longer time spent foraging per day. The accumulated daily food intake did not differ between adults in flocks and in families. We suggest that winter territoriality is a facultative strategy, conditioned by parental experience and habitat availability.     

Effects of supplementary feeding on territoriality, breeding success and survival of pheasants

1. The effects of supplementary food in spring on subsequent pheasant breeding in an intensively farmed area in southern England were assessed by a large-scale, replicated field experiment.
2. Territorial cock pheasants were counted in April, and the breeding success and survival of radio-tagged hens were monitored in six 1-km² plots during 1994 and 1995. Total numbers of young reared and post-breeding pheasant densities were found by August counts. In 1994, three randomly selected plots were supplied with wheat grains via hoppers along woodland edges and hedgerows. The other three plots acted as controls and the treatments were reversed in 1995.
3. The density of cock territories increased significantly in food-supplemented plots (44 ± 8 km−2) in relation to control plots (29 ± 8 km−2), and the presence of hoppers significantly affected the locations of territories. However, similar proportions of territorial males acquired harems in the control and food-supplemented plots. Hen density did not increase and, consequently, the mean harem size was significantly lower with supplementary feeding.
4. Hens given supplementary food did not nest earlier and the number of nests initiated, clutch sizes and the proportion of successful nests did not differ significantly from those of controls. However, hens supplied with supplementary food re-nested significantly more quickly following the loss of a nest or brood.
5. Radio-tagged hens did not rear significantly more young with supplementary feeding. Hen survival was unchanged and post-breeding pheasant densities were no higher.
6. On present evidence, spring feeding cannot be advocated as a management technique to improve the breeding success of pheasants surviving the winter.     

THE SOCIAL BEHAVIOUR OF THE WHITE WAGTAIL MOTACILLA ALBA ALBA WINTERING IN ISRAEL

White Wagtails Motacilla alba wintering in Israel are partly territorial, mostly around human habitations, and partly live in flocks around temporary food sources. Individual birds may spend part of the season (or the day) in the territory and the other part with a flock. Experiments with artificial distribution of food, in a natural habitat, brought about a change from flocking to territorial behaviour. Preliminary observations suggest that in the natural situation the pattern of food distribution may be the proximate factor which regulates the birds' behaviour, by determining whether they have to fight for their food. Pairs are formed on many territories, and may last for long or short periods. Pair formation is initiated by females, who when seeking food appease the territorial males and are able to stay with them on their territories. Females also manifest territorial behaviour. Although pairing in winter territories is similar, in the behaviour involved, to sexual pairing, it is very unlikely that winter pairing continues into, or influences, pairing for breeding. It is suggested that the function of winter pair formation is that it allows two birds to exploit one territory, and that the main advantage is to the female which is the subordinate bird of the pair. This kind of pair-formation may be analogous to non-breeding group territories reported in some other birds.     

Characteristics of Golden‐winged Warbler territories in plant communities associated with regenerating forest and abandoned agricultural fields

In the Appalachian portion of their breeding range, Golden‐winged Warblers (Vermivora chrysoptera) nest in shrubland and regenerating forest communities created and maintained by disturbance. Because populations of Golden‐winged Warblers have exhibited precipitous declines in population throughout their Appalachian breeding range, management activities that create or maintain early successional habitat are a priority for many natural resource agencies and their conservation partners. Within these early successional habitats, however, additional information is still needed concerning the relative importance of different vegetation features in selection of breeding territories by Golden‐winged Warblers. Our objective, therefore, was to use logistic regression to estimate the probability of territory‐level occupancy by Golden‐winged Warblers in north‐central Pennsylvania at two sites, each with its own early successional community, based on vegetation characteristics. Our communities were composed of shrublands and regenerating forest sites resulting from two disturbances: agriculture and forest fire. Despite differences in vegetation structure, portions of both study areas (regenerating forest and old field) supported territorial Golden‐winged Warblers. Probability of territory occupancy by Golden‐winged Warblers increased with percent blackberry (Rubus) cover in the regenerating forest community, and decreased as basal area and distance to microedge increased (i.e., as vegetation patchiness decreased) in both communities. These habitat features have also been found to influence other aspects of Golden‐winged Warbler breeding ecology such as nest‐site selection, pairing success, and territory abundance. Vegetation features influencing Golden‐winged Warbler territory establishment can differ among shrubland and regenerating forest communities, and management decisions and outcomes may be affected by these differences. Our study provides a starting point for a more comprehensive hypothesis‐driven occupancy survey to investigate features of the territories of Golden‐winged Warblers across a broader geographic range and in different vegetation communities.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Habitat type determines the effects of disturbance on the breeding productivity of the Dartford Warbler Sylvia undata

Numerous studies have examined the causes and impacts of human disturbance on birds, but little is known about how these impacts vary among habitats. This is of applied importance both for predicting bird responses to changes in disturbance and in planning how to reduce disturbance impacts. The Dartford Warbler Sylvia undata , a key heathland breeding species, occupies territories in a range of heathland types. Three territory habitat groups were identified: heather-dominated territories, heather territories with significant areas of European Gorse Ulex europaeus and territories containing Western Gorse U. gallii . Productivity was significantly affected by the timing of breeding in all habitats, but disturbance only appeared to have a significant impact on the productivity of birds in heather territories. Disturbance events in heather territories delayed breeding pairs for up to 6 weeks. This significantly decreased both the number of successful broods raised and the average number of chicks fledged per pair. Nests situated close to territory boundaries in heather territories, with high numbers of disturbance events, were more likely to fail outright. It was determined that an average of between 13 and 16 people passing through a heather territory each hour would delay breeding pairs sufficiently to prevent multiple broods.     

Competitive Asymmetries in the Use of Supplementary Food by the Endangered Iberian Lynx (Lynx pardinus)

Background

As a conservation tool, supplementary feeding programs may be directed to specific individuals or sectors of the target population whose productivity or survival is thought to be limited by food scarcity. However, the use of supplemental food by different sex and age classes has received little attention. We studied individual variation in the access of the endangered Iberian lynx (Lynx pardinus) to supplementary food.

Methodology/Principal Findings

From 5349 pictures taken with automatic cameras placed in 25 feeding stations, we identified 28 individuals whose sex and age were known. All individuals known to live in areas subjected to supplementation regularly visited feeding stations. Food consumption was not proportional to expected variations in energy demand within sex and age classes. Food consumption by males was higher than by females, and increased with age, in agreement with a despotic distribution. Food consumption also increased with lynx body mass, and this pattern held for individuals sharing the same breeding territories. The access of inferior competitors increased with the number of feeding stations available within lynx territories.

Conclusions/Significance

All lynx exposed to food supplementation made a regular use of extra food but individuals predicted to be competitively dominant visited stations more frequently than subordinates of the same breeding territory. Our results suggest that insufficient provision of supplementary food could restrict the access of juveniles, or even adult females, to feeding stations. Limited consumption by these target individuals may compromise the efficiency of the supplementary feeding programme at the population level, in endangered species that, as the Iberian lynx, exhibit marked sexual dimorphism in body size.     

Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

Territory Area as a Determinant of Mating Systems

Territoriality is an integral part of breeding behavior in manyanimals. Because the reproductive success of territorial malesis often limited by access to females, breeding males shouldbehave as "area maximizers" when the basis of female choiceis either the abundance of resources within the territory orterritory area per se. Being reproductively more limited byenergy, territorial females should be "energy maximizers." Aseries of simple analytical models of territory area for suchforagers is developed to explore how changes in local food productionand/or local competitor density affect both the probabilityof a territorial male securing more than one mate (polygyny)and the probability of his and his mates' reproductive successincreasing. Two cases are modeled (only males territorial vs.both sexes territorial), each for various sets of assumptionsregarding interactions between food production, feeding efficiency,and competitor density. Concurrent responses in territory area,territory food reserves, net energy gain, and time budgetingprovide testable sets of predictions for each scenario. Whereonly males are territorial (Case I), changes in food productioncan have different (indeed, opposite) effects upon an individualmale's probability of becoming polygynous, depending upon whetherthe basis of female choice is the abundance of food within theterritory or another factor positively correlated with territoryarea. Increases in competitor density usually decrease the probabilityof polygyny regardless of the basis of female choice. Whereboth sexes are territorial and territories overlap intersexually(Case II), the mating system becomes a function of the numberof female territories within each male's territory, which varieswith the ratio of male to female territory areas. In this case,the probability of polygyny occurring will increase if foodproduction for both sexes increases without concurrent increasesin competitor density, and will decrease if competitor densityfor both sexes increases without concurrent increases in feedingefficiency. Few data are presently available to test eitherthese general predictions or numerous sets of secondary predictionstabulated in the text. Available evidence is largely consistentwith the models, but mostly circumstantial. This is becausethe predictions of these and other models of territory areaare strongly assumption dependent, and few published studieshave investigated these assumptions. These analyses demonstratethat to accurately assess the mechanisms by which environmentalfactors affect territory area, and thus mating systems, testsof the underlying assumptions of models are essential.     

Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

Territory occupancy by common loons in response to disturbance,habitat, and intraspecific relationships

Structure and distribution of animal territories are driven by a variety of environmental and demographic factors. A peninsular population of common loons (Gavia immer) nests on lakes in northwestern Montana, but does not occupy all apparently suitable breeding territories, suggesting unexplained limitations on population growth. To evaluate territorial dynamics of breeding loons in Montana, we created and tested occupancy models that evaluated the hypothesized effects of disturbance, habitat, and intraspecific relationships on territory occupancy by common loons in Montana from 2003 to 2007. Model-averaged results indicated that the abundance of feeding lakes within 10 km (i.e., forage quality) and the number of territorial pairs within 10 km (i.e., density of loons) were equally supported and related to probabilities of occupancy. We found substantial support that the population was in a state of equilibrium, with the numbers of occupied territories stable in time, but not space. We also found that density of territorial pairs was related to the likelihood that an existing territory would be abandoned, but did not influence the establishment of new territories, suggesting the presence of territorial pairs could be a stronger indicator of territory quality to loons than physical lake characteristics. Our index of human disturbance was not well-supported compared to other factors. Our results suggest management for stable or growing loon populations could be achieved using long-term monitoring and protection of occupied territorial lakes and nearby feeding lakes, because these factors most influenced the probability of occupancy of surrounding lakes. © 2011 The Wildlife Society.     

Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

THE ECOLOGY OF THE GREENISH WARBLER PHYLLOSCOPUS TROCHILOIDES IN ITS WINTER QUARTERS

The Greenish Warbler is an abundant entirely insectivorous Pafaearctic migrant wintering in the tropical deciduous forests of India and east Asia. A population was studied at one locality in the eastern Ghats of south India over one winter, with a short return the following winter. Individuals maintain, and annually re-occupy, territories throughout their seven or eight months stay. A general account of territorial behaviour is given.
There is much evidence that food is critically short, particularly during the months of December to February and March, which coincides with the end of the dry season. Because of the short duration of the study, there are shortcomings in data presented. Nevertheless there is evidence that territories vary severalfold in quality and individual birds' behaviour and survival probability are correlated with this. It is suggested that territorial behaviour is the best individual strategy for long-term resource management. The presence of food competitors can alter this behaviour, much as changes in food dispersion are known to. The ability to control return time to a patch as a subsidiary function for territoriality is briefly discussed.     

Avian productivity in urban landscapes: a review and meta-analysis

There is an urgent need to thoroughly review and comprehend the effects of urbanization on wildlife in order to understand both the ecological implications of increasing urbanization and how to mitigate its threat to biodiversity globally. We examined patterns in comparative productivity of urban and non-urban passerine birds, using published estimates from paired comparisons, and by reviewing and developing explanations in terms of resources, competitors, predators and other specifically urban environmental factors. The most consistent patterns were for earlier lay dates, lower clutch size, lower nestling weight and lower productivity per nesting attempt in urban landscapes; these were supported by a formal meta-analysis. Nest failure rates did not show consistent patterns across the species considered. We suggest that food availability is a key driver of differences in passerine demography between landscapes. In urban habitats, human-provided food may improve adult condition over winter, leading to earlier lay dates and, in some species, to higher survival and higher breeding densities, but paucity of natural food may lead to lower productivity per nesting attempt. We demonstrate that additional comparative research is needed on a wider range of species, on the effects of natural and human-provided food availability, and on the differences in survival and dispersal between urban and non-urban populations. Importantly, better-targeted research and monitoring is needed in areas that are at greatest threat from urbanization, especially in the developing world.     

Complex effects of habitat loss on Golden Eagles Aquila chrysaetos

The development of commercial forests presents potential threats to large raptors that rely on prey caught in open country. We examined the effect of afforestation of breeding habitat used by a population of Golden Eagles Aquila chrysaetos in Scotland where, over the last 50 years, extensive stands of exotic conifers have been planted. Using data for 31 years on territory occupancy and breeding success, together with spatiotemporally dynamic mapping of forest cover and predicted areas of territory‐use in a Geographical Information System, we examined relationships between forest cover and Eagle ecology at landscape and individual territory scales. Several territories were abandoned during the earliest phases of forest planting, but relatively few were apparently lost to later plantings. Territories with poorer breeding productivity appeared to be more vulnerable to abandonment than territories with better breeding productivity. At the landscape scale, temporal differences in breeding productivity were negatively related to the extent of forest cover, although productivity of individual territories showed no clear relationship with forest cover. Several territories with less than a 5% increase in forest cover experienced reduced productivity; however, territories least constrained by neighbouring pairs of Eagles showed an increase in productivity. Territories experiencing the greatest increases in forest cover showed a greater use of spatially separated nest‐sites by occupying pairs. Hence, pairs that were less constrained by neighbours appeared to compensate for loss of open habitat by shifting their territory‐use, whereas pairs that were more constrained could not compensate for open habitat loss and suffered reduced productivity (and, probably in some cases, abandoned the territory). We suggest that simple guidelines based on the extent and locations of habitat loss are inadequate when predicting effects on large territorial raptors such as Golden Eagles. Consideration should also be given to the ‘quality’ of a territory or occupying pair, as well as the extent to which territory‐use is constrained by neighbouring pairs or other ‘unsuitable habitat’ which may have been affected by previous episodes of open habitat loss.     

Space,density and extra-pair matings have opposing impacts on male and female reproductive success

Many territorial species have a mating system characterized by males establishing home ranges in the breeding grounds prior to females, resulting in males competing for territories and females choosing a mate upon their arrival. It remains unknown, however, how the outcomes of decisions surrounding territory establishment and mate choice are influenced by the spatial configuration of the breeding grounds. We use a spatially explicit, individual-based model to investigate the sex-specific effects of these decisions on reproductive success. In our model, males that arrive earlier obtain higher quality territories and improve their chances for extra-pair copulations. Females can choose their mate to maximize the quality of the male or to attempt to minimize the density of other females near their nesting site to avoid competition. Females therefore face a tradeoff between high-density regions around high-quality males and low-quality males in areas of low competition. Our model predicts a negative correlation between male and female reproductive success under a wide range of conditions when the majority of the territories are on the margins of the breeding area. Most notably, this sexual conflict arises as an edge effect suggesting that fragmentation of breeding habitats could impact the consequences of mate choice in many species with territorial breeding habits.     

Home-ranges and patterns of spatial use in territorial Bonelli's Eagles Aquila fasciata

Bonelli's Eagle Aquila fasciata is one of the rarest birds of prey in Europe, where it has suffered a significant decline in recent decades. We present information on the home-ranges and spatial parameters of 18 Bonelli's Eagles radiotracked in 2002–2006 in Catalonia (northeast Spain) and describe the home-range probability kernel, distances moved, breeding area eccentricity, territorial overlap, nearest neighbour distance and breeding site fidelity, and assess the influence of sex, breeding status, season and geographical area on these parameters. Median home-range according to the minimum convex polygon (MCP) and 95% kernel were 50.3 and 36.1 km², respectively. The median breeding area eccentricity was 1477 m. There was considerable overlap in the home-range of both sexes within pairs (MCP: 71.4% and 95% kernel: 98.5%), indicating close pair bonding and similar foraging patterns. Overlap in home-ranges of up to 15% between neighbouring individuals also occurred and was positively related to breeding pair density. There was no difference in spatial parameters between sexes or with breeding status, but during the non-breeding season Eagles had larger home-ranges and stayed further from nests. The high consistency across birds suggests a pattern of spatial use that is characteristic of this species. The high level of use of breeding areas and their surroundings (50% kernel) throughout the year makes it important that these areas be protected from human disturbance. Additionally, it is necessary that heavily used areas away from nesting sites, which are used for foraging and roosting, are identified, protected and managed in a sustainable fashion.     

FOOD RESOURCES, TERRITORY DENSITY AND REPRODUCTIVE SUCCESS OF AN ISLAND SILVEREYE POPULATION ZOSTEROPS LATERALIS

We examined the relationships between food resources, territory density and some breeding parameters (individual reproductive success and parental age) of a Capricorn Silvereye population during two years' detailed study on Heron Island, Great Barrier Reef. The territorial behaviour of the Silvereyes included dawn song. aggression and distinctive vocalizations. During both years, about 85% of pairs occupied territories although the population sizes differed. The dispersion pattern of territories was not regular and was correlated with the density of figs and human food scraps. Nestlings were fed more figs as they grew, and the parents foraged in fig trees outside their territories. However, only about 60%, of territorial pairs fledged young. Pairs which fledged most young were older, fed more insects to their nestlings, and nested in areas containing more fruiting fig trees per territory. We conclude that food resources were in short supply, and that access to fig trees provided breeding birds with a quick energy source while they searched for insect protein for the young. The data are consistent with the predictions of the ‘sufficient resource‘ hypothesis of the adaptive significance of territoriality but not with its assumption that the principal benefit is the food within the territory.     

Persistence and local extinction of lion prides in the Ngorongoro Crater, Tanzania

African lions (Panthera leo) live in social groups (prides) that exhibit group territorial behavior. Pride persistence is expected to depend on its ability to compete against neighboring prides as well as on average rates of reproduction and survival, thus providing a meaningful measure of intergenerational reproductive success. We used Akaike’s Information Criterion (AIC) to select the best approximating models explaining how demographic variables influenced pride persistence during a 30-year period in the Ngorongoro Crater, Tanzania, and identified landscape factors affecting those demographic variables. Pride persistence to 10 years depended on adult female density (pride size) and cub productivity (the ultimate source of new females). Average age of adult females had a weakly positive influence on pride persistence, while the effect of female mortality was weakly negative. Adult female mortality increased with disease epidemics and in territories with high human disturbance. Cub productivity was highest in territories closest to rivers and only slightly higher near swamps, and also high in areas of higher vegetative cover and high human use. No landscape variable significantly affected female density. The growth and population size of the Crater lions was closely linked to demographic performance of individual prides, while territorial behavior played a key role in mediating the interactive effects of landscape and demography.     

Causes and consequences of pre‐laying weight gain in a food‐caching bird that breeds in late winter

Animals that reside at high latitudes and altitudes year‐round often use cached food to survive over the winter months, but a few species also rely on stored food to sustain them during the breeding season when the nutritional requirements of females are higher than normal. Gray jays Perisoreus canadensis rely on perishable cached food during the winter and females begin breeding in late winter when fresh food is rarely available. To examine pre‐laying patterns of weight gain, as well as the causes and consequences of weight gain among individuals, we weighed females regularly throughout the pre‐laying period. Females began increasing their weight approximately nine days prior to their first egg date, and on average increased their body weight by 25%, which is on par with other bird species that rely on non‐cached food. Final pre‐laying weight was positively influenced by the percent of conifers on territories, providing some support for previous results showing that coniferous trees are better able to preserve cached food. We also found that both final pre‐laying weight and the rate of weight gain were positively related to female age, supporting the hypothesis that female caching ability improves with age. With increasing final weight, females tended to lay larger clutches and hatched more nestlings, despite the fact that final weight was not influenced by weight at the beginning of the weighing period. Our results confirm that gray jays are able to reach breeding condition while relying primarily on food stored before winter, and suggest a novel mechanism by which habitat‐mediated carry‐over effects and female age may influence reproductive performance in a food‐caching animal.