Breeding territory fidelity in a partial migrant, the American dipper Cinclus mexicanus

American dipper Cinclus mexicanus populations are frequently composed of resident individuals that occupy permanent territories year round and migratory individuals that overwinter with residents but migrate to breeding territories on higher elevation creeks each spring. Between 1999 and 2004 we examined how migratory strategy (resident/migratory) and sex differences influence breeding territory fidelity of American dippers occupying the Chilliwack River watershed, British Columbia, Canada. Counter to expectation we found that the migratory strategy of American dippers did not influence whether birds breeding in one year were found on their former breeding territory in the next. Migratory strategy also did not affect the probability that known surviving dippers occupied the same breeding territory in the following year. Males and females were equally likely to be found on their former territory in the following year (females 43%, males 41%) and known survivors had similar levels of breeding territory fidelity (females 74%, males 68%). However, breeding territory fidelity of males and females varied in response to different factors. Surviving female dippers were more likely to be found on their former breeding territory in the subsequent year following a successful breeding attempt than an unsuccessful breeding attempt. Prior reproductive performance did not influence whether surviving male dippers were found on their former breeding territory. Male dippers were more likely to be found on their former territory and, if they survived, have higher breeding territory fidelity when their mate also returned to that same territory. Mate retention also influenced whether females were found on their former territory in the following year but had no effect on the breeding territory fidelity of known survivors. We argue that sex‐specific dispersal decision rules in American dippers are driven by sex differences in the predictability of breeding performance between years and sex differences in how mate retention influences subsequent reproductive success.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

Sex-biased dispersal: a result of a sex difference in breeding site availability

Sex-biased dispersal is often explained by assuming that the resource-defending sex pays greater costs of moving from a familiar area. We hypothesize that sex-biased dispersal may also be caused by a sex bias in breeding site availability. In avian resource-defense mating systems, site availability is often more constrained for females: males can choose from all vacant sites, whereas females are restricted to sites defended by males. Using data on breeding dispersal of a migratory passerine, we show that average number of available breeding options and availability of the previous year's territory was greater for males than females. The female bias in site unavailability may explain the female bias in probability of breeding dispersal because there was no sex bias in dispersal among birds with their previous year's territory available. We suggest that sex biases in the availability of breeding options may be an important factor contributing to observed variation in sex-biased dispersal patterns.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

Patterns in departure phenology and mass gain on African non‐breeding territories prior to the Sahara crossing in a long‐distance migrant

Afro‐Palaearctic migrants are declining to a greater degree than other European species, suggesting that processes occurring in Africa or on migration may be driving these trends. Constraints on food availability on the wintering grounds may contribute to these declines but little is known about when and where these resource constraints may occur. Sufficient resources are particularly important prior to spring migration, when migrants must cross the Sahara Desert. We examined mass gain and departure phenology in a long‐distance Palaearctic passerine migrant to determine the degree to which pre‐migratory fattening occurs in their long‐term non‐breeding territories in the Guinea Savannah region of Africa. We monitored 75 Whinchats Saxicola rubetra for departure from their non‐breeding territories in one spring, and analysed mass data of 377 Whinchats collected over three non‐breeding seasons plus 141 migrating Whinchats caught in April over 8 years, all within the same few square kilometres of human‐modified Guinea Savannah in central Nigeria. Whinchats left their winter territories throughout April, with males departing on average 8 days earlier than females. However, there was no evidence that time of departure from territory was linked to age, body size or mass at capture. Whinchats departed their territories with a predicted mass of 16.8 ± 0.3 g, considerably less than the c. 24 g required for the average Whinchat to cross the Sahara directly. Comparing departure dates with arrival dates in southern Europe showed a discrepancy of at least 2 weeks, suggesting that many Whinchats spend considerable time on pre‐migratory fuelling outside their winter territory prior to crossing the Sahara. Overwintering birds gained mass slowly during February and March (0.03 g/day), and non‐territorial or migrating birds at a much higher rate in April (at least 0.23 g/day), with up to 20% of migrating Whinchats in April potentially having sufficient fuel loads to cross the Sahara directly from central Nigeria. Our results suggest that most Whinchats leave their winter territories to fatten up locally or, possibly, by staging further north, closer to the southern limit of the Sahara. Resource constraints are therefore likely to be particularly focused in West Africa during mid‐April and possibly at staging areas before the crossing of the Sahara Desert.     

Genetic and environmental influences on natal dispersal distance in a resident bird species

We analyzed more than 1,600 dispersal events from two populations of a North American cooperatively breeding woodpecker species to determine what factors influence natal dispersal distance and whether distance traveled affects reproduction later in life. We found significant heritability of natal dispersal distance, in both males and females, indicating substantial additive genetic variance for this behavioral trait. Natal dispersal distance additionally was affected by social and ecological factors: individuals dispersing in their first year of life moved longer distances than those staying on their natal site as helpers for a prolonged time prior to dispersal, and increasing territory isolation led to longer dispersal distances. Successful dispersers incurred fitness costs, with lifetime fledgling production (in both sexes) and lifetime production of recruits to the breeding population (in females only) decreasing with increasing natal dispersal distance. We conclude that natal dispersal distance has a genetic basis but is modulated by environmental and social factors and that natal dispersal distance in this species is (currently) under selection.     

Can Experience Alter the Avian Annual Cycle? Results of Migration Experiments with Indigo Buntings

Field experiments were conducted during 2 years testing whether premature release on their familiar, perennial breeding territories would cause indigo buntings (Passerina cyanea) to delete spring migration from their annual cycle. In the first year, male buntings were captured on their breeding territories in summer and held indoors for the winter on the photoregime that they would have encountered in nature as fall migrants and winter residents. The following spring they were released at the time buntings begin spring migration from the subtropics. Half were released on their breeding territories, to which they would normally have returned; half were released far to the south. Some of the males released on their territories remained and passed into breeding condition without having migrated, while some of those released to the south returned north to their territories. The results suggest that endogenous or photoperiodic mechanisms controlling the timing and occurrence of migration in this species can be overridden by experience. In the second year we tested alternative explanations for these results. Our second year's results seem to indicate (1) that birds that deleted migration in year one did not do so because of unfavourable weather and (2) that the birds that returned to their breeding sites in year one were actually migrating and not simply retracing their outward journey.     

Mechanisms and reproductive consequences of breeding dispersal in a specialist predator under temporally varying food conditions

Individual variation in breeding dispersal has extensive ecological and evolutionary consequences, but the factors driving individual dispersal behaviour and their fitness consequences remain poorly understood. Our data on dispersal events of a rodent‐specialist predator, the Eurasian kestrel Falco tinnunculus, over 20 years in western Finland offers a unique opportunity to explore the mechanisms underlying breeding dispersal behaviour and its reproductive consequences in a wild bird population. Sex, age, body condition and previous breeding success affected breeding dispersal. Dispersal distances were longer in females than in males as well as longer in yearlings than in older individuals. Body condition was positively correlated to breeding dispersal distances, particularly for females. The lowest dispersal distances were recorded for intermediate brood sizes in the year preceding dispersal. Our results highlight sex‐ and environment‐specific consequences of breeding dispersal on reproductive performance. During increase phases of the three‐year vole cycles, males dispersing further had lower reproductive performance after dispersal, whereas in females, long breeding dispersal distances were associated with increased breeding success under all environmental conditions. These results suggest benefits associated to breeding dispersal in females, potentially related to large spatio‐temporal variation in main food abundance and intensity of intra‐specific competition. Breeding dispersal of males was costly during increasing food abundance, indicating the potential fitness benefits of environmental familiarity in this migratory species. Overall, our results indicate that both individual traits and environmental factors interact to shape breeding dispersal strategies in wide‐ranging predator populations under fluctuating food conditions.     

Stepping stones of life: natal dispersal in the group-living but noncooperative speckled warbler

In most cooperatively breeding birds the offspring of one sex, usually male, delays dispersal to remain on the natal territory and helps its parents to rear subsequent young. Thus delayed dispersal could be the first step in the evolution of cooperative breeding. We studied natal dispersal in a population of the group-living speckled warbler, Chthonicola sagittata, based on observations of a colour-banded population over 3 years. Unlike other group-living members of the Acanthizinae, all juvenile males in this population dispersed to settle on foreign territories as subordinates, which do not help rear the young. Speckled warblers showed all the life history traits that are thought to result in a saturated habitat and lead to delayed dispersal: they were sedentary, had high adult survival and had a male-biased sex ratio. However, they differed from other acanthizids in occurring at low density (0.18 birds/ha) on large breeding territories (6-12 ha), with a maximum of two males per territory. This may allow subordinates to live on foreign territories yet avoid aggression from dominants. A benefit of dispersal is that it provides an additional route to gaining a breeding vacancy. Dispersers can acquire vacancies on their new territory or on a neighbour's, but incest avoidance would be likely to constrain nondispersing males to neighbours' territories. A model of relative lifetime success showed that the survival benefits of natal philopatry are unlikely to outweigh this benefit of dispersal.     

Sex biased natal dispersal in a closed, saturated population of Seychelles warblers Acrocephalus sechellensis

The distances that individuals disperse, from their natal site to the site of first breeding and between breeding sites, have important consequences for the dynamics and genetic structure of a population. Nearly all previous studies on dispersal have the problem that, because the study area encompassed only a part of the population, emigration may have been confounded with mortality. As a result long-distance dispersers may have been overlooked and dispersal data biased towards short distances. By studying a virtually closed population of Seychelles warblers Acrocephalus sechellensis we obtained almost unbiased results on several aspects of dispersal. As in the majority of other avian species, natal dispersal distance was female biased in the Seychelles warbler. Female offspring also forayed further from the natal territory in search of breeding vacancies than male offspring. The sex bias in natal dispersal distance did, however, depend on local breeding density. In males, dispersal distance decreased as the number of territories bordering the natal territory increased, while in females, dispersal distance did not vary with local density. Dispersal by breeders was rare and, unlike in most species, distances did not differ between the sexes. We argue that our results favour the idea that the sex bias in natal dispersal distance in the Seychelles warbler is due to inbreeding avoidance and not resource competition or intrasexual competition for mates.     

Annual survival and breeding dispersal of a migratory passerine,the Scissor‐tailed Flycatcher

Knowledge of survival rates is critical for advancing our understanding of the dynamics of populations and here we report apparent annual survival and breeding dispersal of Scissor‐tailed Flycatchers (Tyrannus forficatus) breeding at two sites in southwest Oklahoma (Ft. Sill and Wichita Mountain Wildlife Refuge [WMWR]). Our Cormack‐Jolly‐Seber estimate of apparent adult survival for the period from 2008 to 2105 was relatively low (0.514) compared to estimates for 36 other migratory and socially monogamous passerines breeding in North America, and was independent of sex (males: N = 151; females: N = 119), breeding status (territory holder or floater), body mass, site, year, and precipitation during the year prior to breeding. Although apparent survival did not differ between sites, dispersal (N = 66 individuals) was more common and breeding dispersal distance (BDD) was greater for Scissor‐tailed Flycatchers at Ft. Sill where anthropogenic disturbance was more frequent. BDD also increased with body mass at Ft. Sill (but not at WMWR) and, after accounting for it, BDD at Ft. Sill tended to be greater for birds that failed to breed successfully in the past year. Older birds and males had the longest BDDs at WMWR, and males exhibited a similar tendency at Ft. Sill. We contend that our estimate of apparent survival is low, not because of inherently low survivorship, but, instead, as a consequence of frequent permanent emigration from our population. We also suggest that the greater BDD of older birds (WMWR) and males (both sites) reflects a history of selection for dispersal in response to frequent habitat disturbance. Frequent habitat disturbance, in addition to the opportunity to prospect for territories both before and after breeding, probably enable the earliest spring arrivals (typically older birds and males) to often relocate between years.     

Habitat use throughout migration: linking individual consistency, prior breeding success and future breeding potential

1.?Habitat use can influence individual performance in a wide range of animals, either immediately or through carry-over effects in subsequent seasons. Given that many animal species also show consistent individual differences in reproductive success, it seems plausible that individuals may have consistent patterns of habitat use representing individual specializations, with concomitant fitness consequences. 2.?Stable-carbon isotope ratios from a range of tissues were used to discern individual consistency in habitat use along a terrestrial-aquatic gradient in a long-distance migrant, the Bewick's swan (Cygnus columbianus bewickii). These individual specialisations represented <15% of the isotopic breadth of the population for the majority of individuals and were seen to persist throughout autumn migration and overwintering until aquatic habitats were no longer available. 3.?Individual foraging specialisations were then used to demonstrate two consecutive carry-over effects associated with macroscale habitat segregation: consequences of breeding season processes for autumn habitat use; and consequences of autumn habitat use for future reproductive success. Adults that were successful breeders in the year of capture used terrestrial habitats significantly more than adults that were not successful, revealing a substantial cost of reproduction and extended parental care. Use of aquatic habitats during autumn was, however, associated with increased body condition prior to spring migration; and increased subsequent breeding success in adults that had been unsuccessful the year before. Yet adults that were successful breeders in the year of capture remained the most likely to be successful the following year, despite their use of terrestrial habitats. 4.?Our results uniquely demonstrate not only individual foraging specializations throughout the migration period, but also that processes during breeding and autumn migration, mediated by individual consistency, may play a fundamental role in the population dynamics of long-distance migrants. These findings, therefore, highlight the importance of long-term consistency to our understanding of habitat function, interindividual differences in fitness, population dynamics and the evolution of migratory strategies.     

Relationship between arrival timing and breeding success of intra‐tropical migratory Fork‐tailed Flycatchers (Tyrannus savana)

For migratory birds, early arrival at breeding areas has many benefits, such as acquisition of better territories and mates. This strategy has been found in numerous species breeding at north‐temperate latitudes, but has not been yet reported for intra‐tropical migratory species. We evaluated the relationship between arrival date, initiation of breeding, and breeding success of Fork‐tailed Flycatchers (Tyrannus savana) breeding in southeastern Brazil and overwintering in northern South America. We color‐banded adult flycatchers during three breeding seasons and searched for them during the following breeding seasons. We also monitored nests from construction until either failure or fledging of young. We found that: (1) male Fork‐tailed Flycatchers arrived at the breeding site earlier than females, (2) males that arrived earlier had greater breeding success, and (3) nests where eggs were laid earlier in the breeding season were more likely to be successful than those where eggs were laid later. Male Fork‐tailed Flycatchers appeared to benefit from early arrival at a tropical breeding site, potentially mediated by their ability to acquire a high‐quality territory and mate as early as possible, and by the ability of their mate to begin breeding as early as possible. Breeding success for female Fork‐tailed Flycatchers may be determined primarily by a combination of the arrival date of their mate and how quickly they can begin breeding. Our results suggest that protandry occurs in an intra‐tropical migratory bird and that early arrival of males and early initiation of reproduction by females results in greater reproductive success. A better understanding of the underlying mechanisms that control the timing of migration and reproduction of this and other intra‐tropical migratory species is important for evaluating the challenges they face in light of current and future rapid environmental changes.     

Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.     

Dispersal and philopatry in the European badger, Meles meles

In comparison with other carnivores, European badgers, Meles meles L., show an unusually high degree of natal philopatry. In this paper, we present data on temporary and permanent movements between groups, in both male and female badgers, in a moderately high density population. A relatively small proportion of males dispersed, alone, to neighbouring territories. Dispersing males were larger than those remaining in their natal groups, and following dispersal they had higher testosterone titres and maintained testicular activity for a greater part of the year. Circumstantial evidence suggests that immigrants were the principal breeding males in their new territories. Dispersal was slightly more common in females, which dispersed away from large groups, where their chances of breeding were relatively low. Females dispersed in coalitions of 2–3, over longer distances, to territories occupied by single females. Resident females disappeared following the arrival of the immigrants, suggesting that territory 'takeovers' may have occurred. Members of both sexes also made temporary 'visits' to neighbouring territories, probably to obtain extra-group matings. Comparison with other badger populations suggests that the frequency of male dispersal declines at high population densities. In contrast, there is no effect of density on female dispersal, which occurs only rarely in some other populations. We suggest that the pattern of female/female competition is too complex to be explained solely in terms of variation in population density.     

Influence of Age and Prior Experience on Territorial Behavior and the Costs of Defense in Male Collared Lizards

In polygynous species where males maintain strong interseasonal philopatry to the same breeding territories, older individuals have prior experience defending their areas, whereas younger individuals are defending a territory for the first time. Theoretical and empirical studies predict that under such conditions the cumulative costs of defense may be lower for older experienced males as a consequence of familiarity with environmental and/or social conditions in their particular local habitats. We used quantitative data from focal observational studies and introduction experiments to test the hypothesis that older collared lizard males (Crotaphytus collaris) with prior territorial experience (3 yr+ males) acquire larger territories allowing them to court more, different females, with greater frequency without incurring higher defense costs. Consistent with this hypothesis, 3 yr+ males controlled significantly larger territories and courted more females, without having significantly higher rates of territory patrol, frequencies of advertisement display, or aggressive interactions with same‐sex competitors. Moreover, the intensity of responses to size‐ and age/experience‐matched tethered intruders by 3 yr+ males was lower than that by 2 yr males in their first season of territory defense. Our results support the hypothesis that age/prior occupancy of territories lowers defense costs allowing males to defend larger areas and increase opportunities to court females, perhaps increasing mating opportunities. By contrast, 2 yr males may need to respond more aggressively to intruders because their ownership of territories is tenuous as a consequence of shorter territory occupancy.     

Prospectors combine social and environmental information to improve habitat selection and breeding success in the subsequent year

1.?Because habitats have profound effects on individual fitness, there is strong selection for improving the choice of breeding habitat. One possible mechanism is for individuals to use public information when prospecting future breeding sites; however, to our knowledge, no study has shown prospecting behaviour to be directly linked to subsequent choice of breeding site and future reproductive success. 2.?We collected long-term data on territory-specific prospecting behaviour and subsequent breeding in the short-lived northern wheatear (Oenanthe oenanthe). Non-breeders established prospecting territories (<2 ha) that overlapped the breeding territories of conspecifics. We tested whether: (i) prospectors used social and environmental cues that predicted territory-specific breeding success in the following year, and (ii) the prospecting territory was tightly linked to the subsequent breeding territory of the prospector, and whether this link would be weakened by intraspecific competition with original territory owners if they also survived. 3.?As expected, prospectors were attracted to a combination of site-specific cues that predicted future breeding success, i.e. short ground vegetation, a successfully breeding focal pair and successful close neighbours. 4.?Prospecting behaviour was directly linked to the choice of the following year's breeding territory: 79% of surviving prospectors established a breeding territory at their prospecting site in the following year, with their breeding success being higher than other individuals of the same age. As predicted, fidelity to the prospected site was strongly dependent on whether the original territory owner of the same sex had died or moved. 5.?Our findings suggest that the use of multiple cues reduces the negative impact of stochasticity on the reliability of social cues at small spatial scales (e.g. territories) and hence increases the probability of breeding success in the next year. Also, the use of conspecific attraction (i.e. the preference for breeding aggregations) is selectively advantageous because individuals are more likely to find a vacancy in an aggregation as compared to a solitary site. By extension, we hypothesize that species life-history traits may influence the spatial scale of prospecting behaviour and habitat selection strategies.     

Breeding dispersal in an isolated population of Spotted Owls Strix occidentalis: evidence for improved reproductive output

Breeding dispersal among territorial species is of interest to population biologists because leaving a territory carries fundamental risks to the dispersing individuals, and this may not outweigh the costs of maintaining the territory. Most studies of breeding dispersal have focused on species inhabiting spatially open populations, in which undetected emigration could impart a negative bias to estimates of dispersal. We studied breeding dispersal in an isolated (spatially closed) population of California Spotted Owl Strix occidentalis occidentalis in southern California for 12 years to assess factors that might correlate with breeding dispersal. Twenty‐nine per cent (n = 47) of territorial females and 19% (n = 35) of territorial males dispersed at least once during the study. Annually, 0–13% of the territorial females and 0–12% of the territorial males dispersed. Among a set of a priori and post hoc models related to breeding dispersal, the top a priori model indicated that birds having higher reproductive output relative to the population average were less likely to disperse. A post hoc model based on an index of territory quality was ranked higher than the top a priori model and indicated that birds occupying higher quality territories were less likely to disperse. These two models were correlated and represented short‐ and long‐term reproductive performance, respectively. Birds that dispersed also failed to fledge young in the year prior to dispersal, but the failure to fledge young did not, by itself, explain dispersal. Because Spotted Owls are long‐lived, they may ultimately improve their reproduction by dispersing given that they would have future opportunities to breed over the long term. Birds whose mates are likely to have died tended to improve their reproductive success, whereas the relationship between reproductive success of birds that divorced was less clear. Substantial variation in breeding dispersal was unexplained by our analysis so it is likely that the motivation to disperse was a complex process in this population.