BREEDING COMPETITION IN A PACIFIC SALMON (COHO: ONCORHYNCHUS KISUTCH): MEASURES OF NATURAL AND SEXUAL SELECTION

In the breeding system of Pacific salmon, females compete for oviposition territories, and males compete to fertilize eggs. The natural selection in females and sexual selection in males likely has been responsible for their elaborate breeding morphologies and the dimorphism between the sexes. We quantified direct-selection intensities during breeding on mature coho salmon (Oncorhynchus kisutch), measured for seven phenotypic characters, including three secondary sexual characters. Wild and sea-ranched hatchery coho were used to enhance the range of phenotypes over which selection could be examined. The fish were allowed to breed in experimental arenas where we could quantify components of breeding success as well as estimate overall breeding success. We found that without competition, natural selection acts only on female body size for increased egg production; there is no detectable selection on males for the phenotypic distribution we used. Under competition, the opportunity for selection increased sixfold among females. Natural selection favored female body size and caudal-peduncle (tail) depth. Increased body size meant increased egg production and access to nesting territories. The caudal peduncle, used in burst swimming and nest digging, influenced both successful egg deposition and nest survival. Increasing density increased competition among females, though it did not significantly intensify natural selection on their characters. In males, competition increased the opportunity for selection 52-fold, which was nine times greater than for females. Sexual selection favored male body size and hooked snout length, both characters directly influencing male access to spawning opportunities. Selection on male body size was also affected significantly by breeding density. The ability of large males to control access to spawning females decreased at higher densities reflecting an increase in the operational sex ratio. Further, the relative success of small males, which could sneak access to spawning females, appeared to increase as that of intermediate-sized males decreased. Such disruptive selection may be responsible for the evolution of alternative reproductive tactics in salmon.     

Competitive interactions among raptors in boreal forests

We examined inter-specific interactions among goshawks (Accipiter gentilis), common buzzards (Buteo buteo) and honey buzzards (Pernis apivorus) in western Finland in 1983–1996. Because goshawks are among the largest birds of prey species in boreal forests they may take over the nest of smaller and less-competitive forest-dwelling raptors when searching for suitable places for breeding. Accordingly, more than half of newly established goshawk territories were found on the territories previously occupied by the common buzzard and the honey buzzard. Otherwise, territory sharing between these species was rare. Fledgling production of honey buzzards was not associated with the presence of goshawks, probably owing to the almost 2 months later onset of breeding. This probably decreases competitive interactions between these two species. An intensive interference competition, instead, seemed to be evident between common buzzards and goshawks, because the fledgling production of common buzzards was decreased by 20% as a result of failures during incubation and nestling period in the vicinity (<1 km) of occupied goshawk nests. Similarly, territory occupancy of common buzzards till the next breeding season was significantly reduced in the presence of goshawks. Relatively high proportions of occupied buzzard territories (17%) in the study area were shared by breeding goshawks on the same territory. This suggests that although their diets are dissimilar they inhabit similar habitats and might compete for the available prime nesting habitats within forest landscapes. In addition, goshawks benefit from taking over the complete nests of other raptors, imposing upon the original owners of the nest, because building a large stick nest is probably energetically costly. As a large raptor, the goshawk apparently has a competitive advantage over smaller ones, and may have an ever-increasing impact on smaller birds of prey, if there is a lack of sheltered forests inducing competition for the available nest sites.     

Effects of nest-box density on the behavior of Tree Swallows during nest building

ABSTRACT.   In territorial species, increased density is often linked to an increase in aggressive interactions, which may result in trade-offs between competitive behavior and nest construction. We examined the impact of nesting in areas of high-density versus low-density nest boxes on conspecific interactions and nest-building effort in a population of Tree Swallows ( Tachycineta bicolor ). We also examined whether expected differences in behavior related to variation in nest quality and reproductive success in high-density and low-density areas. No differences in either nest-building behavior or reproductive success were observed between areas of high-density and low-density boxes, but there was a tendency of more frequent behavioral interactions at high density. Similarly, there was a significant difference between pairs defending single nest boxes and those defending multiple nest boxes in the number of interactions with conspecifics. These results suggest that although there may be more competition with conspecifics for Tree Swallows nesting at high density, this does not appear to affect either nest quality or reproductive success.     

An ‘ecological trap’ for yellow warbler nest microhabitat selection

Contrary to assumptions of habitat selection theory, field studies frequently detect ‘ecological traps’, where animals prefer habitats conferring lower fitness than available alternatives. Evidence for traps includes cases where birds prefer breeding habitats associated with relatively high nest predation rates despite the importance of nest survival to avian fitness. Because birds select breeding habitat at multiple spatial scales, the processes underlying traps for birds are likely scale‐dependent. We studied a potential ecological trap for a population of yellow warblers Dendroica petechia while paying specific attention to spatial scale. We quantified nest microhabitat preference by comparing nest‐ versus random‐site microhabitat structure and related preferred microhabitat features with nest survival. Over a nine‐year study period and three study sites, we found a consistently negative relationship between preferred microhabitat patches and nest survival rates. Data from experimental nests described a similar relationship, corroborating the apparent positive relationship between preferred microhabitat and nest predation. As do other songbirds, yellow warblers select breeding habitat in at least two steps at two spatial scales; (1) they select territories at a coarser spatial scale and (2) nest microhabitats at a finer scale from within individual territories. By comparing nest versus random sites within territories, we showed that maladaptive nest microhabitat preferences arose during within‐territory nest site selection (step 2). Furthermore, nest predation rates varied at a fine enough scale to provide individual yellow warblers with lower‐predation alternatives to preferred microhabitats. Given these results, tradeoffs between nest survival and other fitness components are unlikely since fitness components other than nest survival are probably more relevant to territory‐scale habitat selection. Instead, exchanges of individuals among populations facing different predation regimes, the recent proliferation of the parasitic brown‐headed cowbird Molothrus ater, and/or anthropogenic changes to riparian vegetation structure are more likely explanations.     

Joint effects of feeding and breeding behaviour on trophic dimorphism in hummingbirds

A survey of 166 hummingbird species reveals novel associations of bill-length sexual dimorphism (BLSD) with plumage and breeding behaviours. Across all species, female bills become proportionately longer than male bills (higher female-to-male BLSD ratio) as sexual dichromatism increases. However, male bills are proportionately longer (lower female-to-male BLSD ratio) in both lekkers (traditional group display) and clustered breeders (female harems or colonial nests) compared with dispersed breeders. The overall positive association of plumage with BLSD suggests that social status determines priority of access to nectar-providing flowers. Furthermore, the distinctive BLSD associated with breeding aggregations may arise from behaviours that impose constraints on the usual male priority at flowers: female dominance over males around nest colonies and male residence on lek-mating territories. These various factors appear to alter plumage and bill characters of both sexes to produce the range of dimorphisms within the various dispersed and aggregated breeding system categories. Feedback loops caused by ecological consequences of breeding behaviour may alter the evolutionary dynamics of breeding systems, bird-plant interactions, and competing pollinators, as well as help explain the lek paradox.     

Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

Social composition of destination territories and matrix habitat affect red-cockaded woodpecker dispersal

We studied the influence of matrix habitats and destination territory social composition on dispersal choices made by red-cockaded woodpeckers (Picoides borealis). We compared matrix habitats and social conditions for 165 dispersals documented between 1995 and 2005 with similarly distanced destinations that went unselected. We modeled data with discrete choice analysis methods, and ranked models using a model selection approach. Model-averaged parameters indicated that juvenile female, and juvenile and helper male red-cockaded woodpeckers dispersed across matrix forests with characteristics similar to those used for breeding. Models differed for sex and social classes, but results generally indicated that birds chose not to transit forests with greater densities of hardwood trees and young pines, and that dispersal was more likely to occur across forests with more large diameter pine trees. Dispersers of both sexes also chose destinations without non-breeding helpers that might present aggressive social challenges to dispersers, particularly males. Juvenile females and helper males also preferred to disperse to territories that were currently occupied by breeding woodpeckers. Previous studies documented the influence on dispersal of travel distance, natal area social interactions and resources, and resources at dispersal destinations. Our results extend factors that affect dispersal movement to include matrix habitats traversed by dispersers and the social composition of destination territories. As such, results have key implications for promoting population connectivity and conservation management of endangered red-cockaded woodpeckers, as well as other resident territorial birds. © 2011 The Wildlife Society.     

The breeding ecology and behaviour of Wrens Troglodytes troglodytes under primaeval and secondary conditions

The breeding habits of Wrens inhabiting the primaeval forest of Biatowieza National Park, Poland, are described and compared with those of Wrens living in man-transformed habitats of Western Europe.
Significant inter-habitat differences in numbers exist in Biatowieza. On average there are 4-3 territories per 10 ha in ash-alder, 1 -7 territories per 10 ha in oak–hornbeam and 0–4 territories per 10 ha in coniferous habitats. In ash–alder stands the males have smaller territories, build fewer nests and have better pairing success than in oak–hornbeam stands. However, these two habitats do not differ significantly in average clutch size, timing of breeding or nesting success. Over 80% of nests used for breeding in Biatowieza are situated in uprooted trees. Inter-habitat differences in bird numbers reflect differences in distribution and density of such trees.
Low numbers of Biatowieza Wrens cannot be accounted for solely by high mortality away from the breeding grounds. Available data strongly suggest that numbers of both sexes are proximately limited by behavioural interactions on the breeding ground.
In west European Wrens, densities are higher, territories smaller, polygyny more frequent, clutches smaller and production of young higher than in Biatowieza. Differences in production of young result from different rates of nest predation. Indirect evidence suggests that the breeding habits of Wrens once living in the primaeval forests of western Europe were similar to those of Biatowieza birds, and that the differences observed nowadays result to a large extent from man-made transformations of habitat. It is suggested that man's lowering of the predation pressure acting both on proximate (enabling birds to produce more young) and ultimate levels (decreasing relative advantages of dispersed breeding) was the decisive factor allowing an increase in numbers of Wrens in secondary habitats.     

A Young Manakin Knows His Place: Evidence for an Age‐Graded Dominance Hierarchy Among Long‐Tailed Manakins

In lek‐breeding systems where many males gather at display sites, males benefit from the establishment of dominance hierarchies to reduce intrasexual aggression and the associated risk of injuries. Long‐tailed manakins (Chiroxiphia linearis) exhibit an exploded lek‐breeding system wherein the two top‐ranking males at each display site team up to perform elaborate coordinated courtship displays for females. Young males undergo delayed plumage maturation whereby they acquire distinct pre‐definitive plumage patterns each year until they attain definitive plumage in their fifth year. This unique characteristic is thought to have evolved as a status‐signalling mechanism to aid in the establishment of an age‐graded dominance hierarchy in which older males are dominant to younger males. Previous research has shown evidence for such a dominance hierarchy among alpha and beta males; however, the presence of this hierarchy among males of other age classes has never been quantified. In this study, we investigated the presence of an age‐graded dominance hierarchy by determining whether older males direct more aggressive behaviours towards younger males. We also investigated whether status signalling is less clear within age classes than between age classes, by determining whether males within the same age class exhibit more aggression towards each other. We found that older males performed aggressive behaviours towards younger males much more frequently than younger males performed aggressive behaviours towards older males. We also found that some aggressive interactions occurred between males within the same age class more frequently than between males from different age classes. Our study provides some evidence for an age‐graded dominance hierarchy among male long‐tailed manakins of all age classes and also provides some support for the status‐signalling hypothesis. However, further research is needed to conclusively establish the presence of a linear dominance hierarchy among younger male manakins. This research may help us better understand the evolution of complex hierarchical systems in animals.     

Providing nest boxes for hole-nesting birds – Does habitat matter?

Nest boxes are a popular management tool to increase nest site availability for hole-nesting birds, but biological consequences of this technique in different habitats are poorly studied. In our study area in southwestern Estonia, nest boxes for small passerines were set up in deciduous and coniferous woods. Great tits Parus major preferred the food-rich deciduous habitat for breeding, as judged by higher nest-box occupation, earlier egg-laying and larger clutches and eggs. However, in coniferous habitat more and heavier young fledged per nest, and the return rate of both fledglings and adults was higher. We propose two mutually non-exclusive explanations, both related to the maladaptive outcome of the provision of nest boxes: (i) in the preferred habitat, nest boxes caused a supra-optimal breeding density leading to an ecological trap; (ii) boxes drastically improved the non-preferred habitat, but birds were unable to exploit the breeding habitat fully. One should be careful in providing large numbers of artificial nest sites in preferred habitats. Sometimes it would be more preferable to improve less favourable habitats by removing critical constraints.     

Polygyny and nest site selection in the pied flycatcher

Nest site choice, and its influence on mating and breeding success were studied in a population of individually marked pied flycatchers (Ficedula hypoleuca) for 7 years. Some nest boxes were used more frequently than expected from random choice of nest site. Flycatchers bred more successfully in such attractive boxes, irrespective of whether the brood was attended by the male or not. Arriving males and females settled at boxes in descending order of attractiveness. Boxes accepted by females were more attractive than alternative boxes advertised simultaneously by males. Secondary nest boxes of polygynous males tended to be more attractive than boxes at which monogamous males displayed, without success, for secondary mates. Returning males with local breeding experience started to sing at nest boxes earlier in spring than first season males, settled at more attractive boxes, and mated polygynously more often. The results indicate that male mating success should be closely dependent on the male's ability to monopolize attractive nest sites.     

The role of conservative versus innovative nesting behavior on the 25‐year population expansion of an avian predator

Species ranges often change in relation to multiple environmental and demographic factors. Innovative behaviors may affect these changes by facilitating the use of novel habitats, although this idea has been little explored. Here, we investigate the importance of behavior during range change, using a 25‐year population expansion of Bonelli's eagle in southern Portugal. This unique population is almost exclusively tree nesting, while all other populations in western Europe are predominantly cliff nesting. During 1991–2014, we surveyed nest sites and estimated the year when each breeding territory was established. We approximated the boundaries of 84 territories using Dirichlet tessellation and mapped topography, land cover, and the density of human infrastructures in buffers (250, 500, and 1,000 m) around nest and random sites. We then compared environmental conditions at matching nest and random sites within territories using conditional logistic regression, and used quantile regression to estimate trends in nesting habitats in relation to the year of territory establishment. Most nests (>85%, n = 197) were in eucalypts, maritime pines, and cork oaks. Nest sites were farther from the nests of neighboring territories than random points, and they were in areas with higher terrain roughness, lower cover by agricultural and built‐up areas, and lower road and powerline densities. Nesting habitat selection varied little with year of territory establishment, although nesting in eucalypts increased, while cliff nesting and cork oak nesting, and terrain roughness declined. Our results suggest that the observed expansion of Bonelli's eagles was facilitated by the tree nesting behavior, which allowed the colonization of areas without cliffs. However, all but a very few breeding pairs settled in habitats comparable to those of the initial population nucleus, suggesting that after an initial trigger possibly facilitated by tree nesting, the habitat selection remained largely conservative. Overall, our study supports recent calls to incorporate information on behavior for understanding and predicting species range shifts.     

Winter territory prospecting is associated with life‐history stage but not activity in a passerine

Finding a high quality territory is essential for many animals to reproduce successfully. Despite its importance for fitness, we know little about the process of territory prospecting in wild birds, and whether individual traits and behaviours, such as personality, co‐vary with territory prospecting. Here, we use long‐term data from a wild, insular house sparrow Passer domesticus population to test three hypotheses about territory fidelity and prospecting: 1) house sparrows show high territory fidelity between years and also during winter. 2) Individuals will prospect for a breeding territory during their first winter whereas older, more experienced individuals will keep a territory from previous years and will, therefore, show no or reduced winter territory prospecting. 3) More active behavioural types will prospect more than less active behavioural types. We use data from four winters from automatically, daily recorded nest‐box visits of 188 birds of known age. The number of nest‐boxes that each individual visited within each winter was used as a proxy of winter territory prospecting. We show that house sparrows visit multiple nest‐boxes during their first winter, whereas older individuals keep territories year‐round and, potentially because of this, indeed show reduced winter territory prospecting. Activity was not associated with the number of nest‐boxes visited. Further research is needed to investigate whether time of territory and mate acquisition differs among individuals and the possible effect on lifetime fitness.     

Quiet violence: Savannah Sparrows respond to playback‐simulated rivals using low‐amplitude songs as aggressive signals

When animals compete over resources such as breeding territories, they often use signals to communicate their aggressive intentions. By studying which signals are associated with aggressive interactions, we gain a deeper appreciation of animal behaviour. We studied aggressive signalling in male Savannah Sparrows, Passerculus sandwichensis, focusing on signals that precede physical attack against territorial intruders. We simulated intruders using song playback and taxidermic models, and we determined which behaviours were associated with physical attack. Previous studies that have used this approach suggest that many species produce songs of dramatically lower amplitude, or “soft songs,” as signals of aggressive intention. Savannah Sparrows, however, are not known to produce soft songs, and therefore, they provided an interesting system for testing signals that predict attack. Of 93 playback subjects, 23 males attacked the simulated intruder and 70 did not. To our surprise, Savannah Sparrows produced soft songs, and the number of soft songs was a significant predictor of attack on the simulated intruder. Birds also showed a nonsignificant tendency to produce more “chip” calls prior to attack on the simulated intruder, whereas three other measured behaviours (aggressive calls, wing waving and passes over the model) did not predict attack. Our study contributes to the growing body of research on aggressive territorial signals and reveals that soft song is an even more widespread signal of aggression in songbirds than previously recognized.     

Seasonal and Diurnal Changes in Aggression and Territory Size in the Black-headed Gull (Larus ridibundus L.) on Islands in the Middle Reaches of the Vistula River

Seasonal and diurnal changes in territory size and frequency of aggressive interactions of various intensity were examined in colonies of the black-headed gull (Larus ridibundus) on river islands and ponds of central Poland. The highest frequency of the total aggression and the largest territories were recorded before the egg-laying stage, this being related to the establishing of territories, defence of the mate, and defence of nest material. Later in the season both aggression and territory size declined. Unlike the large species of gulls, black-headed gulls did not increase their aggression and territory size during hatching of the young. High-intensity aggressive behaviours (choking, attacking and fighting) were not so frequently displayed as low-intensity aggressive postures (upright, long call, forward) in each phase of the breeding cycle. Presumably this was because the former were more costly in terms of energy. Peak territorial activity occurred in the late afternoon. An increase in the frequency of intense aggressive behaviours (attack, fight) at that time was combined with an increased mobility of birds (departures to and arrivals from foraging sites). A reduced frequency of high-intensity aggressive displays in the period when the largest number of birds was present in the colony was likely to be adaptive.     

Nest box use by woodland dormice (Graphiurus murinus): the influence of life cycle and nest box placement

The use of nest boxes by the woodland dormouse, Graphiurus murinus, was investigated over a 13-month period in a riverine forest of the Great Fish River Reserve, South Africa. We predicted that some characteristics of nest box placement would affect nest box use and that the seasonal pattern of nest box use would be linked to the species' life cycle and physiological and socioecological characteristics. Generalized linear models indicated that the time since nest box installation and nest box height above ground positively affected the frequency and intensity of nest box use. Male and female dormice, as well as adults and juveniles, did not differ in the number of nest boxes used and equally occupied individual nest boxes. The percentage of nest boxes used peaked during spring and summer (breeding period) and dropped during winter (hibernation). However, whereas significantly more males were caught during the mating season (spring), the number of females occupying nest boxes was constant during the year. As female dormice successfully bred in the nest boxes, the observed sexual patterns suggest that (artificial) nest sites represent an important resource for females, whereas females seem to constitute the main resource for males, as predicted by the socioecological model.     

Increased microclimatic variation in artificial nests does not create ecological traps for a secondary cavity breeder,the European roller

Artificial devices are increasingly used in conservation measures to mitigate the disappearance of natural habitats. However, few studies have demonstrated their benefits for the target species, and they may pose a risk of creating ecological traps. This occurs when lower individual fitness is found in artificial habitats that are more attractive than their natural equivalents. In this study, we tested the ecological trap hypothesis on a dense population of European rollers Coracias garrulus breeding in both natural cavities and nest boxes. Our initial prediction was that the more stressful microclimatic conditions of nest boxes would lead to reduced fitness of European rollers, thus creating an ecological trap. The results showed that nest boxes were preferred over natural cavities. Despite significantly more extreme microclimatic conditions in nest boxes, we found similar breeding parameters between artificial and natural nest types. Our results also suggest that European rollers selected the nest boxes which best buffered the temperature, thus avoiding potential ecological traps. Overall our results led to the conclusion that nest boxes do not create ecological traps for European rollers in this study area. However, other species may be more sensitive to microclimatic variations or less able to avoid the least favorable nest boxes. These findings could help to inform the placement of nest boxes in order to reduce extreme temperatures and variation in humidity rates. Future studies could compare nest types for other fitness parameters, such as juvenile body condition or survival. We also recommend the ecological trap hypothesis as a useful framework to evaluate the outcomes of artificial devices used for conservation.     

Predation risk effects on fitness related measures in a resident bird

Predation risk is thought to be highly variable in space and time. However, breeding avian predators may create locally fixed and spatially fairly predictable predation risk determined by the distance to their nest. From the prey perspective, this creates predation risk gradients that potentially have an effect on fitness and behavioural decisions of prey. We studied how breeding avian predators affect habitat selection (nest location) and the resulting fitness consequences in a northern population of resident willow tit ( Parus montanus ). Data included 429 willow tit nests over a four year period in a landscape containing a total of 33 avian predator nests. Willow tit nests were located randomly in the landscape and no predator avoidance in habitat selection or emptying of territories in proximity to predators was observed. Nestling size, however, was positively associated with distance from predator nests (n=252). Nestling mass and wing length were about 4.5% smaller close to predator nests compared to nestlings raised far from predator nests. Tarsus length also exhibited a positive relationship with increasing distance from predator nest but this was limited to habitats of young forests and pine bogs or dense mixed forests (4% increase). It is likely that habitat structural complexity influenced the perception of predation risk in different habitats. Our results indicate that willow tits do not provide reliable cues of predator free habitats for settling migrants. Nonetheless, breeding avian predators may create predictable predation risk in the landscape which is an important factor affecting reproductive success and potentially the demography of prey populations.     

Size-dependent selection on arrival times in sticklebacks: why small males arrive first

Abstract Studies on arrival time to breeding areas show that high-quality males usually arrive first and gain the highest reproductive success. This is generally assumed to be due to phenotype-dependent costs and benefits of early arrival. We show that the opposite arrival order can occur, probably due to selection on poor-quality males to increase their chances of reproduction. In a fish species, the threespine stickleback, Gasterosteus aculeatus , small males arrived before larger males at the breeding grounds. Early arrival was costly because predation risk was at its highest at the start of the season and early territory establishment was selected against, as demonstrated by selection coefficients for territory maintenance and hatching success. Large males probably postponed arrival until females were available to decrease predation risk costs and increase offspring production. An experimental study showed that a delay in arrival of large males does not decrease their probability of reproduction, because large males are able to take over nest sites from small males. Small males, on the other hand, are less likely to establish territories in competition with large males but can pay the costs of early arrival in exchange for the benefit of access to territories. Thus, whereas natural selection favors later arrival, sexual selection through competition for breeding territories favors early arrival in small, competitively inferior males. This results in the benefits of early arrival depending on the competitive ability of the male, which favors size-dependent optimal arrival times.     

Role of breeding and natal movements in lifetime dispersal of a forest‐dwelling rodent

The lifetime movements of an individual determine the gene flow and invasion potential of the species. However, sex dependence of dispersal and selective pressures driving dispersal have gained much more attention than dispersal at different life and age stages. Natal dispersal is more common than dispersal between breeding attempts, but breeding dispersal may be promoted by resource availability and competition. Here, we utilize mark–recapture data on the nest‐box population of Siberian flying squirrels to analyze lifetime dispersal patterns. Natal dispersal means the distance between the natal nest and the nest used the following year, whereas breeding movements refer to the nest site changes between breeding attempts. The movement distances observed here were comparable to distances reported earlier from radio‐telemetry studies. Breeding movements did not contribute to lifetime dispersal distance and were not related to variation in food abundance or habitat patch size. Breeding movements of males were negatively, albeit not strongly, related to male population size. In females, breeding movement activity was low and was not related to previous breeding success or to competition between females for territories. Natal philopatry was linked to apparent death of a mother; that is, we did not find evidence for mothers bequeathing territories for offspring, like observed in some other rodent species. Our results give an example of a species in which breeding movements are not driven by environmental variability or nest site quality. Different evolutionary forces often operate in natal and breeding movements, and our study supports the view that juveniles are responsible for redistributing individuals within and between populations. This emphasizes the importance of knowledge on natal dispersal, if we want to understand consequences of movement ecology of the species at the population level.