Reciprocating dispersal by habitat-selecting white-footed mice

Theories of dispersal driven by density-dependent habitat selection demonstrate that putative examples of source–sink dynamics and balanced dispersal may also be explained by a more general pattern of bi-directional, reciprocating dispersal. Analyses of 19 years of data on dispersal by white-footed mice confirm the theory. Fitness of territorial white-footed mice living in an agricultural mosaic is higher in forest habitat than it is in either edge or fencerows. Density-dependent habitat selection theory predicts that if net emigration by mice flows from the forest to forest-edge during periods of population growth, animals should subsequently move from the edge to forest during population decline. The pattern of mouse dispersal varies between seasons as populations wax and wane in abundance. Mice tend, as predicted, to move from high-density forest habitat into low-density edge during periods of population increase, and from the low-density edge into high-density forest during periods of population decline. Over all years combined, dispersal by white-footed mice was balanced. Each habitat tended to gain as many dispersing individuals as it lost. The results support a conditional dispersal strategy linked to density-dependent habitat selection, but also suggest the possibility of multiple coexisting strategies.     

Environmental networks,compensating life histories and habitat selection by white-footed mice

Summary Analysis of 6 years' data on a population of free-living white-footed mice documents both phenotypic and environmental control of litter size. Litter size was positively correlated with maternal body size. Maternal size depended upon both seasonal and annual variation. Paradoxically, the proportion of small versus large litters varied among habitats independently of the effects of body size. The result is an influence of habitat on life history that yields patterns of reproduction and survival opposite to the predictions of demographic theory. The habitat producing the largest litters had a relatively high ratio of adult/juvenile survival. Litter size was small in the habitat where the adult/juvenile survival ratio was smallest. All of these anomalous patterns can be explained through density-dependent habitat selection by female white-footed mice. Life-history studies that ignore habitat and habitat selection may find spurious correlations among traits that result in serious misinterpretations about life history and its evolution.     

Habitat matching: Alternatives and implications to populations and communities

Summary I evaluate habitat matching rules based on ideal distribution models of density-dependent habitat use. Recent approaches and the ideal free continuous input matching rule on which they depend, are restricted to only those habitats that are jointly occupied across the full range of population sizes. These assumptions may often be inappropriate to field applications of habitat matching. I develop alternatives that can be applied to a wide array of ideal forms of habitat selection, including the ideal free, continuous input example. Input matching can be distinguished from assumptions of consumer-resource models and preemptive habitat use by regressions of density between paired habitats (isodars). Isodars for continuous input models should be linear on a logarithmic scale, while those for consumer-resource models should be linear on an arithmetic scale. Pre-emptive isodars can be distinguished from the others by dramatic non-linearities at both low and high densities. Field data on white-footed mice support the consumer-resource theory. Implications of the rules for population regulation and community organization are highlighted by new models that specify how the fitness of pre-emptive habitat selectors should decline with increasing density. Strong non-linearities produced by comparisons between variable and homogeneous habitats produce reversing source-sink population regulation and a new form of cyclical community dynamics. Variable habitats act as a source of emigrants at low density and a sink for immigrants at high density. Subordinate species may occupy only the variable habitat at both low and high density.     

The cost of habitat selection in prairie voles: an empirical assessment using isodar analysis

The ideal free distribution assumes that habitat selection is without cost and predicts that fitness should be equal in different habitats. If habitat selection has a cost, then individuals should only move to another habitat when potential fitness in the new habitat exceeds that in the source habitat by an amount greater than the cost of habitat selection. We used isodar techniques to assess the cost of habitat selection. In an experimental landscape, we monitored density, movement, and reproductive success of adult female prairie voles, Microtus ochrogaster, in adjacent paired habitats with low and high cover. We tested the following hypotheses: (1) adult female prairie voles exhibited density-dependent habitat selection; (2) the cost of habitat selection was density-independent. Habitat quality based on population density and fitness of adult females was higher in high cover habitats. Net movement was from low cover to high cover habitats. The results indicated that adult female prairie voles exhibited density-dependent habitat selection. Furthermore, there was a significant cost of habitat selection, and the cost was density-independent.     

Incorporating density dependence into the oviposition preference-offspring performance hypothesis

1. Although theory predicts a positive relationship between oviposition preferences and the developmental performance of offspring, the strength of this relationship may depend not only on breeding site quality, but also on the complex interactions between environmental heterogeneity and density-dependent processes. Environmental heterogeneity may not only alter the strength of density dependence, but may also fundamentally alter density-dependent relationships and the preference-performance relationship. 2. Here I present results from a series of field experiments testing the effects of environmental heterogeneity and density-dependent feedback on offspring performance in tree-hole mosquitoes. Specifically, I asked: (i) how do oviposition activity, patterns of colonization and larval density differ among habitats and among oviposition sites with different resources; and (ii) how is performance influenced by the density of conspecifics, the type of resource in the oviposition site, and the type of habitat in which the oviposition site is located? 3. Performance did not differ among habitats at low offspring densities, but was higher in deciduous forest habitats than in evergreen forest habitats at high densities. Oviposition activity and larval densities were also higher in deciduous forests, suggesting a weak preference for these habitats. 4. The observed divergence of fitness among habitats with increasing density may select for consistent but weak preferences for deciduous habitats if regional abundances vary temporally. This would generate a negative preference-performance relationship when population densities are low, but a positive relationship when population densities are high. 5. This study demonstrates that failure to recognize that fitness differences among habitats may themselves be density-dependent may bias our assumptions about the ecological and evolutionary processes determining oviposition preferences in natural systems.     

Hamilton's rule confronts ideal free habitat selection

If individuals occupy habitats in a way that maximizes their fitness, if they are free to occupy the habitats they choose and if fitness declines with population density, then their abundance across habitats should follow an ideal free distribution. But, if individuals are genetically related, this simple fitness-maximization mechanism breaks down. Habitat occupation should obey Hamilton's rule (natural selection favours traits causing a loss in individual fitness as long as they result in an equal or greater gain in inclusive fitness) and depends more on inclusive fitness than it does on individual fitness. We demonstrate that the resulting inclusive-fitness distribution inflates the population density in habitats of poorer inherent quality, creating pronounced source sink dynamics. We also show that density-dependent habitat selection among relatives reinforces behaviours such as group defence and interspecific territoriality, and that it explains many anomalies in dispersal and foraging.     

An appraisal of the fitness consequences of forest disturbance for wildlife using habitat selection theory

Isodar theory can help to unveil the fitness consequences of habitat disturbance for wildlife through an evaluation of adaptive habitat selection using patterns of animal abundance in adjacent habitats. By incorporating measures of disturbance intensity or variations in resource availability into fitness-density functions, we can evaluate the functional form of isodars expected under different disturbance-fitness relationships. Using this framework, we investigated how a gradient of forest harvesting disturbance and differences in resource availability influenced habitat quality for snowshoe hares (Lepus americanus) and red-backed voles (Myodes gapperi) using pairs of logged and uncut boreal forest. Isodars for both species had positive intercepts, indicating reductions to maximum potential fitness in logged stands. Habitat selection by hares depended on both conspecific density and differences in canopy cover between harvested and uncut stands. Fitness-density curves for hares in logged stands were predicted to shift from diverging to converging with those in uncut forest across a gradient of high to low disturbance intensity. Selection for uncut forests thus became less pronounced with increasing population size at low levels of logging disturbance. Voles responded to differences in moss cover between habitats which reflected moisture availability. Lower moss cover in harvested stands either reduced maximum potential fitness or increased the relative rate of decline in fitness with density. Differences in vole densities between harvested and uncut stands were predicted, however, to diminish as populations increased. Our findings underscore the importance of accounting for density-dependent behaviors when evaluating how changing habitat conditions influence animal distribution.     

Density-dependent host selection in ectoparasites: An application of isodar theory to fleas parasitizing rodents

Parasites should make the same decisions that every animal makes regarding fitness reward. They can maximize reproductive success by selection of those habitats that guarantee the greatest fitness output. We consider the host population as a habitat of a parasite population. Consequently, hosts (=habitats) that differ quantitatively or qualitatively will support different numbers of parasites. The nature of habitat selection can be detected by isodars, lines along which habitat selection yields equivalent fitness reward. We applied this approach to study host selection of five fleas, each infesting two desert rodents. Xenopsylla conformis, Xenopsylla ramesis, Nosopsyllus iranus theodori and Stenoponia tripectinata medialis parasitize Gerbillus dasyurus and Meriones crassus. Synosternus cleopatrae pyramidis parasitizes Gerbillus andersoni allenbyi and Gerbillus pyramidum. Three fleas ( X. conformis, X. ramesis and S. c. pyramidis) were able to perceive quantitative (amount of the resource; e.g. organic matter in the nest for flea larvae) and/or qualitative (pattern of resource acquisition; e.g. host defensiveness) differences between hosts. Two other fleas did not perceive between-host differences. X. conformis was a density-dependent host selector that showed sharp selectivity at low density. X. ramesis and S. c. pyramidis were density-independent host selectors with a direct correspondence of density with habitat quality. N. i. theodori and S. t. medialis were non-selectors with no relationship at all between density and host quality. The results of the application of the isodar theory suggest that ectoparasites, like other animals, behave as if they are able to make choices and decisions that favour environments in which their reproductive benefit is maximized.     

Density-dependent habitat selection by brown-headed cowbirds (<Emphasis Type="Italic">Molothrus ater</Emphasis>) in tallgrass prairie

Local distributions of avian brood parasites among their host habitats may depend upon conspecific parasite density. We used isodar analysis to test for density-dependent habitat selection in brown-headed cowbirds (Molothrus ater) among tallgrass prairie adjacent to wooded edges, and prairie interior habitat (>100 m from wooded edges) with and without experimental perches. Eight study sites containing these three habitat treatments were established along a geographical gradient in cowbird abundance within the Flint Hills region of Eastern Kansas and Oklahoma, USA. The focal host species of our study, the dickcissel (Spiza americana), is the most abundant and preferred cowbird host in the prairie of this region. Cowbird relative abundance and cowbird:host abundance ratios were used as estimates of female cowbird density, whereas cowbird egg density was measured as parasitism frequency (percent of dickcissel nests parasitized), and parasitism intensity (number of cowbird eggs per parasitized nest). Geographical variation in cowbird abundance was independent of host abundance. Within study sites, host abundance was highest in wooded edge plots, intermediate in the experimental perch plots, and lowest in prairie interior. Cowbirds exhibited a pattern of density-dependent selection of prairie edge versus experimental perch and interior habitats. On sites where measures of cowbird density were lowest, all cowbird density estimates (female cowbirds and their eggs) were highest near (100 m) wooded edges, where host and perch availability are highest. However, as overall cowbird density increased geographically, these density estimates increased more rapidly in experimental perch plots and prairie interiors. Variation in cowbird abundance and cowbird:host ratios suggested density-dependent cowbird selection of experimental perch over prairie interior habitat, but parasitism levels on dickcissel nests were similar among these two habitats at all levels of local cowbird parasitism. The density-dependent pattern of cowbird distribution among prairie edge and interior suggested that density effects on perceived cowbird fitness are greatest at wooded edges. A positive relationship between daily nest mortality rates of parasitized nests during the nestling period with parasitism intensity levels per nest suggested a density-dependent effect on cowbird reproductive success. However, this relationship was similar among habitats, such that all habitats should have been perceived as being equally suitable to cowbirds at all densities. Other unmeasured effects on cowbird habitat suitability (e.g., reduced cowbird success in edge-dwelling host nests, cowbird despotism at edges) might have affected cowbird habitat selection. Managers attempting to minimize cowbird parasitism on sensitive cowbird hosts should consider that hosts in otherwise less-preferred cowbird habitats (e.g., habitat interiors) are at greater risk of being parasitized where cowbirds become particularly abundant.     

Effects of forest edge on populations of white-footed mice Peromyscus leucopus

Several studies have reported higher densities of white-footed mice in small fragments than in large fragments of eastern deciduous forests. The edge hypothesis states that higher densities in smaller fragments reflect an increase in relative amount of edge habitat, which supports higher densities of mice because of its higher quality. To test this hypothesis we live trapped white-footed mice along edge-to-interior gradients in forest fragments of east-central Illinois. Our results indicated a greater abundance of mice in the forest interior than near the edge, which did not support the edge hypothesis. This pattern could occur because dominant adults hold larger territories of higher quality habitat, thereby reducing density and increasing fitness near the edge (an ideal despotic distribution). We found some evidence of increased reproductive success (juveniles per female) at the edge, but this could reflect density-dependent demographic processes rather than habitat quality. Furthermore, other indicators of dominance (body weight, and reproductive activity) did not show an increase at the edge, and other studies indicate higher prevalence of natural enemies at edges, which could account for lower densities there. Reduced competition from larger rodents and reduced predation could cause higher densities in small fragments but the distributions of competitors and predators do not strongly support these hypotheses. We suggest two additional hypotheses that could account for greater densities in smaller fragments: 1) estimates of high densities could be artifacts of the large effect that a few captures can have on density estimates for very small fragments, and 2) densities in smaller fragments are overestimated because mice use a relatively larger area of surrounding habitat as fragment size decreases.     

Population dynamics in two-patch environments: Some anomalous consequences of an optimal habitat distribution

The effect of dispersal on population size and stability is explored for a population that disperses passively between two discrete habitat patches. Two basic models are considered. In the first model, a single population experiences density-dependent growth in both patches. A graphical construction is presented which allows one to determine the spatial pattern of abundance at equilibrium for most reasonable growth models and rates of dispersal. It is shown under rather general conditions that this equilibrium is unique and globally stable. In the second model, the dispersing population is a food-limited predator that occurs in both a source habitat (which contains a prey population) and a sink habitat (which does not). Passive dispersal between source and sink habitats can stabilize an otherwise unstable predator-prey interaction. The conditions allowing this are explored in some detail. The theory of optimal habitat selection predicts the evolutionarily stable distribution of a population, given that individuals can freely move among habitats so as to maximize individual fitness. This theory is used to develop a heuristic argument for why passive dispersal should always be selectively disadvantageous (ignoring kin effects) in a spatially heterogeneous but temporally constant environment. For both the models considered here, passive dispersal may lead to a greater number of individuals in both habitats combined than if there were no dispersal. This implies that the evolution of an optimal habitat distribution may lead to a reduction in population size; in the case of the predator-prey model, it may have the additional effect of destabilizing the interaction. The paper concludes with a discussion of the disparate effects habitat selection might have on the geographical range occupied by a species.     

Habitat‐dependent population regulation in an irrupting population of long‐nosed bandicoots (Perameles nasuta)

We used isodars to analyse habitat‐dependent population regulation by long‐nosed bandicoots Perameles nasuta during an irruption and subsequent population crash in three habitats (heath, woodland and forest) at Booderee National Park, south‐eastern Australia. Specifically, we aimed to see whether patterns of habitat‐dependent population regulation matched a priori estimates of quantitative and qualitative differences between habitats. We also tested if habitat preference changed between the increasing and decreasing phase of the irruption as predicted by the reciprocating dispersal theory. Quantitative differences in habitat quality were indexed by the relative abundance of the main food of long‐nosed bandicoots (terrestrial invertebrates), while qualitative differences were indexed by the availability of refuge from predation (vegetation understorey density). One index of fitness, body weight, was highest in forest, and lowest in heath, suggesting an ideal despotic model of habitat selection. Over the entire course of the irruption, there was density‐dependent habitat selection with forest and woodland both quantitatively superior to heath. This reflected the overall abundance of invertebrates with highest abundance in woodland and forest and less in heath. Isodar analysis also revealed that although forest was quantitatively better than heath and equivalent to woodland it was qualitatively poorer than either habitat. Heath had a higher density of understorey than woodland and woodland having a higher density of understorey than forest giving crossover population regulation. When the increasing and declining phase of the irruption were analysed separately, no habitat was quantitatively superior to any other during either phase. The lack of switching in preference between habitats from the increasing to the declining phase of the irruption and the virtual absence of any dispersal by adults, does not support the reciprocating dispersal hypothesis.     

How can maladaptive habitat choice generate source‐sink population dynamics?

Several theoretical models have been proposed to describe population dynamics in a spatially heterogeneous environment. The source-sink model is among the most popular. Diffendorfer recently summarized its assumptions and predictions. Given the model reviewed, he argued that source-sink population dynamics arises if dispersal is somehow constrained. I offer an additional mechanism by suggesting that source-sink population dynamics can be generated by anthropogenic changes in landscapes that occur so quickly that organisms no longer make optimal habitat selection decisions. Individuals select the same habitats as their ancestors but these decisions no longer provide high fitness because of human-induced changes in habitat quality, such as increased rates of predation and/or parasitism. Provided that some of the habitats selected are turned by human-induced changes into sink habitats, source-sink population dynamics can emerge.     

The ecological-evolutionary interplay: density-dependent sexual selection in a migratory songbird

Little is understood about how environmental heterogeneity influences the spatial dynamics of sexual selection. Within human-dominated systems, habitat modification creates environmental heterogeneity that could influence the adaptive value of individual phenotypes. Here, we used the gray catbird to examine if the ecological conditions experienced in the suburban matrix (SM) and embedded suburban parks (SP) influence reproductive strategies and the strength of sexual selection. Our results show that these habitats varied in a key ecological factor, breeding density. Moreover, this ecological factor was closely tied to reproductive strategies such that local breeding density predicted the probability that a nest would contain extra-pair offspring. Partitioning reproductive variance showed that while within-pair success was more important in both habitats, extra-pair success increased the opportunity for sexual selection by 39% at higher breeding densities. Body size was a strong predictor of relative reproductive success and was under directional selection in both habitats. Importantly, our results show that the strength of sexual selection did not differ among habitats at the landscape scale but rather that fine-scale variation in an ecological factor, breeding density, influenced sexual selection on male phenotypes. Here, we document density-dependent sexual selection in a migratory bird and hypothesize that coarse-scale environmental heterogeneity, in this case generated by anthropogenic habitat modification, changed the fine-scale ecological conditions that drove the spatial dynamics of sexual selection.     

Dispersal among habitats varying in fitness: reciprocating migration through ideal habitat selection

Current evolutionary models of dispersal set the ends of a continuum where the number of individuals emigrating from a habitat either equals the number of individuals immigrating (balanced dispersal) or where emigrants flow from a source habitat to a corresponding sink. Theories of habitat selection suggest a more sophisticated conditional strategy where individuals disperse from habitats where they have the greatest impact on fitness to habitats where their per capita impact is lower. Asymmetries between periods of population growth and decline result in a reciprocating dispersal strategy where the direction of migration is reversed as populations wax and wane. Thus, for example, if net migration of individuals flows from high- to low-density habitats during periods of population growth, net migration will flow in the opposite direction during population decline. Stochastic simulations and analytical models of reciprocating dispersal demonstrate that fitness, carrying capacity, stochastic dynamics, and interference from dominants interact to determine whether dispersal is balanced between habitats, or whether one habitat or the other acts as a net donor of dispersing individuals. While the pattern of dispersal may vary, each is consistent with an underlying strategy of density-dependent habitat selection.     

Population differences in female resource abundance, adult sex ratio, and male mating success in Dendrobates pumilio

In this study I examined the relationship among abundance ofreproductive resources, population density, and adult sex ratioin the strawberry dart-poison frog, Dendrobates pumilio, andhow these variables in turn influence the mating system, malereproductive success, and sexual selection. I studied the matingbehavior in two populations of D. pumilio living in a primaryand secondary rainforest on the Caribbean slope of Costa Rica.The abundance of tadpole-rearing sites (reproductive resourcesfor females) was approximately 10-fold higher in the secondary forest. Accordingly, the population density was higher and theadult sex ratio was strongly female biased in the secondaryforest, whereas the adult sex ratio was even in the primaryforest. The female-biased sex ratio was associated with a higherlevel of polygyny and higher male mating and reproductive successin the secondary forest. In contrast, the level of polyandrydid not differ between habitats. As expected, the opportunityfor sexual selection on male mating success was lower in thesecondary forest, the habitat with high female density. Inconclusion, my results suggest that ecological variables suchas resource availability have a great impact on the matingsystem and sexual selection through their effect on population structure. Moreover, the results of this study give furtherevidence that the opportunity for sexual selection is influencedby the adult sex ratio and hence by the operational sex ratioin a population.     

Density-dependent habitat selection in muskrats: a test of the ideal free distribution model

Summary Two predictions of the ideal free distribution model, a null hypothesis of habitat selection, were examined using free-ranging muskrats. We rejected the prediction that the proportion of the animals found in each of five habitats was independent of population size. Data on over-winter occupancy of muskrat dwellings tend also to refute the prediction of equal fitness reward among habitats. Habitat type and water-level had a profound effect on the suitability of a site for settlement. We concluded that the observed pattern of muskrat distribution followed more closely an ideal despotic distribution where some individuals benefited from a higher fitness because of resource monopolization. Current theories of density-dependent habitat selection, which assume an ideal free distribution, would not apply to muskrats and possibly to many other mammal species.     

Evidence for density-dependent habitat occupancy at varying scales in an expanding bird population

Understanding factors shaping the spatial distribution of animals is crucial for the conservation and management of wildlife species. However, few studies have investigated density-dependent habitat selection in wild populations in non-equilibrium conditions and at varying spatial scales. Here, we investigated density-dependent habitat selection at varying spatial scales in an increasing white stork Ciconia ciconia population using a long term data set in western France. During a 16-year study period, the breeding population density increased from 0.66 nests per 100 km² to 6.6 nests per 100 km². At the beginning of the colonisation of the area settlement probability of storks was mainly positively affected by grasslands located near wetlands. Areas with intensive or moderately intensive agriculture were extremely unlikely to be occupied by breeding birds. However, selection for the initially preferred habitats faded as stork density increased although the proportions of habitat types remained unchanged. At the same time selection for initially less favoured habitats had increased. Moreover, we found that the spatial scale of selection for each foraging habitat variable was consistent over time. Our results suggest that snapshot analyses of resource selection in populations at high density may be misleading for population conservation or management. In contrast, a longitudinal approach to resource selection can be a valuable tool for understanding resource limitation.     

Spatio-temporal shifts in gradients of habitat quality for an opportunistic avian predator

We used the conceptual framework of the theory of natural selection to study breeding habitat preferences by an opportunistic avian predator, the black kite Milvus migrans. In Europe, black kite populations are mostly found near large networks of aquatic habitats, usually considered optimal for foraging and breeding. We hypothesized that proximity to wetlands could vary among individuals and affect their fitness, and thus be subject to natural selection. We tested the hypothesis first on a population on Lake Lugano (Italian pre‐Alps) which has been monitored for nine years, and then on seven other populations, each studied for four–five years, located along a continuum of habitat from large water bodies to scarce aquatic habitat of any kind. In the Lake Lugano population, black kite abundance was negatively related to distance to the lake in all the nine years of study, consistent with long‐term natural selection. There was evidence of ongoing directional selection on strategic nest location in three of the years, and evidence of stabilizing selection in two years. In eight of the nine years the trend was for a linear increase in fitness with increasing proximity to the lake. At the population level, results were consistent with adaptive habitat choice in relation to the previous year's spatial variation in fitness: higher associations between fitness and distance to the lake (i.e. higher selection gradients) resulted in higher density variations in the following year, in turn related to the availability of fish, the main local prey. The progressive decline of inland pairs and increase in the density of lakeshore pairs caused a directional long‐term trend of declining mean distance to the lake. Breeding near aquatic habitats was associated with higher foraging success, and higher frequency and biomass of prey deliveries to offspring. There was weak evidence of selection in other populations. The inland‐wetland gradient of habitat quality may have been affected by predation risk, as estimated by density of a major predator of adults and nestlings, the eagle owl Bubo bubo. Behavioral decisions at the level of the individual probably translated into population effects on density and distribution at various spatial scales. Populations in optimal habitats showed higher density and produced six times as many young per unit space as those in sub‐optimal habitats.     

Adult survival and reproductive rate are linked to habitat preference in territorial,year‐round resident Song Sparrows Melospiza melodia

Individual animal fitness can be strongly influenced by the ability to recognize habitat features which may be beneficial. Many studies focus on the effects of habitat on annual reproductive rate, even though adult survival is typically a greater influence on fitness and population growth in vertebrate species with intermediate to long lifespans. Understanding the effects of preferred habitat on individuals over the annual cycle is therefore necessary to predict its influences on individual fitness. This is particularly true in species that are resident and territorial year‐round in the temperate zone, which may face potential trade‐offs between habitat that maximizes reproduction and that which maximizes non‐breeding season (‘over‐winter’) survival. We used a 37‐year study of Song Sparrows Melospiza melodia residing territorially year‐round on a small island to examine what habitat features influenced adult over‐winter survival, how site‐specific variation in adult survival vs. annual reproductive rate influenced long‐term habitat preference, and if preferred sites on average conferred higher individual fitness. Habitat features such as area of shrub cover and exposure to intertidal coastline predicted adult over‐winter survival independent of individual age or sex, population size, or winter weather. Long‐term habitat preference (measured as occupation rate) was better predicted by site‐specific annual reproductive rate than by expected over‐winter survival, but preferred sites maximized fitness on average over the entire annual cycle,. Although adult over‐winter survival had a greater influence on population growth (λ) than did reproductive rate, the influence of reproductive rate on λ increased in preferred sites because site‐specific variation in reproductive rate was higher than variation in expected over‐winter survival. Because preferred habitats tended to have higher mean site‐specific reproductive and adult survival rates, territorial birds in this population do not appear to experience seasonal trade‐offs in preferred habitat but are predicted to incur substantial fitness costs of settling in less‐preferred sites.