Density-dependent habitat selection: Testing the theory with fitness data

Summary According to density-dependent habitat selection theory, reproductive success should decline with increased density. Fitness should be similar between habitats if habitat selection follows an ideal free distribution; fitness should be dissimilar between habitats if habitat selection is modified by territorial behavior. I tested these assumptions by examining a variety of fitness estimates obtained from white-footed mice living in nest boxes in forest, forest edge and fencerow habitats in southwestern Ontario. As expected, mean litter size declined with increased population density. Litter sizes, adult longevity and the proportion of adult animals in breeding condition were not significantly different among the three habitats. The success at recruiting at least one offspring to the adult population and the number of recruits per litter were much greater in the forest than in either of the other two habitats. Fitness was thus unequal among habitats and the results confirm both assumptions of density-dependent habitat selection theory for territorial white-footed mice.     

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.     

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.     

Emigration to new habitats by voles: the cost of dispersal paradox

We tested the hypothesis that dispersal and philopatry are components of a mixed evolutionarily stable strategy (ESS). The hypothesis predicts that fitness of dispersers should be equal to that of philopatric individuals. Alternatively, fitness of dispersers could be lower (the resident fitness hypothesis) or greater (the cost of dispersal hypothesis) than that of philopatric individuals. We compared fitness of individuals that moved to new habitats (emigrants) and those that remained within habitat boundaries (residents) in populations of the prairie vole, Microtus ochrogaster, and the meadow vole, M. pennsylvanicus. We established vole populations in four enclosures (). Within each enclosure, voles were free to move between four types of habitats that varied in the availability of supplemental food and the amount of vegetative cover. We analysed two fitness components: the survival rates of all individuals, and pregnancy rates of females. Our study showed that emigrants generally had greater fitness than residents and that the difference in fitness was habitat dependent (i.e. was greater when individuals were emigrating from low-quality habitats than from high-quality habitats). High-food, high-cover habitats were the only habitat types for which fitness of emigrants was lower than that of residents. Similar patterns occurred in both prairie voles and meadow voles. Our results support the cost of dispersal hypothesis.     

Habitat Selection by Prairie Dogs in a Disturbed Landscape at the Edge of Their Geographic Range

ABSTRACT Although habitat attributes of black-tailed prairie dog (Cynomys ludovicianus) colonies have been described for central and northern portions of the species' geographic range, little is known about these associations at the southern edge of this species' distribution. Because high-quality habitats are expected to be scarcer at the edge of the species' geographic range, different patterns of habitat selection might emerge in these populations. We analyzed habitat selection by black-tailed prairie dogs in a human-disturbed mosaic of desert grasslands and shrublands in northwestern Chihuahua, Mexico. We contrasted 151 used and 133 unused habitat units producing 11 case-control logistic regression models to explain site occupancy by prairie dogs with different combinations of environmental variables. Prairie dogs from Chihuahua occupy sites similar in most respects to sites in more northern regions, although these prairie dogs appear to be more tolerant of increased shrub density and reduced herbage cover. We found that site occupancy was best modeled by positive effects of soil moisture level, cover of forbs, cover of unpalatable vegetation, cover of bare ground, and amount of prairie-dog colony area within 1 km and by the inverse of altitude, shrub density, herbage height, and amount of hostile habitat within 1 km. The 2 most significant variables were herbage height and shrub density, which might reflect the prominent role that visibility plays in habitat selection by prairie dogs. In contrast, we found weak evidence that human features have significant impacts on site occupancy by prairie dogs. Our results support the prediction that environmental conditions of sites used by prairie dogs in edge regions partially differ from those observed in more northern latitudes. We suggest that reserve managers focus conservation efforts on areas with short vegetation, low density of shrubs, and high herbage cover, conditions that could be promoted by controlled burns, herbage mowing, and mechanical removal of shrubs.     

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.     

Brown trout spawning habitat selection preferences and redd characteristics in the Credit River, Ontario

Brown trout Salmo trutta redds in the Credit River, Ontario, were enumerated and assessed for physical, location and cover feature characteristics during the 2002 spawning season. Hydraulic habitats were classified on the basis of channel morphology and availability recorded. Combined use and availability data were used to assess habitat selection preferences and test for significant differences. Significant preferences for upstream pool and riffle habitat were found, but all available habitat types were used to some degree. Non‐preferred habitat redds were significantly larger than those located in preferred habitats and more likely to be found in association with woody debris cover. Larger redds were interpreted as indicative of larger fish. The use of non‐preferred downstream habitats by larger fish was further interpreted in the context of overwinter habitat requirements to reflect possible trade‐offs between reproductive and adult over‐winter survival requirements resulting in the selection of habitat for multiple life‐history functions. Results suggest that redd density alone should not be used to infer critical autumn habitat requirements for brown trout.     

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.     

The effects of habitat manipulation on population distribution and foraging behavior in meadow voles

Individuals, free to choose between different habitat patches, should settle among them such that fitness is equalized. Alternatives to this ideal free distribution result into fitness differences among the patches. The concordance between fitnesses and foraging costs among inhabitants of different quality patches, demonstrated in recent studies, suggests that the mode of habitat selection and the resulting fitness patterns may have important implications to the resource use of a forager and to the survival of its prey. We studied how coarse scale selection between habitat patches of different quality and quitting harvest rate in these patches are related to each other and to fine scale patch use in meadow voles (Microtus pennsylvanicus). To demonstrate these relationships, we manipulated habitat patches within large field enclosures by mowing vegetative cover and adding supplemental food according to a 2×2 factorial design. We tracked vole population densities, collected giving‐up densities (GUDs, a measure of patch quitting harvest rate), and monitored the removal of seeds from lattice grids with 1.5 m intervals (an index of fine‐scale space use) in the manipulated habitat patches. Changes in habitat quality induced changes in habitat use at different spatial scales. In preferred habitats with intact cover, voles were despotic and GUDs were low, but increased with the addition of food. In contrast, voles in less‐preferred mowed habitats settled into an ideal free distribution, GUDs were high and uninfluenced by the addition of food. Seed removal was enhanced by the presence of cover but inhibited by supplemental food. Across all treatments, vole densities and GUDs were strongly correlated making it impossible to separate their effects on seed removal rates. However, this relationship broke down in unmowed habitats, where GUDs rather than vole density primarily influenced seed removal by voles. GUDs and seed removal correlated with predation on tree seedlings formerly planted into the enclosures, demonstrating the mechanisms between coarse‐scale habitat manipulations and community level consequences on a forager's prey.     

Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Density-dependent habitat selection between maize cropfields and their borders in two rodent species (Akodon azarae and Calomys laucha) of Pampean agroecosystems

We studied habitat preferences and intra and interspecific density-dependent effects on habitat selection by Akodon azarae and Calomys laucha between maize fields and their adjacent borders, during different developmental stages of the crop. Akodon azarae detected quantitative differences between habitats, using preferentially borders throughout the year, while C. laucha perceived borders and cropfields as quantitatively similar during spring and summer and it detected borders as quantitatively better at the high density period (autumn and winter). These results support the prediction of differential habitat preferences as a model of community organisation at the low density period, while they are consistent with shared habitat preferences during autumn and winter when both species apparently coexist in the better habitat (border). Akodon azarae showed intraspecific density-dependent habitat selection throughout the year, except in spring, while habitat selection by C. laucha was density-dependent in spring, autumn and winter. The effect of interspecific density on habitat selection was detected in both habitats and changed seasonally. The effect of A. azarae over C. laucha by resources exploitation was detected in borders, while competitive effects of C. laucha over A. azarae was observed within cropfields. Both species were more affected by exploitation competition than interference, which was more common in borders than in maize fields. We conclude that seasonally have a profound effect in habitat selection of these species because it changes the intensity of intra and interspecific competition and affects different habitat preferences and basic suitability of habitats. This revised version was published online in November 2006 with corrections to the Cover Date.     

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.     

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.     

The importance of spatial scale in habitat selection by European beaver

We evaluated habitat selection by European beaver Castor fiber L. across a spatial gradient from local (within the family territory) to a broad, ecoregional scale. Based on aerial photography, we assessed the habitat composition of 150 beaver territories along the main water bodies of the Vistula River delta (northern Poland) and compared these data with 183 randomly selected sites not occupied by the species. The beavers preferred habitats with high availability of woody plants, including shrubs, and avoided anthropogenically modified habitats, such as arable lands. Within a single family territory, we observed decreasing woody plant cover with increasing distance from a colony centre, which suggests that beaver habitat preferences depend on the assessment of both the abundance and spatial distribution of preferred habitat elements. We tested the importance of spatial scale in beaver habitat selection with principal coordinates of neighbour matrices analysis, which showed that the geographical scale explained 46.7% of the variation in habitat composition, while the local beaver density explained only 10.3% of this variability. We found two main spatial gradients that were related to the broad spatial scale: first, the most important gradient was related to the largest distances between beaver sites and was independent of woody plant cover and the local beaver site density. The second most important gradient appeared more locally and was associated with these variables. Our results indicate that European beaver habitat selection was affected by different scale‐related phenomena related 1) to central place foraging behaviour, which resulted in the clumped distribution of woody plants within the territory, and 2) local population density and woody plant cover. Finally, 3) habitat selection occurs independently across the largest spatial scale studied (e.g. between watersheds), which was probably due to the limited natal dispersal range of the animals.     

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.     

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.     

Avian fitness consequences match habitat selection at the nest‐site and landscape scale in agriculturally fragmented landscapes

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Habitat selection of breeding brown eared-pheasants (Crossoptilon mantchuricum) in Xiaowutaishan National Nature Reserve, Hebei Province, China

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Nonideal nest box selection by tree swallows breeding in farmlands: Evidence for an ecological trap?

Animals are expected to select a breeding habitat using cues that should reflect, directly or not, the fitness outcome of the different habitat options. However, human‐induced environmental changes can alter the relationships between habitat characteristics and their fitness consequences, leading to maladaptive habitat choices. The most severe case of such nonideal habitat selection is the ecological trap, which occurs when individuals prefer to settle in poor‐quality habitats while better ones are available. Here, we studied the adaptiveness of nest box selection in a tree swallow (Tachycineta bicolor) population breeding over a 10‐year period in a network of 400 nest boxes distributed along a gradient of agricultural intensification in southern Québec, Canada. We first examined the effects of multiple environmental and social habitat characteristics on nest box preference to identify potential settlement cues. We then assessed the links between those cues and habitat quality as defined by the reproductive performance of individuals that settled early or late in nest boxes. We found that tree swallows preferred nesting in open habitats with high cover of perennial forage crops, high spring insect biomass, and high density of house sparrows (Passer domesticus), their main competitors for nest sites. They also preferred nesting where the density of breeders and their mean number of fledglings during the previous year were high. However, we detected mismatches between preference and habitat quality for several environmental variables. The density of competitors and conspecific social information showed severe mismatches, as their relationships to preference and breeding success went in opposite direction under certain circumstances. Spring food availability and agricultural landscape context, while related to preferences, were not related to breeding success. Overall, our study emphasizes the complexity of habitat selection behavior and provides evidence that multiple mechanisms may potentially lead to an ecological trap in farmlands.     

Habitat selection for resting sites by the water opossum (Chironectes minimus) in the Brazilian Atlantic Forest

We evaluated the selection of resting sites occupied by the water opossum Chironectes minimus, between 2004 and 2010, in streams of the Brazilian Atlantic Forest. Fourteen radio-tracked adult (males and females) opossums used natural cavities as resting sites. Opossums selected narrow river stretches and selected their resting sites mainly according to characteristics favoring protection against adverse weather conditions. Likewise, opossums avoided disturbed habitats and established their resting sites in well-preserved riparian forest sites, selecting river sections containing a high density of trees and a high proportion of forest cover between the river banks and 50 m from the river. Besides increasing our knowledge on this species habitat selection, such findings further highlight this species’ sensitivity to human disturbance.