Habitat use of Tibetan Eared Pheasant Crossoptilon harmani flocks in the non-breeding season

Habitat use by Tibetan Eared Pheasant Crossoptilon harmani flocks in shrub vegetation was investigated in the Lhasa area of Tibet during the non-breeding season of 19951996. Home range composition varied considerably among flocks, but stream belts were consistently used as foraging grounds. Slope direction, altitude and vegetation had little effect on habitat selection. In the absence of supplemental food, core range size was positively correlated with flock size, suggesting that food supplementation could support larger flocks. Flocks regularly roosted on the ground at midday at two or three relatively fixed sites within core ranges. At night they used patches of relatively tall, dense vegetation at the year-round sites in areas near cliffs or in hollows. The size of the night-roost site was related to flock size. Our results strongly suggested that both foraging and night-roosting habitats in the shrub environment are crucial to the birds.     

Habitat preferences,seasonal abundance and diets of rodents in Alage,Southern Ethiopia

To understand habitat preferences, seasonal abundance and diets of rodents in wet and dry season surveys were conducted in Alage, Southern Ethiopia. Sherman and snap traps were used to capture rodents from the four habitats: bushland, Acacia woodland, maize and wheat farmlands. A total of 3312 trap nights, from the four trapping habitats, yielded 776 individuals that represented 11 species of rodents. The distribution of rodents varied between habitats and seasons. Wet season rodent abundance was 52.3% while in the dry season it was 47.7%. Seasonal differences in species abundance were insignificant. Bushland habitat had high wet and dry season abundances with 137 and 211 individuals, respectively. Abundance was low in maize farm (57 individuals) in the wet season and wheat farm (10 individuals) in the dry season. Stomach content composition analysis of snap‐trapped rodents from different habitats showed differences between species and across seasons. Six rodent species were recorded as pests on the farmlands in this study area. In conclusion, variation in habitat preferences and diet of rodents in different habitats and across seasons might be due to the role of ground cover and food sources.     

A Hierarchical Analysis of Habitat Selection by Raccoons in Northern Indiana

Abstract: Although numerous studies have examined habitat use by raccoons (Procyon lotor), information regarding seasonal habitat selection related to resource availability in agricultural landscapes is lacking for this species. Additionally, few studies using radiotelemetry have investigated habitat selection at multiple spatial scales or core-use areas by raccoons. We examined seasonal habitat selection of 55 (31 M, 24 F) adult raccoons at 3 hierarchical orders defined by the movement behavior of this species (second-order home range, second-order core-use area, and third-order home range) in northern Indiana, USA, from May 2003 to June 2005. Using compositional analysis, we assessed whether habitat selection differed from random and ranked habitat types in order of selection during the crop growing period (season 1) and corn maturation period (season 2), which represented substantial shifts in resource availability to raccoons. Habitat rankings differed across hierarchical orders, between seasons within hierarchical orders, and between sexes within seasons; however, seasonal and intersexual patterns of habitat selection were not consistent across hierarchical orders of spatial scale. When nonrandom utilization was detected, both sexes consistently selected forest cover over other available habitats. Seasonal differences in habitat selection were most evident at the core-area scale, where raccoon selection of agricultural lands was highest during the maturation season when corn was available as a direct food source. Habitat use did not differ from availability for either sex in either season at the third-order scale. The selection of forest cover across both seasons and all spatial orders suggested that raccoon distribution and abundance in fragmented landscapes is likely dependent on the availability and distribution of forest cover, or habitats associated with forest (i.e., water), within the landscape. The lack of consistency in habitat selection across hierarchical scales further exemplifies the need to examine multiple biological scales in habitat-selection studies.     

How do vigilance and feeding by common cranes Grus grus depend on age,habitat, and flock size?

Animals often spend less time vigilant and more time feeding when foraging in larger groups. This group-size effect does not, however, consider if larger groups differ systematically from smaller ones: Large groups could form in different habitats than small groups or be composed of a different mix of ages or classes than small groups. We examined how habitat differences and flock size and composition explain feeding and vigilance rates in common cranes Grus grus , wintering in holm oak Quercus ilex dehesas of Spain. Flock size and composition were related to habitat type in cranes: flocks formed in areas sown with cereal crops were larger than flocks formed in set aside areas. Vigilance rate depended on habitat but decreased with increasing flock size in a similar way across all habitats. Juveniles were less vigilant than adults and showed little change in vigilance with flock size. Vigilance increased and feeding time decreased over months from November through February. Our results show that vigilance is affected by habitat but that the group size effect on vigilance is not the product of differences between habitats in group size or composition.     

Sexual and seasonal variation in the diet and foraging behaviour of a sexually dimorphic carnivore, the honey badger (Mellivora capensis)

The honey badger, or ratel, Mellivora capensis has not been well studied despite its extensive distribution. As part of the first detailed study, visual observations of nine habituated free-living individuals (five females, four males) were used to investigate seasonal, annual and sexual differences in diet and foraging behaviour. Theory predicts that generalist predators 'switch' between alternative prey species depending on which prey species are currently most abundant, and diet breadth expands in response to decreased availability of preferred food types. There were significant seasonal differences in the consumption of eight prey categories related to changes in prey availability but no seasonal differences in food intake per kg of body mass. As predicted, the cold-dry season diet was characterized by low species richness and low foraging yield but high dietary diversity, while the reverse was true in the hot-dry and hot-wet seasons. In accordance with these predictions, results suggest that the honey badger maintains its intake level by food switching and by varying dietary breadth. Despite marked sexual size dimorphism, male and female honey badgers showed no intersexual differences in prey size, digging success, daily food intake per unit body weight or foraging behaviour. Results do not support the hypothesis that size dimorphism is primarily an adaptation to reduce intersexual competition for food.     

Circannual variation in habitat use of the White-winged Snowfinch Montifringilla nivalis nivalis

High mountain areas are subject to strong seasonal fluctuations, and species inhabiting these particular environments show a high degree of habitat specialization to cope with extreme abiotic conditions. Estimates of habitat use are influenced by the spatial and seasonal scales at which they are evaluated, so studies at multiple scales are important in order to explore adaptive responses to seasonal environments. In the present study, we assessed habitat use of the White-winged Snowfinch Montifringilla nivalis subsp. nivalis (henceforth Snowfinch) during breeding and non-breeding seasons at three different spatial scales (diameters of 100, 250 and 500 m). Although Snowfinches clearly used high-elevation habitats in both seasons, there was evidence that they are less specific during the non-breeding period: the variance explained by habitat and topographical factors was lower in winter than in the breeding season. Moreover, our results suggest that the use of habitat is scale-dependent. This pattern was especially relevant in the breeding season, perhaps because habitat use might be more related to nest-site selection and specific foraging sites to provide food for nestlings. Snowfinches use high mountain habitats throughout the year, probably as a consequence of physiological and morphological specializations typical of high-elevation species, but in winter they show a certain flexibility in habitat use. Snowfinches might thus adopt a flexible specialist strategy. This could represent a trade-off to overcome possible effects on survival, condition and fitness, which can be particularly strong in harsh, unpredictable environments.     

Long-term seasonal variation in the biological traits of benthic-macroinvertebrates in two Mediterranean-climate streams in California, U.S.A.

1. We investigated the seasonal variation of biological traits and the influence of interannual rainfall variability on this pattern. Using long‐term survey data (6–19 years) from an intermittent and a perennial stream in the Mediterranean‐climate region of northern California, we examined 16 fuzzy‐coded biological traits (e.g. maximum size, life cycle duration, and mode of respiration). 2. Seasonal habitat variability is higher in the intermittent stream than in the perennial stream. During the winter and spring wet‐season both streams flood; however, during the summer dry‐season, the intermittent stream forms isolated pools in (occasionally drying completely). 3. Seasonal habitat variability influenced both taxonomic and biological trait composition. Distinct taxonomic communities were present in each season, particularly in the intermittent stream. The intermittent stream also exhibited more seasonal variation in biological traits than the perennial stream. 4. Despite statistically significant seasonal variation, trait composition was relatively stable among seasons in comparison with taxonomic composition and abundance. Taxonomic composition varied considerably between seasons, because of high seasonal and interannual replacement of taxa resulting from seasonal habitat changes. 5. The seasonality of taxonomic composition and abundance was sensitive to interannual rainfall variability. In dry years, the taxonomic composition of communities was more similar between seasons than in wet years, while trait composition was relatively insensitive to rainfall variability. 6. Despite high seasonal variation in abundance and taxonomic composition, biological traits of aquatic macroinvertebrates varied less and exhibited seasonal stability, which may be a result of the unpredictability and harshness of stream environments.     

A review of vertebrate community composition in seasonal forest pools of the northeastern United States

Seasonally-flooded wetlands occur throughout the world and provide important foraging, resting, and breeding habitat for a broad array of organisms. This review summarizes our current understanding of vertebrate community composition at seasonal forest pools in the northeastern United States. These wetlands typically have hydroperiods that range from temporarily flooded to intermittently exposed, which reduces densities of many potential predators (e.g., fish). Current research has shown that pool hydroperiod, canopy closure, vegetation structure within pools, presence of potential predators, and landscape structure surrounding pools are the key factors determining vertebrate diversity at seasonal forest pools. Of 25 species of amphibians in the region, frogs (10 of 12 species) are more likely to breed in seasonal forest pools than salamanders (6 of 13 species). Seven of 10 amphibian species that breed in seasonal forest pools are state-listed as threatened or endangered. Among 27 species of reptiles, 3 of 15 species of snakes, and 6 of 12 species of turtles utilize seasonal pools during at least one stage of their annual cycle. Seasonal forest pools are important foraging and basking habitat for three species of turtles listed as threatened or endangered. Compared to other vertebrate taxa, most species of mammals are habitat generalists, with 50 of 63 mammal species potentially foraging at seasonal pools during part of their annual cycle. Chiroptera (bats; all 9 species) are believed to actively forage at seasonal pools and some Insectivora, particularly Sorex palustris Richardson and S. fumeus (Miller) and Condylura cristata (L.), are detected regularly at seasonal pools. Breeding birds are less likely to utilize seasonal pools than other vertebrate taxa, although 92 of 233 species might forage or breed near seasonal pools. Several species of Anatidae, Rallidae, and some Passeriformes use seasonally flooded pools. All vertebrates that use seasonal forest pools use other habitats during some stage in their life cycle; thus gaining a clear understanding of their habitat requirements is critical to their long-term persistence.     

Functional responses in polar bear habitat selection

Habitat selection may occur in situations in which animals experience a trade-off, e.g. between the use of habitats with abundant forage and the use of safer retreat habitats with little forage. Such trade-offs may yield relative habitat use conditional on the relative availability of the different habitat types, as proportional use of foraging habitat may exceed proportional availability when foraging habitat is scarce, but be less than availability when foraging habitat is abundant. Hence, trade-offs in habitat use may result in functional responses in habitat use (i.e. change in relative use with changing availability). We used logistic and log-linear models to model functional responses in female polar bear habitat use based on satellite telemetry data from two contiguous populations; one near shore inhabiting sea ice within fjords, and one inhabiting pelagic drift ice. Open ice, near the ice edge, is a highly dynamic habitat hypothesised to be important polar bear habitat due to high prey availability. In open ice-polar bears may experience a high energetic cost of movements and risk drifting away from the main ice field (i.e. trade off between feeding and energy saving or safety). If polar bears were constrained by ice dynamics we therefore predicted use of retreat habitats with greater ice coverage relative to habitats used for hunting. The polar bears demonstrated season and population specific functional responses in habitat use, likely reflecting seasonal and regional variation in use of retreat and foraging habitats. We suggest that in seasons with functional responses in habitat use, polar bear space use and population distribution may not be a mere reflection of prey availability but rather reflect the alternate allocation of time in hunting and retreat habitats.     

Habitat use and mating system of the houbara bustard (Chlamydotis undulata undulata) in a semi-desertic area of North Africa: implications for conservation

Studies of the movements and home-ranges of houbara bustards (Chlamydotis undulata undulata) showed sexual and seasonal differences in the use of space, with a polygynous mating system similar to an ‘exploded-lek’ or a ‘resource-defence-polygyny’, that remains undefined. We used the arthropod biomass as an index of the trophic quality of six defined habitats and we radio-tracked 7 females and 13 males to test whether sexual and seasonal variations in habitat use were related to resource availability, and to verify if critical resources for breeding females were monopolised by males. We analysed habitat selection in both sexes separately. We used the habitat type composition of buffer zones around radio-locations to study annual and seasonal habitat selection and to identify preferred habitats, using the chi-square goodness-of-fit test. Habitat use between sexes and between seasons were compared using MANOVA based on log-ratios of habitat proportions. During the year, and in each season, both sexes appeared to be significantly selective for habitats in comparison to their availability. But males avoided esparto grass, while females used all habitats. Habitat use differed between sexes in the breeding season, but not in the non-breeding season. In spring, when food resources were abundant and uniformly distributed in space, males preferred ‘temporarily flooded areas’ and females preferred ‘reg with tall perennials’ that offered both food and cover for brooding. Critical resources were not monopolised by males and the mating system fulfilled the definition of the ‘exploded-lek’. Leks are key sites for reproduction and should be considered as priority areas in further conservation plans.     

A mosaic of opportunities? Spatio-temporal patterns of bat diversity and activity in a strongly humanized Mediterranean wetland

The Baixo Vouga Lagunar (BVL) landscape, in the Portuguese central-west coast, harbours a mosaic of wetland habitat types, interspersed by intensive and extensive agricultural fields, pastures, production forests and urban areas. In this study, we aimed to determine the species composition and the structure of the bat assemblages of the different habitats that constitute this heterogeneous landscape and to investigate seasonal changes in the patterns of bat diversity and activity across habitats. We acoustically sampled bats across 24 sampling sites representative of the eight main habitat types that shape the landscape—Bocage, forests, maize fields, marshlands, reed beds, rice fields, sea rushes and urban settlements. We compared bat richness, diversity and evenness across habitat types and seasons. We analysed habitat-specific and season-specific overall bat activity, and because habitat selection by bats is known to reflect morphological characters, foraging strategies and echolocation call structures, we also analysed the activity of individual species and of eco-morphological guilds. From 1,544 bat-passes recorded, we identified 12 species. Pipistrellus pygmaeus, Pipistrellus pipistrellus and Eptesicus serotinus/Eptesicus isabellinus were the most frequently recorded. Species composition and activity were similar across habitats, whilst exhibiting strong seasonal dynamics within habitats. Our results suggest that the mosaicism of the landscape provides several opportunities for bats, enabling them to explore different resources in distinct habitat patches. However, it may also reflect a forced exploitation of less optimal habitats and resources by bats, due to the scarcity of opportunities provided by fragmented landscapes.     

Dynamics of habitats and macroinvertebrate assemblages in rivers of the Australian dry tropics

1. The dry tropics are characterised by episodic summer rainfall such that the majority of annual river flow occurs in a short period of time. This dryland hydrological cycle leads to variably connected channels and waterholes along the length of a river bed. 2. We investigated the seasonal changes in biophysical characteristics and macroinvertebrate assemblage composition in dry‐tropics rivers at 15 sites on four rivers, each sampled five times (representing one annual hydrological cycle), in the Burdekin catchment, north Queensland, Australia. 3. Assemblages and their temporal trajectories differed among seasons, sites and habitats, even within the same habitat and/or river. Wet season flooding did not appear to ‘reset’ assemblages, with post‐wet season assemblages differing between years. 4. We found no consistent pattern in taxonomic richness over time, and sites within rivers showed no consistent convergence or divergence (i.e. turnover) in macroinvertebrate assemblage composition. However, biophysical variables associated with the rigours of the late dry season had significant effects on macroinvertebrate assemblages, highlighting the variable and often harsh conditions of dry‐tropics rivers. Underlying these patterns were different resistance and resilience traits of invertebrates (such as colonisation and establishment abilities), as well as the local‐scale effects of biophysical variables. 5. The dynamic nature of dryland rivers presents major challenges to monitoring programmes, and our results suggest a more complex scenario for monitoring and management than previously described.     

Context-dependent specialisation drives temporal dynamics in intra- and inter-individual variation in foraging behaviour within a generalist bird population

Individual niche variation is common within animal populations, and has significant implications for a wide range of ecological and evolutionary processes. However, individual niche differences may also temporally vary as a result of behavioural plasticity. While it is well understood how niche variation is affected by changes in resource availability, comparatively little is known about the extent to which individual niche differences may vary within the annual cycle due to internal drivers. Here, we assess how time- and energy-constraints imposed by incubating and brood rearing affect inter- and intra-individual variation in the foraging behaviour of lesser black-backed gulls, a generalist seabird with strong individual niche variation. To this end, we compared daily foraging trips of 22 breeding and 23 non-breeding GPS-tracked adult gulls from two colonies in the Southern Bight of the North Sea over the course of the breeding season. We find that breeding birds, unlike non-breeding ones, did indeed alter their foraging behaviour during the breeding season. Both sexes reduced their searching effort by increasingly revisiting earlier foraging locations, allowing for shorter and more frequent foraging trips. Breeding females also showed pronounced shifts in their habitat use and strongly specialised on urbanised foraging habitats throughout the breeding season. Hence, while individual variation in habitat use remained largely consistent within non-breeders and in breeding males, individual variation among breeding females almost completely disappeared. Female lesser black-backed gulls are on average smaller, and therefore often outcompeted by males for the most profitable food sources. The temporal specialisation on spatially reliable anthropogenic food sources during breeding hence suggests a complex interplay between intrinsic competitive constraints, resource reliability and shifting time- and energy budges in shaping temporal dynamics in individual niche variation within our study population.     

Habitat selection and activity patterns in Alpine mountain hare (Lepus timidus varronis)

We investigated activity patterns and habitat use of 34 radio-tracked mountain hares (Lepus timidus) in the Italian Alps. We first showed that hares were nocturnal and that activity patterns changed seasonally in parallel with circadian rhythms. We predicted that day home ranges will include suitable resting (shelter) habitats, and night home ranges will primarily include suitable foraging habitats. A hare's night-range was larger than its day-range. On average, night and day ranges overlapped by 36%, suggesting that selective pressures affecting space use were, at least partly, different at night than day. Dwarf mountain-pine was the most preferred habitat in all seasons and was selected both for active behaviour (night) and resting (day) and hares avoided the most open habitats. Exploring the effects of season, time of day (day vs. night) and site, we found that habitat use by mountain hares did not differ between seasons or between the active (night) and resting (day) period of circadian cycle. Also, we found no effects of differences in landscape structure (habitat patchiness and heterogeneity) on the patterns of habitat selection. Hares always preferred the dense, forested habitats, which seemed to provide food resources as well as shelter from predators throughout the year.     

Urbanization and the temporal patterns of social networks and group foraging behaviors

Urbanization causes dramatic and rapid changes to natural environments, which can lead the animals inhabiting these habitats to adjust their behavioral responses. For social animals, urbanized environments may alter group social dynamics through modification of the external environment (e.g., resource distribution). This might lead to changes in how individuals associate or engage in group behaviors, which could alter the stability and characteristics of social groups. However, the potential impacts of urban habitat use, and of habitat characteristics in general, on the nature and stability of social associations remain poorly understood. Here, we quantify social networks and dynamics of group foraging behaviors of black‐capped chickadees (N = 82, Poecile atricapillus), at four urban and four rural sites weekly throughout the nonbreeding season using feeders with radio frequency identification of individual birds. Because anthropogenic food sources in urban habitats (e.g., bird feeders) provide abundant and reliable resources, we predicted that social foraging associations may be of less value in urban groups, and thus would be less consistent than in rural groups. Additionally, decreased variability of food resources in urban habitats could lead to more predictable foraging patterns (group size, foraging duration, and the distribution of foraging events) in contrast to rural habitats. Networks were found to be highly consistent through time in both urban and rural habitats. No significant difference was found in the temporal clumping of foraging events between habitats. However, as predicted, the repeatability of the clumping of foraging events in time was significantly higher in urban than rural habitats. Our results suggest that individuals living in urban areas have more consistent foraging behaviors throughout the nonbreeding season, whereas rural individuals adjust their tactics due to less predictable foraging conditions. This first examination of habitat‐related differences in the characteristics and consistency of social networks along an urbanization gradient suggests that anthropic habitat use results in subtle modifications in social foraging patterns. Future studies should examine potential implications of these differences for variation in predation risk, energy intake, and information flow.     

Seasonal habitat selection and space use by a semi‐free range herbivore in a heterogeneous savanna landscape

Understanding factors that influence habitat selection in heterogeneous landscapes is fundamental for establishing realistic models on animal distribution to inform rangeland management. In this study, we tested whether seasonal variation in habitat selection within the home range of a large herbivore was influenced by constraints such as, distances from water and central place using semi‐free range cattle (Bos taurus) as a case study. We also tested whether shifts in space use over time were dependent on spatial scale and on the overall abundance of resources. We predicted that distance from water significantly influenced dry season habitat selection while the influence of the central place on habitat selection was season‐independent. We also predicted that shifts in space use over time were spatial scale‐dependent, and that large herbivores would include more diverse habitats in their home ranges during the dry season, when water and food resources are less abundant. Multinomial logit models were used to construct habitat selection models with distances from water and central place as habitat‐specific constraints. Results showed significant variations in habitat selection between the dry and wet season. As predicted, the effect of distance from central place was season‐independent, while the effect of water was not included in the top dry season models contrary to expectation. A diverse range of habitats were also selected during the dry season including agricultural fields. Results also indicated that shifts in space use were spatial scale dependent, with core areas being more sensitive to changes than the home range. In addition, shifts in space use responded to temporal changes in habitat composition. Overall, our results suggest that semi‐free range herbivores adopt different foraging strategies in response to spatial‐temporal changes in habitat availability.     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

Samango Monkeys (<Emphasis Type="Italic">Cercopithecus albogularis labiatus</Emphasis>) Manage Risk in a Highly Seasonal,Human-Modified Landscape in Amathole Mountains,South Africa

Wild species use habitats that vary in risk across space and time. This risk can derive from natural predators and also from direct and indirect human pressures. A starving forager will often take risks that a less hungry forager would not. At a highly seasonal and human-modified site, we predicted that arboreal samango monkeys (Cercopithecus albogularis labiatus) would show highly flexible, responsive, risk-sensitive foraging. We first determined how monkeys use horizontal and vertical space across seasons to evaluate if high-risk decisions (use of gardens and ground) changed with season, a proxy for starvation risk. Then, during a subsequent winter, we offered equal feeding opportunities (in the form of high-value, raw peanuts) in both gardens and forest to see if this short-term change in food availability and starvation risk affected monkeys’ foraging decisions. We found that during the food-scarce winter, monkeys foraged outside indigenous forest and in gardens, where they fed on exotic species, especially fallen acorns (Quercus spp.), despite potential threats from humans. Nevertheless, and as predicted, when given the choice of foraging on high-value foods in gardens vs. forest during our artificial foraging experiment, monkeys showed a preference for a safer forest habitat. Our experiment also indicated monkeys’ sensitivity to risk in the lower vertical strata of both habitats, despite their previous extensive use of the ground. Our findings support one of the central tenets of optimal foraging theory: that risk of starvation and sensitivity to the variation in food availability can be as important drivers of behavior as risk of predation.     

Density dependent and temporal variability in habitat selection by a large herbivore; an experimental approach

Both density dependent and density independent processes such as climate affect population dynamics in large herbivores. Understanding herbivore foraging patterns is essential to identify the underlying mechanisms behind variation in vital rates. However, very little is known about how animals vary their selection of habitat temporally, alone or in interaction with density during summer. At the foraging scale, we tested using a fully replicated experiment whether domestic sheep Ovis aries stocked at high (80  per  km²) and low (25  per  km²) densities (spatial contrasts) varied their habitat selection temporally over a four year period. We predicted reduced selection of high productivity vegetation types with increasing density, and that seasonal and annual variation in climate would affect this density dependent selection pattern by increasing competition for high quality habitats in late grazing season and in years with poor vegetation development and over time related to vegetation responses to grazing. As predicted from the Ideal free distribution model, selection of high productivity habitat decreased at high density. There was also a marked temporal variation in habitat selection. Selection of the most productive vegetation types declined towards the end of each grazing season, but increased over years both at low and high sheep density. There was only weak evidence for interactions, as selection ratio of highly productive habitats tended to increase more over years at low density as compared to high density. Limited interactive effects of density and annual variation on habitat selection during summer may explain why similar interactions in vital rates have rarely been reported for summer seasons. Our results are consistent with the view that variation in habitat selection is a central mechanism for climate and density related variation in vital rates.