Spatial heterogeneity and scale‐dependent habitat selection for two sympatric raptors in mixed‐grass prairie

Sympatric predators are predicted to partition resources, especially under conditions of food limitation. Spatial heterogeneity that influences prey availability might play an important role in the scales at which potential competitors select habitat. We assessed potential mechanisms for coexistence by examining the role of heterogeneity in resource partitioning between sympatric raptors overwintering in the southern Great Plains. We conducted surveys for wintering Red‐tailed hawk (Buteo jamaicensis) and Northern Harrier (Circus cyanea) at two state wildlife management areas in Oklahoma, USA. We used information from repeated distance sampling to project use locations in a GIS. We applied resource selection functions to model habitat selection at three scales and analyzed for niche partitioning using the outlying mean index. Habitat selection of the two predators was mediated by spatial heterogeneity. The two predators demonstrated significant fine‐scale discrimination in habitat selection in homogeneous landscapes, but were more sympatric in heterogeneous landscapes. Red‐tailed hawk used a variety of cover types in heterogeneous landscapes but specialized on riparian forest in homogeneous landscapes. Northern Harrier specialized on upland grasslands in homogeneous landscapes but selected more cover types in heterogeneous landscapes. Our study supports the growing body of evidence that landscapes can affect animal behaviors. In the system we studied, larger patches of primary land cover types were associated with greater allopatry in habitat selection between two potentially competing predators. Heterogeneity within the scale of raptor home ranges was associated with greater sympatry in use and less specialization in land cover types selected.     

Space use and temporal partitioning of sympatric Tasmanian devils and spotted‐tailed quolls

Sympatric species can minimise interspecific competition by spatial avoidance or by altering their temporal activity to reduce encounter rates. The Tasmanian devil (Sarcophilus harrisii), the largest carnivorous marsupial, coexists with the smaller spotted‐tailed quoll (Dasyurus maculatus) in Tasmania, Australia. Quolls may be susceptible to interspecific competition from devils, because they utilise similar habitats, consume similar prey species and are displaced by devils at food sources. Such competition might cause quolls to spatially or temporally avoid devils. To investigate whether spatial or temporal avoidance occurred, we deployed GPS collars on sympatric devils and quolls and conducted a camera survey at a site in northwest Tasmania where the devil population was not affected by devil facial tumour disease. GPS tracking coincided with the lactation period when devils and quolls had young in dens and continued until weaning occurred. We found little spatial segregation of home range and core area placement between devils and quolls and among devils. Quolls showed more spatial segregation within the sexes than between them. Devils had larger home ranges than quolls. Male devils had larger home ranges than females, but there was no difference in home range size between the sexes of quolls. Females of both species travelled significantly further per night than did males. There was moderate temporal partitioning between the two species: devil activity peaked after dusk and devils remained active until the early morning, while quoll activity showed distinct peaks around dusk and dawn. In conclusion, quolls did not spatially avoid devils but moderate temporal partitioning occurred. It is plausible that quolls are active at different times of the diel cycle to reduce encountering devils, but further studies are needed to resolve the cause of this temporal partitioning.     

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.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

Oviposition habitat selection by container‐dwelling mosquitoes: responses to cues of larval and detritus abundances in the field

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Foraging in a risky environment: a comparison of Bennett's wallabies Macropus rufogriseus rufogriseus (Marsupialia: Macropodidae) and red‐bellied pademelons Thylogale billiardierii (Marsupialia: Macropodidae) in open habitats

Abstract This study aimed to establish whether red‐bellied pademelons (Thylogale billiardierii) and Bennett's wallabies (Macropus rufogriseus rufogriseus) alter their foraging distribution in open habitat, in response to food availability and distance to protective shelter, the latter used as a measure of predation risk. Scat counts were used as a measure of the presence or absence of these macropods over two plantations (Russell and Dunalley). These plantations differed in both their on‐site food and shelter characteristics (the presence or absence of windrows). Logistic regression indicated that at Russell, which had low food availability but the presence of on‐site shelter, probability of scats of both species increased with the percentage cover of both edible and inedible vegetation. The probability of both pademelon and wallaby scats decreased with increasing distance from windrows, but increased with increasing distance from forest at the plantation edge. Logistic regression indicated that at Dunalley, which had high food availability but no on‐site shelter, the probability of scats of both species increased with an increase in the percentage cover of edible vegetation. In relation to predation risk, however, the two species differed in their response. Pademelons exhibited a decrease in scat probability with increasing distance from the forest at the plantation edge, while wallabies showed an increase in scat probability with distance from the forest at the plantation edge. Results indicated some differences in antipredation strategies of the two species, which may be a function of differences in body size.     

Foraging behaviour and habitat partitioning of two sympatric cormorants in Patagonia, Argentina

Radiotelemetry was used to assess the distribution and diving behaviour of Rock Shags Phalacrocorax magellanicus and Red-legged Cormorants Phalacrocorax gaimardi breeding in sympatry, and Rock Shags breeding in isolation. When breeding in sympatry there was little overlap in the foraging locations of the two species, with the highest densities of each species separated by 10 km. Red-legged Cormorants fed significantly closer to the breeding colony than did Rock Shags and undertook shorter foraging trips, making almost twice as many foraging trips per day as Rock Shags. Rock Shags breeding in isolation had a shorter foraging range than the birds breeding in sympatry with Red-legged Cormorants and foraging trip duration was significantly shorter. However, the number of feeding trips per day was similar between areas of sympatry and allopatry. Differences in the foraging ecology of Rock Shags in areas of sympatry and allopatry may be due to interspecific competition, which forces niche differentiation. The distance between foraging sites, the speed of movement of the prey, a species tendency to move into prey-depleted areas and the length of the breeding season (during which the birds are constrained to be in the same area) may play critical roles in determining the extent to which differential area use by competitors is a strategy that benefits both parties.     

Terrestrial activity,movements and spatial ecology of an Australian freshwater turtle,Chelodina longicollis,in a temporally dynamic wetland system

Animal movements, use of space, activity patterns and habitat use are in part determined by proximal factors such as the landscapes they occupy, seasonal or environmental cues and individual attributes such as sex and body size. Using radio‐telemetry and a drift fence, we examined the contribution of these factors to variation in movements, use of space and terrestrial activity in a freshwater turtle, Chelodina longicollis (Testudines: Chelidae), in south‐east Australia. Movements and use of space depended strongly on landscape attributes, while sex and body size were less important. Movements and use of space also varied seasonally and were partly correlated with rainfall. The high overall vagility of turtles, irrespective of sex and adult body size (13.8 ± 2.8 ha (±SE) home range, 2608 ± 305 m total distance moved, 757 ± 76 m range length), probably reflects a common need to be mobile in a landscape characterized by fluctuating resources in temporary wetlands. Use of temporary wetlands also drives C. longicollis into terrestrial habitats for movements between wetlands and extended refuge. Timing of inter‐wetland movements was associated with rainfall, but most notably for immature individuals and for those moving towards temporary wetlands. Movements of adults, evacuation of the drying wetland and periods of extended refuge (i.e. aestivation) were less dependent upon rainfall if at all. We conclude that movements about and use of the landscape by C. longicollis are under the strong influence of several interacting factors such as patch configuration, seasonal and environmental cues, and in part, body size. We argue that such behaviours are also ultimately under selection from the costs and benefits of these behaviours in the context of fluctuating resources.     

Examination of fine‐scale spatial‐temporal overlap and segregation between two closely related congeners Gadus morhua and Gadus ogac in coastal Newfoundland

The spatial and temporal movement patterns of sympatric juvenile Atlantic cod Gadus morhua and Greenland cod Gadus ogac were studied using high‐resolution radio‐acoustic positioning in a coastal area of Newfoundland during the summers of 2009 and 2010. A total of 20 fish (10 G. ogac and 10 G. morhua) were equipped with acoustic transmitters and monitored for periods up to 23 days. Most fishes showed high site fidelity with mean residence times of 12·4 (G. morhua) and 14·4 days (G. ogac). A few individuals showed a transient use of the study area, ranging distances up to c. 4 km. Mean daily home ranges [95% kernel utilization distributions (KUDs)] and core activity areas were significantly larger for G. morhua (3·8 and 0·5 ha) than for G. ogac (2·7 and 0·3 ha). Home ranges were not related to total length (L_T) for G. morhua but showed a weak positive relationship for G. ogac. Gadus morhua occupied larger areas during the day while G. ogac occupied slightly larger areas at night. Mean rates of movement for G. ogac and G. morhua ranged from 0·83 to 1·24 and 0·76 to 1·76 L_T s^?1, respectively, and were highest during crepuscular periods. Overall, G. morhua were wider ranging, moved at faster rates and were active throughout the diel cycle compared to G. ogac of the same size. It is suggested that differential use of space and activity periods plays an important role in the successful coexistence of these two species.     

Living on the edge: Multiscale habitat selection by cheetahs in a human‐wildlife landscape

Animals select habitats that will ultimately optimize their fitness through access to favorable resources, such as food, mates, and breeding sites. However, access to these resources may be limited by bottom‐up effects, such as availability, and top‐down effects, such as risk avoidance and competition, including that with humans. Competition between wildlife and people over resources, specifically over space, has played a significant role in the worldwide decrease in large carnivores. The goal of this study was to determine the habitat selection of cheetahs (Acinonyx jubatus) in a human‐wildlife landscape at multiple spatial scales. Cheetahs are a wide‐ranging, large carnivore, whose significant decline is largely attributed to habitat loss and fragmentation. It is believed that 77% of the global cheetah population ranges outside protected areas, yet little is known about cheetahs’ resource use in areas where they co‐occur with people. The selection, or avoidance, of three anthropogenic variables (human footprint density, distance to main roads and wildlife areas) and five environmental variables (open habitat, semiclosed habitat, edge density, patch density and slope), at multiple spatial scales, was determined by analyzing collar data from six cheetahs. Cheetahs selected variables at different scales; anthropogenic variables were selected at broader scales (720–1440 m) than environmental variables (90–180 m), suggesting that anthropogenic pressures affect habitat selection at a home‐range level, whilst environmental variables influence site‐level habitat selection. Cheetah presence was best explained by human presence, wildlife areas, semiclosed habitat, edge density and slope. Cheetahs showed avoidance for humans and steep slopes and selected for wildlife areas and areas with high proportions of semiclosed habitat and edge density. Understanding a species’ resource requirements, and how these might be affected by humans, is crucial for conservation. Using a multiscale approach, we provide new insights into the habitat selection of a large carnivore living in a human‐wildlife landscape.     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

Comparing the consequences of natural selection,adaptive phenotypic plasticity,and matching habitat choice for phenotype–environment matching,population genetic structure,and reproductive isolation in meta‐populations

Organisms commonly experience significant spatiotemporal variation in their environments. In response to such heterogeneity, different mechanisms may act that enhance ecological performance locally. However, depending on the nature of the mechanism involved, the consequences for populations may differ greatly. Building on a previous model that investigated the conditions under which different adaptive mechanisms (co)evolve, this study compares the ecological and evolutionary population consequences of three very different responses to environmental heterogeneity: matching habitat choice (directed gene flow), adaptive plasticity (associated with random gene flow), and divergent natural selection. Using individual‐based simulations, we show that matching habitat choice can have a greater adaptive potential than plasticity or natural selection: it allows for local adaptation while protecting genetic polymorphism despite global mating or strong environmental changes. Our simulations further reveal that increasing environmental fluctuations and unpredictability generally favor the emergence of specialist genotypes but that matching habitat choice is better at preventing local maladaptation by individuals. This confirms that matching habitat choice can speed up the genetic divergence among populations, cause indirect assortative mating via spatial clustering, and hence even facilitate sympatric speciation. This study highlights the potential importance of directed dispersal in local adaptation and speciation, stresses the difficulty of deriving its operation from nonexperimental observational data alone, and helps define a set of ecological conditions which should favor its emergence and subsequent detection in nature.     

Habitat selection by feral cats and dingoes in a semi‐arid woodland environment in central Australia

Habitat use by feral cats and dingoes was examined within a heterogeneous semi‐arid woodland site in central Australia over 2 years. Density estimates of feral cats based on tracks were higher in mulga habitat than in open habitat. Isodar analysis implied that this pattern of habitat use by feral cats was consistent with the consumer‐resource model of density‐dependent habitat selection, which is an ideal free solution. The reason why mulga supported higher densities of feral cats was unclear. Foraging success of feral cats may be higher in the mulga because the stalk and ambush hunting tactics typically employed by felids are well suited to dense cover. Mulga may also have offered feral cats more protection from dingo predation. Dingo activity was distributed uniformly across habitats. The dingo isodar was statistically non‐significant, suggesting that habitat selection by dingoes was independent of density.     

Population Structure and Ranging Patterns of <Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis> in Zhouzhi National Nature Reserve,Shaanxi, China

We describe the population structure and ranging patterns of a troop of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) based on a study conducted between November 2002 and November 2003 in Zhouzhi National Nature Reserve, Shaanxi Province, China. The troop comprised several 1-male units and an all-male unit. Opportunistic censuses revealed that there were ≥112 individuals in the troop. The adult sex ratio (male vs. female) was 1:3.7. The ratios of adults to immatures and infants to adult females were 1:0.7 and 1:2, respectively. Via a grid system, we estimated the home range of the troop to be 18.3 km², of which 7.4 km² was the core area. The subjects exhibited distinct seasonal ranging patterns. Their movement across the home range was extensive in spring and restricted in autumn. In addition, reuse of quadrats was highest in winter and lowest in spring in comparison with other seasons. The daily path length (DPL) varied from .75 to 5 km, with a mean of 2.1 km. Seasonal analysis showed that DPL is significantly shorter in winter than in spring or summer; however, there is no significant difference between the DPLs of spring and summer or those of spring and autumn. The monkeys occupied elevations 1500–2600 m above sea level; the annual mean of altitudinal range is 2137 m. Contrary to early studies that reported Rhinopithecus roxellana migrates to lower elevations in winter, we found no evidence supporting a seasonal altitudinal shift. Using the highest troop count and home range estimate, and considering the extent of range overlap between neighboring troops, we calculated the population density and biomass of Rhinopithecus roxellana to be 7.2 individuals/km² and 68.3 kg/km², respectively. The temporal and spatial distribution of food resources may be the most important determinant of ranging behavior in Rhinopithecus roxellana, though understanding the relationship between resource distribution and seasonal range use may require further investigation.     

Home range size of adult Indo‐Pacific bottlenose dolphins (Tursiops aduncus) in a coastal and estuarine system is habitat and sex‐specific

This study examined sex‐specific differences in home range size of adult Indo‐Pacific bottlenose dolphins off Bunbury, Western Australia. We applied a new kernel density estimation approach that accounted for physical barriers to movements. A Bayesian mixture model was developed to estimate a sex effect in home range size with latent group partitioning constrained by association data. A post hoc analysis investigated group partitioning relating to the proportion of time spent in open vs. sheltered waters. From 2007 to 2013, photographic‐identification data were collected along boat‐based systematic transect lines (n = 586). Analyses focused on adult dolphins of known sex (sighted ≥ 30 times; n = 22 males and 34 females). The 95% utilization distributions of males varied between 27 and 187 km² (; 94.8 ± 48.15) and for females between 20 and 133 km² (65.6 ± 30.9). The mixture model indicated a 99% probability that males had larger home ranges than females. Dolphins mostly sighted in open waters had larger home ranges than those in sheltered waters. Home ranges of dolphins sighted in sheltered waters overlapped with areas of highest human activity. We suggest that sex differences in home ranges are driven by male mating strategies, and home range size differences between habitats may be influenced by prey availability and predation risk.     

Habitat selection by blue swallows Hirundo atrocaerulea Sundevall, 1850 breeding in South Africa and its implications for conservation

This study investigated the utilization of mist‐belt grassland habitat by the threatened blue swallow and was conducted over three successive breeding seasons in the Blue Swallow Natural Heritage Site at Kaapsehoop, Mpumalanga, South Africa. Blue swallows significantly preferred wetlands over grasslands for foraging. Sufficient foraging habitat must be within a 1.5 km radius from an active nest site. The minimum size of a pair of blue swallows home range should be 333 ha, consisting at a minimum of grasslands (243 ha or 73%) and wetlands (90 ha or 27%). In order to conserve this threatened species, habitat transformation should not take place within a minimum radius of 1.5 km of any blue swallow nest. Due to the critical dependence of the blue swallow on wetland habitat for foraging, any development outside the 1.5 km radius that would affect hydrology and water quality within this range would need to be considered, and any adverse effect mitigated. Rehabilitation of areas to a grassland/wetland mosaic would rather quickly support foraging, and eventually breeding. Although a better understanding of the dynamics between wetlands and adjacent grasslands regarding blue swallow habitat requirements is needed, action can already be taken, based on our results.     

Population Density and Home Range Size of Red‐Rumped Agoutis (Dasyprocta leporina) Within and Outside a Natural Brazil Nut Stand in Southeastern Amazonia1

This study examined whether the population density and home range size of red‐rumped agoutis were affected by the spatial distribution of Brazil nut trees, at the Pinkaiti Research Station, in eastern Amazonia. Agouti densities in a Brazil nut grove were two‐fold higher and home ranges were half the size than those outside it. This indicates that the large supply of Brazil nuts results in higher densities and smaller home ranges of agoutis in this seasonally dry Amazonian forest.     

Honey Buzzard Pernis apivorus nest‐site selection in relation to habitat and the distribution of Goshawks Accipiter gentilis

The selection of a suitable nest‐site is critical for successful reproduction. Species' preferences for nest‐sites have presumably evolved in relation to local habitat resources and/or interactions with other species. The importance of these two components in the nest‐site selection of the Eurasian Honey Buzzard Pernis apivorus was assessed in two study areas in eastern Austria. There was almost no difference in macro‐ and micro‐habitat features between nest‐sites and random plots, suggesting that Honey Buzzards did not base their choice of nest‐site on habitat characteristics. However, nests were placed significantly further from nests of Northern Goshawk Accipiter gentilis than would be expected if nest‐sites had been chosen at random. Furthermore, in one study area Honey Buzzards appeared to favour areas close to human settlements, perhaps indicating a mechanism to avoid Goshawks, which tend to avoid the proximity of humans. No habitat variable was significantly associated with the loss of Honey Buzzard young, but predation was higher in territories closer to breeding pairs of Goshawks at both study sites. Although Honey Buzzards are restricted to nesting in forests, their choice of nest‐site therefore appears to be largely dictated by the distribution of predators. Studies of habitat association may yield misleading results if the effects of predation risk on distribution are not considered.     

Genetic structure,spatial organization,and dispersal in two populations of bat‐eared foxes

We incorporated radio‐telemetry data with genetic analysis of bat‐eared foxes (Otocyon megalotis) from individuals in 32 different groups to examine relatedness and spatial organization in two populations in South Africa that differed in density, home‐range sizes, and group sizes. Kin clustering occurred only for female dyads in the high‐density population. Relatedness was negatively correlated with distance only for female dyads in the high‐density population, and for male and mixed‐sex dyads in the low‐density population. Home‐range overlap of neighboring female dyads was significantly greater in the high compared to low‐density population, whereas overlap within other dyads was similar between populations. Amount of home‐range overlap between neighbors was positively correlated with genetic relatedness for all dyad‐site combinations, except for female and male dyads in the low‐density population. Foxes from all age and sex classes dispersed, although females (mostly adults) dispersed farther than males. Yearlings dispersed later in the high‐density population, and overall exhibited a male‐biased dispersal pattern. Our results indicated that genetic structure within populations of bat‐eared foxes was sex‐biased, and was interrelated to density and group sizes, as well as sex‐biases in philopatry and dispersal distances. We conclude that a combination of male‐biased dispersal rates, adult dispersals, and sex‐biased dispersal distances likely helped to facilitate inbreeding avoidance in this evolutionarily unique species of Canidae.