A review of conspecific attraction for habitat selection across taxa

Many species across taxa select habitat based on conspecific presence, known as conspecific attraction. Studies that document conspecific attraction typically provide social information (i.e., cues that indicate the presence of a given species) and then determine if a given species is more likely to settle at locations where the social information is provided compared to those locations that do not. Although the number of studies examining conspecific attraction has grown in recent years, a comprehensive review has not yet been undertaken. Here, we conducted a review of the literature and found 151 studies investigating conspecific attraction across eight taxa. We found that conspecific attraction is widespread with between 80% and 100% of studies, depending on taxa, documenting positive associations between habitat selection and the presence of conspecific cues. Conspecific attraction has been documented more frequently in bird and fish species with less attention given to invertebrate and mammal species. We use the patterns we found to (a) provide an overview of the current state of research on conspecific attraction and (b) discuss how important factors, such as cue characteristics and life history traits, may play a role in shaping conspecific attraction patterns within and across taxa.     

Species interactions and population density mediate the use of social cues for habitat selection

1. The perspective that populations and communities are structured by antagonistic interactions among individuals has dominated much of ecology. Yet how animals use social information to guide decisions, such as habitat selection, may be influenced by both positive and negative interactions among individuals. Recent theory also suggests that the way animals use social information may be substantially influenced by population density, which alters the potential costs and benefits of such behaviours. 2. I manipulated cues of two competitors, the dominant least flycatcher Empidonax minimus (Baird & Baird) and the subordinate American redstart Setophaga ruticilla (Linnaeus), to assess the use of conspecific and heterospecific cues during habitat selection, and if population density influences these strategies. The experiment consisted of surveying birds during a pre-treatment year, which allows for the control and testing the effect of baseline densities, and a treatment year, in which treatments were applied just prior to settlement. Treatments included broadcasting songs of flycatchers and redstarts, and were compared with controls. 3. When controlling for pre-treatment densities, bird densities, and to a lesser extent arrival dates, during the treatment year suggested that flycatchers were attracted to both conspecific and heterospecific cues during settlement. Furthermore, attraction was strongest for flycatchers in plots with moderate pre-treatment densities. American redstarts were rare in the study area but showed apparent attraction to conspecifics and avoidance of heterospecifics. 4. These results provide experimental evidence for the use of multiple social cues in habitat selection and suggest that heterospecific attraction may operate under broader contexts than originally envisioned. In such instances, nontarget effects can potentially occur when manipulating social cues to elicit settlement in conservation strategies. The impact of population density on the use of social cues shown here can also influence our understanding of metapopulation dynamics by causing complex threshold effects on the likelihood of rescue, which may influence metapopulation stability and the likelihood of local extinction.     

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.     

Evolution in heterogeneous environments: Effects of migration on habitat specialization

Summary Richard Levins introduced fitness sets as a tool for investigating evolution within heterogeneous environments. Evolutionary game theory permits a synthesis and generalization of this approach by considering the evolutionary response of organisms to any scale of habitat heterogeneity. As scales of heterogeneity increase from fine to coarse, the evolutionary stable strategy (ESS) switches from a single generalist species to several species that become increasingly specialized on distinct habitats. Depending upon the organisms' ecology, the switch from one to two species may occur at high migration rates (relatively fine-grained environment), or may only occur at very low migration rates (coarse-grained environment). At the ESS, the evolutionary context of a species is the entire landscape, while its ecological context may be a single habitat.Evolution towards the ESS can be represented with adaptive landscapes. In the absence of frequency-dependence, shifting from a single strategy ESS to a two strategy ESS poses the problem of evolving across valleys in the adaptive surface to occupy new peaks (hence, Sewell Wright's shifting balance theory). Frequency-dependent processes facilitate evolution across valleys. If a system with a two strategy ESS is constrained to possess a single strategy, the population may actually evolve a strategy that minimizes fitness. Because the population now rests at the bottom of a valley, evolution by natural selection can drive populations to occupy both peaks.     

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

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

内蒙古达赉湖地区赤狐生境选择及生境景观特征分析

近年来,干旱、鼠害及人为对草原的过度开发利用等对达赉湖地区赤狐的数量及分布范围造成了很大影响。借助于主成分分析、Bailey’s判别分析、遥感和地理信息系统对达赉湖自然保护区赤狐的生境选择及生境的景观特征进行了研究。在赤狐家域内采用样线法和样方法,共设置10 m×10 m样方245个(实验样方101个,对照样方144个),并测定样方内的7个生境因子:植被类型、隐蔽级、食物丰富度、雪深、距水源距离、距围栏距离、距居民点距离。对样方内数据进行主成分分析得出,食物和隐蔽性是赤狐生境选择的主要因子,距居民点距离和距围栏距离是赤狐生境选择的次要因子,雪深和距水源距离是赤狐在生境选择中未表现出选择和利用的因子。利用Bailey’s方法和基于赤狐的生境分布图的景观统计得出:赤狐偏好选择柳灌丛和芦苇塘两种生境,这两种生境的总面积约为1093.47 km2,占研究区总面积的14.05%;赤狐随机利用河道、典型草原和草甸草原3种生境类型,这3种生境的总面积约为4721.79 km2,占研究区总面积的60.67%。;回避的生境为冰面和沙化草地,总面积约为1968.09 km2,占研究区总面积的25.29%。基于地理信息系统和Fragstats的景观特征分析得出,赤狐最适宜生境的面积最少,斑块数量最多,平均斑块面积最小,平均形状指数最小,平均斑块距离最大;其次是较适宜生境和不适宜生境;一般适宜生境的面积最多,斑块数量最少,平均斑块面积和平均形状指数最大,平均斑块距离最小。赤狐多分布于斑块较大的适宜其生存的生境或分布于由这些斑块形成的生境斑块镶嵌体中。     

Spatial heterogeneity in habitat selection: Nest site selection by greater prairie-chickens

Ecological relationships of animals and their environments are known to vary spatially and temporally across scales. However, common approaches for evaluating resource selection by animals assume that the processes of habitat selection are stationary across space. The assumption that habitat selection is spatially homogeneous may lead to biased inference and ineffective management. We present the first application of geographically weighted logistic regression to habitat selection by a wildlife species. As a case study, we examined nest site selection by greater prairie-chickens at 3 sites with different ecological conditions in Kansas to assess whether the relative importance of habitat features varied across space. We found that 1) nest sites were associated with habitat conditions at multiple spatial scales, 2) habitat associations across spatial scales were correlated, and 3) the influence of habitat conditions on nest site selection was spatially explicit. Post hoc analyses revealed that much of the spatial variability in habitat selection processes was explained at a regional scale. Moreover, habitat features at local spatial scales were more strongly associated with nest site selection in unfragmented grasslands managed intensively for cattle production than they were in fragmented grasslands within a matrix of farmland. Female prairie-chickens exhibited spatial variability in nest site selection at multiple spatial scales, suggesting plasticity in habitat selection behavior. Our results highlight the importance of accounting for spatial heterogeneity when evaluating the ecological effects of habitat components. © 2013 The Wildlife Society.     

Polymorphism in heterogeneous environments,evolution of habitat selection and sympatric speciation: Soft and hard selection models

Summary The adaptation to a variable environment has been studied within soft and hard selection frameworks. It is shown that an epistatically determined habitat preference, following a Markovian process, always leads to the maintenance of an adaptive polymorphism, in a soft selection context. Although local mating does not alter the conditions for polymorphism maintenance, it is shown that, in that case, habitat selection also leads to the evolution of isolated reproductive units within each available habitat. Habitat selection, however, cannot evolve in the total absence of adaptive polymorphism. This represents a theoretical problem for all models assuming habitat selection to be an initially fixed trait, and means that within a soft selection framework, all the available habitats will be exploited, even the less favourable ones.On the other hand, polymorphism cannot be maintained when selection is hard, even when all individuals select their habitat. Here, the evolution of habitat selection does not need any prerequisite polymorphism, and always leads to the exploitation of only one habitat by the most specialized genotype. It appears then that hard selection can account for the existence of empty habitat and for an easier evolution of habitat specialization.     

中国大陆鸟类栖息地选择研究十年

栖息地选择研究一直是鸟类生态学研究的重要内容之一。通过对2001年1月至2010年12月10年期间中国大陆鸟类学家在国内外期刊上发表的鸟类栖息地选择研究的论文进行分析,对我国鸟类栖息地选择研究提出展望。10年间,我国鸟类学家共发表有关鸟类栖息地选择或利用的论文170篇,共涉及到鸟类10目31科73种。在这10年里,中国大陆有关鸟类栖息地选择或利用的文章持续增长。然而,我国鸟类栖息地选择的研究也存在着如下问题:(1)存在栖息地选择和栖息地利用误用的现象,这种现象在10年内并无明显改善;(2)在选择研究方法时,很少考虑个体的可获得性、种群密度及抽样尺度对栖息地选择的影响;(3)大多数的栖息地选择的论文没有对鸟类的栖息地选择行为进行研究,也缺乏对其选择的适合度背景去进行研究。根据这些问题,对我国未来的鸟类栖息地选择研究提出了建议。     

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.     

A review of habitat selection by the swamp rat,Rattus lutreolus (Rodentia: Muridae)

Abstract This review clarifies important points on habitat selection by the Swamp Rat Rattus lutreolus (Rodentia: Muridae), a species that has been the subject of much research in Australia and has provided a useful model for understanding ecological and biological processes. It also provides an opportunity to cite important earlier research, not readily available through electronic search engines, thus bringing it into current literature to avoid its disappearance into Internet obscurity. We comment on some papers in the literature to correct errors detected and to emphasize the importance of due care in all aspects of a research project, including its reporting. We show that both floristic and structural components have been reported as important to an understanding of habitat and microhabitat selection by R. lutreolus and conclude that it is vegetation density that is of paramount importance. Female R. lutreolus are clearly dominant in driving microhabitat selection, occupying the ‘best’ or densest habitats with male R. lutreolus occupying the next best and Pseudomys or other species, where present occupying the remainder. This demonstrates the important role that intraspecific and interspecific competition play in determining habitat selection. Direct predation and the perception of predation risk may also play a role in habitat selection, again perceived to be pushing individuals towards denser vegetation, representing ‘better cover’. Whether these effects operate as bottom‐up or top‐down needs careful consideration. Climatic variables, such as ENSO‐affecting productivity, and related variables such as temperature and humidity may also play important roles in habitat selection, as can disturbance effects such as wildfire. The relative importance of all of these potential determining factors may vary from place to place, particularly when climatic clines are involved.     

Fire,landscape change and models of small mammal habitat suitability at multiple spatial scales

Fire is an important process in many ecosystems, but inappropriate fire regimes can adversely affect biodiversity. We identified a naturally flammable heathy woodland ecosystem where the use of planned fire had increased the extent of older vegetation, and quantified the abundance of two small native mammals in this landscape (silky mouse Pseudomys apodemoides and heath rat P. shortridgei). We defined four time‐since‐fire (TSF) categories representing a 2‐ to 55‐year post‐fire sequence and, on the basis of a habitat accommodation model, predicted that both species would select younger age‐classes over older ones. We also predicted that (i) much of the variance in vegetation structure would remain unexplained by TSF and (ii) statistical models of mammal abundance and occupancy including structural variables as predictors would be better than models including TSF. Pseudomys apodemoides selected 17‐ to 23‐year‐old sites, while there was no evidence that P. shortridgei selected a particular TSF category, findings that were inconsistent with our predictions. In line with our predictions, relatively large portions of the variance in vegetation structure remained unexplained by TSF (adjustedr² for four structural variables: 0.24, 0.29, 0.35 and 0.57), and in three of four cases there was strong evidence that statistical models of mammal abundance and occupancy including structural variables were better than those including TSF. At the site scale (hectares), P. shortridgei abundance was positively related to the cover of dead material at the base of Xanthorrhoea plants and at the trap scale (metres), the trapability of both species was significantly related to vegetation volume at 0–20 cm. Our findings suggest that TSF may not be a good proxy for either vegetation structure or species abundance/occupancy.     

Difference in habitat selection by different morphs of the land snail Arianta arbustorum

The behaviour of morphs of Arianta arbustorum has been investigated using a new technique involving a paint which fades in daylight. The technique was used to study the response of different morphs to exposure to sunlight at a homogeneous woodland habitat and a heterogeneous open habitat. The yellow morph showed a significantly higher rate of fading than the brown morph. A mark, release and recapture programme was used to analyse the association between frequencies of different morphs recaptured and not recaptured. In both habitats, there was a general tendency for the frequency of recapture to increase with increasing age in both morphs. Habitat selection resulting from different behaviour of different morphs of A. arbustorum is discussed. Morph frequencies can be related to habitat, the brown morphs of A. arbustorum are significantly less frequent in the open habitat. The behavioural responses of Cepaea hortensis , present with A. arbustorum in two other populations, to the exposure of sunshine were also studied. Climatic selection by solar energy associated with different morphs, species and habitat are also discussed.     

Speciation via habitat specialization: the evolution of reproductive isolation as a correlated character

Summary A diverse group of theoretical and empirical studies are integrated into a composite model of sympatric speciation via habitat specialization. It is shown that disruptive selection on a continuous distribution of habitat preference can lead to the evolution of prezygotic reproductive isolation as a correlated character. The form of selection eliminates the major theoretical objections to the process of sympatric speciation. The principal difference between this model and the allopatric model of speciation is that the initial barrier to gene flow between subpopulations is produced by the evolution of gaps in the phenotypic distribution of spatial/temporal habitat use, rather than an extrinsic geographical barrier.     

贵州草海越冬黑颈鹤觅食栖息地选择的初步研究

本文报道了利用Friedman非参数统计方法,研究越冬黑颈鹤(Grus nigricollis)在贵州草海对其觅食栖息地选择的结果。在草海的黑颈鹤越冬觅食栖息地可分为5种——莎草草甸、浅水沼泽、草地、玉米地和蔬菜地。黑颈鹤对莎草草甸的选择性最高,对玉米地的选择性最差,对其余类型栖息地的选择随地点的不同而不同。草海的人为活动很多,是影响黑颈鹤利用和选择栖息地的一个重要因素。本文也对改善黑颈鹤越冬栖息地的管理和保护提出了一些建议。     

Matrix quality and disturbance frequency drive evolution of species behavior at habitat boundaries

Previous theoretical studies suggest that a species' landscape should influence the evolution of its dispersal characteristics, because landscape structure affects the costs and benefits of dispersal. However, these studies have not considered the evolution of boundary crossing, that is, the tendency of animals to cross from habitat to nonhabitat (“matrix”). It is important to understand this dispersal behavior, because of its effects on the probability of population persistence. Boundary‐crossing behavior drives the rate of interaction with matrix, and thus, it influences the rate of movement among populations and the risk of dispersal mortality. We used an individual‐based, spatially explicit model to simulate the evolution of boundary crossing in response to landscape structure. Our simulations predict higher evolved probabilities of boundary crossing in landscapes with more habitat, less fragmented habitat, higher‐quality matrix, and more frequent disturbances (i.e., fewer generations between local population extinction events). Unexpectedly, our simulations also suggest that matrix quality and disturbance frequency have much stronger effects on the evolution of boundary crossing than either habitat amount or habitat fragmentation. Our results suggest that boundary‐crossing responses are most affected by the costs of dispersal through matrix and the benefits of escaping local extinction events. Evolution of optimal behavior at habitat boundaries in response to the landscape may have implications for species in human‐altered landscapes, because this behavior may become suboptimal if the landscape changes faster than the species' evolutionary response to that change. Understanding how matrix quality and habitat disturbance drive evolution of behavior at boundaries, and how this in turn influences the extinction risk of species in human‐altered landscapes should help us identify species of conservation concern and target them for management.     

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.     

Measures of giant panda habitat selection across multiple spatial scales for species conservation

Examining ecological processes across spatial scales is crucial as animals select and use resources at different scales. We carried out field surveys in September 2005, March–September 2006, and April 2007, and used ecological niche factor analysis to determine habitat preferences for the giant panda (Ailuropoda melanoleuca) across 4 spatial scales: daily movement, core range, home range, and seasonal elevational migration. We found that giant pandas prefer conifer forest and mixed forest at higher than average elevation (2,157 m) of study area in the 4 scale models. However, we also observed significant scale differences in habitat selection. The strength of habitat preference increased with scale for the 2 disturbed forests (sparse forest and fragmented forest), and decreased with scale for 0–30° gentle slope and south- and north-facing aspect. Furthermore, habitat suitability patterns were scale-dependent. These findings highlight the need to determine species–environment associations across multiple scales for habitat management and species conservation. © 2012 The Wildlife Society.