The attraction of the known: the importance of spatial familiarity in habitat selection in wapiti Cervus elaphus

Most studies of habitat selection by large herbivores focus on the resource availability and interactions with other species, but neglect the importance of an animal being familiar with an area due to past use. Yet, studies of the establishment and retention of territories, home ranges, birth sites, and feeding site choices in experimental settings have shown the importance of spatial familiarity at these scales. We used GPS locations of translocated wapiti Cervus elaphus , resource selection functions (RSF), and time-to-return to examine whether previous site use was important for selection of sites by wapiti in west-central Alberta. To construct RSFs, we used logistic regression that included spatial familiarity (presence of a previous GPS location within a 50-m radius) as well as estimates of herbaceous and shrub biomass, elevation, aspect, slope, and predation risk to wapiti from wolf predation, as dependent variables. We found that previous use had a strong positive relationship with subsequent site use, indicating that wapiti were not avoiding previously visited locations, as would be expected if memory of forage depletion (which we did not measure) determined response to familiar locations. Revisited sites were of higher quality, i.e. had more moderate terrain, higher forage, and lower predation risk, than sites that were not revisited, indicating that the selection of familiar locations was likely not the result of avoidance of unfamiliar locations. Finally, animals demonstrated preference for familiar locations that it had visited most recently, indicating that memory (which would decline with time) of higher site quality, rather than high quality alone, influenced selection for familiar locations. We conclude that spatial familiarity is important not only for large scale processes such as selection of home range and territory, but for smaller scale habitat selection and foraging as well.     

Functional responses in the habitat selection of a generalist mega‐herbivore,the African savannah elephant

Resource selection function (RSF) models are commonly used to quantify species/habitat associations and predict species occurrence on the landscape. However, these models are sensitive to changes in resource availability and can result in a functional response to resource abundance, where preferences change as a function of availability. For generalist species, which utilize a wide range of habitats and resources, quantifying habitat selection is particularly challenging. Spatial and temporal changes in resource abundance can result in changes in selection preference affecting the robustness of habitat selection models. We examined selection preference across a wide range of ecological conditions for a generalist mega‐herbivore, the African savanna elephant Loxodonta africana, to quantify general patterns in selection and to illustrate the importance of functional responses in elephant habitat selection. We found a functional response in habitat selection across both space and time for tree cover, with tree cover being unimportant to habitat selection in the mesic, eastern populations during the wet season. A temporal functional response for water was also evident, with greater variability in selection during the wet season. Selection for low slopes, high tree cover, and far distance from people was consistent across populations; however, variability in selection coefficients changed as a function of the abundance of a given resource within the home range. This variability of selection coefficients could be used to improve confidence estimations for inferences drawn from habitat selection models. Quantifying functional responses in habitat selection is one way to better predict how wildlife will respond to an ever‐changing environment, and they provide promising insights into the habitat selection of generalist species.     

Linking trade-offs in habitat selection with the occurrence of functional responses for moose living in two nearby study areas

A species may modify its relative habitat use with changing availability, generating functional responses in habitat selection. Functional responses in habitat selection are expected to occur when animals experience trade-offs influencing their habitat selection, but only a few studies to date have explicitly linked functional responses to the underlying trade-offs faced by the animals. We used data from 39 female moose fitted with GPS telemetry collars in two nearby study areas in Canada to investigate if moose (1) were faced with a food/cover trade-off in habitat selection, as typically acknowledged in the literature, and (2) showed a functional response in their use of food/cover-rich habitats. We also examined how habitat selection patterns varied seasonally, and between study areas. The occurrence of functional responses varied strongly between study areas, and could not always be related to a measurable food/cover trade-off. Functional responses were observed more often in the study area where the environmental conditions were more severe (colder temperatures, higher precipitations, and lower food availability). Selection coefficients were also less variable among individuals in that study area, suggesting that severe environmental conditions may constrain individuals to a few selection tactics and promote the development of functional responses. Moose reacted to the availability of different habitat types in different seasons, reflecting the changing trade-offs faced by the animals. We found considerable behavioral differences between individuals from two adjacent study areas, and therefore recommend caution when extrapolating habitat selection results. We advocate for the wider use of functional responses to identify critical habitats for a species from a management or conservation perspective.     

The interpretation of habitat preference metrics under use�Cavailability designs

Models of habitat preference are widely used to quantify animal–habitat relationships, to describe and predict differential space use by animals, and to identify habitat that is important to an animal (i.e. that is assumed to influence fitness). Quantifying habitat preference involves the statistical comparison of samples of habitat use and availability. Preference is therefore contingent upon both of these samples. The inferences that can be made from use versus availability designs are influenced by subjectivity in defining what is available to the animal, the problem of quantifying the accessibility of available resources and the framework in which preference is modelled. Here, we describe these issues, document the conditional nature of preference and establish the limits of inferences that can be drawn from these analyses. We argue that preference is not interpretable as reflecting the intrinsic behavioural motivations of the animal, that estimates of preference are not directly comparable among different samples of availability and that preference is not necessarily correlated with the value of habitat to the animal. We also suggest that preference is context-dependent and that functional responses in preference resulting from changing availability are expected. We conclude by describing advances in analytical methods that begin to resolve these issues.     

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.     

Scale dependency and functional response in moose habitat selection

Habitat selection can be influenced by the distribution of the habitat types in the landscape as well as net gain in visiting patches of resources, causing individual variation in habitat selection. Moreover, the hypothesis of functional response in habitat selection predicts that the degree of selection of a resource depends on its relative availability. We used radio-telemetry data from individual moose on an island off the coast of northern Norway to evaluate whether the selection of habitat types at the landscape scale differed from the choice of habitat types within the home range, and investigated the functional response in habitat selection by relating individual habitat selection to home range characteristics. At the landscape scale, moose selected for habitat types that provided both good forage and cover, with small differences between sex and age groups. At the home range scale, all individuals selected habitat types that were associated with cover and low human impact. Habitat selection was not modified by local moose density, but was related to home range size at both spatial scales. Larger home ranges contained larger proportions of non-preferred habitat types compared to smaller home ranges. At the home range scale, the selection for a habitat type decreased with its relative availability, indicating a functional response in habitat selection. This suggests that habitat selection is modified by home range size, which influences the availability of habitat types and shapes individual habitat selection patterns. Our results support previous suggestions that analyses of habitat or resource selection should follow a multi-scale approach. Both the relative availability of habitat types as well as individual variation in home range size should be accounted for in order to disentangle the complex mechanisms that contribute to shape patterns of resource selection in animals.     

High interindividual variability in habitat selection and functional habitat relationships in European nightjars over a period of habitat change

An animal's choice of foraging habitat reflects its response to environmental cues and is likely to vary among individuals in a population. Analyzing the magnitude of individual habitat selection can indicate how resilient populations may be to anthropogenic habitat change, where individually varying, broadly generalist populations have the potential to adjust their behavior. We collected GPS point data from 39 European nightjars (Caprimulgus europaeus) at a UK breeding site where restoration measures have altered large areas of habitat between breeding seasons. We calculated individual habitat selection over four breeding seasons to observe changes that might align with change in habitat. We also analyzed change in home range size in line with change in habitat availability, to examine functional relationships that can represent trade‐offs made by the birds related to performance of the habitat. Individual explained more of the variation in population habitat selection than year for most habitat types. Individuals differed in the magnitude of their selection for different habitat types, which created a generalist population composed of both generalist and specialist individuals. Selection also changed over time but only significantly for scrub habitat (60% decrease in selection over 4 years). Across the population, individual home range size was 2% smaller where availability of cleared habitat within the home range was greater, but size increased by 2% where the amount of open water was higher, indicating the presence of trade‐offs related to habitat availability. These results highlight that using individual resource selection and specialization measures, in conjunction with functional responses to change, can lead to better understanding of the needs of a population. Pooling specialist and generalist individuals for analysis could hide divergent responses to change and consequently obscure information that could be important in developing effective conservation strategies.     

Investigating the Effect of Familiarity on Kin Recognition of Three‐Spined Stickleback (Gasterosteus aculeatus)

The ability to discriminate between related and unrelated individuals has been demonstrated in many species. The mechanisms behind this ability might be manifold and depend on the ecological context in which the species lives. In brood‐caring species, both familiarity and phenotype matching are known to be used in kin recognition. However, results of studies disentangling these two phenomena have proved contradictory. We aimed to broaden our knowledge about the mechanisms of kin recognition using shoaling preferences of three‐spined stickleback (Gasterosteus aculeatus) as a model behavior. In our first experiment, focal fish had the choice to shoal either with kin or unfamiliar non‐kin. In half of the trials, kin groups were composed of familiar individuals, while they were unfamiliar in the other half. Focal fish significantly preferred kin as shoaling partner, a result which was not reinforced by familiarity. In our second experiment, focal fish were given the choice between a shoal of familiar kin and a shoal of unfamiliar kin. Here, focal fish did not show any significant preference. These results indicate that familiarity does not impact stickleback's ability to recognize kin. Furthermore, they show that familiarity does not overrule recognition based on phenotype matching or innate recognition, underlining the importance of these mechanisms. Finally, our results lead to the assumption that individual recognition might play a minor role also in non‐kin‐based preferences for familiars.     

Frequency-dependent food selection by arthropods: a review

We review published studies in which arthropods were presented with a choice of food types at a range of relative densities. Twenty-four of 28 studies showed frequency-independent preference; 11 studies also showed frequency-dependent preference. Both pro- and anti-apostatic selection were found. In the light of these studies we discuss possible behavioural mechanisms which produce such selection; the potential benefits to the predators and the consequences of such selection for diversity both within and between species.     

How mechanisms of habitat preference evolve and promote divergence with gene flow

Habitat preference may promote adaptive divergence and speciation, yet the conditions under which this is likely are insufficiently explored. We use individual‐based simulations to study the evolution and consequence of habitat preference during divergence with gene flow, considering four different underlying genetically based behavioural mechanisms: natal habitat imprinting, phenotype‐dependent, competition‐dependent and direct genetic habitat preference. We find that the evolution of habitat preference generally requires initially high dispersal, is facilitated by asymmetry in population sizes between habitats, and is hindered by an increasing number of underlying genetic loci. Moreover, the probability of habitat preference to emerge and promote divergence differs greatly among the underlying mechanisms. Natal habitat imprinting evolves most easily and can allow full divergence in parameter ranges where no divergence is possible in the absence of habitat preference. The reason is that imprinting represents a one‐allele mechanism of assortative mating linking dispersal behaviour very effectively to local selection. At the other extreme, direct genetic habitat preference, a two‐allele mechanism, evolves under restricted conditions only, and even then facilitates divergence weakly. Overall, our results indicate that habitat preference can be a strong reproductive barrier promoting divergence with gene flow, but that this is highly contingent on the underlying preference mechanism.     

Spatial and behavioural measures of social discrimination by captive male zebra finches: Implications of sexual and species differences for recognition research

All bird species reproduce sexually and individuals need to correctly identify conspecifics for successful breeding. Captive zebra finches are a model system for studying the factors involved in species recognition and mate choice. However, male zebra finches’ behavioural responses in a spatial preference paradigm to a range of estrildid finch species, other than domesticated Bengalese finches, remain unknown. We investigated spatial and display responses of male zebra finch subjects to stimulus females between conspecific and four phylogeographically relevant finch species, in addition to female Bengalese finches. Surprisingly, male subjects did not show consistent spatial association with conspecific over heterospecific females. Overall, as predicted by sexual selection theory, the spatial proximity responses of males were less discriminatory compared to female zebra finches’ responses tested previously using the same paradigm. However, male subjects showed consistently more behavioural displays towards female conspecifics than heterospecifics which were positively related to the behavioural display rates of the respective female stimuli. Some male behavioural responses, other than song, also showed significant differences between the different stimulus species and consistently differed across individual test subjects, with the most individual subject variation seen in choice trials between female conspecific and Bengalese finch stimuli. The results are important for the design and interpretation of future behavioural and neurobiological experiments on species recognition systems using the zebra finch as a model species.     

Looking for sexual selection in the female brain

Female mate choice behaviour has significant evolutionary consequences, yet its mechanistic origins are not fully understood. Recent studies of female sensory systems have made great strides in identifying internal mechanisms governing female preferences. Only recently, however, have we begun to identify the dynamic genomic response associated with mate choice behaviour. Poeciliids provide a powerful comparative system to examine genomic responses governing mate choice and female preference behaviour, given the great range of mating systems: from female mate choice taxa with ornamental courting males to species lacking male ornamentation and exhibiting only male coercion. Furthermore, they exhibit laboratory-tractable preference responses without sexual contact that are decoupled from reproductive state, allowing investigators to isolate mechanisms in the brain without physiological confounds. Early investigations with poeciliid species (Xiphophorus nigrensis and Gambusia affinis) have identified putative candidate genes associated with female preference response and highlight a possible genomic pathway underlying female social interactions with males linked functionally with synaptic plasticity and learning processes. This network is positively correlated with female preference behaviour in the female mate choice species, but appears inhibited in the male coercive species. This behavioural genomics approach provides opportunity to elucidate the fundamental building blocks, and evolutionary dynamics, of sexual selection.     

Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use.     

Direct Familiarity Does Not Alter Mating Preference for Sisters in Male Pelvicachromis taeniatus (Cichlidae)

In social species, individuals who grew up together are usually relatives. Therefore, direct familiarity is normally a reliable kin recognition mechanism that is used in many species to discriminate kin from non‐kin. It has been shown in animals and in humans that familiar individuals are rejected as mating partners in order to circumvent potential costs of inbreeding. Here, we tested whether direct familiarity also leads to inbreeding avoidance behaviour in male Pelvicachromis taeniatus, a small socially monogamous cichlid with biparental brood care. In mate choice experiments, reproductively active males were given the choice between familiar sisters and unfamiliar, unrelated females. In a previous study, both sexes of P. taeniatus had preferred unfamiliar full‐sibs over unfamiliar unrelated individuals as mating partners. Here, we show that direct familiarity does not alter the male preference for closely related females. This result is in accordance with theoretical predictions, that inbreeding can be advantageous under certain conditions, and confirms previous findings, that active inbreeding is an adaptive strategy in P. taeniatus.     

Social familiarity and shoal formation in juvenile fishes

The potential influence of social familiarity in shoal‐choice decisions was investigated in two sympatric species of north temperate fishes, juvenile banded killifish Fundulus diaphanus and juvenile bluegill sunfish Lepomis macrochirus. Groups of socially familiar and unfamiliar conspecifics were formed in the laboratory using wild‐caught fishes. Juvenile F. diaphanus demonstrated a strong preference for familiar conspecific shoalmates, whereas juvenile L. macrochirus exhibited no preference for either unfamiliar or familiar conspecific shoalmates. The differential influence of familiarity on shoalmate choice in juveniles of these two species could be due to their different ecologies, local population densities and life histories.     

根田鼠的熟悉性及其自然动情下的配偶选择

报道了在室内雌鼠自然动情条件下根田鼠两性个体配偶选择的3种熟悉性效应的5组实验结果,包括雌性对熟悉和陌生雄鼠、配偶和陌生雄鼠、配偶和熟悉雄鼠的择偶行为实验 , 雄性对熟悉和陌生雌鼠、配偶和陌生雌鼠的择偶行为实验。在实验前, 将1对性成熟的雌雄鼠 (非亲缘关系) 每天关养8 h 共8 d 建立熟悉性; 配对雌雄鼠至少生育1胎则为配偶关系。在 Y 形迷宫内的30 min 实验中, 两个被选择鼠分别以项链拴在横跨选择箱顶端中央的一根铁丝上, 限制它们在各自箱内活动; 而允许异性选择鼠从中立箱自由进入两个选择箱。记录选择鼠对异性被选择鼠的访问、社会探究、攻击、交配和友好行为的频次和时间。经Wilcoxon 关联样本 T 检验发现, 除交配行为不显著外, 雌性根田鼠均选择熟悉性较高的雄鼠; 而雄鼠不具这种选择性。该结果提示雌雄根田鼠的不同择偶行为也许就是反映该种婚配制度特征的直接行为过程。因此, 在择偶行为中, 雌性根田鼠表现的单配性倾向和雄鼠的多配性倾向, 只能以其婚配制度为一雄多雌制的假设来解释。我们的实验还说明在多配制田鼠中, 一雄多雌制与混交制在雌鼠择偶行为上是不同的, 而与单配制的相似或相近, 故简单地比较多配制与单配制则不能反映田鼠亚科动物婚配制度的多样性。     

Functional responses in habitat selection are density dependent in a large herbivore

Habitat selection is complex due to density dependence and functional responses, defined as variation in relative habitat use depending on availability. In this study we unite these concepts by empirically testing for density‐dependent functional responses in habitat selection using a large herbivore, elk Cervus canadensis manitobensis, as a model species. Theory on density‐dependent habitat selection predicts specialised behaviour when densities are low with a gradual switch to generalist behaviour (more even selection of habitat) as competition intensifies. This suggests that functional responses in habitat selection should be positive when competition is low, but that density may have a negative effect on the functional response due to constraining effects of competition on habitat use and availability. We tested this prediction using data from Global Positioning System (GPS) collared elk (n = 33) and empirical data on spatiotemporal variation in local density during winter in Riding Mountain National Park, Manitoba, Canada (2002–2011). As local density increased, winter home range size decreased and the proportion of mixed forest (providing shelter and forage to elk) used and available within the home range also decreased. Our resource selection function (RSF) revealed clear density‐dependent effects in selection, being strongest (or weakest) for high quality (or low quality) habitat types at lowest observed density leading to more even selection as densities increased. The functional response in mixed forest selection was negatively affected by local density. Increasing availability of mixed forest in the home range was associated with higher selection at low density (positive functional response); no effect of availability on selection at moderate density (no functional response); and lower selection as availability increased at high density (negative functional response). Our study demonstrates that this process is largely driven by the negative effect of density on home range size as it constrains use and availability of habitat.     

The effect of familiarity on aggregation and social behaviour in juvenile small spotted catsharks Scyliorhinus canicula

This study was designed to address whether juvenile small spotted catsharks Scyliorhinus canicula aggregate and to determine whether potential aggregation is underpinned by social preferences for conspecifics. Using controlled and replicated experiments, the role of familiarity as a potential mechanism driving aggregation and social behaviour in this species was considered. Observed S. canicula association data compared to null model simulations of random distributions revealed differences in aggregation under different social contexts. Only familiar juvenile S. canicula aggregated more than would be expected from random distribution across their habitat. Familiarity increased the mean number of groups but did not significantly affect mean group size. Significant preference and avoidance behaviour across all groups were also observed. Furthermore, the strength of social attraction, quantified by the mean association index, was significantly higher in groups containing familiar individuals. Mixed familiar and unfamiliar treatments were also conducted to test for within‐ and between‐group effects, finding high variation across replicates with some groups assorting by familiarity and others not. It is believed that this study is the first to examine experimentally the influence of conspecific familiarity on aggregation behaviour in sharks. These results not only imply a functional benefit to aggregation, but also suggest that persistent social affiliation is likely to influence dispersal following hatching in this small benthic elasmobranch.     

Host‐associated divergence in sympatric host races of the leaf beetle Lochmaea capreae: implications for local adaptation and reproductive isolation

Ecological specialization is widely recognized as a major determinant of the emergence and maintenance of biodiversity. We studied two critical facets of specialization – local adaptation and habitat choice – in the host races of the leaf beetle Lochmaea capreae on willow and birch. Our results revealed that there is asymmetric disruptive selection for host use traits, and host races achieved different adaptive sets of life history traits through association with their host plant. Beetles from each host race exhibited food and oviposition preference for their own host plant. Reciprocal transplant displayed significant variation in host acceptance and performance: all families from the willow race rejected the alternative host plant before initiation of feeding and all died on this host plant. By contrast, all families from the birch race accepted willow for feeding, but they consumed less and performed less well. Intriguingly, families that performed well on birch also performed well on willow, suggesting positive genetic correlation rather than genetic trade‐offs. Our results suggest that the major proximal determinant of host specialization in the willow race is the behavioural acceptance of a plant rather than the toxicity of the food resource. However, in the birch race a combination of behavioural host acceptance and performance may play a role in specialization. Our study sheds light on the mechanisms by which divergent host adaptation might influence the evolution of reproductive isolation between herbivorous populations.     

Habitat Selection in Birds and the Role of Early Experience

The role of early experience in habitat selection in linnets (Acanthis cannabina) was tested in automatic habitat cages. Young linnets were kept in groups under three different habitat conditions: spruce-, foliage- and plastic-perches. In a choice test the members of the spruce- and the foliage-group preferred their familiar habitat whereas the “plastic”-group showed this preference only on the first experimental day and then behaved at random on the following days. In a further aviary experiment the females of the spruce-group constructed nests in small spruce trees, whereas the females of the foliage-group constructed nests in small foliage bushes. The early habitat experience seems strongly to influence later habitat and nest-site selection in the linnets.