Breeding habitat selection behaviors in heterogeneous environments: implications for modeling reintroduction

Animal movement and habitat selection behavior are important considerations in ecology, and remain a major issue for successful animal reintroductions. However, simple rules are often used to model movement or focus only on intrinsic environmental cues, neglecting recent insights in behavioral ecology on habitat selection processes. In particular, social information has been proposed as a widespread source of information for habitat evaluation.
We investigated the role of explicit breeding habitat selection strategies on the establishment pattern of reintroduced populations and their persistence. We considered local movement at the scale of a single population. We constructed a spatially-implicit demographic model that considered five breeding habitat selection rules: 1) random, 2) intrinsic habitat quality, 3) avoidance of conspecifics, 4) presence of conspecifics and 5) reproductive success of conspecifics. The impact of breeding habitat selection was examined for different release methods under various levels of environmental heterogeneity levels, for both long and short-lived monogamous species.
When heterogeneity between intrinsic habitat patch qualities is high, the persistence of reintroduced populations strongly depends on habitat selection strategies. Strategies based on intrinsic quality and conspecific reproductive success lead to a lower reintroduction failure risk than random, conspecific presence or avoidance-based strategies. Conspecific presence or avoidance-based strategies may aggregate individuals in suboptimal habitats. The release of adults seems to be more efficient independent of habitat selection strategy.
We emphasize the crucial role of oriented habitat selection behavior and non ideal habitat selection in movement modeling, particularly for reintroduction.     

The impact of directed versus random movement on population dynamics and biodiversity patterns

An improved understanding of dispersal behavior is needed to predict how populations and communities respond to habitat fragmentation. Most spatial dynamic theory concentrates on random dispersal, in which movement rates depend neither on the state of an individual nor its environment and movement directions are unbiased. We examine the neglected dispersal component of directed movement in which dispersal is a conditional and directional response of individuals to varying environmental conditions. Specifically, we assume that individuals bias their movements along local gradients in fitness. Random movers, unable to track heterogeneous environmental conditions, face source-sink dynamics, which can result in deterministic extinction or increase their vulnerability to stochastic extinction. Directed movers track environmental conditions closely. In fluctuating environments, random movers "spread their bets" across patches, while directed movers invest offspring in habitats currently enjoying propitious conditions. The autocorrelation in the environment determines each strategy's success. Random movers permeate entire landscapes, but directed movers are more geographically constrained. Local information constraints limit the ranges of directed movers and introduce a role for historical contingency in determining their ultimate distribution. These geographic differences have implications for biodiversity. Random movement maintains biodiversity through local coexistence, but directed movement favors a spatial partitioning of species.     

“Perchance to dream?”: Assessing the effects of dispersal strategies on the fitness of expanding populations

Unraveling the patterns of animals’ movements is crucial to understanding the basics of biogeography, tracking range shifts resulting from climate change, and predicting and preventing biological invasions. Many researchers have modeled animals’ dispersal under the assumptions of various movement strategies, either predetermined or directed by external factors, but none have compared the effects of different movement strategies on population survival and fitness. In this paper, using an agent-based model with a landscape divided into cells of varying quality, we compare the ecological success of three movement and habitat selection strategies (MHSSs): (i) Smart, in which animals choose the locally optimal cell; (ii) Random, in which animals move randomly between cells without taking into account their quality; (iii) Dreamer, in which animals attempt to find a habitat of dream whose quality is much higher than that of the habitat available on the map. We compare the short-term success of these MHSSs in good, medium and bad environments. We also assess the effect of temporal variation of habitat quality (specifically, winter harshness) on the success of each MHSS. Success is measured in terms of survival rate, dispersal distance, accumulated energy and quality of settled habitat. The most general conclusion is that while survival rate, accumulated energy and quality of settled habitat are affected primarily by overall habitat composition (proportions of different habitat types in the landscape), dispersal distance depends mainly on the MHSS. In medium and good environments, the Dreamer strategy is highly successful: it simultaneously outperforms the Smart strategy in dispersal distance and the Random strategy in terms of the other metrics.     

Guild mobility affects spider diversity: Links between foraging behavior and sensitivity to adjacent vegetation structure

The movement patterns of species may affect their susceptibility to modified habitat structure. It is likely that sedentary species perceive habitat features at smaller spatial extents compared to mobile species, but there is a lack of experimental research on the effects of fine-scale habitat characteristics on organisms of differing mobility. Spiders display two basic mobility levels based on foraging behavior: web-building species are restricted to specific sites whereas active hunters are mobile. We collected spiders inhabiting sagebrush shrubs with a structurally enhanced, unmodified, or removed understory, to examine (1) whether habitat structure in the immediate vicinity of shrubs affected cursorial and web spiders differently in terms of abundance and species richness and (2) which genera most contributed to changes in community composition. Shrubs without understory had reduced cursorial spider densities and species richness compared to shrubs with added and unmodified understories, whereas web spiders lacked significant responses to treatments. Community-level differences based on relative abundance of genera were detected in cursorial spiders but not in web spiders, despite a strong contribution of the web-building genus Theridion to community dissimilarities. Our results support the hypothesis that sedentary organisms may be sensitive to contiguous habitat at finer spatial scales than cursorial organisms, and highlight the risks associated with only collecting local habitat information when studying mobile species.     

The factors that favor adaptive habitat construction versus non‐adaptive environmental conditioning

Adaptive habitat construction is a process by which individuals alter their environment so as to increase their (inclusive) fitness. Such alterations are a subset of the myriad ways that individuals condition their environment. We present an individual‐based model of habitat construction to explore what factors might favor selection when the benefits of environmental alterations are shared by individuals of the same species. Our results confirm the predictions of inclusive fitness and group selection theory and expectations based on previous models that construction will be more favored when its benefits are more likely to be directed to self or near kin. We found that temporal variation had no effect on the evolution of construction. For spatial heterogeneity, construction was disfavored when the spatial pattern of movement did not match the spatial pattern of environmental heterogeneity, especially when there was spatial heterogeneity in the optimal amount of construction. Under those conditions, very strong selection was necessary to favor genetic differentiation of construction propensity among demes. We put forth a constitutive theory for the evolution of adaptive habitat construction that unifies our model with previous verbal and quantitative models into a formal conceptual framework.     

Optimal strategies and complexity: A theoretical analysis of the anti-predatory behavior of the hare

Predator—prey relationships involving rabbits and hares are widely studied at a long-term population level, while the short-term ethological interactions between one predator and one prey are less well documented. We use a physiologically-based model of hare behavior, developed in the framework of artificial intelligence studies, to analyse its sophisticated anti-predatory behavior. The hares use to stand to the fox in order to inform it that its potential prey is alerted. The behavior of the hare is characterized by specific standing and flushing distances. We show that both hare survival probability and body condition depend on habitat cover, as well as on the ability of the predator to approach—undetected—a prey. We study two anti-predatory strategies, one based on the maximization of the survival probability and the other on the maximization of the body conditions of the hare. Despite the fact that the two strategies are not independent, they are characterized by quite different behavioral patterns. Field estimates of flushing and standing distances are consistent with survival maximization. There exists an optimal anti-predatory strategy, characterized by a flushing distance of 20 m and a standing distance of 30 m, which is optimal in a large set of environmental conditions with a sharp fitness advantage with respect to suboptimal strategies. These results improve our understanding of the anti-predatory behavior of the hare and lend credibility to the optimality approach in the behavioral analysis, showing that even for complex organisms, characterized by a large network of internal constraints and feedback, it is possible to identify simple optimal strategies with a large potential for selection.     

Females roam while males patrol: divergence in breeding season movements of pack-ice polar bears (Ursus maritimus)

Intraspecific differences in movement behaviour reflect different tactics used by individuals or sexes to favour strategies that maximize fitness. We report movement data collected from n = 23 adult male polar bears with novel ear-attached transmitters in two separate pack ice subpopulations over five breeding seasons. We compared movements with n = 26 concurrently tagged adult females, and analysed velocities, movement tortuosity, range sizes and habitat selection with respect to sex, reproductive status and body mass. There were no differences in 4-day displacements or sea ice habitat selection for sex or population. By contrast, adult females in all years and both populations had significantly more linear movements and significantly larger breeding range sizes than males. We hypothesized that differences were related to encounter rates, and used observed movement metrics to parametrize a simulation model of male–male and male–female encounter. The simulation showed that the more tortuous movement of males leads to significantly longer times to male–male encounter, while having little impact on male–female encounter. By contrast, linear movements of females are consistent with a prioritized search for sparsely distributed prey. These results suggest a possible mechanism for explaining the smaller breeding range sizes of some solitary male carnivores compared to females.     

Effects of seasonality on brain size evolution: evidence from strepsirrhine primates

Seasonal changes in energy supply impose energetic constraints that affect many physiological and behavioral characteristics of organisms. As brains are costly, we predict brain size to be relatively small in species that experience a higher degree of seasonality (expensive brain framework). Alternatively, it has been argued that larger brains give animals the behavioral flexibility to buffer the effects of habitat seasonality (cognitive buffer hypothesis). Here, we test these two hypotheses in a comparative study on strepsirrhine primates (African lorises and Malagasy lemurs) that experience widely varying degrees of seasonality. We found that experienced seasonality is negatively correlated with relative brain size in both groups, controlling for the effect of phylogenetic relationships and possible confounding variables such as the extent of folivory. However, relatively larger-brained lemur species tend to experience less variation in their dietary intake than indicated by the seasonality of their habitat. In conclusion, we found clear support for the hypothesis that seasonality restricts brain size in strepsirrhines as predicted by the expensive brain framework and weak support for the cognitive buffer hypothesis in lemurs.     

Conservation implications of the refugee species concept and the European bison: king of the forest or refugee in a marginal habitat?

The concept of refugee species provides a theoretical framework towards increasing the predictive power of the ‘declining population paradigm’ through identifying species which are expected to suffer from a declining population syndrome. Using a simple habitat model as a framework, refugee species are defined as those that can no longer access optimal habitat, but are confined to suboptimal habitats, with consequences of decreased fitness and density, and attendant conservation risks. Refugee species may be difficult to detect in the absence of information on prior habitat use and fitness and their observed ecology will be constrained by the habitat limits forced on them. Identification of refugee species, characterisation of pre‐refugee ecology and the restoration of such species to optimal habitat is critical to their successful conservation. The concept is showcased by addressing the conundrum of a large grazing bovid, the European bison Bison bonasus, being managed as a forest specialist, despite its evolutionary background, dental morphology, neonatal behaviour, diet and microhabitat selection being characteristic of a grazing species inhabiting open, grass‐rich habitats. It is hypothesized that a combination of increasing replacement of open steppe by forest cover after the last postglacial period and increasing human pressure forced bison into forests as a refuge habitat. This process was then reinforced through active management of bison in forests as managers committed themselves to the ‘bison as forest species’ paradigm. A research agenda to test this hypothesis using an experimental approach in the conservation management of European bison by introducing populations into diverse habitat types is suggested.     

Do all inter-patch movements represent dispersal? A mixed kernel study of butterfly mobility in fragmented landscapes

1. In times of ongoing habitat fragmentation, the persistence of many species is determined by their dispersal abilities. Consequently, understanding the rules underlying movement between habitat patches is a key issue in conservation ecology. 2. We have analysed mark-release-recapture (MRR) data on inter-patches movements of the Dusky Large Blue butterfly Maculinea nausithous in a fragmented landscape in northern Bavaria, Germany. The aim of the analysis was to quantify distance dependence of dispersal as well as to evaluate the effect of target patch area on immigration probability. For statistical evaluation, we apply a 'reduced version' of the virtual migration model (VM), only fitting parameters for dispersal distance and immigration. In contrast to other analyses, we fit a mixed dispersal kernel to the MRR data. 3. A large fraction of recaptures happened in other habitat patches than those where individuals were initially caught. Further, we found significant evidence for the presence of a mixed dispersal kernel. The results indicate that individuals follow different strategies in their movements. Most movements are performed over small distances, nonetheless involving travelling between nearby habitat patches (median distance c. 480 m). A small fraction (c. 0·025) of the population has a tendency to move over larger distances (median distance c. 3800 m). Further, immigration was positively affected by patch area (I～A(ζ) ), with the scaling parameter ζ = 0·5. 4. Our findings should help to resolve the long-lasting dispute over the suitability of the negative exponential function vs. inverse-power one for modelling dispersal. Previous studies on various organisms found that the former typically gives better overall fit to empirical distance distributions, but that the latter better represents long-distance movement probabilities. As long-distance movements are more important for landscape-level effects and thus, e.g. for conservation-oriented analyses like PVAs, fitting inverse-power kernels has often been preferred. 5. We conclude that the above discrepancy may simply stem from the fact that recorded inter-patch movements are an outcome of two different processes: daily routine movements and genuine dispersal. Consequently, applying mixed dispersal kernels to disentangle the two processes is recommended.     

Should I Stay or Should I Go? A Habitat-Dependent Dispersal Kernel Improves Prediction of Movement

The analysis of animal movement within different landscapes may increase our understanding of how landscape features affect the perceptual range of animals. Perceptual range is linked to movement probability of an animal via a dispersal kernel, the latter being generally considered as spatially invariant but could be spatially affected. We hypothesize that spatial plasticity of an animal''s dispersal kernel could greatly modify its distribution in time and space. After radio tracking the movements of walking insects (Cosmopolites sordidus) in banana plantations, we considered the movements of individuals as states of a Markov chain whose transition probabilities depended on the habitat characteristics of current and target locations. Combining a likelihood procedure and pattern-oriented modelling, we tested the hypothesis that dispersal kernel depended on habitat features. Our results were consistent with the concept that animal dispersal kernel depends on habitat features. Recognizing the plasticity of animal movement probabilities will provide insight into landscape-level ecological processes.     

Visual fields, eye movements, and scanning behavior of a sit-and-wait predator, the black phoebe (Sayornis nigricans)

Foraging mode influences the dominant sensory modality used by a forager and likely the strategies of information gathering used in foraging and anti-predator contexts. We assessed three components of visual information gathering in a sit-and-wait avian predator, the black phoebe (Sayornis nigricans): configuration of the visual field, degree of eye movement, and scanning behavior through head-movement rates. We found that black phoebes have larger lateral visual fields than similarly sized ground-foraging passerines, as well as relatively narrower binocular and blind areas. Black phoebes moved their eyes, but eye movement amplitude was relatively smaller than in other passerines. Black phoebes may compensate for eye movement constraints with head movements. The rate of head movements increased before attacking prey in comparison to non-foraging contexts and before movements between perches. These findings suggest that black phoebes use their lateral visual fields, likely subtended by areas of high acuity in the retina, to track prey items in a three-dimensional space through active head movements. These head movements may increase depth perception, motion detection and tracking. Studying information gathering through head movement changes, rather than body posture changes (head-up, head-down) as generally presented in the literature, may allow us to better understand the mechanisms of information gathering from a comparative perspective.     

Animal ecology meets GPS-based radiotelemetry: a perfect storm of opportunities and challenges

Global positioning system (GPS) telemetry technology allows us to monitor and to map the details of animal movement, securing vast quantities of such data even for highly cryptic organisms. We envision an exciting synergy between animal ecology and GPS-based radiotelemetry, as for other examples of new technologies stimulating rapid conceptual advances, where research opportunities have been paralleled by technical and analytical challenges. Animal positions provide the elemental unit of movement paths and show where individuals interact with the ecosystems around them. We discuss how knowing where animals go can help scientists in their search for a mechanistic understanding of key concepts of animal ecology, including resource use, home range and dispersal, and population dynamics. It is probable that in the not-so-distant future, intense sampling of movements coupled with detailed information on habitat features at a variety of scales will allow us to represent an animal''s cognitive map of its environment, and the intimate relationship between behaviour and fitness. An extended use of these data over long periods of time and over large spatial scales can provide robust inferences for complex, multi-factorial phenomena, such as meta-analyses of the effects of climate change on animal behaviour and distribution.     

How optimal foragers should respond to habitat changes: a reanalysis of the Marginal Value Theorem

The Marginal Value Theorem (MVT) is a cornerstone of biological theory. It connects the quality and distribution of patches in a fragmented habitat to the optimal time an individual should spend exploiting them, and thus its optimal rate of movement. However, predictions regarding how habitat alterations should impact optimal strategies have remained elusive, with heavy reliance on graphical arguments. Here we derive the sensitivity of realized fitness and optimal residence times to general habitat attributes, for homogeneous and heterogeneous habitats, retaining the level of generality of the MVT. We provide new predictions on how altering travel times, patch qualities and/or relative abundances should affect optimal strategies, and study the consequences of habitat heterogeneity. We show that knowledge of average characteristics is in general not sufficient to predict the change in the average rate of movement. We apply our results to examine the conditions under which the optimal strategies are invariant to scaling. We prove a previously conjectured form of invariance in homogeneous habitats, but show that invariances to scaling are not generic in heterogeneous habitats. We also consider the relative exploitation of patches that differ in quality, clarifying the conditions under which it is adaptive to stay longer on poorer patches.     

Prospecting and dispersal: their eco-evolutionary dynamics and implications for population patterns

Dispersal is not a blind process, and evidence is accumulating that individual dispersal strategies are informed in most, if not all, organisms. The acquisition and use of information are traits that may evolve across space and time as a function of the balance between costs and benefits of informed dispersal. If information is available, individuals can potentially use it in making better decisions, thereby increasing their fitness. However, prospecting for and using information probably entail costs that may constrain the evolution of informed dispersal, potentially with population-level consequences. By using individual-based, spatially explicit simulations, we detected clear coevolutionary dynamics between prospecting and dispersal movement strategies that differed in sign and magnitude depending on their respective costs. More specifically, we found that informed dispersal strategies evolve when the costs of information acquisition during prospecting are low but only if there are mortality costs associated with dispersal movements. That is, selection favours informed dispersal strategies when the acquisition and use processes themselves were not too expensive. When non-informed dispersal strategies evolve, they do so jointly with the evolution of long dispersal distance because this maximizes the sampling area. In some cases, selection produces dispersal rules different from those that would be ‘optimal’ (i.e. the best possible population performance—in our context quantitatively measured as population density and patch occupancy—among all possible individual movement rules) for the population. That is, on the one hand, informed dispersal strategies led to population performance below its highest possible level. On the other hand, un- and poorly informed individuals nearly optimized population performance, both in terms of density and patch occupancy.     

Movement patterns,home range size and habitat utilization of the bluespine unicornfish, <Emphasis Type="BoldItalic">Naso unicornis</Emphasis> (Acanthuridae) in a Hawaiian marine reserve

Synopsis We quantified bluespine unicornfish, Naso unicornis, movement patterns, home range size and habitat preferences in a small Hawaiian marine reserve. Bluespine unicornfish were site-attached to home ranges situated within the reserve boundaries and their movements were aligned with topographic features. Two different diel movement patterns (‘commuting’ and ‘foraying’) were observed. Commuters made crepuscular migrations of several hundred meters between daytime foraging areas and nighttime refuge holes. Foraying fish did not partake in crepuscular migrations and utilized refuge holes both day and night. Two bluespine unicornfish were also nocturnally active. There was little direct evidence of dispersal from the reserve but differences in bluespine unicornfish abundance and size among reef habitat zones were consistent with ontogenetic habitat shifts. The influence of habitat topography on bluespine unicornfish movements suggests that gross habitat characteristics could be used to predict reef fish movements. This could provide a simple method for setting marine reserve boundaries at sites for which empirical fish movement data are unavailable.     

Context-dependent movement behavior of woodland salamanders (Plethodon) in two habitat types

Animal movement is critical to the maintenance of functional connectivity at the landscape scale and can play a key role in population persistence and metapopulation dynamics. The permeability of habitat to animal movement may vary as a result of either differential mortality, physical resistance, or simply the behavioral responses of organisms to perceived habitat quality. Understanding how and when animal movement behavior varies among habitat types is critical for identifying barriers to dispersal and predicting species distributions in relation to landscape features. We conducted an experimental translocation study and compared the movement success and behavioral strategies of plethodontid salamanders in both forest and open-canopy habitat. We found that individuals in closed-canopy forest oriented more strongly towards their home ranges and moved significantly farther on their release night. In spite of the clear differences in movement paths, the ultimate movement success of homing salamanders did not appear to vary with habitat type. Our study contributes to a growing body of literature suggesting the importance of recognizing the context dependence of animal movement behavior. Because the movement rates of displaced salamanders were significantly reduced in open-canopy, dispersal rates of plethodontid salamanders in open-canopy habitat are likely lower than in control forest. Further mechanistic studies focusing on habitat-specific movement behavior and survival costs will be valuable for effectively identifying and mitigating barriers to animal movement.     

Habitat selection by two species of small mammals in the Atlantic Forest,Brazil: Comparing results from live trapping and spool-and-line tracking

Habitat selection by small mammals is usually evaluated using data from live trapping, which provides little information about the movements of individuals. Few studies used movement data or compared the results of different sampling methods to study habitat selection by these animals. We evaluated habitat selection by the rodent Nectomys squamipes and the marsupial Micoureus paraguayanus in the Atlantic Forest of Brazil using the spool-and-line technique. We also determined if percentage of captures reflected the amount of movements in each habitat. Habitat selection was determined comparing use and availability of five habitat types at two spatial scales (movement paths and movement areas) using compositional analysis, which allowed ranking of habitats according to their relative use by animals. The use of available habitat types was non-random for both species at both spatial scales. The two species had contrasting habitat affinities directly related to their particular habits, with N. squamipes using predominantly the stream habitat, and M. paraguayanus using mainly the restinga forest habitat. Patterns of habitat selection were similar at both spatial scales probably due to the small size of movement areas, which may not represent habitat use at a broader scale. For both species, live trapping and movement data provided the same ranking in habitat use, demonstrating that simple capture indices may be used to study habitat selection by these species across different habitat types.     

Group dynamics and landscape features constrain the exploration of herds in fusion-fission societies: the case of European roe deer

Despite the large number of movement studies, the constraints that grouping imposes on movement decisions remain essentially unexplored, even for highly social species. Such constraints could be key, however, to understanding the dynamics and spatial organisation of species living in group fusion-fission systems. We investigated the winter movements (speed and diffusion coefficient) of groups of free-ranging roe deer (Capreolus capreolus), in an agricultural landscape characterised by a mosaic of food and foodless patches. Most groups were short-lived units that merged and split up frequently during the course of a day. Deer groups decreased their speed and diffusion rate in areas where food patches were abundant, as well as when travelling close to main roads and crest lines and far from forests. While accounting for these behavioural adjustments to habitat features, our study revealed some constraints imposed by group foraging: large groups reached the limit of their diffusion rate faster than small groups. The ability of individuals to move rapidly to new foraging locations following patch depression thus decreases with group size. Our results highlight the importance of considering both habitat heterogeneity and group dynamics when predicting the movements of individuals in group fusion-fission societies. Further, we provide empirical evidence that group cohesion can restrain movement and, therefore, the speed at which group members can explore their environment. When maintaining cohesion reduces foraging gains because of movement constraints, leaving the group may become a fitness-rewarding decision, especially when individuals can join other groups located nearby, which would tend to maintain highly dynamical group fusion-fission systems. Our findings also provide the basis for new hypotheses explaining a broad range of ecological patterns, such as the broader diet and longer residency time reported for larger herbivore groups.     

Spatial ecology and habitat use of two-spined blackfish Gadopsis bispinosus in an upland reservoir

The scale and patterns of movement and habitat use are primary considerations in the conservation and management of threatened species. Movement, activity and habitat use of the threatened two-spined blackfish Gadopsis bispinosus were assessed in a small upland reservoir in south-eastern Australia using manual and remote radio-telemetry. Movements and activity of two-spined blackfish (n?=?19) were studied over a 28-day period and exhibited proportionately large directional crepuscular movement and activity with heightened activity continuing throughout the night (although movement was subdued). Two daily movement strategies were observed: movements from diurnal home-shelter habitats (predominantly rock) to macrophytes at night (14 individuals), and occupation of macrophytes during the entire diel period and restricted movement (five individuals). Daily movement strategies were fixed (not plastic) among all individuals, with one exception, for the duration of the study period. Rock, fallen timber and macrophytes were the most commonly used daytime shelter habitat (in order of preference). Although some information exists on movements and habitat use of this species and the congeneric river blackfish G. Marmoratus in lotic environments, we present the first study of movements and habitat use for either species in lentic environments. Given the occupation of lentic environments by this threatened species, the data presented in this study provide insight into the habitat requirements for this species, and offer opportunities for habitat enhancement in existing reservoirs within the species’ geographic range.