Road crossing in bank voles and yellow-necked mice

Roads and highways represent one of the most important anthropogenic impacts on natural areas and contribute to habitat fragmentation, because they are linear features that can inhibit animal movement, thereby causing barrier effects subdividing the populations adjacent to the roads. The paper examines to what extent a narrow (2-lane) and a wide (4-lane) highways represent barriers for two small mammal species: bank volesClethrionomys glareolus Schreber, 1780 and yellow-necked miceApodemus flavicollis Melchior, 1834, and whether displaced rodents are able to return across roads of different widths. The study was performed at four sites in the Czech Republic. The capture-mark-recapture method was used to determine crossing rates. At two sites, the animals captured close to the road were transferred to the other side and released, to compare return movements across the roads with the movements made by the non-transferred animals. We found that the narrow highway did not prevent movement of neither of the species, although voles crossed only after they had been transferred. Wide highways, on the other hand, completely prevened crossing of both species. While the narrow highways acted at individuals level, the wide highways affected the population subvision.     

The role of habitat barriers in woodlot recolonization by small mammals

The immigration and colonization of small mammals into a small depopulated wooded area surrounded by meadows was monitored by live-trapping and then compared with the same processes in an open forest following removal trapping. Rate of recolonization was lower in the depopulated woods than in the continuous forest. Bank voles, Clethrionomys glareolus , predominated the small mammal communities before removal; however, after removal the percentage of bank voles decreased and species diversity increased. When comparing the bank voles from the wooded area with the continuous forest, we found a higher percentage of sexually active animals, the lack of reproducing females and slightly lower body weights. Based on these characteristics the bank voles that recolonized the small woods were regarded as dispersers. The meadow belt surrounding the depopulated wood acted as a "filter" which stopped some bank voles and allowed others to pass through, and also "selected" different species crossing it according to their mobility. Some effects of such habitat barriers on the functioning of metapopulations are discussed.     

Barriers to dispersal of rain forest butterflies in tropical agricultural landscapes

Fragmentation of natural habitats can be detrimental for species if individuals fail to cross habitat boundaries to reach new locations, thereby reducing functional connectivity. Connectivity is crucial for species shifting their ranges under climate change, making it important to understand factors that might prevent movement through human‐modified landscapes. In tropical regions, rain forests are being fragmented by agricultural expansion, potentially isolating populations of highly diverse forest‐dependent species. The likelihood of crossing habitat boundaries is an important determinant of species dispersal through fragmented landscapes, and so we examined movement across rain forest‐oil palm plantation boundaries on Borneo by using relatively mobile nymphalid butterflies as our model study taxon. We marked 1666 individuals from 65 species, and 19 percent (100/527) of recaptured individuals crossed the boundary. Boundary crossing was relatively frequent in some species, and net movement of individuals was from forest into plantation. However, boundary crossing from forest into plantation was detected in less than 50 percent (12/28) of recaptured species and was dominated by small‐sized butterfly species whose larval host plants occurred within plantations. Thus, while oil palm plantations may be relatively permeable to some species, they may act as barriers to the movement of forest‐dependent species (i.e., species that require rain forest habitat to breed), highlighting the importance of maintaining forest connectivity for conserving rain forest species.     

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.     

Gap‐crossing decisions of forest birds in a fragmented landscape

Habitat loss and fragmentation are recognized as primary drivers of biodiversity loss worldwide. To understand the functional effects of habitat fragmentation on bird populations, data on movement across gaps in habitat cover are necessary, although rarely available. In this study, we used call playback to simulate a conspecific territorial intruder to entice birds to move through the landscape in a predictable and directional manner. We then quantified the probability of movement in continuous forest and across cleared gaps for two forest‐dependent species, the grey shrike‐thrush (Colluricincla harmonica) and the white‐throated treecreeper (Cormobates leucophaeus). Fifty‐four playback trials were conducted for each species across distances ranging from 25 to 480 m in continuous forest and 15–260 m across gaps in a forest‐agricultural landscape in southern Victoria, Australia. The probability of movement was significantly reduced by gaps in forest cover for both species. Shrike‐thrushes were six times more likely to move 170 m in continuous forest than to cross 170‐m gaps. The mean probability that treecreepers would cross any gap at all was less than 0.5, and they were three times less likely to move 50 m across a gap than through continuous forest. Both species displayed non‐linear responses to increasing gap distance: we identified a gap‐tolerance threshold of 85 m for the shrike‐thrush and 65 m for the treecreeper beyond which individuals were most unlikely to cross. The presence of scattered paddock trees increased functional connectivity for the shrike‐thrush, with individuals crossing up to 260 m when scattered trees were present. We conclude that gaps in habitat cover are barriers to movement, and that characteristics of the intervening matrix influence landscape permeability.     

Nowhere to run: semi-permeable barriers affect pronghorn space use

Animal movement can mediate the ecological consequences of fragmentation; however, barriers such as fences, roads, and railways are becoming a pervasive threat to wildlife. Pronghorn (Antilocapra americana) habitat in western North America has been fragmented by roads, railways, and fences. Although pronghorn are sensitive to barriers, neither the relative permeability of different barriers to crossing nor their influence on space use have been quantified. We used a large global positioning system (GPS)-collar dataset of pronghorn (n = 1,010 animal-years) in Wyoming, USA, to first quantify the likelihood that pronghorn cross each of 5 different anthropogenic barriers, including fences, county roads, railroads, state highways, and interstate highways (i.e., interstates). Next, we assessed how each barrier influenced pronghorn space use during the winter as indexed by the area occupied, and daily displacement relative to the density of barriers on an individual's winter range. The semi-permeability of the 5 barriers varied substantially, with the interstate being the most severe barrier to pronghorn movement. Pronghorn were >300 times less likely to cross interstates compared to state highways. Although pronghorn space use was rarely influenced by barriers within individual core winter ranges, pronghorn space use was constrained by barriers on the buffered periphery of individual winter ranges. Despite their different permeability to movement, the density of fences and combined interstates and railroads had similarly negative effects on pronghorn space use. Our results illustrate that the degree to which pronghorn avoid crossing barriers may scale up to affect access to habitat. Additionally, our results indicate that the effects of barriers on habitat access are not proportional to their permeability. Our results add to a growing consensus that effective management of mobile species depends on understanding how different kinds of semi-permeable barriers influence access and use of habitats.     

Effects of Barriers and Thermal Refugia on Local Movement of the Threatened Leopard Darter, Percina pantherina

Local, short-term dispersal by the U.S. federally-threatened leopard darter, Percina pantherina, was examined in the field and in the laboratory to assess the possible effects of natural versus man-made barriers on movement. Mark-resight studies were conducted in two summers at sites in the Glover River (southeastern Oklahoma, U.S.A.). At one site, patches of 'preferred' habitat were separated by a natural riffle; at the other site, by a low-water road crossing with culverts. At the Natural Riffle site, darters moved downstream across the riffle, but also moved upstream into deeper water when water temperatures exceeded 29°C in the `preferred' habitat. Use of deeper, cooler waters by this species in late summer suggests that thermal refugia may be important habitats for the long-term management of leopard darters. At the Road Crossing site, all documented movement was in a downstream direction, and at least two darters traversed culverts in the low-water bridge. Laboratory studies of movement across several types of culverts suggested that culverts significantly decrease the probability of movement among habitat patches.     

The effect of landscape structure on dispersal distances of the Eurasian red squirrel

Landscape structure can affect dispersal and gene flow in a species. In urban areas, buildings, roads, and small habitat patches make the landscape highly fragmented and can inhibit movement and affect dispersal behavior. Similarly, in rural forested areas, large open areas, such as fields, may act as barriers to movement. We studied how landscape structure affects natal dispersal distances of Eurasian red squirrels (Sciurus vulgaris) in an urban area and a rural area in Finland, by monitoring juvenile red squirrels with radio telemetry. We observed extremely long dispersal distances—up to 16 km—in the rural study area, but shorter distances—on average only half a kilometer—in the urban study area. The landscape structure affected the eventual dispersal paths; in the rural landscape, dispersers favored spruce dominated areas and avoided fields along their dispersal route, although they occasionally even crossed wide fields. In the urban landscape, squirrels preferred areas with deciduous or coniferous trees. The movement steps made by dispersers were longer in the more hostile landscape compared to forested areas. Despite these effects on movement path, the landscape structure only had a minor effect on straight line dispersal distances moved from the natal nest. In other words, individuals moved longer distances and were likely to circumvent barriers in their path, but this did not affect how far they settled from their natal home. This result indicates that, although landscape structure has obvious effects on movement, it still may have only a small effect on other aspects of the population, for example, gene flow.     

The effect of local anthropogenic habitat heterogeneity on assemblages of carabids (Coleoptera,Caraboidea) endemic to the Alps

We assessed the response of a guild of endemic medium and large-bodied carabid species to the local (fine-grained) habitat heterogeneity induced by man. The study was conducted by means of trapping (without killing and preservative agents) and radio-tracking methods (to collect data on the spatial ecology of the endangered species Carabus olympiae Sella). Habitat differentiation induced by man (a mosaic of beech forests, alpen rose shrubberies and pastures) affected species richness and abundance of the guild, which were significantly higher in forests and/or shrubberies than in pastures. Most species preferred forests and shrubberies, with the only exception of Carabus concolor Fabricius, which preferred open areas. Species distribution was also affected by stone density and cattle grazing intensity. In particular, petrophilous species peaked on the roadside because of the high stone density there (stones amassed during road construction), while species abundances significantly lowered in overgrazed pastures. The compactness of the paths made by C. olympiae individuals (evaluated through a tortuosity index) was significantly higher in beech forests than in alpen rose shrubberies, indicating movements in the latter habitat type were bound by the spatial distribution of shrubs, which imposes a limit on path tortuosity. Our findings suggest that local ground beetle species diversity strongly depends on small-scale anthropogenic variables (namely habitat type, stone density and grazing intensity), and that habitat modifications (namely from forest to shrubbery) may significantly affect species movement patterns. General and local conservation management suggestions are given in conformity with these results.     

Roads as barriers to animal movement in fragmented landscapes

Roads can act as barriers to animal movement through mortality during crossing attempts or behavioral avoidance. This barrier effect has negative demographic and genetic consequences that can ultimately result in local or regional extinction. Here we use radio-telemetry data on three terrestrial vertebrates (eastern massasauga Sistrurus catenatus , eastern box turtle Terrapene carolina and ornate box turtle Terrapene ornata ) to test whether roads acted as barriers to movement. Specifically, we test whether individuals avoided crossing roads by comparing the number of observed crossings with the number of road crossings predicted by randomizations of individual movement paths. All species crossed roads significantly less often than predicted by chance, indicating strong road avoidance. Results of this study showing behavioral avoidance and previous studies on road mortality indicate that roads are strong barriers to these species. High mortality during crossing attempts would select for road avoidance, reducing the number of individuals killed on roads over time but leading to genetically partitioned subpopulations due to a lack of gene flow. In species that are long-lived and late-maturing, negative genetic effects might not be observable over short time-scales, thus placing populations at high risk of extinction because of a failure to detect an incrementally worsening problem. Formulating successful management strategies for many species in decline will require integrating data on road mortality, animal behavior and population genetics in order to understand more clearly the barrier effect of roads.     

Quantifying the activity levels and behavioural responses of butterfly species to habitat boundaries

1. The ability of species' to undergo climate‐driven range shifts across fragmented landscapes depends on their dispersal ability as well as the structure of the landscape. For species' range shifts to occur, individuals must first leave suitable habitat to seek new habitat; this is likely to depend on the rate of movement of individuals within habitat and the likelihood that a boundary is crossed, once it is encountered. For three species of butterfly with contrasting histories of recent range expansion, we examined the propensity of individuals to move within a habitat and their responses to habitat boundaries. 2. We quantified the extent to which Plebejus argus (Linnaeus) (a declining habitat specialist), Aricia agestis (Schiffermuller) (an expanding generalist) and Polymmatus icarus (Rottemburg) (a geographically ubiquitous generalist) crossed habitat boundaries into unsuitable habitat and moved within suitable habitat. The observed movement was then related to individual and environmental conditions. 3. Species differed in their activity levels in accordance within their recent distribution patterns (P. icarus > A. agestis > P. argus). Our results for P. argus suggest that movement may be motivated by nectar‐seeking, and that males generally move more than females. All three species tended to avoid crossing habitat boundaries; however the proportion of individuals crossing habitat boundaries did not differ significantly among species. 4. We conclude that levels of activity within a habitat, which will affect the frequency with which individuals encounter habitat boundaries, rather than behavioural responses to the boundaries, may be important drivers of distribution change.     

Identification of the barrier to gene flow between phylogeographic lineages of the common hamster <Emphasis Type="Italic">Cricetus cricetus</Emphasis>

In anthropogenically disturbed habitats, natural barriers still exist and have to be recognized, as they are important for conservation measures. Areas of phylogeographic breaks within a species are often stabilized in inhospitable regions which act as natural barriers. An area of contact between phylogeographic lineages of the common hamster (Cricetus cricetus) was found in the Małopolska Upland in Poland. A total of 142 common hamsters were captured between 2005 and 2009. All hamsters were genotyped at 17 microsatellite loci and partial sequences of the mitochondrial (mtDNA) control region were obtained. No mixed populations with mtDNA haplotypes of both lineages were found. The distance between marginal populations was about 20 km; no hamsters were found in the area between. A principal components analysis (PCA) was performed on microsatellite data and the greatest change in PC1 scores was found between marginal samples. To define the habitat components responsible for the phylogeographic break, we compared the habitat composition of sites occupied by hamsters with those from which hamsters were absent. We found that hamsters avoided forested areas and sandy soils. The area of the potential barrier was characterized by a high proportion of woodland and unfavorable soils in comparison with neighboring areas inhabited by hamsters. They cannot settle in this area due to their high winter mortality in shallow burrows and high predation in the fields adjacent to forests.     

Combined effects of seed and soil quality drive seedling performance of a late‐successional canopy tree in a tropical forest

Habitat loss and fragmentation affect the structure and functioning of forested ecosystems worldwide, yet we lack an understanding of how species respond to environmental changes. Here, we examined reproductive success and seedling performance of Poulsenia armata (Moraceae) in continuous and fragmented forests of Los Tuxtlas, southern Mexico. We further investigated how maternal habitat and soil conditions manifested in the seedling stage. We determined seed quality and seedling performance by combining isotopic analyses in seed quality with field observations of P. armata fruit production and a common‐‐garden experiment. Soil conditions in forest fragments negatively impacted P. armata reproductive success. Trees of P. armata in forest fragments were smaller in size and produced fewer fruits and smaller seeds with lower quality compared with trees from the continuous forest. The combined effects of maternal habitat and soil conditions determined seedling survival and growth of this tropical tree. Notably, seedlings had restricted plasticity for biomass allocation to roots, limiting the capacity of fragmented populations to compensate for the initial low N content in seeds. Trees in forest fragments at Los Tuxtlas produced offspring competitively inferior and potentially less resilient than counterparts in continuous forest, jeopardizing future persistence of this late‐successional tree species.     

Forests do not limit bumble bee foraging movements in a montane meadow complex

1. Understanding the roles of habitat fragmentation and resource availability in shaping animal movement are integral for promoting species persistence and conservation. For insects such as bumble bees, their movement patterns affect the survival and reproductive potential of their colonies, as well as the pollen flow of plant species. However, the understanding of their mobility or the impact of putative barriers in natural environments is limited due to the technical difficulties of studying wild populations. 2. Genetic mark–recapture was used to estimate the foraging distance, resource use, and site connectivity of two bumble bee species in a montane meadow complex composed of open meadows within a matrix of forest. 3. There was no evidence that forests or changes in landcover function as barriers to the fine-scale movement for either species. Substantially greater colony-specific foraging distances were found for Bombus vosnesenskii (maximum: 1867 m) compared to Bombus bifarius (maximum: 362 m). Despite this difference in absolute range, both species were detected across putative forest barriers at frequencies expected by uninhibited movement. Siblings separated by greater distances were more likely to be foraging on different floral species, potentially suggesting a resource-based motivation for movement. 4. These results suggest that bumble bee foraging patterns are influenced by species-specific differences in movement capacity, with little influence of matrix composition between resource patches. They also support the perspective that habitat conservation for bumble bees should prioritise providing abundant and diverse patches of resources within species-specific movement radii with less emphasis on matrix composition.     

Using animal movement paths to measure response to spatial scale

Animals live in an environment that is patchy and hierarchical. I present a method of detecting the scales at which animals perceive their world. The hierarchical nature of habitat causes movement path structure to vary with spatial scale, and the patchy nature of habitat causes movement path structure to vary throughout space. These responses can be measured by a combination of path tortuousity (measured with fractal dimension) versus spatial scale, the variation in tortuousity of small path segments along the movement path, and the correlation between tortuousities of adjacent path segments. These statistics were tested using simulated animal movements. When movement paths contained no spatial heterogeneity, then fractal D and variance continuously increased with scale, and correlation was zero at all scales. When movement paths contained spatial heterogeneity, then fractal D sometimes showed a discontinuity at transitions between domains of scale, variation showed peaks at transitions, and correlations showed a statistically significant positive value at scales smaller than patch size, decreasing to below zero at scales greater than patch size. I illustrated these techniques with movement paths from deer mice and red-backed voles. These new analyses should help understand how animals perceive and react to their landscape structure at various spatial scales, and to answer questions about how habitat structure affects animal movement patterns.     

Fine-scale movement behaviors of calopterygid damselflies are influenced by landscape structure: an experimental manipulation

We explore the effect of differences in landscape structure, arising from habitat loss, on the fine-scale movement behaviors of two congeneric damselflies – Calopteryx aequabilis and C. maculata . Both species require streams for breeding and naiad development and both often use forest for foraging. We compare movement behaviors across three types of landscape: forested landscapes, where stream and forest habitat are adjacent; partially forested landscapes, where streams and forest habitat are disjunct, and non-forested landscapes, where little to no forest habitat is available. We employ a reciprocal transplant experiment to determine the extent to which movement along and away from streams is influenced by landscape structure and historical behavior or morphological adaptations. For both species, we show that both the propensity to move away from streams and rates of net displacement differ among landscape types. Both species move away from streams on landscapes with high or moderate levels of forest cover but neither moves away from streams on landscapes with little or no forest. Furthermore, C. maculata native to predominantly forested landscapes are more likely to move away from streams, regardless of the landscape structure they encounter, than are individuals native to moderately forested or non-forested landscapes. There was no effect of natal landscape on C. aequabilis . Comparisons with microlandscape studies suggest that there may be some general similarities among the different systems but these are clouded by uncertainty regarding the similarity of the underlying processes responsible for observed behavioral responses to landscape structure. Despite this uncertainty, animal movement behaviors are contingent upon the structure of the broader landscape, regardless of the absolute scale of the landscape.     

Gene flow among native bush rat,Rattus fuscipes (Rodentia: Muridae), populations in the fragmented subtropical forests of south‐east Queensland

Abstract Many natural populations in areas of continuous habitat exhibit some form of local genetic structure. Anthropogenic habitat fragmentation can also strongly influence the dynamics of gene flow between populations. We used eight microsatellite markers to investigate the population genetic structure of an abundant forest species, the Australian bush rat (Rattus fuscipes), in the subtropical forests of south‐east Queensland. Five sites were sampled, allowing pairwise comparisons within continuous habitat and across clearings. Weak, but significant population differentiation and a significant pattern of isolation by distance was detected over the small scale (<10 km) of this study. Fine‐scale analysis at a single site (<1 km) showed a significant correlation between individual female genetic distance and geographical distance, but no similar pattern among male individuals. There was no evidence of increased population differentiation across clearings relative to comparisons within continuous forest. This was attributed to dispersal within corridors of remnant and revegetated habitat between the forested areas. We concluded that an inherently restricted dispersal ability, female philopatry and natural habitat heterogeneity play an important part in the development of genetic structure among populations of R. fuscipes. It is important to understand the relationship between landscape features and the pattern of gene flow among continuous populations, as this allows us to predict the impact of fragmentation on natural populations.     

Effects of Connectivity and Regional Dynamics on Restoration of Small Mammal Communities in Midwestern Grasslands

Grasslands are among the most imperiled North American ecosystems. State Acres for Wildlife Enhancement (SAFE) is a national conservation program that converts agricultural fields into grasslands mainly to improve habitat for high priority wildlife species. To provide a broader assessment of the contribution of the SAFE program to biodiversity in the Midwest region of North America, we evaluated local and landscape constraints to restoration of small mammal communities. We livetrapped small mammals during three summers (2009–2011) on plots that were recently seeded, seeded 1–4 years prior to sampling, or established references (>10 years old). Restoration trajectories for small mammal communities included a shift over time from dominance by the habitat generalist Peromyscus maniculatus (deer mouse) to communities dominated by grassland Microtus species (prairie voles and meadow voles). Vole abundance during the first year following restoration depended on spatial connectivity provided by linear habitats (roadside ditches and grass waterways) within 300 m of the restored grassland. Patch size and seeding type (cool‐season versus warm‐season grasses) were not predictors of early restoration success. In 2011, voles experienced a severe regional decline consistent with multi‐year population cycles. During the crash, most remaining voles occurred on restored SAFE grasslands, but not on established grasslands. This surprising outcome suggests young restoration plots could function as refuges for voles during population declines in agricultural landscapes.     

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.     

Landscape structure influences the use of social information in an insectivorous bat

In anthropogenic landscapes, aerial insectivores are often confronted with variable habitat complexity, which may influence the distribution of prey. Yet, high mobility may allow aerial insectivores to adjust their foraging strategy to different prey distributions. We investigated whether aerial-hunting common noctules Nyctalus noctula adjust their foraging strategy to landscapes with different habitat complexity and assumingly different prey distribution. We hypothesized that the movement behaviour of hunting common noctules and changes of movement behaviour in reaction towards conspecifics would depend on whether they hunt in a structurally poor cropland dominated landscape or a structurally rich forest dominated landscape. We tracked flight paths of common noctules in northeastern Germany using GPS loggers equipped with an ultrasonic microphone that recorded foraging events and presence of conspecifics. Above cropland, common noctules hunted mainly during bouts of highly tortuous and area restricted movements (ARM). Bats switched from straight flight to ARM after encountering conspecifics. In the forested landscape, common noctules hunted both during ARM and during straight flights. The onset of ARM did not correlate with the presence of conspecifics. Common noctules showed a lower feeding rate and encountered more conspecifics above the forested than above the cropland dominated landscape. We conjecture that prey distribution above cropland was patchy and unpredictable, thus making eavesdropping on hunting conspecifics crucial for bats during search for prey patches. In contrast, small scale structural diversity of the forested landscape possibly led to a more homogeneous prey distribution at the landscape scale, thus enabling bats to find sufficient food independent of conspecific presence. This suggests that predators depending on ephemeral prey can increase their foraging success in structurally poor landscapes by using social information provided by conspecifics. Hence, a minimum population density might be obligatory to enable successful foraging in simplified landscapes.