Representing genetic variation as continuous surfaces: an approach for identifying spatial dependency in landscape genetic studies

Landscape genetics, an emerging field integrating landscape ecology and population genetics, has great potential to influence our understanding of habitat connectivity and distribution of organisms. Whereas typical population genetics studies summarize gene flow as pairwise measures between sampling localities, landscape characteristics that influence population genetic connectivity are often continuously distributed in space. Thus, there are currently gaps in both the ability to analyze genotypic data in a continuous spatial context and our knowledge of expected of landscape genetic structure under varying conditions. We present a framework for generating continuous “genetic surfaces”, evaluate their statistical properties, and quantify statistical behavior of landscape genetic structure in a simple landscape. We simulated microsatellite genotypes under varying parameters (time since vicariance, migration, effective population size) and used ancestry (q) values from STRUCTURE to interpolate a genetic surface. Using a spatially adjusted Pearson's correlation coefficient to test the significance of landscape variable(s) on genetic structure we were able to detect landscape genetic structure on a contemporary time scale (≥5 generations post vicariance, migration probability ≤0.10) even when population differentiation was minimal (F_ST≥0.00015). We show that genetic variation can be significantly correlated with geographic distance even when genetic structure is due to landscape variable(s), demonstrating the importance of testing landscape influence on genetic structure. Finally, we apply genetic surfacing to analyze an empirical dataset of black bears from northern Idaho USA. We find black bear genetic variation is a function of distance (autocorrelation) and habitat patch (spatial dependency), consistent with previous results indicating genetic variation was influenced by landscape by resistance. These results suggest genetic surfaces can be used to test competing hypotheses of the influence of landscape characteristics on genetic structure without delineation of categorical groups.     

Landscape configuration drives persistent spatial patterns of occupant distributions

Variation in the density of organisms among habitat patches is often attributed to variation in inherent patch properties. For example, higher quality patches might have higher densities because they attract more colonists or confer better post-colonization survival. However, variation in occupant density can also be driven by landscape configuration if neighboring patches draw potential colonists away from the focal habitat (a phenomenon we call propagule redirection). Here, we develop and analyze a stochastic model to quantify the role of landscape configuration and propagule redirection on occupant density patterns. We model a system with a dispersive larval stage and a sedentary adult stage. The model includes sensing and decision-making in the colonization stage and density-dependent mortality (a proxy for patch quality) in the post-colonization stage. We demonstrate that spatial variation in colonization is retained when the supply of colonists is not too high, post-colonization density-dependent survival is not too strong, and colonization events are not too frequent. Using a reef fish system, we show that the spatial variation produced by propagule redirection is comparable to spatial variation expected when patch quality varies. Thus, variation in density arising from the spatial patterning of otherwise identical habitat can play an important role in shaping long-term spatial patterns of organisms occupying patchy habitats. Propagule redirection is a potentially powerful mechanism by which landscape configuration can drive variation in occupant densities, and may therefore offer new insights into how populations may shift as landscapes change in response to natural and anthropogenic forces.     

Effects of environment,habitat configuration and forest continuity on the distribution of forest plant species

In present day European landscapes many forest plant species are restricted to isolated remnants of a formerly more or less continuous forest cover. The two major objectives of this study were (1) to determine the relative importance of habitat quality (mainly in terms of soil parameters), habitat configuration (patch size and isolation) and habitat continuity for the distribution of herbaceous forest plant species in a highly fragmented landscape and (2) to examine if groups of species with different habitat requirements are affected differently. Deciduous forest patches in northwestern Germany were surveyed for the presence of a large set of forest species. For each patch, habitat quality, configuration and continuity were determined. Data were analysed by Redundancy Analysis with variation partitioning for effects on total species composition and multivariate logistic regression for effects on individual species, for two different data sets (base‐rich and base‐poor forest patches). Overall, we found strong effects of habitat quality (particularly of soil pH, water content and topographic heterogeneity in the base‐rich forest patches; and of calcium content and disturbance in the base‐poor patches), but only relatively weak effects of habitat configuration and habitat continuity. However, a number of species were positively affected by patch area and negatively affected by patch isolation. Furthermore, the relative importance of habitat configuration tended to be higher for species predominantly growing in closed forests compared to species occurring both in the forest and in the open landscape.     

Forest bird diversity in Mediterranean areas affected by wildfires: a multi-scale approach

Fire is a key mechanism creating and maintaining habitat heterogeneity in Mediterranean landscapes by turning continuous woody landscapes into mosaics of forests and shrublands. Due to the long historical role of fires in the Mediterranean, we hypothesised a moderate negative effect of this type of perturbation on forest bird distribution at a landscape level. We conducted point bird censuses in Aleppo pine forest patches surrounded by burnt shrublands and studied the relationships between three ecological groups of bird species (forest canopy species, forest understorey species, and ubiquitous species) and the features of local habitat, whole patch and surrounding landscape. We used a multi-scale approach to assess the effects of landscape variables at increasing spatial scales on point bird richness. Regarding local habitat components, canopy species were positively associated with tall pines while understorey species with the cover of shrubs and plants from holm-oak forests. Forest birds were positively related to patch size and irregular forest shapes, that is, with high perimeter/size ratios. Thus, these species did not seem to perceive edges as low quality but rather favourable microhabitats. We did not detect any negative effect of isolation or cover of woodlands in the landscape on the presence of forest species after local habitat factors had been accounted for. Finally, only local habitat factors entered the model for ubiquitous species. We suggest that mosaic-like landscapes shaped by fires in the Mediterranean basin are not strongly associated with negative effects fragmentation on forest birds other than those related with habitat loss.     

Propagule Limitation,Disparate Habitat Quality,and Variation in Phenotypic Selection at a Local Species Range Boundary

Adaptation to novel conditions beyond current range boundaries requires the presence of suitable sites within dispersal range, but may be impeded when emigrants encounter poor habitat and sharply different selection pressures. We investigated fine-scale spatial heterogeneity in ecological dynamics and selection at a local population boundary of the annual plant Gilia tricolor. In two years, we planted G. tricolor seeds in core habitat, margin habitat at the edge of the local range, and exterior habitat in order to measure spatial and temporal variation in habitat quality, opportunity for selection, and selection on phenotypic traits. We found a striking decline in average habitat quality with distance from the population core, yet some migrant seeds were successful in suitable, unoccupied microsites at and beyond the range boundary. Total and direct selection on four out of five measured phenotypic traits varied across habitat zones, as well as between years. Moreover, the margin habitat often exerted unique selection pressures that were not intermediate between core and exterior habitats. This study reveals that a combination of ecological and evolutionary forces, including propagule limitation, variation in habitat quality and spatial heterogeneity in phenotypic selection may reduce opportunities for adaptive range expansion, even across a very local population boundary.     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

丹顶鹤多尺度生境选择机制——以黄河三角洲自然保护区为例

生境在鸟类生活史中发挥着重要的作用,关系到鸟类的生存和繁衍。由于鸟类对环境变化的响应发生在等级序列空间尺度上,基于多尺度的研究更能深入刻画鸟类-环境之间关系。以丹顶鹤(Grus japonensis)为研究对象,以其迁徙和越冬的重要地区-黄河三角洲自然保护区为研究区域,应用等级方差分解法和等级划分法,分析丹顶鹤与微生境、斑块、景观尺度因子之间的关系,探求丹顶鹤生境选择的主要影响因素和尺度。等级方差分解结果表明,在第1等级水平,景观尺度因子与微生境、斑块尺度因子之间的联合效应大于独立效应,景观尺度因子的独立效应大于微生境和斑块尺度因子;在第2等级水平,景观尺度上的景观组成因子重要性大于景观结构因子,微生境尺度上的植被和水分因子为重要影响因素。等级划分结果表明,景观尺度上,翅碱蓬滩涂、水体面积大小是主要影响因素;微生境尺度上,植被盖度和水深为主要限制因子;在斑块尺度上,斑块类型对丹顶鹤生境选择最为重要。研究认为,在黄河三角洲自然保护区,景观尺度是影响丹顶鹤生境选择的主要尺度,景观尺度因子通过与微生境和斑块尺度因子的独立和联合作用制约着丹顶鹤在保护区的生境选择和空间分布格局。建议加强对翅碱蓬滩涂、芦苇沼泽、水体等湿地生境的保护和管理,规范和控制保护区内人类活动强度。     

Effects of landscape and patch-level attributes on regional population persistence

Species responses are influenced by processes operating at multiple scales, yet many conservation studies and management actions are focused on a single scale. Although landscape-level habitat conditions (i.e., habitat amount, fragmentation and landscape quality) are likely to drive the regional persistence of spatially structured populations, patch-level factors (i.e., patch size, isolation, and quality) may also be important. To determine the spatial scales at which habitat factors influence the regional persistence of endangered Ord's kangaroo rats (Dipodomys ordii) in Alberta, Canada, we simulated population dynamics under a range of habitat conditions. Using a spatially-explicit population model, we removed groups of habitat patches based on their characteristics and measured the resulting time to extinction. We used proportional hazards models to rank the influence of landscape and interacting patch-level variables. Landscape quality was the most influential variable followed by patch quality, with both outweighing landscape- and patch-level measures of habitat quantity and fragmentation/proximity. Although habitat conservation and restoration priorities for this population should be in maximizing the overall quality of the landscape, population persistence depends on how this goal is achieved. Patch quality exerted a significant influence on regional persistence, with the removal of low quality road margin patches (sinks) reducing the risk of regional extinction. Strategies for maximizing overall landscape quality that omit patch-level considerations may produce suboptimal or detrimental results for regional population persistence, particularly where complex local population dynamics (e.g., source-sink dynamics) exist. This study contributes to a growing body literature that suggests that the prediction of species responses and future conservation actions may best be assessed with a multi-scale approach that considers habitat quality and that the success of conservation actions may depend on assessing the influences of habitat factors at multiple scales.     

Patch quality and context, but not patch number, drive multi-scale colonization dynamics in experimental aquatic landscapes

Colonization and extinction are primary drivers of local population dynamics, community structure, and spatial patterns of biological diversity. Existing paradigms of island biogeography, metapopulation biology, and metacommunity ecology, as well as habitat management and conservation biology based on those paradigms, emphasize patch size, number, and isolation as primary characteristics influencing colonization and extinction. Habitat selection theory suggests that patch quality could rival size, number, and isolation in determining rates of colonization and resulting community structure. We used naturally colonized experimental landscapes to address four issues: (a) how do colonizing aquatic beetles respond to variation in patch number, (b) how do they respond to variation in patch quality, (c) does patch context affect colonization dynamics, and (d) at what spatial scales do beetles respond to habitat variation? Increasing patch number had no effect on per patch colonization rates, while patch quality and context were critical in determining colonization rates and resulting patterns of abundance and species richness at multiple spatial scales. We graphically illustrate how variation in immigration rates driven by perceived predation risk (habitat quality) can further modify dynamics of the equilibrium theory of island biogeography beyond predator-driven effects on extinction rates. Our data support the importance of patch quality and context as primary determinants of colonization rate, occupancy, abundance, and resulting patterns of species richness, and reinforce the idea that management of metapopulations for species preservation, and metacommunities for local and regional diversity, should incorporate habitat quality into the predictive equation.     

生境质量对城镇化的时空响应——以长春市为例

生境质量是评价区域生态安全的重要指标,掌握其对城镇化发展的时空响应,有利于新型城镇化的深入推行。本研究基于土地利用数据,在综合利用空间分析及生态模型分析基础上,从网格尺度对长春市景观格局、生境质量及其样带进行时空特征分析,并进一步探讨生境质量在城镇化发展过程中的响应状况。结果表明: 长春市景观斑块密度、边缘密度及Shannon多样性指数低值多分布在西部平原地带,聚合度指数高值区则在市域东部及南部呈现片状格局。2000—2015年,长春市生境质量呈退化趋势且空间异质性显著,大致呈现出“东高西低”分布态势,建设用地扩张及交通基础设施建设对区域生境质量退化起到主导作用;不同样带内生境质量变化存在显著差异性,水域整体变化相对较小,而山脉、城市扩张及交通的变化频度及幅度处于较高水平。坡度及高程等自然因素基本塑造了长春市生境质量整体分布格局,而人口密度、国内生产总值、夜间灯光指数等城镇化要素与生境质量总体呈负相关关系。为缓解城镇化带来的生态压力、促进生境质量的回升,本研究提出防止大黑山脉林区滥砍乱伐、以生态手段修复生境退化地区,提升城镇建成区土地利用效率、促进城镇“精明增长”,设定丘陵地带耕地红线、加强生态基础设施建设等差异化发展策略。     

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.     

Evolutionary responses of dispersal distance to landscape structure and habitat loss

It is generally well understood that some ecological factors select for increased and others for decreased dispersal. However, it has remained difficult to assess how the evolutionary dynamics are influenced by the spatio-temporal structure of the environment. We address this question with an individual-based model that enables habitat structure to be controlled through variables such as patch size, patch turnover rate, and patch quality. Increasing patch size at the expense of patch density can select for more or less dispersal, depending on the initial configuration. In landscapes consisting of high-quality and long-lived habitat patches, patch degradation selects for increased dispersal, yet patch loss may select for reduced dispersal. These trends do not depend on the component of life-history that is affected by habitat quality or the component of life-history through which density-dependence operates. Our results are based on a mathematical method that enables derivation of both the evolutionary stable strategy and the stationary genotype distribution that evolves in a polymorphic population. The two approaches generally lead to similar predictions. However, the evolutionary stable strategy assumes that the ecological and evolutionary time scales can be separated, and we find that violation of this assumption can critically alter the evolutionary outcome.     

Metacommunity diversity depends on connectivity and patch arrangement in heterogeneous habitat networks

Connectivity is critical to the maintenance of biodiversity in fragmented landscapes, but its effects differ depending on the arrangement of linkages within a habitat network. Additionally, heterogeneity in habitat quality within the habitat network can alter patterns of diversity at local and regional scales in the metacommunity. Using a controlled experiment we examined the interactive effects of habitat connectivity, network form (linear vs square), and habitat patch quality on a moss‐inhabiting microarthropod community. We fragmented moss habitat while controlling for habitat loss, and altered habitat patch quality by regulating moisture conditions in landscapes differing in patch arrangement. Habitat patch quality had a significant effect on patterns of species richness, extinction, abundance and biomass. The effects of network form on diversity were strongest in heterogeneous landscapes. Gamma and beta diversity were greatest in continuous and linear landscapes. However, linear habitat networks showed marked patch specific edge effects that were detrimental to diversity under heterogeneous conditions. We provide direct evidence that habitat network structure impacts species community properties through mass effects, that are most evident when heterogeneity in habitat patch quality is present within the network. We conclude that habitat quality at the individual patch level and the distribution of high‐quality habitat within the network are important factors affecting biodiversity in metacommunities.     

The limiting behaviour of a mainland-island metapopulation

Stochastic patch occupancy models (SPOMs) are a class of discrete time Markov chains used to model the presence/absence of a population in a collection of habitat patches. This class of model is popular with ecologists due to its ability to incorporate important factors of the habitat patch network such as connectivity and distance between patches as well as heterogeneity in patch characteristics. We present an asymptotic examination of a simple type of SPOM called the mainland-island model. In this model a single patch called the mainland is connected to a large number of smaller patches called islands and each island is only connected to the mainland. We discuss the limiting behaviour of the SPOM as the number of islands increases and the size of the islands decrease relative to the mainland. We demonstrate that a variety of limiting behaviours is possible depending on the scaling of the island size and on the heterogeneity of habitat quality.     

Importance of patch scale vs landscape scale on selected forest birds

The management and protection of natural areas have primarily occurred in isolation from surrounding land management. The structure of surrounding land cover, however, may be important to the abundance and reproductive success of birds within a habitat patch. We investigated the relative importance of forest patch area, within patch habitat and surrounding landscape forest cover on the abundance of three Neotropical migrant bird species thought to be area-sensitive (ovenbird [ Seiurus aurocapillus ], wood thrush [ Hylocichla mustelina ] and red-eyed vireo [ Vireo olivaceus ]), and on pairing success of the ovenbird. We selected 31 isolated forest patches of differing sizes, and three 80-ha plots in continuous forest each centered within non-overlapping 200-ha landscapes, such that patch area and landscape forest cover were uncorrelated among landscapes. Each study plot was surveyed to estimate abundances of territorial males and ovenbird pairing success. Landscape forest cover ( p <0.05) explained the most variation in ovenbird abundance, while percent deciduous forest cover within patches ( p <0.05) and patch size ( p <0.05) explained the most variation in red-eyed vireo and wood thrush abundance, respectively. Patch size was a significant ( p <0.05) predictor of abundance for all three study species; however, density for all species decreased significantly ( p <0.05) with patch size. Ovenbird pairing success was higher in continuous forest plots than in forest patches ( p =0.018). This study's findings suggest that the relative importance of within patch characteristics, patch size and landscape forest cover varies for different bird species, and that conservation efforts would benefit from the inclusion of all three factors.     

Remnants fragments preserve genetic diversity of the old forest lichen <Emphasis Type="Italic">Lobaria pulmonaria</Emphasis> in a fragmented Mediterranean mountain forest

Fragmentation represents a serious threat to biodiversity worldwide, however its effects on epiphytic organisms is still poorly understood. We study the effect of habitat fragmentation on the genetic population structure and diversity of the red-listed epiphytic lichen, Lobaria pulmonaria, in a Mediterranean forest landscape. We tested the relative importance of forest patch quality, matrix surrounding fragments and connectivity on the genetic variation within populations and the differentiation among them. A total of 855 thalli were sampled in 44 plots (400 m²) of 31 suitable forest fragments (beeches and oaks), in the Sierra de Ayllón in central Spain. Variables related to landscape attributes of the remnant forest patches such as size and connectivity and also the nature of the matrix or tree species had no significant effects on the genetic diversity of L. pulmonaria. Values of genetic diversity (Nei’s) were only affected by habitat quality estimated as the age patches. Most of the variation (76%) in all populations was observed at the smallest sampled unit (plots). Using multiple regression analysis, we found that habitat quality is more important in explaining the genetic structure of the L. pulmonaria populations than spatial distance. The relatively high level of genetic diversity of the species in old forest patches regardless of patch size indicates that habitat quality in a highly structured forest stand determines the population size and distribution pattern of this species and its associated lichen community. Thus, conservation programmes of Mediterranean mountain forests have to prioritize area and habitat quality of old forest patches.     

Information thresholds,habitat loss and population persistence in breeding birds

The combination of spatial structure and non‐linear population dynamics can promote the persistence of coupled populations, even when the average population growth rate of the patches seen in isolation would predict otherwise. This phenomenon has generally been conceptualized and investigated through the movement of individuals among patches that each holds many individuals, as in metapopulation models. However, population persistence can likewise increase as the result of individuals moving among sites (e.g. breeding territories) within in a single patch. Here I examine the latter: individuals making small‐scale informed decisions with respect to where to breed can promote population persistence in poor environments. Based on a simple algebraic model, I demonstrate information thresholds, and predict that greater information use is required for population persistence under lower spatial heterogeneity in habitat quality, all else equal. Second, I implement an individual‐based model to explore prior experience and prospecting on conspecific success within a more complex, and spatially heterogeneous environment. Uniquely, I jointly examine the effects of simulated habitat loss, spatial heterogeneity prior to habitat, and variation in information gathering on population persistence. I find that habitat loss accelerates population quasi‐extinction risk; however, information use reduces extinction probabilities in proportion to the level of information gathering. Per capita reproductive success declines with number of breeding sites, suggesting that information‐mediated Allee effects may contribute to extinction risk. In conclusion, my study suggests that populations in a changing world may be increasingly vulnerable to extinction where patch size and spatial heterogeneity constrain the effectiveness of information‐use strategies.     

Spatial Heterogeneity in Habitat Quality and Cross-Scale Interactions in Metapopulations

Abstract Integration of habitat heterogeneity into spatially realistic metapopulation approaches reveals the potential for key cross-scale interactions. Broad-scale environmental gradients and land-use practices can create autocorrelation of habitat quality of suitable patches at intermediate spatial scales. Patch occupancy then depends not only on habitat quality at the patch scale but also on feedbacks from surrounding neighborhoods of autocorrelated patches. Metapopulation dynamics emerge from how demographic and dispersal processes interact with relevant habitat heterogeneity. We provide an empirical example from a metapopulation of round-tailed muskrats (Neofiber alleni) in which habitat quality of suitable patches was spatially autocorrelated most strongly within 1,000 m, which was within the expected dispersal range of the species. After controlling for factors typically considered in metapopulation studies—patch size, local patch quality, patch connectivity—we use a cross-variogram analysis to demonstrate that patch occupancy by muskrats was correlated with habitat quality across scales ≤1,171 m. We also discuss general consequences of spatial heterogeneity of habitat quality for metapopulations related to potential cross-scale interactions. We focus on spatially correlated extinctions and metapopulation persistence, hierarchical scaling of source–sink dynamics, and dispersal decisions by individuals in relation to information constraints.     

Matrix heterogeneity affects population size of the harvest mice: Bayesian estimation of matrix resistance and model validation

It has been increasingly recognized that landscape matrices are an important factor determining patch connectivity and hence the population size of organisms living in highly fragmented landscapes. However, most previous studies estimated the effect of matrix heterogeneity using prior information regarding dispersal or habitat preferences of a focal organism. Here we estimated matrix resistance of harvest mice in agricultural landscapes using a novel pattern‐oriented modeling with Bayesian estimation and no prior information, and then conducted model validation using data sets independent from those used for model construction. First, we investigated the distribution patterns of harvest mice for approximately 400 habitat patches, and estimated matrix resistance for different matrix types using statistical models incorporating patch size, patch environment, and patch connectivity. We used Bayesian estimation with a Markov chain Monte Carlo algorithm, and searched for appropriate matrix resistance that best explained the distribution pattern. Patch connectivity as well as patch quality was an important determinant of local population size for the harvest mice. Moreover, matrix resistance was far from uniform, with rice and crop fields exhibiting low resistance and forests, creeks, roads and residential areas showing much higher resistance. The deviance explained by this model (heterogeneous matrix model) was much larger than that obtained by the model with no consideration of matrix heterogeneity (homogeneous matrix model). Second, we obtained distribution data from five additional landscapes that were more fragmented than that used for model construction, and used them for model validation. The heterogeneous matrix model well predicted the population size for four out of five landscapes. In contrast, the homogeneous model considerably overestimated population sizes in all cases. Our approach is widely applicable to species living in fragmented landscapes, especially those for which prior information regarding movement or dispersal is difficult to obtain.     

How landscape scale changes affect ecological processes in conservation areas: external factors influence land use by zebra (Equus burchelli) in the Okavango Delta

Most large‐bodied wildlife populations in sub‐Saharan Africa only survive in conservation areas, but are continuing to decline because external changes influence ecological processes within reserves, leading to a lack of functionality. However, failure to understand how landscape scale changes influence ecological processes limits our ability to manage protected areas. We used GPS movement data to calculate dry season home ranges for 14 zebra mares in the Okavango Delta and investigated the effects of a range of landscape characteristics (number of habitat patches, mean patch shape, mean index of juxtaposition, and interspersion) on home range size. Resource utilization functions (RUF) were calculated to investigate how specific landscape characteristics affected space use. Space use by all zebra was clustered. In the wetter (Central) parts of the Delta home range size was negatively correlated with the density of habitat patches, more complex patch shapes, low juxtaposition of habitats and an increased availability of floodplain and grassland habitats. In the drier (Peripheral) parts of the Delta, higher use by zebra was also associated with a greater availability of floodplain and grassland habitats, but a lower density of patches and simpler patch shapes. The most important landscape characteristic was not consistent between zebra within the same area of the Delta, suggesting that no single foraging strategy is substantially superior to others, and so animals using different foraging strategies may all thrive. The distribution and complexity of habitat patches are crucial in determining space use by zebra. The extent and duration of seasonal flooding is the principal process affecting habitat patch characteristics in the Okavango Delta, particularly the availability of floodplains, which are the habitat at greatest risk from climate change and anthropogenic disturbance to the Okavango's catchment basin. Understanding how the factors that determine habitat complexity may change in the future is critical to the conservation of large mammal populations. Our study shows the importance of maintaining flood levels in the Okavango Delta and how the loss of seasonal floodplains will be compounded by changes in habitat configuration, forcing zebra to change their relative space use and enlarge home ranges, leading to increased competition for key resources and population declines.