Distinguishing habitat types and the relative influences of environmental factors on patch occupancy for a butterfly metapopulation

The goals of this study were to examine whether subjective habitat types may be segregated based on environmental variables and whether the relative influences of environmental factors on patch occupancy differ among habitat types. We examined these questions using a metapopulation of the butterfly Speyeria nokomis carsonensis, surveying sites for environmental characteristics and butterfly presence/absence over a 2-year period. Discriminant function analyses correctly classified 84% of sites within habitat types based on environmental variables that explained 82 and 67% of the variance along two ordination axes. Occupancy models for each habitat type demonstrated that the relative influences of environmental factors differed among habitat types. This study broadens the current paradigm of metapopulation dynamics by demonstrating that systems exist in which distinct habitat types can be delineated and differential suites of environmental factors explain occupancy according to habitat type. These findings demonstrate that even for highly specialized species, all habitat areas cannot be assumed to contribute equally to occupancy status, and thus have comparable conservation value. Designing land-use plans and policies based, in part, on careful habitat assessments can have important implications for the conservation of species existing in patchy landscapes, and can help guide reserve design, management actions, and allocation of resources on multiple-use landscapes.     

An improved multi-scale approach to modeling habitat occupancy of northern bobwhite

Predicting species presence requires knowledge of detection of individuals, scale of model variables, model selection uncertainty, and spatial autocorrelation. Our objective was to incorporate recent modeling advances to predict potential habitat occupancy of northern bobwhite (Colinus virginianus). From 15 May–15 August 2008 and 2009, we conducted repeat-visit surveys at 360 sites within Delaware to sample presence of bobwhite. We randomly selected half the data to model scale-dependent relationships of bobwhite presence with metrics of landscape- and site-scale habitat composition and configuration. The final averaged habitat-occupancy model fit the remainder testing dataset with an area under the receiver operating characteristic curve value of 0.62. At the site scale, bobwhite presence was negatively related to interspersion and juxtaposition of early successional habitat (ESH; grassland and shrubland), ESH to forest edge density, and agriculture to forest edge density, though relative effect sizes were weak to moderate after accounting for model selection uncertainty. At the landscape scale, bobwhite presence was negatively related to patch cohesion of human development within 2.5 km and positively related to patch cohesion of ESH within 2.0 km, with both variables exerting strong effects. Bobwhite presence was also weakly and positively related to percentage of shrubland habitat within 1.0 km of the sampling point. We applied our habitat occupancy model to map the predicted presence of breeding bobwhite within the Delmarva Peninsula, USA. The modeling results and distribution map will provide guidance to State and Federal private land management programs in the Mid-Atlantic to identify where habitat management efforts will be most effective. Our methodology can also serve as a basis for future habitat modeling of bobwhite and other grassland–shrubland species across their range. © 2011 The Wildlife Society.     

Long distance dispersal and landscape occupancy in a metapopulation of the cranberry fritillary butterfly

Movements between habitat patches in a patchy population of the butterfly Boloria aquilonaris were monitored using capture-mark-recapture methods during three successive generations. For each data set, the inverse cumulative proportion of individuals moving 100 m distance classes was fitted to the negative exponential function and the inverse power function. In each case, the negative exponential function provided a better fit than the inverse power function. Two dispersal kernels were generated using both negative exponential and inverse power functions. These dispersal kernels were used to predict movements between 14 suitable sites in a landscape of 220 km². The negative exponential function generated a dispersal kernel predicting extremely low probabilities for movements exceeding 1 km. The inverse power function generated probabilities predicting that between site movements were possible, according to metapopulation size. CMR studies in the landscape revealed that long distance movements occurred at each generation, corresponding to predictions of the inverse power function dispersal kernel. A total of 26 movements between sites (up to 13 km) were detected, together with recolonisation of empty sites. The spatial scale of the metapopulation dynamics is larger than ever reported on butterflies and long distance movements clearly matter to the persistence of this species in a highly fragmented landscape.     

Linking habitat use to range expansion rates in fragmented landscapes: a metapopulation approach

Temperature increases because of climate change are expected to cause expansions at the high latitude margins of species distributions, but, in practice, fragmented landscapes act as barriers to colonization for most species. Understanding how species distributions will shift in response to climate change therefore requires techniques that incorporate the combined effects of climate and landscape‐scale habitat availability on colonization rates. We use a metapopulation model (Incidence Function Model, IFM) to test effects of fine‐scale habitat use on patterns and rates of range expansion by the butterfly Hesperia comma. At its northern range margin in Britain, this species has increased its breadth of microhabitat use because of climate warming, leading to increased colonization rates. We validated the IFM by reconstructing expansions in five habitat networks between 1982 and 2000, before using it to predict metapopulation dynamics over 100 yr, for three scenarios based on observed changes to habitat use. We define the scenarios as “cold‐world” (only hot, south‐facing 150–250° hillsides are deemed warm enough), “warm‐world” in which 100–300° hillsides can be populated, and “hot‐world”, where the background climate is warm enough to enable use of all aspects (as increasingly observed). In the simulations, increased habitat availability in the hot‐world scenario led to faster range expansion rates, and to long‐term differences in distribution size and pattern. Thus, fine‐scale changes in the distribution of suitable microclimates led to landscape‐scale changes in population size and colonization rate, resulting in coarse‐scale changes to the species distribution. Despite use of a wider range of habitats associated with climate change, H. comma is still expected to occupy a small fraction of available habitat in 100 yr. The research shows that metapopulation models represent a potential framework to identify barriers to range expansion, and to predict the effects of environmental change or conservation interventions on species distributions and persistence.     

Predator-prey interactions in a nonequilibrium context: the metapopulation approach to modeling "hide-and-seek" dynamics in a spatially explicit tri-trophic system

Predators and prey are usually heterogeneously distributed in space so that the ability of the predators to respond to the distribution of their prey may have a profound influence on the stability and persistence of a predator‐prey system. A special type of dynamics is “hide‐and‐seek” characterized by a high turnover rate of local populations of prey and predators, because once the predators have found a patch of prey they quickly overexploit it, whereupon the starving predators either should move to better places or die. Continued persistence of prey and predators thus hinges on a long‐term balance between local extinctions and founding of new subpopulations. The colonization rate depends on the rate of emigration from occupied patches and the likelihood of successfully arriving at a suitable new patch, while extinction rate depends on the local population dynamics. Since extinctions and colonizations are both discrete probabilistic events, these phenomena are most adequately modeled by means of a stochastic model. In order to demonstrate the qualitative differences between a deterministic and stochastic approach to population dynamics, a spatially explicit tritrophic predator‐prey model is developed in a deterministic and a stochastic version. The model is parameterized using data for the two‐spotted spider mite (Tetranychus urticae) and the phytoseiid mite predator Phytoseiulus persimilis inhabiting greenhouse cucumbers.
Simulations show that the deterministic and stochastic approaches yield different results. The deterministic version predicts that the populations will exhibit violent fluctuations, implying that the system is fundamentally unstable. In contrast, the stochastic version predicts that the two species will be able to coexist in spite of frequent local extinctions of both species, provided the system consists of a sufficiently large number of local populations. This finding is in agreement with experimental results. It is therefore concluded that demographic stochasticity in combination with dispersal is capable of producing and maintaining sufficient asynchrony between local populations to ensure long‐term regional (metapopulation) persistence.     

The relative effects of local management and landscape context on weed seed predation and carabid functional groups

Farmland biodiversity and its associated ecosystem services are affected by agricultural activities at multiple spatial scales. Among these services, the regulation of weeds by invertebrate seed predators has received much attention recently but little is known about the relative effect of local management and landscape context of fields on this process. We monitored seed predation on four common weed species and carabid communities in 28 winter-cereals fields during five consecutive weeks in spring 2010. These fields were situated in contrasted landscape contexts and varied in terms of intensity of pesticide treatments and soil tillage regimes. Weed seed predation was strongly and positively related to the Shannon diversity of (strictly) granivorous carabids and to the activity–density of omnivorous carabids but negatively to the richness of omnivorous carabids. Weed seed predation and granivore diversity were positively related to landscape diversity and the proportion cover of temporary grassland within a 1000 m radius around focal fields and were negatively affected by the intensity of local pesticide treatments. No-till systems sheltered higher diversity of granivorous carabids but did not show higher seed predation rates. We showed that landscape composition factors had a higher relative influence than local practices factors on weed seed predation service. Consequently, weed management strategies should not only consider the management of single fields but also the surrounding landscape to preserve carabid biodiversity and enhance weed seed predation service.     

Dynamic populations in a dynamic landscape: the metapopulation structure of the marsh fritillary butterfly

The marsh fritillary butterfly Euphydryas aurinia is an endangered species in most of northern Europe. We describe the metapopulation structure of E. aurinia in Finland, where the species has declined drastically in the past decades. We found two types of habitat patches suitable for the species: semi-permanent meadows and transient clearcuts in the forest. Patch area was the most significant variable predicting the occurrence of E. aurinia in a habitat patch. The species tended to be found in young rather than old clearcuts, apparently because the vegetation became too high in the latter. We used the incidence function model to simulate the metapopulation dynamics of E. aurinia in its dynamic landscape and discovered that the continued presence of the semi-permanent meadows is essential for the survival of the species in the study area in southeast Finland.     

The role of local populations within a landscape context: defining and classifying sources and sinks

The interaction of local populations has been the focus of an increasing number of studies in the past 30 years. The study of source-sink dynamics has especially generated much interest. Many of the criteria used to distinguish sources and sinks incorporate the process of apparent survival (i.e., the combined probability of true survival and site fidelity) but not emigration. These criteria implicitly treat emigration as mortality, thus biasing the classification of sources and sinks in a manner that could lead to flawed habitat management. Some of the same criteria require rather restrictive assumptions about population equilibrium that, when violated, can also generate misleading inference. Here, we expand on a criterion (denoted "contribution" or Cr) that incorporates successful emigration in differentiating sources and sinks and that makes no restrictive assumptions about dispersal or equilibrium processes in populations of interest. The metric Cr is rooted in the theory of matrix population models, yet it also contains clearly specified parameters that have been estimated in previous empirical research. We suggest that estimates of emigration are important for delineating sources and sinks and, more generally, for evaluating how local populations interact to generate overall system dynamics. This suggestion has direct implications for issues such as species conservation and habitat management.     

The effects of grassland management using fire on habitat occupancy and conservation of birds in a mosaic landscape

Prescribed burning is routinely used to improve grazing in Pyrenean rangelands affected by an overall trend of land abandonment. This study considers the environmental variables influencing habitat occupancy by birds and the consequences of the use of fire in range management for bird conservation. Bird use and habitat structure of 11 cover types, the result of specific management regimes, were monitored for two breeding seasons in a mosaic landscape. Three main gradients of avian composition, corresponding to tree cover, shrub volume and grazing intensity, were identified from canonical correspondence analysis. The structure of the bird community seemed more intensely affected by species-specific selection of cover types than by the birds' use of multiple patches. Out of a total of 10 bird species analysed by a simultaneous confidence intervals procedure, four species with an unfavourable conservation status in Europe (Emberiza cia, Lullula arborea, Saxicola torquata and Lanius collurio) preferred managed grassland. Three types of grassland with shrubs (derived from single or repeated burning) had the highest bird conservation index (taking into account specific status and abundance of the bird assemblage), whereas forests showed middle or low values. The relation (P = 0.054) of this index to the logarithm of the pastoral value (which includes density and grazing quality of grasses) in currently managed cover types suggests that the objectives of grassland recovery by appropriate management practices and those of bird conservation coincide in our study area.     

The importance of local and landscape-scale processes to the occupancy of wetlands by pond-breeding amphibians

Variation in the distribution and abundance of species across landscapes has traditionally been attributed to processes operating at fine spatial scales (i.e., environmental conditions at the scale of the sampling unit), but processes that operate across larger spatial scales such as seasonal migration or dispersal are also important. To determine the relative importance of these processes, we evaluated hypothesized relationships between the probability of occupancy in wetlands by two amphibians [wood frogs (Lithobates sylvaticus) and boreal chorus frogs (Pseudacris maculata)] and attributes of the landscape measured at three spatial scales in Rocky Mountain National Park, Colorado. We used cost-based buffers and least-cost distances to derive estimates of landscape attributes that may affect occupancy patterns from the broader spatial scales. The most highly ranked models provide strong support for a positive relationship between occupancy by breeding wood frogs and the amount of streamside habitat adjacent to a wetland. The model selection results for boreal chorus frogs are highly uncertain, though several of the most highly ranked models indicate a positive association between occupancy and the number of neighboring, occupied wetlands. We found little evidence that occupancy of either species was correlated with local-scale attributes measured at the scale of individual wetlands, suggesting that processes operating at broader scales may be more important in influencing occupancy patterns in amphibian populations.     

Long-term demographic surveys reveal a consistent relationship between average occupancy and abundance within local populations of a butterfly metapopulation

Species distribution models are the tool of choice for large-scale population monitoring, environmental association studies and predictions of range shifts under future environmental conditions. Available data and familiarity of the tools rather than the underlying population dynamics often dictate the choice of specific method – especially for the case of presence–absence data. Yet, for predictive purposes, the relationship between occupancy and abundance embodied in the models should reflect the actual population dynamics of the modelled species. To understand the relationship of occupancy and abundance in a heterogeneous landscape at the scale of local populations, we built a spatio-temporal regression model of populations of the Glanville fritillary butterfly Melitaea cinxia in a Baltic Sea archipelago. Our data comprised nineteen years of habitat surveys and snapshot data of land use in the region. We used variance partitioning to quantify relative contributions of land use, habitat quality and metapopulation covariates. The model revealed a consistent and positive, but noisy relationship between average occupancy and mean abundance in local populations. Patterns of abundance were highly variable across years, with large uncorrelated random variation and strong local population stochasticity. In contrast, the spatio-temporal random effect, habitat quality, population connectivity and patch size explained variation in occupancy, vindicating metapopulation theory as the basis for modelling occupancy patterns in fragmented landscapes. Previous abundance was an important predictor in the occupancy model, which points to a spillover of abundance into occupancy dynamics. While occupancy models can successfully model large-scale population structure and average occupancy, extinction probability estimates for local populations derived from occupancy-only models are overconfident, as extinction risk is dependent on actual, not average, abundance.     

Site and landscape features ruling the habitat use and occupancy of the polecat (Mustela putorius) in a low density area: a multiscale approach

We studied the habitat of the polecat at different scales in a low density area. For this purpose we gathered data on the presence of the species and characterised them by location, home range and landscape scales. Polecats selected areas of high diversity close to, but not in, streams whilst avoiding intensively managed conifer plantations and dense urban areas. Variables determining the presence/absence of the species were found at home range scales, which implies that management and conservation practices for the species should be aimed mainly at this scale. Finally, our results agree with previously published works, which validate GIS-based approaches as a tool for carnivore management in areas with scarce data or in cases of rare species.     

The effects of habitat destruction in finite landscapes: a chain-binomial metapopulation model

We present a stochastic model for metapopulations in landscapes with a finite but arbitrary number of patches. The model, similar in form to the chain-binomial epidemic models, is an absorbing Markov chain that describes changes in the number of occupied patches as a sequence of binomial probabilities. It predicts the quasi-equilibrium distribution of occupied patches, the expected extinction time (τ¯), and the probability of persistence (l¯(x)) to time x as a function of the number N of patches in the landscape and the number S of those patches that are suitable for the population. For a given value of N , the model shows that: (1) τ¯ and l¯(x) are highly sensitive to changes in S and (2) there is a threshold value of S at which τ¯ declines abruptly from extremely large to very small values. We also describe a statistical method for estimating model parameters from time series data in order to evaluate metapopulation viability in real landscapes. An example is presented using published data on the Glanville fritillary butterfly, Meltiaea cinxia , and its specialist parasitoid Cotesia melitaearum . We calculate the expected extinction time of M. cinxia as a function of the frequency of parasite outbreaks, and are able to predict the minimum number of years between outbreaks required to ensure long-term persistence of M. cinxia . The chain-binomial model provides a simple but powerful method for assessing the effects of human and natural disturbances on extinction times and persistence probabilities in finite landscapes.     

The landscape ecological approach in bird conservation: integrating the metapopulation concept into spatial planning

In The Netherlands, fragmentation of (semi)natural ecosystems is regarded as a major nature conservation problem. The current Dutch Nature Conservation Policy Plan proposes a spatial network consisting of existing nature reserves, nature redevelopment areas and corridor zones. One of the objectives is to stop the assumed decline of biodiversity due to fragmentation. In this contribution we show that breeding birds are affected by the spatial distribution of their habitat. We also show how problems due to fragmentation can be solved by integrating landscape ecological research data Into planning procedures. Fragmented bird populations show metapopulation characteristics, dependent on the degree of fragmentation. This can be concluded from pattern studies, in which presence or absence patterns are correlated with spatial characteristics. Metapopulation dynamics were used, and the frequency of local extinction and of recolonization were related to size of habitat patches and spatial position in the landscape. The conclusion is that, depending on the spatial scale, landscape fragmentation is a threat to birds. On the basis of empirical data, statistical and metapopulation models are being developed to be used to evaluate spatial planning scenarios and to support decision making about which scenario is closest to the planning aims.     

集成灰色分析和元胞自动机用于景观动态模拟

针对景观生态系统"复杂性"的特点,采用灰色局势确定邻域转换规则,构建元胞自动机,用于模拟和分析景观生态动态变化.在信息不完全的情况下,提高了元胞自动机景观生态动态模型的可靠性和可行性.以中国云南省怒江流域中段作为实例,确定影响元胞转换规则的三因素主要有元胞邻域的聚集程度,土地适宜程度和人类活动影响程度,根据景观这一类复杂系统的动态变化特征其权重随地理空间位置和时间而变化,同时用Monte Carlo法考虑了模拟时转换的随机性.通过计算显示模拟未来发展的情景总的趋势是符合实际的,不但模拟了微观的景观单元的自组织机制,而且在一定程度上反映了宏观的社会经济因素影响,因而更具有针对性、典型性及准确性.     

Distribution of birds in natural landscape mosaics of old-growth forests in northern Sweden: relations to habitat area and landscape context

We censused breeding birds for three years in natural landscape mosaics of virgin old-growth spruce forest and mire in a large protected forest area in northern Sweden Twenty forest patches, ranging from 0 2 to 17 8 ha in size, were selected in two matrix types, dominated by forest and mire, respectively Patches were very similar with regards to habitat features There was a strong effect of patch area on species richness, but no effect of matrix type Standardization of species richness by rarefaction revealed that small patches (<5 ha) had fewer and large patches (>10 ha) more species than expected Overall distribution of species across patches showed a highly significant nested pattern, indicating that a few habitat generalists occupy all size classes, whereas more demanding species avoid small patches regardless of landscape composition Individual species tended to be distributed evenly across patch classes and no significant edge effect in terms of density of birds was found Our results have bearings on actions to preserve avian diversity in northern boreal forests small patches (<5 ha) provide habitat only for habitat generalists, and therefore larger (>10 ha) patches should be preserved     

Spatially structured metapopulation models: global and local assessment of metapopulation capacity

We model metapopulation dynamics in finite networks of discrete habitat patches with given areas and spatial locations. We define and analyze two simple and ecologically intuitive measures of the capacity of the habitat patch network to support a viable metapopulation. Metapopulation persistence capacity lambda(M) defines the threshold condition for long-term metapopulation persistence as lambda(M)>delta, where delta is defined by the extinction and colonization rate parameters of the focal species. Metapopulation invasion capacity lambda(I) sets the condition for successful invasion of an empty network from one small local population as lambda(I)>delta. The metapopulation capacities lambda(M) and lambda(I) are defined as the leading eigenvalue or a comparable quantity of an appropriate "landscape" matrix. Based on these definitions, we present a classification of a very general class of deterministic, continuous-time and discrete-time metapopulation models. Two specific models are analyzed in greater detail: a spatially realistic version of the continuous-time Levins model and the discrete-time incidence function model with propagule size-dependent colonization rate and a rescue effect. In both models we assume that the extinction rate increases with decreasing patch area and that the colonization rate increases with patch connectivity. In the spatially realistic Levins model, the two types of metapopulation capacities coincide, whereas the incidence function model possesses a strong Allee effect characterized by lambda(I)=0. For these two models, we show that the metapopulation capacities can be considered as simple sums of contributions from individual habitat patches, given by the elements of the leading eigenvector or comparable quantities. We may therefore assess the significance of particular habitat patches, including new patches that might be added to the network, for the metapopulation capacities of the network as a whole. We derive useful approximations for both the threshold conditions and the equilibrium states in the two models. The metapopulation capacities and the measures of the dynamic significance of particular patches can be calculated for real patch networks for applications in metapopulation ecology, landscape ecology, and conservation biology.