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.     

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.     

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.     

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     

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.     

Osmotic and diffusive flows in single-file pores: new approach to modeling pore occupancy states

Background

The relation between osmotic permeability, P_f, diffusion permeability, P_d, and the number of water molecules, N_p, in the single-file membrane pore remains an open question. Theoretical analyses, empirical studies on aquaporins and nanotubes, and molecular dynamics simulations have yet to provide a consensus view.

Results

This paper presents a new combinatorial analysis of the different pore states formed from water molecules and the presence of a vacancy that differs from the several previous combinatorial approaches to analyzing pore states. It is the first such analysis to show that P_f / P_d?=?N_p. It is rooted in the concept of different classes of pore occupancy states, tracer states and tracer exit states, present in the pore. This includes pores with and without a single vacancy. The concepts of knock-on collisions and concerted Brownian fluctuations provide the mechanisms underlying the behaviors of the tracer and vacancy as each moves through the pore during osmotic or diffusive flow. It develops the important role of the knock-on collision mechanism for osmotic flow. An essential feature of the model is the presence, or absence, of a single vacancy in the pore. The vacancy slows down tracer translocation through the pore. Its absence facilitates osmotic flow.

Conclusions

The full pore states and the single vacancy states together with the knock-on and Brownian mechanisms account for the relative values of P_f and P_d during osmotic and diffusive flow through the single-file pore. The new approach to combinatorial analysis differs from previous approaches and is the first to show a simple intuitive basis for the relation P_f / P_d?=?N_p. This resolves a long persisting dichotomy.

     

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.     

Detecting local adaptation in a natural plant-pathogen metapopulation: a laboratory vs. field transplant approach

Antagonistic coevolution between hosts and parasites in spatially structured populations can result in local adaptation of parasites. Traditionally parasite local adaptation has been investigated in field transplant experiments or in the laboratory under a constant environment. Despite the conceptual importance of local adaptation in studies of (co)evolution, to date no study has provided a comparative analysis of these two methods. Here, using information on pathogen population dynamics, I tested local adaptation of the specialist phytopathogen, Podosphaera plantaginis, to its host, Plantago lanceolata at three different spatial scales: sympatric host population, sympatric host metapopulation and allopatric host metapopulations. The experiment was carried out as a field transplant experiment with greenhouse-reared host plants from these three different origins introduced into four pathogen populations. In contrast to results of an earlier study performed with these same host and parasite populations under laboratory conditions, I did not find any evidence for parasite local adaptation. For interactions governed by strain-specific resistance, field studies may not be sensitive enough to detect mean parasite population virulence. Given that parasite transmission potential may be mediated by the abiotic environment and genotype-by-environment interactions, I suggest that relevant environmental variation should be incorporated into laboratory studies of parasite local adaptation.     

Assessing the impact of local habitat variables and landscape context on riparian birds in agricultural,urbanized, and native landscapes

Large tracts of natural habitat are being replaced by agriculture and urban sprawl in Mediterranean regions worldwide. We have limited knowledge about the effects of human activities on native species in these landscapes and which, if any, management practices might enhance the conservation of native biodiversity within them. Through a citizen volunteer bird-monitoring project, we compared bird abundance and species richness in northern Californian riparian zones surrounded by vineyards, urban areas, and natural areas. We assessed both local and landscape-level variables that may enhance native bird diversity in each land use type. We also demonstrate a new statistical approach, generalized estimating equations, to analyze highly variable data, such as that collected by volunteers. Avian abundance was highly correlated with both landscape context and local habitat variables, while avian richness was correlated with local habitat variables, specifically shrub richness, and percent of tree cover. In particular, shrub species richness has a strong positive correlation with riparian-preferring bird species. This suggests that active local management of riparian zones in human-dominated landscapes can increase our ability to retain native bird species in these areas.     

Monitoring arthropods in a tropical landscape: relative effects of sampling methods and habitat types on trap catches

To discuss the challenge of monitoring multi-species responses of tropical arthropods to disturbance, we considered a large dataset (4 × 10⁵ individuals; 1,682 morphospecies representing 22 focal taxa) based on the work of parataxonomists to examine the effects of anthropogenic disturbance on arthropods at Gamba, Gabon. Replication included three sites in each of four different stages of forest succession and land use after logging, surveyed during a whole year with four sampling methods: pitfall, Malaise, flight-interception and yellow pan traps. We compared the suitability of each sampling method for biological monitoring and evaluated statistically their reliability for 118 arthropod families. Our results suggest that a range of sampling methods yields more diverse material than any single method operated with high replication. Multivariate analyses indicated that morphospecies composition in trap catches was more strongly influenced by habitat type than by sampling methods. This implies that for multi-species monitoring, differences in trap efficiency between habitats may be neglected, as far as habitat types remain well contrasted. We conclude that for the purpose of monitoring large arthropod assemblages in the long-term, a protocol based on operating a set of different and non-disruptive traps appears superior in design than summing a series of taxa-specific protocols.     

A landscape genetics approach for quantifying the relative influence of historic and contemporary habitat heterogeneity on the genetic connectivity of a rainforest bird

Landscape genetics is an important framework for investigating the influence of spatial pattern on ecological process. Nevertheless, the standard analytic frameworks in landscape genetics have difficulty evaluating hypotheses about spatial processes in dynamic landscapes. We use a predictive hypothesis-driven approach to quantify the relative contribution of historic and contemporary processes to genetic connectivity. By confronting genetic data with models of historic and contemporary landscapes, we identify dispersal processes operating in naturally heterogeneous and human-altered systems. We demonstrate the approach using a case study of microsatellite polymorphism and indirect estimates of gene flow for a rainforest bird, the logrunner ( Orthonyx temminckii ). Of particular interest was how much information in the genetic data was attributable to processes occurring in the reconstructed historic landscape and contemporary human-modified landscape. A linear mixed model was used to estimate appropriate sampling variance from nonindependent data and information-theoretic model selection provided strength of evidence for alternative hypotheses. The contemporary landscape explained slightly more information in the genetic differentiation data than the historic landscape, and there was considerable evidence for a temporal shift in dispersal pattern. In contrast, migration rates estimated from genealogical information were primarily influenced by contemporary landscape change. We discovered that landscape heterogeneity facilitated gene flow before European settlement, but contemporary deforestation is rapidly becoming the most important barrier to logrunner dispersal.     

Patch occupancy in the endangered butterfly Lycaena helle in a fragmented landscape: effects of habitat quality, patch size and isolation

While there is agreement that both habitat quality and habitat network characteristics (such as patch size and isolation) contribute to the occupancy of patches by any given species, the relative importance of these factors is under debate. This issue is of fundamental ecological importance, and moreover of special concern for conservation biologists aiming at preserving endangered species. Against this background we investigated patch occupancy in the violet copper Lycaena helle, one of the rarest butterfly species in Central Europe, in the Westerwald area (Rhineland-Palatinate, Western Germany). Occupied (n = 102) differed from vacant (n = 128) patches in altitude, size, connectivity, availability of wind shelter, in the abundance of the larval host-plant, in the abundance of a grass species indicating favorable habitat conditions and in the abundance of nitrophilous plants. Overall, patch occupancy was primarily determined by patch size, connectivity and the abundance of the larval host plant, while all other parameters of habitat quality were of subordinate importance. Therefore, our findings suggest that even for extremely sedentary species such as L. helle habitat networks are decisive and—next to the preservation of habitat quality—need to be an integral part of any conservation management for this species.