The influence of patch demographics on metapopulations, with particular reference to successional landscapes

The classical view of metapopulations relates the regional abundance of a species to the balance between the extinction and colonization dynamics of identical local populations. Species in successional landscapes may represent the most appropriate examples of classical metapopulations. However, Levins‐type metapopulation models do not explicitly separate population loss due to successional habitat change from other causes of extinction. A further complication is that the chance of population loss due to successional habitat change may be related to the age of a patch. I developed simple patch occupancy models to include succession and included consideration of patch age structure to address two related questions: what are the implications of changes in patch demographic rates and when is a move to a structured patch occupancy model justified? Age‐related variation in patch demography could increase or decrease the equilibrium fraction of the available habitat occupied by a species when compared to the predictions of an unstructured model. Metapopulation persistence was enhanced when the age class of patches with the highest species occupancy suffered relatively low losses to habitat succession. Conversely, when the age class of patches with the highest species occupancy also had relatively high successional loss rates, extinction thresholds were higher that would be predicted by a simple unstructured model. Hence age‐related variation in patch successional rate introduces biases into the predictions of simple unstructured models. Such biases can be detected from field surveys of the fraction of occupied and unoccupied patches in each age class. Where a bias is demonstrated, unstructured models will not be adequate for making predictions about the effects of changing parameters on metapopulation size. Thinking in successional terms emphasizes how landscapes might be managed to enhance or reduce the patch occupancy by any particular metapopulation     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Metapopulation theory for fragmented landscapes

We review recent developments in spatially realistic metapopulation theory, which leads to quantitative models of the dynamics of species inhabiting highly fragmented landscapes. Our emphasis is in stochastic patch occupancy models, which describe the presence or absence of the focal species in habitat patches. We discuss a number of ecologically important quantities that can be derived from the full stochastic models and their deterministic approximations, with a particular aim of characterizing the respective roles of the structure of the landscape and the properties of the species. These quantities include the threshold condition for persistence, the contributions that individual habitat patches make to metapopulation dynamics and persistence, the time to metapopulation extinction, and the effective size of a metapopulation living in a heterogeneous patch network.     

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.     

The relative contribution of local habitat and landscape context to metapopulation processes: a dynamic occupancy modeling approach

Changes in site occupancy across habitat patches have often been attributed to landscape features in fragmented systems, particularly when considering metapopulations. However, failure to include habitat quality of individual patches can mask the relative importance of local scale features in determining distributional changes. We employed dynamic occupancy modeling to compare the strength of local habitat variables and metrics of landscape patterns as drivers of metapopulation dynamics for a vulnerable, high‐elevation species in a naturally fragmented landscape. Repeat surveys of Bicknell's thrush Catharus bicknelli presence/non‐detection were conducted at 88 sites across Vermont, USA in 2006 and 2007. We used an organism‐based approach, such that at each site we measured important local‐scale habitat characteristics and quantified landscape‐scale features using a predictive habitat model for this species. We performed a principal component analysis on both the local and landscape features to reduce dimensionality. We estimated site occupancy, colonization, and extinction probabilities while accounting for imperfect detection. Univariate, additive, and interaction models of local habitat and landscape context were ranked using AICc scores. Both local and landscape scales were important in determining changes in occupancy patterns. An interaction between scales was detected for occupancy dynamics indicating that the relationship of the parameters to local‐scale habitat conditions can change depending on the landscape context and vice versa. An increase in both landscape‐ and local‐scale habitat quality increased occupancy and colonization probability while decreasing extinction risk. Colonization and extinction were both more strongly influenced by local habitat quality relative to landscape patterns. We also identified clear, qualitative thresholds for landscape‐scale features. Conservation of large habitat patches in high‐cover landscapes will help ensure persistence of Bicknell's thrushes, but only if local scale habitat quality is maintained. Our results highlight the importance of incorporating information beyond landscape characteristics when investigating patch occupancy patterns in metapopulations.     

Metapopulation-level adaptation of insect host plant preference and extinction-colonization dynamics in heterogeneous landscapes

Species living in highly fragmented landscapes typically occur as metapopulations with frequent turnover of local populations. The turnover rate depends on population sizes and connectivities, but it may also depend on the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinxia) in Finland uses two host plant species, which show variation in their relative abundances at two spatial scales: locally among individual habitat patches and regionally among networks of patches. Female butterflies in turn exhibit spatial variation in genetically determined host plant preference within and among patch networks. Emigration, immigration and establishment of new populations have all been shown to be strongly influenced by the match between the host plant composition of otherwise suitable habitat patches and the host plant preference of migrating butterflies. The evolutionary consequences of such biased migration and colonization with respect to butterfly phenotypes might differ depending on spatial configuration and plant species composition of the patches in heterogeneous patch networks. Using a spatially realistic individual-based model we show that the model-predicted evolution of host plant preference due to biased migration explains a significant amount of the observed variation in host plant use among metapopulations living in dissimilar networks. This example illustrates how the ecological extinction-colonization dynamics may be linked with the evolutionary dynamics of life history traits in metapopulations.     

Colonization and extinction dynamics of an annual plant metapopulation in an urban environment

The metapopulation framework considers that the spatiotemporal distribution of organisms results from a balance between the colonization and extinction of populations in a suitable and discrete habitat network. Recent spatially realistic metapopulation models have allowed patch dynamics to be investigated in natural populations but such models have rarely been applied to plants. Using a simple urban fragmented population system in which favourable habitat can be easily mapped, we studied patch dynamics in the annual plant Crepis sancta (Asteraceae). Using stochastic patch occupancy models (SPOMs) and multi‐year occupancy data we dissected extinction and colonization patterns in our system. Overall, our data were consistent with two distinct metapopulation scenarios. A metapopulation (sensu stricto) dynamic in which colonization occurs over a short distance and extinction is lowered by nearby occupied patches (rescue effect) was found in a set of patches close to the city centre, while a propagule rain model in which colonization occurs from a large external population was most consistent with data from other networks. Overall, the study highlights the importance of external seed sources in urban patch dynamics. Our analysis emphasizes the fact that plant distributions are governed not only by habitat properties but also by the intrinsic properties of colonization and dispersal of species. The metapopulation approach provides a valuable tool for understanding how colonization and extinction shape occupancy patterns in highly fragmented plant populations. Finally, this study points to the potential utility of more complex plant metapopulation models than traditionally used for analysing ecological and evolutionary processes in natural metapopulations.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Landscape attributes affecting patch occupancy by howler monkeys (Alouatta palliata mexicana) at Los Tuxtlas, Mexico

Understanding how forest cover is related to patch attributes such as size, shape, and isolation, and how this influences the occurrence of a species in fragmented landscapes is an important question in landscape ecology and conservation biology. To study the effects of fragmentation on patch occupancy by the critically endangered Mexican mantled howler monkey (Alouatta palliata mexicana) in the Los Tuxtlas region of Mexico, we selected three landscapes of ca. 5,000 ha, which differed in their remaining forest cover (24, 11, and 4%). For each landscape, we related patch occupancy to forest cover, patch size and shape, and four isolation parameters. Landscape attributes varied according to forest cover, and the percentage of occupation was greater in landscapes with more forest cover. The attributes affecting the probability of occupancy differed among landscapes. Occupancy was positively related to patch size in all landscapes, but in the northernmost landscape, shape irregularity had a negative effect on occupancy, whereas in the southernmost landscape, occupancy was favored by greater distances to the nearest village. The results show that not only the total amount of forest cover but also patch configuration need to be taken into consideration when designing management strategies for the conservation of the Mexican mantled howler monkey.     

异质种群动态模型:破碎化景观动态模拟的新途径

景观破碎化导致物种以异质种群方式存活,使得基于异质种群动态模拟破碎化景观动态成为可能。异质种群动态模型的发展为景观动态模拟奠定了良好基础。根据空间处理方式的不同,异质种群模型可分为三大类,可不同程度地用于描述破碎化景观动态。(1)空间不确定异质种群模型,假定所有局域种群间均等互联,模型中不包含空间信息,仅能用于景观斑块动态描述;(2)空间确定异质种群模型,假设局域种群在二维空间上以规则格子形式排列,是一种准现实的空间处理方式,可用于景观动态的简单描述;(3)空间现实异质种群模型,包含了破碎化景观中局域种群的几何特征,可直接用于真实景观动态的模拟研究。空间现实的和基于个体的异质种群模型不但是未来异质种群模型发展的主流,也将成为未来破碎化景观动态研究的重要工具。为了更加准确完整地描述破碎化景观动态,不但应该综合运用已有的各种异质种群模型方法,更要引进新模型来刎画多物种、多变量、高维度、复杂连接的破碎化景观格局与过程。     

Functional connectivity as an indicator for patch occupancy in grassland specialists

Habitat specialists living in metapopulations are sensitive to habitat fragmentation. In most studies, the effects of fragmentation on such species are analyzed based on Euclidean inter-patch distances. This approach, however, ignores the role of the landscape matrix. Recently, therefore, functional distances that account for the composition of the landscape surrounding the habitat patches have been used more frequently as indicators for patch occupancy. However, the performance of functional and non-functional connectivity measures in predicting patch occupancy of such species has never been compared in a multi-species approach.Here we evaluate the effect of habitat connectivity on the patch occupancy of 13 habitat specialists from three different insect orders (Auchenorrhyncha, Lepidoptera, Orthoptera) in fragmented calcareous grasslands. In order to calculate functional distances we used four different sets of resistance values and rankings. We then modelled species’ occurrence using both Euclidean and functional (based on least-cost modelling) inter-patch distances as predictors.We found that functional connectivity measures provided better results than the non-functional approach. However, a functional connectivity measure that was based on very coarse land-cover data performed even better than connectivity measures that were based on much more detailed land-use data.In order to take into account possible effects of the landscape matrix on patch occupancy by habitat specialists, future metapopulation studies should use functional rather than Euclidean distances whenever possible. For practical applications, we recommend a ‘simple approach’ which requires only coarse land-cover data and in our study performed better than all other functional connectivity measures, even more complex ones.     

Refugia and connectivity sustain amphibian metapopulations afflicted by disease

Metapopulation persistence in fragmented landscapes depends on habitat patches that can support resilient local populations and sufficient connectivity between patches. Yet epidemiological theory for metapopulations has largely overlooked the capacity of particular patches to act as refuges from disease, and has suggested that connectivity can undermine persistence. Here, we show that relatively warm and saline wetlands are environmental refuges from chytridiomycosis for an endangered Australian frog, and act jointly with connectivity to sustain frog metapopulations. We coupled models of microclimate and infection probability to map chytrid prevalence, and demonstrate a strong negative relationship between chytrid prevalence and the persistence of frog populations. Simulations confirm that frog metapopulations are likely to go extinct when they lack environmental refuges from disease and lose connectivity between patches. This study demonstrates that environmental heterogeneity can mediate host–pathogen interactions in fragmented landscapes, and provides evidence that connectivity principally supports host metapopulations afflicted by facultative pathogens.     

Anthropogenic landscape change promotes asymmetric dispersal and limits regional patch occupancy in a spatially structured bird population

1. Local extinctions in habitat patches and asymmetric dispersal between patches are key processes structuring animal populations in heterogeneous environments. Effective landscape conservation requires an understanding of how habitat loss and fragmentation influence demographic processes within populations and movement between populations. 2. We used patch occupancy surveys and molecular data for a rainforest bird, the logrunner (Orthonyx temminckii), to determine (i) the effects of landscape change and patch structure on local extinction; (ii) the asymmetry of emigration and immigration rates; (iii) the relative influence of local and between-population landscapes on asymmetric emigration and immigration; and (iv) the relative contributions of habitat loss and habitat fragmentation to asymmetric emigration and immigration. 3. Whether or not a patch was occupied by logrunners was primarily determined by the isolation of that patch. After controlling for patch isolation, patch occupancy declined in landscapes experiencing high levels of rainforest loss over the last 100 years. Habitat loss and fragmentation over the last century was more important than the current pattern of patch isolation alone, which suggested that immigration from neighbouring patches was unable to prevent local extinction in highly modified landscapes. 4. We discovered that dispersal between logrunner populations is highly asymmetric. Emigration rates were 39% lower when local landscapes were fragmented, but emigration was not limited by the structure of the between-population landscapes. In contrast, immigration was 37% greater when local landscapes were fragmented and was lower when the between-population landscapes were fragmented. Rainforest fragmentation influenced asymmetric dispersal to a greater extent than did rainforest loss, and a 60% reduction in mean patch area was capable of switching a population from being a net exporter to a net importer of dispersing logrunners. 5. The synergistic effects of landscape change on species occurrence and asymmetric dispersal have important implications for conservation. Conservation measures that maintain large patch sizes in the landscape may promote asymmetric dispersal from intact to fragmented landscapes and allow rainforest bird populations to persist in fragmented and degraded landscapes. These sink populations could form the kernel of source populations given sufficient habitat restoration. However, the success of this rescue effect will depend on the quality of the between-population landscapes.     

Is metapopulation patch occupancy in nature well predicted by the Levins model?

Although the Levins model has made important theoretical contributions to ecology, its empirical support has not been conclusively established yet. We used published colonization and extinction data from 55 metapopulations to calculate their Levins equilibrium patch occupancy. Over all species, there were not significant differences between the observed patch occupancies and the Levins model's estimates. However, invertebrates and vertebrate species with some degree of threat had patch occupancies larger than the model's expectancies. A temporal sampling effect was found for invertebrate species, with departure from the Levins model decreasing as the length of the study period increased. There was a negative relationship between patch occupancy and extinction probability, as expected under the “rescue effect”. The high rates at which invertebrates produce propagules could lead the Levins model to underestimate patch occupancy, whereas the observed patch occupancy of threatened species may be a transient phenomenon that results from extinction probabilities that increase over time. Therefore, the Levins model captures the metapopulation dynamics of a wide range of species in a simple formula whereas its equilibrium point can be used as evidence of metapopulation stability. Although mechanistic models provide more precise and accurate metapopulation predictions, they also can sacrifice the generality and simplicity of the Levins model.     

Range shifting on a fragmented landscape

Projected responses of species' to climate change have so far included few of the factors that are important determinants of species' distributions within its range. In this paper we utilise a spatially explicit cellular lattice, colonisation–extinction model to investigate the effect of habitat loss, fragmentation and species characteristics on range shifting in response to climate change. Contrary to the predictions of patch occupancy in static climate models we show that fragmentation can have a positive effect on species survival when species have high colonisation rates. For species with low colonisation rates aggregative behaviours prevent success on fragmented landscapes at high levels of habitat loss, and range shifting is more successfully achieved where habitat is correlated. At levels of habitat loss near the extinction threshold, less fragmented landscapes can facilitate range shifting even for the best colonisers. We discuss how imposing a climate window may reduce percolation routes and have implications for the area of usable habitat at any given level of habitat availability. We demonstrate the importance of landscape structure for range shifting dynamics and argue that management of reserve networks needs to consider the requirements of species with different life history characteristics.     

The frequency of metapopulations, metacommunities and nestedness in a fragmented landscape

Metapopulation and metacommunity theory may be extremely useful for understanding community composition and species distributions in fragmented landscapes. However, how well particular spatial ecological models represent natural systems is not known and so the general importance of those models is unclear. In three naturally fragmented landscapes in Tasmania, Australia, I sampled beetles from 67 sites, including continuous forest, small forest patches in a sedgeland matrix, streamside forest, and the matrix. Of forty commonly captured beetle species, at least 37% do not form metapopulations of any kind because they do not disperse far enough, disperse too far, or occur in the matrix. Only 7.5% of the species showed distributions consistent with a mainland‐island metapopulation, which was surprising given the mainland‐island structure of the landscape. These patterns were also consistent with a classic metapopulation. Deterministic metapopulations were probably common, involving at least 32% of species. Most predicted metacommunity patterns were not observed because only subsets of the fauna followed a particular metacommunity model. Seven, mostly flying, species were influenced only by habitat quality, supporting the Species‐sorting or Mass‐effects models, and the strong distinction between the beetle fauna of Eucalyptus patches and the matrix suggested an over‐riding influence of a Species‐sorting process. Seven, mostly flightless, species showed a negative relationship with increasing patch isolation, while another six, mostly flying, species showed the opposite pattern, with higher frequency of occurrence in more isolated patches. Both groups were strongly nested. The flying species were probably excluded from less isolated sites through interactions with poor dispersers , providing a novel mechanism leading to a nested distribution. The Patch‐dynamics and Species‐sorting processes were equally consistent with the pattern, emphasizing that these models are relatively simplistic when confronted with real community data. Additional complexity needs to be built into the models to accommodate the diverse patterns observed in natural communities.     

Extinction-colonization dynamics and host-plant choice in butterfly metapopulations

Species living in highly fragmented landscapes often occur as metapopulations with frequent population turnover. Turnover rate is known to depend on ecological factors, such as population size and connectivity, but it may also be influenced by the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinxia) in Finland uses two host-plant species that vary in their relative abundances among distinct habitat patches (dry meadows) in a large network of approximately 1,700 patches. We found no effect of host species use on local extinction. In contrast, population establishment was strongly influenced by the match between the host species composition of an empty habitat patch and the relative host use by larvae in previous years in the habitat patches that were well connected to the target patch. This "colonization effect" could be due to spatially variable plant acceptability or resistance or to spatially variable insect oviposition preference or larval performance. We show that spatial variation in adult oviposition preference occurs at the relevant spatial scale and that the other possible causes of the colonization effect can be discounted. We conclude that the colonization effect is generated by host preference influencing the movement patterns of ovipositing females. Migrant females with dissimilar host preferences have different perceptions of relative patch quality, which influences their likelihood of colonizing patches with particular host composition.     

Thresholds of riparian forest use by terrestrial mammals in a fragmented Amazonian deforestation frontier

Species persistence in fragmented landscapes is intimately related to the quality, structure, and context of remaining habitat remnants. Riparian vegetation is legally protected within private landholdings in Brazil, so we quantitatively assessed occupancy patterns of terrestrial mammals in these remnants, examining under which circumstances different species effectively use them. We selected 38 riparian forest patches and five comparable riparian sites within continuous forest, at which we installed four to five camera-traps per site (199 camera-trap stations). Terrestrial mammal assemblages were sampled for 60 days per station during the dry seasons of 2013 and 2014. We modelled species occupancy and detection probabilities within riparian forest remnants, and examined the effects of patch size, habitat quality, and landscape structure on occupancy probabilities. We then scaled-up modelled occupancies to all 1915 riparian patches throughout the study region to identify which remnants retain the greatest potential to work as habitat for terrestrial vertebrates. Of the ten species for which occupancy was modelled, six responded to forest quality (remnant degradation, cattle intrusion, palm aggregations, and understorey density) or structure (remnant width, isolation, length, and area of the patch from which it originates). Patch suitability was lower considering habitat quality than landscape structure, and virtually all riparian remnants were unsuitable to maintain a high occupancy probability for all species that responded to forest patch quality or structure. Beyond safeguarding legal compliance concerning riparian remnant amount, ensuring terrestrial vertebrate persistence in fragmented landscapes will require curbing the drivers of forest degradation within private landholdings.     

Gap-crossing movements predict species occupancy in Amazonian forest fragments

In fragmented landscapes, species persistence within isolated habitat patches is governed by a myriad of species life‐history, habitat patch and landscape characteristics. We investigated the inter‐specific variation in non‐forest gap‐crossing abilities of an entire tropical forest‐dependent avifauna. We then related this measure of dispersal ability to species life‐history characteristics and occupancy data from 31 variable‐sized forest patches sampled within the same fragmented forest landscape. A total of 5436 gap‐crossing movements of 231 forest‐dependent bird species were observed across ten linear forest gaps of varying widths, adjacent to large areas of undisturbed forest. Species persistence in isolated fragments was strongly linked to gap‐crossing ability. The most capable gap‐crossers were medium to large‐bodied species in the large insectivore, frugivore and granivore guilds, matching the most prevalent subset of species in small forest patches. However, some competent gap‐crossing species failed to occur in small patches, and minimum forest‐patch area requirements were more important in determining patch occupancy for these species. Narrow forest gaps (4–70 m) created by roads and power‐lines may become territory boundaries, thereby eliminating home‐range gap‐crossing movements for many forest species, but permit rarer dispersal events. Wider gaps (>70 m) may inhibit gap‐crossing behaviour for all but the most vagile species. Although patch size and quality may be the most important factors in structuring species assemblages in forest fragments, our results show that the degree of patch isolation and permeability of the surrounding matrix also explain which species can persist in forest isolates. Reducing the number and width of forest‐dividing gaps; maintaining and/or creating forest corridors and increasing matrix permeability through the creation and maintenance of ‘stepping‐stone’ structures will maximise the species retention in fragmented tropical forest landscapes.     

Host selection and responses to forest fragmentation in acorn weevils: inferences from dynamic occupancy models

Prior studies on species‐specific responses to habitat alteration have demonstrated that niche breadth is positively associated with patch occupancy rates in landscapes fragmented by agriculture. However, these studies generally have focused on vertebrates and relied upon data collected at a single point in time, neglecting dynamic processes that could alter inferences. We studied the effects of host selection and forest fragmentation on population dynamics of acorn weevils Curculio, the primary insect seed predators of oaks in North America. Detection/non‐detection data were collected from 174 red and white oaks in 19 forested fragments from 2005–2008. We used dynamic multi‐season site‐occupancy models within a Bayesian framework to explore variation in patch (tree‐level) occupancy dynamics of three species of weevils that vary in their specialization, i.e. their relative selection of red and white oak as hosts: C. pardalis (white oak specialist), C. sulcatulus (generalist) and C. proboscideus (generalist). Contrary to expectations, the specialist exhibited greater estimated rates of occupancy than generalists. However, red oak trees occupied by the white oak specialist appeared to function as sink populations maintained by frequent colonization following local extinction. Specialists also exhibited greater relative variation in occupancy and relative abundance on their host trees among years. Generalists exhibited lower local extinction and colonization rates than the specialist. Occupancy and vital rates of weevils on a host tree increased with acorn production and were significantly influenced by neighborhood forest density. Our results suggest that across much of their range in the eastern United States acorn weevils exist in fragmented, temporally dynamic landscapes, with generalists occurring on a lower proportion of usable trees but buffered by access to more suitable patches and greater patch‐specific survival. More generally, our results demonstrate that estimates of specialization derived from occupancy data may be misleading in the absence of patch‐specific information on vital rates.