Habitat patch size alters the importance of dispersal for species diversity in an experimental freshwater community

Increased dispersal of individuals among discrete habitat patches should increase the average number of species present in each local habitat patch. However, experimental studies have found variable effects of dispersal on local species richness. Priority effects, predators, and habitat heterogeneity have been proposed as mechanisms that limit the effect of dispersal on species richness. However, the size of a habitat patch could affect how dispersal regulates the number of species able to persist. We investigated whether habitat size interacted with dispersal rate to affect the number of species present in local habitats. We hypothesized that increased dispersal rates would positively affect local species richness more in small habitats than in large habitats, because rare species would be protected from demographic extinction. To test the interaction between dispersal rate and habitat size, we factorially manipulated the size of experimental ponds and dispersal rates, using a model community of freshwater zooplankton. We found that high‐dispersal rates enhanced local species richness in small experimental ponds, but had no effect in large experimental ponds. Our results suggest that there is a trade‐off between patch connectivity (a mediator of dispersal rates) and patch size, providing context for understanding the variability observed in dispersal effects among natural communities, as well as for developing conservation and management plans in an increasingly fragmented world.     

Do invasive species undergo metapopulation dynamics? A case study of the invasive Caspian gull,Larus cachinnans,in Poland

Aim The mechanisms of initial dispersal and habitat occupancy by invasive alien species are fundamental ecological problems. Most tests of metapopulation theory are performed on local population systems that are stable or in decline. In the current study we were interested in the usefulness of metapopulation theory to study patch occupancy, local colonization, extinction and the abundance of the invasive Caspian gull (Larus cachinnans) in its initial invasion stages. Location Waterbodies in Poland. Methods Characteristics of the habitat patches (waterbodies, 35 in total) occupied by breeding pairs of Caspian gulls and an equal sample of randomly selected unoccupied patches were compared with t‐tests. Based on presence–absence data from 1989 to 2006 we analysed factors affecting the probability of local colonization, extinction and the size of local populations using generalized linear models. Results Occupied habitat patches were significantly larger and less isolated (from other habitat patches and other local populations) and were located closer to rivers than empty patches. The proximity of local food resources (fish ponds, refuse dumps) positively affected the occurrence of breeding pairs. The probability of colonization was positively affected by patch area, and negatively by distances to fish ponds, nearest habitat patch, nearest breeding colony and to a river, and by higher forest cover around the patch boundaries. The probability of extinction was lower in patches with a higher number of breeding pairs and with a greater area of islets. The extinction probability increased with distances to other local populations, other habitat patches, fish ponds and to refuse dumps and with a higher cover of forest around the patch boundaries. The size of the local population decreased with distances to the nearest habitat patch, local population, river, fish pond and refuse dump. Local abundance was also positively affected by the area of islets in the patch. Main conclusions During the initial stages of the invasion of Caspian gulls in Poland the species underwent metapopulation‐like dynamics with frequent extinctions from colonized habitat patches. The results prove that metapopulation theory may be a useful conceptual framework for predicting which habitats are more vulnerable to invasion.     

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.     

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.     

Does colonization asymmetry matter in metapopulations?

Despite the considerable evidence showing that dispersal between habitat patches is often asymmetric, most of the metapopulation models assume symmetric dispersal. In this paper, we develop a Monte Carlo simulation model to quantify the effect of asymmetric dispersal on metapopulation persistence. Our results suggest that metapopulation extinctions are more likely when dispersal is asymmetric. Metapopulation viability in systems with symmetric dispersal mirrors results from a mean field approximation, where the system persists if the expected per patch colonization probability exceeds the expected per patch local extinction rate. For asymmetric cases, the mean field approximation underestimates the number of patches necessary for maintaining population persistence. If we use a model assuming symmetric dispersal when dispersal is actually asymmetric, the estimation of metapopulation persistence is wrong in more than 50% of the cases. Metapopulation viability depends on patch connectivity in symmetric systems, whereas in the asymmetric case the number of patches is more important. These results have important implications for managing spatially structured populations, when asymmetric dispersal may occur. Future metapopulation models should account for asymmetric dispersal, while empirical work is needed to quantify the patterns and the consequences of asymmetric dispersal in natural metapopulations.     

Observational evidence that maladaptive gene flow reduces patch occupancy in a wild insect metapopulation

Theory predicts that dispersal throughout metapopulations has a variety of consequences for the abundance and distribution of species. Immigration is predicted to increase abundance and habitat patch occupancy, but gene flow can have both positive and negative demographic consequences. Here, we address the eco‐evolutionary effects of dispersal in a wild metapopulation of the stick insect Timema cristinae, which exhibits variable degrees of local adaptation throughout a heterogeneous habitat patch network of two host‐plant species. To disentangle the ecological and evolutionary contributions of dispersal to habitat patch occupancy and abundance, we contrasted the effects of connectivity to populations inhabiting conspecific host plants and those inhabiting the alternate host plant. Both types of connectivity should increase patch occupancy and abundance through increased immigration and sharing of beneficial alleles through gene flow. However, connectivity to populations inhabiting the alternate host‐plant species may uniquely cause maladaptive gene flow that counters the positive demographic effects of immigration. Supporting these predictions, we find the relationship between patch occupancy and alternate‐host connectivity to be significantly smaller in slope than the relationship between patch occupancy and conspecific‐host connectivity. Our findings illustrate the ecological and evolutionary roles of dispersal in driving the distribution and abundance of species.     

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

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

A comparison of patch connectivity measures using data on invertebrates in hollow oaks

This study aimed at comparing six patch connectivity measures by fitting them to field data. We used occupancy data for eight beetle and two pseudoscorpion species from 281 hollow oaks in southeast Sweden. Species occupancy was modelled in relation to tree characteristics and one measure of patch connectivity at a time. For each connectivity measure we searched for the spatial scale that generated the best fit to field data. Connectivity measures that only include occupied patches provided better model fits than those that include all patches. When occupancy data are absent for surrounding habitat patches, information that reflects occurrence probabilities can be included in the connectivity measure. However, in this study incorporation of such information resulted in only a slight improvement of model fit. A frequently used connectivity measure based on the negative exponential function was relatively poor in explaining species’ occurrence; for eight species out of nine a buffer measure was better. A better fit was obtained when the negative exponential function was modified to take into account that habitat patches may “compete” for the immigrants. The spatial scale with the best fit tended to be larger when we used connectivity measures in which dispersal sources are identified with lower precision. Thus, the outcomes from different multiple‐scale studies are not directly comparable if the density of dispersal sources is not measured in the same way. Overall we conclude that buffer measures are useful, as they give good predictions and are easy to understand and use. If a biologically more realistic measure is needed, one that up‐weights the closest patches should be used. Finally, the possibility that habitat patches may compete with each other for immigrants should be considered when selecting a connectivity measure.     

Host–parasitoid dynamics in a fragmented landscape: Holly trees,holly leaf miners and their parasitoids

Many species inhabit fragmented landscapes, where units of resource have a patchy spatial distribution. While numerous studies have investigated how the incidence and dynamics of individual species are affected by the spatial configuration and landscape context of habitat patches, fewer studies have investigated the dynamics of multiple interacting resource and consumer species in patchy landscapes. We describe a model system for investigating host–parasitoid dynamics in a patchy landscape: a network of 166 holly trees, a specialised herbivore of holly (the leaf miner, Phytomyza ilicis (Curtis, 1948)), and its suite of parasitoids. We documented patch occupancy by P. ilicis, its density within patches, and levels of parasitism over a 6-year period, and manipulated patch occupancy by creating artificially vacant habitat patches. Essentially all patches were occupied by the herbivore in each year, suggesting that metapopulation dynamics are unlikely to occur in this system. The main determinants of densities for P. ilicis and its parasitoids were resource availability (patch size and host density, respectively). While P. ilicis is apparently not restricted by the spatial distribution of resources, densities of its parasitoids showed a weaker positive relationship with host density in more isolated patches. In patches where local extinctions were generated experimentally, P. ilicis densities and levels of parasitism recovered to pre-manipulation levels within a single generation. Furthermore, patch isolation did not significantly affect re-colonisation by hosts or parasitoids. Analysing the data at a variety of spatial scales indicates that the balance between local demography and dispersal may vary depending on the scale at which patches are defined. Taken together, our results suggest that the host and its parasitoids have dispersal abilities that exceed typical inter-patch distances. Patch dynamics are thus largely governed by dispersal rather than within-patch demography, although the role of demography is higher in larger patches.     

The importance of forest patch networks for the conservation of the Thorn-tailed Rayaditos in central Chile

Conservation of forest birds in fragmented landscapes requires not only determining the critical patch characteristics influencing local population persistence but also identifying patch networks providing connectivity and suitable habitat conditions necessary to ensure regional persistence. In this study, we assessed the importance of patch attributes, patch connectivity, and network components (i.e., groups of interconnected patches) in explaining the occupancy pattern of the Thorn-tailed Rayadito (Aphrastura spinicauda), a forest bird species of central Chile. Using a daily movement threshold distance, we identified a total of 16 network components of sclerophyllous forest within the study area. Among those components, patch area and vegetation structure-composition were important predictors of patch occupancy. However, the inclusion of patch connectivity and component size (i.e., the area of a network component) into the models greatly increases the models’ accuracy and parsimony. Using the best-fitted model, a total of 33 patches were predicted to be occupied by rayaditos within the study area, but such occupied patches were distributed in only six network components. These results suggest that persistence of rayaditos in central Chile requires the maintenance of large single patches and patch networks providing habitat and connectivity.     

Factors affecting the local abundance of two anemonefishes (Amphiprion frenatus and A. perideraion) around a semi-closed bay in Puerto Galera,the Philippines

Many marine organisms disperse or migrate among habitats, which affects their abundance patterns at individual local habitats. To clarify the factors affecting the distribution patterns of two anemonefishes (Amphiprion frenatus and A. perideraion), we measured the habitat patch size (anemone size), patch isolation (mean distance from other anemones), presence/absence of other anemonefish species, depth, and abundance of the two anemonefishes at each anemone around a semi-closed bay (up to 3.7 km) in Puerto Galera, the Philippines. We assumed that local abundance increases with habitat size and decreases with patch isolation because of greater resource availability and reduced rates of recruitment from other patches. Local abundance of A. frenatus was related to habitat size and the presence of other anemonefish species, whereas that of A. perideraion was affected by the presence of other anemonefish species and water depth. Interspecific competition and/or niche differentiation of habitat can explain the negative relationship between the local abundance of the target species and other anemonefish. Patch isolation was not significant for both species probably because the dispersal rate was not directly proportional to the geographic distance between patches at our study site.     

Process from pattern in the distribution of an endangered leaf beetle

We investigated whether signals of known dispersal processes and habitat patch turnover could be detected in a snapshot of the distribution of the tansy leaf beetle Chrysolina graminis among patches of its host plant tansy Tanacetum vulgare . Beetle occupancy in 1305 patches was analysed using autologistic generalised additive models (GAMs). These model spatial autocorrelation with an autocovariate calculated as the distance-weighted rate of occupancy among neighbouring patches. The autocovariate that best explained beetle occupancy was one which represented the active search for patches during beetle dispersal, included a distance weight that closely matched a previously fitted dispersal kernel and had neighbourhood sizes encompassing ∼95% of known dispersal distances. Autocovariates distinguishing between neighbours on the same and opposite riverbanks outperformed those that did not, revealing the river as a barrier to dispersal. Differentiating between up and downstream autocorrelation did not improve model fit, as is consistent with the beetle's lack of directional bias in dispersal. Habitat connectivity (the extent to which it was surrounded by other patches) did not appear to affect beetle occupancy in the field, while positive effects were found for distributions simulated from the GAM. We argue that this reflects a non-equilibrium distribution driven by slow responses to high rates of habitat patch turnover due to limited dispersal ability. Our findings suggest that presence/absence snapshots can reveal patterns of dispersal and be used to test whether species' ranges are at equilibrium. Such information is important for effective conservation so the possibility of inferring these patterns from distribution data is an appealing one.     

Landscape connectivity dynamics based on network analysis in the Xishuangbanna Nature Reserve,China

Lack of landscape connectivity and habitat loss is major threats to biodiversity and ecosystem integrity in nature reserves aimed at conservation. In this study, we used structural pattern and functional connectivity metrics to analyze the spatial patterns and landscape connectivity of habitat patches for the Shangyong sub-reserve of the Xishuangbanna Nature Reserve from 1970, 1990, and 2000. On the basis of vegetation and land cover data, we applied the equivalent connected area ECA(PC) indicator to analyze the changes in forest connectivity. Four distance thresholds (2, 4, 8, 12 km) were considered to compare the patch importance of connectivity by dECA values. The results showed the declining trends of landscape connectivity measured by ECA(PC) index from 1970 to 2000. The importance of connectivity in each forest patch varied with the increment of dispersal distances at the patch level, and some important habitat patches, which exhibit a potential to enhance landscape connectivity, should be given more attention. The least-cost pathways based on network structure were displayed under four dispersal distances in three periods. The results showed that the number of paths among the fragments of forest patches exhibited radical increases for larger dispersal distances. Further correlation analyses of AWF, ECA (IIC), and ECA (PC) showed the weakest and least-frequent correlations with the structural pattern indices, while H presented more significant correlations with the PD fragmentation metric. Furthermore, Kendall's rank correlations between the forest patch area and functional connectivity indicators showed that dECA (PC) and dAWF indicators should provided the area-based prioritization of habitat patches. Moreover, the low-rank correlations showed that dF and dLCP can be considered as effective and appropriate indicators for the evaluation of habitat features and network patterns.     

Characterizing the importance of habitat patches in maintaining landscape connectivity for Tibetan antelope in the Altun Mountain National Nature Reserve,China

Connectivity of habitat patches is crucial for wildlife dispersal and survival, and identifying patches with high importance for maintaining connectivity can aid effective wildlife management. Knowledge of the habitat distribution of the Tibetan antelope in the Altun Mountain National Nature Reserve, which is essential for connectivity analysis, remains limited. We analyzed potential habitat distribution and priority patches using GIS-based habitat suitability modeling with three weighting factors and evaluated the connectivity of habitat patches under four dispersal distance scenarios. Patches with high habitat suitability covered 25.39 % of the total area, and these patches were selected for connectivity analysis as resource patches. Connectivity analysis indicated that, although the overall probability of connectivity (PC) showed an upward trend with increasing dispersal distance, the importance of each patch varied considerably under different dispersal distance scenarios. Transfer analysis of patch numbers between different importance levels revealed that the number of patches becoming less important was higher than the number of patches becoming more important when dispersal distance increased. In addition, nine patches covering 38.49 % of the suitable habitat area were identified as priority patches, in particular the patches near the Kardun and the Karchuka inspection stations. We also found that the habitat distribution of the Tibetan antelope obtained from the suitability model matched the population distribution determined by a field survey. Correlation analysis between patch area and the percentage of PC index value loss (dPC) revealed that the larger patches in this region were more likely to be important for maintaining connectivity.     

Identifying potential conservation areas in the Cuitzeo Lake basin,Mexico by multitemporal analysis of landscape connectivity

Lake Cuitzeo basin is an important ecological area subjected to strong human pressure on forest covers that are key elements for the long-term support of biodiversity. We studied landscape connectivity changes for the years 1975, 1996, 2000, 2003 and 2008 to identify potential conservation areas in the basin. We modeled potential distributions of the Mexican bobcat (Lynx rufus escuinapae) and the ringtail (Bassariscus astutus) – two terrestrial mammal focal species with contrasting dispersal capacities – and we determined their habitat availability and suitability in the basin. We then identified their optimal habitat patches and produced landscape cumulative resistance maps, estimated least-cost paths (graph theory approach), and elaborated current flow maps (circuit theory approach). For evaluation of landscape connectivity, we applied an integral index of connectivity (IIC) to each study period, and determined individual habitat patch contribution to the overall landscape connectivity. The IIC index had very low values associated with reduced availability of focal species habitat. However, our study showed the conservation importance of the surface of optimal habitat patch areas. The combined application of a graph-based approach and current flow mapping were useful, and complementary both in terms of estimating potential dispersal corridors and identifying high probability dispersal areas. This indicated that landscape connectivity analysis is a useful tool for identification of potential conservation areas and for local landscape planning.     

Host plant density and patch isolation drive occupancy and abundance at a butterfly's northern range margin

Marginal populations are usually small, fragmented, and vulnerable to extinction, which makes them particularly interesting from a conservation point of view. They are also the starting point of range shifts that result from climate change, through a process involving colonization of newly suitable sites at the cool margin of species distributions. Hence, understanding the processes that drive demography and distribution at high‐latitude populations is essential to forecast the response of species to global changes. We investigated the relative importance of solar irradiance (as a proxy for microclimate), habitat quality, and connectivity on occupancy, abundance, and population stability at the northern range margin of the Oberthür's grizzled skipper butterfly Pyrgus armoricanus. For this purpose, butterfly abundance was surveyed in a habitat network consisting of 50 habitat patches over 12 years. We found that occupancy and abundance (average and variability) were mostly influenced by the density of host plants and the spatial isolation of patches, while solar irradiance and grazing frequency had only an effect on patch occupancy. Knowing that the distribution of host plants extends further north, we hypothesize that the actual variable limiting the northern distribution of P. armoricanus might be its dispersal capacity that prevents it from reaching more northern habitat patches. The persistence of this metapopulation in the face of global changes will thus be fundamentally linked to the maintenance of an efficient network of habitats.     

The larval ecology of the butterfly Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant quantity and quality as main predictors of habitat quality

Corresponding to theory, the persistence of metapopulations in fragmented landscapes depends on the area of suitable habitat patches and their degree of isolation, mediating the individual exchange between habitats. More recently, habitat quality has been highlighted as being equally important. We therefore assess the role of habitat area, isolation and quality for the occupancy of larval stages of the regionally threatened butterfly Euphydryas desfontainii occurring in grassland habitats comprising the host plant Dipsascus comosus. We put a special focus on habitat quality which was determined on two spatial scales: the landscape (among patches) and the within-patch level. On the landscape level, occupancy of caterpillars was determined by a presence-absence analysis at 28 host plant patches. On the within-patch level, oviposition site selection was studied by comparing 159 host plants with egg clutches to a random sample of 253 unoccupied host plants within six habitat patches. The occupancy of caterpillars and presence of egg clutches on host plants was then related to several predictors such as patch size and isolation on the landscape level and host plant characteristics and immediate surroundings on the within patch level. On the landscape level, only host plant abundance was related to the presence of caterpillars, while size and isolation did not differ between occupied and unoccupied patches. However, the weak discrimination of larval stages among patches changed on the within-patch level: here, several microclimatic predictors such as sunshine hours and topography, host plant morphology and phenology as well as further potential host plants in the immediate surroundings of the plant chosen for oviposition strongly determined the presence of egg clutches. We strongly suggest promoting the presence of the host plant in topographically and structurally rich habitat patches to offer potential for microclimatic compensation for a species considered threatened by climate change.     

Hygrochastic capsule dehiscence supports safe site strategies in New Zealand alpine Veronica (Plantaginaceae)

Background and Aims

Hygrochasy is a capsule-opening mechanism predominantly associated with plants in arid habitats, where it facilitates spatially and temporally restricted dispersal. Recently, hygrochastic capsules were described in detail for the first time in alpine Veronica in New Zealand. The aim of the present study was to investigate whether hygrochastic capsules are an adaptation of alpine Veronica to achieve directed dispersal to safe sites. We expect that by limiting dispersal to rainfall events, distances travelled by seeds are short and confine them to small habitat patches where both seedlings and adults have a greater chance of survival.

Methods

Dispersal distances of five hygrochastic Veronica were measured under laboratory and field conditions and the seed shadow was analysed. Habitat patch size of hygrochastic Veronica and related non-hygrochastic species were estimated and compared.

Key Results

Dispersal distances achieved by dispersal with raindrops did not exceed 1 m but weather conditions could influence the even distribution of seeds around the parent plant. Compared with related Veronica species, hygrochastic Veronica mostly grow in small, restricted habitat patches surrounded by distinctly different habitats. These habitat patches provide safe sites for seeds due to their microtopography and occurrence of adult cushion plants. Non-hygrochastic Veronica can be predominantly found in large habitats without clearly defined borders and can be spread over long distances along rivers.

Conclusions

The results suggest that hygrochasy is a very effective mechanism of restricting seed dispersal to rainfall events and ensuring short-distance dispersal within a small habitat patch. It appears that it is an adaptation for directed dispersal to safe sites that only exist within the parent habitat.     

Population turnover and habitat dynamics in Notonecta (Hemiptera: Notonectidae) metapopulations

Simple metapopulation models assume that local populations occur in patches of uniform quality habitat separated by non-habitat. However field metapopulations tend to show considerable spatial and temporal variation in patch quality, and hence probability of occupancy. This may have implications for the adequacy of simple metapopulation models in describing and predicting regional population dynamics of natural systems. This study investigated the effects of habitat characteristics on landscape-scale occupancy dynamics of two species of backswimmer (Notonecta, Hemiptera: Notonectidae) in small freshwater ponds. The results demonstrated clear links between habitat, pond occupancy and population turnover, particularly local extinction. There were considerable changes in the habitat of individual ponds between years, but local changes were not spatially correlated and the frequency distribution of habitat conditions at the landscape level remained similar in different years. Stable occupancy levels of Notonecta species appears to result from a balance of the rates of creation and loss of suitable habitat due to spatially uncorrelated habitat change. Systems such as this, where turnover is driven by habitat dynamics, demonstrate the potential value of incorporating the dynamics of habitat change into metapopulation models. Such developments are likely to improve predictions of landscape-scale occupancy dynamics, whilst also allowing patch-level predictions of occupancy, based on local habitat conditions. Received: 18 August 1999 / Accepted: 3 December 1999     

Heterogeneity in patch quality buffers metapopulations from pathogen impacts

Many wildlife species persist on a network of ephemerally occupied habitat patches connected by dispersal. Provisioning of food and other resources for conservation management or recreation is frequently used to improve local habitat quality and attract wildlife. Resource improvement can also facilitate local pathogen transmission, but the landscape-level consequences of provisioning for pathogen spread and habitat occupancy are poorly understood. Here, we develop a simple metapopulation model to investigate how heterogeneity in patch quality resulting from resource improvement influences long-term metapopulation occupancy in the presence of a virulent pathogen. We derive expressions for equilibrium host–pathogen outcomes in terms of provisioning effects on individual patches (through decreased patch extinction rates) and at the landscape level (the fraction of high-quality, provisioned patches), and highlight two cases of practical concern. First, if occupancy in the unprovisioned metapopulation is sufficiently low, a local maximum in occupancy occurs for mixtures of high- and low-quality patches, such that further increasing the number of high-quality patches both lowers occupancy and allows pathogen invasion. Second, if the pathogen persists in the unprovisioned metapopulation, further provisioning can result in all patches becoming infected and in a global minimum in occupancy. This work highlights the need for more empirical research on landscape-level impacts of local resource provisioning on pathogen dynamics.