Incidence and population dynamics of the leaf beetle Gonioctena olivacea in dynamic habitats

In natural as well as in cultural landscapes, disturbance and succession are responsible for the emergence and subsequent disappearance of suitable habitat patches. The dynamics of habitat patches has important consequences for the spatial structure and dynamics of regional populations. However, there are only few studies quantifying both patch dynamics and incidence of insect species in a dynamic landscape over several years. I studied the incidence and population dynamics of the leaf beetle Gonioctena olivacea in a system of dynamic patches of the host plant Scotch broom Cytisus scoparius . The incidence of the beetle was most strongly affected by patch area, whereas connectivity, patch quality, patch age, and landscape context had no or only a minor effect when analysed with logistic regression. The size of local beetle populations was highly fluctuating between the years; however, the population dynamics of the local populations was not synchronous. Adjacent patches did not show higher degrees of synchrony than patches separated by large distances. In the three years of study, local populations became extinct through demographic or environmental stochasticity and patch destruction. Each year >10% of the patches disappeared. The extinction rate of beetles in persistent patches was decreasing with increasing patch area. On the other hand, patches newly emerged and were rapidly colonized by the beetle. The colonization rate depended on patch connectivity. Obviously, Gonioctena olivacea was capable of persisting in this system with high turnover of patches owing to its high dispersal power.     

Experimental beetle metapopulations respond positively to dynamic landscapes and reduced connectivity

Interactive effects of multiple environmental factors on metapopulation dynamics have received scant attention. We designed a laboratory study to test hypotheses regarding interactive effects of factors affecting the metapopulation dynamics of red flour beetle, Tribolium castaneum. Within a four-patch landscape we modified resource level (constant and diminishing), patch connectivity (high and low) and patch configuration (static and dynamic) to conduct a 2(3) factorial experiment, consisting of 8 metapopulations, each with 3 replicates. For comparison, two control populations consisting of isolated and static subpopulations were provided with resources at constant or diminishing levels. Longitudinal data from 22 tri-weekly counts of beetle abundance were analyzed using bayesian Poisson generalized linear mixed models to estimate additive and interactive effects of factors affecting abundance. Constant resource levels, low connectivity and dynamic patches yielded greater levels of adult beetle abundance. For a given resource level, frequency of colonization exceeded extinction in landscapes with dynamic patches when connectivity was low, thereby promoting greater patch occupancy. Negative density dependence of pupae on adults occurred and was stronger in landscapes with low connectivity and constant resources; these metapopulations also demonstrated greatest stability. Metapopulations in control landscapes went extinct quickly, denoting lower persistence than comparable landscapes with low connectivity. When landscape carrying capacity was constant, habitat destruction coupled with low connectivity created asynchronous local dynamics and refugia within which cannibalism of pupae was reduced. Increasing connectivity may be counter-productive and habitat destruction/recreation may be beneficial to species in some contexts.     

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.     

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.     

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.     

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.     

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.     

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.     

Metapopulation dynamics of a beetle species confined to burned forest sites in a managed forest region

Despite increasing awareness of the theoretical importance of habitat dynamics on metapopulations, only a few empirical studies have been conducted. We aimed to increase our understanding of how patch size, dynamics and connectivity affect colonization–extinction dynamics and the occurrence patterns of a beetle (Stephanopachys linearis), which breeds only in burned trees, existing as dynamic habitat patches that have become rare in managed forest landscapes. We assessed species’ presence/absence twice in all known habitat patches (i.e. > 1 ha sites where forest fires had occurred during the previous 2–15 yr) in a 200 × 150 km region of central Sweden, dominated by managed boreal forest. Evaluated over six years, the colonization rate was 47% and the local extinction risk was 65%. Probability of colonization increased with patch size (number of suitable trees in a site) and connectivity to occupied patches within 30 km, and decreased with increasing time since fire. Local extinction risk decreased with habitat patch size but increased, unexpectedly, with connectivity. Occurrence increased with patch size and decreased with increasing time since fire. At a regional scale, S. linearis tracks the fire dynamics by colonising sites with burned trees and by becoming extinct at rates which make the species rare at sites where burnt trees are more than eight years old. In managed boreal forest landscapes, a large proportion of sites may be created by prescribed burning (in our study area: 82%), and consequently human decisions strongly affect the future amount of habitat for fire‐dependent species and its spatial distribution. Stephanopachys linearis uses burned sites more often if more trees are retained and, to some extent, if sites are concentrated in those parts of a region that already support high population densities of the species.     

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     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

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.     

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.     

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.     

Patch occupancy of two hemipterans sharing a common host plant

Aim Two hemipteran species were chosen as a study system for the comparative analysis of patch occupancy and spatial population structure of insects sharing a common host plant. This study tested whether (1) the incidence in the host plant patches differed between the two species, and (2) the two species exhibited a different spatial population structure, i.e. were they affected differentially by isolation and area of the host plant patches. Location The porphyry landscape north of Halle (Saale) in Germany comprising 506 patches of the host plant Brachypodium pinnatum. Methods The host plant patches were surveyed for the two hemipterans. To assess the influence of patch quality on species occurrence the patches were characterized by mean cover abundance of B. pinnatum, type of subsoil, slope, exposure, and shading. The spatial configuration of the patches was considered by patch area and isolation. The influence of the habitat factors and the spatial configuration on the occupancy of the two species was analysed by logistic regression. Results Adarrus multinotatus was found in 441 patches, while Neophilaenus albipennis was found in only 90 patches. While A. multinotatus showed virtually no relationship to the habitat factors, the occupancy of N. albipennis was influenced by subsoil type, cover abundance, and shading. The effects of area and isolation on occupancy of the patches also differed between the two species. The occupancy of N. albipennis was determined largely by area and isolation, whereas in A. multinotatus no considerable effect of spatial configuration was found. Main conclusions The study revealed a marked difference between the two hemipteran species in respect of spatial population structure. Adarrus multinotatus built up a ‘patchy population’, whereas N. albipennis showed a ‘metapopulation’ structure within the same set of patches in the same landscape. Spatial population structure was found to be not only a function of spatial configuration of habitat patches, but population structure differed between the habitat generalist A. multinotatus and the habitat specialist N. albipennis.     

Quantitative predictions for patch occupancy of capercaillie in fragmented habitats

A major conclusion of studying metapopulation biology is that species conservation should favor regional rather than local population persistence. Regional persistence is tightly linked to size, spatial configuration and quality of habitat patches. Hence it is important for the management of endangered species that priority patches can be identified. We developed a predictive model of patch occupancy by capercaillie, a threatened grouse species, based on a single snapshot of data. We used logistic regression to predict patch occupancy as a function of patch size, isolation, connectivity, relative altitude, and biogeographical area. The probability of a patch being occupied increased with patch size and increasing altitude, and decreased with increasing distance to the next occupied patch. Patch size was the most important predictor although occupied patches varied considerably in size. Our model only uses data on the number, size and spatial configuration of habitat patches. It is a useful tool to designate priority areas for conservation, i.e. large core patches with high resilience in habitat quality, smaller island‐patches that still have high probability of being inhabited or becoming recolonised, and patches functioning as “stepping stones”. If capercaillie is to be preserved, habitat suitability needs to be maintained in a functional network of patches that account for size and inter‐patch distance thresholds as found in this study. We suggest that similar area‐isolation relationships are valid for almost any region within the distribution range of capercaillie. The thresholds for occupancy are however likely to depend on characteristics of the respective landscape. The outcome of our study emphasises the need for future investigations that explore the relationship between patch occupancy, matrix quality and its resistance to dispersing individuals.     

Connectivity compensates for low habitat quality and small patch size in the butterfly <Emphasis Type="Italic">Cupido minimus</Emphasis>

Habitat size, habitat isolation and habitat quality are regarded as the main determinants of butterfly occurrence in fragmented landscapes. To analyze the relationship between the occurrence of the butterfly Cupido minimus and these factors, patch occupancy of the immature stages in patches of its host plant Anthyllis vulneraria was studied in the nature reserve Hohe Wann in Bavaria (Germany). In 2001 and 2002, 82 A. vulneraria patches were surveyed for the presence of C. minimus larvae. The occurrence was largely affected by the size of the food plant patches. In a habitat model that uses multiple logistic regression, the type of management and habitat connectivity are further determinants of species distribution. Internal and temporal validation demonstrate the stability and robustness of the developed habitat models. Additionally, it was proved that the colonization rate of C. minimus was significantly influenced by the distance to the next occupied Anthyllis patch. Concerning long-term survival of (meta-) populations in fragmented landscapes, the results show that lower habitat quality may be compensated by higher connectivity between host plant patches. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

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.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.