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

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

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.     

Determinants of distribution and abundance in the clouded apollo butterfly: a landscape ecological approach

Recent studies on the determinants of distribution and abundance of animals at landscape level have emphasized the usefulness of the metapopulation approach, in which patch area and habitat connectivity have often proved to explain satisfactorily existing patch occupancy patterns. A different approach is needed to study the common situation in which suitable habitat is difficult to determine or does not occur in well‐defined habitat patches. We applied a landscape ecological approach to study the determinants of distribution and abundance of the threatened clouded apollo Parnassius mnemosyne butterfly within an area of 6 km² of agricultural landscape in south‐western Finland. The relative role of 24 environmental variables potentially affecting the distribution and abundance of the butterfly was studied using a spatial grid system with 2408 grid squares of 0.25 ha, of which 349 were occupied by the clouded apollo. Both the probability of butterfly presence and abundance in a 0.25 ha square increased with the presence of the larval host plant Corydalis solida the cover of semi‐natural grassland, the amount of solar radiation and spalial autocorrelation in butterfly occurrence. Additionally, butterfly abundance increased with overall mean patch size and decreased with maximum slope angle and wind speed. Two advantages of the employment of a spatial grid system included the avoidance of a subjective definition of suitable habitat patches and an evaluation of the relative significance of different components of habitat quality at the same time with habitat availability and connectivity. The large variation in habitat quality was influenced by the abundance of the larval host plant and adult nectar sources but also by climatological. topographical and structural factors. The application of a spatial grid system as used here has potential for a wide use in studies on landscape‐level distribution and abundance patterns in species with complex habitat requirements and habitat availability patterns.     

Temporal patch occupancy dynamics of the Siberian flying squirrel in a boreal forest landscape

Ability to predict species distribution in a landscape is of crucial importance for natural resource management and species conservation. Therefore, the understanding of species habitat requirements and spatio-temporal dynamics in occurrence is needed. We examined patch occupancy patterns of the Siberian flying squirrel Pteromys volans in northern Finland across a seven year study period. Forest patches dominated by mature spruce ( Picea abies ) in a study area (375 km²) were surveyed to monitor the presence or absence of the flying squirrel. The patch occupancy pattern was dynamic: about half of the habitat patches were occupied at least once during the study period and more patches were colonised than were abandoned. Patches that were continuously occupied (i.e. occupied during all sample periods) were typically of high quality (based on habitat and landscape characteristics), continuously unoccupied patches were usually of low quality, and intermediate quality patches were occupied intermittently. The variables explaining patch occupancy were similar each year, and a statistical model based on data from the year 2000 also predicted occupancy in 2004 with similar accuracy. However, data from a single survey were inadequate for identifying patches used intermittently by flying squirrels. Despite inconsistent occupancy, these patches may be important for the local persistence of flying squirrels. The dynamic occupancy pattern may thus affect estimates of suitable habitat area and identification of functional patch networks for landscape planning. These results emphasise the need for follow-up studies to better understand population patterns and processes in time.     

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.     

Monitoring Golden-Cheeked Warblers on Private Lands in Texas

ABSTRACT A majority of North American breeding habitat for neotropical migrants exists on private lands, requiring monitoring strategies focused on habitat in these private holdings. We outline study designs and protocols using repeated Presence-Absence surveys across a gradient of patch sizes to develop a range-wide monitoring program for the endangered golden-cheeked warbler (Dendroica chrysoparia) in Texas, USA. We surveyed 200–400 point-count locations across approximately 30 private properties annually from 2005 to 2008. We used data from our surveyed patches (n = 147) and the Ψ (occupancy), p (detection), and γ = 1 - ɛ parameterization to estimate patch dynamics and associated detection probabilities for golden-cheeked warblers. Patch size had a strong association with patch occupancy, and all patches >160 ha were predicted to be occupied. We found no evidence that large golden-cheeked warbler populations located on public lands in the vicinity of our study area influenced occupancy dynamics. We conducted simulations across a range of detection probabilities to evaluate potential sample sizes for both standard- and removal-based occupancy modeling. Simulations using parameter estimates from our analysis indicated that removal-based sampling is superior to standard sampling. Based on our results, surveying golden-cheeked warbler presence in oak-juniper (Quercus-Juniperus) patches under a removal modeling framework should be considered as one alternative for range-wide monitoring programs because patch-level monitoring would be necessary to estimate proportion of range occupied. Large contiguous patches are rare across the species’ range; hence, conservation and management of the mosaic of smaller patches within a landscape context would be required for maintaining species viability. Thus, we recommend the identification of areas where smaller, contiguous patches represent a significant portion of the available habitat within the local landscape and targeting these areas for habitat maintenance and improvement.     

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

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

Host-plant patch qualities and presence of a likely competitor species affect the distribution and abundance of a rare British moth, <Emphasis Type="Italic">Cucullia lychnitis</Emphasis>

Understanding the environmental factors that affect the distribution and abundance of rare and threatened species can help identify priority sites for conservation action such as habitat management. The moth Cucullia lychnitis is identified in UK legislation as a priority species for conservation. Its conspicuous larvae feed on a patchily distributed host plant Verbascum nigrum, but are entirely absent from some host plant patches within the species’ range. Environmental variability among patches was investigated to understand factors affecting (i) patch occupancy by C. lychnitis and (ii) abundance of C. lychnitis, where present. Occupancy of individual V. nigrum plants within occupied patches was also investigated. The likelihood of patch occupancy increased with patch size, and decreased with patch isolation. Abundance of C. lychnitis was negatively correlated with patch isolation and with abundance of the weevil Cionus nigritarsis, which exploits the same host plant and may be a competitor. Within occupied patches, larvae of C. lychnitis were significantly more likely to be found on taller plants with more flower spikes; such plants are typical of established rather than newly-created patches. These results may help to guide efforts to conserve C. lychnitis, highlighting the importance of habitat connectivity and indicating potential management actions to promote features positively associated with larval incidence and abundance.     

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.     

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.     

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.     

Landscape context outweighs local habitat quality in its effects on herbivore dispersal and distribution

Past studies with spatially structured herbivore populations have emphasized the primacy of intrinsic factors (e.g., patch quality), patch geometry (e.g., patch size and isolation), and more recently landscape context (e.g., matrix composition) in affecting local population abundance and dispersal rate. However, few studies have examined the relative importance of each factor, or how they might interact to affect herbivore abundance or dispersal. Here, we performed a factorial field experiment to examine the independent and interactive effects of patch quality (plant biomass, leaf protein, leaf phenolics) and matrix composition [mudflat or non-host grass (Bromus inermis)] on planthopper (Prokelisia crocea) emigration from host-plant patches (prairie cordgrass, Spartina pectinata). In addition, a field survey was conducted to examine the relative importance of patch quality, geography, and matrix composition on planthopper occupancy and density. In the experiment, we found that rates of emigration from low and intermediate quality patches were, on average, 21% percent higher for patches embedded in brome than mudflat. In contrast, the emigration rate was unaffected by matrix composition in nutrient-rich patches. Within matrix types, plant quality had little effect on emigration. In the survey, planthopper density and the patch occupancy rate of planthoppers increased nonadditively with increasing patch size and the percentage of the surrounding matrix composed of mudflat. This study suggests that landscape-level factors, such as the matrix, may be more important than factors intrinsic to the patches.     

Effects of patch characteristics and isolation on relative abundance of the scarce heath butterfly <Emphasis Type="Italic">Coenonympha hero</Emphasis> (Nymphalidae)

The scarce heath (Coenonympha hero) is an internationally threatened butterfly in Western Europe, where it occurs primarily on hay fields and abandoned arable land in a small-scale agricultural landscape of south-central Scandinavia. Due to afforestation, this habitat is becoming increasingly fragmented in Sweden, and it can be expected that the scarce heath will decline abruptly when threshold conditions for metapopulation persistence are no longer met. We used stepwise polychotomous logistic regression to compare habitat characteristics and isolation measures for patches that harbour large, small or no populations, respectively, in an area of south-western Sweden. We found that patch area, distance to the nearest large population and amount of Galium spp. explained a significant part of the variation in relative abundance among patches. Distance to nearest large population resulted in a better model to predict occupancy than both distance to the nearest inhabited patch and connectivity, which suggests that primarily large populations act as sources for small satellite populations. Today, sites of three of the eight larger populations in the study area have been planted with spruce or pine and will disappear within 20 years. We argue that the disappearance of these patches may very well lead to rapid extinction of the whole metapopulation system.     

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.     

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.     

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.     

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     

Bayesian state-space modeling of metapopulation dynamics in the Glanville fritillary butterfly

The complexity of mathematical models of ecological dynamics varies greatly, and it is often difficult to judge what would be the optimal level of complexity in a particular case. Here we compare the parameter estimates, model fits, and predictive abilities of two models of metapopulation dynamics: a detailed individual-based model (IBM) and a population-based stochastic patch occupancy model (SPOM) derived from the IBM. The two models were fitted to a 17-year time series of data for the Glanville fritillary butterfly (Melitaea cinxia) inhabiting a network of 72 small meadows. The data consisted of biannual counts of larval groups (IBM) and the annual presence or absence of local populations (SPOM). The models were fitted using a Bayesian state-space approach with a hierarchical random effect structure to account for observational, demographic, and environmental stochasticities. The detection probability of larval groups (IBM) and the probability of false zeros of local populations (SPOM) in the observation models were simultaneously estimated from the time-series data and independent control data. Prior distributions for dispersal parameters were obtained from a separate analysis of mark–recapture data. Both models fitted the data about equally, but the results were more precise for the IBM than for the SPOM. The two models yielded similar estimates for a random effect parameter describing habitat quality in each patch, which were correlated with independent empirical measures of habitat quality. The modeling results showed that variation in habitat quality influenced patch occupancy more through the effects on movement behavior at patch edges than on carrying capacity, whereas the latter influenced the mean population size in occupied patches. The IBM and the SPOM explained 63% and 45%, respectively, of the observed variation in the fraction of occupied habitat area among 75 independent patch networks not used in parameter estimation. We conclude that, while carefully constructed, detailed models can have better predictive ability than simple models, this advantage comes with the cost of greatly increased data requirements and computational challenges. Our results illustrate how complex models can be helpful in facilitating the construction of effective simpler models.     

Consequences of insular population structure: Distribution and extinction of spruce grouse populations

Summary The population structure of the spruce grouse (Canachites canadensis) was studied in the Adirondack Mountains of New York, U.S.A. Twenty-five isolated habitat patches exist and are occupied by spruce grouse, with 7 suitable but unoccupied patches existing at the periphery of the range. The regional distribution and abundance of spruce grouse is correlated with the amount of lowland coniferous forest habitat. Unoccupied patches were significantly smaller and significantly farther from occupied patches than were other occupied patches. For all patches, as distance from the nearest occupied patch increased, the percent of occupied patches decreased linearly. I incorporated birth and death rates for spruce grouse into the MacArthur-Wilson survivorship model which closely predicted the proportion of occupied patches for an average population density (2.8 spruce grouse/100ha). For the same demographic parameters, extinction times were calculated which indicate that the 15 habitat patches of a carrying capacity of 3 female spruce grouse (100 ha) would have an average extinction time of less than 6 years. This in part accounts for the high proportion of these patches which are unoccupied. Extinctions and recolonizations of patches were observed during the study. The patterns of patch occupancy can partially be predicted based on their size, spatial arrangement, and the demographic characteristics of the spruce grouse.     

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.