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.     

Habitat quality and population density drive occupancy dynamics of snowshoe hare in variegated landscapes

The influence of habitat quality and population density on occupancy dynamics may surpass that of traditional metrics of area and isolation, but often this is not considered explicitly in studies of spatially structured populations. In landscapes that are not easily characterized as binary habitat/non‐habitat (e.g. variegated landscapes), this influence may be even more important and occur at both local and landscape levels. It follows that occupancy dynamics may be driven by disparate processes depending on how extinction or colonization relate to habitat quality and population density. We examined the relative influence of area, structural isolation, habitat quality, local population density, and neighborhood population density (i.e. population density in the landscape around a site) on the probability of extinction and colonization of snowshoe hare Lepus americanus across an expansive forest mosaic landscape (encompassing the northern third of Idaho). Habitat quality and population density were highly influential in determining extinction and colonization, whereas patch area and isolation were much less important. Sites with heavier vegetative cover at the site or landscape‐level were more likely to be colonized and less likely to go extinct, and sites with greater local population density in the previous time step had lower probability of extinction. Sites embedded in high density neighborhoods also were less likely to go extinct, but not more likely to be colonized. We found a significant interaction between local and neighborhood population density on extinction in 1 yr, suggesting that the strength of demographic rescue may vary dependent on local site densities. Our results add to a growing literature showing that factors outside of structural metrics of area and isolation are important drivers of occupancy dynamics. Given the multi‐scaled influence of habitat quality and population density on occupancy dynamics, our work also indicates that research on snowshoe hare must extend beyond simply assessing local factors to understand the spatial dynamics of populations.     

Seasonal Differences in Extinction and Colonization Drive Occupancy Dynamics of an Imperilled Amphibian

A detailed understanding of the population dynamics of many amphibian species is lacking despite concerns about declining amphibian biodiversity and abundance. This paper explores temporal patterns of occupancy and underlying extinction and colonization dynamics in a regionally imperiled amphibian species, the Northern leopard frog (Lithobates pipiens) in Alberta. Our study contributes to elucidating regional occupancy dynamics at northern latitudes, where climate extremes likely have a profound effect on seasonal occupancy. The primary advantage of our study is its wide geographic scale (60,000 km²) and the use of repeat visual surveys each spring and summer from 2009–2013. We find that occupancy varied more dramatically between seasons than years, with low spring and higher summer occupancy. Between spring and summer, colonization was high and extinction low; inversely, colonization was low and extinction high over the winter. The dynamics of extinction and colonization are complex, making conservation management challenging. Our results reveal that Northern leopard frog occupancy was constant over the last five years and thus there is no evidence of decline or recovery within our study area. Changes to equilibrium occupancy are most sensitive to increasing colonization in the spring or declining extinction in the summer. Therefore, conservation and management efforts should target actions that are likely to increase spring colonization; this could be achieved through translocations or improving the quality or access to breeding habitat. Because summer occupancy is already high, it may be difficult to improve further. Nevertheless, summer extinction could be reduced by predator control, increasing water quality or hydroperiod of wetlands, or increasing the quality or quantity of summer habitat.     

Reproductive habitat provisioning promotes survival and reproduction of the endangered endemic damselfly <Emphasis Type="Italic">Calopteryx exul</Emphasis>

Effective habitat management is predicted to have positive effects on populations and species of conservation concern. Although studies have shown that ecological processes such as colonization can be promoted after habitat management, we still need more information on the survival and reproductive consequences at the individual level in order to reach positive conservation outcome. Here we assess the effects of reproductive habitat supplementation (host oviposition plant) on survival and mating success of an endangered endemic damselfly, Calopteryx exul, using capture-mark-recapture data. We first determined that the species prefer to oviposit on floating leaves of Potamogeton spp. Based on Cormack-Jolly-Seber modeling, we found that recapture and survival probabilities were positively affected by the number of the host oviposition patches of the host plant. Moreover, we showed a strong positive relationship between adult lifespan and lifetime mating success. Our results suggest that host-plant provisioning for reproduction not only increases the survival of individuals, but also increases the number of matings per lifetime. The procedure of supplying reproductive sites may enhance population growth of threatened odonates and other aquatic insects.     

Evidence-based assessment of butterfly habitat restoration to enhance management practices

The decline in distribution and abundance of biodiversity requires evidence-based guidelines for cost-effective conservation management and systematic quantitative assessments of its effects. We investigated the efficiency of a habitat restoration programme aimed at reducing the risk of extinction of the Iolas blue Iolana iolas (Ochsenheimer, 1816), one of the rarest butterflies of Central Europe. Using occupancy and capture-mark-recapture (CMR) models accounting for probability of detection, we assessed habitat patch occupancy, habitat selection, demography and dispersal with the aim of testing and refining restoration measures. Count surveys performed at 38 plantations dedicated to the species’ unique host plant resulted in an occupancy rate of 50 %, with mostly very low relative abundance indices. The site-occupancy habitat analysis demonstrated that species abundance was best explained by host plant vitality, habitat patch connectivity, and solar radiation. CMR surveys yielded very high catchability (82 %), individual detectability (86 %) rates and limited dispersal capacity. These results form the basis for future efficient count surveys to assess species distribution and abundance. They also provide evidence-based recommendations for improving ongoing habitat restoration: (i) the attractiveness of host plant plantations must be enhanced by promoting mass blossoming, which can be achieved through systematic autumn pruning of the extant plantations; (ii) new plantations should be created in order to fill in the gaps in the landscape matrix, to increase meta-population capacity through improved habitat connectivity. Finally, this study demonstrates the relevance of efficiency tests as an integral, adaptive phase of any conservation research activity.     

高原鼠兔在不同植被覆盖的立地占据、局部灭绝和建群率研究

青藏高原的草地生态系统被野生动物和家畜广泛地利用。大型有蹄类动物的放牧和小型哺乳类动物的挖掘行为,会改变其栖息地的植物群落。基于拉萨市墨竹工卡县日多镇年村牦牛(Bos grunniens)放牧的实践,我们发现高原鼠兔(Ochotona curzoniae)立地占域率、建群和区域灭绝率与栖息地变量之间存在一定的格局关系。为了了解高原鼠兔立地占域率、建群和区域灭绝率与栖息地之间的关系,我们采用多季节性立地占据模型,利用Presence软件对相关数据进行分析。结果表明,高原鼠兔局部灭绝率与植被覆盖度之间存在正相关性,而与建群率之间则呈负相关性。     

Evaluating the local habitat history deepens the understanding of the extinction debt for endangered plant species in semi-natural grasslands

Habitat losses occur non-randomly within human-modified landscapes, resulting in high spatial heterogeneity of local habitat histories. Although local habitat history can modulate the existence of extinction debt (i.e., the number of populations predicted to become extinct) in a landscape, its role in detecting extinction debt has not been examined explicitly. We aimed to compare the detectability of extinction debt among populations of an endangered semi-natural grassland species, Echinops setifer (Compositae), in the grassland landscape of Mt. Aso, Japan. We classified populations into three groups that differed in local habitat history: stable (habitat loss ≤30% since the 1930s), moderately decreased (30% < loss ≤ 90%), and severely decreased (loss >90%). We then evaluated whether the effects of habitat areas during the 1930s and 2000s varied among groups to explain population size by GLMMs and estimated coefficient of explanatory variable by Bayesian MCMC methods. Within the groups, stable group showed significant positive relationships with both past and current habitat areas. The moderately decreased group only showed significant positive relationships with past habitat areas, indicating the existence of extinction debt in these populations. The severely decreased group only showed significant positive relationships with current habitat areas, indicating that they may have already paid their extinction debt because the rate of grassland loss exceeded the extinction threshold. Even within the same landscape, extinction debt varied in response to local habitat history. In spatially heterogeneous landscapes, evaluation of effects of local habitat history can elucidate the habitat-based extinction risks for plant populations.     

Occupancy, colonization and extinction patterns of rabbit populations: implications for Iberian lynx conservation

In Mediterranean ecosystems, rabbits are a key prey species for many predators, such as the Iberian lynx, which is threatened with extinction and has gone extinct locally in several regions of its historical distribution range. One of these regions is Serra da Malcata Nature Reserve, Portugal, which is also currently proposed as a potential site for reintroduction. We intended to investigate annual variation, potential time trends and the effects of management practices on the rabbit population in Serra da Malcata as a model for future potential reintroduction areas. The rabbit population was monitored over 12?years (from 1997 to 2009) by counting latrines along linear transects. These data were used to estimate rabbit occupancy, colonization and extinction patterns using a likelihood-based method including habitat, population and topographic covariate effects. Our results suggest that initial occupancy, when management practices were absent, was driven by the presence of Erica spp. and Cistus ladanifer shrubs and by distance to summits. Site colonization was positively influenced by the presence of edges between shrubs and pastureland and by patterns of rabbit distribution in the previous sampling season. On the other hand, local extinction was negatively influenced by edges. We conclude that the increase in rabbit occupancy and local colonization patterns was clearly associated with management actions (particularly, the creation of pasturelands), although the recovery of the species was noticeably limited by previous patterns of spatial distribution.     

Multilevel landscape utilization of the Siberian flying squirrel: Scale effects on species habitat use

Animals use and select habitat at multiple hierarchical levels and at different spatial scales within each level. Still, there is little knowledge on the scale effects at different spatial levels of species occupancy patterns. The objective of this study was to examine nonlinear effects and optimal‐scale landscape characteristics that affect occupancy of the Siberian flying squirrel, Pteromys volans, in South‐ and Mid‐Finland. We used presence–absence data (n = 10,032 plots of 9 ha) and novel approach to separate the effects on site‐, landscape‐, and regional‐level occupancy patterns. Our main results were: landscape variables predicted the placement of population patches at least twice as well as they predicted the occupancy of particular sites; the clear optimal value of preferred habitat cover for species landscape‐level abundance is a surprisingly low value (10% within a 4 km buffer); landscape metrics exert different effects on species occupancy and abundance in high versus low population density regions of our study area. We conclude that knowledge of regional variation in landscape utilization will be essential for successful conservation of the species. The results also support the view that large‐scale landscape variables have high predictive power in explaining species abundance. Our study demonstrates the complex response of species occurrence at different levels of population configuration on landscape structure. The study also highlights the need for data in large spatial scale to increase the precision of biodiversity mapping and prediction of future trends.     

Prey and tigers on the forgotten trail: high prey occupancy and tiger habitat use reveal the importance of the understudied Churia habitat of Nepal

Tigers are globally threatened and their conservation relies on intact habitat that supports key large prey. The Churia habitat is relatively unknown even though it occupies a significant portion of the forested landscape of the Terai Arc, which stretches over 1000 km in a narrow band across Nepal and India, parallel to the Himalayas. To address this lack of detailed information relevant to tiger conservation, we used sign surveys to estimate occupancy probability for 5 focal prey species of tigers (gaur, sambar, chital, wild pig, and barking deer), and assess tiger habitat use within 537 km² of the understudied Churia habitat in Chitwan National Park (CNP), Nepal. Multi-season occupancy models allowed us to make seasonal (winter vs. summer) inferences regarding changes in occupancy or habitat use based on covariates influencing occupancy and detection. We found that sambar had the largest spatial distribution occupying 431–437 km², while chital had the smallest at 100–158 km² across both seasons. The gaur population showed the most seasonal variation occupying from 413 to 318 km², suggesting their migration out of the Churia in summer and moving in during winter. Wild pigs showed the opposite trend occupying from 444 to 383 km²; suggesting moving into Churia in summer and out in winter. Barking deer were widespread in both seasons (329–349 km²). Tiger habitat use (\({{\hat{\varPsi } }}({\text{SE}})\)) was higher in winter 0.63 (0.11) than in summer 0.54 (0.21), but confidence intervals overlapped and area used was similar across seasons, 337 km² (winter) to 291 km² (summer). Available habitat, distribution of water sources, and human disturbance were the most common variables influencing spatial distribution of prey and habitat use of tigers at different spatial scales. Overall, we found high prey occupancy and tiger habitat use, suggesting the Churia is valuable habitat for ungulates and tigers. Given that this habitat accounts for 639 km² within CNP and 7642 km² across the entire Terai Arc, the Churia should no longer be neglected in global tiger conservation planning.     

A Global Population Genetic Study of Pantala flavescens

Among terrestrial arthropods, the dragonfly species Pantala flavescens is remarkable due to their nearly global distribution and extensive migratory ranges; the largest of any known insect. Capable of migrating across oceans, the potential for high rates of gene flow among geographically distant populations is significant. It has been hypothesized that P. flavescens may be a global panmictic population but no sufficient genetic evidence has been collected thus far. Through a population genetic analysis of P. flavescens samples from North America, South America, and Asia, the current study aimed to examine the extent at which gene flow is occurring on a global scale and discusses the implications of the genetic patterns we uncovered on population structure and genetic diversity of the species. This was accomplished using PCR-amplified cytochrome oxidase one (CO1) mitochondrial DNA data to reconstruct phylogenetic trees, a haplotype network, and perform molecular variance analyses. Our results suggested high rates of gene flow are occurring among all included geographic regions; providing the first significant evidence that Pantala flavescens should be considered a global panmictic population.     

Human disturbance and stage‐specific habitat requirements influence snowy plover site occupancy during the breeding season

Habitat use has important consequences for avian reproductive success and survival. In coastal areas with recreational activity, human disturbance may limit use of otherwise suitable habitat. Snowy plovers Charadrius nivosus have a patchy breeding distribution along the coastal areas on the Florida Panhandle, USA. Our goal was to determine the relative effects of seasonal human disturbance and habitat requirements on snowy plover habitat use. We surveyed 303 sites for snowy plovers, human disturbance, and habitat features between January and July 2009 and 2010. We made multiple visits during three different sampling periods that corresponded to snowy plover breeding: pre‐breeding, incubation, and brood‐rearing and used multi‐season occupancy models to examine whether human disturbance, habitat features, or both influenced site occupancy, colonization (probability of transition from an unoccupied site to an occupied site), and extinction (probability of transition from an occupied site to an unoccupied site). Snowy plover site occupancy and colonization was negatively associated with human disturbance and site extinction was positively associated with human disturbance. Interdune vegetation had a negative effect on occupancy and colonization, indicating that plovers were less likely to use areas with uniform, dense vegetation among dunes. Also, dune shape, beach debris, and access to low‐energy foraging areas influenced site occupancy, colonization, and extinction. Plovers used habitat based on beach characteristics that provided stage‐specific resource needs; however, human disturbance was the strongest predictor of site occupancy. In addition, vegetation plantings used to enhance dune rehabilitation may negatively impact plover site occupancy. Management actions that decrease human disturbance, such as symbolic fencing and signage, may increase the amount of breeding habitat available to snowy plovers on the Florida Panhandle and in other areas with high human activity. The specific areas that require this protection may vary across snowy plover life history stages.     

Occupancy versus colonization–extinction models for projecting population trends at different spatial scales

Understanding spatiotemporal population trends and their drivers is a key aim in population ecology. We further need to be able to predict how the dynamics and sizes of populations are affected in the long term by changing landscapes and climate. However, predictions of future population trends are sensitive to a range of modeling assumptions. Deadwood‐dependent fungi are an excellent system for testing the performance of different predictive models of sessile species as these species have different rarity and spatial population dynamics, the populations are structured at different spatial scales, and they utilize distinct substrates. We tested how the projected large‐scale occupancies of species with differing landscape‐scale occupancies are affected over the coming century by different modeling assumptions. We compared projections based on occupancy models against colonization–extinction models, conducting the modeling at alternative spatial scales and using fine‐ or coarse‐resolution deadwood data. We also tested effects of key explanatory variables on species occurrence and colonization–extinction dynamics. The hierarchical Bayesian models applied were fitted to an extensive repeated survey of deadwood and fungi at 174 patches. We projected higher occurrence probabilities and more positive trends using the occupancy models compared to the colonization–extinction models, with greater difference for the species with lower occupancy, colonization rate, and colonization:extinction ratio than for the species with higher estimates of these statistics. The magnitude of future increase in occupancy depended strongly on the spatial modeling scale and resource resolution. We encourage using colonization–extinction models over occupancy models, modeling the process at the finest resource‐unit resolution that is utilizable by the species, and conducting projections for the same spatial scale and resource resolution at which the model fitting is conducted. Further, the models applied should include key variables driving the metapopulation dynamics, such as the availability of suitable resource units, habitat quality, and spatial connectivity.     

Dispersal and local environment affect the spread of an invasive apple snail (<Emphasis Type="Italic">Pomacea maculata</Emphasis>) in Florida,USA

Dispersal and local environmental factors are major determinants of invasive species distribution. We examined how both dispersal-related geospatial characteristics and environmental factors influence an ongoing invasion of wetlands in a south-central Florida ranchland by non-native apple snails (Pomacea maculata, Ampullariidae). We found P. maculata in 73 (43%) of a random set of 171 wetlands in 2014. We used model selection to evaluate multiple hypotheses of predictors of P. maculata occurrence in 95 wetlands with standing water, including spatially-explicit distances in ditches from wetlands to the presumed entry point, Euclidean (overland) distances, presence/absence of ditches in wetlands, and environmental variables (e.g. pH). We also performed a 5-month field experiment in 20 wetlands to evaluate if snail absence was associated with conditions that limit survival and growth (i.e. unfavorable habitats). Snail occurrence was primarily associated with presence of ditches in wetlands and more neutral wetland pH. These variables more plausibly explained snail occurrence than did Euclidean (overland) distance and minimum ditch travel (rectilinear) distance from propagule sources (a major waterway). Wetland pH best explained survival and growth under the experimental conditions. We found no evidence that prior occupancy by conspecifics affected survival and growth, suggesting that dispersal limitation may contribute to lack of occupancy of wetlands, despite suitable pH. Our study supports man-made conduits as facilitators of dispersal by non-native species, where environmental characteristics (here pH) then also affect colonization within habitats. An understanding of both dispersal mechanisms and local environmental factors is necessary to better predict invasive species distribution.     

Diversity of Odonata (Insecta) from Dholbaha Dam (Distt. Hoshiarpur) in Punjab Shivalik,India

Study on the species diversity of the order Odonata was carried out during 2002 ∼ 2004 at Dholbaha dam, which has a moist deciduous forest surrounding it in district Hoshiarpur, Punjab, India. A total of 30 species belonging to 7 families of order Odonata were recorded during the study period. The family Libellulidae, represented by 18 species was the most dominant followed by Coenagrionidae (6 species), Aeshnidae (2 species) and Calopterygidae, Chlorocyphidae, Euphaeidae and Gomphidae each having 1 species. In terms of total number of individuals, family Libellulidae constituted maximum with 64.36% followed by Coenagrionidae (28.50%), Chlorocyphidae (1.83%), Gomphidae (1.62%), Euphaeidae (1.56%), Calopterygidae (1.38%) and Aeshnidae (0.75%). Pantala flavescens (Fabricius), a migratory species was the most dominant in number of individuals constituting 17.12% of the total. The least dominant species included Anax immaculifrons Rambur (0.38%) and Anax parthenope parthenope (Selys) (0.36%). Shannon-Wiener index of species diversity of Odonata was 2.988 and 3.029 during 2002–2003 and 2003–2004, respectively. Seven new species have also been reported from the Dholbaha dam during this study period thus increasing the total species number of odonates so far recorded from this area from 29 to 36.     

Occupancy modeling of <Emphasis Type="Italic">Parnassius clodius</Emphasis> butterfly populations in Grand Teton National Park,Wyoming

Estimating occupancy patterns and identifying vegetation characteristics that influence the presence of butterfly species are essential approaches needed for determining how habitat changes may affect butterfly populations in the future. The montane butterfly species, Parnassius clodius, was investigated to identify patterns of occupancy relating to habitat variables in Grand Teton National Park and Bridger-Teton National Forest, Wyoming, United States. A series of presence–absence surveys were conducted in 2013 in 41 mesic to xeric montane meadows that were considered suitable habitat for P. clodius during their flight season (June–July) to estimate occupancy (ψ) and detection probability (p). According to the null constant parameter model, P. clodius had high occupancy of ψ?=?0.78?±?0.07 SE and detection probability of p?=?0.75?±?0.04 SE. In models testing covariates, the most important habitat indicator for the occupancy of P. clodius was a strong negative association with big sagebrush (Artemisia tridentata; β = ??21.39?±?21.10 SE) and lupine (Lupinus spp.; β?=???20.03?±?21.24 SE). While P. clodius was found at a high proportion of meadows surveyed, the presence of A. tridentata may limit their distribution within montane meadows at a landscape scale because A. tridentata dominates a large percentage of the montane meadows in our study area. Future climate scenarios predicted for high elevations globally could cause habitat shifts and put populations of P. clodius and similar non-migratory butterfly populations at risk.     

Metapopulation Dynamics of a Burrowing Herbivore Drive Spatio-temporal Dynamics of Riparian Plant Communities

Animals can modify their environment by consumptive and physical activities such as herbivory and soil disturbance. Engineering species may create structures that long outlive them and have lasting impacts on local communities of plants and animals. Water voles, Arvicola amphibious, are rodents that visibly impact riparian plant communities by grazing on surface and root vegetation and excavating long-lasting burrow systems. This species has a metapopulation structure and occurs across patches which are subject to frequent extinction and colonization events, causing spatially heterogeneous disturbances across the landscape. Using a chronosequence of water vole occupancy in the Highlands of Scotland, we show that heterogeneity in plant community composition and structure—both within and between colony patches—was related to cumulative measures of past physical impact: burrow density and time since a patch was last occupied by voles, rather than to current indices of vole occupancy. In our sample of 107 patches monitored over 5 years, no fewer than 31 unique patch occupancy histories were found, each with potentially subtle differences in the accumulated influence of water vole herbivory and engineering. As a result, a patchwork of different plant successional stages occurs across the riparian landscape which is both created and maintained by water vole extinction-colonization dynamics. We propose that the water vole-vegetation system can be described as a metacommunity where dispersal by a higher tropic agent at the landscape scale influences the spatial dynamics of plants at the patch level.     

Determinants of smooth-coated otter occupancy in a rapidly urbanizing coastal landscape in Southeast Asia

Urbanization often has negative impacts on wildlife, nevertheless many species can persist in heavily modified habitats. Understanding factors that promote species persistence in urbanizing landscapes is therefore important for maintaining biodiversity in changing landscapes and may inform more biodiversity-friendly development. We investigated effects of landscape-scale variables on habitat occupancy of Smooth-coated otter (Lutrogale perspicillata) in the Inner Gulf of Thailand. In this internationally important wetland complex, 86% of natural habitats have been altered and are now highly urbanized. We conducted track and sign surveys in 60, 25 km² grid cells encompassing 1,474 km². Within each cell, we quantified the landscape cover types including urban areas, natural habitat (predominately thin ribbons of mangrove along waterways), agriculture, aquaculture, and potential prey availability using a GIS and field surveys. We used occupancy models to identify habitat variables that affected probability of detection and occupancy. Estimated otter occupancy, based on the top model, was 0.33 ± 0.07 (95% CI 0.18–0.48) and detection probability was 0.50 ± 0.05 (95% CI 0.41–0.60). Otter occupancy was positively associated with the proportion of natural habitat and the cover of traditional aquaculture ponds, but negatively associated with agriculture and urban cover. The remaining natural patches appear to act as critical refuges for otter, allowing them to persist in an otherwise heavily transformed landscape. Because aquaculture ponds are likely important prey sources for otters, it may lead to conflict with aquaculture farmers. Further studies of feeding and movement patterns in cooperation with aquaculture farmers would be beneficial for developing detailed management plans for the species in this human dominated landscape.     

Landscape-scale response to local habitat restoration in the regal fritillary butterfly (<Emphasis Type="Italic">Speyeria idalia</Emphasis>) (Lepidoptera: Nymphalidae)

The butterfly Speyeria idalia was once a widespread species in the midwestern United States, but declined precipitously during the 1980s. By the early 1990s in NW Indiana and adjacent NE Illinois, it was reduced to a single site where it persisted as a precarious three deme metapopulation managed using prescribed fire. In 1996, a large-scale high diversity restoration was initiated at the site to restore ecological connectivity for grassland and oak barrens communities across the site. Among many restoration goals, we included restoring connectivity for habitat restricted insect communities across the highly fragmented site. In support of that goal, we included over 620 local genotype plant species in the restoration. S. idalia responded positively to the restoration and was the most abundant butterfly across the site in 2014. Ironically, we incorrectly considered the primary hostplant at the site, Viola bicolor, to be non-native when we initiated the restoration and did not include it in our seed mixes. This weedy annual aggressively expanded into the restoration without our assistance. We believe that we transformed the precarious metapopulation into a well-connected patchy population across the restoration. In addition, the butterfly occupied previously unoccupied habitats far beyond the restoration. This suggests that we have created a regional mainland—island metapopulation structure, with the restoration serving as a source of immigrants that sustains colonization rates at or above the rate of local deme extinction across a landscape that currently spans almost 60 km.     

Environmental and dispersal controls of an annual plant’s distribution: how similar are patterns and apparent processes at two spatial scales?

At scales from microsites to entire ranges, species’ distributions reflect limited adaptation and/or limited dispersal. To what extent are specific distribution patterns and processes similar across scales? We investigated environmental effects—presumed because of adaptation—and independent spatial effects—presumed because of dispersal—on distribution at two scales (landscape patches of approximately 1,300 m², sampled along transects, and 4-m² cells, sampled in contiguous grids within populations) and on individual performance (water status, reproduction) in the California annual, Clarkia xantiana ssp. xantiana. Because water limitation helps set this species’ regional borders, we expected occupancy and performance at smaller scales to correlate with topographic and soil features affecting water relations. At the patch scale, environmental features associated with reduced water stress (i.e., steep slopes that face north; coarse, soft soils; igneous rather than metasedimentary parent rock) predicted occupancy. Spatial aggregation was not detected, but incomplete occupancy of apparently suitable patches indicated that dispersal limits occupancy. At the scale of small cells, relationships between environmental variables, occupancy, density, and performance varied among populations. Associations sometimes resembled those at the patch scale but sometimes opposed them. Spatial aggregation in cell occupancy and/or density occurred in all populations, implying limited dispersal, whereas spatial aggregation of water potential values in some populations might have arisen from spatially structured unmeasured environmental variables. Limited adaptation to drought and limited patch colonization appear to affect patch occupancy in C. xantiana ssp. xantiana, whereas smaller-scale patterns indicate consistent effects of limited dispersal and somewhat variable environmental effects.