Habitat selection by a threatened desert amphibian

Habitat degradation and fragmentation are major drivers of amphibian declines. The loss of environmental features that allow for movement between water sources may be particularly detrimental for amphibians in arid environments. Climate changes will increase the importance of microhabitats to amphibians. Enhancing areas to facilitate movement may be a necessary conservation strategy for many animal species that depend on wetlands, including federally threatened Chiricahua leopard frogs (Lithobates chiricahuensis). Habitat preferences of this frog species are not well understood. We sought to better understand fine‐scale habitat selection, to inform conservation of Chiricahua leopard frogs. We conducted our study on the Ladder Ranch, a privately owned working bison ranch in New Mexico, USA that supports a large proportion of the remaining Chiricahua leopard frogs in the state. We attached radio transmitters to 44 frogs during summer 2014. We located each frog daily for up to 8 weeks (median = 30 days). We assessed fine‐scale habitat selection by comparing characteristics at each frog location and a random location 5 m away using conditional logistic regression. Frogs preferred features that likely reduce desiccation, even after accounting for the presence of water. Frogs selected areas with more low‐lying cover, especially aquatic vegetation and woody debris, a tree overstory, and a mud substrate. We recommend managing potential movement corridors for Chiricahua leopard frogs by ensuring the presence of muddy creek bottoms, woody debris, riparian overstory, low‐lying ground cover, and pools. Microclimates created by these features seem especially valuable given warming temperatures and modified precipitation regimes, resulting in decreased surface water, soil moisture, and vegetation cover. Retaining or creating preferred habitat features and microclimates in areas between water sources may increase connectivity among isolated populations of Chiricahua leopard frogs and could improve persistence and recovery of other water‐obligate species in arid landscapes.     

The importance of ant-rich habitats for the persistence of the Wryneck Jynx torquilla on farmland

The frequency of territory occupancy is a good indicator of territory quality. We studied territory occupancy in a Swiss population of the Wryneck Jynx torquilla , a declining farmland woodpecker, with the aim of identifying key habitat features for conservation management. Both static and dynamic approaches were applied using data on nest-site occupancy of 100 territories from six successive years. The static approach models the probability of territory occupancy; the dynamic approach estimates territory colonization and extinction. Frequently occupied territories were settled earlier in the season, suggesting that they may be of better quality, and birds settling in these territories had higher breeding success. Probability of territory occupancy increased with the area of old pear orchards and decreased with the area of vegetable cultivation. Both the area of old pear orchards and the presence of conspecifics within a territory were positively related to territory colonization, whereas territory extinction was negatively related to habitat heterogeneity. Old pear orchards were characterized by having both the highest density of ant nests and the greatest amount of bare ground. The latter is likely to facilitate access to ant prey. To ensure persistence of Wryneck populations in farmland, heterogeneous habitat matrices with high ant nest density and bare ground should be promoted. Finally, provision of artificial nesting cavities is likely to enhance territory occupancy. Providing that these key resources are present, Wrynecks are likely to persist even in intensively farmed areas.     

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.     

Estimating occupancy dynamics in an anuran assemblage from Louisiana,USA

Effective monitoring programs are designed to track changes in the distribution, occurrence, and abundance of species. We developed an extension of Royle and Kéry's (2007) single species model to estimate simultaneously temporal changes in probabilities of detection, occupancy, colonization, extinction, and species turnover using data on calling anuran amphibians, collected from 2002 to 2006 in the Lower Mississippi Alluvial Valley of Louisiana, USA. During our 5-year study, estimates of occurrence probabilities declined for all 12 species detected. These declines occurred primarily in conjunction with variation in estimates of local extinction probabilities (cajun chorus frog [Pseudacris fouquettei], spring peeper [P. crucifer], northern cricket frog [Acris crepitans], Cope's gray treefrog [Hyla chrysoscelis], green treefrog [H. cinerea], squirrel treefrog [H. squirella], southern leopard frog [Lithobates sphenocephalus], bronze frog [L. clamitans], American bullfrog [L. catesbeianus], and Fowler's toad [Anaxyrus fowleri]). For 2 species (eastern narrow-mouthed toad [Gastrophryne carolinensis] and Gulf Coast toad [Incilius nebulifer]), declines in occupancy appeared to be a consequence of both increased local extinction and decreased colonization events. The eastern narrow-mouthed toad experienced a 2.5-fold increase in estimates of occupancy in 2004, possibly because of the high amount of rainfall received during that year, along with a decrease in extinction and increase in colonization of new sites between 2003 and 2004. Our model can be incorporated into monitoring programs to estimate simultaneously the occupancy dynamics for multiple species that show similar responses to ecological conditions. It will likely be an important asset for those monitoring programs that employ the same methods to sample assemblages of ecologically similar species, including those that are rare. By combining information from multiple species to decrease the variance on estimates of individual species, our results are advantageous compared to single-species models. This feature enables managers and researchers to use an entire community, rather than just one species, as an ecological indicator in monitoring programs. © 2011 The Wildlife Society.     

Colonization and extinction dynamics of a declining migratory bird are influenced by climate and habitat degradation

Uncovering the mechanisms involved in the decline of long‐distance migrants remains one of the most pressing issues in European conservation. Since the 1980s, the British breeding population of Garden Warbler Sylvia borin has declined by more than 25%. Here we use data from repeated bird surveys of woodland sites in the 1980s and in 2003–2004 to show that, although the overall population declined between the two periods, the probability of occupancy for this species increased at high latitudes and decreased at low latitudes. Range shifts such as this arise from a change in the ratio of colonizations to extinctions at the range margins, and we therefore related colonization and local extinction at the patch level to concurrent changes in temperature and habitat. The probability of patch colonization by this species was significantly lower where the percentage cover of vegetation in the understorey had declined, reducing habitat quality for this species. The probability of local extinction was significantly correlated with increasing mean May temperature, which may reflect a change in phenology, making breeding conditions less suitable. Changed regimes of grazing and woodland management could be used to increase habitat suitability and thereby increase colonization probability at the local scale, which may in turn increase the probability of patch occupancy despite future climatic unsuitability.     

The disappearing northern leopard frog (Lithobates pipiens): conservation genetics and implications for remnant populations in western Nevada

Global amphibian declines suggest a major shift in the amount and quality of habitat for these sensitive taxa. Many species that were once widespread are now experiencing declines either in part of or across their historic range. The northern leopard frog (Rana [Lithobates] pipiens] has undergone significant declines particularly in the western United States and Canada. Leopard frog population losses in Nevada are largely due to habitat fragmentation and the introduction of nonnative fish, amphibian, and plant species. Only two populations remain in the Truckee and Carson River watersheds of western Nevada which represents the western boundary of this species range. We used sequence data for an 812 base pair fragment of the mitochondrial NADH dehydrogenase 1 (ND1) gene to support a native origin for western Nevada populations. All frogs had a single haplotype (W07) from the distinct western North America ND1 haplotype clade. Data from seven polymorphic microsatellite loci show that Truckee and Carson River populations are highly differentiated from each other and from leopard frogs collected from eastern Nevada sites. Lack of gene flow among and distinct color morphs among the western Nevada populations likely predates the current geographical isolation. Comparisons with other peripheral L. pipiens populations show western Nevada populations have similar levels of gene diversity despite their contemporary isolation (H(E) 0.411, 0.482). Restoration of leopard frog populations in these watersheds will be challenging given well-entrenched nonnative bullfrog populations and major changes to the riparian zone over the past century. Declines of once common amphibian species has become a major conservation concern. Contemporary isolation of populations on a species range periphery such as the leopard frog populations in the Truckee and Carson rivers further exacerbate extirpation risk as these populations are likely to have fewer genetic resources to adaptively respond to rapidly changing biotic and abiotic environments.     

Extinction, colonization, and species occupancy in tidepool fishes

Despite the increasing sophistication of ecological models with respect to the size and spatial arrangement of habitat, there is relatively little empirical documentation of how species dynamics change as a function of habitat size and the fraction of habitat occupied. In an assemblage of tidepool fishes, I used maximum-likelihood estimation to test whether models which included habitat size provided a better fit to empirical data on extinction and colonization probabilities than models that assumed constant probabilities over all habitats. I found species differences in how extinction and colonization probabilities scaled with habitat size (and hence local population size). However, there was little evidence for a relationship between extinction and colonization probabilities and the fraction of occupied tidepools, as assumed in simple metapopulation models. Instead, colonization and extinction were independent of the fraction of occupied tidepools, favoring a MacArthur-Wilson island-mainland model. When I incorporated declines in extinction probability with tidepool volume in a simple simulation model, I found that predicted occupancy could change greatly, especially when colonization was low. However, the predicted fraction of occupied patches in the simulation model changed little when I incorporated the range of values reported here for extinction and colonization and the rate at which they scale with habitat size. Quantifying extinction and colonization patterns of natural populations is fundamental to understanding how species are distributed spatially and whether metapopulation models of species occupancy provide explanatory power for field populations. Received: 14 March 1997 / Accepted: 21 September 1997     

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 quality and context, but not patch number, drive multi-scale colonization dynamics in experimental aquatic landscapes

Colonization and extinction are primary drivers of local population dynamics, community structure, and spatial patterns of biological diversity. Existing paradigms of island biogeography, metapopulation biology, and metacommunity ecology, as well as habitat management and conservation biology based on those paradigms, emphasize patch size, number, and isolation as primary characteristics influencing colonization and extinction. Habitat selection theory suggests that patch quality could rival size, number, and isolation in determining rates of colonization and resulting community structure. We used naturally colonized experimental landscapes to address four issues: (a) how do colonizing aquatic beetles respond to variation in patch number, (b) how do they respond to variation in patch quality, (c) does patch context affect colonization dynamics, and (d) at what spatial scales do beetles respond to habitat variation? Increasing patch number had no effect on per patch colonization rates, while patch quality and context were critical in determining colonization rates and resulting patterns of abundance and species richness at multiple spatial scales. We graphically illustrate how variation in immigration rates driven by perceived predation risk (habitat quality) can further modify dynamics of the equilibrium theory of island biogeography beyond predator-driven effects on extinction rates. Our data support the importance of patch quality and context as primary determinants of colonization rate, occupancy, abundance, and resulting patterns of species richness, and reinforce the idea that management of metapopulations for species preservation, and metacommunities for local and regional diversity, should incorporate habitat quality into the predictive equation.     

Variability in primary productivity determines metapopulation dynamics

Temporal variability in primary productivity can change habitat quality for consumer species by affecting the energy levels available as food resources. However, it remains unclear how habitat-quality fluctuations may determine the dynamics of spatially structured populations, where the effects of habitat size, quality and isolation have been customarily assessed assuming static habitats. We present the first empirical evaluation on the effects of stochastic fluctuations in primary productivity—a major outcome of ecosystem functions—on the metapopulation dynamics of a primary consumer. A unique 13-year dataset from an herbivore rodent was used to test the hypothesis that inter-annual variations in primary productivity determine spatiotemporal habitat occupancy patterns and colonization and extinction processes. Inter-annual variability in productivity and in the growing season phenology significantly influenced habitat colonization patterns and occupancy dynamics. These effects lead to changes in connectivity to other potentially occupied habitat patches, which then feed back into occupancy dynamics. According to the results, the dynamics of primary productivity accounted for more than 50% of the variation in occupancy probability, depending on patch size and landscape configuration. Evidence connecting primary productivity dynamics and spatiotemporal population processes has broad implications for metapopulation persistence in fluctuating and changing environments.