Drought-mediated extinction of an arid-land amphibian: insights from a spatially explicit dynamic occupancy model.

Understanding how natural and anthropogenic processes affect population dynamics of species with patchy distributions is critical to predicting their responses to environmental changes. Despite considerable evidence that demographic rates and dispersal patterns vary temporally in response to an array of biotic and abiotic processes, few applications of metapopulation theory have sought to explore factors that explain spatiotemporal variation in extinction or colonization rates. To facilitate exploring these factors, we extended a spatially explicit model of metapopulation dynamics to create a framework that requires only binary presence-absence data, makes few assumptions about the dispersal process, and accounts for imperfect detection. We apply this framework to 22 yr of biannual survey data for lowland leopard frogs, Lithobates yavapaiensis, an amphibian that inhabits arid stream systems in the southwestern United States and northern Mexico. Our results highlight the importance of accounting for factors that govern temporal variation in transition probabilities, as both extinction and colonization rates varied with hydrologic conditions. Specifically, local extinctions were more frequent during drought periods, particularly at sites without reliable surface water. Colonization rates increased when larval and dispersal periods were wetter than normal, which increased the probability that potential emigrants metamorphosed and reached neighboring sites. Extirpation of frogs from all sites in one watershed during a period of severe drought demonstrated the influence of site-level features, as frogs persisted only in areas where most sites held water consistently and where the amount of sediment deposited from high-elevation wildfires was low. Application of our model provided novel insights into how climate-related processes affected the distribution and population dynamics of an arid-land amphibian. The approach we describe has application to a wide array of species that inhabit patchy environments, can improve our understanding of factors that govern metapopulation dynamics, and can inform strategies for conservation of imperiled species.

A spatially explicit hierarchical model to characterize population viability.

Many of the processes that govern the viability of animal populations vary spatially, yet population viability analyses (PVAs) that account explicitly for spatial variation are rare. We develop a PVA model that incorporates autocorrelation into the analysis of local demographic information to produce spatially explicit estimates of demography and viability at relatively fine spatial scales across a large spatial extent. We use a hierarchical, spatial, autoregressive model for capture-recapture data from multiple locations to obtain spatially explicit estimates of adult survival (ϕad ), juvenile survival (ϕjuv ), and juvenile-to-adult transition rates (ψ), and a spatial autoregressive model for recruitment data from multiple locations to obtain spatially explicit estimates of recruitment (R). We combine local estimates of demographic rates in stage-structured population models to estimate the rate of population change (λ), then use estimates of λ (and its uncertainty) to forecast changes in local abundance and produce spatially explicit estimates of viability (probability of extirpation, Pex ). We apply the model to demographic data for the Sonoran desert tortoise (Gopherus morafkai) collected across its geographic range in Arizona. There was modest spatial variation in λ^ (0.94-1.03), which reflected spatial variation in ϕ^ad (0.85-0.95), ϕ^juv (0.70-0.89), and ψ^ (0.07-0.13). Recruitment data were too sparse for spatially explicit estimates; therefore, we used a range-wide estimate ( R^  = 0.32 1-yr-old females per female per year). Spatial patterns in demographic rates were complex, but ϕ^ad , ϕ^juv , and λ^ tended to be lower and ψ^ higher in the northwestern portion of the range. Spatial patterns in Pex varied with local abundance. For local abundances >500, Pex was near zero (<0.05) across most of the range after 100 yr; as abundances decreased, however, Pex approached one in the northwestern portion of the range and remained low elsewhere. When local abundances were <50, western and southern populations were vulnerable (Pex  > 0.25). This approach to PVA offers the potential to reveal spatial patterns in demography and viability that can inform conservation and management at multiple spatial scales, provide insight into scale-related investigations in population ecology, and improve basic ecological knowledge of landscape-level phenomena.

Hydrologic variability governs population dynamics of a vulnerable amphibian in an arid environment.

Dynamics of many amphibian populations are governed by the distribution and availability of water. Therefore, understanding the hydrological mechanisms that explain spatial and temporal variation in occupancy and abundance will improve our ability to conserve and recover populations of vulnerable amphibians. We used 16 years of survey data from intermittent mountain streams in the Sonoran Desert to evaluate how availability of surface water affected survival and adult recruitment of a threatened amphibian, the lowland leopard frog (Lithobates yavapaiensis). Across the entire study period, monthly survival of adults ranged from 0.72 to 0.99 during summer and 0.59 to 0.94 during winter and increased with availability of surface water (Z = 7.66; P < 0.01). Recruitment of frogs into the adult age class occurred primarily during winter and ranged from 1.9 to 3.8 individuals/season/pool; like survival, recruitment increased with availability of surface water (Z = 3.67; P < 0.01). Although abundance of frogs varied across seasons and years, we found no evidence of a systematic trend during the 16-year study period. Given the strong influence of surface water on population dynamics of leopard frogs, conservation of many riparian obligates in this and similar arid regions likely depends critically on minimizing threats to structures and ecosystem processes that maintain surface waters. Understanding the influence of surface-water availability on riparian organisms is particularly important because climate change is likely to decrease precipitation and increase ambient temperatures in desert riparian systems, both of which have the potential to alter fundamentally the hydrology of these systems.

Red squirrel middens influence abundance but not diversity of other vertebrates.

Some animals modify the environment in ways that can influence the resources available to other species. Because red squirrels (Tamiasciurus hudsonicus) create large piles of conifer-cone debris (middens) in which they store cones, squirrels concentrate resources that might affect biodiversity locally. To determine whether other animals are attracted to midden sites beyond their affinity for the same resources that attract red squirrels, we assessed associations between middens, mammals, and birds at population and community levels. We surveyed 75 middens where residency rates of red squirrels varied during the previous five years; sampling along this residency gradient permitted us to evaluate the influence of resources at middens beyond the influence of a resident squirrel. At each location, we quantified vegetation, landscape structure, abundance of conifer cones, and midden structure, and used capture-recapture, distance sampling, and remote cameras to quantify presence, abundance, and species richness of mammals and birds. Red squirrels and the resources they concentrated at middens influenced mammals and birds at the population scale and to a lesser extent at the community scale. At middens with higher residency rates of red squirrels, richness of medium and large mammals increased markedly and species richness of birds increased slightly. After accounting for local forest characteristics, however, only species richness of medium-to-large mammals was associated with a red squirrel being resident during surveys. In areas where red squirrels were resident during surveys or in areas with greater amounts of resources concentrated by red squirrels, abundances of two of four small mammal species and two of four bird species increased. We conclude that the presence of this ecosystem modifier and the resources it concentrates influence abundance of some mammals and birds, which may have implications for maintaining biodiversity across the wide geographic range inhabited by red squirrels and other larderhoarding animals.

Spatial and temporal variation in survival of a rare reptile: a 22-year study of Sonoran desert tortoises.

Although many species may be vulnerable to changes in climate, forecasting species-level responses can be challenging given the array of physiological, behavioral, and demographic attributes that might be affected. One strategy to improve forecasts is to evaluate how species responded to climatic variation in the past. We used 22 years of capture-recapture data for Sonoran desert tortoises (Gopherus morafkai) collected from 15 locations across their geographic range in Arizona to evaluate how environmental factors affected spatial and temporal variation in survival. Although rates of annual survival were generally high ([Formula: see text] = 0.92), survival of adults decreased with drought severity, especially in portions of their range that were most arid and nearest to cities. In three locations where large numbers of carcasses from marked tortoises were recovered, survival of adults was markedly lower during periods of severe drought ([Formula: see text] = 0.77-0.81) compared to all other periods ([Formula: see text] = 0.93-0.98). Assuming continued levels of dependency of humans on fossil fuels, survival of adult tortoises is predicted to decrease by an average of 3 % during 2035-2060 relative to survival during 1987-2008 in 14 of the 15 populations we studied. This decrease could reduce persistence of tortoise populations, especially in arid portions of their range. Temporal and spatial variation in drought conditions are important determinants of survival in adult desert tortoises.

Importance of environmental and spatial gradients on patterns and consequences of resource selection.

Strategies to conserve rare species require identifying resources that function as important habitat elements and that promote high demographic performance. Assessing the relative importance of resources, however, can be confounded by natural variation in resource availability and by the hierarchical spatial structure in which resources are organized. Because availability and relative importance of resources often vary across environmental and spatial gradients, we used gradients together with resource selection functions and variance decomposition to assess the relative importance of resources to nest site selection and reproductive performance of Ferruginous Pygmy-Owls (Glaucidium brasilianum). We measured habitat characteristics of 106 nests and paired available sites at five spatial scales across a 220-km gradient of precipitation and vegetation in northwest Mexico, in a region adjacent to the southwestern United States where pygmy-owls have declined to near extinction. Resources explained 76-85% of variation in nest site selection and 21-31% of variation in reproductive performance across all spatial scales combined. Although we found evidence of resource selection at each scale, the magnitude of selection and influence of resources on reproductive performance were greatest where availability of selected resources were low and where temperature extremes and predation risk likely increased the relative importance of these resources. At larger scales, geographic changes in resource use corresponded with changes in availability, whereas at smaller scales, resource use varied little despite changes in availability, suggesting higher specificity and importance of resources at smaller scales. At the smallest scale, owls selected nest cavities with smaller entrances, larger volume, greater height, and orientations that produced cooler microclimates in the hottest regions of the study area; these choices promoted higher reproductive performance. Cavity resources explained more variation in selection and reproductive performance than resources at larger scales, highlighting their importance as conservation targets. High correlation of resource characteristics among spatial scales, however, indicated that selection of resources at small scales depended on characteristics of resources at larger scales. Assessing how resource selection changes in response to underlying variation in resource availability can help prioritize resources most important for conservation and management.