Rick D. Scherer

Effects of Amphibian Chytrid Fungus on Individual Survival Probability in Wild Boreal Toads

Chytridiomycosis is linked to the worldwide decline of amphibians, yet little is known about the demographic effects of the disease. We collected capture-recapture data on three populations of boreal toads (Bufo boreas [Bufo = Anaxyrus]) in the Rocky Mountains (U.S.A.). Two of the populations were infected with chytridiomycosis and one was not. We examined the effect of the presence of amphibian chytrid fungus (Batrachochytrium dendrobatidis [Bd]; the agent of chytridiomycosis) on survival probability and population growth rate. Toads that were infected with Bd had lower average annual survival probability than uninfected individuals at sites where Bd was detected, which suggests chytridiomycosis may reduce survival by 31-42% in wild boreal toads. Toads that were negative for Bd at infected sites had survival probabilities comparable to toads at the uninfected site. Evidence that environmental covariates (particularly cold temperatures during the breeding season) influenced toad survival was weak. The number of individuals in diseased populations declined by 5-7%/year over the 6 years of the study, whereas the uninfected population had comparatively stable population growth. Our data suggest that the presence of Bd in these toad populations is not causing rapid population declines. Rather, chytridiomycosis appears to be functioning as a low-level, chronic disease whereby some infected individuals survive but the overall population effects are still negative. Our results show that some amphibian populations may be coexisting with Bd and highlight the importance of quantitative assessments of survival in diseased animal populations.

ESTIMATION OF TEMPORARY EMIGRATION IN MALE TOADS

Male boreal toads (Bufo boreas) are thought to return to the breeding site every year but, if absent in a particular year, will be more likely to return the following year. Using Pollocks robust design we estimated temporary emigration (the probability a male toad is absent from a breeding site in a given year) at three locations in Colorado, USA: two in Rocky Mountain National Park and one in Chaffee County. We present data that suggest that not all male toads return to the breeding site every year. Our analyses indicate that temporary emigration varies by site and time (for example, from 1992 to 1998, the probability of temporary emigration ranged from 10% to 29% and from 3% to 95% at Lost Lake and Kettle Tarn, respectively). Although the results provide weak evidence that males are more likely to return after a years hiatus, a general pattern of state-dependent temporary emigration was not supported. We also hypothesized relationships between temporary emigration and a number of weather variables. While some competitive models included weather covariates, imprecise and variable estimates of the effects of these covariates precluded fully defining their impact on temporary emigration.

Effects of Weather on Survival in Populations of Boreal Toads in Colorado

Abstract Understanding the relationships between animal population demography and the abiotic and biotic elements of the environments in which they live is a central objective in population ecology. For example, correlations between weather variables and the probability of survival in populations of temperate zone amphibians may be broadly applicable to several species if such correlations can be validated for multiple situations. This study focuses on the probability of survival and evaluates hypotheses based on six weather variables in three populations of Boreal Toads (Bufo boreas) from central Colorado over eight years. In addition to suggesting a relationship between some weather variables and survival probability in Boreal Toad populations, this study uses robust methods and highlights the need for demographic estimates that are precise and have minimal bias. Capture-recapture methods were used to collect the data, and the Cormack-Jolly-Seber model in program MARK was used for analysis. The top models included minimum daily winter air temperature, and the sum of the model weights for these models was 0.956. Weaker support was found for the importance of snow depth and the amount of environmental moisture in winter in modeling survival probability. Minimum daily winter air temperature was positively correlated with the probability of survival in Boreal Toads at other sites in Colorado and has been identified as an important covariate in studies in other parts of the world. If air temperatures are an important component of survival for Boreal Toads or other amphibians, changes in climate may have profound impacts on populations.

The importance of local and landscape-scale processes to the occupancy of wetlands by pond-breeding amphibians

Variation in the distribution and abundance of species across landscapes has traditionally been attributed to processes operating at fine spatial scales (i.e., environmental conditions at the scale of the sampling unit), but processes that operate across larger spatial scales such as seasonal migration or dispersal are also important. To determine the relative importance of these processes, we evaluated hypothesized relationships between the probability of occupancy in wetlands by two amphibians [wood frogs (Lithobates sylvaticus) and boreal chorus frogs (Pseudacris maculata)] and attributes of the landscape measured at three spatial scales in Rocky Mountain National Park, Colorado. We used cost-based buffers and least-cost distances to derive estimates of landscape attributes that may affect occupancy patterns from the broader spatial scales. The most highly ranked models provide strong support for a positive relationship between occupancy by breeding wood frogs and the amount of streamside habitat adjacent to a wetland. The model selection results for boreal chorus frogs are highly uncertain, though several of the most highly ranked models indicate a positive association between occupancy and the number of neighboring, occupied wetlands. We found little evidence that occupancy of either species was correlated with local-scale attributes measured at the scale of individual wetlands, suggesting that processes operating at broader scales may be more important in influencing occupancy patterns in amphibian populations.

Evidence for plasticity in the frequency of skipped breeding opportunities in common toads

Breeding is limited by energetic or environmental constraints and long-lived species sometimes skip breeding opportunities. Environmental conditions may vary considerably across the geographic and elevational range of a species and species that can respond through variation in life history strategies are likely to maintain populations at the extremes of their ranges. The decision to skip breeding enables animals to adjust life history to circumstances, and plasticity in behavior allows implementation of adjustments. Elevational patterns suggest that breeding may be limited physiologically at high elevations (e.g., greater probability of skipped breeding; resources and environmental conditions more variable) in contrast to low elevations (probability of skipping breeding lower; resources and environmental conditions more predictable). We estimated the probabilities of survival and skipped breeding in a high-elevation population of common toads and compared estimates to existing data for common toads at low elevations, and to another toad species inhabiting a similar high elevation environment. Female common toads at high elevations tend to have high probabilities of skipping breeding and survival relative to data for common toads at low elevations, and appear to use a similar strategy of skipping breeding in response to similar environmental constraints as other toads at high elevations. We provide evidence of variability in this aspect of life history for common toads. Understanding variation in life history within widely distributed species is critical. Knowing that certain life history strategies are employed on a continuum informs conservation efforts, especially as impacts of climate change are likely to be different depending on elevation.

Breeding Chorus Indices Are Weakly Related to Estimated Abundance of Boreal Chorus Frogs

Abstract Call surveys used to monitor breeding choruses of anuran amphibians generate index values that are frequently used to represent the number of male frogs present, but few studies have quantified this relationship. We compared abundance of male Boreal Chorus Frogs (Pseudacris maculata), estimated using capture–recapture methods in two populations in Colorado, to call index values derived from automated recordings. Single index values, such as might result from large monitoring efforts, were unrelated to population size. A synthetic call saturation index (CSI), the daily proportion of the maximum possible sum of index values derived from multiple recordings, was greater in larger populations, but the relationship was not highly predictive.

PORTRAIT OF A SMALL POPULATION OF BOREAL TOADS (ANAXYRUS BOREAS)

Abstract Much attention has been given to the conservation of small populations, those that are small because of decline, and those that are naturally small. Small populations are of particular interest because ecological theory suggests that they are vulnerable to the deleterious effects of environmental, demographic, and genetic stochasticity as well as natural and human-induced catastrophes. However, testing theory and developing applicable conservation measures for small populations is hampered by sparse data. This lack of information is frequently driven by computational issues with small data sets that can be confounded by the impacts of stressors. We present estimates of demographic parameters from a small population of Boreal Toads (Anaxyrus boreas) that has been surveyed since 2001 by using capture–recapture methods. Estimates of annual adult survival probability are high relative to other Boreal Toad populations, whereas estimates of recruitment rate are low. Despite using simple models, clear patterns emerged from the analyses, suggesting that population size is constrained by low recruitment of adults and is declining slowly. These patterns provide insights that are useful in developing management directions for this small population, and this study serves as an example of the potential for small populations to yield robust and useful information despite sample size constraints.

Estimates of Survival Probability from Two Populations of Giant Gartersnakes in California’s Great Central Valley

The loss and modification of freshwater ecosystems has led to high rates of imperilment for freshwater species. The Giant Gartersnake (Thamnophis gigas) is among the species that have suffered declines in abundance and spatial distribution and is currently listed as a threatened species by the U.S. government and the State of California. Conservation and management of populations of T. gigas are hampered by a lack of information on its demography. Without estimates of demographic parameters, the status of the population is difficult to characterize, and identifying the parameters to target in management planning is problematic. We used capture–recapture data from two populations in the Great Central Valley of California to estimate annual survival probability. We also evaluated hypothesized causes of variation in survival probability among individuals and among years. Model-selection results for the population in the American Basin indicated that females have a higher survival probability than males and that survival probability and the amount of precipitation between 15 April and 15 May in a year are negatively correlated. Associations with other weather covariates were also supported, but the evidence was weaker. For the population in the Natomas Basin, the model-selection results indicated a positive association between survival probability and the body size of an individual (snout-to-vent length). There was also evidence that females have higher survival probabilities than males, but the support for this effect was weaker. This work fills gaps in our understanding of the demography of T. gigas by providing estimates of survival probability for males and age classes for which estimates were previously not available.

Demography of common toads after local extirpation of co-occurring midwife toads

Estimating demographic parameters like survival or recruitment provides insight into the state and trajectory of populations, but understanding the contexts influencing those parameters, including both biotic and abiotic factors, is particularly important for management and conservation. At a high elevation national park in Central Spain, common toads (Bufo bufo) are apparently taking advantage of the near-extirpation of the midwife toad (Alytes obstetricans), as colonization into new breeding ponds is evident. Within this scenario, we expected demographic parameters of common toad populations to be affected favorably by the putative release from competition. However, we found the population growth rate was negative in 4 of 5 years at the long-standing population; survival probability at the long-standing population and newly-colonised breeding ponds was lower than reported for other toads living at high elevations and the probability of recruitment was inadequate to compensate for the survival rate in maintaining a positive trajectory for either of the breeding ponds. We assessed weather covariates and disease for their contribution to the context that may be limiting the common toad’s successful use of the niche vacated by the midwife toad.

Logistic quantile regression provides improved estimates for bounded avian counts: A case study of California Spotted Owl fledgling production

ABSTRACT Counts of avian fledglings, nestlings, or clutch size that are bounded below by zero and above by some small integer form a discrete random variable distribution that is not approximated well by conventional parametric count distributions such as the Poisson or negative binomial. We developed a logistic quantile regression model to provide estimates of the empirical conditional distribution of a bounded discrete random variable. The logistic quantile regression model requires that counts are randomly jittered to a continuous random variable, logit transformed to bound them between specified lower and upper values, then estimated in conventional linear quantile regression, repeating the 3 steps and averaging estimates. Back-transformation to the original discrete scale relies on the fact that quantiles are equivariant to monotonic transformations. We demonstrate this statistical procedure by modeling 20 years of California Spotted Owl fledgling production (0−3 per territory) on the Lassen National Forest, California, USA, as related to climate, demographic, and landscape habitat characteristics at territories. Spotted Owl fledgling counts increased nonlinearly with decreasing precipitation in the early nesting period, in the winter prior to nesting, and in the prior growing season; with increasing minimum temperatures in the early nesting period; with adult compared to subadult parents; when there was no fledgling production in the prior year; and when percentage of the landscape surrounding nesting sites (202 ha) with trees ≥25 m height increased. Changes in production were primarily driven by changes in the proportion of territories with 2 or 3 fledglings. Average variances of the discrete cumulative distributions of the estimated fledgling counts indicated that temporal changes in climate and parent age class explained 18% of the annual variance in owl fledgling production, which was 34% of the total variance. Prior fledgling production explained as much of the variance in the fledgling counts as climate, parent age class, and landscape habitat predictors. Our logistic quantile regression model can be used for any discrete response variables with fixed upper and lower bounds.