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The need to increase our understanding of factors that regulate animal population dynamics has been catalysed by recent, observed declines in wildlife populations worldwide. Reliable estimates of demographic parameters are critical for addressing basic and applied ecological questions and understanding the response of parameters to perturbations (e.g. disease, habitat loss, climate change). However, to fully assess the impact of perturbation on population dynamics, all parameters contributing to the response of the target population must be estimated.

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.

We assess existing disease mitigation strategies, some in early experimental stages, which focus on the globally emerging chytrid fungus Batrachochytrium dendrobatidis with the view that disease mitigation is an essential component of population management. We discuss the precedent for each strategy in systems ranging from agriculture to human medicine, and the outlook for each strategy in terms of research needs and long-term potential.

Excepts from the introduction to the published papers arising from the 6th World Congress of Herpetology in Manuas.

Introduction (excerpt)to disease symposium publication following the 6th World Congress of Herpetology in Manaus: The goal of the symposium, Amphibian disease: Where do we go from here?, was to foster discussion of “what comes next?” in the efforts of the global scientific community to combat amphibian decline, specifically in the arena of disease. The symposium included presentations from Australia, Denmark, Ecuador, Spain, Kenya and the United States. Topics ranged from laboratory studies to landscape-scale field studies. Although the debate on “what comes next” remains in progress, the symposium prompted discussion and increased international collaborations – including subsequent workshops in Australia and Switzerland. The series of papers from this symposium, featured in this issue, do not provide a roadmap of what comes next, but illustrate the diverse nature of research into amphibian diseases and highlight some of the promising directions being pursued to understand the effects of disease on amphibian populations

1. Estimates of demographic parameters for females, in many organisms, are sparse. This is particularly worrisome as more and more species are faced with high extinction probabilities and conservation increasingly depends on actions dictated by complex predictive models that require accurate estimates of demographic parameters for each sex and species.

2. This study assesses demographic parameters, specifically temporary emigration and survival, for females, a class that has been difficult to investigate historically because of lack of data. Amphibians provide a particularly good example because there is global concern about amphibian decline yet most demographic parameter estimates are based on data from males, which we show can lead to erroneous conclusions.

3. We use 10 years of capture-recapture data from boreal toads (Bufo boreas) and the multi-state open robust design model to provide evidence for the occurrence of skipped breeding opportunities (i.e., temporary emigration) in females. This is the first time that the open robust design model has been applied to an analysis of an amphibian population that we are aware of.

4. We determined that the transition from breeder to non-breeder is obligate and the probability of a non-breeder remaining a non-breeder is 64%, thus temporary emigration is first-order Markovian in nature, where breeding probability is dependent on the previous year’s activity, i.e. if a female did not breed in year one, there is a 36% chance that she will breed in year two. With temporary emigration accounted for, we estimated between-year female survival at 87%.

5. Establishing the occurrence of temporary emigration not only reduces bias in estimates of survival probabilities, but also provides information about expected breeding attempts by females, a critical element in understanding the ecology of an organism and the impacts of outside stressors and conservation actions.

Understanding the distribution of chytridiomycosis, both at global and local scales, is important to controlling its impacts on host species (e.g., biocontrol or eradication) and to managing host amphibian populations (e.g., reintroduction and habitat management). In response to this, efforts to map observations of Batrachochytrium dendrobatidis (Bd) are underway to better understand its distribution and impact on amphibian populations. (e.g www.spatialepidemiology.net\Bd).
While there are many legitimate reasons to sample opportunistically for this pathogen, we question the validity of continuing this sort of exercise without a more cohesive and directed effort to apply the information to management of Bd. We use a recent example (Scalera et al. 2008) and some additional data from Denmark to discuss this question.

The US Geological Survey’s Amphibian Research and Monitoring Initiative (ARMI) is a national project with goals to monitor the status and trends of amphibians, conduct research on causes of declines, and provide information and support to management agencies for conservation of amphibian populations. ARMI activities are organized around extensive inventories and place-based monitoring (such as collaboration with the Greater Yellowstone I&M Network), and intensive population studies and research at selected locations (apex sites). One such is an oxbow pond on the Buffalo Fork near the Black Rock Ranger Station east of Grand Teton National Park. We have been conducting capture-mark-recapture of boreal toads (Anaxyrus boreas) at Black Rock since 2002. In concert with studies of other toad populations in the Rocky Mountains, we have documented a high rate of incidence of the chytrid fungus Batrachochytrium dendrobatidis (Bd) and a negative rate of growth of the toad population, but not the population crash or extinction observed in other populations with high occurrence of Bd.
Long-term observations at other ARMI apex sites have proven invaluable for studying effects of climate change on amphibian behavior, and the Black Rock site has been upgraded with on-site recording of temperature and precipitation, and auditory monitoring of other amphibian species. Continued research at Black Rock will be critical for understanding the interrelated effects of climate and disease on amphibians in the Greater Yellowstone Ecosystem.

Two populations of boreal toads (Bufo boreas) experienced drastic declines in abundance in the late-1990s. Evidence supported the hypothesis of disease (the chytrid fungus, Batrachochytrium dendrobatidis) as the cause of these declines, but other hypotheses had not been evaluated. We used an 11- year capture-recapture data set to evaluate weather and disease as causes of these declines. We developed sets of mathematical models that reflected hypothesized relationships between several weather variables and annual survival rates of adult males in these populations. In addition, models that reflected the possibility that the declines were caused by an introduced fungus were developed. All models were fit to the data and evaluated using a model selection criterion (QAICc). Our analysis provided strong support for the hypothesis of an introduced fungus and little support for the hypothesis that weather conditions caused the declines. Our results also suggest a strong, negative ‘marking effect’ on survival rates of boreal toads. Model-averaged estimates of survival rate are presented.

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 6 weather variables in 3 populations of Boreal Toads (Bufo boreas) from central Colorado over 8 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 https://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.

PSEUDACRIS TRISERIATA (Western Chorus Frog) and RANA SYLVATICA (Wood frog). CHYTRIDIOMYCOSIS. Chytrid fungus (Batrachochytrium dendrobatidis) is a known pathogen of anuran amphibians, and has been correlated with amphibian die-offs worldwide (Daszak et. al. 1999. Emerging Infectious Diseases 5: 735-748). In Colorado, B. dendrobatidis has infected Boreal toads (Bufo boreas)and has been identified on museum specimens of northern leopard frogs (Rana pipiens) (Carey et. al. 1999. Developmental and Comparative Immunology 23: 459-472). We report the first verified case of chytrid fungus in chorus frogs (Pseudacris triseriata) and wood frogs (Rana sylvatica) in the United States.

Chytridiomycosis is an amphibian disease linked with population declines worldwide, yet there is little information about its effects on the demography of infected populations. We used capture-recapture data from two infected populations and one uninfected population of boreal toads (Bufo boreas) in the Rocky Mountains, U.S.A. to examine a priori hypotheses about the effect of the amphibian chytrid fungus (Batrachochytrium dendrobatidis, Bd) on survival probability (Φ) and population growth rate (). Bd-positive toads had lower average annual Φ than Bd-negative individuals at sites where Bd was detected suggesting that chytridiomycosis may reduce survival by 31-42% in wild boreal toads. Bd-negative toads at infected sites had comparable survival probabilities as Bd-negative toads at the uninfected site, further supporting observed differences. Model results indicated weak evidence that environmental covariates (particularly cold temperatures during the breeding season) influenced toad survival. The diseased populations were declining by 5 - 7% per year over the 6 years of this study whereas the Bd-free population had comparatively stable ( ≈ 1) population growth. Our data suggest that Bd is a low-level chronic mortality factor in these toad populations rather than an acute factor causing rapid population declines. These results show how some amphibian populations may be coexisting with the amphibian chytrid fungus and highlight the importance of quantitative assessments of survival in wildlife populations challenged with disease.

Reintroduction is a powerful tool in our conservation toolbox. However, the necessary follow-up, i.e. long-term monitoring, is not commonplace and if instituted may lack rigor. We contend that valid monitoring is possible, even with sparse data. We present a means to monitor based on demographic data and a projection model using the Wyoming toad (Bufo baxteri) as an example. Using an iterative process, existing data is built upon gradually such that demographic estimates and subsequent inferences increase in reliability. Reintroduction and defensible monitoring may become increasingly relevant as the outlook for amphibians, especially in tropical regions, continues to deteriorate and emergency collection, captive breeding, and reintroduction become necessary. Rigorous use of appropriate modeling and an adaptive approach can validate the use of reintroduction and substantially increase its value to recovery programs.

Male boreal toads are thought to return to the breeding site year after year and, if absent in a particular year, be more likely to be present the following year. Using Pollock’s robust design we estimated temporary emigration (the probability a male toad is absent from a breeding site in a given year) at locations in Colorado: Rocky Mountain National Park (n=2) and Chaffee County (n=1). We present data that suggest that male toads do not return to the breeding site every year. Estimates of both temporary emigration parameters ( ’ = probability that a male toad is absent from the breeding site at time i if absent at time i - 1; ” = probability that a male toad is absent at time i if present at time i - 1) varied by site and time (for example, at Lost Lake, the probability of temporary emigration ranged from 10 - 29%). Although we suspected that the probability of temporary emigration would be state dependent (Markovian temporary emigration, e.g. whether a male toad is at the breeding site at time i is affected by whether it was at the breeding site at time i - 1), the data showed greater support for models of random temporary emigration or no emigration. However, some models of Markovian emigration were supported. We also hypothesized relationships between temporary emigration and a number of environmental variables. While some competitive models included environmental covariates, model selection uncertainty precluded fully defining the role of these covariates in temporary emigration.

Amphibian populations continue to be imperiled by the chytrid fungus (Batrachochytrium dendrobatidis). Understanding where B. dendrobatidis (Bd) occurs and how it may be limited by environmental factors is critical to our ability to effectively conserve the amphibians affected by Bd. We sampled 1247 amphibians (boreal toads and surrogates) at 261 boreal toad (Bufo boreas) breeding sites (97 clusters) along an 11o latitudinal gradient in the Rocky Mountains to determine the distribution of B. dendrobatidis and examine environmental factors, such as temperature and elevation, that might affect its distribution. The fungus was detected at 63% of all clusters and occurred across a range of elevations (1030–3550 m) and latitudes (37.6–48.6 o) but we detected it in only 42% of clusters in the south (site elevations higher), compared to 84% of clusters in the north (site elevations lower). Maximum ambient temperature (daily high) explained much of the variation in Bd occurrence in boreal toad populations and thus perhaps limits the occurrence of the pathogen in the Rocky Mountains to areas where climatic conditions facilitate optimal growth of the fungus. This information has implications in global climate change scenarios where warming temperatures may facilitate the spread of disease into previously un- or little-affected areas (i.e., higher elevations). This study provides the first regional-level, field-based effort to examine the relationship of environmental and geographic factors to the distribution of B. dendrobatidis in North America and will assist managers to focus on at-risk populations as determined by the local temperature regimes, latitude and elevation.

Species account for Bufo(Anaxyrus)boreas

Most research to assess amphibian declines has focused on local-scale projects on one or a few species. The Amphibian Research and Monitoring Initiative (ARMI) is a national program in the United States mandated by congressional directive and implemented by the U.S. Department of the Interior (specifically the U.S. Geological Survey, USGS). Program goals are to monitor changes in populations of amphibians across U.S. Department of the Interior lands and to address research questions related to amphibian declines using a hierarchical framework of base-, mid- and apex-level monitoring sites. ARMI is currently monitoring 83 amphibian species (29% of species in the U.S.) at mid- and apex-level areas. We chart the progress of this 5-year-old program and provide an example of mid-level monitoring from 1 of the 7 ARMI regions.

The boreal toad (Bufo boreas) is an endangered species in Colorado and is considered a candidate species for federal listing by the United States Fish and Wildlife Service. Boreal toads are absent from many areas of suitable habitat in the Southern Rocky Mountains of Colorado presumably due to a combination of causes. We moved boreal toads from existing populations and from captive rearing facilities to habitat which was historically, but is not currently, occupied by toads to experimentally examine methods of repatriation for this species.

The Amphibian Research and Monitoring Initiative (ARMI) is an innovative, multidisciplinary program that began in 2000 in response to a congressional directive for the Department of the Interior to address the issue of amphibian declines in the United States. ARMI’s formulation was cross-disciplinary, integrating U.S. Geological Survey scientists from Biology, Water, and Geography to develop a course of action (Corn and others, 2005a). The result has been an effective program with diverse, yet complementary, expertise.

ARMI’s approach to research and monitoring is multiscale. Detailed investigations focus on a few species at selected local sites throughout the country; monitoring addresses a larger number of species over broader areas (typically, National Parks and National Wildlife Refuges); and inventories to document species occurrence are conducted more extensively across the landscape. Where monitoring is conducted, the emphasis is on an ability to draw statistically defensible conclusions about the status of amphibians. To achieve this objective, ARMI has instituted a monitoring response variable that has nationwide applicability. At research sites, ARMI focuses on studying species/environment interactions, determining causes of observed declines, and developing new techniques to sample populations and analyze data. Results from activities at all scales are provided to scientists, land managers, and policymakers, as appropriate.

The ARMI program and the scientists involved contribute significantly to understanding amphibian declines at local, regional, national, and international levels. Within National Parks and National Wildlife Refuges, findings help land managers make decisions applicable to amphibian conservation. For example, the National Park Service (NPS) selected amphibians as a vital sign for several of their monitoring networks, and ARMI scientists provide information and assistance in developing monitoring methods for this NPS effort. At the national level, ARMI has had major exposure at a variety of meetings, including a dedicated symposium at the 2004 joint meetings of the Herpetologists’ League, the American Society of Ichthyologists and Herpetologists, and the Society for the Study of Amphibians and Reptiles. Several principal investigators have brought international exposure to ARMI through venues such as the World Congress of Herpetology in South Africa in 2005 (invited presentation by Dr. Gary Fellers), the Global Amphibian Summit, sponsored by the International Union for Conservation of Nature (IUCN) and Wildlife Conservation International, in Washington, D.C., 2005 (invited participation by Dr. P.S. Corn), and a special issue of the international herpetological journal Alytes focused on ARMI in 2004 (edited by Dr. C.K. Dodd, Jr.).

ARMI research and monitoring efforts have addressed at least 7 of the 21 Threatened and Endangered Species listed by the U.S. Fish and Wildlife Service (California red-legged frog [Rana draytonii ], Chiricahua leopard frog [R. chiricahuensis], arroyo toad [Bufo californicus], dusky gopher frog [Rana sevosa], mountain yellow-legged frog [R. muscosa], flatwoods salamander [Ambystoma cingulatum], and the golden coqui [Eleutherodactylus jasperi]), and 9 additional species of concern recognized by the IUCN. ARMI investigations have addressed time-sensitive research, such as emerging infectious diseases and effects on amphibians related to natural disasters like wildfire, hurricanes, and debris flows, and the effects of more constant, environmental change, like urban expansion, road development, and the use of pesticides.

Over the last 5 years, ARMI has partnered with an extensive list of government, academic, and private entities. These partnerships have been fruitful and have assisted ARMI in developing new field protocols and analytic tools, in using and refining emerging technologies to improve accuracy and efficiency of data handling, in conducting amphibian disease, malformation, and environmental effects research, and in implementing a network of monitoring and research sites. Accomplishments from these endeavors include more than 40 publications on amphibian status and trends, nearly 100 publications on amphibian ecology and causes of declines, and over 30 methodological publications. Several databases have emerged as a result of ARMI and its partnerships; one, a digital atlas of ranges for all U.S. amphibian species, was used by the IUCN to display amphibian distribution maps in the Global Amphibian Assessment Project.

Given the scope of ARMI and the panoply of projects, findings have had implications for policy. Investigations that demonstrate amphibian declines or illuminate causes of declines provide valuable information about habitat management, environmental effects, mechanisms for the spread of disease, and human/amphibian interfaces. This information has been made available to land managers, scientists, educators, Congress and other policymakers, and the public. The support afforded ARMI by Congress has been influential in the program’s development and success. The value of ARMI’s efforts will continue to increase as we are able to extend our studies spatially and temporally to answer critical questions with more confidence. We are using ARMI’s resources efficiently and continuing to develop innovative mechanisms for leveraging resources for maximum effectiveness during challenging financial times.

This report is a 5-year retrospective of the structure, methodology, progress, and contributions to the broader scientific community that have resulted from this national USGS program. We evaluate ARMI’s success to date, with regard to the challenges faced by the program and the strengths that have emerged. We chart objectives for the next 5 years that build on current accomplishments, highlight areas meriting further research, and direct efforts to overcome existing weaknesses.

Tetracycline has been used to batch-mark hatchery-reared fish with some success. It is incorporated into bone and fluoresces when prepared sections are viewed under a microscope with UV light. We investigated its utility for marking large numbers of metamorphic boreal toads, Bufo boreas. We concluded that this is not a useful technique. Marks were ambiguous - exposure to tetracycline was often indistinguishable from autofluorescence in unexposed animals.