Multistate occupancy modeling improves understanding of amphibian breeding dynamics in the Greater Yellowstone Area.

Discerning the determinants of species occurrence across landscapes is fundamental to their conservation and management. In spatially and climatologically complex landscapes, explaining the dynamics of occurrence can lead to improved understanding of short- vs. long-term trends and offer novel insight on local vs. regional change. We examined the changes in occupancy for two species of anurans with different life histories over a decade using hundreds of wetland sites in Yellowstone and Grand Teton National Parks. To account for the joint dynamics of wetland drying and amphibian breeding, we adopted a multistate occupancy model as a means to investigate mechanistic relationships of observed occurrence patterns with climatological drivers of wetland hydrologic variability. This approach allowed us to decompose occupancy dynamics into habitat changes caused by wetland drying and amphibian breeding activity, conditional on available water and previous breeding state. Over our 10-yr time series, we observed considerable variability in climate drivers and the proportion of dry wetlands. Boreal chorus frogs (Pseudacris maculata) were more responsive to changes in wetland inundation status than Columbia spotted frogs (Rana luteiventris), as indicated by higher breeding colonization probabilities under favorable (wet) conditions. Both species had high probabilities of breeding persistence in permanently inundated wetlands with prior breeding. Despite the absence of multi-year drought in our time series, mechanistic relationships described here offer insights on how future climate variation may result in reduced and/or shifted occurrence patterns for pond-breeding anurans in the Greater Yellowstone Area. Further, our modeling approach may prove valuable in evaluating determinants of occurrence for other species that are dependent on wetlands or other dynamic habitats.

Evaluation of a combined macrophyte-epiphyte bioassay for assessing nutrient enrichment in the Portneuf River, Idaho, USA.

We describe and evaluate a laboratory bioassay that uses Lemna minor L. and attached epiphytes to characterize the status of ambient and nutrient-enriched water from the Portneuf River, Idaho. Specifically, we measured morphological (number of fronds, longest surface axis, and root length) and population-level (number of plants and dry mass) responses of L. minor and community-level (ash-free dry mass [AFDM] and chlorophyll a [Chl a]) responses of epiphytes to nutrient enrichment. Overall, measures of macrophyte biomass and abundance increased with increasing concentrations of dissolved phosphorus (P) and responded more predictably to nutrient enrichment than morphological measures. Epiphyte AFDM and Chl a were also greatest in P-enriched water; enrichments of N alone produced no measurable epiphytic response. The epiphyte biomass response did not directly mirror macrophyte biomass responses, illustrating the value of a combined macrophyte-epiphyte assay to more fully evaluate nutrient management strategies. Finally, the most P-enriched waters not only supported greater standing stocks of macrophyte and epiphytes but also had significantly higher water column dissolved oxygen and dissolved organic carbon concentrations and a lower pH. Advantages of this macrophyte-epiphyte bioassay over more traditional single-species assays include the use of a more realistic level of biological organization, a relatively short assay schedule (~10 days), and the inclusion of multiple biological response and water-quality measures.

A Dataset of Amphibian Species in U.S. National Parks.

National parks and other protected areas are important for preserving landscapes and biodiversity worldwide. An essential component of the mission of the United States (U.S.) National Park Service (NPS) requires understanding and maintaining accurate inventories of species on protected lands. We describe a new, national-scale synthesis of amphibian species occurrence in the NPS system. Many park units have a list of amphibian species observed within their borders compiled from various sources and available publicly through the NPSpecies platform. However, many of the observations in NPSpecies remain unverified and the lists are often outdated. We updated the amphibian dataset for each park unit by collating old and new park-level records and had them verified by regional experts. The new dataset contains occurrence records for 292 of the 424 NPS units and includes updated taxonomy, international and state conservation rankings, hyperlinks to a supporting reference for each record, specific notes, and related fields which can be used to better understand and manage amphibian biodiversity within a single park or group of parks.

Assessing the potential for salmon recovery via floodplain restoration: a multitrophic level comparison of dredge-mined to reference segments.

Pre-restoration studies typically focus on physical habitat, rather than the food-base that supports aquatic species. However, both food and habitat are necessary to support the species that habitat restoration is frequently aimed at recovering. Here we evaluate if and how the productivity of the food-base that supports fish production is impaired in a dredge-mined floodplain within the Yankee Fork Salmon River (YFSR), Idaho (USA); a site where past restoration has occurred and where more has been proposed to help recover anadromous salmonids. Utilizing an ecosystem approach, we found that the dredged segment had comparable terrestrial leaf and invertebrate inputs, aquatic primary producer biomass, and production of aquatic invertebrates relative to five reference floodplains. Thus, the food-base in the dredged segment did not necessarily appear impaired. On the other hand, we observed that off-channel aquatic habitats were frequently important to productivity in reference floodplains, and the connection of these habitats in the dredged segment via previous restoration increased invertebrate productivity by 58%. However, using a simple bioenergetic model, we estimated that the invertebrate food-base was at least 4× larger than present demand for food by fish in dredged and reference segments. In the context of salmon recovery efforts, this observation questions whether additional food-base productivity provided by further habitat restoration would be warranted in the YFSR. Together, our findings highlight the importance of studies that assess the aquatic food-base, and emphasize the need for more robust ecosystem models that evaluate factors potentially limiting fish populations that are the target of restoration.

Nutrient limitation of algae and macrophytes in streams: Integrating laboratory bioassays, field experiments, and field data.

Successful eutrophication control strategies need to address the limiting nutrient. We conducted a battery of laboratory and in situ nutrient-limitation tests with waters collected from 9 streams in an agricultural region of the upper Snake River basin, Idaho, USA. Laboratory tests used the green alga Raphidocelis subcapitata, the macrophyte Lemna minor (duckweed) with native epiphytes, and in situ nutrient-limitation tests of periphyton were conducted with nutrient-diffusing substrates (NDS). In the duckweed/epiphyte test, P saturation occurred when concentrations reached about 100 µg/L. Chlorophyll a in epiphytic periphyton was stimulated at low P additions and by about 100 µg/L P, epiphytic periphyton chlorophyll a appeared to be P saturated. Both duckweed and epiphyte response patterns with total N were weaker but suggested a growth stimulation threshold for duckweed when total N concentrations exceeded about 300 µg/L and approached saturation at the highest N concentration tested, 1300 µg/L. Nutrient uptake by epiphytes and macrophytes removed up to 70 and 90% of the N and P, respectively. The green algae and the NDS nutrient-limitation test results were mostly congruent; N and P co-limitation was the most frequent result for both test series. Across all tests, when N:P molar ratios >30 (mass ratios >14), algae or macrophyte growth was P limited; N limitation was observed at N:P molar ratios up to 23 (mass ratios up to 10). A comparison of ambient periphyton chlorophyll a concentrations with chlorophyll a accrued on control artificial substrates in N-limited streams, suggests that total N concentrations associated with a periphyton chlorophyll a benchmark for desirable or undesirable conditions for recreation would be about 600 to 1000 µg/L total N, respectively. For P-limited streams, the corresponding benchmark concentrations were about 50 to 90 µg/L total P, respectively. Our approach of integrating controlled experiments and matched biomonitoring field surveys was cost effective and more informative than either approach alone.

Wetland drying linked to variations in snowmelt runoff across Grand Teton and Yellowstone national parks.

In Grand Teton and Yellowstone national parks wetlands offer critical habitat and play a key role in supporting biological diversity. The shallow depths and small size of many palustrine wetlands in these protected areas and elsewhere make them vulnerable to changes in climate compared with larger and deeper aquatic habitats. Here, we use a simple water balance model to generate estimates of biophysical drivers of wetland change. We then examine the relationship between wetland inundation status and four principal drivers (i.e., temperature, precipitation, evapotranspiration, and runoff) spanning varying meteorological conditions over an 8-year time series from Grand Teton and Yellowstone national parks. We found that models containing snowmelt runoff outperformed models with other meteorological drivers and determined that a higher percentage of surveyed wetlands were dry in years characterized by lower runoff. Our work further shows that wetland drying was widespread across both parks, but sub-regional variations were best described at the hydrologic subbasin-level. Documenting the varying responses of wetlands to meteorological drivers is a necessary first step to identifying which subbasins are most sensitive to recent climatic change and contemplating how future change may alter the distribution of wetlands and their dependent taxa.

Effects of brine contamination from energy development on wetland macroinvertebrate community structure in the Prairie Pothole Region.

Wetlands in the Prairie Pothole Region (PPR) of North America support macroinvertebrate communities that are integral to local food webs and important to breeding waterfowl. Macroinvertebrates in PPR wetlands are primarily generalists and well adapted to within and among year changes in water permanence and salinity. The Williston Basin, a major source of U.S. energy production, underlies the southwest portion of the PPR. Development of oil and gas results in the coproduction of large volumes of highly saline, sodium chloride dominated water (brine) and the introduction of brine can alter wetland salinity. To assess potential effects of brine contamination on macroinvertebrate communities, 155 PPR wetlands spanning a range of hydroperiods and salinities were sampled between 2014 and 2016. Brine contamination was documented in 34 wetlands with contaminated wetlands having significantly higher chloride concentrations, specific conductance and percent dominant taxa, and significantly lower taxonomic richness, Shannon diversity, and Pielou evenness scores compared to uncontaminated wetlands. Non-metric multidimensional scaling found significant correlations between several water quality parameters and macroinvertebrate communities. Chloride concentration and specific conductance, which can be elevated in naturally saline wetlands, but are also associated with brine contamination, had the strongest correlations. Five wetland groups were identified from cluster analysis with many of the highly contaminated wetlands located in a single cluster. Low or moderately contaminated wetlands were distributed among the remaining clusters and had macroinvertebrate communities similar to uncontaminated wetlands. While aggregate changes in macroinvertebrate community structure were observed with brine contamination, systematic changes were not evident, likely due to the strong and potentially confounding influence of hydroperiod and natural salinity. Therefore, despite the observed negative response of macroinvertebrate communities to brine contamination, macroinvertebrate community structure alone is likely not the most sensitive indicator of brine contamination in PPR wetlands.

Quantifying climate sensitivity and climate-driven change in North American amphibian communities.

Changing climate will impact species' ranges only when environmental variability directly impacts the demography of local populations. However, measurement of demographic responses to climate change has largely been limited to single species and locations. Here we show that amphibian communities are responsive to climatic variability, using >500,000 time-series observations for 81 species across 86 North American study areas. The effect of climate on local colonization and persistence probabilities varies among eco-regions and depends on local climate, species life-histories, and taxonomic classification. We found that local species richness is most sensitive to changes in water availability during breeding and changes in winter conditions. Based on the relationships we measure, recent changes in climate cannot explain why local species richness of North American amphibians has rapidly declined. However, changing climate does explain why some populations are declining faster than others. Our results provide important insights into how amphibians respond to climate and a general framework for measuring climate impacts on species richness.

Quantitative evidence for the effects of multiple drivers on continental-scale amphibian declines.

Since amphibian declines were first proposed as a global phenomenon over a quarter century ago, the conservation community has made little progress in halting or reversing these trends. The early search for a "smoking gun" was replaced with the expectation that declines are caused by multiple drivers. While field observations and experiments have identified factors leading to increased local extinction risk, evidence for effects of these drivers is lacking at large spatial scales. Here, we use observations of 389 time-series of 83 species and complexes from 61 study areas across North America to test the effects of 4 of the major hypothesized drivers of declines. While we find that local amphibian populations are being lost from metapopulations at an average rate of 3.79% per year, these declines are not related to any particular threat at the continental scale; likewise the effect of each stressor is variable at regional scales. This result - that exposure to threats varies spatially, and populations vary in their response - provides little generality in the development of conservation strategies. Greater emphasis on local solutions to this globally shared phenomenon is needed.

The Shifting Climate Portfolio of the Greater Yellowstone Area.

Knowledge of climatic variability at small spatial extents (< 50 km) is needed to assess vulnerabilities of biological reserves to climate change. We used empirical and modeled weather station data to test if climate change has increased the synchrony of surface air temperatures among 50 sites within the Greater Yellowstone Area (GYA) of the interior western United States. This important biological reserve is the largest protected area in the Lower 48 states and provides critical habitat for some of the world's most iconic wildlife. We focused our analyses on temporal shifts and shape changes in the annual distributions of seasonal minimum and maximum air temperatures among valley-bottom and higher elevation sites from 1948-2012. We documented consistent patterns of warming since 1948 at all 50 sites, with the most pronounced changes occurring during the Winter and Summer when minimum and maximum temperature distributions increased. These shifts indicate more hot temperatures and less cold temperatures would be expected across the GYA. Though the shifting statistical distributions indicate warming, little change in the shape of the temperature distributions across sites since 1948 suggest the GYA has maintained a diverse portfolio of temperatures within a year. Spatial heterogeneity in temperatures is likely maintained by the GYA's physiographic complexity and its large size, which encompasses multiple climate zones that respond differently to synoptic drivers. Having a diverse portfolio of temperatures may help biological reserves spread the extinction risk posed by climate change.