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Range-wide salamander densities reveal a key component of terrestrial vertebrate biomass in eastern North American forests

Characterizing the population density of species is a central interest in ecology. Eastern North America is the global hotspot for biodiversity of plethodontid salamanders, an inconspicuous component of terrestrial vertebrate communities, and among the most widespread is the eastern red-backed salamander, Plethodon cinereus . Previous work suggests population densities are high with significant geographic variation, but comparisons among locations are challenged by lack of standardization of methods and failure to accommodate imperfect detection. We present results from a large-scale research network that accounts for detection uncertainty using systematic survey protocols and robust statistical models. We analysed mark–recapture data from 18 study areas across much of the species range. Estimated salamander densities ranged from 1950 to 34 300 salamanders ha −1 , with a median of 9965 salamanders ha −1 . We compared these results to previous estimates for P. cinereus and other abundant terrestrial vertebrates. We demonstrate that overall the biomass of P. cinereus , a secondary consumer, is of similar or greater magnitude to widespread primary consumers such as white-tailed deer ( Odocoileus virginianus ) and Peromyscus mice, and two to three orders of magnitude greater than common secondary consumer species. Our results add empirical evidence that P. cinereus , and amphibians in general, are an outsized component of terrestrial vertebrate communities in temperate ecosystems.

Biology Letters

Using expert knowledge to support Endangered Species Act decision‐making for data‐deficient species

Many questions relevant to conservation decision making are characterized by extreme uncertainty due to lack of empirical data and complexity of the underlying ecological processes, leading to a rapid increase in the use of structured protocols to elicit expert knowledge. Published ecological applications often employ a modified Delphi method, where experts provide judgments anonymously and mathematical aggregation techniques are used to combine judgments. The Sheffield Elicitation Framework (SHELF) differs in its behavioral approach to synthesizing individual judgments into a fully specified probability distribution for an unknown quantity. This study demonstrates the remote use of the SHELF protocol for an extinction risk assessment of three subterranean aquatic species petitioned for listing under the US Endangered Species Act. Experts were provided an empirical threat assessment for each known locality using video conferencing and asked for judgments on the probability of population persistence over four generations using online submission forms and R‐shiny apps available through the SHELF package. Despite large uncertainty for all populations, results reveal key differences between species’ risk of extirpation based on spatial variation in dominant threats, local land use and management practices, and microhabitat use. The resulting probability distributions provide decision makers with a full picture of uncertainty that is consistent with the probabilistic nature of risk assessments, and discussions during the behavioral aggregation stage clearly document dominant threats (e.g., development, timber harvest, animal agriculture, and cave visitation) and their interactions with local cave geology and species’ habitat preferences. Our virtual implementation of the SHELF protocol demonstrates the flexibility of this approach for conservation applications operating on budgets and timelines that can limit in‐person meetings of geographically dispersed experts.

Conservation Biology

Forest Vegetation Monitoring Protocol for National Parks in the North Coast and Cascades Network

Plant communities are the foundation for terrestrial trophic webs and animal habitat, and their structure and species composition are an integrated result of biological and physical drivers (Gates, 1993). Additionally, they have a major role in geologic, geomorphologic and soil development processes (Jenny, 1941; Stevens and Walker, 1970). Throughout most of the Pacific Northwest, environmental conditions support coniferous forests as the dominant vegetation type. In the face of anthropogenic climate change, forests have a global role as potential sinks for atmospheric carbon (Goodale and others, 2002). Consequently, knowledge of the status of forests in the three large parks of the NCCN [that is, Mount Rainier (MORA), North Cascades (NOCA), and Olympic (OLYM) National Parks] is fundamental to understanding the condition of Pacific Northwest ecosystems. Diverse climate and soil properties across the Pacific Northwest result in a variety of forest types (Franklin and Dyrness, 1973; Franklin and others, 1988; Henderson and others, 1989, 1992). The mountainous terrain of Mount Rainier, North Cascades, and Olympic National Parks create steep elevational and precipitation gradients within and among the parks: collectively, these parks span from sea level to more than 4,200 m; and include areas with precipitation from 90 to more than 500 cm. The resulting forests range from coastal rainforests with dense understories and massive trees draped with epiphytes; to areas with drought-adapted Ponderosa pines; to high-elevation subalpine fir forests interspersed with meadows just below treeline (table 1). These forests, in turn, are the foundation for other biotic communities constituting Pacific Northwest ecosystems.

Techniques and Methods

An initial assessment of plankton tow detection probabilities for dreissenid mussels in the western United States

Early detection of dreissenid mussels ( Dreissena polymorpha and D. rostriformis bugensis ) is crucial to mitigating the economic and environmental impacts of an infestation. Plankton tow sampling is a common method used for early detection of dreissenid mussels, but little is known about the sampling intensity required for a high probability of early detection using the method. We used implicit dynamic occupancy models to estimate plankton tow detection probabilities of dreissenid mussels from a long-term data set containing plankton tow samples collected across central and western United States. We fit models using a) the entire data set, including water bodies with unknown occupancy status in addition to heavily infested water bodies, b) a data subset that included water bodies with paired water temperature data, and c) a data subset that included water bodies with lower dreissenid densities. For the entire data set, we found that estimated detection probabilities varied by water body size and ranged from approximately 0.10 to 0.86. For the water temperature subset, we observed the same pattern between detection probability and water body size as we did for the full data but additionally found that the estimated detection probabilities were much higher when water temperatures were above 12 °C. For the lower dreissenid density subset, we found that the estimated probability of detecting dreissenid mussels with a single aggregated plankton tow sample was near zero. Given these estimates, we conclude that the number of aggregated plankton tow samples taken per water body in the data is far fewer than the number needed to ensure a high probability of detecting dreissenid mussels, especially if they are at low densities. We summarize the analyses with a discussion of plankton tow sampling protocol changes needed to improve estimates of dreissenid detection probabilities.

western United States

Refining sampling protocols for cavefishes and cave crayfishes to account for environmental variation

Subterranean habitats support a diverse array of organisms and represent imperative habitats in many conservation strategies; however, subterranean habitats are one of the most difficult environments to study. Accounting for variable sampling detection is necessary to properly evaluate conservation options for rare species such as karst and other groundwater organisms. New sampling methods, such as environmental DNA, show promise to improve stygobiont detection; however, sources of sampling bias are poorly understood. Therefore, our objective was to determine factors affecting detection probability of both visual and environmental DNA (eDNA) surveys for cavefishes and cave crayfishes. We sampled 40 sites across the Ozark Highlands ecoregion in Arkansas, Missouri, and Oklahoma, USA using visual and eDNA surveys. We used occupancy modeling to estimate the detection probability of the two taxa using both survey methods under varying environmental conditions. Overall, eDNA sampling resulted in higher detection probability for cavefishes when compared to visual surveys, whereas visual surveys typically had higher detection probability for cave crayfishes. Greater water volume at the time of sampling was related to lower detection using visual surveys for both taxa, but there was no relationship between eDNA detection and water volume. Detection probability of both cavefishes and crayfishes was higher using visual surveys when sampling units were classified by coarse rather than fine substrate, whereas detection of cave crayfishes surveyed using eDNA was higher in coarse substrate environments. Detection of cavefishes and cave crayfishes was higher via eDNA sampling when water was flowing, but similar sampling conditions resulted in lower detection using visual surveys. Our results indicate detection should be considered when sampling stygobionts even if using traditional visual surveys. Environmental DNA is a useful tool; however, the limitations we identified indicate eDNA for these taxa currently are not adequate to replace traditional surveys in subterranean environments.

Arkansas, Missouri, Oklahoma

The potential of semi-structured citizen science data as a supplement for conservation decision-making: Validating the performance of eBird against targeted avian monitoring efforts

Methods are being developed to capitalize on citizen science data for research and monitoring, but these data are rarely used within established decision-making frameworks of wildlife agencies. Citizen science data are often collected at higher resolution and extent than targeted monitoring programs, and may provide complementary information. Here, we demonstrate that carefully filtered semi-structured citizen science observations, when paired with targeted survey data, can produce ecological predictions at higher resolution and extent than targeted surveys alone, and both datasets can represent complementary aspects of species' ecology. We present case studies demonstrating how citizen science data can enhance or supplement decision-making of government and conservation organizations. First, we show how the continuous spatial coverage of citizen science projects, when coupled with targeted surveys, can improve estimates of metrics used by the U.S. Fish and Wildlife Service in regulatory processes to estimate population size, and inform take limits of federally managed species nationwide. Second, we show that the spatial coverage of citizen science accommodates dynamic avian space use patterns during key times of the year, relative to standardized monitoring protocols carried out by the Illinois Natural History Survey. Lastly, we demonstrate that citizen science information can replicate estimates of migratory chronologies for the Illinois Natural History Survey and the U.S. Fish and Wildlife Service for some waterfowl species, and in some contexts can supplement missing data on abundance. These findings illustrate the value of integrating validated information from semi-structured citizen science into the current evidence base used to justify, inform, and evaluate conservation decision-making.

California, Illinois, Iowa, Missouri

The Amphibian Research and Monitoring Initiative (ARMI): 5-year report

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.

Scientific Investigations Report

Occurrence of fecal-indicator bacteria and protocols for identification of fecal-contamination sources in selected reaches of the West Branch Brandywine Creek, Chester County, Pennsylvania

The presence of fecal-indicator bacteria indicates the potential presence of pathogens originating from the fecal matter of warm-blooded animals. These pathogens are responsible for numerous human diseases ranging from common diarrhea to meningitis and polio. The detection of fecal-indicator bacteria and interpretation of the resultant data are, therefore, of great importance to water-resource managers. Current (2005) techniques used to assess fecal contamination within the fluvial environment primarily assess samples collected from the water column, either as grab samples or as depth- and (or) width-integrated samples. However, current research indicates approximately 99 percent of all bacteria within nature exist as attached, or sessile, bacteria. Because of this condition, most current techniques for the detection of fecal contamination, which utilize bacteria, assess only about 1 percent of the total bacteria within the fluvial system and are, therefore, problematic. Evaluation of the environmental factors affecting the occurrence and distribution of bacteria within the fluvial system, as well as the evaluation and modification of alternative approaches that effectively quantify the larger population of sessile bacteria within fluvial sediments, will present water-resource managers with more effective tools to assess, prevent, and (or) eliminate sources of fecal contamination within pristine and impaired watersheds. Two stream reaches on the West Branch Brandywine Creek in the Coatesville, Pa., region were studied between September 2002 and August 2003. The effects of sediment particle size, climatic conditions, aquatic growth, environmental chemistry, impervious surfaces, sediment and soil filtration, and dams on observed bacteria concentrations were evaluated. Alternative approaches were assessed to better detect geographic sources of fecal contamination including the use of turbidity as a surrogate for bacteria, the modification and implementation of sandbag bacteria samplers, and the use of optical brighteners. For the purposes of this report, sources of bacteria were defined as geographic locations where elevated concentrations of bacteria are observed within, or expected to enter, the main branch of the West Branch Brandywine Creek. Biologic sources (for example, waterfowl) were noted where applicable; however, no specific study of biologic sources (such as bacterial source tracking) was conducted. Data indicated that specific bacterial populations within fluvial sediments could be related to specific particle-size ranges. This relation is likely the result of the reduced porosity and permeability associated with finer sediments and the ability of specific bacteria to tolerate particular environments. Escherichia coli (E. coli) showed a higher median concentration (2,160 colonies per gram of saturated sediment) in the 0.125 to 0.5-millimeter size range of natural sediments than in other ranges, and enterococcus bacteria showed a higher median concentration (61,830 colonies per gram of saturated sediment) in the 0.062 to 0.25-millimeter size range of natural sediments than in other ranges. There were insufficient data to assess the particle-size relation to fecal coliform bacteria and (or) fecal streptococcus bacteria. Climatic conditions were shown to affect bacteria concentrations in both the water column and fluvial sediments. Drought conditions in 2002 resulted in lower overall bacteria concentrations than the more typically wet year of 2003. E. coli concentrations in fluvial sediment along the Coatesville study reach in 2002 had a median concentration of 92 colonies per gram of saturated sediment; in 2003, the median concentration had risen to 4,752 colonies per gram of saturated sediment. Symbiotic relations between bacteria and aquatic growth were likely responsible for increased bacteria concentrations observed within an impoundment area on the Coatesville study reach. This reach showed evidence of elevated aquatic growth and sharp increases in E. coli concentrations from upstream to downstream through the impoundment area in both 2002 and 2003. In 2003, E. coli concentrations within the waters column increased from 940 colonies per 100 milliliters upstream to 6,000 colonies per 100 milliliters at the dam crest. Given that these bacteria likely resulted from natural bacterial regrowth, the use of E. coli as an indicator of fecal contamination was severely impaired. Variable environmental conditions along the West Branch Brandywine Creek made the common field-chemical parameters of specific conductance, temperature, pH, and dissolved oxygen ineffective and (or) impossible to use for the determination of inputs of fecal contamination. Extreme variations in chemical gradients commonly were related to the urban/industrial signature of the watershed. For example, during base-flow sampling in 2002, specific-conductance values exceeding 1,000 microsiemens per centimeter observed in effluent from a local steel mill. This effluent raised the specific conductance within the West Branch Brandywine from just above 200 microsiemens per centimeter upstream from the outfall to just below 500 microsiemens per centimeter downstream from the outfall. These chemical gradients also, likely, had an effect on the initial colonization of bacteria, the formation of biofilms, and the persistence of certain types of bacteria along the study reach. Data collected in 2003 indicated that nutrients increased during both base-flow and stormflow conditions along the Coatesville study reach. For example, during base-flow sampling in 2003, 20 pounds of phosphorus was shown to enter the West Branch Brandywine Creek along the Coatesville study reach. The largest contributors to this base-flow nutrient load were likely two wastewater-treatment facilities adjacent to the study reach. During stormflow sampling in 2003, 480 pounds of phosphorus was shown to enter the West Branch Brandywine Creek along the Coatesville study reach. Data, along with other research, indicated the largest contributor to this stormflow nutrient load was likely remobilized sediment originating from a large dam impoundment. These elevated nutrient concentrations were considered sufficient to promote accelerated aquatic growth along the reach. Data collected in 2003 showed that wastewater constituents entered the West Branch Brandywine Creek largely from urban storm-sewer systems. Samples from the primary storm sewer for the city of Coatesville had detections for 20 of 69 wastewater constituents. These constituents included both strong and weak indicators of fecal contamination and generally indicated the storm-sewer system along the Coatesville study reach was a likely source of fecal-indicator bacteria and fecal contamination under base-flow conditions. By comparison, 5 constituents were detected in samples from the upstream end of the reach, and 10 constituents were detected in samples from the downstream end of the reach. During stormflow, numbers of detections were similar along the entire length of the study reach-five in samples from the upstream end, eight in samples from the center of the reach, and seven in samples from the downstream end of the reach. These data indicate that point sources (such as culverts and pipes, septic systems, and wastewater-treatment facilities) are not likely the origin of bacteria contamination during stormflow. The bacteria concentrations observed during stormflow events probably result from remobilized sessile bacteria stored within fluvial sediments. In this case, these bacteria should not be considered indicators of current fecal contamination. Impervious surfaces were found to increase bacteria concentrations along the West Branch Brandywine Creek because contaminated runoff from impervious areas generally flows into, and is concentrated within, the confines of the local storm-sewer system. During 2002, storm-sewer outfalls draining impervious areas were associated with all major locations of elevated bacterial concentrations (greater than 1,200 colonies per gram of saturated sediment) in fluvial sediments. During 2003, wetter conditions and overall bacteria concentrations higher than in 2002 resulted in point sources of bacterial contamination becoming less pronounced; however, the storm-sewer system, draining adjacent impervious areas, was still observed to be the primary source of bacteria along the reach. Where stormwater and (or) other runoff from these areas was allowed to infiltrate and (or) flow through wetland and riparian buffers, bacteria concentrations were not observed to be elevated above background levels commonly observed throughout similar areas of the same reach. Two run-of-the-river dams along the Coatesville study reach were evaluated for their effects on observed bacterial concentrations. These dams were shown to have greater or lesser effects on bacterial concentrations depending on the size of the structure and the capacity of the structure to impede flows. The smaller upstream dam had an approximate height of 3 feet and showed little observed effect on measured turbidity values; these data indicated that the dam did not effectively impede the flow of water or sediment within the West Branch Brandywine Creek. Consequently, this small dam did not show any observed effect on bacterial concentrations either upstream or downstream of the structure. The larger dam, near the middle of the reach, had an approximate height of 20 feet and showed greater effects on both turbidity and bacteria concentrations. The capacity of the larger dam to impede flows, combined with nutrients entering the reach, resulted in increased biologic activity throughout the impoundment area. Within this larger impoundment, enterococcus bacteria populations were observed to decrease sharply and E. coli bacteria populations were observed to increase sharply as flow approached the dam crest. All bacteria levels were then observed to drop to background levels, in both the water column and fluvial sediment, immediately downstream from the dam crest. Additional study is required to determine the cause for this rapid die off. Turbidity was assessed as a potential surrogate for E. coli bacteria. Regression analysis indicated higher turbidity levels usually can indicate higher concentrations of bacteria (R2 = 0.67), but the relation was too sporadic on the West Branch Brandywine Creek to use turbidity as a surrogate for estimated bacteria concentrations. Evaluation of data from individual base-flow and stormflow events resulted in variable and generally poor statistical relations between E. coli bacteria and turbidity (R2 values ranged from 0.02 to 0.94). Sandbag samplers were used in 2003 to determine their suitability for the assessment of fecal contamination. Sandbag samplers rely on the ability of bacteria to attach to surfaces and use the larger sessile bacteria populations instead of the more commonly used planktonic bacteria populations. E. coli bacteria concentrations observed in the sandbag samplers, after 1 week in place, were similar to those found within natural sediments collected concurrently. Enterococcus bacteria concentrations within the same sandbag samplers were not similar, and were generally lower, than those observed within the natural sediments. This discrepancy was likely because sand within the samplers was sieved to a size that was likely too coarse for enterococcus bacteria to persist. Optical-brightener samplers were installed along with each sandbag sampler. Optical brighteners are additives used in common household detergents; therefore, detection of optical brighteners, along with elevated fecal-indicator bacteria concentrations, strongly indicates a link to humans. Positive results for optical brighteners were detected only at the outfalls of two sewage-treatment facilities; because of treatment of the effluent from these facilities, these samples did not have elevated bacteria concentrations. The lack of additional positive results was largely because this method is not sensitive to low concentrations of optical brighteners.

Pennsylvania