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Amphibians and reptiles of Guyana, South America: illustrated keys, annotated species accounts, and a biogeographic synopsis

Guyana has a very distinctive herpetofauna. In this first ever detailed modern accounting, based on voucher specimens, we document the presence of 324 species of amphibians and reptiles in the country; 148 amphibians, 176 reptiles. Of these, we present species accounts for 317 species and color photographs of about 62% (Plates 1–40). At the rate that new species are being described and distributional records are being found for the first time, we suspect that at least 350 species will be documented in a few decades. The diverse herpetofauna includes 137 species of frogs and toads, 11 caecilians, 4 crocodylians, 4 amphisbaenians, 56 lizards, 97 snakes, and 15 turtles. Endemic species, which occur nowhere else in the world, comprise 15% of the herpetofauna. Most of the endemics are amphibians, comprising 27% of the amphibian fauna. Type localities (where the type specimens or scientific name-bearers of species were found) are located within Guyana for 24% of the herpetofauna, or 36% of the amphibians. This diverse fauna results from the geographic position of Guyana on the Guiana Shield and the isolated highlands or tepuis of the eastern part of the Pantepui Region, which are surrounded by lowland rainforest and savannas. Consequently, there is a mixture of local endemic species and widespread species characteristic of Amazonia and the Guianan Region. Although the size of this volume may mislead some people into thinking that a lot is known about the fauna of Guyana, the work has just begun. Many of the species are known from fewer than five individuals in scientific collections; for many the life history, distribution, ecology, and behavior remain poorly known; few resources in the country are devoted to developing such knowledge; and as far as we are aware, no other group of animals in the fauna of Guyana has been summarized in a volume such as this to document the biological resources. We briefly discuss aspects of biogeography, as reflected in samples collected at seven lowland sites (in rainforest, savanna, and mixed habitats below 500 m elevation) and three isolated highland sites (in montane forest and evergreen high-tepui forest above 1400 m elevation). Comparisons of these sites are preliminary because sampling of the local faunas remains incomplete. Nevertheless, it is certain that areas of about 2.5 km2 of lowland rainforest can support more than 130 species of amphibians and reptiles (perhaps actually more than 150), while many fewer species (fewer than 30 documented so far) occur in a comparable area of isolated highlands, where low temperatures, frequent cloudiness, and poor soils are relatively unfavorable for amphibians and reptiles. Furthermore, insufficient study has been done in upland sites of intermediate elevations, where lowland and highland faunas overlap significantly, although considerable work is being accomplished in Kaieteur National Park by other investigators. Comparisons of the faunas of the lowland and isolated highland sites showed that very few species occur in common in both the lowlands and isolated highlands; that those few are widespread lowland species that tolerate highland environments; that many endemic species (mostly amphibians) occur in the isolated highlands of the Pakaraima Mountains; and that each of the isolated highlands, lowland savannas, and lowland rainforests at these 10 sites have distinctive faunal elements. No two sites were identical in species composition. Much more work is needed to compare a variety of sites, and especially to incorporate upland sites of intermediate elevations in such comparisons. Five species of sea turtles utilize the limited areas of Atlantic coastal beaches to the northwest of Georgetown. All of these are listed by the International Union for the Conservation of Nature as being of global concern for long-term survival, mostly owing to human predation. The categories of Critically Endangered or Endangered are applied to four of the local sea turtles (80%). It is important to protect the few good nesting beaches for the sea turtles of Guyana. We have documented each of the species now known to comprise the herpetofauna of Guyana by citing specimens that exist in scientific collections, many of which were collected and identified by us and colleagues, including students of the University of Guyana (UG). We also re-identified many old museum specimens collected by others in the past (e.g., collections of William Beebe) and we used documented publications and collection records of colleagues, most of whom have been working more recently. We present dichotomous keys for identifying representatives of the species known to occur in Guyana, and we present brief annotated species accounts. The accounts provide the current scientific name, original name (with citation of the original description, which we personally examined in the literature), some outdated names used in the recent past, type specimens, type localities, general geographic distribution, examples of voucher specimens from Guyana, coloration in life (and often a color photograph), and comments pointing out interesting subjects for future research.

Proceedings of the Biological Society of Washingto

The potential influence of changing climate on the persistence of salmonids of the inland west

The Earth's climate warmed steadily during the 20th century, and mean annual air temperatures are estimated to have increased by 0.6°C (Intergovernmental Panel on Climate Change, 2007). Although many cycles of warming and cooling have occurred in the past, the most recent warming period is unique in its rate and magnitude of change (Siegenthaler and others, 2005) and in its association with anthropogenic emissions of greenhouse gases (Intergovernmental Panel on Climate Change , 2007). The climate in the western United States warmed in concert with the global trend but at an accelerated rate (+0.8°C during the 20th century; Saunders and others, 2008). The region could also prove especially sensitive to future changes because the relatively small human population is growing rapidly, as are demands on limited water supplies. Regional hydrological patterns are dominated by seasonal snow accumulation at upper elevations. Most of the region is relatively dry, and both terrestrial and aquatic ecosystems are strongly constrained b y water availability (Barnett and others, 2008; Brown and others, 2008). Stream environments are dynamic and climatically extreme, and salmonid fishes are the dominant elements of the native biodiversity (McPhail and Lindsey, 1986; Waples and others, 2008). Salmonids have broad economic and ecologic importance, but a century of intensive water resource development, nonnative fish stocking, and land use has significantly reduced many populations and several taxa are now protected under the Endangered Species Act (Thurow and others, 1997; Trotter, 2008). Because salmonids require relatively pristine, cold water environments and are often isolated in headwater habitats, members of this group may be especially vulnerable to the effects of a warming climate (Keleher and Rahel, 1996; Rieman and others, 2007; Williams and others, 2009). Warming during the 20th century drove a series of environmental trends that have profound implications for many aspects of salmonid habitat, including disturbance regimes such as wildfire, and unfavorable changes to thermal and hydrologic properties of aquatic systems. Warmer air temperatures have been associated with decreased winter snow accumulations, have accelerated snowmelt, and have advanced the timing of peak runoff by several days to weeks across most of western North America (Stewart and others, 2005; Barnett and others, 2008). Less snow and earlier runoff decrease aquifer recharge, make less water available for groundwater inputs to streams, and are contributing to widespread decreases in summer low flows (Stewart and others, 2005; Rood and others, 2008; Luce and Holden 2009). Interannual variability in stream flow is increasing, as is the persistence of multi-year extreme conditions (McCabe and others, 2004; Pagano and Garen 2005). In many areas of western North America, flood risks have increased in association with warmer temperatures during the 20th century (Hamlet and Lettenmaier, 2005). Streams where midwinter temperatures are near freezing have proven especially sensitive to increased flooding because of associated transitional hydrological patterns (mixtures of rainfall and snowmelt) and propensity for occasional rain-on-snow events to rapidly melt winter snowpack and generate large floods (Hamlet and Lettenmaier, 2005). Stream temperatures in many areas are increasing (Peterson and Kitchell, 2001; Morrison and others, 2002; Bartholow, 2005; Kaushal and others, 2010), due to both air temperature increases and reduced summer flows that make streams more sensitive to warmer air temperatures (Isaak and others, 2010). In recent decades, wildfires have become more common across much of the western United States during periods of more frequent droughts (Westerling and others, 2006; Hoerling and Eischeid, 2007), and local stream temperature can increase in postfire environments (Gresswell, 1999; Dunham and others, 2007). Fire-related temperature increase within streams is commonly a transient phenomenon, lasting only until riparian vegetation has recovered (Gresswell, 1999); however, ongoing climate change could preclude recovery to higher stature, prefire vegetation types in some areas (McKenzie and others, 2004; van Mantgem and Stephenson, 2007), resulting in a loss of critical riparian shading. Additionally, when wildfires occur in steep mountain topographies, the vegetation that stabilize s soils on hillslopes is often killed and landslides become more prevalent (Gresswell, 1999). Landslides int o stream channels form debris flows composed of sediment slurries and dead trees that can scour channels to bedrock and further exacerbate stream heating, delay recovery of riparian areas, or extirpate fish populations (Gresswell, 1999; May and Gresswell, 2003; Dunham and others, 2007). Changes in stream environments will shift habitat distributions, sometimes unpredictably, in both time and space for many salmonid fishes. Water temperature fundamentally influences aquatic ecosystem health because distribution, reproduction, fitness, and survival of ectothermic organisms are inextricably linked to the thermal regime of the environment. Historically, research has focused on defining lethal thermal limits of salmonids (Eaton and others, 1995; Selong and others, 2001; Todd and others, 2008); however, water temperature is known to be important in biological processes at a variety of spatial scales and levels of biological organization (Rahel and Olden, 2008; McCullough and others, 2009). For instance, trout are affected directly by water temperature through feeding, metabolism, and growth rates, and indirectly by factors such as prey availability and species interactions (Wehrly and others, 2007; Rahel and Olden, 2008). Where cold water temperatures currently limit habitat suitability and distributions of some species (for example, at the highest and most northerly distributional extents; Nakano and others, 1996; Coleman and Fausch, 2007), a warming climate may gradually increase the quality and extent of suitable habitat. Over time, previously constrained populations are expected to expand into these new habitats and increase in number. Some evidence suggests this may already be happening in Alaska, where streams in recently deglaciated areas are being colonized by emigrants from nearby salmon and char populations (Milner and others, 2000). Unfortunately, many of the sensitive salmonid species that are often the focus of western managers are unlikely to benefit from future water temperature increases. Warmer stream temperatures will facilitate invasion by nonnative species that are broadly established in downstream areas into upstream areas where they will compete with native species (Rieman and others, 2006; Rahel and Olden, 2008; Fausch and others, 2009). In other cases, warmer stream temperatures will render thermally suitable habitats unsuitable in downstream areas and effect net losses of habitat because upstream distributions are often constrained by streams that are too small or steep (Hari and others, 2006; Isaak and others, 2010). Both scenarios are realistic for fish species like bull trout ( Salvelinus confluentus ) (Rieman and others, 2006; Rieman and others, 2007), the various subspecies of cutthroat trout ( Oncorhynchus clarkii ) (Williams and others, 2009), Gila trout ( Oncorhynchus gilae gilae ) (Kennedy and others, 2008), and Apache trout ( Oncorhynchus gilae apache ) (Rinne and Minckley, 1985; Carmichael and others, 1993). As native species are increasingly confined to smaller and more isolated habitats by a gradually warming climate, the effects of wildfires (whether related to lethal changes in water quality during a fire, channel debris flows, or chronic postfire warming ) could have greater proportional effects on remaining habitats (for example, Brown and others, 2001; Rieman and others, 2007). If these changes were accompanied by additional hydrologic alterations associated with changes to the magnitude, frequency, duration, timing, and rate of change of discharge patterns (Jager and others, 1999; Henderson and others, 2000), populations may begin to lose some of their historic resilience and become ever more susceptible to local extirpations. As dramatic and extensive as climatic and environmental trends are for salmonid habitats, global climate models (GCMs) project that many of these trends will continue and even accelerate until at least the middle of the 21st century (Intergovernmental Panel on Climate Change, 2007). Current projections suggest mean annual air temperatures will increase by an additional 1–3°C, and early indications are that climate trajectory is at the higher end of this range (Pittock, 2006; Raupach and others, 2007). Although predicted changes vary considerably, even the most conservative estimates suggest a warming rate that will be twice that observed during the 20th century. Projections for the midcentury are most certainly due to the effects of greenhouse gases already emitted or predicted in the short term, uncertainties of the effects of longer-term greenhouse gas emissions, short-term climate cycles, and process errors associated with climate models (Cox and Stephenson, 2007). Projections of changes in total precipitation are less certain than those for air temperatures, but most GCMs project relatively small changes in the Northwest, with the exception of slightly drier summer periods (Mote and others, 2008; Karl and others, 2009). In the Southwest, however, significant decreases (such as 15–30 percent ) are projected during most periods of the year, and this area is one of the few for which Intergovernmental Panel on Climate Change (2007) precipitation projections have a high level of certainty (Hoerling and Eischeid, 2007; Karl and others, 2009). Clearly, managers of native salmonids in the wester n United States should consider adjusting management strategies to accommodate a warmer and possibly drier future (Williams and others, 2009). Tools are needed to forecast where important changes may occur and how conservation efforts should be prioritized. In this Open-File Report, we document our initial efforts in this regard for 10 species and subspecies of inland trout and Montana Arctic grayling ( Thymallus arcticus ) across the western United States.

Arizona, California, Colorado, Idaho, Montana, Nev

A Framework for Long-term Ecological Monitoring in Olympic National Park: Prototype for the Coniferous Forest Biome

This report is the result of a five-year collaboration between scientists of the U.S. Geological Survey Forest and Rangeland Ecosystem Science Center, Olympic Field Station, and the natural resources staff of Olympic National Park to develop a comprehensive strategy for monitoring natural resources of Olympic National Park. Olympic National Park is the National Park Serviceʼs prototype monitoring park, representing parks in the coniferous forest biome. Under the umbrella of the National Park Serviceʼs prototype parks program, U.S. Geological Survey and Olympic National Park staffs are obligated to: develop strategies and designs for monitoring the long-term health and integrity of national park ecosystems with a significant coniferous forest component. design exportable monitoring protocols that can be used by other parks within the coniferous forest biome (i.e., parks having similar environments), and create a demonstration area and ʻcenter of excellenceʼ for assisting other parks in developing ecological monitoring programs. Olympic National Park is part of the North Coast and Cascades Network, a network of seven Pacific Northwestern park units created recently by the National Park Serviceʼs Inventory and Monitoring Program to extend the monitoring of ʻvital signsʼ of park health to all National Park Service units. It is our intent and hope that the monitoring strategies and conceptual models described here will meet the overall purpose of the prototype parks monitoring program in proving useful not only to Olympic National Park, but also to parks within the North Coast and Cascades Network and elsewhere. Part I contains the conceptual design and sampling framework for the prototype long-term monitoring program in Olympic National Park. In this section, we explore key elements of monitoring design that help to ensure the spatial, ecological, and temporal integration of monitoring program elements and discuss approaches used to design an ecosystem-based monitoring program. Basic monitoring components include ecosystem drivers, (e.g., climate, atmospheric inputs, human pressures), indicators of ecosystem integrity (e.g., biogeochemical indicators), known threats (e.g., impacts of introduced mountain goats), and focal or ʻkeyʼ species (e.g., rare or listed species, Roosevelt elk). Monitoring system drivers and key indicators of ecosystem integrity provide the long-term baseline needed to judge what constitutes ʻunnaturalʼ variation in park resources and provide the earliest possible warning of unacceptable change. Monitoring effects of known threats and the status of focal species will provide information useful to park managers for dealing with current park issues. In Part I we describe the process of identifying potential indicators of ecological condition and present conceptual models of park ecosystems. In addition we report results from several workshops held in conjunction with Olympic National Park aimed at identifying potential indicators of change in the parkʼs ecosystem. First, we describe the responses of Olympic National Park staff to the generic question, “What is the most important resource to monitor in Olympic National Park and why?” followed by the responses from resource and land managers from areas adjoining the park. We also catalogue the responses of various expert groups that we asked to help identify the most appropriate system drivers and indicators of change in the Olympic National Park ecosystems. Results of the workshops provided the justification for selecting basic indicators of ecosystem integrity, effects of current threats to park resources, and focal resources of parks to detect both the currently evident and unforeseeable changes in park resources. We conclude Part I by exploring several generic statistical issues relevant to monitoring natural resources in Olympic National Park. Specifically we discuss trade-offs associated with sampling extensively versus sampling intensively in smaller geographic regions and describe a conceptual framework to guide development of a generic sampling frame for monitoring. We recommend partitioning Olympic National Park into three zones of decreasing accessibility to maximize monitoring efficiency. We present examples of how the generic sampling frame could be used to help ensure spatial integration of individual monitoring projects. Part II of the report is a record of the potential monitoring questions and indicators identified to date in our workshops. The presentation is organized according to the major system drivers, components, and processes identified in the intermediate-level working model of the Olympic National Park ecosystem. For each component of the park system, we develop the need and justification for monitoring, articulate park management issues, and describe key resources and ecosystem functions. We also present a pictorial conceptual model of each ecological subsystem, identify monitoring questions, and list potential indicators for each monitoring question. We conclude each section by identifying linkages of indicators to other ecological subsystems in our general ecosystem model, spatial and temporal contexts for monitoring (where and how often to monitor), and research and development needs. Part II represents the most current detailed listing of potential indicators—the material for subsequent discussions of monitoring priorities and selection of indicators for protocol development. Collectively, the sections of this report contain a comprehensive list of the important monitoring questions and potential indicators as well as recommendations for designing an integrated monitoring program. In Part I, Chapter 6 we provide recommendations on how to proceed with the important next steps in the design process: establishing priorities among the many possible monitoring questions and indicators, and beginning to research and design effective long-term monitoring protocols.

Information and Technology Report

Understanding the influence of predation by introduced fishes on juvenile salmonids in the Columbia River Basin: Closing some knowledge gaps. Interim Report of Research 2010

Project overview Predation on juvenile salmonids by fish in the Columbia River Basin (CRB) has impacted salmon survival and is a topic that has received considerable attention over the last three decades. Some of the earliest and most detailed research focused on the food habits, consumption rates, abundance, and distribution of predaceous northern pikeminnow Ptychocheilus oregonensis , smallmouth bass Micropterus dolomieu , walleye Sander vitreus , and channel catfish Ictalurus punctatus in John Day Reservoir (Beamesderfer and Rieman 1991; Poe et al. 1991; Vigg et al. 1991). This group of researchers also estimated the loss of juvenile salmonids to predation by some of these predators (Rieman et al. 1991). Since this pioneering effort, others have evaluated various aspects of predation-related mortality on juvenile salmonids in the CRB, focusing mostly on northern pikeminnow and smallmouth bass (e.g., Tabor et al. 1993; Zimmerman 1999; Naughton et al. 2004). Perhaps the most significant finding coming from this body of research was that the native northern pikeminnow was the dominant predator of juvenile salmonids in the CRB. Indeed, Beamesderfer et al. (1996) estimated that northern pikeminnow consumed about 16 million (8%) of the estimated 200 million juvenile salmonids emigrating annually in the CRB, far surpassing the consumption of smallmouth bass, walleye, and channel catfish combined. Because of this, large-scale management fisheries (i.e., the northern pikeminnow management program, or NPMP; see Rieman and Beamesderfer 1990; Beamesderfer et al. 1996) have been implemented in the CRB since 1990 to achieve a 10%–20% exploitation rate on northern pikeminnow and reduce predation on juvenile salmonids. The NPMP has been a success, resulting in up to 38% potential reductions in predation (Friesen and Ward 1999; Knutsen and Ward 1999; Ward and Zimmerman 1999;). In contrast to the NPMP, Oregon and Washington state fish and wildlife agencies manage and enhance recreational fisheries for smallmouth bass and walleye by implementing size and harvest limit regulations. Recently, many biologists and fish managers have become concerned about the impact of non-native predaceous fishes on juvenile salmonid survival. For example, Poe et al. (1994) warned that smallmouth bass, walleye, and channel catfish were expanding their populations in some areas, that these fish could be significant predators on juvenile salmonids, and that they may compete with northern pikeminnow for common prey items, resulting in higher consumption rates of salmonids by the native predator. Sanderson et al. (2009) reported that the impact of non-indigenous species (including piscivorous fishes) on salmon survival within the CRB can be severe and suggested that managing nonindigenous species may be imperative for salmon recovery. Assessing the current ecological impacts of introduced fishes throughout the CRB will fill information gaps associated with their impact on salmonid survival and contribute to the description of CRB food webs. In response to these recent concerns about the potential predatory impact of non-native piscivores on salmon survival, the Bonneville Power Administration (BPA) and the Columbia Basin Fish and Wildlife Authority (CBFWA) co-hosted a workshop to address predation on juvenile salmonids in the CRB by non-native fish (Halton 2008). The purpose of the workshop was to review, evaluate, and develop strategies to reduce predation by non-native fishes on juvenile salmonids. In the end, discussion at the workshop and at subsequent meetings considered two potential ideas to reduce predation by non-native fish on juvenile salmonids; (1) understanding the role of juvenile American shad Alosa sapidissima in the diet of non-native predators in the fall; and (2) the effects of localized, intense reductions of smallmouth bass in areas of particularly high salmonid predation. In this report, we describe initial efforts to understand the influence of juvenile American shad as a prey item for introduced predators in the middle Columbia River. Our first objective, addressed in Chapter 1, was to evaluate the efficacy of nonlethal methods to describe the physiological condition of smallmouth bass, walleye, and channel catfish from late summer through late fall. Such information will be used to understand the contribution of juvenile American shad to the energy reserves of predaceous fish prior to winter. In Chapter 2, we describe the results of some limited sampling to document the food habits of smallmouth bass, walleye, and channel catfish in three reservoirs of the middle Columbia River during late fall. Collectively, we hope to increase our understanding of the contribution of juvenile American shad to the diets of introduced predators and the contribution of this diet to their energy reserves, growth, and perhaps over-winter survival. Managers should be able to use this information for deciding whether to control the population of American shad in the CRB or for managing introduced predaceous fish in the CRB.

Oregon, Washington

Southern Salish Sea Habitat Map Series: Admiralty Inlet

In 2010 the Environmental Protection Agency, Region 10 initiated the Puget Sound Scientific Studies and Technical Investigations Assistance Program, designed to support research in support of implementing the Puget Sound Action Agenda. The Action Agenda was created in response to Puget Sound having been designated as one of 28 estuaries of national significance under section 320 of the U.S. Clean Water Act, and its overall goal is to restore the Puget Sound Estuary's environment by 2020. The Southern Salish Sea Mapping Project was funded by the Assistance Program request for proposals process, which also supports a large number of coastal-zone- and ocean-management issues. The issues include the recommendations of the Marine Protected Areas Work Group to the Washington State Legislature (Van Cleve and others, 2009), which endorses a Puget Sound and coast-wide marine conservation needs assessment, gap analysis of existing Marine Protected Areas (MPA) and recommendations for action. This publication is the first of four U.S. Geological Survey Scientific Investigation Maps that make up the Southern Salish Sea Mapping Project. The remaining three map blocks to be published in the future, located south of Admiralty Inlet, are shown in figure 1. Puget Sound is a deep, fjord-type estuary covering an area of 2,330 km 2 in the Pacific Northwest region of the United States (fig. 1). It is connected to the ocean by the Strait of Juan de Fuca, a turbulent passage approximately 160 km in length and 22 km wide at its west end, expanding to over 40 km wide at its east end (Thomson, 1994). During the Pleistocene, the area was occupied several times by lobes of continental ice, resulting in a complex basin-fill of glacial and interglacial deposits that are locally as thick as 1100 m (Johnson and others, 2001). The last glaciation, called the Fraser glaciation, began after 28,800±740 14 C yr B.P. when ice started a slow expansion (Clague, 1981). At peak advance the westward Juan de Fuca lobe reached the edge of the continental shelf through the Juan de Fuca Strait shortly before 14,460±200 14 C yr B.P. (Herzer and Bornhold, 1982). The southward Puget lobe advanced to its terminal position in Puget Sound by around 14,150 14 C yr B.P. (Porter and Swanson, 1998). Ice retreated from its maximum to northern Whidbey Island by 13,650±350 14 C yr B.P. (Dethier and others, 1995). Retreating glaciers resulted in a thick sequence of ice-contact, glacial-marine sediment, and early post-glacial sediments (Linden and Schurrer, 1988). These deposits have experienced the effects of a marine transgression followed by regression, resulting in a sea-level several tens of meters lower than the present day (Linden and Schurrer, 1988). A second transgression brought sea level to about the present level by around 5,470±120 14 C yr B.P. (Clague and others, 1982) establishing the present oceanographic and geologic environment Puget Sound is separated into four interconnected basins; Whidbey, Central (Main), Hood Canal, and South (Thomson, 1994). The Whidbey, Central, and Hood Canal basins are the three main branches of the Puget Sound estuary and are separated from the Strait of Juan de Fuca by a double sill at Admiralty Inlet. The Admiralty Inlet map area includes the Inlet and a portion of the Whidbey Basin (fig. 1). The shallower South Basin is separated by a sill at Tacoma Narrows and is highly branched with numerous finger inlets. Flow within Puget Sound is dominated by tidal currents of as much as 1 m/s at Admiralty Inlet, reducing to approximately 0.5 m/s in the Central Basin (Lavelle and others, 1988). The lack of silt and clay-sized sediments in the Admiralty Inlet map area is likely a result of the strong currents (see Ground-Truth Studies for the Admiralty Inlet Map Area, sheet 3). The subtidal component of flow reaches approximately 0.1 m/s and is driven by density gradients arising from the contrast in salty ocean water at the entrance and freshwater inputs from stream flow (Lavelle and others, 1988). The total freshwater input to Puget Sound is approximately 3.4 x 10 6 m 3 /day, primarily from the Skagit River (Cannon, 1983). The subtidal circulation mostly consists of a two-layered flow in the basins with fresher water exiting at the surface and saltier water entering at depth (Ebbesmeyer and Cannon, 2001). In general, surface waters flow north and deeper waters flow south; variations arise from wind effects that can drive a surface current in the same direction as the wind, and a baroclinic response in the lower layer to about 100-m depth (Matsuura and Cannon, 1997). Oceanographic properties are influenced by temporal forcing parameters such as reduced stream flow during the 2000-01 drought that increased surface salinity and decreased differences between surface and bottom waters (Newton and others, 2003). On offshore seismic-reflection profiles, Pleistocene strata (excluding latest Pleistocene glacial and post-glacial deposits) form a distinct seismic unit, bounded below by pre-Tertiary or Tertiary basement and above by typically flat-lying latest Pleistocene to Holocene deposits that fill in erosional or depositional relief (Johnson and others, 2001). Cores from central Puget Sound have accumulation rates that range from 85 to 1200 mg/cm 2 /yr, or 0.12 to 2.4 cm/yr; the highest accumulation rates are near the southern end of central Puget Sound (Carpenter and others, 1985). Carpenter and others (1985) un-weighted arithmetic mean of accumulation rates for central Puget Sound deeper stations is 480±340 (± one standard deviation) mg/cm 2 /yr. Lavelle and others (1985) also found rates as high as 1200 mg/cm 2 /yr over the past approximately 70 years in cores in the Central Basin off of and north and south of Elliott Bay. Puget Sound basin rates are comparable to rates in midshelf silt deposits on the Washington coast north of the Columbia River (Nittrouer and others, 1979). The deep subtidal (in other words, below SCUBA depths) habitats of Puget Sound are relatively poorly known. A few subtidal surveys exist for several habitat types from the 1960s and 1970s (reviewed in Dethier, 1990), using grab and box core data. The Dethier (1990) review divides habitat up into Coast and Marine Ecological Classification Standard (CMECS) substrate, water column energy, and depth zones but does not attempt to map these habitats, rather it is an inventory of habitats found in the area and the flora and fauna associated with each habitat. The approach of the Southern Salish Sea Mapping project is to create highly detailed seafloor maps through collection, integration, interpretation, and visualization of swath sonar data (the undersea equivalent of satellite remote-sensing data in terrestrial mapping), acoustic backscatter, seafloor video, seafloor photography, and bottom-sediment sampling data. This approach is based in part on methods presented and data collection and product needs identified at the Washington State Seafloor Mapping Workshop (Washington State Seafloor Mapping Workshop Steering Committee, 2008), attended by coastal and marine managers and scientists. The map products display seafloor geomorphology and substrate, and identify potential marine benthic habitats. It is emphasized that the more interpretive habitat and geology maps rely on the integration of multiple, new high-resolution datasets and that mapping at small scales would not be possible without such data. Oceanographic current and wave data is not included in this analysis, however, the accompanying geographic information system (GIS) data set is designed and intended to be combined with oceanographic and biologic data sets assembled by others in the future and some of the GIS data has already been incorporated in the unpublished Nature Conservancy Benthic Habitats of Puget Sound database. This publication includes four map sheets, explanatory text, and a descriptive pamphlet. Each map sheet is published as a portable document format (PDF) file. ESRI ArcGIS compatible geotiffs (for example, bathymetry) and shapefiles (for example video observation points) will be available for download in the data catalog associated with this publication (Cochrane, 2015). An ArcGIS Project File with the symbology used to generate the map sheets is also provided. For those who do not own the full suite of ESRI GIS and mapping software, the data can be read using ESRI ArcReader, a free viewer that is available at http://www.esri.com/software/arcgis/arcreader/index.html .

Washington

Evaluating management alternatives for Wyoming elk feedgrounds in consideration of chronic wasting disease

Executive Summary The authors used decision and modeling analyses to evaluate management alternatives for a decision on whether to permit Cervus canadensis (elk) feeding on two sites on Bridger-Teton National Forest, Dell Creek and Forest Park. Supplemental feeding of elk could increase the transmission of chronic wasting disease (CWD) locally and disease spread regionally, potentially impacting elk populations over time with wider implications for Odocoileus hemionus (mule deer) and Odocoileus virginianus (white-tailed deer) populations and hunting, tourism, and regional revenue. Supplemental feeding is thought to improve overwinter elk survival and reduce the commingling of elk with cattle during months when brucellosis transmission risk is highest. We worked with the U.S. Department of Agriculture Forest Service to identify their fundamental objectives and associated performance metrics related to this feedground decision. We then developed disease and habitat selection models to quantify the effect of four management alternatives on select performance metrics. The four alternatives were to continue to permit feeding, phaseout permits to feed in three years, permit feeding on an emergency basis, or stop permitting feeding. In this report, we present methods and summarized results on disease and habitat selection models and summaries of other performance metrics analyzed by BIO-WEST, Inc. and Cirrus Ecological Solutions as part of an Environmental Impact Statement. Data from Wyoming Game and Fish Department (WGFD) supported the assumption that supplemental elk feeding allows for larger elk populations in a region. We documented that herd units (HU) without feedgrounds had 23 percent lower densities of elk per area of winter range when compared against HUs with feedgrounds, after accounting for differences in sightability of elk during counts on and off feedgrounds. Thus, throughout our analyses, we assumed feedground closures would reduce elk carrying capacity resulting in an average decline of previously fed elk population segments by 23 percent (5th and 95th percentiles = [11 percent, 35 percent]) by year 20. Most of that decline occurred within the first few years after a feedground ceases to operate. We used a panel of CWD experts to help estimate CWD trans-mission in fed and unfed elk population segments. In aggregate, the expert panel estimated that median values of direct and indirect transmission of CWD are expected to be 1.9 and 4 times higher, respectively, in fed elk populations compared to unfed elk. We used these disease transmission estimates in combination with local elk demographic rates and carrying capacity estimates to project disease and population dynamics. In year 20, we predicted CWD prevalence would increase to 42 percent (5th and 95th percentiles = [29 percent, 55 percent]), and 13 percent (5th and 95th percentiles = [4 percent, 26 percent]) on average for fed and unfed elk population segments, respectively, given a starting prevalence of 1.6 percent. The prevalence estimates for the unfed elk population segments are in the range of previous observations of CWD in elk in the western United States. The average CWD prevalence from 2016 to 2018 in the unfed elk population of Wind Cave National Park in South Dakota was 18 percent overall but up to 30 percent in some regions (Sargeant and others, 2021). Meanwhile, CWD prevalence in the Iron Mountain and Laramie Peak elk herds in Wyoming from 2016 to 2018 was 14 percent and 7 percent, respectively, despite being present since at least 2002 (Wyoming Game and Fish Department, 2020b). From 2016 to 2020, elk that were fed at Dell Creek and Forest Park constituted on average 12–20 percent of the total elk on their respective HUs. As a result, the differences between management alternatives are modest when considering the closure of only one feedground on a HU. The no feeding alternative for Forest Park resulted in a CWD prevalence of 17 percent (SD = 7 percent) in the Afton HU compared to 20 percent (SD = 7 percent) with continued feeding by year 20. In the Upper Green River HU, no feeding on Dell Creek resulted in a CWD prevalence of 27 percent (SD = 6 percent) compared to 30 percent (SD = 5 percent) with continued feeding. In terms of disease-associated mortality, we predicted the closure of Forest Park and Dell Creek feedgrounds would reduce the total number of CWD mortalities by 9 percent in the Upper Green River HU and 26 percent in the Afton HU during the 20-year timespan. Our spatial analyses predicted that management alternative effects vary by HU as a function of private property and other wildlife winter ranges proximity relative to feedground location. The predicted number of elk abortions on private land, as a proxy for brucellosis risk to cattle, may increase by 8–21 percent in the absence of feeding at Dell Creek and Forest Park. Eight feedgrounds are located on Bridger-Teton National Forest, all of which have permits that have expired or will expire prior to 2028. In addition, WGFD could change their management of feedgrounds given new information; therefore, we also assessed the cumulative effects of continued feeding, phaseout, and no feeding management alternatives across five HUs south of Jackson, Wyoming (Afton HU, Fall Creek HU, Piney HU, Pinedale HU, and Upper Green River HU). These five HUs ranged from about 41 to 85 percent of the elk herd using feedgrounds, which corresponded to a CWD prevalence at year 20 of 23–34 percent if all feedgrounds in those five HUs remained open relative to 12 to 14 percent if all feedgrounds were closed. We predicted feedground closures may result in immediate reductions in population size relative to alternatives that continue feeding (for example, continued feeding and emergency feeding alternatives); however, over longer periods of time, CWD-associated mortality leads to larger population reductions. The no feeding alternative resulted in higher elk population sizes compared to the continued feeding alternative after about 10 years of implementation. Delayed action under a phaseout alternative resulted in increasing the CWD prevalence to 20 percent relative to 12 to 14 percent, on average, without feeding on HUs with a large population of fed elk such as the Upper Green River HU. Summarizing our cumulative results across all five of the analyzed HUs, we predicted continued feeding will lead to fewer elk by year 20 (mean = 8,300, standard deviation [SD] = 740) compared to no feeding at U.S. Department of Agri-culture Forest Service sites (10,700, SD = 890). The closure of all feedgrounds was projected to result in the largest elk populations at year 20 (12,500, SD = 980). No feeding at all sites also resulted in the largest cumulative harvest of 57,700 (SD = 2,600) compared to 51,100 (SD = 3,800) for continued feeding at all current feedground sites on the five HUs. Continued feeding also resulted in the lowest brucellosis costs to producers ($194,600, SD = $11,500) compared to no feeding on all feedgrounds ($243,000, SD = $13,700). Assuming moderate reductions in hunter interest because of increasing CWD prevalence in elk, we predicted that no feeding resulted in regional revenues generated by hunting activities of $190 million (SD = $10 million) compared to $173 million (SD = $10 million) for continued feeding over the 20-year timeframe. Recent CWD detections in mule deer and elk in Grand Teton National Park has elevated the importance of the cur-rent decision on whether, and how, to permit elk feeding on Dell Creek and Forest Park and the management of the other feedgrounds. Aggressive male harvest has slowed, but not stopped, the increasing prevalence of CWD in mule deer (Conner and others, 2021). It is unclear whether harvest management can be an effective tool to slow the spread of CWD in elk. There are also no effective treatments or vaccines for CWD, and it is unlikely that any will be developed that can be easily deployed in the near future. Thus, reducing artificial aggregations is one of the few management approaches suggested by the Western Association of Fish and Wildlife Agencies (Almberg and others, 2017). Future surveillance and monitoring can be designed to resolve uncertainties that can improve future decision-making. If feedgrounds close, research could quantify elk population reductions in the absence of feeding, the redistribution of fed elk to other places, or the consequences of elk movement on private property. If feedgrounds remain open, research could assess how rapidly CWD spreads in artificial aggregations of elk; however, surveillance programs would need to be designed with sufficient power to detect initial changes of CWD prevalence. Delaying action on feedground management was projected to be costly. Results of the phaseout alternative relative to the no feeding alternative suggested a 3-year delay was enough for substantial long-term changes in CWD prevalence. The long-term persistence of infectious CWD prions in the environment suggests that feedground management decisions may have long-lasting consequences. Our results indicated tradeoffs in the ability of a management agency to achieve all their objectives, and all management alternatives resulted in significant reductions in elk population size. This report contains the foundational elements for formal decision analysis methods, which can be implemented to help decision makers transparently evaluate the consequences of decision alternatives and identify the set of actions that best achieve agency and stakeholder priorities.

Wyoming