Species Status Assessment (SSA) Report

For the

Peaks of Otter Salamander (Plethodon hubrichti)

February 21, 2019

Version 1.1

Photo Credit: J.D. Wilson

U.S. Fish and Wildlife Service Northeast Region

Acknowledgements

The research for this document was prepared by Rose Agbalog (U.S. Fish and Wildlife Service (USFWS—Southwestern Virginia Field Office) with technical assistance from Krishna Gifford (USFWS—Northeast Regional Office).

We greatly appreciate the assistance of the following individuals who provided helpful information and/or review of the draft document:

John (J.D.) Kleopfer (State Herpetologist—Virginia Department of Game and Inland Fisheries) Anthony Tur (USFWS—Northeast Regional Office) Meagan Kelhart (formerly, USFWS—Headquarters) Cindy Schulz (USFWS—VAFO) Norman Reichenbach (Liberty University) David Marsh (Washington and Lee University) Jean Brennan (USFWS—Science Applications)

We also thank our peer reviewers:

Fred Huber (formerly USFS—George Washington and Jefferson National Forest) Joseph C. Mitchell (Florida Museum of Natural History—University of Florida) Thomas K. Pauley (Marshall University) Olivia LeDee (USGS—Northeast Climate Adaptation Science Center) Adrianne Brand (USGS—Patuxent Wildlife Research Center)

Suggested reference: U.S. Fish and Wildlife Service. 2019. Species status assessment for the Peaks of Otter Salamander (Plethodon hubrichti). Version 1.1. February 2019. Hadley, MA. 84 pages

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Table of Contents EXECUTIVE SUMMARY ...... 5 CHAPTER 1 INTRODUCTION...... 7 1.1 Background ...... 7 1.2 Analytical Framework ...... 7 CHAPTER 2 SPECIES INFORMATION ...... 10 2.1 Taxonomy and Genetics ...... 10 2.2 Species Description ...... 10 2.3 Range, Distribution, and Abundance ...... 11 2.3.1 Historical and Current Range ...... 11 2.3.2 Distribution and Abundance ...... 13 2.4. Life History ...... 14 2.4.1 Behavior ...... 14 2.4.2 Reproduction ...... 15 2.4.2 Feeding ...... 16 2.5 Individual Requirements (Ecological Setting and Habitat Needs) ...... 16 2.6 Population Needs ...... 18 2.7 Species Needs ...... 19 CHAPTER 3 FACTORS INFLUENCING THE SPECIES ...... 20 3.1 Mature Hardwood Forests ...... 22 3.1.1 Timber Harvest ...... 22 3.1.2 Fire ...... 26 3.1.3 Firewood Collection ...... 27 3.1.4 Defoliation and Tree Damage by Insects ...... 28 3.1.5 Tree Damage by Extreme Weather Events ...... 29 3.1.6 NPS Maintenance Work ...... 29 Summary of 3.1 Mature Hardwood Forests ...... 30 3.2 Barriers ...... 30 3.2.1 Roads ...... 30 3.2.2 Streams ...... 31

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3.2.3 Summary of Barriers ...... 32 3.3 Hybridization ...... 32 3.4 Competition ...... 32 3.4.1 Summary of Competition ...... 34 3.5 Infectious Disease ...... 34 3.6 Changing Climate Conditions ...... 35 3.6.1 Summary of Changing Climate Conditions ...... 40 3.7. Other Influences Considered ...... 40 3.7.1 Collection ...... 40 3.7.2 Predation ...... 40 3.8 Other Conservation Actions ...... 41 3.9 Summary of Factors Influencing the Species ...... 41 CHAPTER 4 – CURRENT CONDITION ...... 42 4.1 Methodology ...... 42 4.1.1 Density of Salamanders ...... 43 4.1.2 Reproductive Output ...... 44 4.1.3 Habitat Quality ...... 44 4.1.4 Allopatry/Sympatry ...... 44 4.1.5 Habitat Management ...... 44 4.1.6 Overall Condition ...... 44 4.2 Current Condition ...... 45 4.2.1 Resiliency ...... 45 4.2.2 Redundancy ...... 47 4.2.3 Representation ...... 48 CHAPTER 5 FUTURE SCENARIOS ...... 49 5.1 Methodology ...... 49 5.2 Future Scenarios ...... 49 5.2.1 Scenario 1 ...... 50 5.2.2 Scenario 2 ...... 53 5.2.3 Scenario 3 ...... 56 5.3 Summary of Species Viability ...... 59

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CHAPTER 6 KEY UNCERTAINTIES ...... 61 REFERENCES CITED ...... 62 APPENDIX A Current Condition Methodology ...... 72 A1. Survey Data ...... 72 A2. Demographic Parameters ...... 73 Number of Salamanders ...... 73 Reproductive Output ...... 74 A3. Habitat Parameters ...... 75 Habitat Quality ...... 75 Sympatry ...... 78 Habitat Management ...... 79 Overall Condition ...... 81 APPENDIX B Future Conditions ...... 82

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EXECUTIVE SUMMARY In 1957, Gordon Thurow first described the Peaks of Otter salamander. It is a slender, lungless woodland salamander that is dark brown with abundant brassy metallic spots or blotches. Adults range from a body length of 40 to 58 millimeter (mm) (1.57 to 2.28 inches (in), and likely live up to 8-10 years of age in the wild.

It is a narrow ranging endemic species that historically and currently occurs mainly in forested habitats of the mountaintops and high elevation areas between Flat Top Mountain and White Oak Ridge in Bedford, Botetourt and Rockbridge Counties, Virginia. The salamander’s range is approximately 45 square miles (mi2) (less than 120 square kilometers (km2)), and is almost entirely restricted to the Glenwood Ranger District of the George Washington and Jefferson National Forests and primarily between mile 77 and 84 of the National Park Service’s (NPS) Blue Ridge Parkway, with some limited occurrences on adjacent private lands. It is presumed extirpated from the Visitor Center AU at the southern edge of its range, however it is possible that the Peaks of Otter salamander still inhabits the area but has been undetected in recent surveys. Primary influences to the Peaks of Otter salamander’s viability currently and in the future are (1) activities that disrupt or remove the forest canopy, understory vegetation, and cover objects (e.g., primarily timber harvest); (2) competition with red-backed salamanders; and (3) changing climate patterns of increasing temperatures and changes in precipitation patterns.

In projecting the future viability of the Peaks of Otter salamander, three scenarios were considered and each scenario was projected 20 years and 80 years into the future. In Scenario 1, current influences remain constant into the future; in Scenario 2, negative influences decrease due to management efforts increasing into the future; and in Scenario 3, negative influences increase into the future and most are exacerbated by changing climactic conditions. Current and future projections for each Analysis Unit (AU) are summarized in table ES-1. This table helps to summarize the viability of the Peaks of Otter salamander currently, and into the future. Viability is supported by having multiple (redundancy), self-sustaining (resiliency) AUs distributed throughout the geographical extent of the species range and in the breadth of suitable elevations (representation).

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Table ES-1. Summary of AUs with current and future conditions under each scenario.

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CHAPTER 1 INTRODUCTION 1.1 Background This report summarizes the results of a Species Status Assessment (SSA) conducted by the U.S. Fish and Wildlife Service (USFWS or Service) for the Peaks of Otter salamander (Plethodon hubrichti). In 2012, we received a petition to list 53 species of amphibians and reptiles, including the Peaks of Otter salamander, as either endangered or threatened, and to designate critical habitat, under the Endangered Species Act of 1973, as amended (Act) (Center for Biological Diversity 2012, entire). On September 18, 2016, the Service found that the petition presented substantial scientific or commercial information indicating that the listing of the Peaks of Otter salamander may be warranted (80 FR 56423). Thus, we conducted a SSA to compile the best scientific and commercial data available regarding the species’ biology and factors that influence the species’ viability. 1.2 Analytical Framework The SSA report, the product of conducting a SSA, is intended to be a concise review of the species’ biology and factors influencing the species, an evaluation of its biological status, and an assessment of the resources and conditions needed to maintain long-term viability. The intent is for the SSA report to be easily updated as new information becomes available, and to support all functions of the Endangered Species Program. As such, the SSA report will be a living document upon which other documents, such as listing rules, recovery plans, and 5-year reviews, would be based if the species warrants listing under the Act.

This SSA report for the Peaks of Otter salamander is intended to provide the biological support for the decision on whether or not to propose to list the species as threatened or endangered and if so, whether or not to propose designating critical habitat. The process and this SSA report do not represent a decision by the Service whether or not to list a species under the Act. Instead, this SSA report provides a review of the best available information strictly related to the biological status of the Peaks of Otter salamander. The listing decision will be made by the Service after reviewing this document and all relevant laws, regulations, and policies, and a decision will be announced in the Federal Register.

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Figure 1. Species Status Assessment Framework

Using the SSA framework (figure 1), we consider what a species needs to maintain viability by characterizing the biological status of the species in terms of its Resiliency, Redundancy, and Representation (Smith et al. 2018, entire). For the purpose of this assessment, we generally define viability as the ability of the species to sustain populations in natural ecosystems within a biologically meaningful timeframe: in this case, 20 to 80 years. We chose 20 to 80 years because the available data allow us to reasonably predict the potential significant effects of stressors and conservation management within the range of the species within this timeframe. This is also consistent with the time scale for which we have previous data available on the species.

Resiliency, Redundancy, and Representation are defined as follows:

Resiliency means having sufficiently large populations for the species to withstand stochastic events (arising from random factors). We can measure resiliency based on metrics of population health, for example population size, if that information exists. Resilient populations are better able to withstand disturbances such as random fluctuations in birth rates (demographic stochasticity), variations in rainfall (environmental stochasticity), or the effects of human activities.

Redundancy means having a sufficient number of populations for the species to withstand catastrophic events (such as a rare destructive natural event or episode involving many populations). Redundancy is about spreading the risk and can be measured through the duplication and distribution of populations across the range of the species. Generally, the greater the number of populations a species has distributed over a larger landscape, the better it can withstand catastrophic events.

Representation means having the breadth of genetic makeup of the species to adapt to changing environmental conditions. Representation can be measured through the genetic diversity within and among populations and the ecological diversity (also called environmental variation or diversity) of populations across the species’ range. The more representation, or diversity, a species has, the more it is capable of adapting to changes (natural or human caused) in its environment. In the

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absence of species-specific genetic and ecological diversity information, we evaluate representation based on the extent and variability of habitat characteristics within the geographical range.

The decision whether to list a species is based not on a prediction of the most likely future for the species, but rather on an assessment of the species’ risk of extinction. Therefore, to inform this assessment of extinction risk, we describe the species’ current biological status and assess how this status may change in the future under a range of scenarios to account for the uncertainty of the species’ future. We evaluate the current biological status of the species by assessing the primary factors negatively and positively affecting the species to describe its current condition in terms of Resiliency, Redundancy, and Representation (together, the 3Rs). We then evaluate the future biological status by describing a range of plausible future scenarios representing a range of conditions for the primary factors affecting the species and forecasting the most likely future condition for each scenario in terms of the 3Rs. As a matter of practicality, the full range of potential future scenarios and the range of potential future conditions for each potential scenario are too large to individually describe and analyze. These scenarios do not include all possible futures, but rather include specific plausible scenarios that represent examples from the continuous spectrum of possible futures.

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CHAPTER 2 SPECIES INFORMATION 2.1 Taxonomy and Genetics Peaks of Otter salamander (Plethodon hubrichti) is classified in the order Caudata and the family Plethodontidae. Thurow described the species in 1957 (Thurow 1957, entire). The species was defined morphologically by Highton in 1962 and biochemically by Highton and Larson in 1979 (entire). Highton’s taxonomic description has been widely accepted (Wiens et al. 2016, p. 2586). Using electrophoresis, Highton (1999, p. 43) analyzed geographic protein variation in 24 genetic loci of 5 previously unstudied species of the Plethodon cinereus group (P. hoffmani, Peaks of Otter salamander, P. nettingi, P. richmondi, and P. shenandoah). He found that nine of the salamanders within this species group appeared to have been derived from a common ancestor at about the same time in the Pliocene Epoch, but that all of the populations within each species cluster as a monophyletic group (i.e., all share a common ancestor) (Highton 1999, p. 43). A later phylogenetic study of the Plethodon cinereus group indicated that Peaks of Otter salamander has low haplotype diversity and is a well-defined species as compared to other similar groups of plethodontids (Sites et al. 2004, pp. 103–104). Sites et al.’s (2004, entire) study was based on individuals collected from throughout the species’ range (n=17) and the results did not detect genetic partitioning throughout the population. Because the taxonomy is widely accepted by species experts and researchers, and genetic studies have shown the Peaks of Otter salamander to be a distinct species within the Plethodon cinereus group, we accept the existing taxonomy. 2.2 Species Description The Peaks of Otter salamander is a slender, lungless woodland salamander and, like other Plethodontids, has no aquatic life stages (Thurow 1957, p. 59; Spotila 1972, p. 96). All respiration occurs through their skin and mucous membranes in their mouth and throat. Adults snout-vent length (SVL) ranges from 40 to 58 millimeters (mm) (1.57 to 2.28 inches (in)) with a total body length (includes tail) of 85 to 131 mm (3.35 to 5.16 in) (Highton 1986, p. 393.1; Dodd 1979, p. 384). This species has 20 trunk vertebrae and, usually, 19 costal grooves (Highton 1986, p. 393.1; Thurow 1957, p. 59). The body is dark brown with abundant brassy metallic spots or blotches that occasionally form an irregular dorsal (back) stripe, or, less commonly, larger, white spotting (figure 2a). In adults, the venter (underside) is plain dark gray to black (Bury et al. 1980, p. 30) and there are white spots on the sides. There is no sexual dimorphism in the species except during the breeding season when males have a large mental gland (see section 2.4.2 Reproduction) (Pague and Mitchell 1991, p. 436 In Terwilliger 1991). Kniowski and Reichenbach (2006, p. 332) observed that neonates are dark brownish-gray in coloring with no distinct marks or spots, which is in contrast to Petranka’s (1998, p. 335) observations that hatchlings have a distinct dorsal stripe consisting on reddish spots. Other reports note that juveniles have small red spots on the dorsum (Highton 1962 In Highton 1986, p. 393.1; Dodd 1979, p. 384). The red-backed salamander has similar life history characteristics and overlaps with the Peaks of Otter salamander in some areas of their ranges. Red-backed salamanders can be distinguished from the Peaks of Otter salamander morphologically. There are typically two color morphs of red- backed salamanders. The striped or red-backed morph has a broad, straight edged, orangish red or

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red dorsal stripe that extends from the head to the tail, the sides are dark and the venter is mottled with black and white. The unstriped morph is similar but lacks the red dorsal stripe, and the entire dorsum is dark and the animals are unicolored (Petranka 1998, p. 335).

Figures 2a, 2b, and 2c. Adult, subadult, and hatchling Peaks of Otter salamanders. Figure 2a (top left): adult, photo credit: John White. Figure 2b (top right): subadult, photo credit: A.B. Kniowski. Figure 2c (lower left): hatchlings, photo credit: A.B. Kniowski. 2.3 Range, Distribution, and Abundance 2.3.1 Historical and Current Range The Peaks of Otter salamander is a high elevation, range-restricted endemic that historically and currently occurs primarily in forested habitats between Flat Top Mountain and White Oak Ridge in Bedford and Botetourt counties of Virginia (Highton 1986, p. 391.1; Dodd 1979, p. 384; Pague and Mitchell 1991, p. 436 In Terwilliger 1991), and Thunder Ridge in Rockbridge County, Virginia (Thurow 1957, p. 60; Pague et al. 1992, p. 9). The salamander’s range is approximately 45 square miles (mi2) (less than 120 square kilometers (km2)), and is almost entirely restricted to the Glenwood Ranger District of the George Washington and Jefferson National Forests and primarily between mile 77 and 84 of the National Park Service’s (NPS) Blue Ridge Parkway, with some limited occurrences on adjacent private lands (Dodd 1979, p. 384; Pague and Mitchell 1990, p. 5; Mitchell et al. 1996, p. 15; USFWS 1997, p.7; Marsh 2018b) (table 1).

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Figure 3. Historical and current range of the Peaks of Otter salamander. The gray outline represents the known outer boundaries of the species’ range, based on the best available survey data. The Peaks of Otter salamander’s range is a combination of Federal and private land. Approximately 92 percent of the species’ range is on federally owned land, the majority of which is owned and managed by the U.S. Forest Service (USFS) (table 1).

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Table 1. Estimated land ownership in known occupied Peaks of Otter salamander habitat. Range estimate is closely tied to survey data and does not account for topography. Total acreage calculated is less than what is defined in the literature.

Land Owner Approximate Approximate Percent of Number of Number of Landownership Acres Hectares U.S. Forest Service 13,134 5,315 76%

Blue Ridge Parkway (NPS) 2,676 1,083 16%

Private Land (entire parcel) 1,456 589 8%

Total 17,248 6,980 100%

2.3.2 Distribution and Abundance Within the species’ range, the Peaks of Otter salamander occurs at mid to higher elevations, with varying densities. The best available information suggests that the species is restricted to elevations above 443 meters (m) (1,453.41 feet (ft)) (Mitchell et al. 1996, p. 15). However, the majority of individuals occur above 760 m (2,493 ft) (Bury et al. 1980, p. 30; Petranka 1998, p. 364), with elevations greater than 650 m (2,133 ft) considered to be optimal (Reichenbach and Brophy 2017, p. 14). The best available information indicates that surface-active (SA) (see section 2.4 Life History) salamander densities peaked between elevations ranging from 900 to 1100 m and then decreased rapidly with a decrease in elevation. Survival rates were also found to decrease with elevation, and the decreases were correlated with increases in temperature and a decrease in relative humidity associated with lower elevations. For elevations above the optimum, a shorter active season may have contributed to decreases in SA salamander density and relative egg output (Reichenbach and Brophy 2017, p. 1). The Peaks of Otter salamander has been documented to be abundant within its known range, leading some researchers to conclude that the species is “healthy” and stable. Pague and Mitchel (1987, p. 310) noted that it is abundant in many localities within its restricted range, while Pague and Mitchell (1990, p. 8) agreed with reports from Bury et al. (1980, p. 30) that the species is seasonally abundant where found. Reichenbach and Sattler (2007, p. 626) found that within its range, the Peaks of Otter salamander dominated the terrestrial salamander community, making up 94.8 percent of the species observed. A variety of occurrence data were used for analysis in this SSA. Dates of survey data ranged from 1962 to 2017, and came from a variety of sources including the USFS, Virginia Department of Game and Inland Fisheries (DGIF), various researchers and educational institutions. The methodology for data collection and documentation varied among data sources and are generally not consistent with one another. See Appendix A for a more detailed explanation of survey data.

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2.4. Life History The Peaks of Otter salamander has four discrete stages of its life cycle: eggs, hatchling, juveniles, and adults (figure 4). There is no estimate for how long the species may live in the wild, but based on similar salamander species like red-backed salamanders (LeClair et al. 2006, p. 266), 8 to 10 years may be a reasonable assumption. Mean generation time for Plethodon jordani, was calculated to be 9.8 years (Hairston 1983, p. 1030). Snider and Bowler (1992, p. 7) found that Peaks of Otter salamanders could live over 19 years in captivity where food was consistently available and habitat resources were ideal. This is likely much longer than animals would naturally live in the wild.

Figure 4. Life stages of the Peaks of Otter salamander.

2.4.1 Behavior As previously discussed, the Peaks of Otter salamander does not have an aquatic life stage, and its entire life cycle is reliant upon environmental moisture because it requires moist skin for respiration (Terwilliger 1991, p. 420). When environmental conditions are favorable, this species will spend time on the surface to forage and search for mates. However, it also spends a considerable amount of time in the subsurface environment to overwinter, hide from predators, lay eggs, and escape unfavorable temperature, precipitation, and relative humidity conditions (Spotila 1972, pp. 121– 122; Keen 1979, pp. 686–687). Peaks of Otter salamanders can often be found under wet leaf-litter, rocks, logs and other cover objects (Bury et al. 1980, p. 30; Martof et al. 1980, p. 88; Pague and Mitchell 1991, p. 437 In Terwilliger 1991). Surface and subsurface behavior patterns are seasonal as well as, in the active season, contingent upon adequate environmental conditions. See section 2.5 Individual Requirements (Ecological Setting and Habitat Needs) below for more details.

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This species is primarily nocturnal, although individuals can be found during the day in deeply shaded habitat or under rocks, logs, branches, and leaf litter (Dodd and Linzey 1979, p. 384; Pague 1989, p. 11; Reichenbach and Brophy 2017, p. 8). Surface activity peaks between 2100 and 2300 hours in the spring and from 2000 to 2300 hours in the fall, and is generally associated with leaf litter moisture due to recent rainfall (Kramer et al. 1993, p. 433). The majority of surface-active salamanders collected from June through September were observed on vegetation, primarily ferns (Kramer et al. 1993, p. 433). Abiotic factors including moon phase and precipitation are significant predictors of movement (Goff 2015, pp. 23–27). In addition to moving underground to escape predators, the Peaks of Otter salamander has other defense mechanisms. Individuals may defend themselves against predators by becoming immobile, dropping their tail (i.e., tail autonomy), coiling and releasing noxious secretions (Dodd et al. 1974, pp. 90–91). Home range size of the Peaks of Otter salamander is typically less than 1 square meter (m2)(10.8 square feet (ft2)), while the median area is 0.6 m2 (6.5 ft2)(Kramer et al. 1993 p. 433). According to Goff (2015, p. 11) Peaks of Otter salamanders move an average total distance of 1.41 m (4.6 ft), while being tracked over a two month period. 2.4.2 Reproduction In both males and females, secondary sexual characteristics become more pronounced with age, even after sexual maturity, which occurs near a length of 38 mm from snout to posterior margin of the hind limbs (Thurow 1957, p. 62). Males develop a large mental gland (Martof et al. 1980, p. 88) under the chin during the breeding season that produces pheromones to increase female receptivity. Mating occurs in fall and spring, but eggs are not laid until late May or June (Pague 1989, p. 11; Kniowski and Reichenbach 2006, p. 322). Eggs are fertilized internally by means of a spermatophore (Dodd 1979 p 384; Pague 1989, p. 11) and laid together in nests under logs or rocks (Petranka 1998, p. 339; Kniowski and Reichenbach 2006, p. 332). Females produce 1 to 15 eggs (mean of 8.5 (95 percent confidence interval (CI) 8.2 to 8.9)) (Reichenbach and Brophy 2017, p. 11) and have an average clutch size of 10 (Martof et al. 1980, p. 88). Peaks of Otter salamander females guard their nests by coiling around the nest, which helps protect the eggs from predators and may minimize egg dehydration (Petranka 1998, p. 339; Kniowski and Reichenbach 2006, p. 332). Reproductive output is affected by surface-active salamander density, percent gravid females, and the number of eggs per female, which is directly correlated with the salamander’s body mass and elevation (Reichenbach and Brophy 2017, p. 11). The number of eggs per female appears to increase with elevation, and then levels off. Reichenbach and Brophy (2017, p. 11) found that a maximum of 12 eggs per female were produced at 1000 m (3,280.84 ft) elevation, but that value then decreased slightly above 1000 m (3,280.84 ft) elevation. The authors hypothesized that reproduction at higher elevation sites is primarily biennial, because only approximately half (40 to 60 percent) of the observed sexually mature females at higher elevations were gravid. These observations were similar to trends noted in red-backed salamanders (Takahashi and Pauley 2010, p. 90). At the lower elevation sites, in contrast, Reichenbach and Brophy (2017, p. 11) hypothesized that some of the Peaks of Otter salamanders produced eggs annually because the

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majority (67 to 80 percent) of observed sexually mature females were gravid. The overall effect with decline in elevation, even with more frequent reproduction, was a decrease in reproductive output (Reichenbach and Brophy 2017, p. 13). The best available information indicates that growth rates, eggs/female, and reproductive output decreased with elevation, and the decreases were correlated with increases in temperature and a decrease in relative humidity associated with lower elevations (Reichenbach and Brophy 2017, p. 1). The best available information indicates the Peaks of Otter salamander exhibits a 1:1 sex ratio (Reichenbach and Brophy 2017, p. 11).

2.4.2 Feeding Peaks of Otter salamanders are insectivores that feed on a variety of invertebrate prey items including ants (Hymenoptra), ticks and mites (Acarina), springtails (Collembolan) and earthworms (Haplotaxida) (Mitchell et al. 1996, p. 17; Arif et al. 2007, p. 848). Foraging for food occurs primarily on the ground surface and vegetation on foggy, humid or rainy nights (Jaeger 1978, p. 686; Pague and Mitchell 1991, p. 436 In Terwilliger 1991; Kramer et al. 1993, p. 431). The timing and extent of feeding relies heavily on the environment. Plethodontids, like the Peaks of Otter salamander, are increasingly limited in their foraging as environmental moisture decreases (Spotila 1972, pp. 121-122; Keen 1979, pp. 686–687). When the soil surface, leaf litter and vegetation are dry, plethodontids retreat to underground refugia where little feeding occurs (Feder 1983, p. 298). Plethodontid salamanders have adapted different mechanisms for surviving long periods without eating. The low metabolism of plethodontids allows their caloric reserves to sustain them for long periods, and they may also divert energy stores from reproduction to metabolism maintenance when necessary (Feder 1983, p. 304). The lives of plethodontids may consist of long periods of inactivity interspersed with brief periods of activity and energy processing when thermal and hydric conditions permit. Key specializations like low metabolic rate, relatively large energy stores and resistance to starvation may enable plethodontids to survive indefinite periods between unpredictable periods of feeding. The absence of energetically costly “adaptations” like ventilation, thermoregulation, maintenance of a high performance cardiovascular system, and sustained high levels of physical activity, may be a partial explanation of the extraordinarily low energy requirements of plethodontids (Feder 1983, p. 291). 2.5 Individual Requirements (Ecological Setting and Habitat Needs) We evaluate the individual needs of the Peaks of Otter salamander in terms of the resource needs that are necessary to complete each stage of the life cycle, including eggs, hatchlings, juveniles, and adults. All life stages of the Peaks of Otter salamander are dependent upon the maintenance of suitably moist, thermally appropriate microhabitats.

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Table 2. Resource needs by Peaks of Otter salamander life stage.

Resources Eggs Hatchlings Juveniles Adults Moderate temperatures (soil and air) S* S S S Moist environment/relative S S S B*, S humidity/precipitation Moist, soft soil/humus, leaves F*, S F, S F, S Available prey F F F Underground refugia S S S S Logs, rocks, subsurface areas, leaves S F, S F, S B, F, S Optimal elevation B * Resource Functions: B = breeding; F = feeding; S = sheltering

Peaks of Otter salamander habitat is characterized by mature Appalachian hardwood forests, which consist of late successional deciduous oak (Quercus) and maple (Acer) woodlands and rhododendron (Rhododendron) thickets (Thurow 1957, p. 63; Bury et al. 1980, p. 30; Pague and Mitchell 1991, p. 436 In Terwilliger 1991). The salamander may be present in virtually all age classes of forest provided that sufficient moisture is present, but presence and abundance is most often associated with more mature stands of hardwood forests (Pague and Mitchell 1990, p. 7). The Peaks of Otter salamander may be absent from drier south and west facing slopes within its range at lower elevations (Pague and Mitchell 1990, p 5–8). The Peaks of Otter salamander is not associated with forests where pine is dominant, but has been documented at sites where hemlock was codominant (Pague and Mitchell 1990, p. 5). Plethodontid salamanders have low energy requirements and are susceptible to desiccation. Since lungless salamanders use cutaneous respiration (i.e., breathe through their skin), they depend on cool, moist microhabitats for survival. Suitably moist and thermally equable microhabitats are almost always available deep in burrows, rotted logs and rock crevices, while the exposed microhabitats that are suitable for foraging and courtship (e.g., leaf litter, soil surface, rock faces and the surface of vegetation) are likely to be ephemeral in both space and time (Feder 1983, p. 305). In wet weather, most exposed microhabitats will be available if the air is still; however, wind, low humidity, extreme temperatures, or drought will eventually limit their access to these microhabitats. When the habitat becomes hot and/or dry they retreat to underground refugia where soil moisture and temperature are more suitable and stable. During spring and fall, when evaporation rates are lowest and soils are cool and moist, salamanders are most active and found in their highest numbers (Pague 1989, p. 11). Surface activity is generally associated with leaf litter moisture due to recent rainfall. Kramer et al. (1993, p. 433) found that the total percentage of surface-active Peaks of Otter salamanders on a given site visit was consistently between 3 and 5 percent of the total population of surface-active salamanders. In late August and September, that percentage increased to approximately 10 percent of the total population of surface-active salamanders. Taub (1961, p. 697) noted that captures of red-backed salamanders were more frequent in the spring and fall, and that Plethodon are distributed vertically

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within the soil throughout the year, with distribution mostly affected by water table level and temperature.

2.6 Population Needs We evaluate the population needs of the Peaks of Otter salamander in terms of what is required for self-sustaining populations. The measure of resiliency is based on a population’s ability to withstand or recover from environmental or demographic stochastic events, such as changes in air and soil temperature or moisture, or the amount of above or below ground refugia. We evaluate resiliency in terms of resources and/or the circumstances that are necessary to maintain population abundance, distribution, population growth rates, and reproduction (see section 4.2). These may vary if populations inhabit different ecological settings. Resiliency is positively related to population size and growth rate and may be influenced by connectivity among populations/subpopulations. Generally speaking, populations need abundant individuals within habitat patches of adequate area and quality to maintain survival and reproduction in spite of disturbance.

There is little information available regarding the demographic or genetic processes that define the spatial structure of Peaks of Otter salamander populations. It is possible that due to the species’ narrow range, the availability and quality of habitat within its range, the absence of barriers to dispersal, and the species’ distribution within the available habitat (Kramer et al. 1993, p. 433), the Peaks of Otter salamander most likely functions as one population with some inherent metapopulation structure. Because we lack specific information about biologically meaningful populations or subpopulations, for purposes of analyzing the status of the salamander in this SSA report, we defined the species’ “population” as analysis units (AUs). If the Peaks of Otter salamander consists of a single population, then the AUs could be structurally equivalent to subpopulations. We delineated the AUs by clustering the best available survey information in combination with topography and landscape characteristics (see section 4.1).

The following conditions are needed to support self-sustaining AUs: Habitat Quality Like other terrestrial salamanders, the population size of the Peaks of Otter salamander is likely dependent on the habitat availability and density of suitable habitat. Population may vary depending upon the quality of habitat factors including, soil depth, soil temperature, soil moisture, aspect, slope angle, underground shelter availability, nest site availability, surface cover sites for territories, prey quality and abundance, predator abundance, and presence of competitors (Buhlmann et al. 1988; Dodd 1981; Wicknick 1995 In Mitchell et al. 1996, p. 18). Other important factors include connectivity, cover objects, soil moisture, humidity, ground temperature and elevation. Salamander Density and Abundance Due to inconsistent survey data (section 2.3.2 Distribution and Abundance) and the species’ cryptic nature, there is no overall population size estimate available for the Peaks of Otter salamander, however density estimates for the Peaks of Otter salamander were reported to be approximately 4.5

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salamanders/m2 (Kramer et al. 1993, p. 432) in optimal habitat conditions. These estimates are within the range of estimates for the closely related red-backed salamanders, which include 0.05 salamanders/m2 (Test and Bingham, 1948), 0.9/m2 (Heatwole 1962), 2.1/m2 (Jaeger 1979), 2.2/m2 (Jaeger 1980), and 0.21/m2 (Burton and Likens 1975) (In Kramer et al. 1993, p. 433). Several studies have provided density estimates based on the number of salamanders observed on the surface. Reichenbach and Sattler (2007, p. 626–628) compared surface-active salamanders in treated areas (i.e., timber harvest) and reference sites and found similar densities. Densities averaged 0.21/m2 while in reference site plots (no timbering activities) the mean density of surface- active salamanders was 0.28 m2. They suggest that surface counts explained the variability found in population size estimates and therefore was a good predictor of population size, and also that mean surface counts can be used to estimate population size under both closed and open canopy forests (p. 627). While we are unaware of the precise optimum level of abundance or densities needed, we do assume, based on general principles of conservation biology, that AUs containing larger numbers and densities of Peaks of Otter salamanders are likely to be more resilient than AUs containing fewer numbers and densities of salamanders. 2.7 Species Needs We evaluate the species’ needs in terms of the resources and/or the circumstances that support the redundancy and representation of the species. Specific to the Peaks of Otter salamander, redundancy is evaluated by the presence of multiple, self-sustaining AUs distributed throughout its range, and representation is evaluated based on both the presence self-sustaining of AUs distributed across the breadth of the species’ range and in AUs that span a diversity of elevational gradients (see sections 2.4.2 Reproduction and 4.3.3 Representation) In summary, the viability of the Peaks of Otter salamander is supported by having multiple (Redundancy) self-sustaining (Resiliency) AUs distributed throughout the geographical extent of its range along varying suitable elevations (Representation).

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CHAPTER 3 FACTORS INFLUENCING THE SPECIES In this chapter, we evaluate the past, current, and future influences that are affecting or could be affecting the current and future condition of the Peaks of Otter salamander throughout all or some of its range. These influences are summarized in a conceptual model (figure 5) and discussed in more detail in the section below.

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Figure 5. Summary of factors influencing the Peaks of Otter Salamander’s viability. Red arrows and a minus sign (-) indicate a negative effect, blue arrows and a plus sign (+) indicate a positive effect, and grey lines and a question mark (?) indicate an uncertain effect.

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3.1 Mature Hardwood Forests Activities or ecological processes that disrupt or remove the forest canopy, understory vegetation, and cover objects can have significant effects to the Peaks of Otter salamander because they reduce or eliminate moist areas (e.g., leaf litter, humus layers, rotting logs) necessary for the salamander’s breeding and sheltering, and limits foraging habitat and connectivity between the salamander’s subpopulations (Pough et al. 1987 p. 2; Terwilliger 1991, p. 420; Mitchell et al. 1996 p. 15). These activities or processes include timber harvest and firewood collection, and defoliation due to insects, fire and extreme weather events.

3.1.1 Timber Harvest Generally, timber harvests can cause drier soil, loss and drying of leaf litter, and loss of fine woody debris- changes that decrease forests’ ability to sustain salamander populations (Welsh and Droege 2001, pp. 560–562) because removing the forest canopy exposes the ground surface to more sunlight and wind, two ecological factors that heat and dry microhabitat features (Terwilliger 1991, p. 420). According to O’Donnell et al. (2015, p. 767), indirect effects of timber harvest can: “reduce survival of salamanders (Petranka et al.1993, 1994; Homyack and Haas 2009), limit surface activity (Johnston and Frid 2002, Homyack et al. 2011, Hocking et al.2013), and induce emigration from the harvested area (Ash and Bruce 1994, Semlitsch et al.2008, Peterman et al.2011), which have been expressed as the mortality, retreat, and evacuation hypotheses (Semlitsch et al.2009). Many studies that have reported terrestrial salamander declines after timber harvest implicate higher ground temperatures and decreased soil moisture due to canopy cover removal (e.g., Petranka et al.1993, 1994; deMaynadier and Hunter 1995; Semlitsch et al.2009; Tilghman et al. 2012; Homyack and Haas 2013).” Timber harvest is permitted within specific areas of the Peaks of Otter salamander’s range. However, where, how often, and by what method the timber harvest occurs is managed on Federal land by the USFS (2004, Chapter 3 p. 129) and in accordance with the USFS’, NPS’, and USFWS’ Habitat Conservation Agreement (HCA) for the Peaks of Otter salamander (USFWS 1997, entire). There is no restriction on timber harvest or other activities within the 8 percent (1,456 ac (589 ha)) of private land within the species’ range. The HCA (USFWS 1997, entire) outlines two area types: the ‘primary conservation area’, where commercial timber harvest is not allowed, and the ‘secondary conservation area’, where commercial timber harvest is allowed. The primary conservation area represents areas that are primarily unsuitable for timber harvest and consists of approximately 39 percent of the Peaks of Otter salamander’s range (USFWS 1997, p. 11). Activities such as trail construction and dispersed camping may be permitted in the primary conservation area, provided they do not negatively affect the salamander. Activities within the secondary conservation areas must be conducted in a manner that minimizes effects to the species (USFWS 1997, p. 11). The USFS has designated the Peaks of Otter salamander Habitat Conservation Areas which includes approximately 7,700 ac (3,116 ha) within the Glenwood Ranger District (USFS 2004, chapter 3 p. 129). Within the HCA, the USFS has designated additional zones (primary and secondary) related to how the Ranger District manages the habitat. These zones can be overlain by the HCA’s primary and secondary conservation areas and activities within the zones are implemented using different management prescriptions aimed at limiting impacts to the salamander

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and its habitat. These prescriptions allow timber harvest, limit timber harvest, or do not allow timber harvest within the prescription area. When prescriptions overlap, it is the most conservative prescription that takes precedence. For example, when two areas overlap in a secondary conservation zone, a “no timber harvest” prescription will take precedence (Kirk 2018b).

Figure 6. Primary and secondary conservation areas and USFS management units within the Peaks of Otter salamander’s range. Within the primary habitat areas, totaling 2,400 ac (971 ha), no timber harvest is permitted. The landscape consists of closed forest canopy with cool, moist habitats that include ground cover like rocks, down and decaying logs and leaf litter. Soils are deep and uncompacted, there is ample organic matter and small invertebrates are plentiful as food for the salamander. The terrain is steep and rocky and generally unsuitable for timber production (USFS 2004, chapter 3 pp. 129, 131). Within the secondary habitat areas, totaling 5,300 ac (2,145 ha), shelterwood cuts and thinning are permitted, but clear cutting is prohibited (USFS 2004, chapter 3 pp. 129, 131). However, a portion of the USFS management prescription that overlaps with the secondary conservation zone is categorized as a 12.A Remote Backcountry Recreation (~830 ac (336 ha)) area where timber harvests are prohibited (USFS, 2004, Chapter 3 p. 188). As a result, within the secondary habitat area, only shelterwood cuts and thinning are allowed in the 4,470 ac (1,809 ha) area. In the

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secondary conservation areas, the landscape consists of mid to late-successional forest communities with newly regenerated and younger aged forest stands dispersed throughout the area. Cool, moist habitats that include ground cover like rocks, down and decaying logs and leaf litter are maintained and restored. Soils are deep, uncompacted and contain high densities of organic matter allowing salamanders the opportunity to burrow. Small invertebrates are plentiful as food for the salamander (USFS 2004, Chapter 3 p. 132).

Table 3. Area where timber harvest is permitted.

Habitat Designation Number of Acres Where Number of Hectares Where Timber Harvest is Timber Harvest is Permitted Permitted Primary Conservation Area 0 0 Secondary Conservation 4,470 1,809 Area Private 1,456 589

In the secondary habitat areas, spaces without vegetation like roads, trails and utility rights-of-way are minimized, although trail and road reconstruction and new parking facilities are permitted (USFS 2004, Chapter 3 p. 133). Management within the secondary habitat area may include prescribed fire and reduction of fuel hazard (p. 135). There are many standards in place under the Timber Management section of the Jefferson National Forest Management Plan. These standards allow shelterwood cuts and thinning to take place but work to reduce impacts to the salamander’s habitat. Some of these standards include: • limiting the area to be harvested to 100 ac (40.47 ha) per year; • retaining large woody debris on the ground, leaving a minimum of 50 square feet (ft2) (4.64 square meter (m2))of basal area per acre; • limiting second entry to at least 15 years from first harvest; • excluding harvest areas immediately adjacent to areas known to have low Peaks of Otter salamander densities; • suspending timber harvest during periods with the greatest amount of Peaks of Otter salamander surface activity; and • monitoring to determine if the objectives of maintaining Peaks of Otter salamander populations are being met (USFS 2004, Chapter 3 p. 136).

The Forest Management plan states that studies will continue to gather information on how long it takes for Peaks of Otter salamander populations to return to densities equivalent to those in adjacent mature stands, which will help to modify management practices, if necessary (USFS 2004, Chapter 3 p. 137). Monitoring was done to determine effects of different types of timber harvest on the Peaks of Otter salamander population in the Parkers Gap area. Baseline data were gathered in 1993 and subsequent monitoring activities occurred in 1994, 1995, 1997, 1999, 2001 and 2005 (Reichenbach and Sattler, 2005 p. 2; Reichenbach and Sattler, 2007 p. 623). Timber harvest activities have not occurred in any other areas designated as Peaks of Otter salamander conservation areas.

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The effects of clearcutting and shelterwood cuts on the Peaks of Otter salamander have been examined in a number of studies (Sattler 1998, entire; Sattler and Reichenbach 1998, entire; Reichenbach and Sattler 2005, entire; USFS 2005, p. 78–81; Reichenbach and Sattler 2007, entire; Marsh 2014, entire). Because clearcuts are not allowed in either the primary or secondary habitat conservation areas, which cover the federally owned areas (92 percent) of the salamander’s range, only shelterwood cuts will be evaluated in this SSA. It is possible that timber harvest, including clearcutting methods, may be used on the 8 percent of the species’ range on privately owned lands, but we are uncertain if and to what extent this has occurred in the past or will occur in the future. In the past 21 years (1997 to 2018) of implementing the HCA, approximately 156 ac (63.13 ha) of the Peaks of Otter salamander habitat have been harvested. Harvests have been limited to the Parkers Gap area. In that time, several studies documented the effects of timber harvest on the species. For example, Reichenbach and Sattler (2005, entire; 2007, entire) evaluated pre-timbering salamander abundance, then sampled periodically for 12 years following timber harvest. The pattern of change over time for the number of Peaks of Otter salamander at reference sites was not significantly different from shelterwood cut sites (Reichenbach and Sattler 2007, p. 625). In qualitative assessments, canopy and ground cover for sites receiving a shelterwood cut treatment were similar to reference sites, with canopy closure averaging 84.4 percent and dead leaf cover on the forest both averaging 69.8 percent after 5 years of treatment (p. 627). Marsh (2014, pp. 2–3) found that number of Peaks of Otter salamanders decline following shelterwood harvest, with most of the declines farther out into the harvested zone. These patterns are coincident with changes in soil temperature, though the relationship between temperature and salamander abundance may not be causal. These results differed from what Sattler and Reichenbach (1998) found which could be due to site-specific differences or differences in the size of harvest units, which were larger in the Marsh study. Marsh stated that although the study covered only 5 years post-harvest, they expect that salamander numbers will increase again as forest begins to regrow (Marsh 2014, p. 3). Tilghman et al. (2012, p. 1) performed a meta-analysis of the effects of canopy removal on terrestrial salamanders and found that salamander numbers almost always declined following timber removal, but salamanders were never extirpated and populations typically increased as forests regenerated. Mitchell et al. (1996, p. 18) documented that the number of Peaks of Otter salamanders in shelterwood sites were on average 10 to 66 percent lower than in adjacent mature sites, and noted that different levels of cuts had dramatically different effects on the salamander because of the interactions between the amount of basal area left after a shelterwood cut and the quality of habitat before and after the timber operations. Homyack and Hass (2013, p. 362) found that silvicultural regimes that employ multiple entries within a rotation, typical of shelterwood cuts, have the potential to negatively affect terrestrial salamanders in southern Appalachian forests. They suggest that shelterwood cuts that repeatedly reduce salamander populations will require a longer period for population recovery, or may permanently suppress populations. The effects of multiple stand entries within a rotation on soil compaction and soil erosion could also have long-term negative effects on salamanders.

Studies have also analyzed prey in the stomach contents of Peaks of Otter salamander and found that there was no statistical difference in the amount of hard-bodied prey (e.g., beetles and other insects) among different timber treatments, but they did find a statistical difference in soft-bodied collembolans (e.g., springtails) (Mitchel et al. 1996, p. 17). Multiple statistical tests showed that

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salamanders from mature stands ate more collembolans than salamanders from 12 to 18 year old clearcuts and the 2 to 4 year old shelterwood cuts, suggesting that prey resources differ over different treatment areas and that higher quality prey may be more readily available in mature sites as compared to sites with clearcuts or shelterwood cuts. Different timber management practices may cause small-scale differences in growth, diet, reproduction, and population recruitment. 3.1.1.1 Summary of Timber Harvest Broad scale timber harvest can affect Peaks of Otter salamander density due to affects from canopy removal resulting in the heating and drying of the ground surface. Clearcutting is not permitted on federally owned land, which is 92 percent of the species range, of which 37 percent (5,905 ac) of the species’ range may be subject to rotating shelterwood harvest practices and the remaining 63 percent of the species’ range is protected from all forms of timber harvest. Habitat management within the federally owned area of the species’ range is governed by the 1997 HCA (USFWS 1997, entire) and USFS’ Forest Management Plan (2004, entire). We have no information to suggest that these management plans will not continue to be implemented in the future, however because current management allows for some timber harvest, we will evaluate timber harvest in our future condition analysis. 3.1.2 Fire Prescribed fire is used as a habitat management tool within the Peaks of Otter salamander’s range (USFS 2004, Chapter 3 p. 131). Implementation of prescribed fire and wildfire suppression activities is guided by specific management prescriptions that are intended to minimize effects to the Peaks of Otter salamander. These guidelines (USFS 2004 p. 137 Chapter 3) include:

• 8E2-029 Ensure firefighter and public safety as the first priority. Secondly, protect property and natural and cultural resources based on the relative values to be protected. Wildland fire response is suppression with initial attack to minimize acreage burned. Suppression strategies will strive to minimize soil disturbance, as well as canopy and cover loss. • 8E2-030 Minimize the use of soil-disturbing mechanized equipment when suppression can be achieved with other methods. Avoid moist habitats during line construction when fire conditions allow. • 8E2-031 Rehabilitate all firelines as quickly as possible through reseeding and dragging cover objects into the line. • 8E2-032 Prescribed fires are permitted predominantly on drier sites supporting rare plants or unique natural communities. Prescription for fire will ensure low mortality of canopy vegetation and low risk of escape. Stand replacing prescribed fires are not allowed. • 8E2-033 Monitor effects of prescribed fire on Peaks of Otter salamander populations following prescribed fires. • 8E2-034 When their use cannot be avoided, locate disked/bladed/plowed firelines outside of moist habitats.

The Peaks of Otter salamander, like other salamander species, may be directly or indirectly affected by a fire, depending on the timing, extent, and severity of the event. We are unaware of any data that have documented the effects of fire on the Peaks of Otter salamander. In a literature review for another salamander species, one researcher concluded:

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“indirect fire effects are thought to have more influence on terrestrial salamanders because they do not have the capability to emigrate as quickly or as far as many other terrestrial vertebrates (Kleeberger and Werner 1982, Ousterhout and Liebgold 2010). Salamanders may effectively become trapped in a fire-disturbed landscape, which could involve reduced prey availability, fewer cover objects, and decreased soil moisture (Russell et al.1999, Pilliod et al.2003). Prescribed fires generally decrease leaf litter and duff (i.e., decomposed organic material) depths, can combust or desiccate downed wood that salamanders use as refugia, and may lead to higher temperatures at ground level (Harmon and Franklin 1986, Bury et al.2002, Pilliod et al.2003, Cummer and Painter 2007, Matthews et al. 2010). Salamanders may respond by spending more time underground, which could reduce foraging and breeding opportunities and lead to decreased survival” (In O’Donnell et al. 2015, p. 767).

While the Peaks of Otter salamander could be affected by the use of prescribed fire, wildfire or wildfire suppression activities, we are uncertain whether this has happened in the past. There have been two fires documented within the salamanders’ range, one in 1999 and one in 2000. Both of these fires were less than 2 acres each (Wright 2019). The possibility exists for wildfire events to increase in the future as temperature and precipitation patterns shift (see 3.6 Changing Climate Conditions). Because the species’ range is relatively small, we acknowledge that if a severe, wide spread wildfire event happens, it could have catastrophic consequences for the species. Because of the uncertainty about the extent of occurrence and effects of fire on the Peaks of Otter salamander, we are not evaluating it further in our resiliency and viability analysis, but recognize that the risk of such an event occurring remains even though it is not explicitly incorporated into our future scenarios (see Chapter 5). 3.1.3 Firewood Collection Because salamanders rely on surface objects for shelter, foraging and nest sites, the collection of downed wood from the forest floor for use as firewood can remove important habitat features and lead to drying of the soil by increasing exposure to sunlight and air. Firewood collection is addressed in the Jefferson National Forest Management Plan (USFS 2004, Chapter 3 p. 137), which states that personal use of cutting dead and down firewood is permitted only within 100 ft (30.48 m) of roads. Although firewood collection could remove important cover objects for the salamander or result in death due to trampling, restricting firewood collection to road corridors concentrates and limits impacts to the population.

Using this 100 ft (30.48 m) buffer, we estimate that firewood collection is permitted to occur on approximately 1,072 total ac (433.82 ha) within Peaks of Otter salamander’s range on USFS land. In addition, firewood collection for campfires is also permitted in the immediate vicinity of campgrounds and picnic areas (NPS 2018). Because the only campground and picnic area within Peaks of Otter salamander’s range is near the Blue Ridge Parkway Visitor Center, a relatively well- developed area, permitted firewood collection is limited to a few dozen acres.

The collection of firewood may affect individual salamanders if the activity occurs where the species is present. We have no information to indicate that this activity has an AU-level or species- level effect. Therefore, we are not analyzing it further in our resiliency and viability analysis.

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3.1.4 Defoliation and Tree Damage by Insects

Defoliation of the forest canopy and tree mortality by insects like gypsy moths (Lymantria dispar dispar), emerald ash borer (Agrilus planipennis) and hemlock woolly adelgid (Adelges tsugae) can create openings in the canopy and change the microclimate of an area. Tree mortality may lead to changes in fire behavior, exotic plant invasions, slope instability and changes in hydrology (NPS 2018). European and Asian gypsy moth caterpillars feed on several hundred species of trees and shrubs including oaks, which make up nearly 40 percent of forests in the southern Appalachians (NPS 2018), including the Peaks of Otter salamander’s habitat.

According to Chazel (2005, p. 4), surveyors attempted to confirm a population of Peaks of Otter salamander that was documented in 1987 on a “steep hemlock-birch slope.” The surveyors located the hemlock stand, but noted that the hemlocks appeared to be heavily damaged by pests, likely the hemlock wooly adelgid. They did not find any Peaks of Otter salamander or other salamanders in the immediate area, suggesting that changes to the canopy affected the presence of Peaks of Otter salamander.

The treatment of forest pests through pesticide use can effect forest species including the Peaks of Otter salamander. Pesticides like Diflubenzuron, a synthetic arthropod growth regulator, and Bacillus thuringiensis kurstaki (BTK), an entomopathogenic bacterium, are used to control gypsy moths. Pesticides can directly affect salamander body mass, tail fat, follicle production, and hatchling success and also alter the prey base. Reardon (1995 pp. 44–46) looked at effects of Diflubenzuron, also called Dimilin 4, on red-backed salamanders in treated and untreated watersheds over 5 years. Although the study did not find significant differences in body fat, it did find that there was an increase in the consumption of hard-bodied prey. The author also suggested that future research should look at effects to juveniles and include at least 10 years post-treatment. A similar study by Sutton et al. (2005 In Strazanac and Butler 2005, pp. 130–131) examined the potential effects of BTK and found that BTK had little or no effect upon terrestrial and aquatic salamander density, species richness, or feeding ecology.

Chemical spraying is addressed in the Jefferson National Forest Management Plan’s 8E2-013 prescription, which states “biological pesticide controls of gypsy moth, hemlock woolly adelgid, and other detrimental species are permitted with full consideration of the effects on the salamanders, their microhabitat, and their prey. Non-target species-specific chemical insecticides are not permitted under the Management Plan” (USFS 2004, chapter 3 p. 135). According to Wright (2018a), the USFS has not treated for Hemlock woolly adelgid with biological or chemical methods in or near the range of the Peaks of Otter salamander. We are unaware of any information that indicates pesticide spraying has previously affected the Peaks of Otter salamander. While it is possible that the Peaks of Otter salamander could be affected by the use of pesticide spraying activities, we are uncertain whether this has happened in the past; therefore, we are not evaluating it further in our resiliency and viability analysis.

The possibility exists for increased occurrences of forest canopy defoliation due to one or more forest pests. Because the species’ range is relatively small, we acknowledge that if a severe, wide spread defoliation event happens, it could have catastrophic consequences for the species. However, we are unaware of information that can predict the frequency and extent of these events

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on the Peaks of Otter salamander’s habitat. Therefore, we are not evaluating it further in our resiliency and viability analysis, but recognize that the risk of such an event occurring remains even though it is not explicitly incorporated into future scenarios (see Chapter 5).

3.1.5 Tree Damage by Extreme Weather Events

Damage to forests by extreme weather events can alter the structure and condition of forests by defoliating the canopy or causing tree mortality or morbidity. Changes to the canopy and structure of a forest can alter the microclimate of an area. As in the previous section, tree mortality may also lead to changes in fire behavior, exotic plant invasions, slope instability and changes in hydrology (NPS 2018).

In early 1998, an ice storm occurred in the George Washington-Jefferson National Forest, causing extensive damage and considerably reducing the forest canopy in some areas (Reichenbach and Sattler 2000, p. 1). According to Kirk (2018d), in areas with severe damage, the forest was reduced to early successional, shrubby habitat, thus altering the structure and function of the forest in affected areas.

Effects of drought-induced mortality can influence micro and macro level ecological processes. It is difficult to determine the extent to which mortality is caused by droughts because there are many factors that can contribute to declining health prior to drought conditions. Berdanier and Clark (2016, p.21) found that for most species, drought stress contributes to deteriorating health over multiple years and that drought vulnerability may be increased by interactions with pathogens, fungi, insects or habitat. Because there are many interacting factors that affect tree mortality and morbidity, the risk of tree mortality from drought stress threatens forest structure and function at multiple time scales (Berdanier and Clark 2016, p. 17).

The potential exists for extreme weather events to occur in the future. Because the species’ range is relatively small, we acknowledge that if a severe event occurs and results in damages to large areas of suitable habitat, that event could have catastrophic consequences for the species. However, we are unaware of information that can predict reliably the frequency and extent of these events on the Peaks of Otter salamander’s habitat. Therefore, we are not evaluating it further in our resiliency and viability analysis, but recognize that the risk of such an event occurring remains, even though it is not feasible to explicitly incorporate (i.e., quantify) in the representative future scenarios (see Chapter 5).

3.1.6 NPS Maintenance Work

Within the NPS’ Blue Ridge Parkway, although there are no specific guidance documents for management considerations in areas where Peaks of Otter salamander is known to occur, maintenance work avoids Peaks of Otter salamander habitat or is performed during times of the year when the salamanders are inactive and underground (Cherry 2017; Cherry 2018). Most of the work in the Peaks of Otter area is focused on maintaining existing facilities, which are in developed areas of the Parkway including the campground, visitor center, and lodge. The NPS also maintains vistas areas to keep views from the Parkway open. These areas have been previously disturbed, are continuously maintained to clear trees and shrubs, and therefore are not considered suitable habitat.

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The NPS estimates that there are 10.6 ac (4.29 ha) of land within the Peaks of Otter area that are maintained as part of the vista project (Teague 2018). Maintenance of previously disturbed vista areas is not expected to have future impacts on the Peaks of Otter salamander and will not be incorporated into future scenarios (see Chapter 5).

Summary of 3.1 Mature Hardwood Forests

Activities or ecological processes that disrupt or remove the forest canopy, understory vegetation, and cover objects can have significant effects to the Peaks of Otter salamander because they reduce or eliminate moist areas (e.g., leaf litter, humus layers, rotting logs) necessary for the salamander’s breeding and sheltering, and limits foraging habitat and connectivity between the salamander’s AUs. These activities or processes include timber harvest and firewood collection, and defoliation or tree mortality due to insects, fire, and extreme weather events. Of these, timber harvest has been the primary influence, but is currently being managed, and expected to be managed in the future, through the HCA. Firewood collection, where it occurs in proximity to the species, may have an effect on individuals, but we have no information to indicate that this has an AU-level or species level effect. Defoliation due to insects, fire, and weather events has the potential to affect multiple AUs or the species as a whole, but we have no methods to reliably project these events.

3.2 Barriers Physical barriers to movement such as roads and streams can limit dispersal and movement of amphibian species, alter ecosystem processes and result in direct mortality to individuals. Habitat fragmentation by roads and dams can create several small populations from what were formerly large ones (Terwilliger 1991, p. 420).

3.2.1 Roads Roads can have both direct and indirect effects on amphibians (Jochimsen 2004, p. 2). Direct effects involve injury or mortality due to vehicle strikes or from road construction, and presumably maintenance activities. Indirect effects include habitat loss, fragmentation, and alteration of ecosystem processes at both fine and broad scales (physical, chemical, and biological). These changes may influence the behavior, survival, growth, and reproductive success of individual animals; cumulatively resulting in population-level consequences, also there may be cumulative effects on different species, influencing overall species richness and diversity in an area.

An individual’s vulnerability to road mortality is influenced by dispersal ability as well as the spatial scale and frequency of movements. More vagile species are more likely to suffer from road mortality (Jochimsen 2004, p. 8). Although there are caveats in drawing conclusions about the conservation of one species based on data from another, in the absence of detailed genetic studies on terrestrial salamanders like the Peaks of Otter salamander, data from red-backed salamanders are likely the best surrogate available (Marsh et al. 2008, entire). Wide logging roads and narrow paved roads are documented to be partial barriers to salamander movement (Gibbs 1998; deMaynadier and Hunter 2000 In Marsh et al. 2008). Roads decrease movement by 25 to 75 percent but not eliminating it entirely (Marsh et al. 2008, p. 605).

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Goff (2015, pp. 23–27) examined movement behavior of Peaks of Otter salamanders and red- backed salamanders in three treatments: paved roads, gravel roads and hiking trails and compared treatments to controls to assess movement behavior based on fragmentation type. He found that Peaks of Otter salamanders moved an average total distance of 1.41 m (4.6 ft) (Goff 2015, p. 11), but the project did not detect a significant effect of treatment or distance from the treatment edge on individual foraging movements. Instead, abiotic factors including moon phase and precipitation were significant predictors of movement. In addition, Chazel (2005, p. 4) documented the species crossing gravel and asphalt roads during rainy nights and crossing wet, grassy roads during the day, as well as observing juveniles in open grassy areas along unpaved roads. Salamanders living in close proximity to barrier edges may have adapted to living in such conditions and, therefore, did not show changes in foraging behavior as compared to salamanders farther from a treatment edge. Because salamander abundance is generally lower near road edges, Goff (2015, p. 23) suggests that individuals living in these areas would most likely have lower competition with other individuals and potentially higher food and cover availability.

There is some uncertainty about the extent to which wider paved roads may affect the Peaks of Otter salamander. For example, Terwilliger (1991, p. 420) states that the Peaks of Otter salamander “appears to have been split into two isolated populations by the construction of the Blue Ridge Parkway, which they apparently do not cross.” However, Kirk (2018a) and Croy (2018) received a report of the species crossing the Blue Ridge Parkway on a rainy day. The Parkway likely inhibits genetic exchange in the Peaks of Otter salamander, but does not completely restrict it (Kirk 2018a). This conclusion is consistent with Marsh et al.’s (2008, p. 609) finding, based on microsatellite data, that the Blue Ridge Parkway did not appear to be affecting genetic population structure of the red-backed salamanders.

3.2.2 Streams We have no information on how streams affect the Peaks of Otter salamander’s dispersal capability, but such data do exist for the red-backed salamander. Red-backed salamanders use terrestrial habitats in all phases of their life history, and they generally prefer upland areas to streamside habitats. When red-backed salamanders do colonize streamside habitats, they may be rapidly displaced by larger, more stream-adapted salamanders. Thus, both biotic and abiotic factors may result in stream habitats forming barriers to terrestrial salamander dispersal (Marsh et al. 2007, p. 320). Marsh et al. (2007, pp. 324–325) used displacement experiments and analysis of microsatellite markers to investigate the barrier effects of low-order streams on red-backed salamanders. Results for both the displacement experiment and the population genetic study suggest that streams act as partial barriers to dispersal and gene flow in red-backed salamanders. Streams reduced the return rates of salamanders and increased genetic differentiation among subpopulations. The displacement experiment suggested that the barrier effects of streams might be particularly strong for smaller salamanders since salamanders returning across the stream were significantly larger than salamanders returning through the forest (Marsh et al. 2007, p. 324). Because streams appear to be only partial barriers to dispersal, it is unlikely that isolation by a single, small stream would lead to substantial genetic differentiation and local adaptation. However, large streams or multiple small streams have the potential to lead to differentiation in red- backed salamanders given the broad geographic range and low dispersal rates (Marsh et al. 2007, p. 325).

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While these data are informative, it may not be wholly applicable to the Peaks of Otter salamander. Unlike the wider ranging red-backed salamander, most of Peaks of Otter salamander’s range is along mountaintops and in high elevation areas. The streams within its range are headwater streams and small second order streams. Given their size, it is unlikely that these streams are a significant barrier to movement or dispersal of Peaks of Otter salamander.

3.2.3 Summary of Barriers The best available information indicates that some roads and streams may be partial barriers to dispersal of individual salamanders. However, it is unlikely that the roads and streams within the Peaks of Otter salamander’s range are a barrier to gene flow and having an AU-level or species level effect. Therefore, we will not be further considering barriers in our resiliency and viability analysis.

3.3 Hybridization In general, hybridization (inbreeding between genetically unique species) is recognized as a potential driver of extinction because it can lead to the production of less fit hybrids or the replacement of one or both species with hybrids (Page et al. 2018 p.1). The area occupied by Peaks of Otter salamander is surrounded by the morphologically and ecologically similar red-backed salamander. While hybridization among closely related plethodontids is well known, it is unclear whether these two species are hybridizing. According to Wicknick (1995, p. 18), the two species seem to be “ecological equivalents” and “are not known to hybridize (Highton, 1972).” Page et al. (2018, p. 1), used microsatellite markers to investigate evidence of hybridization between Peaks of Otter salamander and red-backed salamanders. Researchers found that hybridization was either infrequent or non-existent (Marsh 2018a), and allelic richness and heterozygosity are substantially higher in Peaks of Otter salamander than red-backed salamanders. These findings suggest that Peaks of Otter salamander may have evolved in situ in this region for longer than red-backed salamanders, and that, despite its large geographic range, the red-backed salamanders is a comparatively recent invader. Although not detailed in the literature, it is possible that historic land management has influenced this dynamic. If hybridization between Peaks of Otter salamander and red-backed salamanders is occurring, it likely is happening at levels (i.e., individuals) such that it has not had a species’ level affect, therefore hybridization will not be considered in the current condition analysis (see chapter 4) or directly in our future condition analysis (Chapter 5). 3.4 Competition Competition for resources may occur between the red-backed salamander and the Peaks of Otter salamander but the overall effect to either species is largely unknown. As explained above, the red- backed salamander’s range encompasses the Peaks of Otter salamander’s entire range. According to Kniowski and Reichenbach (2009, p. 285) “These two species are similar in several ways including: (1) foraging on foggy or rainy nights, sometimes on vegetation (Jaeger 1978; Kramer et al. 1993); (2) consuming soft-bodied prey (Mitchell et al. 1996, Arif et al. 2007); (3) limited movements having home ranges typically < 1 m2 (Kramer et al. 1993; Mathis 1991); (4) use of leaf litter, rocks, logs and soil microhabitats (Taub 1961; Kramer et al. 1993; Wicknick 1995); and (5)

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defending sites (Jaeger et al. 1982; Wicknick 1995). They are also similar in size with adult red- backed salamanders ranging from 6.5–12.5 cm (2.5–4.9 in) total length compared with 8–13 cm (3.1–5.1 in) for Peaks of Otter salamander (Petranka 1998).” Because the two species have similar behaviors and size, this may lead to interspecific competition in sympatric (overlapping) areas and may limit the distribution of Peaks of Otter salamander, especially when the red-backed salamander displays agonistic behavior and territoriality (Kniowski and Reichenbach 2009, p. 285).

In a 1974 paper, Jaeger stated that Peaks of Otter salamander appeared “to be in the process of being displaced” by the red-backed salamander (1974, p. 38). Reichenbach and Brophy (2017, p. 1) hypothesized that in certain allopatric (non-overlapping) areas, Peaks of Otter salamander’s range may be restricted because of the red-backed salamander. In addition to potential displacement or range restriction, densities of Peaks of Otter salamanders are lower in sympatric areas compared to allopatric areas. Kniowski and Reichenbach (2009, p. 285) found that density for Peaks of Otter salamander in sympatry with red-backed salamander was 0.6/m2, which is lower than previously recorded for Peaks of Otter salamander in allopatry (1.6–3.3/m2). While the density of Peaks of Otter salamander in sympatry with red-backed salamander was 0.6/m2, the combined density for both species at sites (1.1 salamanders/m2), is comparable to nearby densities for Peaks of Otter salamander in allopatry noted above. The combined density suggests the species split available resources when sympatric (Kniowski and Reichenbach (2009, p 292). Kniowski also found that in addition to lower adult densities in sympatry, Peaks of Otter salamander has depressed growth rates as compared to allopatric areas (2006, p. 20–21).

Some research suggests that competition in sympatric areas may not be affecting the Peaks of Otter salamander, and some studies suggest that the sympatric areas may have stabilized (Kniowski and Reichenbach 2009, p. 292; Aasen and Reichenbach 2004, entire). According to Wicknick (1995, pp. 71–72), competition between the Peaks of Otter and red-backed salamanders for cover objects was weakly supported and experimental evidence did not support that competition is occurring. Results from Arif et al. (2007, pp. 848, 851–852) suggest that aggressive interactions with Peaks of Otter salamander restrict the distribution of red-backed salamanders and not the converse, whereas Peaks of Otter salamander appears to be limited by abiotic climatic factors. They found that a lack of niche-partitioning of diet in sympatry suggests exploitative competition is unlikely to be an important factor, and that interference competition might be important in sympatric communities; but they state that competition would limit the geographic expansion of red-backed salamanders, not Peaks of Otter salamander, since Peaks of Otter salamanders are more aggressive than red- backed salamanders.

Elevation may limit the distribution of Peaks of Otter salamander in areas where it is allopatric with red-backed salamanders, due to physiological factors such as intolerance to the higher temperatures and lower humidity associated with lower elevations (Reichenbach and Brophy 2017, p. 7). Grant et al. (2018, p. 7559), using a modeling framework that directly accounts for the presence of the red-backed salamander, found that climate rather than competition, is a chief determinant of the lower elevational range limit of P. Shenandoah, another narrow range endemic in the Plethodon genus. Thurow (1957, p. 64), suggested that the red-backed salamander is more resistant to increasing warmth and evaporation rates than Peaks of Otter salamander, which might be expected from its range and habitat. These characteristics could result in red-backed salamander outcompeting Peaks of Otter salamander if future climate conditions favor red-backed salamanders,

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but direct competition with red-backed salamander may not increase the extinction risk for Peaks of Otter salamander under future climate conditions. In addition, if Peaks of Otter salamanders are restricted by climate, any timbering in the contact zone would likely favor the red-backed salamander. This assumption is based on research suggesting that timbering “may increase temperature and decrease moisture on the forest floor (Covington 1981; Ash 1995), and that the red-backed salamander is likely more resistant to increased temperatures and evaporation rates (anecdotal information in Thurow 1957; Arif et al. 2007)” (Kniowski and Reichenbach 2009, p. 292).

Simberloff (1982, p. 241) stated that when species compete with one another, “effects are usually moderated by other factors (e.g., weather, predators, pathogens) that keep populations below levels at which exclusion would occur, or else each competitor is favored in a different set of times and/or places and this fact combined with normal individual movements keep all species in the system. Interspecific competition is as likely to be by interference as by exploitation, and is frequently affected by biological idiosyncrasies of the individual species. Chance plays a major role in many potentially competitive interactions, and there is good evidence that many species that do compete with one another do so rarely or intermittently, and at most times their population dynamics are governed by other forces.” 3.4.1 Summary of Competition The best available information indicates that competition with the red-backed salamander may be affecting individual Peaks of Otter salamanders, however it is unclear as to what extent competition is affecting the species overall. Research has indicated that densities of Peaks of Otter salamanders are lower in areas where they are sympatric with red-backed salamanders as compared to allopatric areas, suggesting that the two species may be competing for certain resources like prey, nesting sites or good foraging habitat. Because changes to climate or weather may favor one species and resource availability could change over time, we will evaluate potential effects of competition on species viability in our future condition analysis.

3.5 Infectious Disease The highly infectious amphibian pathogens Batrachochytrium salamandrivorans (Bsal), Batrachochytrium dendrobatidis (Bd) and ranavirus pose significant threats to a number of amphibians worldwide. In a study by Martel et al. (2014, pp. 630–631), researchers screened more than 5,000 amphibians from across four continents and combined experimental assessment of pathogenicity with phylogenetic methods to estimate the threat that this infection poses to amphibian diversity. Forty-one out of forty-four of the Western Palearctic salamanders (Salamandridae and Plethodontidae) rapidly died after infection with Bsal. The propensity of Bsal to infect these species was confirmed by its ability to successfully invade the skin of several urodelan (newt and salamander), but none of the anuran (toad and frog), species. The outcome of exposure of three lineages of plethodontids (a family comprising 66 percent of global urodelan diversity) to Bsal ranged from a lack of any detectable infection (Gyrinophilus), to transient skin invasion (Plethodon) and lethal infection (Hydromantes), making it likely that other species in this large family are vulnerable.

The Bsal, Bd and ranavirus diseases have not been detected in Peaks of Otter salamanders, and they have not been documented to effect other amphibians within Peaks of Otter salamander’s range.

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Dr. Matthew Becker at Liberty University in Virginia is currently researching emerging infectious diseases in Peaks of Otter salamander and will be looking for these diseases over the next few years (Becker 2017). According to Muletz et al. (2014, p. 4), Bd has been present in the Appalachians at least since the 1970s and in laboratory infection trials Plethodon salamanders are susceptible to Bd and some die from the infection. Although Highton (2005, pp. 34–46) had documented that widespread declines of terrestrial Plethodon salamanders occurred by the 1980s throughout the central Appalachian Mountains, the cause of the perceived decline is unknown and it is unlikely that known amphibian pathogens (e.g. Bd, Bsal and ranavirus) are the cause (Muletz et al. 2014, p. 6).

Given the species’ small range, if a highly infectious disease spread to the Peaks of Otter salamander’s range, it could be catastrophic for the species. Because, however, these diseases have not been detected in Peaks of Otter salamanders, research indicates that known amphibian pathogens are unlikely to be the cause for past declines in Plethodon populations, and we are unaware of information that can predict reliably the frequency and extent of a disease outbreak in the Peaks of Otter salamander’s habitat, we are not evaluating disease in our resiliency and viability analysis. We do recognize, however, that the risk of such an event occurring remains even though it is not explicitly incorporated into our future scenarios (see Chapter 5).

3.6 Changing Climate Conditions The frequency and intensity of extreme heat and heavy precipitation events are increasing in most continental regions of the world, and are consistent with expected physical responses to a warming climate. The frequency and intensity of extreme high temperature events are virtually certain to increase in the future as global temperature increases. Extreme precipitation events will very likely continue to increase in frequency and intensity throughout most of the world. Observed and projected trends for some other types of extreme events, such as floods, droughts, and severe storms, have more variable regional characteristics (Wuebbles et al. 2017, entire). The primary sources of climate data and predictions used in this assessment were the two most recent U.S. Global Change Research Program (USGCRP) reports: Third National Climate Assessment (NCA3, Melillo et al. 2014, entire report) and Fourth National Climate Assessment (NCA4; Wuebbles et al. 2017, entire). The NCA4 used the World Climate Research Programme’s (WCRP’s) Coupled Model Intercomparison Project (CMIP5) and primarily discussed two pathways: the highest radiative forcing pathway (Representative Concentration Pathway (RCP8.5) and the medium-low radiative forcing scenario (RCP4.5). For detailed descriptions of these scenarios, see Hayhoe et al. (2017, pp. 135–149). As part of NCA4, state summaries were prepared by the National Oceanic and Atmospheric Administration (Runkel et al. 2017, entire) with maps of the projections using the RCP8.5 path. We primarily use RCP8.5 in our analysis based on current trend data in global emissions (Jackson et al. 2017, entire), the long-lasting influence of greenhouse gases already in the atmosphere (Collins et al. 2013, p. 1102–1105; Mauritsen and Pincus 2017, entire), and analysis of expected emissions through 2050 (U.S. Energy Information Agency 2017, entire). The USGCRP stated with very high confidence that the observed increase in global carbon emissions over the past 15 to 20 years has been consistent with higher scenarios such as RCP8.5 (Wuebbles et al. 2017, p. 152–153; Carter et al. 2018, p. 751). It is therefore reasonable to conclude that changes from now through

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mid-century will also be closer to RCP8.5 than to RCP4.5. We however did use RCP4.5 in one of the future scenarios (see Section 5.2.2) to incorporate the suite of plausible future scenarios. The following temperature and precipitation graphs were created from the U.S. Resilience Climate Toolkit website. The Peaks of Otter salamander’s range occurs in Bedford, Botetourt and Rockbridge counties of Virginia. Because these counties had identical historical observations and the same predicted changes to annual average daily mean temperature and annual total precipitation to 2100, we used results from Botetourt County for our analysis.

Figure 7. Low emissions scenario Annual Average Daily Mean Temperature for Botetourt Co. Historical observations to 2100.

Figure 8. High emissions scenario Annual Average Daily Mean Temperature for Botetourt Co. Historical observations to 2100.

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Figure 9. Low emissions scenario Annual Total Precipitation for Botetourt Co. Historical observations to 2100.

Figure 10. High emissions scenario Annual Total Precipitation for Botetourt Co. Historical observations to 2100.

According to future predictions, Virginia is projected to experience a range of impacts from climate change from sea-level rise along the coast to increasing air and water temperatures and changes to precipitation patterns. These climate change impacts will directly affect wildlife and their habitats as well as exacerbate already existing stressors (Kane et al. 2013, p. 1). Salamanders are susceptible to the effects of changing climates due to their restrictive physiological requirements (e.g., respiratory mechanism) and low vagility (i.e., patterns of movement) (Sutton et al. 2015, p. 1). Temperature and precipitation are the climatological variables that would most affect habitat and resource needs of the Peaks of Otter salamander. These variables could affect many different components of the habitat or resources that Peaks of Otter salamanders need to persist on the landscape. Research indicates that temperature has a controlling effect on the range size and dispersal capability of Plethodon salamanders and climate-based models may be appropriate tools to evaluate these species’ distributions (Milanovich et al. 2010, p. 2). According to Welsh and Droege (2001, pp. 560–561):

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“it is in the litter and upper soil layers that moisture is most important to terrestrial salamanders and many other hydrophilic forest-dwelling organisms. Because they must continuously maintain moist skin in order to respire, salamanders require constant contact with moist soil or litter. Surface activity, where the majority of feeding and mating occurs (Houck and Verrell 1993; Tilley and Bernardo 1993), requires high relative humidity. Even while out during high moisture conditions on the surface, salamanders are in a dehydrating situation and must return to litter and subterranean refugia for complete hydration (Jaeger 1978). Changes in forests due to disturbance or climatic change that result in a drying of upper soil and litter layers decreases the capacity of the forest to support salamanders. A tall, multilayered canopy buffers weather extremes (i.e., wind, insolation, and fluctuations in temperature), providing stable, within-stand microclimates (Chen et al. 1999). The loss of canopy, or creation of large openings, reduces or removes this buffering element, and much of the evapotranspired moisture that maintains high relative humidity within the interior of the forest is lost. Increased wind effects, greater solar radiation reaching the forest floor, and temperature extremes reduce the available moisture on the forest floor (Chen et al. 1999). These factors shorten the period of suitable environmental conditions for surface activity by salamanders and other moisture-dependent biota of the forest floor.”

According to the National Climate Assessments (NCA), in the Southeastern region, which includes the Peaks of Otter salamander’s range, climate models project increases in both temperature and extreme precipitation events for both RCP4.5 and RCP8.5 (Carter et al. 2014 p. 389–399; Carter et al. 2018, pp. 745, 751). Climate change is expected to intensify the hydrologic cycle and increase the frequency and severity of extreme events like drought and heavy rainfall, and these changes are expected to transform many ecosystems in the Southeast (Carter et al. 2018, pp. 768–776). Authors of the NCA have high confidence in the documentation that projects increases in air temperatures (but not in the precise amount) and associated increases in the frequency, intensity, and duration of extreme heat events; in addition, the projections for increases in temperature are more certain in the Southeast than projections of changes in precipitation, although both are expected to increase (Carter et al. 2014, pp. 397–417). Increases in the frequency, intensity, and duration of extreme heat events could limit the amount of time Peaks of Otter salamanders are able to be surface-active to forage for food and mate. Precipitation is also predicted to increase during all months of the year (see figures 9 and 10), potentially resulting in enhanced surface conditions. These enhanced conditions could prove favorable by extending the foraging and mating seasons. However, increased precipitation may not completely counter the effects of increased temperatures, because even during high moisture surface conditions, salamanders are in a dehydrating situation and must return to litter and subterranean refugia to rehydrate. Changes to feeding and mating behavior could also result in lowered reproductive output, less fit individuals, and fewer individuals over time.

Reichenbach and Brophy (2017, p. 1) found that Peaks of Otter salamander had a significantly lower critical thermal maximum (CTM) and higher dehydration rates relative to red-backed salamanders. For example, the CTM least square means was 33.3 °C and 34.4 °C (91.9 °F and 93.9 °F) for the Peaks of Otter salamander and red-backed salamander, respectively, and the dehydration rates for a mean salamander mass of 2.3 grams was 16.3 and 15.2 mg/cm2 * h (mass loss per respiratory surface area per hour) for the Peaks of Otter salamander and red-backed salamander, respectively (p. 12). Similarly, Thurow (1957, p. 64), suggested that the red-backed salamander is

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more resistant to increasing warmth and evaporation rates than Peaks of Otter salamander. These characteristics could result in red-backed salamanders outcompeting Peaks of Otter salamander, if future climate conditions favor red-backed salamanders. A competition study between the red- backed salamander and Shenandoah salamander (Plethodon shenandoah) under four different climate scenarios suggested that resource use was normalized under future climate conditions and that direct competition may not directly increase the extinction risk for Shenandoah salamander under future climate conditions (Dallalio et al. 2017, p. 194). This may also be true for Peaks of Otter salamanders, but the effects of climate change and interactions with red-backed salamanders on Peak of Otter salamanders at an individual and population level are currently unknown.

Climate change will affect abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents, and interactions between the different agents may amplify disturbances (Seidl et al. 2017, p. 395). Changes in climate conditions can also affect the composition and structure of forests, where more heat tolerant native or nonnative species may tend to migrate and replace historically dominant tree species (Kane et al. 2013, pp.17–18). Changing forest composition could have significant impacts on a number of important issues, including wildfire, water quality, invasive species, and botanical diseases. Under changing climate conditions, invasive species such as the gypsy moth may become more prevalent in the State and others may become more destructive (Hellmann et al. 2008; Virginia Invasive Species Working Group, 2012 In Kane et al. 2013, p. 49).

Region-wide projected changes in temperature and precipitation will likely impact microclimate environments differentially across a landscape. Microclimatic variables, particularly solar radiation, ground surface temperature, and soil temperature, are highly sensitive to changes in the overstory canopy and exhibit relatively high spatial and temporal variability within a forest (Reifsnyder et al. 1971 and Chen and Franklin 1997 In Chen et al. 1999, p. 289). Elevation, slope and aspect, have direct effects on temporal and spatial patterns of microclimate and because microclimates influence complex biological processes in a forest, it is difficult to predict how those interactions may change over time.

Work by Sutton et al. (2015, entire), where they modeled predicted changes in climate niche and refugia for salamanders in the northeast, indicates that salamanders in the Plethodontidae family will lose a greater proportion of their climate niche compared to other families. Although Peaks of Otter salamanders were not included in the analysis, they found that three of the five species predicted to lose the greatest proportion of their respective climate niche are forest-dwelling, endemic species that occupy the highest elevation environments in their respective ranges, like the Peaks of Otter salamander (p. 19). However, Sutton et al. (2015, p. 6) found that the Central Appalachian ecoregion is projected to provide refugia for the greatest number of salamander species under both the current and projected climate scenarios. Landscapes that provide relatively consistent local temperature and moisture conditions in the face of climate change offer the greatest potential as habitat and climate refugia for vulnerable species.

Future climate change is expected to alter the thermal humidity in high elevation habitats. Cloud moisture and cloud base height (CBH) can influence abiotic factors like humidity and microhabitat. Richardson et al. (2003, entire) found that CBH increased across the Appalachians over a 26-year period (1973 to 1999), although there was a slightly negative trend at the Roanoke, VA station,

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which is the closest station to the Peaks of Otter salamander’s range. Grant et al. (2018, p. 7559) suggest that for P. shenandoah, even a small increase in CBH will greatly reduce the species’ total occupied extent. A rising CBH could lead to a range reduction for Peaks of Otter salamanders, especially along the lower elevations at the edge of their range.

It is possible that the predicted increase in rainfall will counter the effects of increased temperature in some areas. Research in the Great Smokey Mountains National Park aimed to address the “downscaling” of climate models applies in a smaller scale geographical context (Fridley 2009, entire). Fridley found that near-stream locations are warmer in the winter and cooler in the summer, and sites of low elevation more closely track synoptic weather patterns than do wetter high-elevation sites. These findings suggest a strong interplay between near-ground heat and water balances and indicates that the influence of past and future shifts in regional temperatures on biota may be buffered by soil moisture surfeits from high regional rainfall. Because the Appalachian Mountains are highly topographically variable, it is unlikely that predicted changes in temperature and precipitation will occur evenly across the landscape. 3.6.1 Summary of Changing Climate Conditions

The effects of changing climate conditions are likely acting upon the species currently by altering characteristics of the habitat including prey, temperature and moisture, however it is unclear if these changing conditions are affecting individuals, or if there is an AU-level or species level effect. Changes to temperature and precipitation are projected to continue into the future and will affect habitat and resource needs of the Peaks of Otter salamander. These changes may also exacerbate already existing stressors. Because changing climate conditions are projected to continue into the future and increase under certain forecasts, we will evaluate changing climate in our future condition analysis.

3.7. Other Influences Considered 3.7.1 Collection Collection and research projects that affect the Peaks of Otter salamander are permitted through the Virginia Department of Game and Inland Fisheries and through special use permits granted by the USFS. Although the number of animals allowed to be taken from the wild is limited, collection affects individual salamanders. We have no information to indicate that this activity has an AU- level or species-level effect. Therefore, we will not discuss it further in our resiliency and viability analysis. 3.7.2 Predation Natural predators of the Peaks of Otter salamander have not been reported, but probably include ring-necked snakes (Diadophis punctatus), garter snakes (Thamnophis sirtalis), turkeys (Meleagris gallopavo), shrews (Blarina spp., Sorex spp.), skunks (Mephitis spp., Spilogale spp.), opossums (Didelphis virginianus) and black bears (Ursus americanus) (USFWS 1997, p. 5; Messenger et al. 2011 p. 582; Kleopfer 2018). Bradshaw (2010, p. 55) found that with edge effects created by fragmentation in forested habitat, snakes and other predators are better able to locate salamanders that occur in the interior of the forest; direct sunlight heating the ground and rocks in these edge areas is more attractive to snakes compared to cover objects in the forest interior that receive

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indirect sunlight. We are unaware of any information indicating that predation of the Peaks of Otter salamander is occurring, or likely to occur, at other than natural levels (i.e., individuals). 3.8 Other Conservation Actions The Peaks of Otter salamander is listed as a Tier I Species of Greatest Conservation Need in the Virginia Wildlife Action Plan and as a G2/S2 species by the Virginia Division of Natural Heritage (VDGIF 2015, Chapter 3 p. 45; Roble 2016, p. 12). The USFS recognizes the Peaks of Otter salamander as a sensitive species (USFS 2004, chapter 3 p. 129). State, Federal, and academic entities have carried out a number of research projects involving the Peaks of Otter salamander. The USFS has funded research led by biologists at Liberty University to determine the impacts of different types of timbering on the species. There have also been a number of studies that have aimed to describe the species’ life history, distribution, population density and to determine physiological responses to the environment and other salamanders. Vincent Farallo at Virginia Polytechnic Institute will conduct a future investigation into habitat niche modeling. 3.9 Summary of Factors Influencing the Species The primary influences to the Peaks of Otter salamander’s viability currently and in the future are (1) activities that disrupt or remove the forest canopy, understory vegetation, and cover objects (e.g., primarily timber harvest); (2) competition with red-backed salamanders; and (3) changing climate patterns of increasing temperatures and changes in precipitation patterns. Timber harvest is currently being managed and is expected to be managed in the future, through the HCA. The effects of competition with the red-backed salamanders are largely unknown; however, literature has shown that densities of Peaks of Otter salamanders are lower in sympatric areas than in allopatric areas. While it is unknown if or to what extent climate change is currently affecting the species, it may have deleterious effects in the future. All three of these factors will be considered in the future condition analysis. Predation and collection are not currently, or expected to be, having a major effect on the species. Other factors (e.g., fire wood collection, roads, predation, prescribed fire) may be having an individual effect, but are not expected to rise to the level of affecting AUs or the species as a whole. Additional factors (e.g., wildfire, defoliation of forest canopy by invasive pests) are not currently affecting individuals, AUs, or the species. While we have no method to project when and where those events may occur in the future, we do recognize that they have the potential to be catastrophic for the Peaks of Otter salamander (e.g., unknown risk of occurrence, but high risk of extinction should they occur). Infectious amphibian pathogens like Bsal, Bd or ranavirus are a potentially catastrophic threat to the Peaks of Otter salamander. Because we lack information on the potential future emergence or spread of disease, we did not incorporate this threat in forecasting future conditions of the species. This remains a gap in our analyses that should be addressed as new information is obtained, especially as we anticipate that the level of disease risk is likely to increase in the future.

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CHAPTER 4 – CURRENT CONDITION 4.1 Methodology To assess the current condition of Peaks of Otter salamander, we looked at a number of parameters influencing demographics and habitat. Analysis Units (AUs) were delineated by topography and landscape characteristics, as well as clusters of species’ observations. There are a total of 20 AUs that span the range of the species (table 4). We assumed that natural breaks in topography and gaps in observation data indicate meaningful boundaries or disruptions in connectivity between units. However, we do not assume that those boundaries are barriers to movement, rather that they represent a mechanism by which to evaluate the species in the absence of a biological population substructure.

Table 4. Peaks of Otter salamander Analysis Units.

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Figure 11. Spatial depiction of Peaks of Otter salamander Analysis Units (AUs).

We used the best available data to assess the demographic and habitat parameters and then scored the AUs according to a condition category. The demographic parameters used were the density of salamanders and reproductive output (based on elevation). The habitat parameters used were habitat quality, allopatry/sympatry, and habitat management. In general, if demographic or habitat metrics for a given unit were considered poor they were categorized as “low” condition, and if those metrics were considered optimal for the species’ viability they were categorized as “high” condition, with “medium,” and “medium/high” conditions in between. See Appendix A for more details. 4.1.1 Density of Salamanders To determine the density of Peaks of Otter salamanders in each AU, the number of individuals per m2 was calculated based on the best available survey data. Values higher than 2/m2 were considered “high,” values between 1.1/m2 and 2/m2 were considered “medium/high,” values between 0.06/m2 and 1/m2 were considered “medium” and values between present and 0.05/m2 were considered “low.”

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4.1.2 Reproductive Output Although Peaks of Otter salamander’s range is relatively small, research has found that there are certain behavioral differences between salamanders at different elevations throughout the range (see 2.3 Range, Distribution and Abundance and 2.4.2 Reproduction). Based on information from Reichenbach and Brophy (2017, p. 13), elevations between 800 m and 1,000 m were considered “high” condition, elevations above 1,000 m were considered “medium/high” condition, elevations between 600 m and 900 m were considered “medium” condition, and elevations between 400 m and 600 m were considered “low” condition. Reproductive output for current condition is scored based solely on elevation; because we do not have available information, regarding climate conditions at the time surveys were conducted. Therefore, we are using the elevation as a proxy for assumed climatic conditions. However, for future conditions, both elevation (which does not vary over time) and climate conditions will be factored into the condition score for this metric. 4.1.3 Habitat Quality Habitat quality reflects the on-the-ground conditions necessary for Peaks of Otter salamander to breed, feed, and shelter. We considered forest type and percent canopy closure as surrogate measures for habitat quality. Habitat quality is presumed to be in “high” condition if there is high canopy cover and intact, productive hardwood forests. We assumed that if forest type and canopy closure were high quality, then there would be adequate surface cover, litter and underground refugia; moist, well-developed soils; abundant, high quality prey; moderate soil and air temperature. If canopy cover was low and forest type was poor quality, we assumed that there will not be adequate surface cover, litter and underground refugia; soils would be dry and compacted; prey would be limited or poor quality; soil and air temperature would not be moderated; and relative humidity would not be adequate and habitat quality was considered to be in “low” condition. 4.1.4 Allopatry/Sympatry As discussed above in section 3.4 Competition, many studies have attempted to determine the extent and effect of sympatry of Peaks of Otter salamander with red-backed salamander. If there has been no documentation of sympatry within the AU, the unit was considered in “high” condition. If Peaks of Otter salamander is in sympatry with red-backed salamander within the unit and there were many observations of red-backed salamander within or near the unit boundary, it was considered to be in “low” condition. 4.1.5 Habitat Management Areas within primary habitat conservation areas, areas that were located within a prescription that did not allow timber harvest, or were on NPS land were considered to be in “high” condition. Areas within USFS primary or secondary habitat areas were considered to be in “medium/high” condition. Areas within a secondary habitat conservation area with limited timbering were considered “medium” condition. Areas within a secondary habitat area and with timber harvest allowed, areas that fell outside of HCA primary or secondary conservation areas, or USFS or NPS managed areas were considered “low condition” because any form of timbering is permitted on private property. 4.1.6 Overall Condition The different population and habitat parameters for each AU were ranked as high, medium/high, medium or low and these ranking received a score from 4 to 1 respectively. No metric was

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weighted differently than the others. The scores were then added together for each AU and were used to calculate overall condition of high, medium/high, medium or low and correlated with the colors of green, yellow, orange and red, respectively, in the tables and maps. 4.2 Current Condition We define viability as the ability of the species to sustain a healthy population within a biologically meaningful timeframe. Using the SSA framework, we describe the species’ current viability in terms of Resiliency, Redundancy, and Representation. 4.2.1 Resiliency Resiliency describes the ability of populations to withstand stochastic events. We can measure resiliency based on metrics of population health. Highly resilient populations are better able to withstand disturbances such as random fluctuations in birth rates (demographic stochasticity), variations in rainfall (environmental stochasticity), or the effects of anthropogenic activities. Small populations distributed over a limited range, like the Peaks of Otter salamander, are limited in their ability to rebound from stochastic events. For the purpose of this SSA, population factors and habitat factors in each AU were used to help provide a measure of resiliency across the salamander's current range. The AUs were ranked according to the following overall condition categories: high, medium/high, medium, and low (table 5 and figure 12).

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Table 5. The Peaks of Otter salamander’s AUs and their corresponding population and habitat parameters and current condition.

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Figure 12. Spatial depiction of the AUs and current condition scores. Considering both demographic and habitat parameters, of the 20 AUs, 2 are in medium condition, 13 are in medium/high condition and 4 are in high overall condition. The Visitor Center AU is likely extirpated. The last confirmed observation at this location was in 1987, and additional surveys in 1990 at the same location had negative results (DGIF Virginia Fish and Wildlife Information Service (VAFWIS) 2018). There have also been surveys near Sharp Top Mountain and Peaks of Otter salamanders were not found. Observation data of red-backed salamanders from the 1990s indicate that some surveys were conducted near the Visitor Center, picnic shelter and campground area, but there is no indication of search effort and it is possible that Peaks of Otter salamanders are present but have not been detected (DGIF VAFWIS 2018). Although the population of the Peaks of Otter salamander is small and geographically restricted, which makes it vulnerable to stochastic events, the salamander is presumed to be abundant throughout most of the AUs, and most of the habitat is in high or medium/high condition. The Peaks of Otter salamander currently has good resiliency across its range. 4.2.2 Redundancy Redundancy refers to the number of populations of a species and their distribution across the landscape, reflecting the ability of a species to survive catastrophic events. The greater the number

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of populations/subpopulations, and the more widely they are distributed, the lower the likelihood a single catastrophic event will cause a species to become extinct. The Peaks of Otter salamander is a narrow endemic with a naturally limited range. The salamander occurs between 400 and 1300 m (1,312 and 4,265 ft) in elevation in mature hardwood forest habitat. Except for the Visitor Center AU, the species appears currently to be well distributed and presumed abundant throughout its historical range. Because the Peaks of Otter salamander has 19 AUs in medium or better condition, we consider the species currently to have good redundancy across its range.

4.2.3 Representation Representation refers to the breadth of genetic or environmental diversity within a species and reflects the ability of a species to adapt to changing environmental conditions. The greater the diversity, the more successfully a species should be able to respond to changing environmental conditions. Although there is a lack of genetic information about the Peaks of Otter salamander, the species is presumed to be well distributed throughout its range. While we do not know the species' genetic diversity, the Peaks of Otter salamander does exhibit a behavior difference in its reproductive strategy across elevations. This behavioral difference may be important to the species' ability to adapt to future changes in environmental conditions. Morphological, behavioral, geographical, and niche diversity is often used as a surrogate for genetic diversity if such data are lacking. The Peaks of Otter salamander currently has good representation (number of AUs at varying elevations) across its range.

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CHAPTER 5 FUTURE SCENARIOS 5.1 Methodology As discussed in Chapter 1, for the purpose of this assessment, we define viability as the ability of the species to sustain AUs in the wild over time (in this case, 20 to 80 years, based on the range of best available climate model forecasts and the effects of conservation management). Using the SSA framework, we describe the species’ viability by characterizing the status of the species in terms of its Resiliency, Redundancy, and Representation. We have considered Peaks of Otter’s life history characteristics, identified the habitat and analysis unit requisites needed for viability (Chapter 2), reviewed the factors that may be driving the historical, current, and future conditions of the species (Chapter 3), and estimated the current condition of those needs through the lens of the 3Rs (Chapter 4). Next, we predict the Peaks of Otter salamander’s future conditions to inform us of the viability of the species. We used the demographic and habitat information to predict how the 19 extant AUs will respond to the primary factors likely to influence the species’ condition in the future. These influencing factors included habitat quality, sympatry with red-backed salamanders and climate change. Our analysis is limited to three future scenarios, which are representative examples from the potential range of plausible scenarios, and that describe how these stressors to the species may drive changes from current conditions. 5.2 Future Scenarios Projections of Peaks of Otter salamander Resiliency, Redundancy and Representation were forecasted using two timelines, approximately 20 and 80 years out (2040 and 2100). These timelines were chosen to correspond to the range of climate model forecasts and timeframes during which the effects of any applicable conservation management can continue to be implemented and realized, and are a reasonable timeframe for the species to respond to potential changes on the landscape. The 2100 timeline represents a potential longer-term trajectory for the species, but with a lower confidence in the outcome than in the 2040 projection.

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Table 6. Summary of Future Scenario Influencing Factors and How They Change Compared To Current Condition.

Influencing Scenario 1 Scenario 2 Scenario 3 Factor Climate RCP RCP 8.5 RCP 4.5 RCP 8.5 Climate Effects ↑↑a temperature and ↑a temperature and ↑↑a temperature and variation in variation in variation in precipitation precipitation precipitation Competition Same Same ↑↑a, exacerbated by changing climate conditions Habitat Same ↑↑a ↓a Management Habitat Quality Same ↑a ↓a , exacerbated by changing climate conditions and decreased habitat management Reproductive Same in 2040 Same ↓a Output projection, ↓a in 2100 projection Conservation Same ↑a Same Actions a: ↑ increase, ↑↑ greater increase, ↓ decrease, ↓↓ greater decrease, Same- continue as in Current Condition 5.2.1 Scenario 1 Under the Continuation Scenario, densities of the Peaks of Otter salamander remain the same or decrease slightly. To analyze potential effects of climate change, we used projected impacts based on the RCP 8.5 emissions scenario, as the latest data indicate that this is the current trajectory (Brown and Caldeira 2017, entire). Climate conditions will become warmer and there will be more variation in precipitation (Carter et al. 2014, pp. 398–399). Prey availability and quality, suitable nesting, foraging and breeding habitat may be diminished, affecting salamander densities and reproductive output of individuals within the population. Sympatry with red-backed salamanders will continue to occur at the periphery of the Peaks of Otter salamanders’ range, but the presence of red-backed salamanders will not negatively affect the number of individuals of Peak of Otter salamanders more than they are in the Current Condition. Infectious diseases like Bsal, Bd or ranavirus may be present in Plethodons in Virginia but are not shown to affect the number of individuals of Peak of Otter salamanders. Therefore, known amphibian pathogens remain unlikely causes for declines in these Plethodon populations. Although forest condition may be influenced by increasing temperatures and variable precipitation, habitat quality throughout the range will remain intact and forests will continue to have high canopy cover along with abundant surface objects and moist, well-developed soils. Maintaining high quality, mature, intact forests will help to mitigate for the impacts of climate change on habitat. Because of projected increases in temperature and variability in precipitation patterns, by 2100, habitat suitability in each AU will decline and climate may begin to limit the species’ range, but

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will not eliminate any AUs. By 2100, reproductive output is expected to decline in the lower elevation AUs. Further, the distribution of reproductive output is likely to shift upslope by the end of the century (Brand 2019). However, as described in Section 2.4.2, there also appears to be an upper elevational limit to optimal reproductive output (Reichenbach and Brophy 2017, p. 11). Thus, it is difficult to predict reliably how an anticipated future upslope shift will be expressed on the landscape over time. Under Scenario 1, optimal reproductive output may occur at the highest elevations of the species’ range as temperatures increase and resources become more scarce at lower elevations; however, we are uncertain if, or to what extent, this this shift will occur. Therefore, we did not include changes to reproductive output at higher elevations in the corresponding condition table or overall resiliency projections. Habitat management will continue as is, and management actions will continue to be implemented in ways that minimize or avoid impacts to the salamander. The USFS does not have plans to harvest timber in the Peaks of Otter salamander conservation area for at least 5 years, and the Jefferson National Forest Plan is not currently scheduled for any revisions (Wright 2018b and Kirk 2018c). The Habitat Conservation Agreement and forest management prescriptions continue to be implemented and active habitat management limits effects to the Peaks of Otter salamander. The collection of firewood within the species range will continue, but it will be limited to the campground area and within 100 ft of roads. In addition to the continued habitat management, the VDGIF will continue to coordinate with the NPS and USFS on state issued Scientific Collection permit applications that include the Peaks of Otter salamander. Table 7. Scenario 1, 2040 projection.

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Table 8. Scenario 1, 2100 projection.

5.2.1.1 Resiliency Considering population and habitat parameters under the Continuation Scenario for the 2040 projection, of the 20 AUs, 2 are in medium condition, 13 are in medium/high condition and 4 are in high overall condition while the Visitor Center AU is extirpated. In the 2100 projection, of the 20 AUs, 4 are in medium condition, 12 are in medium/high condition and 3 are in high overall condition while the Visitor Center AU is extirpated. Although the population of the Peaks of Otter salamander remains small and distributed over a limited range, reducing the salamanders’ ability to rebound from stochastic events, the salamander remains abundant throughout its range and most of the habitat remains in high to medium condition. In the continuation scenario, the Peaks of Otter salamander has good resiliency across its range.

5.2.1.2 Redundancy In the continuation scenario for both 2040 and 2100, salamander density in most of the AUs (19 of 20) is considered high to medium condition, and the population is spread across the range where different habitat types and elevations are being utilized. Under this scenario, the Peaks of Otter salamander has good redundancy across its range.

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5.2.1.3 Representation In the continuation scenario, the population is well distributed throughout the range at different elevations and within different Ecological Zones (figure 15). Under this scenario, the Peaks of Otter salamander has good representation across its range.

5.2.2 Scenario 2 Under Scenario 2, densities of the Peaks of Otter salamander remain the same or increase slightly but not beyond carrying capacity of the available habitat. The effects of climate change are forecast to proceed along the RCP 4.5 scenario, taking into account the refined projections of global temperature change proposed by Brown and Caldeira (2017, p. 47). As discussed in Section 3.5, whereas proceeding along RCP 8.5 is more likely, RCP 4.5 is still plausible and is used in the Fourth National Climate Assessment (Hayhoe et al. 2017, pp. 135–149). Thus, under Scenario 2 in the timelines for 2040 and 2100, increases in temperature and precipitation are less severe than is projected under Scenario 1. Climate conditions will become warmer and there will be more variation in precipitation, but effects to localized habitat will be mitigated as forests will remain intact and in good condition with high canopy cover. Changing conditions may alter prey availability and quality, suitable nesting, foraging and breeding habitat, affecting salamander densities and reproductive output of individuals within the population, but to a lesser extent than in Scenarios 1 and 3. Sympatry with red-backed salamanders will continue to occur at the periphery of the Peaks of Otter salamanders’ range, but the presence of red-backed salamanders will not negatively affect the number of individuals of Peak of Otter salamanders. In both the 2040 and 2100 timelines, sympatry will have less of an effect on the Peaks of Otter salamander so conditions in AUs with sympatry are improved. Infectious diseases like Bsal, Bd or ranavirus may be present in Plethodons in Virginia but are not shown to affect the number of individuals of Peaks of Otter salamanders and known amphibian pathogens remain unlikely causes for declines in these Plethodon populations. Habitat quality will remain high with a high percentage of canopy cover, abundant surface objects, and moist, well-developed soils will remain undisturbed. Maintaining high quality, mature, intact forests will help to mitigate the impacts of climate change on habitat. Habitat management will continue, but will be enhanced to benefit the Peaks of Otter salamander throughout its range. Secondary conservation zones will be managed more closely in line with primary conservation zones and timber harvest will be limited. Areas previously harvested for timber will be allowed to grow back into mature hardwood stands and will not be cut in the future. Because certain AUs will experience enhanced management for the benefit of Peaks of Otter salamander, the habitat condition metric in these AUs improve. Enhanced management includes allowing areas harvested for timber to regenerate back into suitable habitat for Peaks of Otter salamander. The collection of firewood within the species range will also continue to be limited to reduce impacts to suitable habitat. In addition to the enhanced habitat management, the VDGIF will continue to coordinate with the NPS and USFS on State issued scientific collection permit applications that include the Peaks of Otter salamander. Under Scenario 2, some individuals move into the Visitor Center AU, and become reestablished in the area. Natural reestablishment of Peaks of Otter salamanders into this AU is plausible because

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the habitat and elevation between this area and the area near Flat Top Mountain where Peaks of Otter salamanders have been detected (within 0.5 miles of the Visitor Center AU), is also suitable habitat with similar Ecozone characteristics and high canopy cover. Because the habitat quality in this AU is in medium condition with numerous observations of red-backed salamanders, the density of Peaks of Otter salamanders will remain in low condition.

Table 9. Scenario 2, 2040 projection.

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Table 10. Scenario 2, 2100 projection.

5.2.2.1 Resiliency Considering population and habitat parameters under Scenario 2 for both 2040 and 2100, all of the AUs are in medium, medium high or high overall condition. The Visitor Center AU changes to medium condition because it is reestablished, albeit at low density, within an area that has good quality habitat parameters. Although the population of the Peaks of Otter salamander remains small and distributed over a limited range, reducing the salamanders’ ability to rebound from stochastic events, the salamander remains abundant throughout its range and most of the habitat remains in high or medium/high condition. In Scenario 2, the Peaks of Otter salamander has good resiliency across its range. 5.2.2.2 Redundancy In Scenario 2, for both 2040 and 2100 projections, salamander density in most of the AUs (19 of 20) is considered high, medium/high or medium condition, and the population is distributed across the range where different habitat types and elevations are being utilized. The Visitor Center AU is in low condition as some individuals have immigrated into this AU, becoming reestablished. Under this scenario, the Peaks of Otter salamander has improved redundancy across its range. 5.2.2.3 Representation In Scenario 2, the population continues to be well distributed throughout the range at varying elevations and in different Ecological Zones (figure 15). Under this scenario, the Peaks of Otter salamander has good representation across its range.

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5.2.3 Scenario 3 Under Scenario 3, densities of the Peaks of Otter salamander will decrease due to a number of factors. Scenario 3 assumes that climate change will continue to occur at the rate it is currently being observed and the emissions trajectories will remain at the current level (RCP 8.5). Climate conditions will become warmer and there will be more variation in precipitation, which may reduce prey availability and quality, suitable nesting, foraging and breeding habitat. Competition with red-backed salamanders will increase as resources become more limited. Areas along the periphery of the Peaks of Otter range where they currently are sympatric with red-backed salamanders will be impacted more than areas in the core of the range. Hybridization between Peaks of Otter salamanders and red-backed salamanders could occur in sympatric areas, although this will likely continue to effect only individuals, not entire AUs. As conditions change into the future, hybridization could become a more important influencing factor, but because there are significant uncertainties about its effects, hybridization is not explicitly incorporated into the future condition. Infectious amphibian pathogens like Bsal, Bd or ranavirus may be present in Plethodons in Virginia, but are not shown to effect Peak of Otter salamanders and remain unlikely causes for declines in Plethodon populations. Habitat management will continue, but not to the extent that the management efforts protect important habitat for the Peaks of Otter salamander throughout its range. Under Scenario 3, secondary conservation areas will be managed in a way that maximizes allowable timber harvest under the HCA and Forest Management Plan. Because the HCA will continue to be implemented, overstory removal will not occur until approximately 20 years after the initial harvest, so habitat quality will have declined further in the 2100 projection. Additionally, the collection of firewood within the species range will continue and collection is maximized, reducing cover objects along roads and near campgrounds. The VDGIF will continue to coordinate with the NPS and USFS on State issued scientific collection permit applications that include the Peaks of Otter salamander. Forest condition will be negatively influenced by increasing temperatures and variable precipitation. Habitat quality will be degraded and there will be a reduction of canopy cover and surface objects, leading to drier, more exposed soils. Soils will be poorer and more compacted from timber harvest operations, limiting quality and availability of underground habitat and refugia for the Peaks of Otter salamander. The reduction in habitat quality will be exacerbated by changing climate conditions, leading to a reduction in habitat condition and overall condition in many AUs in both the 2040 and 2100 projections. Changing climate will also begin to limit the species’ range but will not eliminate any AUs. Reproductive output will decline in the lower elevation AUs in both the 2040 and 2100 projections. Further, the distribution of reproductive output is likely to shift upslope by the end of the century (Brand 2019). However, as described in Section 2.4.2, there also appears to be an upper elevational limit to optimal reproductive output (Reichenbach and Brophy 2017, p. 11). Thus, it is difficult to predict reliably how an anticipated future upslope shift will be expressed on the landscape over time. Under Scenario 3, optimal reproductive output may occur at the highest elevations of the species’ range as temperatures increase and resources become more scarce at lower elevations; however, we are uncertain if, or to what, extent this this shift will occur. Therefore, we did not include changes to reproductive output at higher elevations in the corresponding condition table or overall resiliency projections.

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An overall decline in habitat condition and an increase in competition for resources will affect salamander densities and reproductive output of individuals within the population, which will also lead to a reduction in overall condition in many AUs.

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Table 11. Scenario 3, 2040 projection.

Table 12. Scenario 3, 2100 projection.

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5.2.3.1 Resiliency Considering population and habitat parameters under the Scenario 3 for the 2040 projection, of the 20 AUs, 3 are in medium condition, 13 are in medium/high condition and 3 are in high overall condition while the Visitor Center AU is extirpated. In the 2100 projection, of the 20 AUs, 5 are in low condition, 2 are in medium condition and 9 are in medium/high condition, 3 are in high condition while the Visitor Center AU is extirpated. The population of the Peaks of Otter salamander remains small and distributed over a limited range, reducing the salamanders’ ability to rebound from stochastic events. The salamander is less abundant throughout its range and the habitat is in medium and medium/high condition. In the continuation scenario, the Peaks of Otter salamander has reduced resiliency across its range. 5.2.3.2 Redundancy In the continuation scenario for the 2040 projection, salamander density in 19 of the AUs is considered high, medium/high or medium condition. In the 2100 projection, salamander density in 14 of the AUs is considered high, medium/high or medium condition, while 5 AUs have low density and 1 remains extirpated. In Scenario 3, the population remains spread across the range where different habitat types and varying elevations are being utilized, although the overall condition of some AUs is reduced. Under this scenario, the Peaks of Otter salamander continues to have good redundancy across its range. 5.2.3.3 Representation In Scenario 3, the population is well distributed throughout the range but with lower densities of salamanders in some AUs. Under this scenario, the Peaks of Otter salamander retains representation across its range. 5.3 Summary of Species Viability This assessment describes the viability of the Peaks of Otter salamander in terms of resiliency, representation, and redundancy by using the best available commercial and scientific information. We used these parameters to describe current and potential future conditions regarding the species’ viability, and to address the uncertainty associated with potential future impacts and how they will affect the species’ resource needs, we assessed potential future conditions using three plausible scenarios. These scenarios were based on a variety of negative and positive influences on the species across its range, allowing us to predict potential changes in population and habitat parameters.

Except for the Visitor Center AU, the Peaks of Otter salamander continues to occupy most of its known historical range. The species is well distributed throughout its range, across varying elevations and habitat types, and it appears that there are some local adaptations, which may be important to the species' ability to adapt to future changes in environmental conditions. The species currently has good representation, redundancy and resiliency.

Future Viability

Estimates of current and future resiliency for the Peaks of Otter salamander are good, and most AUs are in medium or high condition. There are a number of potential threats that could negatively affect demographics or habitat, including habitat degradation or loss, competition, hybridization and

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disease, all which may also be exacerbated by changing climatic conditions. These factors were important for the assessment of future viability, but the extent to which these factors are currently affecting the species is largely unknown and it is difficult to determine their effects in the future. Because conservation measures that protect the species and its habitat are currently being implemented and have shown to be effective, it is likely that the species will remain resilient throughout its range into the foreseeable future. Given current and future projections of resiliency, the Peaks of Otter salamander is not particularly vulnerable to extirpation from stochastic events. If resiliency is maintained, the species will also have good representation and redundancy into the future.

Table 13. AUs with current and future conditions under each scenario.

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CHAPTER 6 KEY UNCERTAINTIES ● Competition—We are uncertain the extent to which competition with red-backed salamanders is affecting the population of Peaks of Otter salamanders. The literature is variable on this topic; however, we do recognize that in sympatric areas the densities of Peaks of Otter salamanders is lower than in allopatric areas. This uncertainty may have led us to under or overestimated viability. ● Survey data—All survey data, including recent and historical data, were used. There is a paucity of recent range wide survey data. If salamanders were historically present in an area and there were no recent survey data to confirm, they were considered still to be present in that area. This could have resulted in an overestimation of salamander density or presence within each AU, thus overestimating viability. ● Red-backed salamander data—Data from the Masterpoints2005, Masterpoly2005, and the DGIF VAFWIS database were used to determine the presence of red-backed salamanders in each AU. It is possible that are other data sources specific to red-backed salamander species’ observations, that if so, they were not used in the SSA analysis because they were not available to us. This could have resulted in an underestimation of red-backed salamander presence and an underestimation of the influence of sympatry in certain AUs; and, thus, an overestimation of the future viability of Peaks of Otter salamanders in some AUs. ● Visitor Center AU—There are historical observation data from 1987 of Peaks of Otter salamanders near the campground. Other surveys in the area in the 1990s found only red- backed salamanders, but there is no indication of the level of effort used during those surveys. Surveys have been conducted near Sharp Top Mountain and no P. hubrichti were found. It is possible that Peaks of Otter salamanders are present but have not been detected since the 1987 survey, thus underestimating viability. ● Thunder Ridge AU—There were two confirmed occurrences of Peaks of Otter salamanders in the Thunder Ridge AU in 1991. Both Marsh (2018b) and Huber (2019) have said that they visited this area and have never seen Peaks of Otter salamanders. It is possible that Peaks of Otter salamanders have been extirpated from this area, thus overestimating current and future viability. ● Disease—Disease is a potentially catastrophic threat to the Peaks of Otter salamander population. Because we lack information on the potential future emergence or spread of disease, we did not include this threat in forecasting future conditions. This remains a gap in our analyses that should be addressed as new information is obtained, especially as we anticipate that the level of disease risk is likely to increase in the future. Lack of inclusion in future scenarios overestimates the predicted viability. ● Reproductive output—Because we are unsure if or how optimal reproductive output may shift upslope as climate conditions change, we did not incorporate changes to reproductive output in the higher elevation AUs in the future condition tables and thus it was not incorporated into overall resiliency projections. Future viability could have been overestimated. ● Climate change—Because the direct effects of climate change spatially and temporally are unknown, especially when considering the effect on overall suitable habitat relative to the species’ critical thermal maximums and drying tolerance, future viability could be under or overestimated.

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REFERENCES CITED Aasen, Garth and Norman Reichenbach. 2004. Is the red-backed salamander (Plethodon cinereus) encroaching upon populations of the Peaks of Otter Salamander (P. hubrichti)? Catesbeiana 24:17–20.

Arif, Saad, Dean C. Adams and Jill Wicknick. 2007. Bioclimatic modelling, morphology, and behavior reveal alternative mechanisms regulating the distribution and two parapatric salamander species. Evolutionary Ecology Research 9:843–854.

Becker, Matthew. 2017. Personal communication April 18, 2017, between M. Becker, Liberty University, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Berdanier, Aaron B. and James S. Clark. 2016. Multiyear drought-induced morbidity preceding tree death in southeastern U.S. forests. Ecological Applications 26(1):17–23.

Bradshaw, Casey Renee. 2010. Effect of snake populations on salamanders as a result of forest fragmentation. Thesis Dissertation, Marshall University, Huntington, WV.

Brand, Adrianne. 2019. Personnel communication February 14, 2019, between A. Brand USGS Patuxent Wildlife Research Center, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Brown, Patrick T., and Ken Caldeira. 2017. Greater future global warming inferred from Earth’s recent energy budget. Nature 552:45–62.

Bury, Bruce, C., Kenneth Dodd Jr., and Gary M. Fellers. 1980. Conservation of the amphibia of the United States: a review. U.S. Fish and Wildlife Service Resource Publication. Washington, D.C.

Carter, L. M., J. W. Jones, L. Berry, V. Burkett, J. F. Murley, J. Obeysekera, P. J. Schramm, and D. Wear. 2014. Chapter 17: Southeast and the Caribbean. In Climate Change Impacts in the United States: The Third National Climate Assessment, J. M. Melillo, Terese (T.C.) Richmond, and G. W. Yohe, Eds., U.S. Global Change Research Program, Washington, DC, USA.

Carter, L. M., A. Terando, K. Dow, K. Hiers, K.E. Kunkel, A. Lascurain, D. Marcy, M. Osland, and P. Schramm. 2018. Chapter 19: Southeast. In Impacts, Risks, and Adaptation in the United States: The Fourth National Climate Assessment, Volume II, D.R. Reidmiller, C.W. Avery, D.R. Easterling, K.E. Kunkel, K.L.M. Lewis, T.K. Maycock, and B.C. Stewart, Eds., U.S. Global Change Research Program, Washington, DC, USA.

Center for Biological Diversity, The Alabama Rivers Alliance, The Clinch Coalition, Dogwood Alliance, The Gulf Restoration Network, Tennessee Forests Council, and The West Virginia Highlands Conservancy. 2012. Petition to List 404 Aquatic, Riparian and Wetland Species from the Southeastern United States as Threatened or Endangered Under the Endangered Species Act. April 20, 2010. 1145 pp.

62

Chazal, Anne C. 2005. Results of Surveys for the Peaks of Otter Salamander (Plethodon hubrichti) in 2005. Natural Heritage Technical Report 06-05. Virginia Department of Conservation and Recreation, Division of Natural Heritage. Richmond, VA 13 pp.

Chen, Jiquan, Sari C. Saunders, Thomas R. Crow, Robert J. Naiman, Kimberley D. Brosofske, Glenn D. Mroz, Brian L. Brookshire, and Jerry F. Franklin. 1999. Microclimate in forest ecosystem and landscape ecology: variations in local climate can be used to monitor and compare the effects of different management regimes. Bioscience 49(4):288–297.

Cherry, Robert. Personal communication November 1, 2017, between R. Cherry, National Park Service Blue Ridge Parkway, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Cherry, Robert. Personal communication August 1, 2018, between R. Cherry, National Park Service Blue Ridge Parkway, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Collins, M., R. Knutti, J. Arblaster, J.-L. Dufresne, T. Fichefet, P. Friedlingstein, X. Gao, W.J. Gutowski, T. Johns, G. Krinner, M. Shongwe, C. Tebaldi, A.J. Weaver, and M. Wehner, 2013. Long-term climate change: Projections, commitments and irreversibility. In Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. T.F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Doschung, A. Nauels, Y. Xia, V. Bex, and P.M. Midgley, Edition. Cambridge University Press. 1029–1136.

Croy, Carol. 2018. Personal communication on August 28, 2018, between C. Croy, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Dallalio, Eric E., Adrianne B. Brand, and Evan H. Campbell Grant. 2017. Climate-Mediated Competition in a High-Elevation Salamander Community. Journal of Herpetology 51(2):190–196

Dodd, Kenneth C. Jr., Judith A. Johnson and Edmund D. Brodie Jr. 1974. Noxious skin secretions of an eastern small Plethodon, P. netting hubrichti. Journal of Herpetology 8(1):89–92.

Dodd, Kenneth C. Jr., and Donald W. Linzey (Ed.). 1979. Peaks of Otter salamander. Proceeding of the Symposium on Endangered and Threatened Plants and Animals of Virginia. Virginia Polytechnic Institute and State University, Blacksburg, Virginia.

Feder, Martin E. 1983. Integrating the ecology and physiology of Plethodontid salamanders. Herpetologica 39(3):291–310.

Fridley, Jason D. 2009. Downscaling climate over complex terrain: high finescale (<1000 m) spatial variation of near-ground temperatures in a montane forested landscape (Great Smokey Mountains). American Meteorological Society 48:1033–1049.

63

Gifford, Matthew E., and Kenneth H. Kozak. 2012. Islands in the sky or squeezed at the top? Ecological causes of elevational range limits in montane salamanders. Ecography 35:193– 203.

Goff, Cory Benjamin. 2015. Effects of Roads and Trails on Peaks of Otter Salamander (Plethodon hubrichti) and Eastern Red-backed Salamander (Plethodon cinereus) Movement Behavior. A thesis submitted to the Graduate College of Marshall University in partial fulfillment of the requirements for the degree of Master of Science in Biological Sciences: Organismal, Evolutionary and Ecological Biology.

Grant, Evan H. Campbell, Adrianne B. Brand, Stephan F.J. Wekker, Temple R. Lee and John E.B. Wofford. 2018. Evidence that climate sets the lower elevation range limit in a high- elevation endemic salamander. Ecology and Evolution 8:7553-7562.

Hairston, Nelson G. 1983. Growth, survival and reproduction of Plethodon jordani: trade-offs between selective pressures. Copeia 4:1024–1035.

Hayhoe, K., J. Edmonds, R.E. Kopp, A.N. LeGrande, B.M. Sanderson, M.F. Wehner, and D.J. Wuebbles, 2017: Climate models, scenarios, and projections. In Climate Science Special Report: Fourth National Climate Assessment, Volume I. D.J. Wuebbles, D.W. Fahey, K.A. Hibbard, D.J. Dokken, B.C. Stewart, and T.K. Maycock, Eds. U.S. Global Change Research Program, pp. 133-160, doi:10.7930/J0WH2N54.

Highton, Richard. 1962. Revision of North American salamanders of the genus Plethodon. Bulletin Florida State Museum 6:235–367.

Highton, Richard. 1986. Plethodon hubrichti. Catalogue of American Amphibians and Reptiles: 1– 2.

Highton, Richard. 1999. Geographic protein variation and speciation in the salamanders of the Plethodon cinereus group with descriptions of two new species. Herpetologica 55(1):43–90.

Highton, Richard. 2005. Declines of Eastern North American woodland salamanders (Plethodon). In: Lannoo M (ed.) Amphibian declines: the conservation status of United States species. University of California Press, Berkeley, CA 34–46.

Highton, Richard, and Allan Larson. 1979. The genetic relationship of the salamanders of the genus Plethodon. Systematic Zoology 28(4):579–599.

Homyack, Jessica A. and Carola A. Haas. 2013. Effects of repeated-stand entries on terrestrial salamanders and their habitat. Southeastern Naturalist 12(2):353–366.

Huber, Fred. 2019. Personal communication on January 29, 2019, between F. Huber, USFS (retired), and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

64

Jackson, R. B., C. Le Quéré, R. M. Andrew, J. G. Canadell, G. P. Peters, J. Roy, and L. Wu. 2017 Warning signs for stabilizing global CO2 emissions. Environmental Research Letters 12:1– 4.

Jaeger, Robert G. 1972. Plant climbing by salamanders: periodic availability of plant-dwelling prey. Copeia 4:686–691.

Jaeger, Robert G. 1978. Competitive exclusion: comments on survival and extinction of species. Bioscience 24(1)33–39.

Jaeger, Robert G., D. Kalvarsky, and N. Shimizu. 1982. Territorial behavior of the red-backed salamander: expulsion of intruders. Animal Behavior 30:490–496.

Jochimsen, Denim M., Charles R. Peterson, Kimberley M. Andrews, and J. Whitfield Gibbons. 2004. A literature review of the effects of roads on amphibians and reptiles and the measures used to minimize those effects. Final Report. Idaho Fish and Game Department and USDA Forest Service.

Kane, A., T.C. Burkett, S. Kloper, and J. Sewall. 2013. Virginia’s Climate Modeling and Species Vulnerability Assessment: How Climate Data Can Inform Management and Conservation. National Wildlife Federation, Reston, Virginia.

Keen, W. Hubert. 1979. Feeding and activity patterns in the salamander Desmognathus ochrophaeus (Amphibia, Urodela, Plethodontidae). Journal of Herpetology 13:461–467.

Kirk, Dawn. 2018a. Personal communication on August 28, 2018, between D. Kirk, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Kirk, Dawn. 2018b. Personal communication on August 29, 2018, between D. Kirk, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Kirk, Dawn. 2018c. Personal communication on October 26, 2018, between D. Kirk, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Kirk, Dawn. 2018d. Personal communication on December 20, 2018, between D. Kirk, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Kleopfer, John D. 2018. Personal communication November 14, 2018, between J. Kleopfer, Virginia Department of Game and Inland Fisheries, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Kniowski, Andrew. 2006. The ecology of the Peaks of Otter salamander (Plethodon hubrichti) in the contact zone with the red-backed salamander (P. cinereus). Thesis. Liberty University, Lynchburg, Virginia.

65

Kniowski, Andrew and Norman Reichenbach. 2006. Plethodon hubrichti (Peaks of Otter Salamander) reproduction. Herpetological Review 37:332.

Kniowski, Andrew and Norman Reichenbach. 2009. The ecology of the peaks of Otter salamander (Plethodon hubrichti) in sympatry with the eastern red-backed salamander (Plethodon cinereus). Herpetological Conservation and Biology 4(3):285–294.

Kramer, Peter, Norman Reichenbach, Michael Hayslett, and Paul Sattler. 1993. Population dynamics and conservation of the Peaks of Otter salamander, Plethodon hubrichti. Journal of Herpetology 27(4):431–435.

Leclair, Maria Helena, Marc Levasseur, Raymond Leclair Jr. 2006. The life-history traits of Plethodon cinereus in the northern parts of its range: variations in population structure, age and growth. Herpetologica 62(3):265–282.

Marsh, David M., Robert B. Page, Teresa J. Hanlon, Rachael Corriton, Elizabeth C. Little, David E. Seifert and Paul R. Cabe. 2007. Effects of roads on patterns of genetic differentiation in red- backed salamanders, Plethodon cinereus. Conservation Genetics 9:603–613.

Marsh, David M., R.B. Page, T.J. Hanlon, H. Bareke, R. Corritone, N. Jetter, N.G. Beckman, K. Gardner, D.E. Seifert, and P.R. Cabe. 2008. Ecological and genetic evidence the low-order streams inhibit dispersal by red-backed salamanders (Plethodon cinereus). Canadian Journal of Zoology 85:319–327.

Marsh, David. 2014. Report on Peaks of Otter salamander (P. hubrichti) monitoring in Parker’s Gap timber units. Unpublished report submitted to the George Washington and Jefferson National Forests February 9, 2014. Washington and Lee University. Lexington, VA.

Marsh, David. 2018a. Personal communication on August 15, 2018, between D. Marsh, Washington and Lee University, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Marsh, David. 2018b. Personal communication on December 19, 2018, between D. Marsh, Washington and Lee University, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Martel, A., M. Blooi, C. Adriaensen, P. Van Rooij, W. Beukema, M. C. Fisher, R. A. Farrer, B. R. Schmidt, U. Tobler, K. Goka, K. R. Lips, C. Muletz, K. R. Zamudio, J. Bosch, S. Lötters, E. Wombwell, T. W. J. Garner, A. A. Cunningham, A. Spitzen-van der Sluijs, S. Salvidio, R. Ducatelle, K. Nishikawa, T. T. Nguyen, J. E. Kolby, I. Van Bocxlaer, F. Bossuyt, F. Pasmans. 2014. Recent introduction of a Chytrid Fungus Endangers Western Palearctic Salamanders. Science 346(6209):630–631.

Martof, Bernard S., Joseph R. Bailey and William M. Palmer. 1980. Amphibians and reptiles of the Carolinas and Virginia. The University of North Carolina Press, Chapel Hill, NC.

66

Mathis, A. 1991. Territories of male and female terrestrial salamanders: costs, benefits, and intersexual spatial associations. Oecologia 86:433–440.

Mauritsen, Thorsten and Robert Pincus. 2017. Committed warming inferred from observations. Nature Climate Change 7:652–655.

Melillo, Jerry M., Terese (T.C.) Richmond, and Gary W. Yohe. 2014. Climate Change Impacts in the United States: The Third National Climate Assessment. U.S. Global Change Research Program. 841.

Messenger, Kevin R., Nathan A. Shepard, and Michael A. Yirka. 2011. Plethodon nettingi (Cheat Mountain Salamander) predation. Herpetological Review 42(4):582–583.

Milanovich, Joseph R., William E. Peterman, Nathan P. Nibbelink, and John C. Maerz. 2010. Projected loss of salamander diversity hotspot as a consequence of projected global climate change. PLOS ONE 5(8)1–10.

Mitchell, Joseph C., Jill A. Wicknick, and Carl D. Anthony. 1996. Effects of timber harvesting practices on Peaks of Otter salamander (Plethodon hubrichti) populations. Amphibian and Reptile Conservation 1(1):15–19.

Muletz, Carly, Nicholas M. Caruso, Robert C. Fleicher, Roy W. McDiarmid, and Karen R. Lips. 2014. Unexpected rarity of the pathogen Batrachochytrium dendrobatidis in Appalachian Plethodon salamanders: 1957–2011. PLOS ONE 9(8):1–7

National Park Service. 2018. Blue Ridge Parkway Firewood Policy FAQ. Accessed on August 6, 2018. https://www.nps.gov/blri/planyourvisit/firewood-faqs.htm.

O’Donnell, Katherine M., Frank R. Thompson III, and Raymond D. Semlitsch. 2015. Prescribed fire and timber harvest effects on terrestrial salamander abundance, detectability, and microhabitat use. The Journal of Wildlife Management 79(5):766–775.

Page, Robert, Clair Connarroe, Diana Quintanilla, Andriea Palomo, Joshua Solis, and David Marsh. 2018. Hybridization and genetic variation in the range-restricted Peaks of Otter salamander, Plethodon hubrichti. Accessed on August 15, 2018. https://app.oxfordabstracts.com/events/264/submissions/42382/abstract-book-view

Pague, Chris A. 1989. King of the Mountain. Virginia Wildlife 50:8–13.

Pague, Chris and Joseph C. Mitchell. 1987. The Status of Amphibians in Virginia. Virginia Journal of Science. 38(4):304–318.

Pague, Chris and Joseph C. Mitchell. 1990. The distribution of the Peaks of Otter salamander (Plethodon hubrichti). A report to the Jefferson National Forest, Roanoke, VA.

67

Pague, Chris and Joseph C. Mitchell. 1991. Peaks of Otter salamander. In Terwilliger, Karen. 1991. Virginia’s Endangered Species. Proceedings of a Symposium. Department of Game and Inland Fisheries. McDonald and Woodward Publishing Company. Blacksburg, VA.

Pague, Chris, Michael Hayslett, and Paul Kramer. 1992. Field notes: Plethodon hubrichti (Peaks of Otter salamander) VA; Rockbridge County, Blue Ridge Parkway at Thunder Ridge overlook, 9.1 km S of Natural Bridge Station. 9 April 1991. Catesbiana 12(1):9.

Petranka, James W. 1998. Salamanders of the United States and Canada. Smithsonian Institute Press, Washington, D.C.

Pough, F. Harvey, Ellen M. Smith, Donald H. Rhodes, and Andres Collazo. 1987. The abundance of salamanders in forest stands with different histories of disturbance. Forest Ecology Management. 20:1–9.

Reardon, Richard C. 1995. Effects of diflubenzuron on non-target organisms in broadleaf forested watersheds in the northeast. National Center of Forest Health Management, USDA, Forest Service Handbook.

Reichenbach, Norman and Tim R. Brophy. 2017. Natural history of the Peaks of Otter salamander (Plethodon hubrichti) along an elevational gradient. The Herpetological Bulletin 141:7–15.

Reichenbach, Norman and Paul Sattler. 2000. Response of Peaks of Otter salamander populations to tree ice damage. Unpublished report submitted to the George Washington and Jefferson National Forests. Roanoke, VA.

Reichenbach, Norman and Paul Sattler. 2005. Effects of timbering on Plethodon hubrichti Final Report (2005). Unpublished report submitted to the George Washington and Jefferson National Forests. Roanoke, VA.

Reichenbach, Norman and Paul Sattler. 2007. Effects of timbering on Plethodon hubrichti over twelve years. Journal of Herpetology 41(4):622–629.

Richardson, Andrew D., Ellen G. Denny, Thomas G. Siccama, and Xuhui Lee. 2003. Evidence for a rising cloud ceiling in eastern North America. Journal of Climate 16:2093–2098.

Roble, Steven M. 2016. Natural Heritage Resources of Virginia: Rare Animals. Natural Heritage Technical Report 16-07. Virginia Department of Conservation and Recreation, Division of Natural Heritage, Richmond, Virginia.

Runkel, J., K. Kunkel, L. Stevens, S. Champion, B. Stewart, R. Frankson, and W. Sweet, 2017: Virginia State Climate Summary. NOAA Technical Report NESDIS 149-VA, p. 4.

Sattler, Paul. 1998. Peaks of Otter salamander study: Movements of Plethodon hubrichti in response to timber harvesting. Unpublished report submitted to the George Washington and Jefferson National Forests. Roanoke, VA.

68

Sattler, Paul and Norman Reichenbach. 1998. The effects of timbering on Plethodon hubrichti: short term effects. Journal of Herpetology 32:399–404.

Seidl, Rupert, Dominik Thom, Markus Kautz, Dario Martin-Benito, Mikko Peltoniemi, Giorgio Vacchiano, Jan Wild, Davide Ascoli, Michal Petr, Juha Honkaniemi, Manfred J. Lexer, Volodymyr Trotsiuk, Paola Mairota, Miroslav Svoboda, Marek Fabrika, Thomas A. Nagel and Christopher P.O. Reyer. 2017. Forest disturbance under climate change. Nature Climate Change 7: 395–402.

Simberloff, Daniel. 1982. The status of competition theory in ecology. Annals of Zoology 19:241– 253

Simon, Steven A. 2013. Ecological zones on the Jefferson National Forest study area: first approximation. Unpublished report submitted to the Jefferson National Forest. Roanoke, VA.

Sites, Jack W. Jr., Mariana Morando, Richard Highton, Fred Huber, and Robin E. Jung. 2004. Phylogenteic Relationships of the Endangered Shenandoah Salamander (Plethodon shenandoah) and Other Salamanders of the Plethodon cinereus Group (Caudata: Plethodontidae). Journal of Herpetology 38(1):96–105.

Smith David R., Nathan L. Allan, Conor P. McGowan, Jennifer A. Szymanski, Susan R. Oetker, and Heather M. Bell. 2018. Development of a species status assessment process for decisions under the U.S. Endangered Species Act, Journal of Fish and Wildlife Management 9(1):302–320; e1944-687X. doi:10.3996/052017-JFWM-041

Snider, Andrew T. and J. Kevin Bowler. 1992. Longevity of reptiles and amphibians in North American collections. Herpetological Circular 21:1–39.

Spotila, James R. 1972. Role of temperature and water in the ecology of lungless salamanders. Ecological Monographs 42(1):95–125.

Strazanac, John S. and Linda Butler. 2005. Long term evaluation of the effects of Bacillus thuringiensis kurstaki, gypsy moth nucleopolyhedrosis virus product Gypchek, and Entomophaga maimaiga on nontarget organisms in mixed broad-leaf forests in the central Appalachians. Division of Plant and Soil Sciences, West Virginia University, Morgantown, WV.

Sutton, William B., Kyle Barrett, Allison T. Moody, Cynthia S. Loftin, Phillip G. deMaynadier and Priya Nanjappa. 2015. Predicted changes in climate niche and climate refugia of conservation priority salamander species in the northeastern United States. Forests 6:1-26.

Takahashi, Mizuki K. and Thomas K. Pauley. 2010. Resource allocation and life history traits of Plethodon cinereus at different elevations. The American Midland Naturalist 163(1):87–94.

69

Taub, Frieda B. 1961. The distribution of the red-back salamander, Plethodon c. cinereus, in the soil. Ecology 24:681–198.

Teague, Bambi. Personal communication October 31, 2018, between B. Teague, National Park Service Blue Ridge Parkway, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Terwilliger, Karen. 1991. Virginia’s Endangered Species. Proceedings of a Symposium. Department of Game and Inland Fisheries. McDonald and Woodward Publishing Company. Blacksburg, VA.

Thurow, Gordon R. 1957. A new Plethodon from Virginia. Herpetologica 13(1):59–66

Tilghman, Joseph M., Shane W. Ramee, and David M. Marsh. 2012. Meta-analysis of the Effects of Canopy Removal on Terrestrial Salamander Populations of North America. Biological Conservation 152:1–9.

U.S. Energy Information Agency. 2017. Annual Energy Outlook 2017 with projections to 2050. Available at https://www.eia.gov/outlooks/aeo/pdf/0383(2017).pdf

U.S. Fish and Wildlife Service. 1997. Habitat Conservation Assessment for the Peaks of Otter Salamander (Plethodon hubrichti). U.S. Fish and Wildlife Service, Unpublished report.

U.S. Forest Service. 2004. Jefferson National Forest Management Plan. Forest Service Southern Region. Roanoke, Virginia.

U.S. Forest Service. 2005. George Washington and Jefferson National Forests Detailed Monitoring and Evaluation Report for Fiscal Year 2004. Resource Management Plan. Forest Service Southern Region. Roanoke, Virginia.

Virginia Department of Game and Inland Fisheries. 2015. State Wildlife Action Plan. Available at http://bewildvirginia.org/wildlife-action-plan/pdf/2015-Virginia-Wildlife-Action-Plan.pdf

Virginia Department of Game and Inland Fisheries. 2018. Virginia Fish and Wildlife Information Service. Richmond, VA. Available from: http://vafwis.org/fwis.

Welsh, Hartwell H., and Sam Droege. 2001. A case for using Plethodontid salamanders for monitoring biodiversity and ecosystem integrity of North American forests. Conservation Biology 15(3):558–569.

Wicknick, Jill. 1995. Interspecific competition and territoriality between a widespread species of salamander and a species with a limited range. Ph.D. dissertation. University of Southwestern Louisiana, Lafayette, Louisiana.

70

Wiens, John J., Tag N. Engstrom and Paul T. Chippindale. 2016. Rapid diversification, incomplete isolation, and the “speciation clock” in the North American salamanders (genus Plethodon): testing the hybrid swarm hypothesis of rapid radiation. Evolution 60(12):2585–2603.

Wright, Daniel. 2018a. Personal communication on October 11, 2018, between D. Wright, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Wright, Daniel. 2018b. Personal communication on October 26, 2018, between D. Wright, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Wright, Daniel. 2019. Personal communication on February 27, 2019, between D. Wright, U.S. Forest Service, and Rose Agbalog, U.S. Fish and Wildlife Service, Abingdon, Virginia.

Wuebbles, D.J., D.R. Easterling, K. Hayhoe, T. Knutson, R.E. Kopp, J.P. Kossin, K.E. Kunkel, A.N. LeGrande, C. Mears, W.V. Sweet, P.C. Taylor, R.S. Vose, and M.F. Wehner, 2017: Our globally changing climate. In Climate Science Special Report: Fourth National Climate Assessment, Volume I. D.J. Wuebbles, D.W. Fahey, K.A. Hibbard, D.J. Dokken, B.C. Stewart, and T.K. Maycock, Eds. U.S. Global Change Research Program, entire.

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APPENDIX A Current Condition Methodology A1. Survey Data Studies to document presence/absence and to quantify numbers or densities of Peaks of Otter salamander have occurred for decades since the species was first recognized by Thurow. Chazal (2005, p. 1) reviewed the available spatial data and performed surveys at the extreme northwest and southwest portions of Peaks of Otter salamander’s range in an attempt to map the microdistribution of the species. As of 2018, the outer range limits of the species were fairly well defined, but the patchiness of the species’ distribution within the range had not been determined.

Table 14. Species observation data used for analysis. Data name Owner Type Date Range Number of Records*

Masterpoly2005 USFS polygon 1962-2001 101

Masterpoints2005 USFS point 1993-2005 501

Masterlines2005 USFS line 1994-1996 23

DGIF VAFWIS VADGIF point 1987-2014 247

POS_16May2014 USFS point 2014 3

POS-RB points Marsh Marsh point 2016-2017 21

*Some records contain more than one observation. Several species observations associated with research projects have been recorded and documented. Many of these projects were focused on determining the impacts of timber harvest activities on salamanders, so observations are typically focused in areas where timbering activities (see Chapter 3) had or would potentially occur. Additional surveys to determine the impacts of roads, competition with red-backed salamanders, and how specific environmental factors relate to distribution patterns, have also contributed to species observation data. Most of these data were given to the U.S. Forest Service (USFS) to compile and manage and were separated by vector data type (point, line, polygon). This dataset is called Master (point, line or polygon) 2005. Some observation data were submitted to the Virginia Department of Game and Inland Fisheries and incorporated into their Virginia Fish and Wildlife Information Service (VAFWIS) database. Dates associated with each dataset varied widely. A small number of the occurrence data had a null value for the date. Other dates ranged from 1962 to 2017. There were a large number of observations from 2005 during the time that Chazel was doing her work, and a large number of

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polygon shapefiles were from surveys from 1989 to1990 done by Pague and Mitchell to define the range and distribution of Peaks of Otter salamander. Some surveys indicated the number of Peaks of Otter salamander while others only indicated presence. Many of the polygons from the Pague and Mitchell data did not indicate exact numbers of Peaks of Otter salamander, but gave an estimate of salamanders present per square meter (m2). Some survey data also indicated the number or presence/absence of red-backed salamanders. Sources of data for red-backed salamander occurrence included the Masterpoints2005, Masterpoly2005 and the DGIF VAFWIS database. These data are displayed in Figure 17. A2. Demographic Parameters Number of Salamanders To estimate the number of individuals present within each analysis unit, available species observation data in both GIS point and polygon shapefiles were used. For the polygon data, if the attributes specified the survey area and the number of Peaks of Otter salamander detected, then those values were used to determine how many Peaks of Otter salamander were present per m2 and those values were included in a new column. If the attributes indicated a number of Peaks of Otter salamander per m2, those values were added to the new column. If a survey area was not defined, and a number of Peaks of Otter salamander was indicated, then the calculate geometry function in ArcMap was used to find the area of the polygon and the number of Peaks of Otter salamander was divided by the area to get the number of Peaks of Otter salamander per m2. If only presence was indicated, then it was assumed that there was 1 location of 1 Peaks of Otter salamander per 1 m2. In the Masterpoint2005 data, an individual point was taken for each Peaks of Otter salamander found. If more than one salamander was found at a location that was reflected in the attribute data. Because there was no information given on the search radius used for each point location and the accuracy was less than 20 m, the number of salamanders was calculated as n/1 m2. In the DGIF VAFWIS data, an individual point was taken for each Peaks of Otter salamander found and there was no information indicating that there were more than one salamander found at each location. Because there was no information given on the search radius used for each point location or the accuracy, the number of salamanders was assumed to be 1/1 m2. For the POS_16_May2014 data there were only three records, one was of red-backed salamander and two were Peaks of Otter salamander. All of these records fell within the White Oak Ridge AU. One record indicated one Peaks of Otter salamander and the other indicated two Peaks of Otter salamanders. Because there was no information given on the search radius used for each point location or the accuracy, the number of salamanders was assumed to be the number of Peaks of Otter salamander/1 m2. The POS-RB points Marsh data indicated presence of both Peaks of Otter salamander and red-backed salamander but did not indicate numbers of salamanders. Because there was no information given on the search radius used for each point location or the accuracy, the number of salamanders was assumed to be 1/1 m2 for each record with Peaks of Otter salamander. Point data from Jill Wicknick’s (1995) thesis work was available but because of the nature of the data, they were not used to estimate the number of individuals. The study aimed to identify effects of timber harvest on Peaks of Otter salamander. The same transects were surveyed in 1994, 1995, and 1996, with 1994 preharvest data and 1996 postharvest data. In many of the locations, the number of salamanders decreased over the 2 years. Because of what the data were used for, and because those occurrences largely overlapped with existing data or more current information, we

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omitted it from the estimation; however, it was included in the range maps. Because these data were originally line shapefiles, to display the data, the centerpoint of each line was used as the location point. To determine the density of salamanders in each AU, all of the densities of the points and polygons within each unit were added together, then that number was divided by the size of the AU in meters. Values between greater than 2.01/m2 were considered “high,” values between 1.01/m2 and 2/m2 were considered “medium/high,” values between 0.06/m2 and 1/m2 were considered “medium” and values between presence and 0.05/m2 were considered “low.” To comply with terms stated in the HCA and protect the locations of the salamander, a map with location data are not included in this document. Reproductive Output Although Peaks of Otter salamander’s range is relatively small, research indicates there are certain behavioral differences between salamanders at different elevations throughout the range. According to Reichenbach and Brophy (2017, p. 13) “at lower elevation sites, salamanders might rarely have environmental conditions that allow them to forage on vegetation and, consequently, they forage more frequently in forest litter or under rocks and logs where foraging efficiency is reduced. Eventually, at certain low elevations, varying based on local conditions such as site aspect and proximity to streams, environmental conditions at the surface might force Peaks of Otter salamander to remain underground most of the time, thus creating an unsustainable energy budget (Spotila, 1972; Gifford and Kozak, 2012). This physiological stress, from reduced foraging efficiency with decline in elevation, may have translated into reduced reproductive output with decline in elevation.” Salamander density also decreased primarily with decline in elevation, and was likely related to habitat (poor habitat quality), temperature (thermal stress from high temperatures), relative humidity (RH) (water stress from dehydration when RH is too low), and potentially differing predator and prey communities with change in elevation. Along the elevational gradient in the Peaks of Otter area, with decline in elevation, it got warmer (1 °C with every decline in elevation of 204 m (669 ft)) and drier (1 percent decrease in RH with a decline in elevation of 120 m (394 ft)) (Reichenbach and Brophy 2017, p. 13). Amphibians living at high elevations “tend to produce fewer clutches per year, larger absolute clutches and larger eggs relative to those living at lower elevations” (Morrison and Hero, 2003 In Reichenbach and Brophy 2017, p. 13). For Peaks of Otter salamander, this physiological stress from reduced foraging efficiency with decline in elevation, may have translated into reduced reproductive output with decline in elevation (Reichenbach and Brophy 2017, p. 13). Results from the study suggested that salamanders at higher elevation sites, up to the optimum elevation, produced more eggs per female. The number of eggs per female Peaks of Otter salamander increased directly with mass and elevation to a maximum of 12 eggs per female at 1000 m in elevation and then decreased slightly above 1000 m in elevation. The number of eggs per female ranged from 1 to 12 with a mean of 8.5 (Reichenbach and Brophy 2017, p. 11). However, with increase in elevation, percent gravid females decreased. At the higher elevation sites in this study, reproduction was primarily biennial as suggested by percent gravid females ranging from 40 to 60 percent. At the lower elevation sites, in contrast, at least some of the salamanders produced eggs annually since percent gravid females ranged from 67 to 80 percent (Takahashi and Pauley 2010 In

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Reichenbach and Brophy 2017, p. 13). The overall effect with decline in elevation, even with more frequent reproduction, was a decrease in reproductive output (Reichenbach and Brophy 2017, p. 13) (figure 13).

Figure 13. Graph of relative egg output per 1000 m2 (combining multiple regression models for number of eggs/female, percent gravid, and SA density) taken from Reichenbach and Brophy (2017, p. 12). Based on information from Reichenbach and Brophy (2017, p. 13), elevations between 800 m and 1000 m (2,625-3,281 ft) were considered “high” condition, elevations between 1000 m and 1200+m (3,282-3,937+ft) were considered “medium/high” condition, elevations between 600 m and 800 m (1,969-2,625 ft) were considered “medium” condition and elevations between 300 m and 600 m (984-1,969 ft) were considered “low” condition. Reproductive output for current condition is scored based solely on elevation because we do not have available information regarding climate conditions at the time surveys were conducted. Therefore, we are using the elevation as a proxy for assumed climatic conditions. However, for future conditions, both elevation (which does not vary over time) and climate conditions will be factored into the condition score for this metric. A3. Habitat Parameters Habitat Quality Habitat quality reflects conditions necessary for Peaks of Otter salamander to reproduce, forage, and shelter. We considered forest type and percent canopy closure as surrogate measures for habitat quality. We used Ecological Zone data developed for the Jefferson National Forest to determine ecological zone type, and canopy cover from the National Land Cover Dataset (NLCD 2011) to determine amount of closed canopy within each unit. Ecological Zone data were generated using landform and environmental variables, and the relationship between Ecological Zone and environments were analyzed using MAXENT 3.2.1 (Simon 2013, pp. 6–7).

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Figure 14. Environmental variables evaluated in Ecological Zone models, taken from Simon 2013 (p. 7). Habitat quality is presumed to be “high” condition in the AU if there is high canopy cover and intact, productive hardwood forests. We assumed that if forest type and canopy cover are high quality, then there will be adequate surface cover, litter and underground refugia; moist, well- developed soils; abundant, high quality prey; suitable soil and air temperature; and suitable relative humidity. If canopy cover is low and forest type is poor quality, we assumed that there would not be adequate surface cover, litter and underground refugia; soils would be dry and compacted; prey

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would be limited or poor quality; soil, ambient temperatures and relative humidity would not be suitable for the Peaks of Otter salamander to breed, feed and shelter. AUs with poor habitat quality were given a “low” condition.

Figure 15. Map of AUs and Ecological Zones.

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Figure 16. Map of AUs and Canopy Cover.

Sympatry Kniowski and Reichenbach (2009, p. 285) attempted to determine the extent and effect of sympatry of Peaks of Otter salamander with red-backed salamander. They found that density for Peaks of Otter salamander in sympatry with red-backed salamander was lower than previously recorded for Peaks of Otter salamander in allopatry, suggesting that the presence of red-backed salamander affects resource availability or results in fewer Peaks of Otter salamander in an area than would otherwise occur in allopatry. Data from the Masterpoints2005, Masterpoly2005 and the DGIF VAFWIS database were used to determine the presence of red-backed salamanders in each AU. If there has been no documentation of sympatry within the AU, the unit was considered in high condition. If Peaks of Otter salamander is sympatric with red-backed salamander within the AU and there were many observations of red-backed salamander within or near the unit boundary, it was considered “low” condition.

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Figure 17. Map of AUs and red-backed salamander occurrences.

Habitat Management Within the NPS’ Blue Ridge Parkway, although there are no specific guidance documents for management considerations in areas where Peaks of Otter salamander is known to occur, maintenance work avoids Peaks of Otter salamander habitat or is performed during times of the year when the salamanders are inactive on the surface (Cherry 2017; Cherry 2018). Most of the work conducted by the NPS in the Peaks of Otter area is focused on maintaining existing facilities (i.e., campground, visitor center and lodge) which are in already developed areas of the Blue Ridge Parkway. According to the Jefferson National Forest Management Plan, within the primary Peaks of Otter salamander habitat area, no timber harvest is permitted (USFS 2004, pp. 129, 131). Within the secondary habitat area, shelterwood cuts, or thinning is permitted and no clear cutting is allowed (USFS 2004, pp. 129, 131). Standards outlined in the Timber Management section of the Jefferson National Forest Management Plan allow thinning to take place but work to ensure that the salamander’s habitat will remain intact. Areas without vegetation like roads, trails and utility rights-of-way are minimized, although trail and road reconstruction and new parking facilities are

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permitted (USFS 2004, p. 133). Management within the secondary habitat area may also include prescribed fire and reduction of fuel hazard (USFS 2004, p. 135). In additional to the Peaks of Otter salamander habitat conservation zones, there are many management prescriptions implemented within the species range. These prescriptions allow timber harvest, limit timber harvest, or do not allow timber harvest within the prescription area. When prescriptions overlap, it is the most conservative prescription that takes precedence. For example, a no timber harvest prescription in backcountry recreation takes precedence over a secondary conservation zone that allows timber harvest (Kirk 2018b). Analysis Units that are within a primary Peaks of Otter salamander habitat conservation area, were located within a prescription that did not allow timber harvest, or were on Park Service land were considered “high” condition because they are being managed for optimal habitat conditions for the Peaks of Otter salamander, while AUs that are managed as both primary and secondary habitat conservation areas were considered “medium/high” condition. AUs within secondary habitat conservation areas with limited timbering were considered “medium” condition, while AUs that are within a secondary habitat conservation area with timber harvest allowed, or are outside of USFS or NPS managed land were considered “low” condition because according to Kniowski and Reichenbach, (2009, p. 292) any form of timbering is permitted on private property.

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Figure 18. Map of AUs and habitat management. Overall Condition The different population and habitat parameters for each AU were ranked as high, medium/high, medium or low and these ranking received a score from four to one respectively. No metric was weighted differently than the others. The scores were then added together for each AU and were used to calculate overall condition of high, medium/high, medium or low. The colors in the tables and maps used to indicate conditions in the current and future condition tables were green, yellow, orange and red, respectively.

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APPENDIX B Future Conditions

Figure 19. Spatial depiction of the AUs and Scenario 1 2040 condition scores.

Figure 20. Spatial depiction of the AUs and Scenario 1 2100 condition scores.

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Figure 21. Spatial depiction of the AUs and Scenario 2 2040 condition scores.

Figure 22. Spatial depiction of the AUs and Scenario 2 2100 condition scores.

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Figure 23. Spatial depiction of the AUs and Scenario 3 2400 condition scores.

Figure 24. Spatial depiction of the AUs and Scenario 3 2100 condition scores.

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