2008 NORTHEASTERN NATURALIST 15(4):595–606 The Appalachian Inferno: Historical Causes for the Disjunct Distribution of Plethodon nettingi (Cheat Mountain Salamander) Thomas K. Pauley* Abstract - The original Picea rubens (Red Spruce) forest in West Virginia covered approximately 1.5 million acres, most of which was eliminated between 1870 and 1920 by clear-cutting and conflagrations. The total range of Plethodon nettingi (Cheat Mountain Salamander) was confined within this Red Spruce forest. Fires burned the duff and soil to the bedrock in many places, thus eliminating salamander habitats. It is hypothesized that Cheat Mountain Salamanders were eradicated throughout much of their range, and only areas with large emergent rocks or boulder fields provided refugia where they survived. Introduction Prior to arrival of European lumber crews in the mid- to late 1800s, the high elevations of the Allegheny Mountains in eastern West Virginia were covered with approximately 1.5 million acres of majestic Picea rubens Sarg. (Red Spruce) forest. By 1900, the forest had been reduced to 225,000 acres (West Virginia Conservation Commission 1908). This near elimination of the Red Spruce forest has been attributed to improved technologies for cutting, hauling, and manufacturing timber (Lewis 1998). Stickel (1923) summed up destruction of the original forest by stating that lumbermen followed a policy of “cut clean and then clear out” (from Lewis 1998). Before large-scale lumbering activities began, accidental wildfires and “hacking” (a technique used to remove trees by girdling with a hatchet or an axe) had destroyed the original forest in some locations. A few years later, after the trees had died, farmers would return to the area and burn the dead trees and undergrowth in order to clear the land for grazing. Hacking was practiced prior to the Civil War in the Red Spruce areas of eastern and southern portions of Randolph and northern Pocahontas counties (Clarkson 1964, Hopkins 1908). Possibly the first major destructive fire in the original forest occurred in 1863 when fire escaped from a campfire of Confederate scouts on the Roaring Plains section of Dolly Sods in Randolph County. This fire burned the summit and sides of the Allegheny Mountains from Tucker County through Grant, Pendleton, and Randolph counties to the head waters of the Greenbrier River (West Virginia Conservation Commission 1908). These events were the beginning of what would lead to the destruction of the original *Department of Biological Sciences, Marshall University, Huntington, WV 25755; pauley@marshall.edu. 596 Northeastern Naturalist Vol. 15, No. 4 Red Spruce forest, the habitat of the federally protected Plethodon nettingi (Green) (Cheat Mountain Salamander). I have studied the Cheat Mountain Salamander, an eastern, small woodland salamander, for 30 years and have found about 80 disjunct populations in an area that extends approximately 92 km north to south and 31 km east to west encompassed within five counties in the high Alleghenies of eastern West Virginia (Pauley 2007). Their habitat typically consists of forests dominated by Red Spruce and Betula alleghaniensis Britt. (Yellow Birch), where the forest floor is covered with Bazzania trilobata (L.) S. Gray (Threelobed Bazzania, a liverwort). Such habitats are associated with emergent rocks, boulder fields, or narrow ravines lined with Rhododendron maximum L. (Great Rhododendron) (Pauley 2005). Clearcuts, such as those that occurred in the high elevations of eastern West Virginia in the late 1800s, are detrimental to forest salamander populations (Ash 1988, Mitchell et al. 1996, Petranka et al. 1993, Sattler and Reichenbach 1998). Clear-cutting opens the forest floor to the drying effects of sun and wind resulting in a dry humus or litter layer that provides fuel for wildfires. In the once abundant Red Spruce forests in eastern West Virginia, leaf-litter decay was slow due to the acidic nature of spruce needles, which resulted in the formation of a thick layer of partially decayed needles and twigs several feet deep (Brooks 1911). Most fires that occurred in the clearcut areas of the original forest were started by sparks from Shay locomotives, sawmills, campfires, and the careless disposal of hot ashes from fireboxes of tenders (Fansler 1962, West Virginia Conservation Commission Report 1908). With the highly combustible properties of dry spruce needles, fires started quickly, and the extreme heat likely reached deep into the rock layers beneath the soil where salamanders took refuge. Because forest fires tend to draw enormous amounts of oxygen from both above and below the surface of the ground (Michael 2002), many species of salamanders may have died due to lack of oxygen before the heat reached their refuge. A thick, moist litter layer on the forest floor is essential to maintain healthy woodland salamander populations (Ash 1997). Moist litter is used as the main foraging venue of woodland salamanders (Jaeger 1978, 1980), as refugia for all age classes, and for cutaneous respiration (Ash 1995). Nests of several species of woodland salamanders, including Cheat Mountain Salamanders, are associated with leaf litter where they are found in and under logs, under bark on logs, and under surface rocks (Green and Pauley 1987). Clearcuts and fires can destroy this layer and make such disturbed areas inhospitable for forest salamanders. The purpose of this paper is to suggest a hypothesis for the link between the fragmented distribution of the Cheat Mountain Salamander, a threatened endemic (Federal Register 1989) woodland salamander, and historical disturbance and destruction of the high-elevation forests due to clearcutting 2008 T.K. Pauley 597 and subsequent fires that occurred from 1870 through 1960 in eastern West Virginia. Methods Salamander surveys were conducted during daylight hours from May 1976 to October 2006 and involved turning cover objects such as rocks and logs. In most cases, I only searched within 48-h following a rain event. Data recorded included species, size class (i.e., juvenile, subadult, adult), gender, cover object, and general habitat characteristics including dominant plant species and presence of emergent rocks or boulder fields. I searched in numerous sites in each of four study areas. If multiple sites with Cheat Mountain Salamanders were within the typical home-range size of Plethodon cinereus (Green) (Eastern Red-backed Salamander) (Kleeberger and Werner 1982), a species similar in size to the Cheat Mountain Salamander, I considered these multiple Cheat Mountain Salamander sites to be one population. If sites were farther apart than could be contained in the typical home range of the Eastern Red-backed Salamander, I considered them separate Cheat Mountain Salamander populations. In this paper, I present species richness and abundance in four areas within the geographic range of the Cheat Mountain Salamander that were impacted by clearcuts and fires. Because the Cheat Mountain Salamander is a federally protected species, exact locations of populations are not provided. Results and Discussion During the 30 years of this study, I searched about 1300 sites and examined 22,389 forest-dwelling salamanders throughout the entire range of the Cheat Mountain Salamander (Table 1). Of these, only about 10.0% were Cheat Mountain Salamanders, found at approximately 80 disjunct locations. In 1996, a colleague and I conducted detailed inventories where Table 1. Salamander species and numbers observed from 1976 through 2006 throughout the range of the Cheat Mountain Salamander. Species Number observed Desmognathus ochrophaeus (Allegheny Mountain Dusky Salamander) 7110 Eurycea bislineata (Green) (Northern Two-lined Salamander) 95 Gyrinophilus p. porphyriticus (Green) (Northern Spring Salamander) 41 Hemidactylium scutatum (Temminck and Schlegel in Von Siebold) 77 (Four-toed Salamander) Plethodon cinereus (Eastern Red-backed Salamander) 9880 Plethodon glutinosus (Northern Slimy Salamander) 987 Plethodon nettingi (Cheat Mountain Salamander) 2229 Plethodon wehrlei (Wehrle’s Salamander) 1746 Notophthalmus v. viridescens (Red-spotted Newt) 224 Total 22,389 598 Northeastern Naturalist Vol. 15, No. 4 I had observed one population of Cheat Mountain Salamanders 25 years earlier during a less extensive search. We found that Cheat Mountain Salamanders were actually in 12 disjunct sites that were all on or adjacent to emergent rocks. After reviewing habitat characteristics of the 60 known populations for Cheat Mountain Salamanders at that time (1996), at least 86.7% (n = 52) were associated with emergent rocks or small boulder fields (Fig. 1). Rocks may have been associated with some or all of the remaining eight populations; however, because in the earlier years of my work I did not understand the positive association between rocks and the occurrence of Cheat Mountain Salamanders, I did not record the presence or absence of these structures. From historical accounts, forest stands supporting all but one of the approximately 80 known populations (Gaudineer Scenic Area) have been cut since 1870 and, in most cases, clearcut. Historical records show that many isolated Cheat Mountain Salamander populations associated with emergent rocks and boulder fields today, existed in areas that were clearcut and burned between 1870 and 1960. Here, I describe four areas as examples where Cheat Mountain Salamanders are confined to microhabitats where forests were burned during the initial cutting of the virgin forests. These areas include: Spruce Knob (Pendleton County), Blackwater Canyon (Tucker County), Dolly Sods (Grant and Tucker counties), and Bald Knob (Pocahontas County) (Fig. 2). Spruce Knob (Pendleton County) Spruce Knob (Pendleton County), the highest elevation in West Virginia, is perhaps the epitome of what the devastation must have been like during the logging of the virgin forest and the subsequent burning of the litter and soil. The 1899–1900 biennial report to the State Board of Agriculture stated that clear-cutting methods employed by timbering companies removed trees of all sizes at a rate of 30 to 50 acres per day, which left Spruce Knob a desolate place (West Virginia Conservation Commission 1908). Stickel (1923) acknowledged that after the initial logging in the Spruce Knob area there was not a single tree left standing and the area was one of the best examples of wasteful and destructive logging (From Lewis 1998). A.B. Brooks, the state’s leading conservationist and Director of the West Virginia Geological Survey in the early 1900s, described the duff of the original forest as a collection of spruce needles, leaves, mosses, and lichens that decomposed over centuries to form a layer of soil one to three feet thick. This highly flammable, dry organic matter was exposed to sunlight due to removal of the trees and was subject to ignition by the slightest spark. According to an eyewitness, the most formidable fire in the Spruce Knob area swept the eastern side of the Allegheny Mountains around the headwaters of Big Run in Pendleton County located just west 2008 T.K. Pauley 599 Figure 1. Examples of an emergent rock and boulder field where Plethodon nettingi (Cheat Mountain Salamander) may have survived forest clearcuts and fires and where they frequently occur today. 600 Northeastern Naturalist Vol. 15, No. 4 of Spruce Knob. Flames were reported to exceed the tallest “pines” (Red Spruce) and advanced 10 miles an hour. Brooks (1911) concluded that perhaps 1000 years would not be enough time to replace the humus of the soil that fire had destroyed. After clearcuts and several fires in the Spruce Knob area, F.E. Brooks visited the region and reported that forests were found east of the Allegheny Front at Spruce Knob and into Virginia where fires had not occurred, but to the west, most of the countryside was a wasteland of Pteridium aquilinum (L.) Kuhn (Bracken Fern) that covered the ground from which almost every trace of the original forest had been destroyed by fires (Brooks 1908). The disjunct microdistribution of Cheat Mountain Salamanders in the Spruce Knob area today attests to the near obliteration of the original forest by fires. Nearly 100 years after the fires, I surveyed 58 sites Figure 2. Range of Plethodon nettingi (Cheat Mountain Salamander) with burn sites. 2008 T.K. Pauley 601 in suitable Cheat Mountain Salamander habitat (i.e., areas within a Red Spruce and Yellow Birch forest stand) and found that they occurred in only eight disjunct populations in the Spruce Knob area. These salamanders were discovered in boulder fields on the upper slopes and on a ridge that stretched from the highest elevational point northeast. Many of these sites surveyed over 30 years were outside of the boulder field in the most severely burned area, and no Cheat Mountain Salamanders have been observed in these sites. Other species of salamanders such as Eastern Red-Backed Salamanders, Plethodon wehrlei Fowler and Dunn (Wehrle’s Salamander), P. glutinosus (Green) (Northern Slimy Salamander), Desmognathus ochrophaeus Cope (Allegheny Mountain Dusky Salamander), and Notophthalmus viridescens viridescens (Rafinesque) (Red-Spotted Newt, Red Eft) were found throughout areas in and outside of boulder fields and in areas devastated by fires (Table 1). Blackwater Canyon (Tucker County) One the most detailed chronologies is of a forest fire that occurred in the Blackwater Canyon, 4.8 km north of Hendricks, Tucker County. Fansler (1962), an eyewitness to the fire, described the conditions that led to this destructive fire, “As long as the trees stood, the humus soil remained shaded and damp and would not burn, but when the trees were cut the sun came in and dried the soil to the highly combustible peat. When fires occurred, the peat burned and smoldered for months and only a heavy and protracted snow would extinguish it.” Fansler watched a fire start in the Blackwater Canyon on 30 May 1914. He stated “The sky over Hendricks was lighted with the reflection of the blaze to such an extent that one could sit on the platform of Harvey’s store at midnight and read the afternoon paper without any other illumination.” Snows finally extinguished the fire on 30 November 1914, after burning six months. Prior to the 1914 fire, Fansler reported a fire that burned 7000 acres on the north side of the Canyon in 1910 above the Western Maryland Railway. I searched 27 sites on the north side of the Canyon, which contains habitat characteristic of Cheat Mountain Salamanders. Of these sites, I found Cheat Mountain Salamanders in only three populations: one small locality in a boulder field in the floodplain and two populations among large emergent rocks along the rim of the Canyon. Species found in the burned areas included Eastern Red-backed Salamander, Northern Slimy Salamander, and Allegheny Mountain Dusky Salamander. In many locations on the north side of the Canyon, the soil was thin and dry, and no salamanders were discovered. Dolly Sods (Tucker County and Grant County) Dolly Sods was logged between the 1880s and the 1920s, and numerous fires burned the slash and the centuries of accumulated humus. In July 1930, one fire destroyed 24,000 acres in the middle of the Sods (Turner 602 Northeastern Naturalist Vol. 15, No. 4 2001). Fires smoldered for months and burned the forest and ground cover down to bare rock. Old charred stumps and charcoal fragments can be still be found under the surface in places like Dobbins Slashings, which is located west of Bear Rocks (Allard and Leonard 1952). Today, in many locations, the Red Spruce forest has been replaced by hardwood tree species, such as Acer rubrum L. (Red Maple) and Fagus grandifolia Ehrh. (American Beech). Rocks that were once covered with 0.5 m or more of humus are now exposed to sunlight, making the area uninhabitable for woodland salamanders (Fig. 3). The first recorded fire in the Dolly Sods region was in 1863 when a campfire of Confederate scouts escaped and ignited the forest in the Roaring Plains and Flat Rock Plains areas (West Virginia Conservation Commission 1908). Today, these areas consist of xeric habitat with numerous exposed rocks. In this region of Dolly Sods, I have surveyed 10 sites since 1979. Cheat Mountain Salamanders were found in one locality in a boulder field along South Fork, a habitat that could have provided shelter for them during the 1863 fire. In addition to the surveys in the Roaring Plains and Flat Rock Plains areas, I have conducted salamander inventories in 36 sites throughout the rest of Dolly Sods. Many sites have been searched several times during the years, and I have located Cheat Mountain Salamanders in only four sites between Fisher Spring Run and Roaring Plains. This area consists of boulder fields Figure 3. Rocks exposed after fires destroyed the humus and soil at Dobbins Slashings near the Allegheny Front at Dolly Sods. 2008 T.K. Pauley 603 with ravines and sufficient soil deposits to support woodland salamanders. The most intensely burned area was probably between Fisher Spring Run and Mount Storm Lake. I surveyed 20 sites in this area and only found one Eastern Red-backed Salamander. Bald Knob (Pocahontas County) Bald Knob was clearcut in 1902, leaving dry duff and dead spruce tops and branches that burned in 1904. The area was cut again between 1950 and 1958 (Clarkson 1990), followed by a storm that felled a large number of spruce trees during the winter of 1979–1980 (Clarkson 1990). Brooks (1948) reported nests and adults of Cheat Mountain Salamanders at Bald Knob in 1940. From 1979–1990, I searched 12 sites on and around Bald Knob and found Cheat Mountain Salamanders in only one site associated with emergent rocks on the west slope. The summit had a new growth of Red Spruce with the liverwort Bazzania trilobata, but I did not find Cheat Mountain Salamanders. Of the remaining 11 sites, I found four Eastern Red-backed Salamanders in one site; no salamanders were found in the other 10 sites. Some Cheat Mountain Salamanders may have survived the initial cut and burn on Bald Knob, but succumbed to the subsequent cuts in the 1950s and in 1979–1980. Conclusions Cheat Mountain Salamanders appear to be associated with either emergent rocks or boulder fields. Cool moist spots under rocks serve as refugia where salamanders may survive clearcuts and fires. All Cheat Mountain Salamander populations observed since 1996 have been found in these rocky microhabitats. Cheat Mountain Salamanders survived cutting and burning of the original forest only around large rocks, boulder fields, and narrow ravines protected by thick growths of Great Rhododendron. There are areas with emergent rocks and boulder fields within the range of the Cheat Mountain Salamander where the salamanders do not occur (Pauley 2007). Fires in such areas may have been too severe for Cheat Mountain Salamanders to survive in these relatively safe refugia. Competition for moist spots with Allegheny Mountain Dusky Salamanders and for food and nesting sites with Eastern Red-backed Salamanders currently limits Cheat Mountain Salamanders to rocky microhabitats (Green and Pauley 1987; B.A. Pauley 1998; T.K. Pauley 1980, 2005). Adams et al. (2007) suggested that Eastern Red-backed Salamanders have morphological and behavioral flexibility that may allow them to adapt more quickly to local environmental conditions. This flexibility perhaps gives Eastern Red-backed Salamanders the competitive advantage over Cheat Mountain Salamanders in becoming re-established in a recovering forest. Allegheny Mountain Dusky Salamanders, a species that deposits eggs in stream banks and seeps rather than the terrestrial habitats where 604 Northeastern Naturalist Vol. 15, No. 4 Cheat Mountain Salamanders nests are located (Green and Pauley 1987, Pauley et al. 2006), could survive in streams and seeps during cutting events and fires, thus providing them a survival advantage. The broader ranges of both Eastern Red-backed Salamanders and Allegheny Mountain Dusky Salamanders suggest they are more capable of adapting to environmental extremes than Cheat Mountain Salamanders. Roads, ski slopes, hiking trails, rights-of-way, and developments currently impact all known Cheat Mountain Salamander populations (Pauley 2005), by opening the forest floor and limiting Cheat Mountain Salamanders to small, disjunct populations. 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