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. Fortunately, some populations of Cheat
Mountain Salamanders described in this paper are within the boundaries of
the Monongahela National Forest and are protected from future logging and
developments.
Acknowledgments
I thank the many students who helped me with the field work during the last 30
years. Most were from Salem College and Marshall University. I acknowledge and
appreciate the financial support provided by the United States Forest Service, United
States Fish and Wildlife Service, and West Virginia Division of Natural Resources
that allowed me to conduct many of the original surveys. I thank Edwin Michael,
Ronald Lewis, Craig Stihler, Jessica Wooten, and Jayme Waldron for valuable comments
on the manuscript.
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