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Comparison of Relative Abundance and Microhabitat of

Desmognathus organi (Northern Pygmy Salamander) and
Desmognathus wrighti (Southern Pygmy Salamander) in ....

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2018
2018 Southeastern Naturalist
SOUTHEASTERN NATURALIST Vol. 17, No. 1
17(1):141–154
C.R. Rossell Jr., Ivy C. Haas, L.A. Williams, and S.C. Patch

Comparison of Relative Abundance and Microhabitat of

Desmognathus organi (Northern Pygmy Salamander) and
Desmognathus wrighti (Southern Pygmy Salamander) in
North Carolina
C. Reed Rossell Jr.1,*, Ivy C. Haas1, Lori A. Williams2, and Steven C. Patch3

Abstract - There are currently no quantitative studies describing the habitat of Desmogna-
thus organi (Northern Pygmy Salamander) and D. wrighti (Southern Pygmy Salamander).
We investigated the relative abundance and microhabitat selection of Northern Pygmy
Salamander and Southern Pygmy Salamander in 3 forest types of different elevations
across the mountains of North Carolina during the summer of 2015. We conducted 2-h
time-constrained searches at 4 sites in Picea rubens (Red Spruce)–Abies fraseri (Fraser Fir)
forests, northern hardwood forests, and mountain cove forests for each species. We quanti-
fied microhabitat characteristics at each pygmy salamander location and at a corresponding
random location 2–30 m away. We captured a total of 98 pygmy salamanders (D. organi =
41, D. wrighti = 57): 52 in spruce–fir forests, 26 in northern hardwood forests, and 20 in
cove forests, and recorded 655 other salamanders representing 15 species. Relative abun-
dance of pygmy salamanders was greater in spruce–fir forests than in cove forests, but was
not significantly different between spruce–fir and northern hardwood forests. Microhabitat
did not differ between Northern Pygmy Salamander and Southern Pygmy Salamander for
any of the variables examined, except for soil moisture, which was greater at Northern
Pygmy Salamander locations and may have been a spurious result. We observed pygmy
salamanders almost exclusively beneath wood cover-objects, and size of cover objects did
not differ from the size available in the surrounding environments. Total area of small-sized
down woody debris (DWD) and total area of large-sized DWD were the only variables
associated with the presence of pygmy salamanders, suggesting that pygmy salamanders
avoid predation and interspecific competition by selecting sites that minimize encounters
with larger salamanders.

Introduction
Desmognathus organi Crespi, Brown, and Rissler (Northern Pygmy Salaman-
der) and D. wrighti King (Southern Pygmy Salamander) are rare endemic species
generally associated with high-elevation forests of the southern Appalachian Moun-
tains (Crespi et al. 2010, Petranka 1998). In 2010, D. wrighti was split into 2 species
based on genetic, morphometric, and ecological measurements (Crespi et al. 2010).
The range of the 2 species is divided by the French Broad River, with Northern
Pygmy Salamander occurring north and east of the river, and Southern Pygmy

1
Department of Environmental Studies, University of North Carolina at Asheville, Ashe-
ville, NC 28804. 2North Carolina Wildlife Resources Commission, 177 Mountain Laurel
Lane, Fletcher, NC 28732. 3Department of Mathematics, University of North Carolina at
Asheville, Asheville, NC 28804. *Corresponding author - crrossell@aol.com.
Manuscript Editor: Max Nickerson

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Salamander occurring south and west of the river (Crespi et al. 2010). Both species
occur in highly localized populations restricted to the most prominent mountains
of western North Carolina, eastern Tennessee, and southwestern Virginia (Crespi
et al. 2003, 2010; Petranka 1998). Both Northern Pygmy Salamander and Southern
Pygmy Salamander are listed as Federal Species of Concern, with global rankings
of G3 (vulnerable; LeGrand et al. 2014). Populations in North Carolina, Virginia,
and Tennessee are considered vulnerable to extinction, with fewer than 100 occur-
rences documented in each state (LeGrand et al. 2014, Roble 2013, Withers 2009).
In North Carolina, both species are listed as State Rare (LeGrand et al. 2014) and
are considered Species of Greatest Conservation Need in the North Carolina Wild-
life Action Plan (NCWRC 2015). Threats to the species include loss and fragmenta-
tion of habitat, environmental perturbations that reduce canopy cover such as acid
rain deposition, outbreaks of insects including Choristoneura fumiferana Clemens
(Spruce Budworm) and Adelges piceae Ratzeburg (Balsam Woolly Adelgid); and
over-collecting for the pet industry (Hammerson 2004).
Pygmy salamanders are one of the smallest and most terrestrial of all Des-
mognathus species (Petranka 1998). They are unique in that embryos completely
absorb their gills before hatching (Organ 1961a, Petranka 1998). Pygmy sala-
manders are thought to have evolved in high-elevation Picea rubens Sarg. (Red
Spruce)–Abies fraseri (Pursh) Poir (Fraser Fir) forests (Crespi et al. 2003, 2010),
where they are most commonly found (Hairston 1949, King 1936, Organ 1961b),
but they also are known to occur in deciduous forests as low as 762 m eleva-
tion (Bruce 1977, Rubin 1971, Tilley and Harrison 1969). However, reports of
low-elevation occurrences are based on only a few observations (Bruce 1977,
Hairston 1949, Rubin 1971, Tilley and Harrison 1969). Several authors have
speculated that these low-elevation populations may be warm-tolerant, post-
glacial relicts that have persisted since the Pleistocene Era (Bruce 1977, Rubin
1971, Tilley and Harrison 1969). Currently, little is known about the relative
abundance of these lower-elevation populations, particularly as they compare to
populations at higher elevations.
To date, there are no quantitative studies describing the habitat of Northern
Pygmy Salamander and Southern Pygmy Salamander. Most of the available infor-
mation on their habitat is general, and either comes from studies documenting the
vertical and horizontal distribution of Desmognathus species in the mountains of
North Carolina (Hairston 1949, Organ 1961b) or from observational reports (e.g.,
Bruce 1977, King 1936, Rubin 1971, Tilley and Harrison 1969). In general, pygmy
salamander habitat is reported as mature forests in areas far from seeps and streams
(Hairston 1949, Organ 1961b, Petranka 1998). However, some females reportedly
nest in seeps and streambanks during the spring and early summer (Bruce 1977, Or-
gan 1961a, Petranka 1998, Rossell et al. 2016), and aggregations have been found
in streambanks in the fall and early winter (Bruce 1977, Organ 1961b). Pygmy Sala-
manders are known to utilize a variety of cover objects including logs, bark, rocks,
and moss (Bruce 1977, King 1936, Hairston 1949, Tilley and Harrison 1969), and
King (1936) indicated that they prefer small cover-objects versus larger ones. We

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C.R. Rossell Jr., Ivy C. Haas, L.A. Williams, and S.C. Patch

have made similar field observations (C. Rossell Jr. and L. Williams; pers. observ.).
However, preference for a habitat attribute cannot be determined with certainty un-
less the attribute is examined in terms of its use versus availability (Rossell et al.
2006, 2013).
In the genus Desmognathus, microhabitat associations can be used to predict
the macrohabitat of individual species (Krzysik 1979). Crespi et al. (2010) used
climate models as supporting evidence for the taxonomic revision of D. wrighti.
Their ecological-niche models indicated that localities of pygmy salamanders in
areas north of the French Broad River were generally colder and drier than those
south of the river. They hypothesized that unique environmental niche space was
produced where divergent selective pressures, coupled with isolation, diversified
the lineage of the species. However, this hypothesis remains untested because the
habitat of Northern Pygmy Salamander and Southern Pygmy Salamander has not
been quantified or compared. In this study we provide quantitative data on relative
abundance and microhabitat of these taxa across 3 forest types of different eleva-
tions in the mountains of North Carolina. In addition, we compare microhabitat
attributes used by both species and determine whether those attributes differ from
those randomly available in the surrounding environment.

Methods
We conducted our study from 29 June to 25 August 2015. We selected our study
sites from North Carolina Wildlife Resource Commission (NCWRC) historical oc-
currence records (Fig. 1). Occurrence records were divided into categories based on
3 major forest types at different elevations: spruce–fir forests at >1645 m, northern
hardwood forests at 1310–1645 m, and mountain cove forests at <1310 m (Schafale
2012). Schafale (2012) provided complete floral descriptions of the forest types.
In brief, spruce–fir forests are considered high-elevation forests dominated by Red
Spruce and Fraser Fir, and also may include Betula alleghaniensis Britt. (Yellow
Birch), Acer spicatum Lam. (Mountain Maple), and Sorbus americana Marshall
(Mountain Ash). Shrubs are often sparse, but may be dense, and herbaceous layers
vary from sparse to dense. Northern hardwood forests are mid-elevation forests
dominated by mesophytic hardwoods including Yellow Birch, Acer saccharum
Marshall (Sugar Maple), and Fagus grandifolia Ehrh. (American Beech). Shrub
layers range from fairly sparse to moderately dense, and herbaceous layers are
generally dense and moderately diverse. Mountain cove forests are low-elevation
forests dominated by mesophytic species including Liriodendron tulipifera L. (Tu-
lip Poplar), Prunus serotina Ehrh. (Black Cherry), Fraxinus americana L. (White
Ash), and Tsuga canadensis (L.) Carr. (Eastern Hemlock). Shrub and herbaceous
layers range from sparse to dense, and some herbaceous layers are highly diverse
(Schafale 2012).
We sampled 4 sites in each of the forest types for both Northern Pygmy Sala-
mander and Southern Pygmy Salamander. We chose sites within each forest type
that were as comparable in elevation as possible between the 2 species. We alter-
nated sampling of each species within 1–4 d of each other to minimize effects of
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changing weather patterns. We also sampled low-elevation sites first to increase our
chances of finding animals active on the surface as the season progressed.
We located pygmy salamander sites using GPS coordinates of occurrence re-
cords. We conducted a 2-h time-constrained search at each site (Corn and Bury
1990). A total of 4 individuals skilled at salamander surveys participated in
searches, and usually 2–3 individuals worked a site at any given time. All search
efforts were equal between sites. Researchers were careful to avoid overlap in areas
searched, and search time was paused during all captures of pygmy salamanders.
Searching for salamanders entailed looking under all potential cover-objects that
were at least 3 cm wide x 5 cm long, the smallest size considered suitable to conceal
an adult pygmy salamander (total length [TL] of adult pygmy salamander = 3.7–5.1
cm, Petranka 1998). During searches, we counted all salamanders and identified
them to species when possible. When we found a pygmy salamander, we paused the
search time to place the animal in a clean plastic bag with moss or leaf litter to pre-
vent its desiccation and flagged the location before resuming the search. Once the
search period expired, we determined the age and sex of each pygmy salamander
selected for habitat analyses (see details below). We based age determinations on

Figure 1. Location of study sites of Desmognathus organi (Northern Pygmy Salamander)

and D. wrighti (Southern Pygmy Salamander) based on historic occurrence records in North
Carolina, June–August 2015.
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the TL of animals (juveniles: TL < 3.7 cm, adults: TL ≥ 3.7 cm) and sex determina-
tion on the presence of a U-shaped mental gland in males and, in some cases, the
observation of eggs in gravid females (Petranka 1998). We released each animal at
its capture site immediately after processing.
We collected habitat data immediately after each time-constrained search.
At each pygmy salamander location (herein, referred to as occupied location),
we recorded type of cover object (rock, round wood, bark, etc.), and measured
cover-object dimensions (maximum length and width), and distance to water
(up to 30 m). We measured soil pH and soil moisture under the cover object at a
depth of 2 cm below the surface using a Kelway soil pH and moisture meter (Kel
Instruments, Wyckoff, NJ). We also measured the soil surface temperature under
the cover object using a Taylor digital thermometer (Taylor Precision Products,
Las Cruces, NM). We estimated percent canopy cover of vegetation >0.5 m in
height using a densitometer (Geographic Resource Solutions, Arcata, CA) held
0.5 m above the cover object. We used a 0.25-m2 quadrat frame centered over the
cover object to estimate percent cover of plants <0.5 m in height; percent ground
cover in each of 5 categories (bare soil, woody debris, leaf litter, rock, and wa-
ter); and percent substrate composition in each of 5 size-classes (modified from
Rosgen 1996): soil and sand (<2 mm), gravel (2–65 mm), cobble (66–250 mm),
boulder (>250 mm), and bedrock. We used a 1-m–radius circular plot centered on
the cover object to measure the density of down woody debris (DWD). We used
a Biltmore stick to measure the length and width of each piece of DWD that met
the minimum size criteria for cover objects (as described above); only portions
of DWD that were touching the ground and within the plot were measured. If we
captured more than 5 animals during a time-constrained search, we limited sam-
pling to 5 randomly selected pygmy salamander locations per site.
To determine whether habitat attributes at occupied locations differed from
those randomly available in the surrounding environment, we identified a corre-
sponding random location by finding a cover object 2–30 m from each occupied
location. We employed a random numbers table to generate a compass bearing (0–
360°) and a distance (2–30 m) from each occupied location to locate cover objects.
We considered a cover object to be suitable if it met the minimum size criteria (as
described above) and had no pygmy salamander under it. In the event there was no
suitable cover object at the random point, we followed the compass bearing until
a suitable object was found. Once we selected a cover object, we collected habitat
data using the same methods as described above. We collected data at each random
location immediately after collecting data at the occupied location.
We compared the total counts of pygmy salamanders between the 3 forest types
and between Northern Pygmy Salamander and Southern Pygmy Salamander jointly
using a negative binomial model for total counts at each site. We obtained the esti-
mated means of pygmy salamanders (i.e., relative abundance) for each forest type
and each species by reverse-transforming the estimates given by the model, which
utilized a log-link function. We employed the Tukey–Kramer procedure applied
to the negative binomial model to perform pairwise comparisons between forest

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types. To test for differences in microhabitat variables between the 2 species, we

combined data for the 3 forest types for each species because of limited sample size.
Cover-object size may be an important habitat attribute to Desmognathus species
(Keen 1982, Krzysik 1979); thus, we divided DWD into 2 categories: small DWD
(width < 6 cm) and large DWD (width ≥ 6 cm). Cutoff between categories was based
on the maximum TL of adult pygmy salamanders (5.1 cm; Petranka 1998). We then
calculated means of habitat variables for occupied and random locations for each
species and compared them using independent-sample t tests with the Satterthwaite
approximation of degrees of freedom. We ran conditional logistic regression mod-
els with a stepwise procedure to compare habitat variables of occupied and random
locations to determine if pygmy salamanders used any habitat attribute differently
than was randomly available. We set an alpha of P < 0.05 to retain variables in the
model. We used a separate conditional logistic regression model to determine if
pygmy salamanders used smaller cover-objects than were randomly available in the
surrounding habitat. We considered all statistical comparisons significant at alpha
< 0.05. We considered an attribute selected if its use differed from its random avail-
ability. We performed all statistical analyses in Statistical Analysis System (SAS,
version 9.2; SAS Institute, Cary, NC).

Results
We captured a total of 98 pygmy salamanders (41 Northern Pygmy Salamanders,
57 Southern Pygmy Salamanders) during 24, two-hour time-constrained searches.
The estimated mean number of pygmy salamanders for all forest types combined
did not differ between Northern Pygmy Salamander and Southern Pygmy Salaman-
der (χ2 = 1.19, df = 1, P = 0.276). We found 52 pygmy salamanders in spruce–fir
forests, 26 in northern hardwood forests, and 20 in cove forests. The estimated
mean number of pygmy salamanders was significantly greater in spruce–fir forests
(estimated mean = 6.38; 95% CI = 4.20, 9.69) than cove forests (estimated mean =
2.45; 95% CI = 1.43, 4.22; P = 0.017), but was not significantly different between
spruce–fir and northern hardwood forests (estimated mean = 3.26; 95% CI = 1.98,
5.36; P = 0.107), or between northern hardwood and cove forests (P = 0.729). In
addition to pygmy salamanders, we recorded 655 other salamanders representing
15 species: 173 in spruce–fir forests, 252 in northern hardwood forests, and 230 in
cove forests (Appendix 1). Appendix 1 provides the relative abundance of salaman-
ders at Northern Pygmy Salamander and Southern Pygmy Salamander sites in each
forest type.
We collected habitat data at 73 pygmy salamander locations (36 Northern
Pygmy Salamander locations, 37 Southern Pygmy Salamander locations) in 32
spruce–fir forests, 24 northern hardwood forests, and 17 cove forests (elevation
range of all sites = 966–1937 m). The sex and age composition of Northern Pygmy
Salamander and Southern Pygmy Salamander representing the habitat data were
similar between the 2 species (Table 1). The total sex ratio (males:females) of the
sample population was 1.04 (24 males, 23 females), with adults comprising 66%
(n = 47) of the population and juveniles 34% (n = 24) (Table 1). Summary statistics
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of the habitat variables at occupied and random Northern Pygmy Salamander and
Southern Pygmy Salamander locations are provided in Tables 2 and 3, respective-
ly. None of the habitat variables differed between occupied locations of Northern
Pygmy Salamander and Southern Pygmy Salamander (all P > 0.05, Table 2),
except for soil moisture which was significantly greater at Northern Pygmy Sala-
mander locations (P = 0.005, Table 2). There were also no differences in any of the
habitat variables between random locations (all P > 0.05, Table 3), except canopy
cover, which was significantly less at Northern Pygmy Salamander locations (P =
0.039, Table 3). The stepwise logistic-regression models indicated that the only
variables significantly associated with pygmy salamander locations were total area
of small DWD (P = 0.0004) followed by total area of large DWD (P = 0.015); for
all other variables, P > 0.05, (Table 4). The odds-ratio estimates for total area of
small and large DWD were 1.398 and 1.052, respectively. This result indicates that
the odds of a pygmy salamander being present at a location increases by factors of
1.398 and 1.052, respectively, as area of small DWD and area of large DWD in-
crease by 100 cm2.

Table 1. Age and sex composition of Pygmy Salamanders represented in the habitat analysis. Data are
missing for 1 Northern Pygmy Salamander and 1 Southern Pygmy Salamander.

AttributeA Northern Pygmy Salamander Southern Pygmy Salamander

Juvenile 12 12
Adult 23 24
Male 13 11
Female 10 13

Table 2. Summary statistics of habitat variables at occupied locations at historic occurrence sites of
Northern Pygmy Salamander and Southern Pygmy Salamander in North Carolina, June–August 2015.
Data presented are for occupied locations of Northern Pygmy Salamander (n = 36) and Southern Pyg-
my Salamander (n = 37) in 32 spruce–fir forests, 24 northern hardwood forests, and 17 cove forests.

Historic occurrence sites of

Northern Pygmy Southern Pygmy
Salamander Salamander
(n = 12) (n = 10)
Variable Mean SD Mean SD t value df P-value
Soil surface temperature ( C) 16.9
o
1.7 17.3 2.0 -0.51 17.8 0.618
Soil moisture (%) 59.9 12.2 41.9 13.6 3.18 18.2 0.005
Soil pH 6.5 0.4 6.6 0.2 -0.42 14.5 0.683
Distance to water (m) 26.8 7.7 25.5 8.2 0.36 18.7 0.721
Plant cover (%) 19.1 15.8 16.1 13.4 0.48 19.9 0.640
Canopy cover (%) 75.0 11.9 83.4 7.9 -1.96 19.1 0.065
Soil (%) 67.0 33.9 85.1 25.0 -1.43 19.8 0.167
Gravel/cobble (%) 0.0 0.0 2.5 5.6 -1.38 9.0 0.202
Boulder (%) 30.9 35.2 12.5 23.6 1.46 19.2 0.151
Other substrate (%) 2.1 5.8 0.0 0.0 1.24 11.0 0.241
Area of small DWD (cm2) 695.0 417.3 489.0 263.9 1.41 18.8 0.176
Area of large DWD (cm2) 2458.1 5539.7 1580.1 928.7 0.54 11.7 0.599

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Cover objects used by pygmy salamanders consisted of either round wood (n =

59), bark (n = 13), or leaf litter (n = 1). We detected no pygmy salamanders under
rocks or other types of cover objects. The size of cover objects used by pygmy
salamanders (mean = 384.3 cm2, SD = 476.1) did not differ from those randomly
available in the surrounding habitat (mean = 496.0 cm2, SD = 646.9, P = 0.21).
Table 3. Summary statistics of habitat variables recorded at random locations at historic occurrence
sites of Northern Pygmy Salamander and Southern Pygmy Salamander in North Carolina, June–Au-
gust 2015. Data presented are for random locations of Northern Pygmy Salamander (n = 36) and
Southern Pygmy Salamander (n = 37) that correspond to occupied locations in 32 spruce–fir forests,
24 northern hardwood forests, and 17 cove forests.

Historic occurrence sites of

Northern Pygmy Southern Pygmy
Salamander Salamander
(n = 12) (n = 10)
Variable Mean SD Mean SD t value df P-value
Soil surface temperature ( C) 16.9
o
1.3 17.5 2.1 -0.81 14.9 0.428
Soil moisture (%) 48.5 12.3 42.8 8.8 1.26 19.6 0.224
Soil pH 6.4 0.3 6.5 0.2 -1.04 19.8 0.309
Distance to water (m) 27.2 6.5 25.3 7.8 0.63 17.5 0.539
Plant cover (%) 30.7 19.3 23.5 19.6 0.86 19.2 0.402
Canopy cover (%) 72.1 21.0 86.9 7.2 -2.28 14.0 0.039
Soil (%) 86.1 24.0 87.3 18.2 -0.13 19.9 0.897
Gravel/cobble (%) 2.1 5.0 3.8 8.0 -0.59 14.5 0.565
Boulder (%) 13.5 26.5 6.5 11.6 0.83 15.7 0.417
Other substrate (%) 0.8 2.9 0.5 1.6 0.34 17.6 0.736
Area of small DWD (cm2) 255.7 175.5 291.2 197.0 -0.44 18.3 0.664
Area of large DWD (cm2) 797.0 716.0 1254.9 1084.6 -1.14 15.1 0.271

Table 4. Results of conditional logistic-regression models with a stepwise procedure comparing

habitat variables of occupied and random locations of pygmy salamanders in North Carolina, June–
August,2015. Habitat data of Northern Pygmy Salamander (n = 36) and Southern Pygmy Salamander
(n = 37) locations were combined for analysis.

P-value
With area of With area of
Variable 1-variable small DWD small and large DWD
Soil surface temperature (oC) 0.871 0.348 0.779
Soil moisture (%) 0.011 0.047 0.054
Soil pH 0.209 0.383 0.182
Distance to water (m) 0.950 0.366 0.304
Plant cover (%) 0.021 0.177 0.266
Canopy cover (%) 0.306 0.228 0.574
Soil (%) 0.043 0.055 0.081
Gravel/cobble (%) 0.232 0.345 0.516
Boulder (%) 0.016 0.039 0.067
Other substrate (%) 0.891 0.907 0.476
Area of small DWD (cm2) 0.0004 - -
Area of large DWD (cm2) 0.023 0.015 -

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Discussion
Searches of natural cover-objects have been shown to be a reliable index for
absolute population size of terrestrial salamanders (Smith and Petranka 2000). The
finding that the overall relative abundance of Northern Pygmy Salamander and
Southern Pygmy Salamander did not differ, suggests that the size of the popula-
tions at our study sites was comparable, despite the fact that we were unable to
find Southern Pygmy Salamander at 2 sites: a low-elevation cove forest and a
mid-elevation northern hardwood forest. All sites were similar in that they were
relatively intact stands of mature forests. The sex and age composition of Northern
Pygmy Salamander and Southern Pygmy Salamander also were similar between
the 2 species (Table 1). The total sex ratio (males:females) was 1.04, and the total
age ratio (adults:juveniles) was 1.96, in contrast to the findings of Organ (1961b),
who reported a total sex ratio of 1.62 and a total age ratio of 0.95 for populations
of Northern Pygmy Salamander in the Balsam Mountains of southwestern Virginia.
Pygmy salamander populations in our study appear to be relatively vigorous com-
pared to those studied by Organ (1961b) in the late 1950s.
We captured twice as many pygmy salamanders in spruce–fir forests than in
northern hardwood forests and 2.6 times as many than in cove forests, which sug-
gests that the most robust populations occur in spruce–fir forests. This result agrees
with Hairston (1949) and Organ (1961b), who reported higher densities of pygmy
salamanders in spruce–fir forests. However, we found that the estimated mean
number of pygmy salamanders in spruce–fir forests was not significantly different
from that in northern hardwood forests. This finding should be viewed with caution
because of the low power of the statistical test due to the small sample size and high
variability of the number of animals captured between sites within a forest type.
The relative abundance of pygmy salamanders in cove forests was not signifi-
cantly different from populations in northern hardwood forests; thus, population
size of pygmy salamanders in some low-elevation cove forests may be comparable
to those in northern hardwood forests. We also observed this pattern in a low-
elevation cove site in the Great Smoky Mountains National Park that had the 5th
highest capture rate (n = 7) of the 24 sites in our study. Low-elevation populations
also may be relatively long-lived and persistent when the habitat remains intact, as
exemplified by the successful capture of pygmy salamanders at 7 of the 8 historical
low-elevation sites, including a 44-year-old record of Southern Pygmy Salamander
in the Great Smoky Mountains National Park (same site as mentioned above). It
also is worth noting that the low-elevation site that failed to yield any pygmy sala-
manders was a Southern Pygmy Salamander site impacted by a heavy infestation of
Adelges tsugae Annand (Hemlock Woolly Adelgid). Although no cause and effect
can be determined from 1 observation, we suspect that a reduced canopy from the
loss of numerous large Eastern Hemlock trees may be the reason for the apparent
demise of this population. Currently, the effects of Hemlock Woolly Adelgid on
salamander populations are unknown. However, variables that could affect sala-
manders, including coarse woody debris, herbaceous cover, and light levels, have
been shown to significantly increase in Hemlock stands with canopy decline caused
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C.R. Rossell Jr., Ivy C. Haas, L.A. Williams, and S.C. Patch

by the Hemlock Woolly Adelgid (Cleavitt et al. 2008). Therefore, further study is
needed to investigate the impacts of Hemlock Woolly Adelgid on pygmy salaman-
ders in low-elevation cove forests.
Strong evidence suggests that predator avoidance and interspecific competi-
tion with other salamanders shape the habitat selection of Desmognathus species
in the southern Appalachian Mountains (e.g., Bruce 1996, Hairston 1986, Krzysik
1979, Southerland 1986). We recorded 655 salamanders inhabiting the same ar-
eas as pygmy salamanders, with the majority either Desmognathus or Plethodon
species (Appendix 1). The total number of salamanders observed in each of the
3 forest types far exceeded the total number of pygmy salamanders in each for-
est type (Appendix 1). These findings suggest that the general habitat of pygmy
salamanders is suitable to a variety of salamander species, and that pygmy sala-
manders are exposed to potential predation by and interspecific competition from
other salamanders.
The findings that none of the habitat variables differed between Northern Pygmy
Salamander and Southern Pygmy Salamander, except for soil moisture, suggests
that these 2 species use similar types of microhabitats (Table 2). We are uncertain
of the reason why soil moisture levels would be higher at Northern Pygmy Sala-
mander locations compared to Southern Pygmy Salamander locations; perhaps it is
a spurious result given the small sample size. Therefore, additional data are needed
to validate this result. Nonetheless, our results do indicate that the microhabitat of
pygmy salamanders may be characterized by: relatively high soil moisture, neutral
soil pH, moderate plant cover, a closed canopy, moderate amounts of exposed soil,
and large amounts of small- and large-sized DWD (Table 2). In addition, micro-
habitat was not associated with surface water; 78% (n = 57) of pygmy salamander
locations were located >30 m from seeps or streams. Although analyses of micro-
habitat associations have not been previously reported for pygmy salamanders, our
results generally agree with those of others who have reported that pygmy salaman-
ders utilize relatively moist areas in undisturbed forests (Hairston 1949, King 1936,
Tilley and Harrison 1969). In addition, Hairston (1949) and Organ (1961b) both
noted that pygmy salamander habitat is often found far from water, and suggested
that this is an adaptation to minimize potential predation from larger predatory
salamanders that reside closer to water.
Our results indicate that total amount of small DWD was the most important
habitat variable related to the presence of pygmy salamanders followed by total
amount of large DWD (Table 4). The highly significant association of small DWD
in the final regression model also highlights the importance of this attribute to
pygmy salamanders (Table 4). Cover objects protect salamanders from stressful
environmental conditions because they provide a cool, moist refuge from predators
and a possible source of invertebrate prey (Heatwole 1962, Jaeger 1980, Mathis
1990). Density of cover objects has been reported as an attractive habitat attribute
for several salamander species. For example, Grover (1998) reported that the abun-
dance of Plethodon glutinosus Green (Northern Slimy Salamander) and P. cinereus
Green (Eastern Red-backed Salamander) was greater in plots with a high density of

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cover objects than in plots with a low density of cover objects. Numerous authors
also have suggested that salamanders differentially use cover objects relative to
their body length to avoid intra- and interspecific encounters (e.g., Colley et al.
1989, Keen 1982, Kryzysik 1979, Southerland 1986). Therefore, we suggest that
pygmy salamanders prefer microhabitats with greater amounts of small DWD to
minimize potential encounters with larger salamanders. It also should be noted that
pygmy salamanders may select microhabitats with higher soil-moisture levels; soil
moisture was significant in the first step of the regression model when not adjust-
ing for other variables (P = 0.011), and was almost significant in the last step of the
regression model after adjusting for small and large DWD (P = 0.054) (Table 4).
However, additional research is needed to verify this assertion.
Pygmy salamanders use a variety of cover objects including logs, rocks, moss,
and leaf litter (Caruso 2016, Hairston 1949, King 1936, Organ 1961b, Tilley and
Harrison 1969). We found pygmy salamanders almost exclusively under wood cov-
er-objects, consisting of either round wood (81%) or bark (18%). These specimens
included 3 gravid females under small round wood and 3 under bark, with 5 located
>30 m from water and one 9.7 m from water. Caruso (2016) reported that the ma-
jority of cover objects used by Southern Pygmy Salamander in the Great Smoky
Mountains National Park consisted of round wood (82%), followed by rocks (14%)
and bark (4%). Caruso (2016) also reported that terrestrial Desmognathus species
preferred wood cover-objects over rock cover-objects. This preference for wood is
likely related to the higher moisture levels maintained under wood cover than rock
cover (Grover 1998).
The finding that size of cover objects used by pygmy salamanders did not differ
from the size of cover objects randomly available suggests that pygmy salamanders
showed no preference for small-sized cover objects. However, this finding should
be viewed with caution, as observations during our field work suggest that indi-
vidual pieces of small-sized DWD were much more prevalent across the forest floor
than pieces of large-sized DWD. Therefore, any preference pygmy salamanders
might have for smaller-sized cover objects may not be apparent in our analysis.
However, data from Caruso (2016) indicate that the mean size of cover objects
used by Southern Pygmy Salamander were consistently the smallest among 11 spe-
cies of salamanders in the Great Smoky Mountains National Park. That finding is
consistent with those of King (1936) that pygmy salamanders are found most often
under small rather than large cover-objects.
The results of our study indicate numerous similarities between populations of
Northern Pygmy Salamander and Southern Pygmy Salamander among the 3 for-
est types investigated. Crespi et al. (2010) purported that the environmental niche
space between the 2 species is unique because of different climatic conditions on
either side of the French Broad River. Our findings that none of the microhabitat
variables differed between the 2 species for either the occupied (with the excep-
tion of soil moisture, see discussion above; Table 2) or random locations (with the
exception of percent canopy cover, both means >72%; Table 3) do not support this
assertion, and suggest that the environmental niche space is commensurate between

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the 2 species. Atypical weather did not likely affect results of our study because
average temperatures and total precipitation for August 2015 were comparable to
historic averages at several weather stations in the northern and southern study
areas (US Climate Data 2017a,b). In view of these findings, we suggest further
studies are needed to investigate the effects of incorporating microhabitat data into
ecological-niche models that rely solely on climatic data. In addition, more research
is needed across different seasons and years to validate our conclusion of similar
environmental-niche space between the 2 species.

Acknowledgments
We thank Bill and Peggy Steiner for providing a scholarship to I. Haas to help fund her
participation in this research, C. Lawson and K. Parker for field assistance, and I. Rossell
for helpful comments on the manuscript. We also thank K. Weeks and A. Boynton, with the
NCWRC’s Wildlife Diversity Program, for administrative and project support.

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Appendix 1. Total number of salamanders recorded at historic occurrence sites of Northern

Pygmy Salamander and Southern Pygmy Salamander in 3 forest types in North Carolina,
June–August 2015. Totals are based on number of animals counted during four 2-h time-
constrained searches in each forest type at Northern Pygmy Salamander and Southern
Pygmy Salamander sites. C = cove, NH = northern hardwood, and SF = spruce–fir.

Forest Type
Salamander species C NH SF
Northern Pygmy Salamander Sites
Desmognathus organi Crespi, Browne, and Rissler (Northern 7 14 20
Pygmy Salamander)
Desmognathus carolinensis Dunn (Carolina Mountain Dusky 72 4 14
Salamander)
Plethodon montanus Highton & Peabody (Gray-cheeked Salamander) 54 34 19
Plethodon cylindraceus (Harlan) (White-spotted Slimy Salamander) 3 1 1
Plethodon yonahlossee Dunn (Yonahlossee Salamander) 1 0 0
Notophthalmus v. viridescens (Rafinesque) (Red-spotted Newt) 2 0 0
Eurycea wilderae Dunn (Blue Ridge Two-lined Salamander) 25 4 8
Unknown spp. 0 3 0

Southern Pygmy Salamander

Desmognathus wright King (Southern Pygmy Salamander) 13 12 32
Desmognathus fuscus (Rafinesque) (Dusky Salamander) 0 6 0
Desmognathus santeetlah Tilley (Santeetlah Dusky Salamander) 1 8 5
Desmognathus imitator Dunn (Imitator Salamander) 0 1 0
Desmognathus ocoee Nicholls (Ocoee Salamander) 7 8 42
Desmognathus spp. 2 0 18
Plethodon metcalfi Brimley (Southern Gray-cheeked Salamander) 30 156 32
Plethodon jordani Blatchley(Red-cheeked Salamander) 30 4 2
Plethodon teyahalee Hairston (Southern Appalachian Salamander) 1 1 0
Plethodon shermani Stejneger (Red-legged Salamander) 0 6 0
Plethodon serratus Grobman (Southern Redback Salamander) 0 1 0
Eurycea wilderae Dunn (Blue Ridge Two-lined Salamander) 2 2 12

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