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2014 Vol. 13, No. 1
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2014 SOUTHEASTERN NATURALIST 13(1):166–175
The Land Snails of White Oak Sinks, Great Smoky
Mountains National Park, Tennessee
Daniel A. Douglas1,*, Daniel C. Dourson2, and Ronald S. Caldwell3
Abstract - Land snails are an essential component of forested ecosystems, but natural histories
for many species are not known, and distribution records are incomplete. This study
examined the land snails present at White Oak Sinks, Great Smoky Mountains National
Park, TN and encompasses surveys that took place over a 9-year period. We collected 58
species during this survey, with 17 new county records for Blount County and several species
with affinities for other regions. Due to the growing threats to biodiversity, a great deal
of attention needs to be given to organisms that are poorly studied. These results highlight
the growing need for further study of land snails.
Introduction
Terrestrial gastropods (i.e., land snails) are often overlooked and understudied,
with the result that basic natural history information as well as distributional
records are lacking in the current literature. Hubricht (1985) presents the most
detailed distributional records for the land snails of the eastern United States;
however, these records are incomplete for many species, especially micro-snails
(<5 mm in diameter). Yet the loss of global biodiversity among these taxa is
increasing (Lydeard et al. 2004). Because there are increased pressures (e.g.,
climate change, pollution [such as acid rain], deforestation) on many flora and
fauna, it is crucial to identify biodiversity hotspots and work to protect and properly
manage these areas.
Land snails are ever-present components of forested ecosystems in the southeastern
United States and play a role in many ecological processes. Decomposition
rates of coarse woody debris are accelerated in their presence (Kappes 2005,
Kappes et al. 2006), and they assist in the detoxification of forest soils through calcium
(Ca) sequestration (Pearce 2008). Land snails also serve as food and a source
of Ca for many vertebrate species and often effect the distribution of these organisms
in forests (Graveland and van der Wal 1996, Graveland et al. 1994, Harper and
Guynn 1999). Because of these characteristics, the presence of land snails is a vital
component of basic forest ecology.
The Great Smoky Mountains National Park (GSMNP) is recognized worldwide
as a center for biodiversity due to high levels of precipitation, varying geology,
and diverse vegetative communities (Nichols and Langdon 2007). The land-snail
1Department of Biological Sciences, Eastern Kentucky University, 521 Lancaster Avenue,
Richmond, KY 40475. 2Belize Foundation for Research and Environmental Education, PO
Box 129, Punta Gorda, Belize. 3Department of Biology and Cumberland Mountain Research
Center, Lincoln Memorial University, 6965 Cumberland Gap Parkway, Harrogate, TN
37752. *Corresponding author - dnl.douglas83@gmail.com.
Manuscript Editor: Lance Williams
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2014 Vol. 13, No. 1
fauna of GSMNP has been poorly studied with the earliest reports coming from the
beginning of the 20th century (Ferriss 1900, Pilsbry 1900). In the latter half of the
1900s, a more detailed study was initiated in which species diversity among forest
types was positively correlated with heterogeneity of canopy tree species (Getz
1974). Recent surveys have included descriptions of new species (Dourson 2012)
and highlighted species assemblages present in high-elevation habitats that are currently
under siege from anthropogenic activities (Dourson and Langdon 2012).
Nonetheless, basic survey work is still needed for this area to document all species
present as well as describe those previously unknown to science before any
adverse effects of climate change alter the species composition. In 1998, GSMNP
initiated an All Taxa Biodiversity Inventory (ATBI) to document all the flora and
fauna present within its boundaries. The results of the surveys discussed in this
manuscript were carried out as part of the ATBI.
Methods
Study site
The results reported here encompass surveys that took place in 1998, 2006, and
2007 in White Oak Sinks (WOS), GSMNP, Blount County, TN. WOS is located
in the Blue Ridge Mountains ecoregion (Level III) (Griffith et al. 1998). This is
an area of second-growth forests predominately composed of Quercus prinus L.
(Chestnut Oak) and Quercus (oak)-Carya (hickory) forests (Jenkins 2007). The geology
of the area is dominated by karst topography with Jonesboro, Kingsport, and
Lenoir limestone, as well as Mascot, Longview, Chapultepec, and Copper Ridge
dolomite (Miller 1974, USGS 1968).
Field methods
We sampled land snails using an opportunistic design based on knowledge of
habitat requirements. Two techniques were utilized during these surveys. The first
technique was used to survey for macro-snails (>5 mm in diameter) and entailed
25 timed searches of 30 person minutes. Because most species of land snails are
habitat specialists, investigators searched specific microhabitats which included
under leaf litter, rocks, downed woody material (DWM), exfoliating bark present
on standing snags and downed woody debris, hollow trees and forks of trees where
detritus had accumulated, damaged trees excreting sap, cliff lines, caves, and rock
talus features. The second sampling technique was used to search for micro-snails
and included the collection of organic material just beneath leaf litter on the forest
floor (i.e., duff) as well as detritus from DWM, forked trees, and rock talus. Organic-
material samples were placed into liter-sized bags and dried for a two-week
period. We then sorted samples using no. 4, 10, 16, and 35 soil sieves and identified
specimens to species with the aid of a dissecting microscope.
Identification and taxonomy
All identifiable shells were assigned to species using Burch (1962), Pilsbry (1940,
1946, 1948) and the reference collection at the Cumberland Mountain Research
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Center, Lincoln Memorial University, Harrogate, TN. New county records were
determined using Hubricht (1985), and taxonomy followed Turgeon et al. (1998).
Results
This study yielded 58 species (several having state and global rankings), representing
13 families, 31 genera, and 17 new county records. Compared to other
areas that have been surveyed in GSMNP, the WOS has the highest land-snail
species richness (Appendix 1). The faunal assemblage collected at WOS included
species with affinities for areas elsewhere in the eastern US as well as a species,
Vertigo tridentata Wolf, 1870 (Honey Vertigo), that has never been collected in
GSMNP before. Of the 17 new county records discovered, including the previously
mentioned species, 23.5% had affinities for regions that are farther north.
The remaining county records were species that are widely distributed, but have
never been detected in Blount County, TN. Of the total species found, 15.5% were
species normally found in other ecoregions of Tennessee and/or other portions of
the southeastern United States. Aside from new county records, the survey also
recorded a proposed new species of Stenotrema and several specimens of what was
once thought to be Carychium mexicanum Pilsbry, 1891 (Southern Thorn), a species
endemic to the deep south, but which is now classified as Carychium exile I.
Lea, 1842 (Ice Thorn) (Weigand et al. 2011).
Discussion
The high species richness present at WOS is likely due to geology and Ca content
of the soil/leaf litter. Calcium carbonate is a known limiting factor for many
species of land snails because it is a requirement for several biological processes
(Burch and Pearce 1990). Availability of this mineral is also known to drive snail
diversity (Hotopp 2002). Though soil Ca was not quantified in this study, the
karst geology at WOS should allow for high soil Ca content, thus providing this
critical mineral and allowing for many species to be present. High soil Ca would
also allow trees that are known (e.g., Cornus florida L. [Flowering Dogwood];
Nation 2007) and hypothesized (e.g., Juglans nigra L. [Black Walnut], Juglans
cinerea L. [Butternut], and Tilia spp. [basswood]; D.A. Douglas, D.C. Dourson,
and R.S. Caldwell, pers. obervs.) “calcium-pumps” (i.e., continuously cycle Ca
from belowground up to their leaves and then back to the soil through leaf fall),
to be present. This level of Ca availability would create an opportunity for many
species, including rare species and those with affinities for other regions, to be
present and warrants further study. In places with generally low overall levels
of Ca, like much of the southern Appalachians, forested areas that have high Ca
availability become land-snail diversity hotspots, such as WOS. However, areas
thought to lack Ca availability (i.e., high elevations) at GSMNP have also yielded
high species richness, though it was hypothesized to be a product of Ca availability
made possible by deposition of plant material through leaf abscission (Dourson
and Langdon 2012). The prevalence of several species that are known only to ocSoutheastern
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2014 Vol. 13, No. 1
cur in the presence of high levels of Ca (Appendix 1) indicates that Ca is available
in large quantities at WOS. Due to the Ca availability and high species richness
found at WOS, this area is a land-snail biodiversity hotspot in the GSMNP, as
well as the southern Appalachian region.
Aside from determining that WOS is a diversity hotspot, another significant
outcome of this study was the discovery of several species of land snails that have
affinities for other portions of the eastern United States. The southern Appalachian
region was divided into an eastern division (i.e., Blue Ridge) and a western division
(i.e., Cumberland Plateau) in the early 1900s (Pilsbry 1900), and it has long since
been thought that the Ridge and Valley ecoregion of east Tennessee was a barrier
for dispersal from either division. However, small, isolated pockets in the Ridge
and Valley where species from both divisions coexist have been recently discovered
(Douglas et al. 2010) suggesting that these species may be more widely distributed
than once thought. Over the past 30 years, several studies have documented species
typical of the Blue Ridge to be present on the Cumberland Plateau (Dourson
and Beverly 2008, Hubricht et al. 1983). These findings suggest that at one point
in time, the Ridge and Valley, or the area that would become the Ridge and Valley,
was not a barrier to dispersal. Our findings at WOS strengthen the argument against
the hypothesis that the Ridge and Valley has not always been a barrier for dispersal
and suggest that 1) these species became isolated from one another as the Ridge and
Valley formed and/or 2) these species are more widely distributed across the Ridge
and Valley. Thus, future studies should focus on locating these populations thereby
bridging the gaps in distribution records.
One explanation of the high diversity found at WOS may be attributable to the
effect that merging of ecoregions has on shaping species composition and diversity
of an area. Similar diversity, richness, and affinities have been reported from localities
in Kentucky where two or more ecoregions come together (Dourson 2007,
Dourson and Beverly 2008). While WOS is completely within the Blue Ridge
ecoregion, it is positioned at the edge against the Ridge and Valley. We report similar
findings as the two previously mentioned studies from Kentucky. These findings
suggest that the merger of multiple ecoregions provides habitat characteristics, geology,
possible forest structure, and availability of various other resources required
to generate high land-snail biodiversity.
Of the 17 new county records found for Blount County at WOS, 14 were microsnails.
Though land snails in general are poorly studied, micro-snails are the more
understudied of the two size classes. Our findings help fill in gaps in distribution
records, as do most studies of land snails, but it also highlights an urgent need for
further study. Micro-snails show great promise in terms of use as indicators of ecosystem
health and past land-use histories (Douglas et al. 2013). They are also an
often overlooked foundational component of many ecosystems, particularly in the
deciduous forests of the eastern United States.
Presently, species are being lost at an alarming rate. This trend, coupled with
the uncertainty that revolves around climate change and its effects on biodiversity
(Bellard et al. 2012), underlines biologists’ need to recognize that many species
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2014 Vol. 13, No. 1
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of neglected organisms, like land snails, could be lost before they are truly understood,
and emphasizes the need for basic natural history surveys and data-driven
research to help identify the important roles that land snails play in an ecosystem.
Proposed new species
While it is easy to hypothesize that undescribed land snails exist in remote
mountain ranges of the Southern Appalachians, it is doubtful that new species are
within easy reach. However, in a well-studied area like WOS, there has remained
at least one species of land snail, yet to be described. This species was misidentified
by past collectors in their surveys of the park. Specimens of this undescribed
species can be found in the GSMNP archive collections, mislabeled as S. stenotrema
(Pfeiffer, 1842) (Inland Slitmouth) (D.C. Dourson, pers. observ.). However,
the undescribed species is typically not found alongside the Inland Slitmouth (a
species with which it was synonymized) (D.C. Dourson, pers. observ.), although
they do occasionally co-occur. Given these circumstances, one might speculate
why this species has remained ambiguous. The undescribed species does not
come close to other Stenotrema species found in the GSMNP in terms of basic
shell interpretation, especially when it, the Inland Slitmouth, S. hirsutum (Say,
1817) (Hairy Slitmouth), S. magnifumosum (Pilsbry, 1900) (Appalachian Slitmouth),
S. altispira (Pilsbry, 1894) (Highland Slitmouth), S. depilatum (Pilsbry,
1895) (Great Smoky Slitmouth) and S. pilula (Pilsbry, 1900) (Pygmy Slitmouth)
are examined together. In terms of shell morphology, the undescribed species
stands closest to S. macgregori (Dourson, 2011) (Fraudulent Slitmouth) and, to
a lesser degree, S. angellum Hubricht, 1958 (Kentucky Slitmouth). All three species
have similar but isolatable shell characteristics. However, the Fraudulent
Slitmouth and the Kentucky Slitmouth have primary distributions in the Pine
Mountain and Bluegrass regions (respectively) of Kentucky and are isolated from
the GSMNP by the Valley and Ridge provinces of Tennessee and western Virginia.
For this reason, they do not seem likely allied. The proposed species will be
described in a future manuscript (D.C. Dourson, unpubl. data).
Acknowledgments
All authors wish to acknowledge Discover Life in America for initiating and supporting
this research as well as K. Langdon and B. Nichols of the National Park Service for
allowing us to collect samples from the site. D.A. Douglas wishes to thank the Cumberland
Mountain Research Center for providing transportation during this study, and D.A.
Douglas and D.C. Dourson wish to thank the Great Smoky Mountains Institute at Tremont
for providing housing during the 2007 surveys. J. Dourson aided in the collection of specimens
and leaf litter samples.
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Appendix 1. Land snail species found at White Oak Sinks, Great Smoky Mountains National Park. Status, affinity, and habitat designations are as follows:
CO = common, LR = limited range, UN = uncommon, BR = Blue Ridge, CP = Cumberland Plateau, NOR = Northern, OA = Ouachita, OZ = Ozark, RV =
Ridge and Valley, WID = widespread, Cal = calciphyle, DWM = downed woody material, GEN = generalist, LL = leaf litter, MAR = marginal (i.e., field
edges, roadsides, railroad tracks), and ROC = rock talus. State and Global rankings are as follows: S2 = very rare and imperiled within the state; S3 =
vulnerable, rare, and uncommon in the state; G2 = imperiled, high risk of extinction; G3 = vulnerable, moderate risk of extinction; G4 = apparently secure,
uncommon. Multiple Global ranks (i.e., G#G#) indicate a range of uncertainty in the status of species (Master et al. 2012). The table was compiled using
information available in Hubricht (1985), Master et al. (2012), and Withers (2009).
Farmily/species Common name Status Affinity Habitat County record State/Global Rank
Carychiidae
Carychium clappi Hubricht, 1959 Appalachian Thorn CO WID LL
Carychium exiguum (Say, 1822) Obese Thorn CO NOR LL X
Carychium exile I. Lea, 1842 Ice Thorn CO NOR LL X
Carychium nannodes G.H. Clapp, 1905 File thorn CO WID LL
Cionellidae
Cionella morseana (Doherty, 1878) Appalachian Pillar CO WID LL
Discidae
Anguispira mordax (Shuttleworth, 1852) Appalachian Tigersnail LR BR, CP DWM
Discus nigrimontanus (Pilsbry, 1924) Black Mountain Disc LR BR, CP, OZ LL
Discus patulus (Deshayes, 1830) Domed Disc CO WID DWM, LL
Haplotrematidae
Haplotrema concavum (Say, 1821) Gray-foot Lancetooth CO WID LL
Helicarionidae
Euconulus dentatus (Sterki, 1893) Toothed Hive CO WID LL
Euconulus trochulus (Reihhardt, 1883) Silk Hive CO WID LL
Guppya sterkii (Dall, 1888) unnamed CO WID
Helicodiscidae
Helicodiscus singleyanus (Pilsbry, 1889) Smooth Coil LR WID MAR X
Philomycidae
Pallifera dorsalis (A. Binney, 1885) Pale Mantleslug LR NOR LL X
Philomycus carolinensis (Bosc, 1802) Carolina Mantleslug CO WID LL
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Farmily/species Common name Status Affinity Habitat County record State/Global Rank
Polygyridae
Appalachina chilhoweensis (Lewis, 1870) Queen Crater LR BR, CP LL, ROC
Fumonelix christyi (Bland, 1860) Glossy Covert LR BR LL S2, G3
Inflectarius rugeli (Shuttleworth, 1852) Deep-tooth Shagreen CO WID DWM, LL
Mesodon clausus (Say, 1821) Yellow Globelet CO WID Cal, MAR
Mesodon normalis (Pilsbry, 1900) Grand Globelet LR BR LL
Mesodon thyroidus (Say, 1816) White-lip Globe CO WID GEN
Mesodon zaleatus (A. Binney, 1837) Toothed Globe CO WID ROC
Patera perigrapta (Pilsbry, 1894) Engraved Bladetooth CO WID DWM, LL
Stenotrema pilula (Pilsbry, 1900) Pigmy Slitmouth LR BR DWM, LL S3, G3G4
Undescribed Stenotrema sp. unnamed LR BR GEN
Stenotrema stenotrema (Pfeiffer, 1842) Inland Slitmouth CO WID GEN
Triodopsis tridentata (Say, 1816) Northern Threetooth CO WID GEN
Xolotrema denotatum (Ferussac, 1821) Velvet Wedge CO WID DWM
Pomatiopsidae
Pomatiopsis lapidaria (Say, 1817) Slender Walker CO WID Cal, GEN
Punctidae
Punctum minutissimum (I. Lea, 1841) Small Spot CO WID LL X
Punctum vitreum (H.B. Baker, 1930) Glass Spot UN WID LL X
Pupillidae
Collumella simplex (Gould, 1841) unnamed CO WID LL X
Gastrocopta armifera (Say, 1821) Armed Snaggletooth CO WID Cal, MAR X
Gastrocopta contracta (Say, 1822) Bottleneck Snaggletooth CO WID GEN X
Gastrocopta corticaria (Say, 1816) Bark Snaggletooth CO WID Cal, DWM
Gastrocopta pentodon (Say, 1821) Comb Snaggletooth CO WID Cal, DWM X
Vetigo gouldii (A. Binney, 1843) Variable Vertigo CO NOR LL
Vertigo tridentata Wolf, 1870 Honey Vertigo CO WID LL X
Strobilopsidae
Strobilops aeneus Pilsbry, 1926 Bronze Pinecone CO WID DWM
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Farmily/species Common name Status Affinity Habitat County record State/Global Rank
Zonitidae
Glyphyalinia carolinensis (Cockrell, 1890) Spiral Mountain Glyph LR BR, CP LL
Glyphyalinia indentata (Say, 1823) Carved Glyph CO WID LL
Glyphyalinia pentadelphia (Pilsbry, 1900) Pink Glyph LR BR LL X S2, G2G3
Glyphyalinia wheatleyi (Bland, 1883) Bright Glyph CO WID LL
Hawaii miniscula (A. Binney, 1840) Minute Gem CO WID GEN X
Mesomphix andrewsae (Pilsbry, 1895) Mountain Button LR BR LL
Mesomphix cupreus (Rafinesque, 1831) Copper Button CO NOR LL
Mesomphix inornatus (Say, 1821) Plain Button CO NOR DWM, LL X
Mesomphix perlaevis (Pilsbry, 1900) Smooth Button CO WID LL
Paravitrea multidentata (A. Binney, 1840) Dentate Supercoil CO WID LL X
Paravitrea petrophila (Bland, 1883) Cherokee Supercoil LR CP, OA LL X
Paravitrea placentula (Shuttleworth, 1852) Glossy Supercoil LR BR, CP LL
Paravitrea umbilicaris (Ancey, 1887) Open Supercoil LR BR LL X S2, G2
Striatura meridionalis (Pilsbry and Ferriss, 1906) Median striate CO WID LL
Ventridens acerra (J. Lewis, 1870) Glossy Dome LR BR, CP LL
Ventridens collisella (Pilsbry, 1896) Sculptured Dome LR BR, CP, RV LL
Ventridens gularis (Say, 1822) Throaty Dome CO WID LL
Zonitoides arboreus (Say, 1816) Quick Gloss CO WID DWM
Zonitoides elliotti (Redfield, 1856) Green Dome LR BR, CP DWM, LL