Southeastern Naturalist D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 166 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 Southeastern Naturalist 167 D.A. Douglas, D.C. Dourson, and R.S. Caldwell 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 Southeastern Naturalist D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 168 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 Naturalist 169 D.A. Douglas, D.C. Dourson, and R.S. Caldwell 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 Southeastern Naturalist D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 170 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. Literature Cited Bellard, C., C. Bertelsmeier, P. Leadley, W. Thuiller, and F. Courchamp. 2012. Impacts of climate change on the future of biodiversity. Ecology Letters 15:365–377. Burch, J.B. 1962. How to Know the Eastern Land Snail. 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Bethesda, MD. 526 pp. United States Geological Survey (USGS). 1968. Geologic map and structure sections of the Great Smoky Mountains National Park and Vicinity, Tennessee and North Carolina. Washington, DC. (map scale: 1:125,000). Weigand, A.M., A. Jochum, M. Pfenninger, D. Steinke, and A. Klussmann-Kolb. 2011. A new approach to an old conundrum—DNA barcoding sheds new light on phenotypic plasticity and morphological stasis in microsnails (Gastropoda, Pulmonata, Carychiidae). Molecular Ecology Resources 11:255–265. Withers, D.I. 2009. Tennessee natural heritage program guide to the rare animals of Tennessee. Tennessee Department of Environment and Conservation, Division of Natural Areas, Nashville, TN. 72 pp. Southeastern Naturalist 173 D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 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 Southeastern Naturalist D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 174 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 Southeastern Naturalist 175 D.A. Douglas, D.C. Dourson, and R.S. Caldwell 2014 Vol. 13, No. 1 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