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A Survey of Terrestrial Gastropods of the Sipsey Wilderness (Bankhead National Forest), Alabama
Janet Waggoner, Stephanie A. Clark, Kathryn E. Perez, and Charles Lydeard

Southeastern Naturalist, Volume 5, Number 1 (2006): 57–68

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2006 SOUTHEASTERN NATURALIST 5(1):57–68 A Survey of Terrestrial Gastropods of the Sipsey Wilderness (Bankhead National Forest), Alabama JANET WAGGONER1, STEPHANIE A. CLARK1,*, KATHRYN E. PEREZ1, AND CHARLES LYDEARD1 Abstract - A survey of the terrestrial mollusks of the Sipsey Wilderness Area, Bankhead National Forest, in northwestern Alabama was conducted from August 2003 to May 2004. A total of 15 sites were sampled across a number of different habitat and vegetation types found within the area. A total of 50 species were found, representing 14 families and 30 genera, including 58 new county and 2 new state records. This represents a significant increase in the known diversity of the area based on a preliminary survey conducted in the 1960s, which yielded only six species. The current survey highlights the need for more detailed survey work across Alabama and the southeastern United States. Introduction Terrestrial gastropods are a ubiquitous component of terrestrial ecosystems of the eastern United States. Distribution maps of the 523 native species and subspecies of terrestrial snails of the eastern United States were presented by Hubricht (1985). The distribution maps are based primarily upon the data presented in the taxonomic monographs published by Henry A. Pilsbry (1939, 1940, 1946, 1948) and the extensive material collected by Leslie Hubricht (deposited and available on-line at the Field Museum of Natural History, Chicago, IL). Alabama is considered one of the states in the eastern United States in which collecting effort has been most intensive (Archer 1939, Hubricht 1965). From 1903 to 1919, Herbert H. Smith, the first curator of the Alabama Museum of Natural History, collected land and freshwater mollusks throughout the state (Clapp 1920). Smith’s land-snail collection provided the basis for a monograph on the terrestrial shell-bearing mollusks of Alabama by Bryant Walker (1928). During the 1960s, Leslie Hubricht conducted numerous terrestrial gastropod surveys throughout the state. These collections provide a strong foundation and historical benchmark for our knowledge of the malacofauna of Alabama. Despite the pioneering efforts of these early naturalists, there is still a considerable amount to be learned about the distribution of the malacofauna of the state. For example, many early collection efforts were opportunistic or even superficial, and without extensive field notes, it is difficult to judge whether the absence of a species at a locale is due to it not being found at that locale or due to insufficient collecting methods. Historical museum records are critically important for evaluating and monitoring changes in species composition through time (Mikkelson and 1University of Alabama, Department of Biological Sciences, Biodiversity and Systematics, Box 870345, Tuscaloosa, AL 35487. *Corresponding author - saclark@bama.ua.edu. 58 Southeastern Naturalist Vol. 5, No. 1 Bieler 2001, Ponder et al. 2001); therefore it is essential that the baseline data fully represents the malacofauna of the region. In order to extend our knowledge of the malacofauna of Alabama, we conducted a survey of the terrestrial mollusks of the northern half of the Bankhead National Forest, concentrating on the Sipsey Wilderness Area. This area was chosen as it is relatively undisturbed compared to the southern half of the Bankhead National Forest and because L. Hubricht had collected in the area previously (= Site 1). We hoped a comparison with this earlier collection would give us an indication of the historical terrestrial gastropod diversity. Study Area The study area is located in the north western section of the William B. Bankhead National Forest (NF) and encompasses the Sipsey Wilderness Area of northern Alabama (Fig. 1). The Bankhead NF is approximately 182,000 acres and the Sipsey Wilderness Area is approximately 25,000 acres in area. The bulk of the NF lies in the Southwestern Appalachian Ecoregion with its northern borders crossing into the Interior Plateau Ecoregion (US Environmental Protection Agency 2005, Omernik 1987). The Southwestern Appalachian Ecoregion is comprised of temperate broadleaf and mixed forests that cover the plateaus and rolling hills west of the Appalachian Figure 1. Map showing the location of the William B. Bankhead National Forest in northern Alabama. The insert shows the location of the 15 sample sites within the Bankhead National Forest and the shaded area is the Sipsey Wilderness Area. 2006 J. Waggoner, S.A. Clark, K.E. Perez, and C. Lydeard 59 Mountains and is considered a “globally outstanding” terrestrial ecoregion by the World Wildlife Fund based on it harbouring some of the richest and most endemic land-snail, amphibian, and herbaceous plant biotas in the United States and Canada (Ricketts et al. 1999). We chose sites within the NF because, other than impacts from silvicultural practices and road building, it is largely a protected area. Materials and Methods A total of 15 randomly chosen sites were surveyed for terrestrial gastropods (Table 1; Fig. 1) between October 2003 and March 2004. At least one Table 1. Sampling sites located in the northern half of the Bankhead National Forest, including the Sipsey Wilderness Area. Site Location Latitude Longitude Date 1 E. side of Sipsey River; along walking track N. of CR 60, Bankhead NF, Winston Co. 34°17'12"N 87°23'56"W 11-Oct-03 2 Along an old track, about 0.5 miles N. of Tennessee Valley Divide Rd, 0.6 miles E.of Cheatham Rd, Bankhead NF, Lawrence Co. 34°23'32"N 87°18'27"W 12-Oct-03 3 Along banks of small trib. of Brushy Creek, ≈ 0.4 mi. E. of Bushy Creek Rd, opposite Pine Torch Church, Bankhead NF, Lawrence Co. 34°19'31"N 87°18'40"W 12-Oct-03 4 About 2.5 miles W. of CR 6, on CR 5, road to bow-hunting camp, Bankhead NF, Lawrence Co. 34°18'39"N 87°23'38"W 19-Oct-03 5 About 1 mile W. of CR 6 on CR 5, Bankhead NF., Lawrence Co. 34°18'51"N 87°22'21"W 19-Oct-03 6 Parker Branch at CR 60, 0.2 miles E. of junction with FR210 (= CR 60), Bankhead NF, Winston Co. 34°16'51"N 87°29'30"W 19-Oct-03 7 Off FR 210, 0.6 miles S. of junction with FR 203, Bankhead NF, Winston Co. 34°20'33"N 87°31'09"W 19-Oct-03 8 Northern side of FR 213, E. of FR 203, Bankhead NF, Winston Co. 34°23'29"N 87°28'16"W 19-Oct-03 9 Off western side of FR 208, W. of Flannagin Creek, Bankhead NF, Lawrence Co. 34°20'27"N 87°23'33"W 19-Oct-03 10 Bluff, above Borden Creek at FR 208, Bankhead NF, Lawrence Co. 34°19'48"N 87°22'39"W 19-Oct-03 11 About 0.2 miles along track S. of FR 203, about 1 mile W. of Tedford Creek, Bankhead NF, Lawrence Co. 34°20'05"N 87°29'01"W 21-Mar-04 12 Off FR 203, 0.8 miles W. of Tedford Creek, Bankhead NF, Lawrence Co. 34°20'23"N 87°29'05"W 21-Mar-04 13 Off FR 203, 0.5 miles W. of Tedford Creek, Bankhead NF, Lawrence Co. 34°20'21"N 87°28'44"W 21-Mar-04 14 Trailhead of Thompson Creek at end of FR 203, Bankhead NF, Lawrence Co. 34°20'28"N 87°28'14"W 21-Mar-04 15 Flannagin Creek at FR 208E, Bankhead NF, Lawrence Co. 34°20'20"N 87°23'17"W 21-Mar-04 60 Southeastern Naturalist Vol. 5, No. 1 Table 2. Rank order and frequency of summed specimens. Number of Percent of New county records New state Taxon Rank Samples specimens specimens Lawrence Winston records Mesomphix globosus (MacMillan, 1940) 1 12 109 7.42 X Gastrodonta interna (Say, 1822) 2 12 94 6.40 Haplotrema concavum (Say, 1821) 3 11 123 8.37 X Mesodon normalis (Pilsbry, 1900) 4 11 75 5.11 X Triodopsis tridentata (Say, 1816) 5 10 43 2.93 X X Glyphyalinia cryptomphala (Clapp, 1915) 6 10 33 2.25 X Patera perigrapta (Pilsbry, 1894) 7 9 64 4.36 X X Zonitoides arboreus (Say, 1816) 8 9 61 4.15 X Stenotrema barbigerum (Redfield, 1856) 9 9 43 2.93 Mesodon thyroidus (Say, 1816) 10 9 35 2.38 Ventridens pilsbryi Hubricht, 1964 11 9 30 2.04 Glyphyalinia indentata (Say, 1823) 12 9 23 1.57 Stenotrema stenotrema (Pfeiffer, 1842) 13 7 33 2.25 X X Carychium exile Lea, 1842 14 6 180 12.25 X X Glyphyalinia wheatleyi (Bland, 1883) 15 6 48 3.27 X Striatura meridonalis (Pilsbry & Ferriss, 1906) 16 6 31 2.11 X X Mesomphix latior (Pilsbry, 1900) 17 6 23 1.57 X X Euconulus dentatus (Sterki, 1893) 18 6 20 1.36 X X Mesomphix capnodes (Binney, 1857) 19 6 14 0.95 X X Gastrocopta contracta (Say, 1822) 20 5 38 2.59 X X Xolotrema obstrictum (Say, 1821) 21 5 32 2.18 X X Strobilops aeneus Pilsbry, 1926 22 5 18 1.23 X Philomycus carolinianus (Bosc, 1802) 23 5 11 0.75 X Inflectarius inflectus (Say, 1821) 24 4 43 2.93 Oligyra orbiculata Say, 1818 25 4 8 0.54 X 2006 J. Waggoner, S.A. Clark, K.E. Perez, and C. Lydeard 61 Table 2, continued. Number of Percent of New county records New state Taxon Rank Samples specimens specimens Lawrence Winston records Punctum minutissimum (Lea, 1841) 26 4 7 0.48 X X Glyphyalinia wetherbyi (Cockerell, 1900) 27 4 4 0.27 Millerelix plicata (Say, 1821) 28 3 45 3.06 X Triodopsis vulgata Pilsbry, 1940 29 3 20 1.36 X X Punctum vitreum (Baker, 1930) 30 3 11 0.75 X X Lobosculum pustuloides (Bland, 1858) 31 3 4 0.27 X Strobilops labyrinthica (Say, 1817) 32 3 3 0.20 X Carychium nannodes Clapp, 1905 33 2 75 5.11 X Cochlicopa morseana (Doherty, 1878) 34 2 22 1.50 X X Anguispira alternata (Say, 1816) 35 2 6 0.41 X Gastrocopta pentadon (Say, 1821) 36 2 5 0.34 X Glyphyalinia carolinensis (Cockerell, 1890) 37 2 4 0.27 X X Glyphyalinia praecox (Baker, 1930) 38 2 4 0.27 X Hawaiia minuscula (Binney, 1841) 39 2 3 0.20 X X Helicodiscus parallelus (Say, 1817) 40 2 3 0.20 X X Ventridens intertextus (Binney, 1841) 41 2 3 0.20 X X Paravitrea multidentata (Binney, 1840) 42 2 2 0.14 X Discus patulus (Deshayes, 1830) 43 1 5 0.34 X Ventridens demissus (Binney, 1843) 44 1 4 0.27 X Euchemotrema fasciatum (Pilsbry, 1940) 45 1 2 0.14 Vertigo gouldii (Binney, 1843) 46 1 1 0.07 X X Punctum smithi Morrison, 1935 47 1 1 0.07 X Mesomphix pilsbryi (Clapp, 1904) 48 1 1 0.07 Paravitrea capsella (Gould, 1805) 49 1 1 0.07 X Mesodon zaletus (Binney, 1837) 50 1 1 0.07 X Total 1469 100.00% 32 26 2 62 Southeastern Naturalist Vol. 5, No. 1 sample per major habitat type was collected from sites accessible from forestry roads and along walking trails. Sampling included hand searching for specimens on site on the ground, in the leaf litter, under bark, under logs, and under stones, and taking a 5-litre bag of leaf litter to be sorted in the lab. An average of about an hour was spent at each site. All shells were identified to species using the keys provided in: Burch 1962; Emberton 1988, 1991; Hubricht 1961, 1962a, 1962b, 1964, 1976, 1978; and Pilsbry 1939, 1940, 1946, 1948. All specimens have been deposited in the University of Alabama Scientific Collections Facility. The taxonomic nomenclature used follows Emberton (1988, 1991), Schileyko (2002, 2003), and Turgeon et al. (1998). Results and Discussion The 15 samples yielded a total of 1469 identifiable shells, representing 50 taxa (Appendix 1). A total of 58 new county records were recorded, with 26 for Winston and 32 for Lawrence counties, and 2 new state records were established (Table 2). While no species of the families Succineidae or Pomatiopsidae were found during the current survey, it is very likely that they are present as suitable habitat does occur within the study area. Therefore, as more extensive surveys of the area are completed, the total number of species inhabiting the area will likely be increased. The rank order was also calculated for each species for the entire study (Table 2). Mesomphix globosus, Gastrodonta interna, and Haplotrema concavum were ranked 1st, 2nd, and 3rd, respectively. The following five taxa are each represented by a single specimen and received the lowest rank: Vertigo gouldii, Punctum smithi, Mesomphix pilsbryi, Paravitrea capsella, and Mesodon zaletus. The molluscan assemblage of taxa found is typically associated with upland woodland areas, with deep layers of leaf litter and fallen timber on the ground (Burch, 1955, Hubricht 1985). Three species found during this survey are considered by Hubricht (1985) to be mainly associated with limestone outcrops or soils (calciphiles): Oligyra orbiculata , Lobosculum pustuloides, and Gastrocopta pentadon. As our specimens were found in woodland habitats, it suggests that the soils may have some limestone influences. A search of the on-line database of the Chicago Field Museum of Natural History (http://www.fmnh.org) showed that L. Hubricht collected in the study area in 1961. His collection effort yielded only 6 species: Glyphyalinia cryptomphala, Glyphyalinia indentata, Haplotrema concavum, Stenotrema barbigerum, Mesodon thyroidus , and Mesodon normalis. Our examination of the same general area yielded 24 species. As little information is available about the collection effort employed or the condition of the site at the time of Hubricht's visit, the difference in the number of species found nonetheless is significant. Therefore, investigators should exercise care in interpreting historical data (e.g., absence of a species may not necessarily equal true absence). 2006 J. Waggoner, S.A. Clark, K.E. Perez, and C. Lydeard 63 Our survey clearly indicates that there is a tremendous amount of information to be gleaned from thorough biotic surveys and inventories of terrestrial gastropods in the region. Due to their limited dispersal capabilities, terrestrial gastropod species can make excellent study organisms for evolutionary (e.g., Hugall et al. 2003) and ecological studies (Coney et al. 1982, Cook 2001, Heller 2001, Neck 1990, Nekola and Smith 1999), as well as inferring paleoclimates and habitats (Theler et al. 2004). In addition, most species of terrestrial mollusks are restricted to particular vegetation and soil types (Burch 1955, Clark 2004, Hotopp 2002, Riggle 1976), and changes in species composition can be used to gauge impacts of both human and non-human factors effecting the environment. From a conservation stand-point, better informed assessments of environmental impacts of various human-related activities or affects of climate change can be assessed when sufficient baseline data exists. Although terrestrial gastropod species are typically not considered keystone species for most ecosystems, the decline in their abundance has been linked to decline in song birds (Graveland et al. 1994). In addition to biotic surveys and inventories, most terrestrial gastropod taxa have not been thoroughly treated taxonomically and systematically. Since Pilsbry’s classic work (1939, 1940, 1946, 1948), there has been no thorough systematic treatment of the United States land snail fauna; however, there have been smaller revisions, a few state field guides, and checklists (e.g., Bequaert and Miller 1973; Emberton, 1988, 1991; Metcalf and Smartt 1997; Roth and Sadeghian 2003). Acknowledgments We are grateful to Alison Cochran and Tommy Counts of Bankhead National Forest for their encouragement and support. Thanks to Patricia West, Andrew, Emily, and Catherine Lydeard for assistance in the field. Thanks to John Slapcinsky of the Florida Museum of Natural History, who confirmed identifications and provided hospitality during two recent visits by S. Clark to Gainesville. This project was supported in-part by a small contract from the US Forest Service to C. Lydeard, the University of Alabama’s Department of Biological Sciences and Centre for Freshwater Studies. Kathryn Perez was supported by a NSF IGERT fellowship (DGE- 9972810 to Dr. Amelia Ward) and S. Clark was supported by the University of Alabama Office of Sponsored Programs and NSF (DEB-0128964 to C. Lydeard). Literature Cited Archer, A.F. 1939. The distribution of land mollusks of Alabama from their probable centers of origin. The Nautilus 52(4):112–115. Bequaert, J.C., and W.B. Miller. 1973. The mollusks of the arid southwest, with an Arizona checklist. The University of Arizona Press, Tucson, AZ. Burch, J.B. 1955. Some ecological factors of the soil affecting the distribution and abundance of land snails in eastern Virginia. The Nautilus 69(2):62–69. Burch, J.B. 1962. How to Know the Eastern Land Snails. Wm. C. Brown, Dubuque, IA. 214 pp. 64 Southeastern Naturalist Vol. 5, No. 1 Clapp, G.H. 1920. Herbert Huntington Smith. The Nautilus 33(4):136–141. Clark, S.A. 2004. Native snails in an urban environment: Conservation from the ground up. Pp. 78–81, In D. Lunney and S. Burgin (Eds.). Urban Wildlife: More Than Meets the Eye. Royal Zoological Society of New South Wales, Mosman, Australia. Coney, C.C., W.A. Tarpey, J.C. Warden, and J.W. Nagel. 1982. Ecological studies of land snails in the Hiwassee River basin of Tennessee, USA. Malacological Review 15:69–106. Cook, A. 2001. Behavioural Ecology: On doing the right thing, in the right place, at the right time. Pp. 447–488, In G.M. Barker (Ed.). The Biology of Terrestrial Mollusks. CAB International, Oxon, UK. Emberton, K.C. 1988. The genitalic, allozymic, and conchological evolution of the eastern North American Triodiopsinae (Gastropoda: Pulmonata: Polygyridae). Malacologia 28(1–2):159–273. Emberton, K.C. 1991. The genitalic, allozymic, and conchological evolution of the Tribe Mesodontini (Pulmonata: Stylommatophora: Polygyridae). Malacologia 33(1–2):71–178. Graveland, J., R. van der Wal, J.H. van Balen, and A.J. van Noordwijk. 1994. Poor reproduction in forest passerines from decline of snail abundance on acidified soils. Nature 368:446–448. Heller, J. 2001. Life history strategies. Pp. 413–446, In G.M. Barker (Ed.). The Biology of Terrestrial Mollusks. CAB International, Oxon, UK. Hotopp, K.P. 2002. Land snails and soil calcium in Central Appalachian mountain forest. Southeastern Naturalist 1:27–44. Hubricht, L. 1961. Eight new species of land snails from the southern United States. Nautilus 75(1):26–33, plate 4. Hubricht, L. 1962a. New species of Helicodiscus from the eastern United States. The Nautilus 75(3):102–107, plates 7–9. Hubricht, L. 1962b. Mesomphix vulgatus and its allies. The Nautilus 76(1):1–7, plate 1. Hubricht, L. 1964. The bidentate species of Ventridens (Stylommatophora: Zonitidae). Malacologia 1(3):417–426. Hubricht, L. 1965. The land snails of Alabama. Sterkiana 17:1–5. Hubricht, L. 1976. Five new species of land snails from the eastern United States. Malacological Review 9:126–130. Hubricht, L. 1978. Thirteen new species of land snails from the southeastern United States with notes on other species. Malacological Review 10:37–52. Hubricht, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana, Zoology 24:1–191. Hugall, A., J. Stanisic, and C. Moritz. 2003. Phylogeography of terrestrial gastropods: The case of the Sphaerospira lineage and history of Queensland Rainforests. Pp. 270–301, In C. Lydeard and D.R. Lindberg (Eds.). Molecular Systematics and Phylogeography of Mollusks. Smithsonian Books, Washington, DC. Metcalf, A.L., and R.A. Smartt. 1997. Land snails of New Mexico: A systematic review. Bulletin of the New Mexico Museum of Natural History and Science 10:1–69. Mikkelson, P.M., and R. Bieler. 2001. Marine bivalves of the Florida Keys: Discovered biodiversity. Geological Society of London, Special Publication 177:247–257. Neck, R.W. 1990. Ecological analysis of the living mollusks of the Texas panhandle. American Malacological Bulletin 8:9–18. 2006 J. Waggoner, S.A. Clark, K.E. Perez, and C. Lydeard 65 Nekola, J.C., and T.M. Smith. 1999. Terrestrial gastropod richness patterns in Wisconsin carbonate cliff communities. Malacologia 41(1):253–269. Omernik, J.M. 1987. Ecoregions of the conterminous United States. Map (scale 1:7,500,000). Annals of the Association of American Geographers 77(1):118–125. Pilsbry, H.A. 1939. 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Riggle, R.S. 1976. Quantitative examination of gastropod and soil relationships in oak-hickory forest in the lower Illinois Valley region. Sterkiana 62:1–17. Roth, B., and P.S. Sadeghian. 2003. Checklist of the land snails and slugs of California. Santa Barbara Museum of Natural History, Santa Barbara, CA. Contributions in Science Number 3. 81 pp. Schileyko, A.A. 2002. Treatise on recent terrestrial pulmonate molluscs. Part 8. Punctidae, Helicodiscidae, Discidae, Cystopeltidae, Euconulidae, Trochomorphidae. Ruthenica, Supplement 2:1035–1166. Schileyko, A.A. 2003. Treatise on recent terrestrial pulmonate molluscs. Part 10. Ariophantidae, Ostracolethidae, Ryssotidae, Milacidae, Dyakiidae, Staffordiidae, Gastrodontidae, Zonitidae, Daudebardiidae, Parmacellidae. Ruthenica, Supplement 2:1309–1466. Theler, J. L. D.G. Wyckoff, and B.J. Carter. 2004. The Southern Plains Gastropod Survey: The distribution of land snail populations in an American grassland environment. American Malacological Bulletin 18:1–21. Turgeon, D.D., J.F. Quinn, A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 16:1–536. US Environmental Protection Agency. 2005. Level III Ecoregions. (Based on Omernik 1987). Available at: http://www.epa.gov/wed/pages/ecoregions/ level_iii.htm. Walker, B. 1928. The terrestrial shell-bearing Mollusca of Alabama. Miscellaneous Publications of the Museum of Zoology, University of Michigan 18:1–180. 66 Southeastern Naturalist Vol. 5, No. 1 Appendix 1. List of species and their locations. Sites Species 1 2 3456789101112131415 Helicinidae Oligyra orbiculata Say, 1818 X X X X Ellobiidae Carychium exile Lea, 1842 X X X X X Carychium nannodes Clapp, 1905 XX Cochlicopidae Cochlicopa morseana (Doherty, 1878) X X Pupillidae Gastrocopta contracta (Say, 1822) X X X X X Gastrocopta pentodon (Say, 1821) X X Vertigo gouldii (Binney, 1843) X Strobilopsidae Strobilops aeneus Pilsbry, 1926 X X X X X Strobilops labyrinthica (Say, 1817) X X X Philomycidae Philomycus carolinianus (Bosc, 1802) X X X X X Patulidae Anguispira alternata (Say, 1816) XX Discus patulus (Deshayes, 1830) X Helicodiscidae Helicodiscus parallelus (Say, 1817) X X Punctidae Punctum minutissimum (Lea, 1841) X X X X Punctum smithi Morrison, 1935 X Punctum vitreum (Baker, 1930) X X X Gastrodontidae Gastrodonta interna (Say, 1822) X XXXXXX XXXXX Striatura meridonalis (Pilsbry & Ferriss, 1906) X X X X X X 2006 J. Waggoner, S.A. Clark, K.E. Perez, and C. Lydeard 67 Sites Species 1 23456789101112131415 Gastrodontidae Ventridens intertextus (Binney, 1841) X X Ventridens demissus (Binney, 1843) X Ventridens pilsbryi Hubricht, 1964 X X X X X X X X X Zonitoides arboreus (Say, 1816) X X X X X X X X X Euconulidae Euconulus dentatus (Sterki, 1893) X X X X X X Haplotrematidae Haplotrema concavum (Say, 1821) X X XXXX X XXXX Zonitidae Glyphyalinia carolinensis (Cockerell, 1890) X X Glyphyalinia cryptomphala (Clapp, 1915) X X X X X X X X X X Glyphyalinia indentata (Say, 1823) X X X X X X X X X Glyphyalinia praecox (Baker, 1930) XX Glyphyalinia wetherbyi (Cockerell, 1900) X X X X Glyphyalinia wheatleyi (Bland, 1883) X X X X X X Hawaiia minuscula (Binney, 1841) X X Mesomphix capnodes (Binney, 1857) X X X X X X Mesomphix globosus (MacMillan, 1940) X X XXXXX XXXXX Mesomphix latior (Pilsbry, 1900) X X X X X X Mesomphix pilsbryi (Clapp, 1904) X Paravitrea capsella (Gould, 1805) X Paravitrea multidentata (A. Binney, 1840) XX Polygyridae Euchemotrema fasciatum (Pilsbry, 1940) X Inflectarius inflectus (Say, 1821) X X X X Lobosculum pustuloides (Bland, 1858) X X X Mesodon normalis (Pilsbry, 1900) X X X X X X X X X X X Mesodon thyroidus (Say, 1816) X X X X X X X X X Mesodon zaletus (A. Binney, 1837) X 68 Southeastern Naturalist Vol. 5, No. 1 Sites Species 1 23456789101112131415 Polygyridae Millerelix plicata (Say, 1821) X X X Patera perigrapta (Pilsbry, 1894) X X X X X X X X X Stenotrema barbigerum (Redfield, 1856) X X X X X X X X X Stenotrema stenotrema (Pfeiffer, 1842) X X X X X X X Triodopsis tridentata (Say, 1816) XXXXX XXXXX Triodopsis vulgata Pilsbry, 1940 X X X Xolotrema obstrictum (Say, 1821) X X X X X