An Inventory of the Land Snails and Slugs (Gastropoda: Caenogastropoda and Pulmonata) of Knox County, Tennessee Author(s): Barbara J. Dinkins and Gerald R. Dinkins Source: American Malacological Bulletin, 36(1):1-22. Published By: American Malacological Society https://doi.org/10.4003/006.036.0101 URL: http://www.bioone.org/doi/full/10.4003/006.036.0101

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Amer. Malac. Bull. 36(1): 1–22 (2018)

An Inventory of the Land Snails and Slugs (Gastropoda: Caenogastropoda and Pulmonata) of Knox County, Tennessee

Barbara J. Dinkins1 and Gerald R. Dinkins2

1Dinkins Biological Consulting, LLC, P O Box 1851, Powell, Tennessee 37849, U.S.A [email protected] 2McClung Museum of Natural History and Culture, 1327 Circle Park Drive, Knoxville, Tennessee 37916, U.S.A.

Abstract: Terrestrial mollusks (land snails and slugs) are an important component of the terrestrial ecosystem, yet for most species their distribution is not well known. This study was a comprehensive inventory of terrestrial mollusks in Knox County, Tennessee; an area within the Valley and Ridge physiographic region rich in calcareous soils and bluffs. Knox County supports a total of 151 species, including 70 newly reported to the county and 15 newly reported to Tennessee. Sixteen species non-native to North America were found, mostly in urban/ residential habitats. Limestone/sandstone bluffs were found to have the highest diversity of terrestrial mollusks.

Key words: Terrestrial Gastropoda, Knox County, Tennessee, Ridge and Valley physiographic region

Land snails and slugs (Gastropoda: Caenogastropopoda provided distributional information on Pupillidae in North and Pulmonata) are a vital component of the terrestrial ecosys- America, and for Knox County they reported 7 species. Except tem. As a group, they serve many roles: they can be detritivo- for Lutz (1950), our survey is the fi rst inventory of terrestrial rous, herbivorous, or predaceous and be an important source mollusks in Tennessee that focused on an individual county. of calcium and food for other animal groups, including insects, Herein, we use the term “land snail” for all species of terrestrial birds, reptiles and amphibians, and small mammals (Ingram mollusks, i.e., including slugs. 1950, Martin et al. 1951, Speiser 2001, Allen 2004, Symondson 2004). In the Netherlands, passerine birds had signifi cantly lower eggshell quality in areas of low snail density (Graveland MATERIALS AND METHODS 1996). Some salamander species rely on land snails as a calcium source and their density is linked to the density of land snails Study area (Harper and Guyen 1999). Empty land snail shells are often Knox County is situated in the eastern third of Tennessee used as shelter by other invertebrates (Taylor et al. 1977, and is entirely within the Southern Appalachian Valley and Dourson 2010, pers. obs.). Snails are present and often abun- Ridge physiographic region, a 60–90 km wide belt of northeast- dant in most terrestrial habitats in eastern U.S.A. and the southwest trending ridges and valleys extending from Maryland Southeast supports an extraordinarily high diversity, as shown south to Georgia and Alabama. Knox County is situated by Coney et al. (1982), Hubricht (1985), Dourson and Freeman approximately midway between the Kentucky and Georgia bor- (2006), Dourson (2007, 2010, 2013), Caldwell and Dourson ders and has a total area of 1,360 km2, of which 1317 km2 is land (2008), and Douglas et al. (2010, 2014). and 43 km2 is water. Knox County’s land area is 54% tree can- Specimens of terrestrial mollusks from Knox County, opy, 28% open space, 13% impervious surface, and 5% bare Tennessee have been archived in several institutions, and were ground (American Forests 2002). The average annual tempera- collected as early as the 1800’s by a number of notable research- ture and precipitation for Knoxville (Knox County’s largest ers, including Mrs. George Andrews, George H. Clapp, Leslie city), is 15.3° C and 122.4 cm, respectively (Your Weather Service Hubricht, Arnold Ortmann, Henry Pilsbry, Victor Sterki, and 2017). Signifi cant land features in Knox County include House Bryant Walker. Several publications have dealt with the occur- Mountain (the county’s highest point located in its northeast rence of land snails in Tennessee. Lutz (1950) surveyed corner), Bays Mountains complex, McAnnally Ridge, Beaver Claiborne County and reported 20 species. Bogan et al. (1982) Ridge, Copper Ridge, Bullrun Ridge and several large cliffs bor- reported on the distribution of Polygyridae in Tennessee and dering the Holston, French Broad, and Tennessee Rivers. reported 75 species. Hubricht (1985) presented distributional information on land snails in eastern U.S.A. counties and pro- Sampling sites vided a map for each species. For Knox County, he reported After reviewing USGS topographic maps and the soil 76 species. Hubricht’s maps were a compilation of published survey of Knox County (Roberts et al. 1955), and using our sources, and combined his collecting efforts with specimens col- knowledge of the study area, we chose 116 sites to sample for lected by others but verifi ed by him. Nekola and Coles (2010) land snails (Fig. 1). Fieldwork was conducted in 2015-2017. 1 2 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018

On each of the different signifi cant land features we chose human-made debris, and wetland edges. A hand-held Global sites within various habitats and slope aspects, including wet- Positioning System device was used to record the coordinates lands, fl oodplains, rock bluffs and forested slopes. Although for the geographic center of each site. Sites ranged from 400 we concentrated our survey efforts on limestone-rich public to 1600 m2. Each site was characterized using a generalized areas and forested parks, we included a number of private list of habitat categories: Seep/wetland/fl oodplain, pine forest/ properties and urbanized settings (e.g., abandoned and occu- cedar grove, open disturbed, mature oak/hickory, sandstone/ pied parcels in downtown Knoxville) to document the diver- limestone outcrops and river bluff, urban/residential, cave, dis- sity of land snails across a wide range of habitats. turbed second growth forest, and disturbed sandstone/limestone outcrop (Appendix 1). Soil samples returned to the laboratory Sampling methods and processing were dried and sifted through a series of four sieves with open- At each site, one to three person-hours were spent assessing ings measuring 4.75 mm to 0.50 mm. The larger factions of the habitat and searching for medium to large land snails; several soil and fi ne woody debris samples were examined using a 10X liters of soil and fi ne woody material were gathered at various lens and the smaller factions were examined for the presence of places across the site, composited into a single sample, and microsnails (<5mm at the greatest dimension) using a dissecting returned to the laboratory. Fine-scale habitats examined at most microscope. Land snails encountered opportunistically (i.e., not survey sites included large tree trunks, the underside, surface and associated with a survey site) were included in the county species interior of coarse woody debris, large rocks and rock outcrops, list. At some sites, snails were encountered that were identifi ed to

Figure 1. Map of Knox County survey sites. LAND SNAILS OF KNOX COUNTY, TENNESSEE 3 the genus level only; identifi cation to the species level could not in CMNH and/or FLMNH. We document 70 species previ- be reached because of shell condition or age. These were included ously unreported in Knox County. Most of these new county in the species total for the site if they represented the only mem- records are common elsewhere in east Tennessee (e.g., 51 of ber of the genus present. Specimens collected in this study were these have been recorded in at least one adjacent county). archived at the University of Tennessee, McClung Museum of Our list of new county records includes three species we Natural History and Culture. found live or fresh dead but which previously were known We used a number of references to identify specimens, from Tennessee only in the fossil record (Hubricht 1985): notably Pilsbry (1940, 1946, 1948), Burch (1962), Kerney and Gastrocopta holzingeri (Sterki, 1889), Cochlicopa lubrica Cameron (1979), Emberton (1988, 1991), Grimm et al. (2009), (Müller, 1774), Lucilla singleyanus (Pilsbry, 1889). It should Dourson (2010, 2013), and Nekola and Coles (2010). We be noted two of our new county records that are based on compared problematic specimens to material at the Chicago museum specimens: Stenotrema edgarianum (I. Lea, 1841) Field Museum (FMNH), Carnegie Museum of Natural and Ventridens coelaxis (Pilsbry, 1899) may be questionable History (CMNH), the Ohio State Museum of Biological based on imprecise locality data, published distributional Diversity (OSUM), University of Michigan Museum of information, and habitat preference. This is discussed in the Zoology (UMMZ), and the McClung Museum of Natural species accounts. Fifteen species are new state records; all are History and Culture (MMNHC). Additionally, selected prob- common in the southeastern U.S.A. and have been reported lematic specimens were sent for verifi cation to Dan Dourson from a state adjacent to Tennessee. Eleven of the state records (Belize Foundation for Research and Environmental Education), are species not native to North America. John Slapcinsky (Florida Museum of Natural History Land snails are widespread in Knox County, and were [FLMNH]), Amy and Wayne Van Devender (Appalachian found at all 116 sites. The three most commonly encountered State University), and Francisco Borrero (Animal and Plant species were Striatura meridionalis (Pilsbry and Ferriss, 1906), Health Inspection Service [APHIS]). County and state records Infl ectarius infl ectus (Say, 1821), and Zonitoides arboreus (Say, were determined using Bogan et al. (1982), Hubricht (1985), 1816), occurring at 57%, 56%, and 54% of all sites, respec- Dourson (2013), and pertinent literature. We searched the tively. Twenty-six (26) species were found at only a single site. database of several institutions for Knox County records. The most speciose site was a limestone/sandstone bluff com- Only published or verifi ed records were considered for the plex on Texas Valley Road in the northeast corner of the list of Knox county species. Some species found in our fi eld county (Site 9, N = 45 species). In general, limestone/sand- survey representing a potentially new county record also were stone bluff habitat was the most speciose. found in one or more museum databases. Due to time con- Including museum records, we found 16 nonindigenous siderations, we did not examine these specimens or ask the species. Except for Arion subfuscus (Draparnaud, 1805) and museum curator for verifi cation. Species in a museum data- Vertigo pygmaea (Draparnaud, 1801), all non-native species base not found in our fi eld survey representing a potential were found exclusively in urban, residential or otherwise new county record were examined by us or verifi ed by the anthropogenically disturbed sites. collection manager. We followed Turgeon et al. (1998) for genus and species names with three exceptions: We followed Taxonomic Notes on Selected Taxa Gargonimy (2011) for Ambigolimax valentianus, Roth (2003) Allopeas cf. clavulinum (Potiez and Michaud, 1838): Allopeas for Cochlicopa, and Coles and Walsh (2006) for Daedalochila. clavulinum is an introduced snail documented in Pennsylvania, For family designations we followed MolluscaBase (2018) Massachusetts, and Florida (Pilsbry 1946, Dundee 1971). with one exception. Following Pokryszko et al. 2009, we Several live immature specimens were found in a degraded retained Columella and Gastrocopta in Vertiginidae as this roadside area in Knoxville (Site 65) and were verifi ed by group is presently under study (Nekola and Coles 2016). F. Borrero as generally matching A. clavulinum. Anguispira Morse 1864 spp.: Hubricht (1985) reported four species of Anguispira from Knox County: Anguispira RESULTS alternata (Say, 1816), Anguispira mordax Kutchka, 1938, Anguispira knoxensis (Pilsbry, 1899), and Anguispira jessica Approximately 10,630 specimens were collected in this Kutchka, 1938. We did not fi nd A. mordax or A. knoxensis in survey. Our survey results combined with museum speci- this survey. Individuals resembling A. strongylodes/jessica mens and published literature revealed the land snail fauna in were sent to J. Slapcinsky (FLMNH) for identifi cation. He Knox County comprises 151 species in 60 genera (Table 1). separated them into two species we tentatively refer to as Our list for Knox County includes eight species not found in Anguispira cf. strongylodes (Pheiffer, 1854) and Anguispira cf. this survey but reported by Hubricht (1985) and/or Bogan jessica. Future studies are needed to reveal the taxonomic et al. (1982), and eight species not found by us but cataloged placement of these taxa. We found A. cf. strongylodes at eight 4 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 DO,DF,MF,MX,OD,PC,SL,UR,WT DF,DO,MF,PC,SL,WT Habitat Codes CV,DO,MF,PC,SL,WT, DO,MF,MX,OD,SL,UR,WT catalogue numbers). Site numbers are d river bluffs, UR = Urban/residential, unty record or the museum curating andstone/limestone outcrops, MF = Mature = State record. 2 = County record; 1 112,113,115,116 55–57,62,65,66,68,74,75,77–84,86,89,94,96,99,102–104,109, 79,81–83,86–88,92,93,99,101–104,109,116 57–59,77, Site Numbers, Reference or Museum Collection 38213,13,2751P621,38,631,3,4,7,9,22,24,29–31,35–38,43,45,50–53,56,58,61,63,64,66, 68–70,74–77,79,91,93,99,102–104,109,113,116 3411,15,64,86 MX WT OD MF CM 62.398949,103 UR MX,OD,UR 10,11,86,891–4,9,10,15,16,18,22–27,29,30,35,37,42–52,59,60,73,77,79, 81–83,86,89,91–95,99,100,102,104,106,109,113,115,116 SL MX,WT Unknown MF,SL WT 1 1,2 1,2 1 1 1,2 1 1 1 1,2 1,2 1 (Pheiffer, 1854) 1,4,37,101,103,109,112,113 MF,SL (C.B. Adams, 1842) (Sterki, 1889) (Pilsbry, 1924) (Say, 1816) 3,4,9,24,33–35,38,39,46,47,55,57,58,75,77–82,86,93,94 DO,MF,MX,OD,PC,SL,WT (Say, 1822) 1,4,7,8,9,11,14,15,21,22,24,29,31,33–35,37,38,45–47,48,50,52, (Say, 1822) ( Say, 1821) 20,35,52,53,56,61,63,66,65,69,73,76–78,84,85 DO,OD,PC,SL,WT (Pilsbry, 1899) Hubricht (1985) Unknown Kutchka, 1938 99 MF (Porro, 1838)* 21,53,66,69 MX,OD,UR oodplains. Species non-native to North American are indicated by an asterisk (*). (Doherty, 1878) 1,4,7,9,26,45,50,55,57,59,99,101,103,104,115 MF,SL,WT (Ferussac, 1821)* (Say, 1822) (Say, 1816) 3-6,8–10,29,36,44,45,51,86,103,116 MF,SL,WT (Gould, 1840) 31,52,53,56,64,65,69,76,77,78 DO,OD,UR,WT (Shuttleworth, 1952) Hubricht (1985) Unknown Normand, 1852* (Sterki, 1909) 64,78,82,84,85 MF,MX,OD,PC (Muller, 1774)* Hubricht, 1959 4,8,9,10,18,45 PC,MF,SL,WT I. Lea, 1842 Férussac, 1816* (Draparnaud, 1805)* (Nilsson, 1923)* jessica strongylodes (Deshayes, 1830) 1,3,5,8,9,17,19,22,24,26,29,32,37,39,44–46,50,51,55, (Harper, 1881) cf. cf. List of land snail taxa Knox County, Tennessee. For those species not found at any study sites, the reference to its Co Gastrocopta procera Gastrocopta tappaniana Carychium clappi Carychium exiguum Gastrocopta contracta Anguispira knoxensis Anguispira mordax Arion hortensis Arion intermedius Arion subfuscus Bradybaena similaris Cochlicopa lubrica Cochlicopa lubricella Cochlicopa morseana Gastrocopta armifera Gastrocopta clappi Gastrocopta pentodon Anguispira alternata Anguispira Anguispira Discus bryanti Discus nigrimontanus Discus patulus Carychium exile Arion fasciatus Gastrocopta holzingeri Gastrocopta corticaria

Ellobiidae

Camaenidae Cionellidae Gastrocoptidae Discidae record is provided (CM and FLMNH refer respectively to Carnegie Museum of Natural History Florida Table 1. Taxon described in Appendix 1. Habitat designations are as follows: CV = Caves, DF Disturbed second growth forest, DO s hardwood forest, MX = Mixed habitats, OD Open disturbed, PC Pine forest/cedar groves, SL Sandstone//limestone outcrops an WT = Seeps/wetlands/fl Arionidae

LAND SNAILS OF KNOX COUNTY, TENNESSEE 5 DF,DO,MF,OD,PC,SL,WT DF,DO,MF,MX,OD,PC,SL,WT WT,PC,OD,MF,SL,DF,DO,MX Habitat Codes DF,DO,MF,MX,OD,SL,UR,WT DF,DO,MF,OD,PC,SL,WT DF,DO,MF,MX,OD,SL,WT 101–105,109,112–116 89,91,93–95,99,101–105,109,112,113,115 103,109,111 96,102, 74–77,79,81,93,99–101,103,116 Site Numbers, Reference or Museum Collection 23393WT 26,35,51,77,79 55,57–59,70–76,78,81,87,89,90,91,95,96,99,103,115 35 CV 1,4,5,7–9,24,25,29,37,42,44,45,50,91,93,99,103,104,114–116CM 66340 DF 62,68,70,72,73,75,77–84,87–89,91,93–96,99, 3,4,7-9,11,16,18,19,22,24,27,28–30,34–38,43,45–51, 53,55,57,60, 91,116 MF,SL CM 1110953,6,8,13–15,21,25–29,31,33,38–42,52,54,63,64, 66,67,69,79,100 DF,DO,MF,MX,OD,SL,UR,WT 30,38,41,51,54–,58,59,61,63–65,70,72,73,75–79,81–83,86–90, DO,MF,SL,WT 92,96,97,106,113 SL Unknown Unknown MF 1 1 1 1 1 1 1 1 1 1 (G.H. Clapp, 1915) 9,10,23–25,35,37,47,50,79,81–85,91,93,94,109,114–116 CV,DO,MF,PC,SL,WT nseque, 1831) nseque, 45,50,55,77,81,82,85,91,93,94,115,116 DO.MF,PC,SL (Say, 1823) 1,3,5–7,9,13–15,18–21,24,26,27,29,35,39,42–45,50–53,55,58,59, (Bland, 1883) 1,2,4–10,16,17,19,22,24–26,30,33,35,38,39,45,47,50,51, (G.H. Clapp, 1908) 34,35,109,112 SL

(A. Binney, 1841) 17,59 MF (Pilsbry, 1900) Clapp, 1905 8–10,12,23,24,26,45,47,81,93,99,102,103 CV,MF,SL,WT (W.G. Binney, 1857) (H.B. Baker, 1930) 1,35,45,47,55,59,70,80 MF,PC,SL (Phillips, 1941) 4,70,71,72,99,105 MF,SL Hubricht, 1966 1,13,38,40,52,102 MF,MX,WT (Pilsbry, 1899) 59,99 MF Hubricht, 1968 (Say, 1822) Hubricht (1985) Unknown (Gould, 1841) (A. Binney, 1843) (Reinhardt, 1883) 3,12,24,32,35,37,47,71,74,76,77,81,95,99,104,105,109,116 DO,MF,SL,WT (Say, 1821) 38,64,75,77 DO,MX,SL (Rafi (Pilsbry, 1896) 1,4,12,17,23,33,37,45,49,66,77–79,81–83,85,93,101,103,109,111,112 SL,CV,DF,DO,MF,OD,PC (Pilsbry, 1899) Call, 1897 (Sterki, 1893) 8,9,17,22,24,25,35,36,49,55,78,83,91,94,99,102,103,105,109,112 DO,MF,OD,SL,WT (H.B. Baker, 1930) 4,7,24,112,114 SL 1,2 (Say, 1822) 15,17,18,24,34,36,38,43,51,56–58,66,71,78–83,88,89,94, (J. Lewis, 1870) (Müller, 1774) (W.G. Binney, 1892) 5,9,24,94,101,104,114–116, DO,MF,SL (Say, 1821) F. G. Thompson and (Dall, 1888) 1,2,4,5,7–9,12,16–19,24–26,29,35–37,39,45–50,59,77–79,81,82, (Continued) Ventridens lasmodon Gastrodonta interna Glyphyalinia cryptomphala Glyphyalinia indentata Ventridens intertextus Ventridens lawae Dryachloa dauca Euconulus chersinus Euconulus dentatus Euconulus trochulus Guppya sterkii Glyphyalinia solida Glyphyalinia wheatleyi Mesomphix cupreus Mesomphix perlaevis Nesovitrea electrina Striatura meridionalis Ventridens coelaxis Ventridens collisella Ventridens demissus Ventridens gularis Euconulus fulvus Ventridens acerra Ventridens ligera Carychium stygium Carychium nannodes Glyphyalinia luticola Glyphyalinia praecox Glyphyalinia rhoadsi Glyphyalinia rimula Glyphyalinia lewisiana Mesomphix capnodes Table 1.

Gastrodontidae Euconulidae H. G. Lee, 1960 (Pilsbry and Ferriss, 1906)

Taxon

6 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 DF,DO,MF,OD,SL,UR,MX,PC,WT CV,DF,DO,MF,PC,SL,WT Habitat Codes DF,MF 94,96, 99,101–105,109,111 55–57,59,70,73–75,81–85,87,88,91,93, 99, 55–57,59,70,73–75,81–85,87,88,91,93, 101–05,107,109,113–116 44,45,47,51,52,54–59,61,63–67,70,74, 76,79,81,86–88,93, 44,45,47,51,52,54–59,61,63–67,70,74, 21,828,25,26,83,85,93 MF,OD MF,PC,SL,WT Site Numbers, Reference or Museum Collection 38,507,8,31,32CM 62.39896P41,7,32,104,109344,9,35,52,65,66,74,91,99,104,105,109,113 Unknown 21,31,69 MF,SL MF,MX 21,63P2,P321,63,64,69, MF,SL 4,5,8–10,115 MF,SL 38,395, 91P5,9,17 CV 17,39 SL MX,OD,UR OD,UR DF,UR MX,OD,UR MF,SL,WT DF,MX MF OD,MF,SL 1 1 1 1 1 1 1 1,2 1,2 * 1 1 1 1 1 1 1 * nesque, 1820 nesque, (Férussac, 1822)* (J. Lewis, 1875) (Bosc, 1802) 9,29,39,79,101 MF,SL (A. Binney, 1840) (Hubricht, 1951) . (Beck, 1837) (Say, 1821) 1,3–7,9,12,16–19,23–25,29,30,33–35,37,42,44,45,47,49,50, Hubricht, 1962 (Say, 1817) 27 WT Rafi (Müller, 1774)* (Shuttleworth, 1852) 46,50,81–83,109,112,113 MF,PC,SL (Bland, 1883) 4,9,55,83,109,112 SL (Walker and Pilsbry, 1902) (Say, 1816) 1,3,6,8,9,12–15,17–19,21,22,24,26,27,29,30,32,34–37,39,41,42, Hubricht, 1962 9,29,42,44,50,94,99,109 DO,MF,SL (Pilsbry, 1889) Say, 1818 Hubricht (1985) Unknown (Gould, 1851) 29,37,45,50,79,81,101,103,104,109,116 MF,SL Hubricht, 1964 Hubricht, 1963 24 SL Hubricht, 1962 (Linnaeus, 1758)* CM 62.30312, FLMNH 096665 Unknown (A. Binney, 1842) Linneaus, 1758 (Müller, 1774) 15,21,27,31,69 OD,MX,UR,WT H.B. Baker, 1929 F.C. Baker, 1939 capsella exuolaris aff. Schrank 1803 sp (Continued) Oxychilus draparnaudi Paravitrea multidentata Haplotrema concavum Cepaea nemoralis Oligyra orbiculata Helicodiscus barri Deroceras reticulatum Ventridens theloides Vitrinizonites latissimus Zonitoides arboreus Helicodiscus multidens Helicodiscus notius Helicodiscus parallelus Lucilla scintilla Lucilla singleyanus Deroceras laeve Ambigolimax valentianus Limax maximus Galba Paravitrea blarina Paravitrea petrophila Paravitrea placentula Megapallifera mutabilis Pallifera dorsalis Pallifera fosteri Philomycus carolinianus Paravitrea capsella Paravitrea Philomycus fl Ventridens pilsbryi Oxychilidae Haplotrematidae Helicidae Helicinidae Helicodiscidae

Limacidae Lymnaeidae Philomycidae

Taxon

Table 1. LAND SNAILS OF KNOX COUNTY, TENNESSEE 7 DF,DO,MF,MX,OD,PC,SL,UR,WT DF,DO,MF,MX,OD,PC,SL,UR,WT Habitat Codes SL CV,DF,DO,MF,MX,OD, SL,UR,WT DF,MF,MX,SL,WT . (1982) Unknown et al . (1982) Unknown et al 70–72,74–86,88–91,93,94,98–101,104,109,110,112–115 81–83,85–88,93–95,98,99,107,109,113–116 9,999SL 9,42,44,50,81–83,85,CM 62.398891114,75 70,73–78,83,95,96,98,105,109,112–114 21 MF,PC,SL MF,SL 1,4–9,19,24–26,29,31,32,34,35,37,42,47,55,59,75,79,83,87,93, 96,101–104,109,112,113 FLMNH 502048 Unknown 3,13–15,20,21,27,31,33,38,39,52,53,82,94,97,100,111,113 DF 1MF DF,DO,MF,MX.OD,SL,WT SL 35 Unknown OD Site Numbers, Reference or Museum Collection 1 1 1 1 1 1 1 1 1,2 1 1 1 (Say, 1824) (J. Lewis, 1870) (I. Lea, 1839) Hubricht (1985) Unknown (Walker and Pilsbry, 1902) 95 SL (I. Lea, 1841) (Bland, 1861) (Shuttleworth, 1852) (Pfeiffer, 1842) 1,4,9,12,22,24,26,32,33,35,38,39,42,50,51,65,71,74,75,77–79, (I. Lea, 1834) (Pilsbry, 1898) 34,35,74,105,115 MF,SL (Hubricht, 1965) Hubricht (1985) Unknown (Ferussac, 1821) 4,7,9,10,29,37,75,105 MF,SL,WT Hubricht, 1958 (Say, 1817) 3,4,10–12,22,25,26,51,52,79,84,86,88,89,93,98 DF,MF,PC,SL,WT (I. Lea, 1830) 37,44,45,49,50,55,58,59,73,74,93,103 DO,MF,SL (Bland, 1873) (Say, 1816) 1,4,5,7,9,12,19,23–26,29,101–104,114,116 CV,DF,MF,SL Hubricht, 1953 P1 MX (Say, 1821) 9,24,33,55,91,104,109 DO,MF,SL (Say, 1821) 1,3,4,6,8,9,17,19,25,26,30,33–39,41,42,45–47,49–52,55,57,59, 65,67, (Dall, 1885) (Gould, 1841) (Say, 1817) 23,24,35,104, CV,MF,SL (Say, 1816) 5–7,17,34,51,55,59,78,81,83,85,94,105,109,111,113,115 DF,DO,MF,PC,SL (Say, 1821) 4,74 SL (Pilsbry, 1900) Pilsbry, 1940 1,9,25,26,29,32,39,44,45,50,55,57,59,81,91,115, MF,SL fraternum (Say, 1821) 4,8,9,34,35,37,49,55,59,74,75,77,83,93,103,105,109,112,113 DO,MF,SL (Shuttleworth, 1852) 12,29,44,109 MF,SL (Pilsbry, 1894) Hubricht (1985) and Bogan (Muhlfeld, 1816) (Say, 1821) 4,29,31,37,78,85,93,98,109 DF,MF,MX,OD,PC,SL,WT (Say, 1825) CM 62.18256, 98025, and Bogan (A. Binney, 1837) 4,6,19,25,55,75 DF,SL cf. (Say,1821) 4,7,9,14,23–25,27,29,33–35,37–39,42,43,51,52,54–56,63,65–67, ectus angellum

(Continued) ectarius kalmianus ectarius ectarius rugeli ectarius ectarius downieanus ectarius infl ectarius Triodopsis hopetonensis Infl Triodopsis complanata Infl Mesodon clausus Mesodon elevatus Mesodon normalis Triodopsis fallax Patera appressa Philomycus venustus Allogona profunda Euchemotrema Daedalochila plicata Infl Infl Mesodon thyroidus Mesodon zaletus Neohelix albolabris Stenotrema Stenotrema spinosum Stenotrema stenotrema Triodopsis tennesseensis Triodopsis tridentata Triodopsis vulgata Xolotrema caroliniense Xolotrema denotatum Pomatiopsis lapidaria Appalachina chilhoweensis Hawaiia alachuana Philomycus togatus Patera perigrapta Polygyra cereolus Stenotrema edgarianum Daedalochila troostiana Fumonelix wheatleyi

Polygyridae Pomatiopsidae Pristilomatidae

Taxon Table 1.

8 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 DF,DO,MF,MX,OD,SL,WT Habitat Codes DO,SL,WT DF,DO,MF,MX,OD,PC,SL,WT DF,DO,MF,MX,PC,SL,UR,WT DF,DO,MF,MX,OD,PC,SL,WT 74,75,77–79,81,83,88,91–95,101–105,109,112,116 9,18,24,103,10424,103,104,6521,31,56,63,65,6615,211,12,21,47,612154,64,686321,56,63,67–69,761,3,4,12,21,35,38,51,55,75,83,92,93,99,102,1034,8,9,18,21,26,30,33–35,37,39,55,56,64,74,77–79,83, 99,103,109,112 MF,PC,SL 89,95,96, 3,4,9,12,19,24,35,38,45–48,50,55,88,96,99,102–105,109,112 MF,SL 15,43,108 DO,MX,OD,UR 35,103,1043,15,28,55,6743,67,75,109 UR MF,OD DF,MF,MX,PC,SL,WT MF,MX,OD,SL,WT OD,WT OD DF,MX,OD OD,F,DO UR WT MF,SL DF,DO,SL,WT, 81–83,87–89,91,93,96,99,103,104,106,109,112 Site Numbers, Reference or Museum Collection 3,7,15,17,18,20,24,28,31,34,37,47,56,61,65,66,74–77,80,82,83, 102,109,113 DF,DO,MF,MX,OD,PC,SL,WT 1,32,34,74,1093,12,14,22,30,34,35,37,38,42,43,45,47,51,52,64,75–81, 91,93,96,105,109 83,84,88, MF,SL 1,2 * 1 1,2 1 1 1 1 1 1 1,2 1 1 1 1,2 1 1,2 1 1 1 (I. Lea, 1841) 4,9–11,14,18,19,24,25,29,33,35,42,46,47,59,64,69,73,74,79, (Webb, 1953) (Potiez and Michaud, 1838) 1 (Say, 1817) Pilsbry, 1900 1–10,12,19,22,24–26,28–30,34–36,38,45,48–50,54,55,57,60,62, (Pilsbry and Ferriss, 1906) Draparnaud, 1801* Sterki, 1893 4,7,34,35,37,55,69,109 SL,UR (A. Binney, 1841) (Say, 1817) Sterki, 1893 15,21,31,52,53,56,64 DO,MX,OD,WT (C.B. Adams, 1841) 20,21,52,53,56,63,65,71,76,84 DO,MF,OD,PC,UR,WT, (Gould, 1840) (Müller, 1774) (I. Lea, 1834) Wolf, 1870 (H.B. Baker, 1930) (Sterki, 1890) (Draparnaud, 1801)* Pilsbry, 1926 1,3,5–7,9,19,24,29,34,36,45,55,57–59,70,74,75,89,94,99,109,115 DF,DO,MF,SL,WT I. Lea, 1864 Hubricht (1985) Unknown (Müller, 1774) Morrison, 1935 (Sterki, 1890) (A. Binney, 1843) (Gould, 1840) 11,15,43,52,86,100 WT Schmacker & Boettger, 1891* Say, 1822 clavulinum cf. (Continued) Strobilops texasianus Allopeas Opeas pyrgula Catinella oklahomarum Novisuccinea ovalis Oxyloma retusum Succinea forsheyi Testacella haliotidea Vallonia costata Vallonia excentrica Vallonia perspectiva Vallonia pulchella Columella simplex Vertigo gouldii Vertigo ovata Vertigo parvula Vertigo pygmaea Punctum minutissimum Punctum blandianum Punctum vitreum Vertigo tridentata Hawaiia minuscula Vertigo milium Vertigo oscariana Punctum smithi Pupoides albilabris Strobilops aeneus Strobilops labyrinthicus Subulinidae Succineidae Testacellidae Valloniidae Vertiginidae

Punctidae

Taxon Table 1.

Pupillidae Strobilopsidae LAND SNAILS OF KNOX COUNTY, TENNESSEE 9 sites (Sites 1, 4, 37, 101, 103, 109, 112, 113), all of which were (Nystrand and Granström 1997). It also was the dominant mature hardwood forest or sandstone/limestone bluff. Anguispira slug found in recently burned black spruce forest in New- cf. jessica was found only at McAnnaly Ridge (Site 99), a mature foundland and has the ability to affect forest regeneration hardwood forest. In general, both species were associated and biodiversity (Moss and Hermanutz 2010). We have with decomposing wood. observed this species in remote forests of northern Minnesota Arion fasciatus Nilsson, 1923: This non-native species is and southern Ontario, where it was the only slug species wide ranging in North America and is found in a variety of encountered. habitats including grasses, shrubs, hedgerows and forests of Bradybaena similaris (Férussac, 1821): This species is maple/ash/elm (Chichester and Getz 1973). Its distribution known from the Gulf coastal states and is a native of south- stretches across Canada and it is well established in the north- eastern Asia (White-McLean 2011). It has invaded Australia, eastern U.S.A. and California. Elsewhere, A. fasciatus has Europe, and North and South America. Numerous live indi- been reported from maize fi elds in Ohio and Kentucky, and viduals were found in May at a landscape nursery business in the Great Smoky Mountain National Park in North Carolina Knoxville outside the greenhouses (Site P6). Because they (Burch 1962, Hammond and Byers 2002, Dourson 2010, were found during a warm season, we are unsure if the spe- 2013). We found this species at Maxwell Spring (Site 38), a cies is surviving year-round. Besides being an important agri- mature forested area near a busy road. cultural pest, B. similaris can be carrier of Angiostrongylus Arion hortensis Férussac, 1816: In North America, A. cantonensis, a parasitic nematode that causes a disease called hortensis has been reported from Canada and a number of human eosinophilic meningoencephalitis (Carvalho et al. eastern states including Delaware, Kentucky, Pennsylvania 2012). This disease typically occurs in Asia but there are doc- and Virginia (Burch 1962, Pearce and Bayne 2003, Dourson umented cases in Florida, Louisiana, and Texas (Iwanowicz 2010, Hotopp et al. 2013). In Virginia it has been found in et al. 2015, Stockdale Walden et al. 2017). At least fi ve other counties adjacent to Tennessee. Its preferred habitat includes species we found in Knox County, Ventridens demissus cultivated areas as well as deciduous and coniferous forest (A. Binney, 1843), Zonitoides arboreus, Limax maximus (Linneaus, (Chichester and Getz 1973). We found this species in shaded 1758), Ambigolimax valentianus (Férussac, 1822) and Deroceras brushy habitat of a residential area (Site 21). This non-native laeve (Müller, 1774) also have the ability to carry this parasite species is very similar to another invasive species, Arion dis- (Senanayake et al. 2003, Asata et al. 2004, Lv et al. 2008, tinctus Mabille, 1868. The identifi cation of A. hortensis was Stockdale Walden et al. 2017). verifi ed by F. Borrero of APHIS. Cochlicopa lubricella (Porro, 1838): This European spe- Arion intermedius (Normand 1852): Reported from cies has colonized southeastern Canada and northeastern Canada, New York, California and as far south as Kentucky U.S.A., and has been reported in Virginia (Hotopp et al. and Virginia, this non-native species is found primarily in 2013). It is generally found in disturbed areas. In Knox disturbed habitats (Burch 1962, Forsyth 2004, Dourson 2010, County, we found specimens adjacent to a landscape nursery Hotopp et al. 2013). We found A. intermedius in second- business in Powell (Site 21) and in three park-like habitats in growth fl oodplain forests and a former pasture at the edge of Knoxville (Sites 53, 66, 69). There is disagreement among a spring (Sites 3, 13, 27). specialists over the separation of this species from Cochlicopa Arion subfuscus (Draparnaud, 1805): This invasive spe- lubrica (Müller, 1774), another species introduced from cies was found on a forested slope adjacent to the Third Creek Europe (Welter-Schultes 2012). We found both species in Greenway within the city limits of Knoxville (Site 51). Arion Knox County and followed Pearce and Drescher (2017) in subfuscus has been introduced throughout northeastern separating the two using shell size (>2.29 mm diameter = U.S.A, where it thrives in natural woodlands as well as devel- C. lubrica, <2.19 mm diameter = C. lubricella). oped areas (Chichester and Getz 1973, Blanchard and Getz Deroceras laeve (Müller, 1774): Although thought by 1979). It appears to be increasing its southern range as there many to be the only native Deroceras in North America, some are records in Virginia and North Carolina (Grimm et al. researchers believe the species originated in Europe (Dourson 2009, Hotopp et al. 2013). It has been noted as a pest of maize 2013, D. Robinson pers. comm.). It is a widely distributed and soybeans in Ohio (Hammand and Byers 2002). Previous garden pest throughout the eastern U.S.A. (Burch 1962) and to this survey, it had not been reported in Tennessee. The was previously known to occur in Knox County (McCracken success of this species in woodland areas of the Third Creek and Selander 1980). We found this slug in residential and Greenway may be a threat to local native slugs. Although this urban areas (Sites 15, 21, 27, 31, 69). site is surrounded by disturbed/residential areas, the occur- Deroceras reticulatum (Müller, 1774): This European rence of A. subfuscus in a mature hardwood forest is cause for slug is an important agricultural pest throughout North concern. Arion subfuscus is an important predator of Pinus America and in many places worldwide (Burch 1962), how- sylvestris (Linnaeus, 1753) seedlings in northern Sweden ever we found no published records of this species in Tennessee. 10 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018

In Knox County, this slug is common in fl oodplains and resi- in a low, scrubby area (Site 111). We separated it from dential and urban areas (Sites 21, 31, 69). Mesodon clausus (Say, 1821) using shell characteristics. Dryachloa dauca Thompson and Lee, 1980: Described Limax maximus Linneaus, 1758: This species is now found from Florida, this distinctive snail has also been found in on every continent except Antarctica (Grimm et al. 2009). In Alabama, and within the last few years has been collected in North America, it is distributed widely in Canada, has been the southern Piedmont of North Carolina where it occurs reported from Mexico, and resides in much of the U.S.A. in degraded habitats along railroads and in relatively undis- including Alaska, Oklahoma and Hawaii. In Tennessee, it has turbed, species-rich areas (A. Van Devender, pers. comm.). It been reported around old homestead sites in the Great Smoky is unlikely this species has been overlooked by earlier collec- Mountains National Park (Dourson 2013) and our discussions tors in the eastern U.S.A. and it’s rapidly expanding range with other malacologists indicate it has resided in urban and suggests it may not be native to North America (H. Lee, pers. residential environments of Knox County for many years. comm.). We found one individual in a second growth forest Limax maximus is a large, yellowish-grey slug readily recog- in Knoxville (Site 39). nized by having distinct black spots on the mantle and spots Galba Schrank 1803 sp.: Four species of Galba are known often coalescing into two or three lines on the body. A total of to occur in Tennessee (Johnson et al. 2013). Galba typically three individuals were found in two urban/residential areas occurs on the edges of ditches and ponds and can be semi- (Sites P2, P3). Both specimens at Site P3 (a parking lot) were aquatic, as long as the area is moist or fl ooded sporadically pale, uniformly yellowish grey, and devoid of black markings. (P. Johnson, pers. comm.). It can survive dry periods for Pilsbry (1948) refers to a rare form of L. maximus that matches more than a year in aestivation (Kendall, 1949). Although it is our specimens. Our identifi cation was made based on internal not typically included in a list of terrestrial snails, we found anatomy and confi rmed by F. Borrero (APHIS). Galba sp. on a mature forested slope approximately 25 m Opeas pyrgula Schmacker and Boettger, 1891: An intro- from a small stream and on the edge of an ephemeral creek duced species common in urban areas, it is known from (both in Site 82). Other specimens were found on the edge of Pennsylvania, Virginia, and most of the Atlantic and Gulf a landscape nursery (Site 21). Coast states of the eastern U.S.A. (Dundee 1971, Hotopp Gastrocopta holzingeri (Sterki, 1889): This species was et al. 2013). We found O. pyrgula in six urban locations (Sites found on Saltpeter Bluff, a limestone/sandstone bluff over- 21, 31, 56, 63, 65, 66). A literature search produced no previ- looking the Tennessee River (Site 34). It has been found in ous Tennessee records. limestone of the same geological origin in the Shenandoah Pallifera dorsalis (A. Binney, 1842): This species ranges Valley of western Virginia (J. Nekola, pers. comm.). The only from eastern Canada and the northeastern U.S.A. to Tennessee previous record of this species in Tennessee is a fossil from (Chichester and Getz 1973, Dourson 2013). At Site 91, a Jackson County near the Kentucky border (Hubricht 1985). mature forest on a karst ridgetop, individuals we encountered Glyphyalinia cryptomphala (G. H. Clapp, 1915) and had a rusty or brownish foot margin along its entire length. Glyphyalinia solida (H. B. Baker, 1930): For these two similar Because this color pattern was subtle, we identifi ed them as species, Hubricht (1985) reported that shell and anatomical P. dorsalis rather than Pallifera ohioensis (Sterki, 1908) which differences do not overlap. Our survey produced specimens is rare and has been reported from only Ohio, Indiana and clearly matching each description. To differentiate between one county in North Carolina (Dourson 2013). Chichester immature shells, the diameter was measured at 4.5 whorls; and Getz (1973) described a similar color form for P. dorsalis immature shells with a diameter of 4.5 mm or less were in their treatment of terrestrial slugs of northeastern North recorded as G. cryptomphala, and immature shells measuring America, but did not discuss P. ohioensis. We found speci- 5.5 mm or more were reported as G. solida. mens without a rusty foot margin at Site 5, a karst forested Hawaiia alachuana (Dall, 1885): Hubricht (1985) con- slope on Copper Ridge. sidered this to be a calciphile, and his distribution map Paravitrea capsella (Gould, 1851): We found individu- showed sporadic occurrence over much of the eastern and als that clearly were Paravitrea capsella and at least three midwestern U.S.A. Three main clusters are apparent: 1) unnamed forms. Hubricht (1985) considered P. capsella to be Tennessee, Virginia, Maryland and West Virginia; 2) eastern a complex of species that share a similar shell form but with Oklahoma and nearby Arkansas; 3) peninsular Florida. We distinct anatomical differences. The three unnamed forms found H. alachuana only at Site 35 on Saltpeter Bluff, a lime- conchologically resembled P. capsella, but each uniquely var- stone cliff complex adjacent to the Tennessee River. ied morphologically from the species description. These dif- Infl ectarius downieanus (Bland, 1861): This species is ferences included size, presence of teeth in juveniles and usually found on fl at-topped mountains where the soil is adults, and genital structure. We lumped these forms into a slightly acidic (Hubricht 1985). A single specimen was col- group we call Paravitrea aff. capsella. As they did not co-occur lected at Seven Islands State Park near the French Broad River with P. capsella, we listed Paravitrea aff. capsella separately. LAND SNAILS OF KNOX COUNTY, TENNESSEE 11

Polygyra cereolus (Mühlfeld, 1816): Native to Florida, and number and strength of ribs as they pass into the basal this species has extended its range to all the Gulf States, the area, showed a clear distinction. Both species were found in Carolinas, Oklahoma, Kentucky and Hawaii (Pilsbry 1940, sites associated with rocky boulders or outcrops. Perez and Cordeiro 2008, Bergey et al. 2014). It can be a pest Triodopsis fallax (Say, 1825): Several lots of this species species on legumes and possibly other crops (Kalmbacher were collected in Knox County in the late 1800s and early et al. 1979). We found this species near the periphery of a 1900s by G. H. Clapp and V. Sterki and cataloged at CMNH; landscape plant nursery (Site 21). we verifi ed two of these lots. This species was also reported Stenotrema angellum Hubricht, 1958: Hubricht (1985) in Knox County by Bogan et al. (1982). Hubricht (1985) reported S. angellum in three north-central Tennessee coun- reported T. fallax from three east Tennessee counties: Blount, ties: Fentress, Clay and Jackson. It has now been found in Hamilton and Monroe. We did not fi nd T. fallax in our study. eight additional Tennessee counties, including three in the Triodopsis hopetonensis (Shuttleworth, 1852): This is a Valley and Ridge physiographic region of East Tennessee: common snail of urban, residential and disturbed habitats. Hancock, Hawkins, and Cocke (H. Lee, pers. comm.). We According to Hubricht (1985) this species has been intro- found this species to be common in Knox County (33 sites in duced in eastern Tennessee, Alabama, and Mississippi. We various habitats). Specimens we encountered were smaller have frequently encountered this species at interstate rest than the typical diameter described by Hubricht (1958) (9.5– areas in Virginia, North Carolina, and South Carolina. In 10.0 vs. 10.0–11.0mm). In a few of the UMMZ lots cataloged as Knox County it was common in 19 urban or otherwise dis- Stenotrema stenotrema (Pfeiffer, 1842) we also found S. angellum. turbed sites, often with Ventridens demissus (Binney, 1843). We suspect this may be typical of other collections, as these Ventridens coelaxis (Pilsbry, 1899): This species is charac- two species are often syntopic in Knox County. terized as a narrow endemic occurring in 13 counties at the Stenotrema edgarianum (Lea, 1841): The sole Knox convergence of Tennessee, Virginia, and North Carolina County record for this species is an undated specimen at (Dourson 2013, Hotopp et al., 2013). Some of these counties FLMNH collected by Mrs. G. Andrews who actively collected occur wholly or partially in the Valley and Ridge, Blue Ridge, in the late 1800’s and early 1900’s (Pilsbry 1900) and was a or the Cumberland Mountain physiographic provinces. resident of Knox County. Only state and county of origin are Hubricht (1985) reported V. coelaxis is usually found on provided for the Knox County specimen in FLMNH. All mountainsides at higher elevations, which would imply the other records for this species are restricted to the Cumberland species is generally in areas within the Blue Ridge or Cumberland Plateau, the physiographic region west of the Valley and Mountains. Our Knox County record of V. coelaxis is from an Ridge. Given this, we question the locality attributed to the undated lot in the CMNH collection. This lot was donated to Knox County specimen in the FLMNH collection. CMNH in the early 1900’s. No specifi c locality within Knox Striatura meridionalis (Pilsbry and Ferriss, 1906): This County was given. This record is interesting, maybe ques- was the most commonly encountered species in Knox County tionable, considering the absence of high elevation habitats in (66 sites). Striatura meridionalis is a microsnail that inhabits Knox County. leaf litter in a broad range of habitats. Coney et al. (1982) Vertigo pygmaea (Draparnaud, 1801): This species has a found this to be the most abundant snail in a survey of the spotty distribution mainly in the northeastern U.S.A. It is a Hiawasee River Basin. non-native species from western Eurasia (Welter-Schultes Strobilops labyrinthicus (Say 1817) and Strobilops texasianus 2012). In Tennessee it had previously been recorded in (Pilsbry and Ferriss, 1906): These species can be diffi cult to Cumberland County (Nekola and Coles 2010). We found this separate. According to Dourson (2015), S. labyrinthicus is species in a wide range of habitats, including wetland, lime- separated from S. texasianus by having weaker ribs that fade stone river bluff, and an urban rubble slope (Sites 3, 15, 28, as they pass into the basal area and a smaller shell height 55, 67). (1.8 mm vs 2 mm). Rib counts, placement of the fi ve basal lamellae, and the number of ribs on the last whirl (i.e., > 50 ribs on the last whirl = S. labyrinthicus) also are useful features DISCUSSION (T. Pearce, pers. comm). We found both species in Knox County. Strobilops labyrinthicus occurred at fi ve sites (Sites 9, Of the 15 new state records we documented in Knox 18, 24, 103, 104) and S. texasianus occurred at three of those County, only four are species native to North America: sites (Sites 24, 103, 104). Signifi cant variation was present in Oxyloma retusum (I. Lea, 1834), Polygyra cereolus (Muhlfeld, position of the basal lamellae, but shell condition often made 1816), Vallonia costata (Müller, 1774) and Vallonia pulchella this character diffi cult to discern. Specimens we identifi ed as (Müller, 1774). All four were found in urban/residential S. texasianus were generally taller but this was not consistent. areas, suggesting they were introduced to Tennessee by Other diagnostic characters, such as width of the umbilicus anthropogenic means. Vallonia costata and V. pulchella are 12 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 generally more northern in distribution, although V. pulchella that includes the Valley and Ridge and the Blue Ridge physio- is also distributed in many western states (Pilsbry 1949, graphic regions, Coney et al. (1982) reported 86 species. With Hubricht 1985). Polygyra cereolus appears to be increasing 151 species, Knox County has a land snail fauna that is diverse its distribution in a northerly direction. Pilsbry (1940) pre- even for the southeastern U.S.A; in fact Knox County has sented numerous records of P. cereolus from Florida but in no more species than has been reported for the entire state of other states. More recently it has been found in several states Louisiana (140 species) (Minton and Perez 2005). Although a including Hawaii. number of species we found are not native to North America, Eleven percent of the Knox County snail fauna are not we report 136 native species. Much of this diversity was found native to North America. Many of the non-native species we on high bluffs along rivers and creeks and in relatively undis- documented in Knox County are considered widespread in turbed forested slopes. While this was a non-quantitative the U.S.A. and likely have been present in Tennessee for study, our survey results indicate that steep slopes and rock decades, but there has been no account of their occurrence in bluffs may be an important predictor of high snail diversity the published literature. The spread of non-native land snails (see Appendix 1). Similarly, Coney et al. (1982) found slope is of concern because they have the potential to cause harm and snail diversity were positively correlated and suggested to agriculturally important crops and to native species slope effect may be indirect; physical characteristics associ- (Robinson and Slapcinski 2005). Several of the non-native ated with slope, such as moisture levels, forest and rock types, species we found in Knox County: Bradybaena similaris and higher calcium availability, may be more infl uential. One (Férussac, 1821), Cochlicopa lubricella, Cochlicopa lubrica, of the more interesting areas we surveyed was Saltpeter Bluff, Deroceras reticulatum and Oxychilus draparnaudi (Beck, a high and nearly inaccessible cliff overlooking the Tennessee 1837), as well as Florida native P. cereolus were found on or River on the southeast edge of Knox County. Two sites on adjacent to landscape plant nurseries. Plant nurseries appear this bluff (Sites 34 and 35, with a combined approximate area to play a role in the spread of non-native land snail species of 2.5 ha) produced a total of 40 species, including several of (Bergey et al. 2014). With the exception of Arion subfuscus, all the rarest species in this survey. Three species were found non-native species we found were in urban, residential or dis- only on this cliff: Hawaiia alachuana, Lucilla scintilla turbed areas. Arion subfuscus has been invading forests in (H.B. Baker, 1929), and Gastrocopta holzingeri. Three other Canada and the northern U.S.; its appearance in a mature Saltpeter Bluff species, Vertigo parvula Sterki, 1890, Gastro- woodland in Knox County is troubling because of the poten- copta lewisiana (G.H. Clapp, 1908) and Punctum smithi tial for competition with native land snail species. Morrison, 1935, were found at only a few other locations. The Microsnails (<5mm at greatest dimension) comprised most speciose area was a forested cove that included three 38% of the Knox County land snail fauna; 46% of our micros- sites in the northeast corner of the county (Sites 9, 10, 11; nail species represent new records for Knox County. Dourson N = 48 species). This area (approximately 5 ha) included a (2007) suggested this component of the southeastern land mature forest and adjacent emergent wetland in the vicinity snail fauna is under reported, and he pointed out that in of an undisturbed chert gravel spring issuing from the base of Hubricht’s 1985 distributional atlas, larger snail species were a sandstone/limestone cliff. better represented. Hubricht (1985) also commented on gaps A review of the published historical land snail fauna in in the distribution for many species, which he attributed to the eight counties adjacent to Knox County produced the fol- differences in collecting effort. This strongly suggests some of lowing number of species (in brackets) by county: Blount the previous collecting efforts in the eastern U.S.A. did not [120], Sevier [105], Roane [54], Anderson [54], Grainger include sampling of soil and leaf litter, or perhaps only minor [43], Union [30], Loudon [17], Jefferson [11] (Hubricht attention was given to this microhabitat. This may explain 1985, Nekola and Coles 2010, Dourson 2013). It is not sur- why one of the most widespread species we found, Gastrocopta prising that Blount and Sevier Counties have the greatest contracta (Say, 1822), a snail 2.2–2.5 mm long, had not previ- putative diversity because about one-third of both counties ously been recorded in Knox County. have been subject to recent land snail surveys as part of the All Compared to the remainder of the U.S.A. the Southeast Taxa Biodiversity Inventory in the Great Smoky Mountains supports a high diversity of land snails. A variety of factors are National Park (Dourson 2013, Douglas et al. 2014). Further, associated with this diversity, including topography, geology Blount and Sevier Counties are in the Valley and Ridge and and geological history, soil chemistry, and climate. In central the Blue Ridge physiographic regions, and areas where mul- Kentucky, Dourson (2007) found a 2 ha site supported at tiple physiographic regions merge have been shown to pro- least 86 species of land snails. This relatively small site encom- vide habitat characteristics, geology, forest structure, and passed the junction of three physiographic regions rich in availability of various other resources required to generate limestone outcrops and calcareous soil. Within the lower high land snail diversity (Dourson 2007, Dourson and Beverly Hiwassee River basin in southeastern Tennessee, in an area 2008, Douglas et al. 2014). Most of Jefferson County is in the LAND SNAILS OF KNOX COUNTY, TENNESSEE 13

Valley and Ridge physiographic region, and a small part of provided information on new Tennessee records for Stenotrema the southeast corner of the county is in the Blue Ridge (Soil angellum. David Etnier and Charley Saylor shared valuable Survey Staff 2002a). This information suggests the known information regarding interesting habitats. Numerous private land snail fauna for Jefferson County is wildly underreported, land owners generously permitted access to sites: Bill and Sarah and to a lesser degree the same may be true of the other counties Alexander, Tim and Valerie Baumann, Jeff and Vicki Chapman, adjacent to Knox County, especially those spanning multiple Willa Essie, Laurence B. Estes, III, Lynn and Edgar Faust, physiographic provinces, e.g., Anderson and Roane – both Dylan Jones and Erica Lyon, Helen Lucas, Brian Mize, Alex are mainly in the Valley and Ridge with a small portion on Osmond, Ruby Paterson, Bonny Rogers, Stephanie Walker and their western side in the Cumberland Plateau (Soil Survey Billy W. Wallace. Art Bogan (North Carolina State Museum) Staff 1978, 2002b). and Stephanie Clark provided references and general encour- This study revealed Knox County’s land snail fauna is agement. Kristin Irwin cataloged specimens in the early stages remarkably more diverse than was previously known. The ear- of this survey. Nick Gladstone contributed information on liest records of land snails in Knox County date to the late snails occurring in Knox County caves. Assistance with collect- 1800’s, but by that time period, all old-growth forest in the ing permits was provided by Doug Bataille (Knox County Parks county had been cleared. Loss of old growth habitat may have and Recreation), Rusty Boles (Tennessee Wildlife Resources affected the land snail community. Thus, Knox County’s com- Agency), Paul James (Ijams Nature Center) and Roger McCoy plete historical record may never be known. To address this (TDEC). Thanks to Carol Evans and Legacy Parks Founda- question future research could compare the land snails of old- tion for the ongoing efforts to conserve our Knox County open growth forests in nearby Great Smoky Mountains National spaces for snails and people, too. Dinkins Biological Consulting, Park to adjacent forest communities similar in elevation, geol- LLC. funded this study. We appreciate our reviewers, Daniel ogy and physiography to those found in Knox County. Douglas and Tim Pearce, who greatly improved this publica- Over the last several years, the municipal governments of tion. Finally, a special thanks to Amy and Wayne Van Devender Knoxville and Knox County, in collaboration with non- who provided specimens, spent numerous hours helping with governmental organizations, have actively promoted the pres- identifi cations and answering questions, and gave enthusiastic ervation and use of urban wilderness for outdoor recreation. encouragement in all aspects of this study. We applaud this effort and hope areas set aside for protection will serve as refugia for native fl ora and fauna in the face of LITERATURE CITED the county’s increasing human population. Allen, J. A. 2004. Avian and mammalian predators of terrestrial gas- tropods. In: G. M. Barker, ed., Natural Enemies of Terrestrial ACKNOWLEDGMENTS Molluscs. CABI Publishing, Oxford, UK. Pp. 1–36. American Forests. 2002. Urban ecosystem analysis Knox County, Our interest in land snails was initiated by David Withers Tennessee. Available at: http://www.systemecology.com/4_Past_ of the Tennessee Department of Environment and Conserva- Projects/AF_KnoxCounty.pdf 22 January 2018. Asato, R., K. Taira, M. Nakamura, J. Kudaka, K. Itokazu, and M. tion (TDEC) and Geoff Call of the U.S. Fish and Wildlife Service Kawanaka. 2004. Changing epidemiology of Angiostrongyliasis who urged one of us (GRD) to become involved with a study cantonensis in Okinawa Prefecture, Japan. Japanese Journal of of Anguispira picta, a federally threatened land snail in Franklin Infectious Disease 57: 184–186. County, Tennessee. From there, our interest in this unique Bergey, E. A., L. L. Figueroa, C. M. Mather, R. J. Martin, E. J. Ray, group of animals grew. A number of people have patiently an- J. T. Kurien, D. R. Westrop, P. and Suriyawong. 2014. Trading swered questions regarding problematic species, distribution, in snails: Plant nurseries as transport hubs for non-native spe- and collecting techniques: Francisco Borrero and David cies. Biological Invasions 16: 1441-1451. Robinson (APHIS), Dan Dourson, Paul Johnson (Alabama Blanchard, D. and L. L. Getz. 1979. Arion subfuscus in southeastern Aquatic Biodiversity Center), Jeff Nekola (University of New Michigan. The Nautilus 93: 36–37. Mexico), and Harry Lee. Several curators provided access to Bogan, A. C., C. Coney, and W. A. Tarpley. 1982. Distribution of the their mollusk collection, verifi ed identifi cations, and generously Polygyridae (Mollusca: Pulmonata) in Tennessee. Journal of the Tennessee Academy of Science 57: 16–22. loaned specimens for comparison, especially Jochen Gerber Bouchet, P. and J. Rocroi. 2005. Classifi cation and nomenclature of (FMNH), Taehwan Lee (UMMZ), Tim Pearce (CMNH), John gastropod families. Malacologia 47: 1–397. Slapcinsky (FLMNH), and Tom Watters (OSUM). We were Burch, J. B. 1962. How to know the Land Snails. W. C. Brown Com- assisted in the fi eld by Jamie, T.C., and Zack Dinkins, Bannon pany Publishers, Iowa. Evans, Tim Evans, Mackenzie Hodges, Teresa Petit, Teresa Sex- Caldwell, R.S. and D. C. Dourson, 2008. Rare Land Snails of the ton, Trevor Tapscott, and Hailey Vincent. Zack Dinkins provided Cherokee National Forest. Miscellaneous Publication No. 1 of valuable assistance interpreting site geology. Bob Winters Cumberland Research Center, Lincoln Memorial University. 14 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018

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The Medical Journal of Australia 179: 430–431. 16 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 r No.Sp. elevation (ELV), and number of -84.0220 -84.0220 -84.0220 -84.0202 -84.0204 -84.0220 -83.9550 -83.9544 -83.8648 -83.8669 -83.8675 -83.9009 -83.9638 Latitude/ Longitude (decimal degrees) /limestone outcrops and river bluffs, (O) = 0–40% canopy, C cave. Sites P1 orest, DO = Disturbed sandstone/limestone ELV (m) CC ASP MF NE F 305 36.0975 27 WT SSE F 335 36.0955 4 SL NEMF F 311SL F NW 36.1489 FWT 293 NW 433 F 36.1481WT 23 360 36.1671 O 36.1666 20 348 44 36.1673 13 7 WT MSL 256 NWMF M SE 36.0850SL 268 F NE F 305 36.0861 24 268 36.0854 36.0839 38 13 14 MF NWT F 366 O 36.0670 311 36.0701 16 6 HAB ne (1955). et al. ne sandy loam from maple/oak; fi acidic sandstone forest, oak/hickory; mesic; fi sandy loam hardwoods, beech/oak; silty clay loam forest, beech/oak; karst; silty clay loam hardwoods, beech/maple/oak; cherty silty loam gravel; mixed hardwoods, oak/beech; cherty silty loam silt loam sycamore/oak; alluvial silt loam mixed hardwoods, beech/sycamore forest, hemlock/oak; silty clay loam hardwood forest, beech/sycamore; silty clay loam limestone outcrops; English ivy; cherty silt loam, walnut/cedar; recently pasture; alluvial silt loam Steep ravine in a seep mixed hardwood Floodplain; mixed hardwoods, Limestone cliff facing creek; Steep slope; karst, mixed hardwood Moderate slope; karst; mixed Moderate slope; mixed hardwoods; In grasses near spring and under Habitat oodplains. Aspect abbreviated by direction: if blank, area was level, mixed or in a cave. Location: LDB = Left descending rive Diggs Gap Hollow, Hollow Hollow Hollow of SR 441, Hole Spring Location Description of land snail survey sites in Knox County, Tennessee, including habitat (HAB), aspect (ASP), canopy coverage (CC), Date 1 6NOV2016 Bull Run Ridge, Diggs Gap Steep slope; mixed hardwood forest, 2 6NOV2016 Bull Run Ridge, W of 3 30MAR20164 Bull Run Valley, E of Stillhouse 30MAR20165 Copper Ridge, E of Stillhouse 30MAR20166 Copper Ridge, E of Stillhouse 30MAR20167 Copper Ridge, W of Stillhouse 6FEB20178 Bull Run Valley, S of Hill Rd 6FEB20179 Limestone cliff facing creek; mixed Bull Run Valley, S of Hill Rd 21MAY201610 Copper Ridge, Texas Valley Rd Moderate slope; mixed hardwood 21MAY2016 Steep wooded karst slope; mixed Copper Ridge, Texas Valley Rd11 21MAY2016 Edge of woodland spring with chert 12 Copper Ridge, Texas Valley Rd 22JAN2017 Edge of pond; mesic; cattails/grasses; Black Oak Ridge, 2 km E 13 4OCT2015 Beaver Valley, Emory Rd, Blue Appendix 1. Site UR = Urban/residential, WT Seeps/wetlands/fl species (No. Sp.). Canopy designations are abbreviated as follows: Full (F) = 60–100% canopy, Medium (M) 40–60% Open through P6 represent opportunistic collections. Habitat designations are as follows: CV = Caves, DF Disturbed second growth f outcrops, MF = Mature hardwood forest, MX Mixed habitats, OD Open disturbed, PC Pine forest/cedar groves, SL Sandstone/ bank, RDB = right descending river bank. Soil information is from Roberts LAND SNAILS OF KNOX COUNTY, TENNESSEE 17 No.Sp. -84.1231 -84.1418 -84.2045 -84.2463 -84.2300 -84.0175 -84.0186 -84.0492 -84.0562 -84.0599 -84.0601 -84.0595 -84.0595 -83.9642 -83.9995 -84.0179 -84.0175 -84.0163 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP MF SEE F 262 35.9469 26 WTDF N F F 287 299WT 35.9816MX 35.9548 F O 10 244 274 5 35.8898 35.8800 13 13 DO O 357CV 36.0212 C 360 6 36.0491 9 DF NW OOD 357WT 36.0179 M 305 FSL WSL 308 36.0160 16 F NWT 36.0473 F 329 24 F 294 36.0478 14 274 36.0449 36.0450 32 18 20 DFWTWT M M 311MF 299 F NW 36.0701PC F 311 36.0455 NW F 317 36.0203 317 8 36.0206 15 36.0213 6 14 13 HAB rst full rst oodplain; alluvial oodplain; eld; grasses; limestone/mudstone ne sandy loam from acidic sandstone mixed hardwood maple/oak; cherty silt loam forest; mixed alluvium growth forest; eroding cherty silt loam silt loam forest edges, fence rows; mixed soils outcrops; silty clay loam dark room around debris, under plastic forest; alluvial silt loam near cave opening; silty clay loam mixed hardwoods, beech/sycamore mixed hardwood forest, beech/ sycamore; silt loam karst; clay loam grasses; alluvial silt loam oak/hickory/poplar; mesic silty clay loam forest, oak/poplar; silty clay loam karst; silty clay loam fi Floodplain; recent pasture, second growth Moderate slope; disturbed second Highly disturbed area; mixed samples; Residential lawn, near landscape business; Low swale; mesic mixed hardwood Cave; twilight section and in fi Mature mixed hardwood; karst, Limestone bluff over forested spring; Mesic soils around spring; karst; Second growth on recent pasture; Habitat eld Edges of wetland; mixed hardwoods, oodplain, of Pellissippi Pkwy around the truck stops Emory and Karns Valley Rds W of Clinton Hwy Campground Rd Valley Cave Bell Campground Rd Bell Campground Rd Bell Campground Rd Blue Hole Spring Location (Continued) Date 28 5JUN201529 Hardin Valley Rd, 1.5 km E 1OCT201530 Beaver Valley, Couch Mill Rd 3AUG201531 Steep karst slope facing Beaver Creek; Hickory Creek Rd, along creek 17APR2016 Hines Valley, I 75 Watt Rd Exit, Scrubby forest in fl 20 22APR201521 Beaver Ridge, NW base 5NOV201622 West Beaver Creek Dr, 150 m 23DEC201523 Copper Ridge, 600 m E of Bell Open fi 27MAR201624 Copper Ridge, Brushy 19SEP201525 Copper Ridge; 650 m E of 30MAR201626 Copper Ridge, Fowler Spring, 30MAR2016 Copper Ridge, Fowler Spring, 27 19SEP2015 Beaver Valley fl 14 4OCT2015 Beaver Valley, Emory Rd. near 15 26MAR201616 Beaver Valley, Old Powell Airfi 12MAR2016 Beaver Ridge, NW base17 14MAR201618 Beaver Ridge, NW slope 1APR201519 Woodland spring, mixed hardwoods, Beaver Ridge, NW base 22APR2015 Moderately steep slope; mixed hardwood Beaver Ridge, along power line Open area along power line; Mixed pine forest, cedar, few hardwoods; Appendix 1. Site 18 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 No.Sp. -84.2336 -84.1162 -84.1655 -84.1649 -84.1636 -84.0641 -84.0386 -84.0214 -83.9935 -83.9928 -83.9925 -83.9931 -83.9524 -83.9470 -83.9506 -83.9395 -83.9491 -83.9403 -83.9216 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP MF NW FDO 305 F 35.8753 287 35.9243 9 14 SL S M 274 35.8076 25 SL NNWMX F 274 F 35.8502 274 35.8851 25 24 SL SMF F F 244DF 305 35.8073 E 35.8077 F 311 38 10 35.8989 18 HAB WT SDF 0 SMF N 250 FWT F 287 35.8871 262 35.8885 O 35.8893 262 2 35.8898 7 15 13 MF NE F 34 35.8968 26 MF NMF F NEPC F 262 WMF 276 M NWT 35.8926 250 F 35.8945SL 262 35.8975 F N 11 256 35.8985 F 27 35.8925 280 12 20 35.8978 7 11 eld; oodplain, some low vegetation; hickory/oak; low limestone outcrops; cherty silt loam hardwoods, oak/sycamore oak/cedar boulders; mixed hardwoods mixed samples; mesic and dry soils sandstone outcrops; cherty silt loam hickory; silty clay loam forest, hickory/oak clay loam calcareous sandstone present; clay loam beech/maple; calcareous sandstone present; clay loam clay loam beech/maple; sandstone outcrops; sandy clay loam hickory; calcareous sandstone; loam hickory; calcareous sandstone; loam silty clay loam forest, oak/hickory; karst; loam bank; beech/oak; limestone rocks; silty clay loam sandstone; mixed hardwoods, beech/hickory/oak; loam Limestone cliff; some mature hardwoods, Very steep slope; mixed hardwoods; some Ridgetop; mixed hardwood forest, oak/ Moderate slope; karst; mixed hardwood Habitat Grassy fl Steep slope; mixed hardwood forest, oak/ Steep slope; mixed hardwood forest, oak/ Cedar grove; limestone/shale outcrops; Moderate slope; mixed hardwood Mixed hardwood forest along stream Mesic bluff of layered calcareous Tennessee R Mile 610 Tennessee R. Mile 610 Tennessee R. mile 610 Rudder Ln Duncan Br near S entrance S of Knob Cr. N side E side E side S of Knob Cr. Location (Continued) Date 32 10OCT2016 Blackoak Ridge, at Watt Rd Steep slope; mixed hardwood forest, 33 12FEB201734 Webb School of Knoxville 17NOV2015 Saltpeter Bluff, RDB of Disturbed area in a sinkhole; karst; mixed 35 8DEC201536 Salt Peter Bluff, RDB of 8DEC201537 Above Saltpeter Bluff, near 7NOV201638 Keller Bend of Tennessee R. 8JUN201539 Maxwell Spring, Northshore Rd 8NOV201540 Steep forested slope; large limestone Karst; mixed hardwood forest near spring; Rudder Oaks Way and 17MAY2015 Tennessee R. Mile 635, RDB Moderate slope; open grassy fi Appendix 1. Site 41 17MAY201542 Tennessee R Mile 636, RDB 2AUG201543 S of Duncan Rd and Br Moderate slope; mixed hardwood forest; 2AUG2015 Steep slope; mixed hardwood forest, 44 S of Duncan Rd; along 4SEP201545 Brown Mtn., I.C.King Park, 12SEP201546 Brown Mtn., I.C. King Park, 7OCT201547 Brown Mtn., I.C.King Park, 24DEC201548 Brown Mtn., I.C.King Park, 12MAR2016 Brown Mtn., I.C. King Park, 49 30MAR2016 Brown Mtn., I.C. King Park, 50 24MAY2016 Brown Mtn., S of Hendrix Ln Steep slope; mixed hardwoods, LAND SNAILS OF KNOX COUNTY, TENNESSEE 19 No.Sp. -83.9635 -83.9572 -83.9413 -83.9423 -83.9363 -83.9311 -83.9299 -83.9280 -83.9232 -83.9210 -83.9211 -83.9098 -83.9334 -83.9229 -83.9215 -83.9253 -83.9215 -83.9168 -83.9122 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP MF NWWT F 280 FDF 35.9484 287 NESL F 35.9513 NDO 250 20 M 35.9505 259 21 M 259 35.9408 35.9443 6 30 15 MF SW F 290 35.9413OD 13 O 329 35.9409 6 MF NEWT F 320 FDF S 35.9399 268 FMX 35.9393 307 21 F 35.9457OD 276 6 DO N 35.9575 M 3 M 274 293 35.9750 15 35.9679 13 7 DO N F 244 35.9414 10 HAB oodplain oodplain with oodplain owers, grasses; loam limestone sinkhole; cherty silt loam mixed hardwoods, sycamore; mesic alluvial silt loam waste area around dumps and buildings; exotic vegetation and weeds, beech/privot next to a busy road; exotic vegetation forest, beech/maple; calcareous sandstone; loam forest, oak/hickory; clay loam hemlock; alluvial loam wildfl eroded soils; some loam and limestone bluff; cherty silt loam; close to urban area wall; silty clay loam next to a parking lot forest, maple/oak; disturbed; trashy; calcareous sandstone outcrops; loam Moderate slope; mixed hardwood Moderate slope; mixed hardwood Urban; trashy; shrubs; grassesUrban, degraded; weedy UR NW M 274 UR 35.9626 S M 256 13 35.9567 13 Habitat R. Mile 646, LDB, R. Mile 646, LDB and Grand St, Knoxville near Second Cr. Location (Continued) Date 51 12SEP2015 Third Creek Greenway Moderate slope; mixed hardwood forest; 52 12SEP201553 Third Creek Greenway 5MAY201554 Third Creek Greenway 5MAY2015 Bamboo stand in fl 55 Third Creek Greenway 26JUL201556 Open grassy slope along roadway Tennessee R. Mile 646, LDB 14MAR201757 Neyland Dr , TN R. Mile 646 Mixed samples in urban park, 24OCT2015 Limestone cliff; mixed hardwoods OD Forested area S of Tennessee Open, exposed limestone outcrops E O 250 35.9529 9 Appendix 1. 58 24OCT20159 Forested area S of Tennessee 22AUG201560 High Ground Park 22AUG201561 High Ground Park 22AUG201562 High Ground Park 2JUN201563 Moderate slope; mixed hardwood Woodlawn Pike, Graystone Church 15MAR2016 Second growth forest; gullied 64 Block around the RR track Along a small wooded stream; beech/ 7MAY2015 Open meadow occasional trees; native 65 Second Creek Greenway 21NOV201666 Neyland Dr, under RR bridge 15MAR201667 Mixed sample; wooded fl Old Grey Cemetery, Tyson St 15MAR201668 Jackson Ave and Gay St Parklike with some trees; along stone 15MAR201669 First Presbyterian Cemetery 15MAR2016 Blount Mansion, Knoxville Urban; steep, trashy, rocky outcrop Urban, parklike; large trees Lawn and shrubs; grasses OD UR M 293 M 35.9647 305 35.9631 5 12 Site 20 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 No.Sp. -83.8762 -83.8780 -83.8492 -83.8442 -83.8470 -83.8561 -83.9491 -83.9355 -83.9411 -83.8603 -83.8596 -83.8626 -83.8677 -83.8741 -83.8752 -83.8731 -83.8497 -83.8491 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP PC NEMF FMF 287 SE F 35.9394SL F 274 NEPC 256 M 35.9330PC 35.9374 248 8 EWT O O 35.9438 248 27 287 M 21 35.9409 256 35.9329 28 35.9311 11 13 14 MF NE F 287 35.9371 26 SL M 250 35.9739OD O 10 293 35.9451 21 MF NW FMF E 388MF F 36.0012 N 388 FSL 36.0082 341 12 NESL 36.0039 F NEDO F 244 7 DO 262 SE M 35.9733 9 F 259 35.9557 290 35.9511 21 35.9464 24 15 25 MF W F 262 35.9303 9 HAB ne sp.; sp.; exposed Vinca Vinca sp. Vinca ne sandy loam ne sandy loam some limestone limestone rocks; silty clay loam of a lime-making operation; dominated by cedar forest, oak/maple; acidic sandstone, fi oak/maple; acidic sandstone; fi sandy loam mixed hardwoods forested slope, oak/sycamore pond; disturbed area marble blocks forest, oak/maple; acidic sandstone; fi oak/hickory; limestone boulders Moderate slope; mixed hardwoods with cedar, invasive Limestone/sandstone cliff; mixed oak/sycamore hardwoods soil with limestone soil with limestone Mixed hardwood forest, hickory/maple; stream bank, limestone; alluvial silt loam Moderate slope, mixed hardwood Steep forested slope, mixed hardwoods Limestone boulders and sandstone ledges; Limestone bluff and steep hardwood Limestone/marble outcrops facing quarry Old marble quarry; mesic; limestone/ maple/hickory, sandstone/limestone hardwoods; Moderate slope; mixed hardwood limestone outcrops limestone Habitat NE side middle of park R, RDB TN River mile 651, LDB Meades Quarry Ross Quarry SW side Burnett Creek Broad R Mile 1.5, LDB tributary to Tennessee R. Location (Continued) Date 81 23MAY201582 Hastie Park 15AUG2015 Burnett Creek Rd and 83 9MAY201684 Forks of the River NWA, French 9MAY201685 Forks of the River NWA 16AUG201586 Hilly mixed hardwood forest, On trail S of Burnett Ck Rd 16AUG2015 S of Burnett Ck Rd, unnamed Open cedar grove; moss covered rocky Open cedar grove; moss covered rocky 71 6SEP201572 Sharp Ridge Mem. Park, 10OCT2015 Sharp Ridge Mem. Park, 73 29MAR201574 Holston R Park 26OCT201675 Holston R Park, 17NOV201576 Ijams Nature Center, 17NOV201577 Ijams Nature Center, 17NOV2015 Mixed hardwood forest; karst; 78 Ijams Nature Center, 17NOV2015 Ijams Nature Center79 21OCT2015 Marie Myers Park80 21OCT2015 Dry alkaline valley; historic location Marie Myers Park Moderate slope; mixed hardwoods, Moderate slope; cedar and mixed 70 6SEP2015 Sharp Ridge Mem. Park, 87 16AUG2015 S of Burnett Ck Rd Mixed hardwood forest, hickory/maple; Appendix 1. Site LAND SNAILS OF KNOX COUNTY, TENNESSEE 21 No.Sp. -83.8526 -83.8529 -83.8287 -83.8251 -83.8334 -83.8311 -83.7975 -83.8365 -83.8278 -83.8296 -83.8592 -83.8203 -83.8198 -83.7624 -83.7647 -83.7725 -83.7710 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP MF E F 274 35.9241 14 SL SWSL F SWDF M 259 SWDF M 268 SW 35.9894 262 F 35.9972 274 36.0145 10 36.0223 14 2 5 WT Flat MDF 274 NEE FMF 35.9244 SW 259WT F 35.9295MF 335 F 13 SDO 35.9317 280 F 3 35.9269 274 M 274 19 35.9269 35.9660 6 MF NE 27 FWT 18 MF 366 O 36.0633WT 305 M SE 354MF F 36.0653 NW 28 372 36.1051 FMF SE 36.1055 518 7 F 36.1088 15 518 19 36.1064 30 27 HAB ne sandy ne ed ne sandy loam ne sandy loam ne sandy loam ne sandy loam silty clay loam alluvial silt loam hardwoods, oak hardwoods, sycamore luvial loam disturbed hardwood forest hardwood forest, oak/maple; alluvial clay loam maple; loam 40% canopy forest, oak/hickory; acid sandstone; fi wet pasture; mesic silt loam forest, oak/hickory; sandstone; fi mesic, standing water; sandstone; fi oak/hickory; sandstone boulder; fi hickory; sandstone boulder; fi loam forest, oak/hackberry; limestone, maple; stream bank; limestone; Second growth forest; entire slope covered with an unidentifi herbaceous ground cover Karst ridgetop; mixed hardwood forest, oak/hickory; loam Steep ravine; in a seep; mixed Shaded ravine; mixed hardwood forest, oak/ Sandstone boulder forest, oak/cedar; Steep slope; mixed hardwood Floodplain and edge of creek, open, Moderate slope; mixed hardwood Mature mixed hardwood forest; Steep slope; mixed hardwood forest, Steep slope; mixed hardwood forest, oak/ Habitat Rd Rd Kennedy Rds E of Ellistown Rd E of Ellistown Rd Natural Area, 1km E of Idumea Rd Natural Area, SE base Natural Area, NW side Natural Area, SE side Fordtown Rd Ck Rd Location (Continued) Date Appendix 1. 88 20NOV2016 Sevierville Pike Moderate slope; mixed hardwood 89 20NOV2016 Sevierville Pike90 9MAY201691 Burnett Ck Rd, N of 9MAY201692 Brown Mtn, NE of Burnett 15MAY2016 Mixed hardwood forest, hickory/ Brown Mtn, base at Fordtown 93 15MAY201694 Brown Mtn, base at Fordtown 2APR201695 Thorn Grove Pike and 9MAY201596 Holston R Mile 3, LDB 9MAY201597 Holston R Mile 4, LDB 9MAY201598 Holston R Mile 5, LDB 11FEB2017 Limestone cliff; some mixed 99 Love Ck Rd 12NOV2016 Limestone cliff; some mixed McAnnally Ridge, 100 Disturbed second growth forest; eroded al- 12NOV2016101 McAnnally Ridge, 5MAR2016 House Mountain State 102 19MAR2016 Moderate slope; limestone boulders; House Mountain State 103 19MAR2016 House Mountain State 104 19MAR2016 House Mountain State Site 22 AMERICAN MALACOLOGICAL BULLETIN 36 · 1 · 2018 No.Sp. -83.7232 -83.7264 -83.6898 -83.6915 -83.6970 -83.6950 -83.6938 -83.6878 -83.6961 -83.7441 -83.7447 -83.7792 -84.0414 -84.0175 -84.0203 -84.0917 -83.9160 -83.9006 Latitude/ Longitude (decimal degrees) ELV (m) CC ASP SL SEE M 274 36.0556 18 HAB WTDO MWT 259 O O 274 36.0548 317 35.9781 35.9543 5 2 1 SL S F 262 35.9515 40 WT O 207 35.9504 3 MF E FMX 290DF 35.9215 NW O F 293 323 36.0193 19 OD 36.0206 M 1 335 1 35.9488 1 DFSL M ESL 262 F NE 35.9499SL 290 M W 274 35.9492 F 35.9435 7 305MF NW 18 35.9218 F 20 415 35.9072 13 18 UR O 274 35.9493 1 ne oodplain; ne sandy loam ne sandy loam ood plain; weedy, open, ne sandy loam ne sandy loam forest, oak/beech mesic; alluvial silt loam shaly clay loam open; silt loam beech/oak/hickory open; alluvial silt loam from micaceous material sandy loam forest residential yard under a board; silty clay loam calcareous sandstone alluvial silt loam from micaceous material maple; sandstone river bluff; fi hardwoods; fi from acid sandstone hickory; acid sandstone outcrops; fi hickory; acid sandstone, fi acid sandstone; fi a greenhouse. Habitat Limestone cliff; mixed hardwoods, Steep mixed hardwood slope, oak/ Limestone/sand stone bluff; mixed Steep mixed hardwood slope, beech/ Steep mixed hardwood slope, beech/ Steep mixed hardwood slope, oaks; Parking lot in the rain UR O 317 36.0262 1 eld Gap, eld French Broad R. mile 15, RDB Broad R Mile 17.5, RDB R Mile 15, RDB Gap, E side W side N of Dry Hollow Rd Bayless Ln Location Davenport Rd, Open, disturbed grassy bank; near (Continued) Date 105 11FEB2017 Holston R mile 16, RDB Limestone bluff; mixed hardwood 106 11FEB2017 Saylor Ford Rd Wetland/fl Site 107 2APR2016108 Midway Rd and Tuckahoe Ck 2APR2016109 Seven Islands State Park 15OCT2015 At a road cut into shaly bank; Seven Islands State Park, Edge of pond; grasses, scrubby trees; 110 2APR2016 Seven Islands State Park111 15OCT2015 Seven Islands State Park112 2APR2016 Edge of pond/wetland; grasses; Seven Islands State Park, French 113 Disturbed, scrubby fl 3APR2016 Bays Mtn, French Broad 114 3APR2016 Bays Mtn, Porterfi 115 3APR2016 Bays Mtn, Porterfi 116 12FEB2017P1 Bays Mtn, Finger Ridge, 8NOV2017P2 Powell Blvd 14MAR2016 West Beaver Creek Dr,P3 12SEP2015P4 W Beaver Creek Dr and 20MAY2016P5 Mixed hardwood forest at edge of a Mud Flat Cave 13MAR2016P6 Sidewalk next to mixed hardwood Fort Dickerson Park, 26MAR2017 Parklike; mixed hardwoods; loam from Cave; limestone CV C 262 35.8875 1 Appendix 1.