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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.
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 -
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.
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
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).
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).
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66 Southeastern Naturalist Vol. 5, No. 1
Appendix 1. List of species and their locations.
Species 1 2 3456789101112131415
Oligyra orbiculata Say, 1818 X X X X
Carychium exile Lea, 1842 X X X X X
Carychium nannodes Clapp, 1905 XX
Cochlicopa morseana (Doherty, 1878) X X
Gastrocopta contracta (Say, 1822) X X X X X
Gastrocopta pentodon (Say, 1821) X X
Vertigo gouldii (Binney, 1843) X
Strobilops aeneus Pilsbry, 1926 X X X X X
Strobilops labyrinthica (Say, 1817) X X X
Philomycus carolinianus (Bosc, 1802) X X X X X
Anguispira alternata (Say, 1816) XX
Discus patulus (Deshayes, 1830) X
Helicodiscus parallelus (Say, 1817) X X
Punctum minutissimum (Lea, 1841) X X X X
Punctum smithi Morrison, 1935 X
Punctum vitreum (Baker, 1930) X X X
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
Species 1 23456789101112131415
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
Euconulus dentatus (Sterki, 1893) X X X X X X
Haplotrema concavum (Say, 1821) X X XXXX X XXXX
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
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
Species 1 23456789101112131415
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