Conservation, Biology, and Natural History of Crayfishes from the Southern US
2010 Southeastern Naturalist 9(Special Issue 3):63–78
Distribution and Conservation Standing of West Virginia
Crayfishes
Zachary J. Loughman1,* and Stuart A. Welsh2
Abstract - The diversity of crayfishes in West Virginia represents a transition between
the species-rich southern Appalachian faunas and the depauperate crayfish
diversity in the northeastern United States. Currently, 22 described species occur
in the state, of which 6 are given S1 status, and 3 are introduced species. One species,
Orconectes limosus (Spinycheek Crayfish) is considered extirpated within
the past decade. Imperiled species include Cambarus veteranus (Big Sandy Crayfish),
Cambarus elkensis (Elk River Crayfish), Cambarus longulus (Atlantic Slope
Crayfish), and Cambarus nerterius (Greenbrier Cave Crayfish). Three species—O.
virilis (Virile Crayfish), Orconectes rusticus (Rusty Crayfish), and Procambarus
zonangulus (Southern White River Crawfish)—have introduced populations within
the state. Procambarus acutus (White River Crawfish) occurs in bottomland forest
along the Ohio River floodplain, and is considered native. Several undescribed
taxa have been identified and currently are being described. A statewide survey was
initiated in 2007 to document the current distribution and conservation status of
crayfishes in West Virginia.
Introduction
Crayfish distributions in West Virginia are a reflection of past geologic
events (Hobbs 1969) and recent anthropogenic events (Jezerinac et al. 1995).
Since the Pleistocene Epoch, the Appalachian region has experienced large
temperature changes (Pielou 1991), observed the rise and fall of proglacial
lakes (Jacobson et al. 1988, Morgan 1994), and seen drainage modifications
(Hack 1969, Hocutt 1979) that influenced distribution patterns of aquatic
species, including crayfishes (Hocutt et al. 1978, Jezerinac et al. 1995).
The crayfish fauna of West Virginia consists of 22 species and 4 genera.
The most diverse genus is Cambarus with 13 species, followed by
Orconectes with 6 species, Procambarus with 2 species, and 1 species of
Fallicambarus (Table 1). The fauna is distributed across 4 physiographic
regions, and 5 major drainage areas (Figs. 1, 2; Table 1). From a diversity
standpoint, West Virginia’s fauna represents a transition between the diverse
southern Appalachian faunas and the depauperate crayfish diversity of the
northeastern United States.
Geographic distributions of West Virginia crayfishes were previously
reviewed by Hobbs (1969) and Jezerinac et al. (1995), but changes in
1Department of Natural Sciences and Mathematics, West Liberty University, West
Liberty, WV 26074. 2US Geological Survey, West Virginia Cooperative Fish and
Wildlife Research Unit, 322 Percival Hall, West Virginia University, Morgantown,
WV 26506. *Corresponding author - zloughman@westliberty.edu.
64 Southeastern Naturalist Vol. 9, Special Issue 3
Table 1. Distribution of West Virginia crayfishes by major river drainage and physiographic province in West Virginia (see Figs. 1, 2); Potomac River (P), Monongahela
River (M), Ohio River direct drains (O), James River (J), Kanawha River (K), Ohio River basins southwestern West Virginia (S), Ridge and Valley (RV),
Allegheny Mountains (AM), and Appalachian Plateau (AP). XH indicates historic records. Global (Gl) and state (St) conservation status ranks (WVNHP 2007) are
as follows: G5 = secure, G3 = vulnerable, G2 = imperiled, S5 = secure, S3 = vulnerable, S1 = critically imperiled, I = introduced, and U = unrankable.
Species P M O J K S RV AM AP Gl St
Cambarus (Cambarus) b. bartonii (Fabricius) (Common Crayfish) X X X X X G5 S5
C. (C.) b. cavatus Hay (Appalachian Brook Crayfish) X X X X G5 S5
C. (C.) carinirostris Hay (Rock Crawfish) X X X X X G5 S5
C. (C.) sciotensis Rhoades (Teays River Crayfish) X X X X G5 S5
C. (Hiatacambarus) chasmodactylus James (New River Crayfish) X X X G5 S3
C. (H.) elkensis Jezerinac and Stocker (Elk River Crayfish) X X G2 S1
C. (H.) longulus Girard (Atlantic Slope Crayfish) X G5 S1
C. (Jugicambarus) dubius Faxon (Upland Burrowing Crayfish) X X X X X X X X X G5 S3
C. (J.) monongalensis Ortmann (Blue Crawfish) X X X X X X X G5 S5
C. (Punticambarus) nerterius Hobbs (Greenbrier Cave Crayfish) X X G2 S1
C. (P.) robustus Girard (Big Water Crayfish) X X X X G5 S5
C. (P.) veteranus Faxon (Big Sandy Crayfish) XH X G3 S1
C. (Tubericambarus) thomai Jezerinac (Little Brown Mudbug) X X X X G5 S5
Fallicambarus (Creaserinus) fodiens (Cottle) (Digger Crayfish) X X G5 S1
Orconectes (Crockerinus) obscurus (Hagen) (Allegheny Crayfish) X X X X X X X G5 S5
O. (C.) sanbornii (Faxon) (Sanborn's Crayfish) X X X X G5 S5
O. (Faxonius) limosus (Rafinesque) (Spinycheek Crayfish) XH XH G5 S1
O. (Gremicambarus) virilis Hagen (Virile Crayfish) X X X X X G5 I
O. (Procericambarus) cristavarius Taylor (Spiny Stream Crayfish) X X X X G5 S5
O. (P.) rusticus (Girard) (Rusty Crayfish) X X X G5 I
Procambarus (Ortmannicus) acutus (Girard) (White River Crawfish) X X G5 U
P. (O.) zonangulus Hobbs and Hobbs (Southern White River Crawfish) X X G5 I
2010 Z.J. Loughman and S.A. Welsh 65
distribution warrant a revised and updated consideration of range changes
and conservation status. The majority of collecting occurred in the mid-
1970s, and only limited collecting occurred following the publication of
Jezerinac et al. (1995). Channell (2004) examined GAP models for prediction
of occurrence of C. veteranus in the southern coal fields. Jones and
Eversole (2005) investigated the life history of C. elkensis.
Recently, anthropogenic activities have altered distributions and habitat
of crayfishes. Bait-bucket introductions of nonnative crayfish species
and degradation of stream habitat and water quality from various land-use
practices have impacted native forms. Several habitat types have been
intensively studied. Z.J. Loughman (unpubl. data) surveyed the Ohio
River floodplain and determined the conservation status and natural history
of species. During this work, P. acutus was added to the state’s fauna
(Loughman 2007a). Invasive crayfishes of the Kanawha and Potomac
river systems were studied by C. Swecker (Marshall University, Huntington,
WV, unpubl. data), as well as the conservation status of O. limosus
(Swecker et al. 2010). Loughman et al. (2009) contributed distribution
records, life-history notes, and conservation concerns for West Virginia’s
fauna. Since the Jezerinac et al. (1995) treatise on West Virginia's crayfish
fauna was published, Taylor (2000) described O. cristavarius, previously
Figure 1. West Virginia physiographic provinces.
66 Southeastern Naturalist Vol. 9, Special Issue 3
known in West Virginia as O. spinosus, and Thoma and Jezerinac (1999)
elevated C. carinirostris to species status.
Threats to native species include nonnative species, reduced water quality,
and habitat degradation, which decrease population size and reduce distribution
of wide-ranging species. Species with small ranges can be extirpated since
source populations cannot be reintroduced after improvements (Lodge et al.
2000a, b). In West Virginia, two species with limited ranges are vulnerable
to mineral extraction: C. elkensis, an endemic restricted to the headwaters of
the Elk River, and C. nerterius, an endemic restricted to the karst system of the
Greenbrier River drainage. Bait-bucket introductions of crayfishes from other
states and across drainages within West Virginia represent pressing concerns
for native crayfish stocks. Bait-bucket introductions likely explain three nonnative
crayfishes in West Virginia: P. zonangulus, O. rusticus, and O. virilis;
the latter two are considered invasive. Orconectes limosus, a species with
restricted range in the Potomac River drainage, was possibly extirpated following
the introduction of O. virilis (Loughman et al. 2009, Swecker et al. 2010).
Populations of C. veteranus, a species with a restricted range in southern West
Virginia, were likely extirpated from habitat degradation associated with mining
(Jones et al. 2010).
Figure 2. Major West Virginia river basins.
2010 Z.J. Loughman and S.A. Welsh 67
Methods
This paper revises the distribution and conservation status of West Virginia
crayfishes. We conducted a thorough review of the research on the crayfishes of West Virginia. Conservation status of West Virginia’s 22 species is
considered at the global and state level (Table 1). Species distributions are presented
under genera headings, followed by conservation status and concerns.
Additionally, we summarize impacts of water quality and habitat modification
that pose current threats to crayfish populations in West Virginia.
Results and Discussion
Composition of fauna
Genus Cambarus. Cambarus is the most diverse genus within West Virginia
(13 species, 59% of described species). Five subgenera of Cambarus
occur within the state’s borders (i.e., Cambarus, Hiatacambarus, Jugicambarus,
Puncticambarus, and Tubericambarus).
The subgenus Cambarus includes C. bartonii bartonii, C. bartonii
cavatus, C. carinirostris, and C. sciotensis. Cambarus b. bartonii occurs
throughout the Atlantic Slope (Potomac and James River systems; Jezerinac
et al. 1995). Habitats preferred by C. b. bartonii include headwater streams
and seeps. Currently C. b. bartonii is stable in West Virginia (Table 1).
Cambarus b. cavatus currently is under taxonomic review by several investigators,
and likely will be elevated to species status (Roger Thoma, Ohio
State University, Columbus, OH, pers. comm.). Several morphological characters,
as well as geographic barrier to gene flow, separate this taxon from C.
b. bartonii. Cambarus b. cavatus occurs in headwaters streams and wetlands
throughout central and southern portions of the Ohio River direct drains,
western portions of the Kanawha River, and southwestern Ohio River basins.
Unlike C. b. bartonii, C. b. cavatus often burrows in seepage wetlands and
hardpan banks associated with small streams.
Cambarus carinirostris occurs in the Monongahela, central portions
of the Kanawha, and north-central regions of the Ohio River direct drains
(Jezerinac et al. 1995). The type locality for this species is Gandy Creek,
Osceola, Randolph County (Faxon 1914). Headwater streams are typical
C. carinirostris habitats, but the species also occurs in larger streams in
the absence of large cambarids. Schwartz and Meredith (1960), as well as
the authors, have found C. carinirostris to be common in the Cheat River
watershed of the Monongahela River drainage. Individuals from Monongahela
River populations differ phenotypically from those in Ohio River direct
drains (Loughman et al. 2009). The extent of this difference is currently being
investigated by the primary author. Within West Virginia, populations of
C. carinirostris are stable (Table 1)
Cambarus sciotensis is the largest member of the subgenus Cambarus in
West Virginia. Unlike the previous 3 taxa of secondary burrowers, C. sciotensis
is a tertiary burrower of moderate to large streams, and occurs in the
68 Southeastern Naturalist Vol. 9, Special Issue 3
southwestern Ohio River basins and the Kanawha River system (Jezerinac
et al. 1995). Within these basins, C. sciotensis dwells in mainstems unless
large sympatric crayfish are present. When this is the case, C. sciotensis
often reaches highest densities in smaller tributaries (Z. Loughman, pers. observ.).
Cambarus sciotensis in West Virginia, and throughout its range, likely
represents a species complex. Several distinct phenotypes of this species are
found throughout West Virginia, and future work will likely recognize these
phenotypes as distinct taxa (Loughman et al. 2009).
The subgenus Hiaticambarus includes 3 taxa in West Virginia: C. chasmodactylus,
C. elkensis, and C. longulus (Jezerinac et al. 1995). All members
of the subgenus are tertiary burrowers. The most prevalent of these taxa, C.
chasmodactylus, was described from the upper Greenbrier River drainage,
Randolph County, WV (James 1966). Cambarus chasmodactylus inhabits
streams in the Greenbrier River system across high and low elevations, and
is the largest native crayfish in West Virginia. Currently the species is given
a status of G5 and S3 (Table 1). Surveys performed in summer 2008 by the
authors indicate stable populations within the Greenbrier River drainage.
Cambarus elkensis, an endemic of the Elk River, occurs in tributaries
and mainstem sections of the upper Elk River, where it burrows through
loose cobble and under rock slabs. Currently, the region harboring C. elkensis
is relatively undisturbed (Z. Loughman and S. Welsh, pers. observ.).
The population is vulnerable to environmental perturbations given the small
restricted range (Jezerinac et al. 1995, Loughman et al. 2009). Reasons for
range restriction of this species are unknown, but no apparent physical factors
prevent range expansion to the western portions of the basin. Currently,
the status of C. elkensis is listed as G2 and S1 (Table 1).
Cambarus longulus has the most restricted range within West Virginia of
any Hiaticambarus, occurring in the small portion of the James River system
that lies in West Virginia (Jezerinac et al. 1995). Loughman et al. (2009)
commented on conservation concern for this species, specifically the Potts
Creek populations. Survey efforts by the authors during the summer of 2008
found that this species is stable throughout the basin. Cambarus longulus
appear to be resistant to anthropogenic disturbances. Densities of this species
were highest in Sweet Springs Creek, the most degraded stream present
in the drainage. Currently C. longulus is listed as S1 due to the small area
occupied by this taxon within the state borders (Table 1).
West Virginia has two members of the subgenus Jugicambarus: C. dubius
and C. monongalensis. Cambarus dubius was described from Terra Alta, WV
(Faxon 1914). Several phenotypes of this primary burrower occur in West
Virginia, and it will likely be elevated to species status in the future. Individuals
from populations in the Allegheny Mountains have orange carapaces
with cream venters, whereas individuals from other portions of the state have
orange venters with black dorsums and orange chelae or blue carapaces with
orange chelae. Monroe County individuals are entirely blue with truncated
chelae; populations in reaches of the Kanawha River near the confluence
2010 Z.J. Loughman and S.A. Welsh 69
with the Ohio are also blue. Habitats preferred by this species are varied,
ranging from high-elevation wetlands to road side ditches in the plateau.
Currently this taxon appears to be stable in West Virginia (Table 1).
Cambarus monongalensis is disjunctively distributed in West Virginia,
with one population on the Allegheny Plateau and another at higher elevation
in the Allegheny Mountains (Jezerinac et al. 1995). Cambarus monongalensis
occupies habitats similar to C. dubius, but the two species rarely occur
sympatrically. Currently C. monongalensis is listed as S3 (Table 1). This
listing likely should be increased given a high frequency of occurrence in a
wide range of habitats (Z. Loughman, pers. observ.).
Puncticambarus is represented in West Virginia by C. nerterius, C. robustus,
and C. veteranus. Cambarus nerterius is endemic to the Greenbrier
River drainage and is not found outside of WV borders (Jezerinac et al.
1995). The species occurs in caves in the karst areas throughout southern
portions of the Greenbrier River basin. It is the only known troglobytic crayfish occurring in West Virginia. Very little is known about the biology of C.
nerterius, making future efforts to study the natural history of this species a
priority. Currently, C. nerterius is listed as G2 (very rare and imperiled) and
S1 (Table 1).
Cambarus robustus occurs throughout the Kanawha River basin, southwestern
Ohio River basins, and central portions of the Ohio River direct
drains. As the predominant large cambarid of West Virginia, it is locally
common in larger ordered streams. Preferred habitats include stream pools,
leaf packs, and slab boulders. Currently, populations of C. robustus are
stable in West Virginia (Table 1).
Cambarus veteranus occurs in two geographically separate populations
in West Virginia, with one population in the Bluestone River system and a
second in the Guyandotte River system. Unlike C. robustus, C. veteranus is
not stable, and is likely the most imperiled crayfish in the state. Recent survey
efforts have been unsuccessful in locating individuals from sites that
historically harbored this species. Impacts to the region include mining,
straight piping, and channelization. Future efforts will focus on determining
the resident status of this species. Mining efforts in the region are the
primary means of imperilment for this species. The type locality of C. veteranus
is Indian Creek, Baileysville, Wyoming County (Faxon 1914).
Cambarus thomai is the sole member of the subgenus Tubericambarus in
West Virginia. This primary burrower is common in bottomlands associated
with the Ohio River, and throughout central portions of the state. Wetlands,
marshes, and roadside ditches are additional habitats utilized by C. thomai.
Basins holding C. thomai populations include Ohio River direct drains, Kanawha
River system, and southwestern Ohio River basins. Cambarus thomai
was recently discovered in Moncove Lake, Monroe County, the farthest east
the species is known to occur in West Virginia (Z. Loughman, pers. observ.).
Recent survey efforts have resulted in expansion of its known range eastward
to the foothills of the Allegheny Mountains. This taxon has not been found
70 Southeastern Naturalist Vol. 9, Special Issue 3
in high elevations (>762 m); introduced populations have been documented
in western Maryland (Loughman 2007b).
Genus Orconectes. Four native orconectids historically occurred in West
Virginia, comprising 18% of West Virginia’s crayfish fauna (Jezerinac et al.
1995). All Orconectes in West Virginia are tertiary burrowers, and include
three subgenera (Crockerinus, Faxonius, and Procericambarus). Crockerinus
is the most diverse subgenus, with two species occurring in West
Virginia (i.e., O. sanbornii and O. obscurus). Orconectes sanbornii, a Teays
River relict, is present throughout portions of the Kanawha River system,
southwestern Ohio River Basins, and southern portions of the Ohio River
direct drains. Throughout this region, O. sanbornii occurs in small and large
streams. Habitat use in streams include pools, eddies, vegetative beds, and
leaf packs. Hobbs and Fitzpatrick (1962) described O. s. erismorophorous
from Cranes Nest Creek, Wirt County. Jezerinac et al. (1995) questioned the
validity of this subspecies, indicating problems were present in determining
identity using morphological traits. Taylor et al. (2007) did not recognize
Orconectes s. erismorophorous within a list of North American crayfishes;
however, further study is warranted to determine specific status.
The distribution of O. obscurus in West Virginia, except Greenbrier River
populations, is allied with the Potomac and ancient Pittsburgh River systems
(Taylor and Hardman 2002). Orconectes obscurus occurs throughout the
Potomac River drainage, Monongahela River drainage, higher elevations
in the Greenbrier River drainage, and central and northern portions of the
Ohio River direct drains. Orconectes obscurus is ubiquitous in moderate to
large stream systems in these regions. Both O. sanbornii and O. obscurus are
stable in West Virginia (Table 1).
Faxonius is represented in West Virginia by a single species, O. limosus.
Recent survey efforts by the authors and Swecker et al. (2010) have been unsuccessful
in securing specimens. Extreme eastern sections of West Virginia’s
portion of the Potomac were historically occupied by this species (Jezerinac et
al. 1995). This region has recently been invaded by the nonnative crayfish, O.
virilis. Both Swecker et al. (2010) and Z. Loughman visited all known historic
sites, as well as sites possessing habitat parameters unique to O. limosus. No
specimens were found; O. virilis was the only crayfish captured at all sites. In
neighboring Maryland, O. virilis has been responsible for range reduction in
O. limosus as well (Schwartz et al. 1963). Given the amount of effort invested
in securing specimens, and the number of individual efforts, Loughman et al.
(2009) considered the species as extirpated in West Virginia.
Orconectes cristavarius is the single member of the subgenus Procericambarus
within West Virginia. This species is prevalent throughout the
southwestern Ohio River basins and James River drainage, and sporadically
distributed throughout portions of the Kanawha River system. Prior to the
species description (Taylor 2000), this taxon was referred to as O. spinosus
(Jezerinac et al 1995). Habitats occupied by C. cristavarius include detritus
beds, pool thalwegs, and slab boulders in medium to large-sized streams.
2010 Z.J. Loughman and S.A. Welsh 71
Genus Fallicambarus. Fallicambarus is represented by a single species
in West Virginia, F. (Creaserinus) fodiens (Jezerinac et al. 1995). This
species is a primary burrower limited to the preglacial Marietta River valley
situated in the vicinity of the Kanawha River/ Ohio River confluence
(Jezerinac and Stocker 1987). Only four populations of this species have
been found in West Virginia, all of which occur in large tracts of either bottomland
forest or bottomland swamps. Greenbottom Wildlife Management
Area currently holds the state’s most robust populations. Fallicambarus
fodiens is a primary burrower during the spring, summer, and fall season.
During late winter and early spring, ephemeral wetlands are used by this
species in West Virginia (Z.J. Loughman, pers. observ.). Causes of imperilment
are unknown at this time, but could include competitive exclusion
with native burrowing species (such as C. thomai), habitat degradation,
wetland draining, and increased industrial use of the region (Z.J. Loughman,
pers. observ.). Fallicambarus fodiens has conservation status listings
as G5 and S1 (Table 1).
Genus Procambarus. The genus Procambarus is represented in West
Virginia by two closely related species: P. acutus and P. zonangulus. Procambarus
acutus is the single native Procambarus species occurring in West
Virginia. This species was recently discovered in a bottomland forest along
the Kanawha River floodplain near Point Pleasant (Loughman 2007a). Populations
within this wetland system are robust. Additional populations were
discovered in a wetland associated with a residential park and a roadside
ditch. Further survey efforts in the region are needed to document the species
range in West Virginia.
Procambarus zonangulus was first documented in Sleepy Creek Wildlife
Management Lake and two isolated wetlands associated with fish hatcheries
in Berkeley County. These collections represent the first record of the species
in West Virginia (Z. Loughman, pers. observ.). These populations likely
represent a bait-bucket introduction, and appear to be isolated to the lake (Z.
Loughman, pers. observ.). This species was recently added to Maryland’s
state fauna, indicating a possible point of entry into West Virginia (Kilian et
al. 2010)
Conservation concerns
Impacts to West Virginia crayfish are regional in scope. A discussion
of the major conservation concerns include invasive species, water quality
impacts and habitat degradation associated with land development, urban
sprawl, and extractive industries.
Invasive crayfish species. Three introduced crayfishes occur in West
Virginia, of which two are considered invasive. Populations of P. zonangulus
in the eastern panhandle appear restricted to Sleepy Creek Lake and
two wetlands in Morgan County. Given the potential for expansion of these
introduced populations, additional studies are warranted in this region. Both
invasive species in West Virginia are orconectids, O. rusticus and O. virilis,
and likely are a result of bait-bucket introductions.
72 Southeastern Naturalist Vol. 9, Special Issue 3
Procambarus clarkii (Girard) (Red Swamp Crayfish) was previously
reared within ponds for aquaculture production in Mason County, and some
of these populations have persisted after termination of aquaculture efforts.
These populations have not escaped from the ponds, and because of this, are
not considered part of West Virginia’s fauna by the authors. Procambarus
clarkii, however, is expected to invade eastern panhandle waterways via
introduced populations persisting in Maryland (Kilian et al. 2010).
Orconectes rusticus was first documented in West Virginia in Fourpole
Creek by Lawton (1979). Other surveys documented O. rusticus in the Little
Kanawha River basin (Jezerinac et al. 1995), Kanawha River mainstem
(C. Swecker, pers. comm.), Ohio River backwaters in Marshall and Wetzel
counties (Z.J. Loughman, pers. observ.), and several impoundments throughout
Twelvepole Creek basin (Z.J. Loughman, unpubl. data). Fourpole Creek
populations have received the most attention, given their close proximity to
Marshall University (Loughman et al. 2009).
The invasion of O. virilis has primarily occurred in the eastern panhandle
(Schwartz et al. 1963), though the species initially was documented in 1970
from southern portions of the New River in Summers County (Jezerinac et
al. 1995). Jezerinac et al. (1995) first documented O. virilis in the eastern
Panhandle. Also, O. virilis was recorded in Point Pleasant and Ohio River
backwaters in Saint Mary’s, Pleasant County (Z.J. Loughman, pers. observ.).
Surveys by the author have also located several additional populations in
state impoundments, including Summerville Lake, Nicholas County and
Moncove Lake, Monroe County.
Invasive orconectid species are known to eliminate native species through
hybridization or competitive exclusion (Lodge 2000a, b). Hybridization associated
with these taxa has been documented elsewhere (Kilian et al. 2010,
Taylor and Schuster 2005), but does not appear to be a conservation concern
in West Virginia given the lack of hybrids in collections. Competitive
exclusion of native species by nonnative invasive species possibly explains
extirpation or population reduction of some native crayfish populations. In
2007, the authors surveyed West Virginia’s portion of the Potomac River
drainage for O. limosus. Neither O. limosus nor sympatric O. obscurus populations
were observed and may be extirpated. Habitat, historically including
native species, was provided by boulders and sandy bottomed pools. These
streams were solely inhabited by O. virilis.
During our surveys, O. obscurus was collected from Justicia sp. (water
willow) beds and detrital beds in pools, but none were collected from beneath
cover objects in the stream. Orconectes obscurus often took refuge
under boulders and small slabs while escaping from collectors’ pursuit, but
immediately were pushed into the open by O. virilis. Our general observations
support a competitive exclusion hypothesis, but further quantitative
studies are needed to examine the importance of competition in shaping
crayfish communities in West Virginia. Orconectes rusticus in Fourpole
Creek possibly eliminated O. sanbornii and C. b. cavatus populations via
competition (Loughman et al. 2009).
2010 Z.J. Loughman and S.A. Welsh 73
Nonnative crayfish populations occur in many impoundments within West
Virginia, possibly as a result of bait-bucket introduction. Further studies are
needed to address the impacts of nonnative crayfishes on native populations
within impoundments. Once introduced into an impoundment, a crayfish
could possibly disperse throughout the watershed, but we have not observed
dispersal in most nonnative populations within small impoundments. Highgradient
stream habitats, such as those found throughout West Virginia, may
inhibit dispersal of some nonnatives from impoundments (Loughman 2010).
Maude and Williams (1983) found that large O. virilis lack the ability to hold
position in high-velocity microhabitats. Future studies, however, are needed
to document the distribution and dispersal abilities of nonnative crayfishes
in West Virginia.
Development and urban sprawl. Development and urban sprawl are not often
considered as major impacts to West Virginia waterways, but have recently
become major conservation concerns in some regions. Areas impacted by
development include Potomac River drainage streams and rivers in Jefferson,
Berkeley, and Morgan counties, associated with urban sprawl from Washington,
DC. Land and stream manipulation linked with urban sprawl and land
development increase stream sedimentation and water temperatures, destruction
of riparian corridors, channelization, and in extreme cases, stream course
alterations. Several streams in the Potomac and Shenandoah basins have witnessed
enough manipulation to completely alter natural stream process and
flows. The invasive crayfish, O. virilis, has thrived in the disturbed environments
of the Potomac River watershed in West Virginia. The combination of
increased development and invasive species work synergistically to extirpate
native crayfish stocks (Lodge 2000a, b). Other regions experiencing elevated
development rates include Morgantown, Monongalia County and the Teays
Valley corridor between Charleston and Huntington. The ultimate impact of
development on crayfishes in these regions needs further study.
Extractive industries. Extractive industries in West Virginia, such as coal,
oil, gas, and timber, have impacted crayfish habitats. Stream sedimentation
increases due to land use disturbance associated with extractive industries,
and negatively impacts aquatic invertebrates (Angradi 1999, Schofield et al.
2004, Waters 1995, Wood and Armitage 1997). Stream sedimentation alters
the composition of benthic stream habitat (Waters 1995), which is the primary
habitat of stream crayfishes. In addition to stream sedimentation, coal
mining causes water quality degradation, such as changes in pH, metals, and
specific conductivity (Dick et al. 1983, Pond et al. 2008, Starnes 1983). The
level of mining disturbance on crayfish habitats is, in part, linked to the type
of mining operation (e.g., underground mining or surface mining, including
mountaintop-removal mining).
The impact of acidification on crayfish physiology is not well understood.
Gallaway and Hummon (1991) studied the impact of acidification from acid
mine drainage on C. b. cavatus in southwestern Ohio. Crayfishes not acclimated
to acidic conditions died quickly following exposure to acidic waters.
74 Southeastern Naturalist Vol. 9, Special Issue 3
Cambarus b. cavatus collected from acidic streams tolerated increases in
acidification during inter-molt periods, but perished during ecdysis events
under elevated acidification.
In addition to acid mine drainage, acid precipitation is another source
of stream acidification in poorly buffered watersheds of West Virginia. In
a study of acidified streams, Seiler and Turner (2004) found lower growth
rates and higher abundances of crayfish, possibly owing to a lack of fish
predators. Observations by the authors indicate that acidification of streams
has adversely effected crayfishes in West Virginia. Acid mine drainage is
prevalent throughout northern portions of the Cheat River basin. Schwartz
and Meredith (1960) surveyed this region during the 1950s and found several
streams with AMD that lacked crayfish populations. These same streams
were surveyed in 2008 by the authors. Acid mine drainage was prevalent,
and zero crayfishes were collected (Z.J. Loughman, unpubl. data).
Surface mining, including contour surface mining and mountaintopremoval
mining, and the associated impacts has altered physical habitat and
the chemical integrity of surface and groundwater (Rogowski et al. 1977).
Fragile seep environments with specific groundwater hydrologies are fractured
(Fields 2003, Vörösmarty et al. 2000). These habitats are important for
montane burrowing crayfish populations (Jezerinac et al. 1995). The mining
process often alters stream courses, and small headwaters are buried during
mountaintop-removal mining (Hartman et al. 2005, Pond et al. 2008). Secondary
and primary burrowing crayfish species occupying forested ravines
in proximity to surface mines are buried during this process. In addition to
impacts at the mining site, the loss of headwater streams during mountaintop-
removal mining has downstream impacts on habitat, water quality, and
energy inputs (Vannote et al. 1980). The range of C. veteranus occurs within
the heavily-mined region of a few southern counties, and the population is
possibly extirpated. Channell (2004) indicated that historic sites of C. veteranus
display elevated siltation, stream manipulation, and channel scouring
from flooding.
Mining operations influence crayfish habitats through leachates associated
with overburden piles and through removal of vegetation prior to
mining. Overburden piled in the headwaters of drainages leach coal fines and
toxic compounds downstream, where they have cumulative effects within
the watershed. Many mining efforts have a series of slurry ponds situated
in headwaters, which catch heavy metals and chemicals. Slurry ponds can
leach chemicals into the watershed or experience catastrophic dam failure,
resulting in inundation of the basin with concentrated coal toxins (Frey et
al. 2001). Much of the vegetation of the region is clear cut prior to mining
efforts. Vegetation represents an important means of water absorption during
increased periods of precipitation. The removal of vegetation and the
reduced water retention rates of overburden lead to an increase in stream
flows, flooding, and silt loads (Hanna 1964, Swift and Swank 1981, Webster
et al. 1992).
2010 Z.J. Loughman and S.A. Welsh 75
Acknowledgments
We would like to thank the many West Liberty University students and Oglebay
Institute interns that assisted us in the field. Financial support for West Virginia
Crayfish surveys was provided by the West Virginia Division of Natural Resources.
Additional Financial support was provided by Oglebay Institute and West Liberty
University. Reference to trade names does not imply endorsement of commercial
products by the US Government. The publication of this manuscript was supported,
in part, by the US Geological Survey Cooperative Research Unit Program, including
the West Virginia Cooperative Fish and Wildlife Research Unit.
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