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. Literature Cited Angradi, T.R. 1999. Fine sediment and macroinvertebrate assemblages in Appalachian Streams: A field experiment with biomonitoring applications. Journal of the North American Benthological Society 18:49–66. Channell, K.B. 2004. Implementation of a spatial-temporal focus to predict habitat locations and distribution of Cambarus veteranus. M.Sc. Thesis. Marshall University, Huntington, WV. 88 pp. Dick, W.A., J.V. Bonta, F. Haghiri, and J.R. Page. 1983. Stream water quality of two small watersheds as affected by surface coal mining. Journal of Environmental Quality 12:351–358. Faxon, W. 1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zoology, with descriptions of new species and subspecies to which is appended a catalogue of the known species and subspecies. Memoirs of the Museum of Comparative Zoology at Harvard College 40:351–427. Fields, S. 2003. The Earth's open wounds: Abandoned and orphaned mines. Environmental Health Perspectives 111:154–161. Frey, K.J., D.P. Michaelson, and W.L. Davis. 2001. Impacts of the Martin County coal slurry spill on fishery resources in eastern Kentucky streams: A case study. Southeastern Association of Fish and Wildlife Agencies 55:95–104. Gallaway, M.S., and W.D. Hummon. 1991. Adaptation of Cambarus bartonii cavatus to Acid Mine Polluted Waters. Ohio Journal of Science 91:167–171. Hack, J.T. 1969. The area, its geology: Cenozoic development of the southern Appalachians. Pp. 1–17, In P.C. Holt (Ed.). The Distributional History of the Biota of the Southern Appalachians Part 1: Invertebrates. Research Division Monograph 1, Virginia Polytechnic Institute, Blacksburg, VA. Hanna, G.P. 1964. The relation of water to strip-mine operation. Ohio Journal of Science 64:120–124. Hartman, K.J., M.D. Kaller, J.W. Howell, and J.A. Sweka. 2005. How much do valley fills influence headwater streams? Hydrobiologia 532:91–102. Hobbs, H.H., Jr. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pp. 93–178, In P.C. Holt (Ed.). The Distributional History of the Biota of the Southern Appalachians Part 1: Invertebrates. Research Division Monograph 1, Virginia Polytechnic Institute, Blacksburg, VA. Hobbs, H.H., and J.F. Fitzpatrick, Jr. 1962. A new crayfish of the Propinquus group of the genus Orconectes from the Ohio drainage system in West Virginia. Proceedings of the Biological Society of Washington 75:207–214. Hocutt, C.H. 1979. Drainage evolution and fish dispersal in the central Appalachians: Summary. Bulletin of the Geological Society of America 90:129–130. 76 Southeastern Naturalist Vol. 9, Special Issue 3 Hocutt, C.H., R.F. Denoncourt, and J.R. Stauffer, Jr. 1978. Fishes of the Greenbrier River drainage, West Virginia, with drainage history of the central Appalachians. Journal of Biogeography 5:59–80. Jacobson, R.B., D.P. Elston, and J.W. Heaton. 1988. Stratigraphy and magnetic polarity of the high-terrace remnants in the upper Ohio and Monongahela Rivers in West Virginia, Pennsylvania, and Ohio. Quaternary Research 29:216–232. James, H.A. 1966. Range and variations of subspecies of Cambarus longulus (Decapoda, Astacidae). Proceedings of the United States National Museum 119:1–24. Jezerinac, R.F., and G.W. Stocker. 1987. Fallicambarus (Creaserinus) fodiens in West Virginia: A new state record. Ohio Journal of Science 87:46–47. Jezerinac, R.F., G.W. Stocker, and D.C. Tarter. 1995. The Crayfishes (Decapoda: Cambaridae) of West Virginia. Bulletin of the Ohio Biological Survey, New Series 10:1–193. Jones, D.R., and A.G. Eversole. 2005. Life-history characteristics of the Elk River Crayfish, Cambarus elkensis, in Left Fork Holly River, Hacker Valley, West Virginia. Final Report, West Virginia Division of Natural Resources, West Virginia Nongame Wildlife and Natural Heritage Program, Elkins, WV. 94 pp. Jones, T.G., K.B. Channel1, S.E. Collins, J. Enz, and C.M. Stinson. 2010. Possible extirpation of Cambarus veteranus (Big Sandy Crayfish) from West Virginia. Southeastern Naturalist 9(Special Issue 3):165–174. Kilian, J.V., A.J. Becker, S.A. Stranko, M. Ashton, R.J. Klauda, J. Gerber, and M. Hurd. 2010. The status and distribution of Maryland Crayfishes. Southeastern Naturalist 9(Special Issue 3):11–32. Lawton, S.M. 1979. A taxonomic and distributional study of crayfishes (Decapoda: Cambaridae) of West Virginia with diagnostic keys to the species of the genera. M.Sc. Thesis. Marshall University, Huntington, WV. Lodge, D.M., C.A. Taylor, D.M. Holdich, and J. Skurdal. 2000a. Nonindigenous crayfishes threaten North American freshwater biodiversity: Lessons from Europe. Fisheries 25(8):7–20. Lodge, D.M., C.A. Taylor, D.M. Holdich, and J. Skurdal. 2000b. Reducing impacts of exotic crayfish introductions: New policies needed. Fisheries 25(89):21–23. Loughman, Z.J. 2007a. First record of Procambarus (Ortmannicus) acutus (White River Crayfish) in West Virginia, with notes on its natural history. Northeastern Naturalist 14:495–500. Loughman, Z.J. 2007b. Cambarus (T.) thomai in Maryland: Conservation implications of an introduced burrowing crayfish population. Freshwater Crayfish News 29:3. Loughman, Z.J. 2010. Crayfishes of Maryland: Conservation and natural history. Southeastern Naturalist 9(Special Issue 3):33–62. Loughman, Z.J. In press. Crayfishes of the West Virginia Portion of the Ohio River Floodplain: Conservation and Natural History. AuthorHouse Press, Bloomington, IN. Loughman, Z.J., T.P. Simon, and S.A. Welsh. 2009. West Virginia crayfishes (Decapoda:Cambaridae): Observations on distribution, natural history, and conservation. Northeastern Naturalist 16(2):225–238. Maude, S.H., and D.D. Williams. 1983. Behavior of crayfish in water currents: Hydrodynamics of eight species with reference to their distribution patterns in southern Ontario. Canadian Journal of Fisheries and Aquatic Science 40:68–77. 2010 Z.J. Loughman and S.A. Welsh 77 Morgan, S.A. 1994. Depositional facies associated with Lake Monongahela. M.Sc. Thesis. West Virginia University, Morgantown, WV. 135 pp. Pielou, E.C. 1991. After the Ice Age: The Return of Life to Glaciated North America. University of Chicago Press, Chicago, IL. 376 pp. Pond, G.J., M.E. Passmore, F.A. Borsuk, L. Reynolds, and C.J. Rose. 2008. Downstream effects of mountaintop coal mining: Comparing biological conditions using family- and genus-level macroinvertebrate bioassessment tools. Journal of the North American Benthological Society 27:717–737. Rogowski, A.S., H.B. Pionke, and J.G. Broyan. 1977. Modeling the impact of strip mining and reclamation processes on quality and quantity of water in mined areas: A review. Journal of Environmental Quality 6:237–244. Schofield, K.A., C.M. Pringle, and J.L. Meyer. 2004. Effects of increased bedload on algal- and detrital-based stream food webs: Experimental manipulation of sediment and macroconsumers. Limnology and Oceanography 49:900–909. Schwartz, F.J., and W.G. Meredith. 1960. Crayfishes of the Cheat River watershed, West Virginia and Pennsylvania. Part I. Species and localities. Ohio Journal of Science 60:40–54. Schwartz, F.J., R. Rubleman, and J. Allison. 1963. Ecological population expansion of the introduced crayfish Orconectes virilis. Ohio Journal of Science 63:265–273. Seiler, S.M., and A.M. Turner. 2004. Growth and population size of crayfish in headwater streams: Individual- and higher-level consequences of acidification. Freshwater Biology 49:870–881. Starnes, L.B. 1983. Effects of surface mining on aquatic resources in North America. Fisheries 8(6):2–4. Swecker, C.D., T.G Jones, K. Donahue II, D. Mckinney, and G.D. Smith. 2010. The extirpation of Orconectes limosus (Spinycheek Crayfish) populations in West Virginia. Southeastern Naturalist 9(Special Issue 3):155–164. Swift, L.W., Jr., and W.T. Swank. 1981. Long-term responses of streamflow following clearcutting and regrowth. Hydrological Sciences Bulletin 26:245–256. Taylor, C.A. 2000. Systematic studies of the Orconectes juvenilis complex, with description of two new species. Journal of Crustacean Biology 20(1):132–152. Taylor, C.A., and M. Hardman. 2002. Phylogenetics of the Crayfish Subgenus Crockerinus, Genus Orconectes (Decapoda: Cambaridae), Based on Cytochrome Oxidase I. Journal of Crustacean Biology 22:874–881. Taylor, C.A., and G.A. Schuster. 2005. Crayfishes of Kentucky. Illinois Natural History Survey Bulletin 28. 220 pp. Taylor, C.A., G.A. Schuster, J.E. Cooper, R.J. DiStefano, A.G. Eversole, P. Hamr, H.H. Hobbs III, H.W. Robsion, C.E. Skelton, and R.F. Thoma. 2007. A reassessment of the conservation status of crayfishes of the United States and Canada after 10+ years of increased awareness. Fisheries 32:372–389. Thoma, R.F., and R.F. Jezerinac. 1999. The taxonomic and zoogeography of Cambarus bartonii carinirostris (Crustacea:Decapoda: Cambaridae). Proceedings of the Biological Society of Washington 112:97–105. Vannote, R.L., G.W. Minshall, K.W. Cummins, J.R. Sedell, and C.E. Cushing. 1980. The river continuum concept. Canadian Journal of Fisheries and Aquatic Sciences 37:130–137. Vörösmarty, C.J., P. Green, J. Salisbury, and R.B. Lammers. 2000. Global water resources: Vulnerability from climate change and population growth. Science 289(5477):284–288. 78 Southeastern Naturalist Vol. 9, Special Issue 3 Waters, T.F. 1995. Sediment in streams: Sources, biological effects, and control. American Fisheries Society Monograph 7, Bethesda, MD. 251 pp. Webster, J.R., S.W. Golladay, E.F. Benfield, J.L. Meyer, W.T. Swank, and J.B. Wallace. 1992. Catchment disturbance and stream response: An overview of stream research at Coweeta Hydrologic Laboratory. Pp. 231–252, In P.J. Boon, P. Calow, and G.E. Petts (Eds.). River Conservation and Management. John Wiley and Sons,Edinburgh, Scotland, UK. 484 pp. Wood, P.J., and P.D. Armitage. 1997. Biological effects of fine sediment in the lotic environment. Environmental Management 21:203–217. West Virginia Natural Heritage Program (WVNHP). 2007. Rare, threatened, and endangered animals. West Virginia natural Heritage Program, West Virginia Division of Natural Resources, February 2007. Available online at http://www.wvdnr. gov/wildlife/documents/animals2007.pdf. Accessed 15 January 2009.