2009 NORTHEASTERN NATURALIST 16(2):225–238 West Virginia Crayfishes (Decapoda: Cambaridae): Observations on Distribution, Natural History, and Conservation Zachary J. Loughman1,2,*, Thomas P. Simon3, and Stuart A. Welsh4 Abstract - West Virginia’s crayfishes have received moderate attention since publication of Jezerinac et al.’s (1995) monograph of the state fauna. Survey efforts were initiated over the summers of 2006 and 2007 to gather voucher material for the Indiana Biological Survey’s Crustacean Collection. These collections have provided new information regarding the distribution, natural history, life history, taxonomy, and conservation status of Cambarus (Cambarus) carinirostris, C. (C.) bartonii cavatus, C. (C.) sciotensis, C. (Hiaticambarus) chasmodactylus, C. (H.) elkensis, C. (H.) longulus, C. (Jugicambarus) dubius, C. (Puncticambarus) robustus, Orconectes (Procericambarus) cristavarius, and O. (P.) rusticus. Orconectes (Faxonius) limosus has apparently been extirpated from West Virginia and should be removed from the state’s list of extant crayfishes. Introduction The first published mention of West Virginia crayfishes appears to have been Faxon’s (1914) report of two taxa referred to as Cambarus bartonii and Cambarus dubius. In more recent years, several surveys have been conducted in various ecological regions, including the Cheat River drainage (Schwartz and Meredith 1960) and the Ohio River bottomlands (Loughman, in press). The last formal statewide effort was made by Jezerinac and colleagues during the summers of 1987 through 1989, and the resultant monograph (Jezerinac et al. 1995) listed 21 crayfish taxa in West Virginia. Since that time, Thoma and Jezerinac (1999) elevated Cambarus (Cambarus) bartonii carinirostris Hay (Rock Crayfish), to species status, and Taylor (2000) described Orconectes (Procericambarus) cristavarius Taylor (Spiny Stream Crayfish), which in West Virginia previously had been assigned to Orconectes (Procericambarus) spinosus Bundy. Loughman (2007) added Procambarus (Ortmannicus) acutus Girard (White River Crayfish) to the state’s fauna, increasing the number of known taxa to 22. Efforts to identify conservation concerns and potential state records since the publication of Jezerinac et al. (1995) have been sporadic. Endemic species such as Cambarus (Hiaticambarus) elkensis Jezerinac and Stocker (Elk River Crayfish), have received further attention, but most taxa have 1Department of Natural Sciences and Mathematics, Campus Service Center Box 139, West Liberty State College, West Liberty, WV 36074. 2Biology Department, Indiana Statue University, Terre Haute, IN 47809. 32364 E. Linden Hill Drive, Bloomington, IN 47401. 4US Geological Survey, West Virginia Cooperative Fish and Wildlife Research Unit, PO Box 6125, Morgantown, WV 26506. *Corresponding author - zloughman@westliberty.edu. 226 Northeastern Naturalist Vol. 16, No. 2 remained understudied. Since the 1980s, conservation problems, including land-use changes, stream degradation, species invasions, and habitat loss, have occurred in areas with diverse crayfish populations throughout the state, but the ultimate impacts of these problems remain poorly understood. In an effort to help remedy this situation, our paper documents noteworthy distribution records, natural- and life-history information, and conservation concerns for 10 of the state’s 22 currently recognized crayfish taxa. Methods The majority of collections were performed in the summers of 2006 and 2007 in the Allegheny Mountain and Appalachian Plateau physiographic provinces of West Virginia. Sites were chosen based on historical records and ease of access. Collection methods included trapping with minnow traps, seining, snorkeling, hand collecting, and burrow excavation. A minimum effort of 10 seine hauls or 60 minutes total collecting time was performed at each site. Field work was completed by Z.J. Loughman and field assistants. Total carapace length (TCL) of all vouchered animals was measured with digital calipers to the nearest 0.1 mm. Ovigerous females’ eggs were counted, and egg diameters were measured to the nearest 0.01 mm. Each species account includes specific locations, followed by new insights into aspects of each species’ biology. Most collections are curated at the Indiana Biological Survey (INBS) Crustacean Collection, Bloomington, Indiana. The following abbreviations are used in the text: TCL = total carapace length, TBL = total body length, M-I = Form I male, M-II = Form II male, F = female, Ovig F = ovigerous female, Juv = juvenile, SR = state route, CR = county road, ints = intersection, and Cr = Creek. Results and Discussion Cambarus (Cambarus) carinirostris Hay, 1914 (Rock Crawfish) Localities Hancock County: (1) Kings Creek, CR 11 2.2 mi E of CR 11/SR 2 ints. 23 April 2007. 5 F, 1 M-I, 1 Juv. (2) Tomlinson Run immediately downstream of Tomlinson Run Lake spillway. 17 October 2005. 2 F, 3 M-I. Fayette County: (3) Keeney Creek, 0.2 mi S from CR 82/ US 60 ints. 19 July 2006. 3 F, 1 Ovig F, 2 M-II. (4) Brackens Creek, 0.78 mi N of US 60/ SR 42 ints on SR 42, 19 July 2006. 3 F, 2 M-II, 1 M-I, Marshall County: (5) Long Run, 0.66 mi S of Long Run/Fish Creek confluence. 20 July 2005. 1 F, 1 M-II. (6) Fish Creek, adjacent to Lynn Camp bridge in Lynn Camp. 30 October 2005. 2 F, 1 M-I. Monroe County: (7) Kitchen Creek, SR 3 crossing 2.1 mi W of Centennial. 18 July 2006. 1 F, 2 M-II. Nicholas County: (8) Collision Creek, CR 41-7 crossing 1.5 mi N of Pool. 19 July 2006. 6 F, 6 M-II, 2 M-I. (9) Summersville Lake at Dive Shop Marina, 19 July 2006. 1 F. (10) Cranberry River in Monongahela National Forest adjacent to Big Rock Campground. 17 July 2006. 1 F, 1 M-II. (11) North Fork of Cherry River adjacent to SR 39 at Four 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 227 Season Lodge. 16 July 2007. 4 F, 4 M-II. (12) Gauley River, adjacent to SR 20 0.1 mi E of Upperglade. 17 July 2007. 1 F. Ohio County: (13) Wiedman Run, West Liberty State College Campus Woods trail head, 23 September 2007. 4 F, 2 Ovig F, 5 M-II, 3 M-I. (14) Unnamed tributary along discovery trail in Oglebay Park 0.25 mi W of Schrader Environmental Education Center, 16 July 2005. 34 F, 2 Ovig F, 13 M-II, 8 M-I. Preston County: (15) Snowy Creek immediately downstream of Terra Alta Lake spillway. 20 June 2006. 7 F, 2 M-II. (16) Cheat River at the confluence of Salt Lick Creek and Cheat River. 22 June 2006. 2 F, 3 M-II. Webster County: (17) Laurel Fork of Holly River in Holly River State Park campground, 17 July 2006. 3 F, 1 M-II, 1 M-I. (18) Roadside ditch along CR 7 1.2 mi from SR 20/CR 7 ints. 17 July 2006. 1 F. Distribution and natural history All of the above sites represent previously unrecorded distribution records for C. carinirostris (Fig. 1). Phenotypic variation in color pattern was obvious in this species in West Virginia, with animals from the Allegheny Mountains dominated by greenish hues, and those from the Appalachian Plateau exhibiting beiges, pinks, and oranges. Morphological differences also were observed between specimens from these regions; their taxonomic significance is being investigated by Z.J. Loughman. Allegheny Mountain populations had elongate chelae, highly sculptured movable fingers, and acuminate rostrums. Appalachian Plateau populations had non-elongate, almost truncated chelae, reduced fixed finger sculpturing, and non-acuminate rostrums. Phylogenetic analysis is currently being performed to determine if future taxonomic investigations are warranted. The life history of C. carinirostris has received moderate attention in the literature, and recent observations add to our knowledge. Developed glair glands were observed in females captured at sites 1 and 15 in April and June. Seven ovigerous females were collected and released at site 14 on 16 July 2005. Others had also been collected in July in previous years. Ortmann (1906) in Pennsylvania and Jezerinac et al. (1995) and Thoma and Jezerinac (1999) in West Virginia also reported ovigerous specimens in July, indicating this as the period of egg extrusion in the species. Two females carrying pleopodal instars were collected at site 13 on 23 September 2007 (Table 1). Neonate TCLs for the 37.4-mm female ranged from 5.6 to 6.3 mm (mean = 5.9 mm). Form I males were collected in April, July, September, and October, perhaps indicating a lack of seasonality associated with this male sexual stage. Cambarus carinirostris has been considered a secondary burrower, and previous descriptions of its behavior (Jezerinac et al. 1995) did not indicate a propensity for extreme burrowing in this species. However, collections made at sites 1, 6, and 17 all involved digging the species from complex burrows. Burrows consisted of central shafts with depths ranging from 0.4 to 1.1 m, terminating in an enlarged resting chamber. Each chamber had multiple tunnels radiating from the bottom, and in burrows at sites 6 228 Northeastern Naturalist Vol. 16, No. 2 and 17, there were additional enlarged galleries. When provoked, animals sequestered themselves in the deepest regions of the burrow and required manual extraction. Cambarus (Cambarus) bartonii cavatus Hay, 1903 (Appalachian Brook Crayfish) Localities Raleigh County: (1) Retention Ponds present on Eccles contour strip mine 1.3 mi E of Lilly Mountain. 9 April 2005. 3 F, 1 M-II, 4 M-I. (2) Lake Stephen Spillway 0.2 mi downstream from Lake Stephen Dam. 19 July 2006. 1 F. Distribution and natural history Cambarus b. cavatus was previously unrecorded from Raleigh County, although it had been recorded by Jezerinac et al. (1995) from the Coal River Basin (Fig. 1). At site 2, the species was found in headwater streams, but it was collected from surface-mine retention ponds at site 1. These mine ponds were created in the mid-1970s by damming a second-order stream. Such Figure 1. Collection sites for the subgenus Cambarus. 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 229 lentic habitats are atypical for this species, and it is likely that the population at that site descended from the original stream dwellers. All animals collected from the ponds were taken in minnow traps, and 96% of them were large, sexually mature individuals. No burrows or cover objects (rocks, logs, etc.) were in the pond, but C. b. cavatus was observed using decaying Typha latifolia L. (Broad-leaved Cattail) for both cover and forage. Cambarus (Cambarus) sciotensis Rhoades, 1944 (Teays River Crayfish) Localities Greenbrier County: (1) Anthony Creek 0.25 mi N of Blue Bend Public swimming area. 18 July 2006, 1 F and 17 July 2007, 1 F, 1 M-II. Nicholas County: (2) Twenty Mile Creek at CR 20-21 crossing. 20 July 2007. 6 F, 2 M-II. (3) Cherry River, SR 20 crossing in Holcomb. 17 July 2006. 3 F, 4 M-II and 16 July 2007, 2 F, 2 M-II. (4) Cranberry River in Monongahela National Forest adjacent to Big Rock campground. 17 July 2006. 6 F, 2 M-II, 1 M-I. (5) North Fork of Cherry River adjacent to SR 39 at Four Season Lodge. 16 July 2007. 1 Juv, 4 F, 4 M-II. (6) Gauley River, adjacent to SR 20 0.1 mi E of Upperglade. 17 July 2007. 1F. (7) Meadow River at Meadow River/Youngs Creek confluence, 19 July 2006. 2 F, 2 M-I. (8) Peters Creek along SR 129 0.1 mi S of Drennen. 20 July 2006. 6 F, 1 Ovig F, 4 M-II, 3 M-I. (9) Peters Creek at SR 129/SR 39 ints. 20 July 2006. 2 F, 2 M-II. Wayne County: (10) Twelve Pole Creek immediately downstream of Dixon Dam in Dixon. 9 October 2004. 1 M-I. Webster County: (11) Grassy Creek at SR 20 crossing adjacent to Thomas Coger logging yard. 16 July 2007. 2 M-II. Distribution, natural history, and taxonomy Cambarus sciotensis has a broad range over central Appalachia and likely represents a species complex. Sites at which Cambarus sciotensis was observed during this study are displayed in Figure 1. Our morphological analysis determined that three distinct phenotypes occur in West Virginia. Variation included palmar tubercles, and spines on the postorbital ridges and rostra. At sites 2–9, all in the Gauley River basin, specimens had a single row of tubercles on the mesial surface of the palm, a narrow rostrum, and prominent spines on the postorbital ridge. Those from sites 1 and 10, in the greater Kanawha River basin, had two rows of palmar tubercles, narrower rostrums, and reduced postorbital ridge spines. This condition was also observed in C. sciotensis populations in Virginia and West Virginia by Hobbs et al. (1967) and Jezerinac et al. (1995). Populations in the upper Elk River basin were the most distinct, possessing rostral spines, double rows of palmar tubercles, and narrow rostra. Future investigations will attempt to elucidate the taxonomic status of these phenotypes in West Virginia. Cambarus sciotensis was found in moderate to large, wadeable streams, but exhibited a preference for larger streams and rivers. In large streams, the 230 Northeastern Naturalist Vol. 16, No. 2 species is a tertiary burrower. Microhabitats used by the species included slab boulders, amorphous boulders, boulder complexes, and leafpacks in pool thalwegs. At sites 2–9, large form I males were found under slab boulders in high-velocity riffles. There was no evidence of habitat specialization among neonates, juveniles, and adults. On 20 July 2006, an ovigerous female was collected at site 8 (Table 1). Egg diameters ranged from 1.7–2.2 mm (mean = 2.0 mm). The female was found under a large slab boulder downstream of a small rapid. Water velocity beneath the boulder was low, and the female was sequestered in a well-constructed burrow composed of various-sized gravels and cobbles. On 9 October 2004, a form I male with a TCL of 60.2 mm and a TBL of 114.6 mm was collected at site 10. It represents a new size record for the species from throughout its range. Taylor and Schuster (2005) noted the maximum TBL for C. sciotensis throughout its range as 105.0 mm, and the largest West Virginia specimen reported by Jezerinac et al. (1995) measured 54.0 mm TCL. Cambarus (Hiaticambarus) chasmodactylus James, 1966 (New River Crayfish) Localities Greenbrier County: (1) Greenbrier River at Renick public stream access 0.3 mi S of Renick. 17 July 2007. 4 F, 2 M-I. (2) Anthony Creek 0.25 mi N of Blue Bend Public swimming area. 17 July 2006, 12 F, 9 M-II, 4 M-I and 17 July 2007, 8 F, 6 M-II, 4 M-I. (3) Second Creek at CR 219-1 crossing 0.25 mi W of 219. 18 July 2007. 7 F, 5 M-II. Table 1. Reproductive biology for ovigerous crayfishes collected during this study. Species Date County Stream TCL Condition Cambarus (C.) carinirostris 23 Sept. 2007 Ohio Wiedman Run 37.4 50, 2nd instars; fully developed ** ** ** 54.3 12, 2nd instars; fully developed Cambarus (C.) sciotensis 20 July 2006 Nicholas Peters Creek 39.7 111 attached ova Cambarus (H.) chasmodactylus 1 Sept. 2007 Greenbrier Anthony Creek 38.0 57 attached ovum Orconectes (P.) rusticus 29 April 2006 Cabell Four Pole Creek 22.8 138 attached ova ** ** ** 25.7 237 attached ova ** ** ** 25.7 121 attached ova ** ** ** 26.9 136 attached ova ** ** ** 30.7 76 attached ova ** ** ** 33.2 172 attached ova ** ** ** 33.6 185 attached ova ** ** ** 33.8 256 attached ova **Data same as above cell. 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 231 Distribution and natural history In West Virginia, C. chasmodactylus occurs in streams and rivers of the Greenbrier River basin. Very little is known about its natural history in the state (Jezerinac et al. 1995). We visited historic sites (1–3) in the Greenbrier basin and gathered detailed natural history data (Fig. 2). Anthony Creek (site 2) was visited on the same date in 2006 and 2007 to determine if cohorts of the same TCLs were present in successive years. Specimens were found under various substrate items ranging from small cobbles to large slate slabs. Adults were always collected under large amorphous boulders or sandstone slabs, particularly in mid-channel environments, and preferred moderate to low stream velocities in larger order streams and rivers. Adults were not gregarious, but younger animals did occur together with them. Juveniles were collected primarily from stream edges and not the channel proper. In several instances, O. cristavarius were found alongside adult C. chasmodactylus. Mean adult TCL was 34.5 mm, with females (n = 29) averaging 37.3 mm, form I males (n = 10) averaging 47.0 mm, and form II males (n = 20) averaging 30.3 mm. TCL ranged from 23.9–54.9 mm for females, 38.6–51.7 mm for form I males, and 23.4–46.1 mm for II males. Total carapace length ranged from 23.1 mm–54.1 mm (n = 37) in 2006 collections and 24.2 mm–51.7 mm (n = 34) in 2007. Both sexes were among the largest animals; large TCL is not a trait limited to form I males in this species. Sex ratio in both years was 1:1. The following TCL cohorts (mm) repeated between 2006 and 2007 and likely represent important age classes: 24 (n = 6), 31 (n = 7), 38 (n = 10), and 47 (n = 5). A captive female measuring 38.0 mm TCL extruded eggs on 1 September 2007. The total complement numbered 57, and the mean diameter was 3.7 mm. This count conforms to those of counts for previous ovigerous females of similar size collected in West Virginia (Jezerinac et al. 1995). Cambarus (Hiaticambarus) elkensis Jezerinac and Stocker, 1993 (Elk River Crayfish) Localities Webster County: (1) Laurel Fork of Holly River in Holly River State Park Campground. 17 July 2006, 3 F, 1 M-I and 16 July 2007, 4 F, 9 M-II. (2) Hodam Creek 0.2 mi S of SR 20/CR 20-18 ints. 17 July 2006. 2 Juv. Distribution and natural history Cambarus elkensis is endemic to headwaters in the Elk River basin of West Virginia (Jezerinac and Stocker 1993). Site 2 above is a new distribution record (Fig. 2). The population there was sympatric with Cambarus (Puncticambarus) robustus, utilizing similar microhabitats and preferring mid-channel slab boulders. The species was also found at the type locality (site 1) in 2006 and 2007. This population appeared stable and was undergoing recruitment with multiple age cohorts. 232 Northeastern Naturalist Vol. 16, No. 2 Cambarus (Hiaticambarus) longulus Girard, 1852 (Atlantic Slope Crayfish) Localities Monroe County: (1) Back Creek 1.1 mi W of Sweet Springs. 18 July 2006. 7 F, 14 M-II. Natural history and conservation Most of the range of this species lies in Virginia and North Carolina, and its range in West Virginia is restricted to the James River basin in Monroe County (Jezerinac et al. 1995). The collection from Back Creek (Fig. 2) is the largest single collection (n = 21) of the species made at one time in West Virginia. Specimens were found primarily under small slabs and boulders in riffles and plunge pools. Interstitial spaces in stream substrate were used by all sizes; adults were encountered in areas of high velocity in plunge pools, and juveniles were found in thalwegs and leafpacks in areas of low velocity. Four size cohorts were present in this sample, and prominent TCLs (mm) were 15 (n = 7), 17 (n = 4), 23 (n = 3), and 27 (n = 3). The Figure 2. Collection sites for subgenera Hiaticambarus, Jugicambarus, and Puncticambarus. 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 233 male:female sex ratio was 2.75:1. Prior to our sampling, the population at Potts Creek reported by Jezerinac et al. (1995) was considered stable and abundant (Jennifer Wykle, WVDNR, Elkins, WV, pers. comm.). Collecting efforts at this site in 2006, however, resulted in zero specimens. Several anthropogenic impacts on the stream were noted, including increased siltation, channelization, eutrophication, and acidification. Considering the current condition of this stream and the extremely restricted range of C. longulus in West Virginia, the species should be monitored to determine its conservation status. Cambarus (Jugicambarus) dubius Faxon, 1884 (Upland Burrowing Crayfish) Localities Cabell County: (1) Seep 0.25 mi S of Ritter Park tennis courts in Huntington. 12 April 2005. 1 M-I. Distribution and natural history Cambarus dubius is a polymorphic species found throughout the Appalachian Mountains, with three distinct phenotypes occurring in West Virginia (Jezerinac et al. 1995). These include an orange morph found in the Allegheny Mountains, a blue morph found in the western part of the Appalachian Plateau and Greenbrier River Valley, and a black/orange morph found in the central Appalachian Plateau and southern portions of the Ridge and Valley. The specimen reported above was the black/orange morph, the first found in Cabell County in the Lower Ohio basin (Fig. 2). Blue specimens have been found in this system within 2 km of the Cabell County site (Jezerinac et al. 1995). The black/orange individual was taken from a burrow in an isolated forest seep. The terminal burrow depth was 1.1 m, ending in a resting chamber that bifurcated into 2 ancillary tunnels, both of them packed with detritus that showed signs of mastication. Cambarus (Puncticambarus) robustus Girard, 1852 (Big Water Crayfish) Localities Hancock County: (1) Kings Creek, CR 11 2.2 mi E of CR 11/SR 2 ints. 23 April 2007. 5 F, 1 M-I, 1 Juv. (2) Tomlinson Run immediately downstream of Tomlinson Run Lake spillway. 17 October 2005. 2 F, 3 M-I. Fayette County: (3) Painter Creek adjacent to SR 612 5.1 mi W of US 19/ SR 612 ints. 19 July 2006. 3 F, 2 M-II. Marshall County: (4) Long Run, 0.66 mi S of Long Run/Fish Creek confluence. 20 July 2005. 1 F, 1 M-II. (5) Fish Creek, adjacent to Lynn Camp bridge in Lynn Camp. 30 October 2005. 2 F, 1 M-I. Raleigh County: (6) Lake Stephen Spillway, 0.2 mi downstream from Lake Stephen Dam. 19 July 2006. 7 F, 2 M-II. Wayne County: (7) Twelve Pole Creek immediately downstream of Dixon Dam 234 Northeastern Naturalist Vol. 16, No. 2 in Dixon. 9 October 2004. 1 M-II. Webster County: (8) Hodam Creek 0.2 mi S of SR 20/CR 20-18 ints. 17 July 2006. 3 F, 4 M-II. Distribution and natural history Cambarus robustus has a broad range in eastern North America, and the taxonomic status of populations in North Carolina and Tennessee is under review (J.E. Cooper, NC Museum of Natural Sciences, Raleigh, NC, pers. comm.). It is widely distributed in West Virginia (Jezerinac et al. 1995), but the eight localities listed above are new records (Fig. 2). Most notable of these are sites (Sites 1, 2, 4, and 5), from the Upper Ohio North basin, the Upper Ohio South basin, and the Coal River basin. The populations in these Upper Ohio basins are the northernmost known in the state, although Jezerinac et al. (1995) indicated that “a lack of suitable habitat” existed there. Interestingly, the species was absent from major streams separating the Fish Creek system in Marshall County and the Kings Creek/Hodam Creek system in Hancock County, although adequate habitat does exist within the region. It is possible that these populations are separated due to different recolonization paths following Pleistocene glaciation. Similar distribution gaps are present within the range of Cambarus (Tubericambarus) thomai (Brown Mudbug) (Jezerinac, 1993) in both of the Upper Ohio basins (Z.J. Loughman, unpubl. data). Although C. robustus is essentially a lotic species, Lake Stephens in Raleigh County contained a large lentic population. Specimens were collected from the lake proper and from the spillway. Within the spillway stream, they were limited to plunge pools, which probably helped them withstand periods of drawdown. Initial colonization of the lake likely occurred via bait-bucket introduction. This is the only population of this species known to occur in a reservoir in West Virginia, although such populations are known to occur rarely elsewhere in North America (Hamr and Berrill 1985). In larger order streams, C. robustus was found under slab boulders, and were mostly collected in runs and glides. Orconectes (Faxonius) limosus (Rafinesque, 1817) (Spinycheek Crayfish) Conservation Orconectes limosus has apparently been extirpated from West Virginia and should be removed from the state’s list of extant crayfishes. Concerted efforts by J.Z. Loughman and S.A. Welsh in the summer of 2007 produced no specimens. All historic sites were surveyed, as well as additional suitable habitat in the Potomac River drainage. Considerable efforts to find this species had been undertaken by other researchers in the summers of 2005 and 2006, with the same negative results (C. Swecker and T. Jones, Marshall University, Huntington, WV, pers. comm.). Factors that probably contributed to the extirpation of O. limosus from West Virginia include invasive species and increased rates of urban development within its former range. Populations of invasive Orconectes (Gremicambarus) 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 235 virilis (Hagen) (Virile Crayfish) now dominate the Potomac River drainage. This species was present at all sites surveyed in Bat Creek and Opequon Creek watersheds in 2006 and 2007, which were the last areas historically known to harbor O. limosus. The invasive species was found with Orconectes (Crockerinus) obscurus in Bat and Opequon creeks, and also occurred in Cherry Run. Previous research in Maryland (Schwartz et al. 1963) documented that O. virilis invasions ultimately led to localized extirpations of O. limosus and C. b. bartonii through increased competition for food and refugia. In addition to O. virilis invasions, increased urban development and its associated impacts on aquatic systems (i.e., siltation, channelization, chemical pollution) have occurred in Bat Creek and Cherry Run systems. These combined factors have apparently caused the extirpation of O. limosus in West Virginia. Orconectes (Procericambarus) cristavarius Taylor, 2000 (Spiny Stream Crayfish) Localities Greenbrier County: (1) Greenbrier River at Renick public stream access 0.3 mi S of Renick. 17 July 2007. 3 F, 1 M-II. (2) Anthony Creek 0.25 mi N of Blue Bend Public swimming area. 18 July 2006, 1 F, 1 M-II, 2 M-I, and 17 July 2007, 2 F, 1 M-II, 4 M-I. (3) Second Creek at CR 219-1 crossing 0.25 mi W of 219. 18 July 2007. 9 F, 8 M-II, 4 M-I. Monroe County: (4) Kitchen Creek at SR 3 crossing in Gap Mills. 18 July 2006. 1 F, 2 M-II. Nicholas County: (5) Twenty Mile Creek at CR 20-21 crossing. 20 July 2007. 4 F, 5 M-II. (6) Gauley River, adjacent to SR 20 0.1 mi E of Upperglade. 17 July 2007. 2 F. (7) Peters Creek along SR 129 0.1 mi S of Drennen. 20 July 2006. 1 M-II. (8) Peters Creek at SR 129/SR 39 ints. 20 July 2006. 2 F, 2 M-II. Raleigh County: (9) Lake Stephen Spillway 0.2 mi downstream from Lake Stephen Dam. 19 July 2006. 4 M-II. Webster County: (10) Grassy Creek at SR 20 crossing adjacent to Thomas Coger logging yard. 16 July 2007. 2 F, 2 M-II. (11) Strouds Creek at CR 11 crossing 1.4 mi N of Camden on the Gauley. 18 July 2007. 4 F, 1 M-II. Distribution and natural history Orconectes cristavarius ranges across the Central Appalachians, and prior to its description (Taylor 2000), the West Virginia populations had been assigned to O. spinosus (Jezerinac et al. 1995). The range of these populations encompasses the Big Sandy and Kanawha river basins, and within the eastern portion of the latter it was more widespread than previously thought. Sites 1 and 3 in the Greenbrier, sites 10 and 11 in the Elk, and site 9 in the Coal are new distribution records (Fig. 3). In addition, sites 5–8 and 11 are first records for the Gauley River system, where O. cristavarius was the dominant orconectid. Prior to our recent surveys, Orconectes (Crockerinus) sanbornii sanbornii (Faxon) (Sanborn Crayfish) was recorded by Jezerinac et al. (1995) as inhabiting the Gauley River system, but we found no specimens. 236 Northeastern Naturalist Vol. 16, No. 2 Orconectes cristavarius was found in various-sized cobbles in moderate- to low- velocity streams. Interstitial spaces between large cobbles and boulders were preferred microhabitats, as were detritus beds in pool thalwegs. At site 3, large numbers were seen in backwater pools during late summer. Several size groups were using these pools, with evidence of habitat specialization, where juveniles preferred shallow pools and adults were seen in deeper water. Orconectes (Procericambarus) rusticus (Girard, 1852) (Rusty Crayfish) Localities Cabell County: (1) Four Pole Creek at 4th street crossing in Huntington. 29 April 2005. 1 F, 8 Ovig F, 10 M-I. Life history Orconectes rusticus has been steadily spreading throughout West Virginia since Lawton (1979) published the first records of the species’ occurrence. Populations are now known from Marshall County (Loughman, in press), throughout the Lower Kanawha River system (C. Swecker, pers. comm.), and the Lower Ohio River Basin (Jezerinac et al. 1995). Despite hundreds Figure 3. Collection sites for the genus Orconectes. 2009 Z.J. Loughman, T.P. Simon, and S.A. Welsh 237 of individuals having been collected, very little life-history information has been published for any of these populations. Eight ovigerous females were found under small slabs and benthic debris at the above locality on 29 April 2004. Egg counts ranged from 76 to 256 (mean = 165; Table 1). There was no statistical correlation between TCL and egg number (r2 = 0.0899). On two occasions, three ovigerous females were found under the same cover. At night, form I males were seen cruising the stream bottom, mostly in runs and pools with ample substrate debris and leaf packs. Several mating efforts were observed in situ, and form I males readily copulated with females in collecting buckets. Four Pole Creek is severely impacted by solid surface runoff, specifically nutrient inputs, but these impacts apparently are not detrimental to the O. rusticus population. No native species (O. s. sanbornii and C. b. cavatus) were found in the creek, indicating local extirpation of the native population stocks. Acknowledgments We would like to thank Nicole Garrison, Christopher Hearn, Natalie Mancusso, Cody Rosettii, and Christopher Vopal for assistance in the field, and Sarah Brammer, Melinda Kreisburg, Nicole Berrisford, and James Harvey for reviewing earlier drafts of the manuscript. Special thanks are due to Dr. John Cooper whose review of this manuscript has increased its value greatly. Indiana Biological Survey provided funds for completion of field work and museum space for specimens. Thanks are also expressed to Oglebay Institute, West Liberty State College, and the West Virginia Division of Natural Resources for providing additional financial support. Literature Cited 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. Harvard College 40(8):351–427. Hamr, P., and M. Berrill. 1985. The life histories of north-temperate populations of the crayfish Cambarus robustus and Cambarus bartoni. Canadian Journal of Zoology 63:2313–2322. Hobbs, H.H., Jr., P.C. Holt, and M. Walton. 1967. 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