Conservation, Biology, and Natural History of Crayfishes from the Southern US 2010 Southeastern Naturalist 9(Special Issue 3):79–126 Distribution, Status, and Life-history Observations of Crayfishes in Western North Carolina Jeffrey W. Simmons1,2,* and Stephen J. Fraley1,3 Abstract - Approximately 390 native North American crayfish species are known, representing nearly two-thirds of the world’s crayfish fauna. The majority of these species occur in the southeastern United States. North Carolina supports a substantial proportion of that diversity with 41 described indigenous crayfish species, 12 of which are endemic, and 3 introduced species, many of which are of significant conservation interest. In the late 1990s, the North Carolina Wildlife Resources Commission (NCWRC) began a focused effort to inventory and establish baselines for monitoring populations of both native stream-dwelling and burrowing crayfishes and invasive non-native species. During 2004–2005, that effort was completed for the Hiwassee, Little Tennessee, Savannah, French Broad, Watauga, New, Catawba, and Broad river basins in western North Carolina. Twenty-four stream-dwelling species and 5 burrowing species were collected from 199 stream sites and 58 burrowing sites. New records for many species, including a new river basin record for Cambarus reduncus (Sickle Crayfish), and new county records for C. howardi (Chattahoochee Crayfish), C. dubius (Upland Burrowing Crayfish), and C. nodosus (Knotty Burrowing Crayfish), were determined during these surveys. Small range expansions were documented for Orconectes virilis (Virile Crayfish; not native to North Carolina) and for Procambarus acutus (White River Crawfish; introduced outside its native range in North Carolina). We failed to detect the non-native O. rusticus (Rusty Crayfish), and P. clarkii (Red Swamp Crawfish) at or near previously reported localities. Observations of life-history traits, such as reproductive condition, fecundity, and habitat use were recorded. Specimens were also provided to crayfish taxonomists to help resolve certain taxonomic problems and to assist in the completion of new species descriptions. Data collected during this and previous NCWRC inventories, as well as data obtained from the North Carolina State Museum of Natural Sciences, North Carolina Division of Water Quality, Ohio State University, and other cooperators, were incorporated into a detailed GIS database. This database was used to identify data gaps to guide sampling efforts and to assess species and population status. In the future, this GIS database should provide a useful tool in monitoring the status of native crayfish populations and the spread of invasive species, and informing conservation and management decision making. Introduction In North Carolina, there are presently 41 described indigenous crayfish species (12 of which are endemic), at least 5 putative undescribed species, 1North Carolina Wildlife Resources Commission, 1701 Mail Service Center, Raleigh, NC 27699. 2Current address - Tennessee Valley Authority, 1101 Market Street, PSC- 1X-C, Chattanooga, TN 37402. 3Current address - North Carolina Wildlife Resources Commission, 50 Trillium Way, Clyde, NC 28721.*Corresponding author - jwsimmons0@ tva.gov. 80 Southeastern Naturalist Vol. 9, Special Issue 3 and 3 introduced species (Cooper 2005). North Carolina’s Western Region (Region), as defined by the North Carolina Wildlife Resources Commission (NCWRC), includes the Hiwassee, Little Tennessee, Savannah, French Broad, Watauga, New, Catawba, and Broad river basins (Fig. 1). The Hiwassee, Little Tennessee, Savannah, French Broad, Watauga, and New river basins are contained within the Blue Ridge physiographic province. The Catawba and Broad river basins are primarily within the Piedmont Plateau physiographic province; the headwaters of these river basins are within the Blue Ridge. This region is presently known to support 26 described indigenous crayfish species (four of which are endemic), at least 5 putative undescribed species, and 4 introduced species (Table 1). The Region is part of an area that supports the greatest crayfish diversity in the world, and many of the species and populations found here are priorities for conservation (Butler 2002, NCWRC 2005a, Taylor et al. 2007). Native crayfish serve vital roles in aquatic and terrestrial ecosystems as predators, scavengers, and prey for fishes and other wildlife (Fullerton 2002, Hobbs 1993). An understanding of crayfish taxonomy, ecology, distribution, and abundance is necessary for resource managers to determine relative conservation status and to develop effective monitoring and management strategies. Great strides have been made in recent years to improve our understanding of North Carolina’s crayfish fauna, but much information is still lacking (Clamp 1999, Cooper 2002). Basic alpha taxonomy of North Carolina’s crayfish fauna is still incomplete. Nine new species were described since 2000, and another 5 forms are believed to be distinct species that are presently being described or are awaiting further investigation and formal description (sensu Cooper Figure 1. Counties, river basins, and major waterways of the Western Region of North Carolina as defined by the NCWRC. 2010 J.W. Simmons and S.J. Fraley 81 2005). For many of these taxa, voucher material from various life stages are needed to aid further understanding of taxonomic resolution, distribution, and conservation status. While the distributions by river basin of most of North Carolina’s crayfish fauna are generally known, detailed distribution and abundance data necessary for assessment of population trends and conservation status are lacking. More than any other factor, Clamp (1999) found that inadequate sampling and a lack of analysis of available data hindered our knowledge of the status of freshwater and terrestrial crustaceans in North Carolina. Because available survey data are limited spatially and temporally and have not been consolidated from multiple sources, it is difficult to determine long-term crayfish population trends. Consequently, the effects of modern widespread habitat changes on the Region’s fauna are difficult to assess. Non-native crayfish introductions may pose the greatest threat to native crayfish populations through potential displacement of indigenous species (Lodge et al. 2000). Wild populations of at least 4 non-native species may occur in the Region (Cooper 2005, Cooper and Armstrong 2007). Orconectes virilis (Virile Crayfish), O. rusticus (Rusty Crayfish), Procambarus acutus (White River Crawfish), and P. clarkii (Red Swamp Crawfish), have been collected from the Catawba River basin; the upper Broad River and Little Tennessee River basins; the French Broad, Broad, and Watauga river basins; and the Broad River basin, respectively. The possession and transport of Virile and Rusty Crayfish was recently banned in North Carolina (NCWRC 2005b). The NCWRC began to inventory crayfishes during the 1990s to establish baselines for monitoring populations of rare native and invasive non-native species. Previous surveys conducted by NCWRC within the Region were reported by Fullerton (2002; French Broad and Savannah river basins), Marsh (1998; Catawba [Linville] River basin), and McGrath (1993, 1997, 1998; French Broad [Nolichucky], Hiwassee, and New river basins, respectively). These data are centrally stored in the NCWRC Aquatics Database and are readily accessible; however, much data from the Region are scattered elsewhere. Considerable information, including voucher specimens and additional distribution data, is available from the North Carolina Museum of Natural Sciences (NCMNS) in Raleigh. More information is available from other sources such as publications, reports, and other museums. Consolidation and analysis of all available data from the Region would improve assessment of the quantity and quality of existing information, identification of spatial and temporal data gaps, and longterm monitoring and population-trend assessment. The objectives of this study were to: (1) acquire and consolidate as much available crayfish collection data as possible from the Region into a GIS database to help identify distribution information gaps; (2) survey areas lacking information to further understand the distribution of burrowing and stream-dwelling crayfishes in the Region and to establish baselines for population monitoring; (3) collect specimens of putative undescribed species and 82 Southeastern Naturalist Vol. 9, Special Issue 3 Table 1. Native (N) and non-native (NN) crayfish taxa by western North Carolina river basin (Catawba = C, Broad = B, Little Tennessee = LT, Hiwasee = H, French Broad = FB, New = N, Watauga = W, and Savannah = S), including putative undescribed species discussed in species accounts. Endemic taxa (*) are only in a specific river basin and only in North Carolina. “Near endemics” (**) are narrowly distributed and present in adjacent river basins or states. Status ranks are explained in Table 2; a superscript (F) indicates a Federal species of concern. Status River basin Global State AFS NC Species C B LT H FB N W S rank rank status status Cambarus (Cambarus) bartonii (Fabricius) (Common Crayfish) N N N N N N N N G5 S5 CS NS C. (C.) eeseeohensis Thoma (Grandfather Mountain Crayfish) N* G1 S1 T SR C. (C.) howardi Hobbs and Hall (Chattahoochee Crayfish) N N G3 S3 CS SR C. (C.) lenati Cooper (Broad River Stream Crayfish) N* G2 S2 T SR C. (C.) sp. A N N N N GU S2S3 NS W C. (Depressicambarus) latimanus (LeConte) (Variable Crayfish) N G5 S5 CS NS C. (D.) reduncus Hobbs (Sickle Crayfish) N N G4G5 S3 CS NS C. (Hiaticambarus) chasmodactylus James (New River Crayfish) N G4 S3 CS W C. (H.) longirostris Faxon (Longnose Crayfish) N N G5 S3S4 CS NS C. (cf. H.) sp. nov. N* NR NR NS NS C. (Jugicambarus) asperimanus Faxon (Mitten Crayfish) N N N N N N N G4 S3 CS NS C. (J.) carolinus (Erichson) (Red Burrowing Crayfish) N N G4 S2 CS W C. (J.) dubius Faxon (Upland Burrowing Crayfish) N N N N N G5 S3 CS NS C. (J.) nodosus Bouchard and Hobbs (Knotty Burrowing Crayfish) N G4 S2 CS SR C. (J.) tuckasegee Cooper and Schofield (Tuckasegee Stream Crayfish) N* G1G2 S1S2 T SR C. (Puncticambarus) brimleyorum Cooper (Valley River Crayfish) N* G3G4 S3 V SR 2010 J.W. Simmons and S.J. Fraley 83 Table 1, continued. Status River basin Global State AFS NC Species C B LT H FB N W S rank rank status status C. (P.) chaugaensis Prins and Hobbs (Chauga Crayfish) N** G2 S2 T SC C. (P.) georgiae Hobbs (Little Tennessee Crayfish) N** G2 S2S3 E SC C. (P.) hiwasseensis Hobbs (Hiwassee Crayfish) N** G3G4 S3S4 V W C. (P.) hobbsorum Cooper (Rocky River Crayfish) N G3G4 S3S4 CS NS C. (P.) johni Cooper (Carolina Foothills Crayfish) N** N** G3 S3 CS SR C. (P.) parrishi Hobbs (Hiwassee Headwater Crayfish) N** G2 S1 E SCF C. (P.) reburrus Prins (French Broad Crayfish) N** N** N** G3 S3 CS SRF C. (P.) robustus Girard (Big Water Crayfish) N N N G5 S3 CS NS C. (P.) spicatus Hobbs (Broad River Spiny Crayfish) N** G3 S2 V SC C. (P.) sp. C N N G4 S4 CS NS C. (P.) sp. nov. N** N** NR NR NS NS C. (Tubericambarus) acanthura Hobbs (Thornytail Crayfish) N G4 S1 CS SR O. (Procericambarus) sp. nov. N* NR S2? NS SR Orconectes (P.) cristavarius Taylor (Spiny Stream Crayfish) N G5 S3 CS W O. (P.) rusticus Girard (Rusty Crayfish) NN NN G5 SNA (SE) CS NS O. (Gremicambarus) virilis Hagen (Virile Crayfish) NN G5 SNA (SE) CS NS Procambarus (Ortmannicus) acutus (Girard) (White River Crawfish) N NN NN G5 S5 CS NS P. (Scapulicambarus) clarkii (Girard) (Red Swamp Crawfish) NN NN G5 SNA (SE) CS NS 84 Southeastern Naturalist Vol. 9, Special Issue 3 other problematic forms to aid in resolving taxonomic problems; (4) record observations of life-history traits as opportunities allow; and, (5) update the GIS database with our collection data to produce more complete species range maps and provide a tool for further conservation status assessments and development of monitoring strategies. Methods Site selection We sought to address the most immediate needs for understanding crayfish distributions, taxonomy, and conservation status throughout the Region, and complete baseline data collection during 2004 and 2005. Locations of most 2004 field surveys were based on recommendations from Dr. John E. Cooper, Research Curator of Crustaceans at the NCMNS, to fulfill the most immediate needs for specimens and distribution information for potential undescribed species. During January 2005, all available crayfish locality data from the NCMNS for the 8 western river basins were obtained, and locality data that lacked latitude/longitude coordinates were georeferenced. Additional collection data were also obtained from the North Carolina Division of Water Quality (NCDWQ) and Ohio State University. All of these data were incorporated into a GIS database, along with all existing crayfish data from the NCWRC Aquatics Database. ArcGIS™ software was used to analyze these data spatially and temporally by species and river basin. Priority areas for surveys in 2005 were directed by lack of adequate or recent survey data. Stream systems where non-native crayfishes were known from earlier collections were surveyed to assess their present status and that of sympatric native species. Burrowing crayfish sample sites were located with the help of private landowners, other field researchers, and by investigating appropriate habitats when encountered in the field. A press release requesting private landowner assistance in identifying the presence of crayfish burrows on their land was sent to local newspapers throughout the Region, posted on the NCWRC website, and published in Wildlife in North Carolina magazine. Eighty-two landowners responded from the Region. These reported localities were compared to previously identified priority areas, and survey efforts were focused on areas where they overlapped. Collection techniques Stream-dwelling crayfishes were collected during crayfish-specific surveys and incidentally during other aquatic surveys. Dip nets and seines were the primary equipment used during crayfish-specific surveys. Root wads, undercut banks, woody debris piles, leaf packs, substrate exposed after flipping slab rocks, and cobble runs and riffles were sampled by sweeping a dip net through cover or by using collectors’ feet to disturb substrata immediately upstream from a dip net or seine. During mussel and fish surveys, 2010 J.W. Simmons and S.J. Fraley 85 crayfish were collected by hand while diving or snorkeling or netted when electrofishing. Baited minnow traps were occasionally used with an assortment of baits and were placed generally in deeper, unwadeable runs and pools. Sampling effort during crayfish-specific surveys was recorded as the total number of rock flips, kicks, etc., as well as an estimate of total stream length surveyed. Burrowing crayfish were primarily collected by excavating burrows with a shovel and by hand. Sampling effort was recorded as the number of burrows excavated. Pit-fall traps and nocturnal surface collections were conducted on a few occasions. Data collection For both stream-dwelling and burrowing populations sampled, the presence or absence of more than one year class was recorded, determined by size range and morphological characteristics. Total carapace length (TCL) was measured to the nearest 0.1 mm using digital calipers. Clutch size of ovigerous females was determined by counting all attached eggs. Male reproductive condition, determined by condition of the gonopods, was recorded as juvenile, form I, or form II. When distinctive, life colors were described and often photographed. Notes were made on habitat where species were collected. For burrowing species, the number of burrows present at a site, burrow type (primary burrow or secondary/tertiary burrow), burrow morphology (number of connected surface entrances if known, approximate depth to water table, chimney structure), and the distance of burrows from a flowing or standing water body were recorded. Crayfish were preserved in 95% ethanol, and voucher specimens were deposited at the NCMNS. Specimens were identified using Hobbs (1972, 1989) and Cooper (2004a,b). Identifications were verified by J.E. Cooper. Nomenclature of species described prior to 1989 followed Williams et al. (1989). Nomenclature of recently described species followed the original description (Cooper 2000, 2001, 2006a, 2006b; Cooper and Schofield 2002; Thoma 2005). Undescribed or putative taxa are denoted as per Cooper (2005; J.E. Cooper, NCMNS, pers. comm.). Common names followed Taylor et al. (2007). Additional specimens and locality data were obtained from cooperators who also collected crayfish from the Region during 2004 and 2005. All data collected or obtained from other sources were entered into the NCWRC Aquatics Database and were added to the regional crayfish GIS database. Results During 2004 and 2005, 241 sites were surveyed for crayfish (Fig. 2). Of these, 199 were surveyed for stream-dwelling species and 58 for primary burrowing species (some were at the same locality as stream site). Of the 58 sites sampled for primary burrowing species, 53 were reported by private landowners. In total, 57 stream and burrowing sites were sampled in the French Broad River basin, 56 in the Little Tennessee River basin, 48 in the Catawba River basin, 34 in the Broad River basin, 20 in the Hiwassee 86 Southeastern Naturalist Vol. 9, Special Issue 3 River basin, 14 in the New River basin, 9 in the Watauga River basin, and 2 in the Savannah River basin. Five burrowing and 23 stream-dwelling native species were collected. New distribution information was gained for several species. Cambarus reduncus (Sickle Crayfish) was collected for the first time from the Broad River basin. New county records were collected for 3 species: C. dubius (Upland Burrowing Crayfish), C. nodosus (Knotty Burrowing Crayfish), and C. howardi (Chattahoochee Crayfish). Other minor expansions of previously reported distributions were recorded for several species, and are described in species accounts. The non-native O. virilis was collected at 2 localities during our efforts in the Catawba River basin. One of those localities was in the White Creek stream system, where Virile Crayfish was not previously known to occur. Procambarus acutus was collected outside of its native range in the Region from Price Lake, a previously known locality in the Watauga River basin. White River Crawfish were also identified from specimens collected by a cooperator at new localities in the Pigeon River system (French Broad River basin), a short distance from where it was previously known to occur. Two other non-native species known to have been introduced in the Region, P. clarkii and O. rusticus, were not found at sites at or near localities where they were collected in the past. Populations of these species may not have become established after initial introductions, or nascent populations may be at such low densities that we failed to detect them. At least for the short term, our results with regard to the spread of invasive non-native species appear to be good news; however, several endemic species are potentially threatened if further expansion of these non-native species occurs. Figure 2. NCWRC crayfish collection localities in the Western Region of North Carolina during 2004–2005, representing 241 sites. 2010 J.W. Simmons and S.J. Fraley 87 While our primary goal was to complete baseline data collection to aid in future trend assessments, distribution trends are suggested by some of the data assembled here. For example, recent surveys within the known range of C. reburrus (French Broad Crayfish) yielded few collections. The French Broad River basin is experiencing rapid human population growth and land-use conversion that may affect this species. Other species in the Region, especially those with narrow geographic ranges or specific habitat requirements, may be vulnerable to habitat alterations that often accompany increased human population and land development. A number of species are identified in species accounts as priorities for monitoring based on their small native range, the relative threat from present and anticipated habitat disturbance, or if available survey data suggest decline. Species Accounts The following species accounts summarize distribution, status, and lifehistory and habitat observations for each species known from the Region. Conservation status and rank, if assigned, are listed for each species. See Table 2 for further explanation of species rank and status criteria. Regional range maps for each species, including data collected during this 2004–2005 NCWRC inventory, as well as all other occurrence data assembled during this effort, are displayed spatially in Figures 3–27. Figure 1 provides an overview of the Western Region river basins and counties and should be referred to for geographical reference for Figures 3–27. Cambarus (Cambarus) bartonii (Fabricius) (Common Crayfish; Fig. 3) Distribution and status. Common Crayfish are found from New Brunswick, Canada to northern Georgia and include a variety of forms (Hobbs Figure 3. Collection localities for Cambarus (Cambarus) bartonii (Common Crayfish) in the Western Region of North Carolina. 88 Southeastern Naturalist Vol. 9, Special Issue 3 Table 2. Definitions of conservation rank and status for crayfish species in North Carolina. “Rank” is a numerical scale of the rarity of a species. “Status” indicates the degree of protection, if any, of a species based on rarity. AFS = American Fisheries Society Endangered Species Committee. AFS status information from Taylor et al. (2007). Category/ Rank or status Definition Global rank Based on population size and trend data from throughout the species range, assigned by Nature Serve (www.natureserve.org). G1 Critically imperiled—at very high risk of extinction due to extreme rarity (often 5 or fewer populations), very steep declines, or other factors. G2 Imperiled—at high risk of extinction due to very restricted range, very few populations (often 20 or fewer), steep declines, or other factors. G3 Vulnerable—at moderate risk of extinction due to a restricted range, relatively few populations (often 80 or fewer), recent and widespread declines, or other factors. G4 Apparently secure—Uncommon but not rare; some cause for long-term concern due to declines or other factors. G5 Secure—common; widespread and abundant. GU Unrankable—currently unrankable due to lack of information or due to substantially conflicting information about status or trends; more information is needed. NR Not ranked State Rank Assigned by the North Carolina Natural Heritage Program as the best available scientific and objective assessment of a species’ rarity at the state level (LeGrand et al. 2008). S1 Critically imperiled—critically imperiled in North Carolina due to extreme rarity (often 5 or fewer occurrences) or because of some factor(s) such as very steep declines making it especially vulnerable to extirpation from the state. S2 Imperiled—imperiled in North Carolina because of rarity due to very restricted range, very few populations (typically 20 or fewer), steep declines, or other factors making it very vulnerable to extirpation from the state. S3 Vulnerable—vulnerable to extirpation in North Carolina due to a restricted range, relatively few populations (typically 100 or fewer), recent and widespread declines, or other factors. S4 Apparently secure—mmany occurrences within the state (100–1000 populations). S5 Secure—common, widespread, and abundant in the state (>1000 populations). SNA (SE) Not applicable—a conservation status rank is not applicable because the element is not a suitable target for conservation due to exotic origin. NR Not ranked 2010 J.W. Simmons and S.J. Fraley 89 Table 2, continued. Category/ Rank or status Definition Federal status Designated by the US Fish and Wildlife Service. There are no federally listed endangered or threatened crayfish in North Carolina. FSC Federal species of concern—a species of concern as designated by the US Fish and Wildlife Service. Further research and field study are needed to determine the conservation status. Future listing may or may not be warranted. Such species are the pool from which future candidates for listing will be drawn. Receives no protection under the US Endangered Species Act. State status Species listed as endangered, threatened, and special concern are designated and protected by the NCWRC per Article 25 of Chapter 113 of the NC General Statutes (1987). Those listed as significantly rare or on the Watch List are designated and tracked by the NCNHP, but have no protection. No crayfish in NC have been listed as endangered or threatened by the NCWRC. SC Special concern—any species of wild animal native or once-native to North Carolina which is determined by the Wildlife Resources Commission to require monitoring but which may be taken under regulations adopted under the provisions of (Article 25). SR Significantly rare—any species which has not been listed by the NC Wildlife Resources Commission as an Endangered, Threatened, or Special concern species, but which exists in the state in small numbers and has been determined by the NC Natural Heritage Program to need monitoring. Significantly Rare species include “peripheral” species, whereby North Carolina lies at the periphery of the species' range. W Watch list—any other species believed to be of conservation concern in the state because of scarcity, declining populations, threats to populations, or inadequacy of information to assess its rarity. NS No status assigned AFS status Based on the best available distributional and status information for species range wide. No state or federal protection is afforded by these designations. E Endangered—a species or subspecies in danger of extinction throughout all or a significant portion of its range. T Threatened—a species or subspecies likely to become endangered throughout all or a significant portion of its range. V Vulnerable—a species or subspecies that may become endangered or threatened by relatively minor disturbances to its habitat and deserves careful monitoring of its abundance and distribution. CS Currently Stable—a species or subspecies whose distribution is widespread and stable and is not in need of immediate conservation management actions. NS No status assigned 90 Southeastern Naturalist Vol. 9, Special Issue 3 1981). Likewise, populations assigned to this species in North Carolina are a complex of various forms (J.E. Cooper, pers. comm.). They are found throughout most river basins in the Region, except the Broad and Catawba river basins where they are primarily found in the western headwaters along the Eastern Continental Divide. This species was collected from 94 of the 199 stream sites sampled in 2004–2005. In the broad sense, this species appears to be stable in North Carolina; however, further taxonomic revisions could reveal more vulnerable forms. Ecology and life-history observations. Form I males were collected in April, May, June, August, September, October, November, and December in water temperatures ranging from 7–20 °C. Adults collected from the New River basin were much smaller than those collected elsewhere; the largest form I male from the New River basin was 26 mm TCL, while form I males from other river basins ranged from 26–42 mm TCL. Ovigerous females were collected in April, May, and June in water temperatures ranging from 14–15 °C. Number of ova ranged from 85–104. Common Crayfish are sympatric with most other stream-dwelling species found in the Region. Habitat observations. The Common Crayfish is the most widespread species in the mountains of North Carolina, where it occurs in large rivers to small, high-elevation mountain seeps. It appears to be a habitat generalist, found under rocks, beneath undercut banks, in detritus and woody debris piles, and occasionally in shallow burrows. However, some occurrences from disparate habitats may represent separate but presently unrecognized taxa. Cambarus (Cambarus) eeseeohensis Thoma (Grandfather Mountain Crayfish; Fig. 4) Distribution and status. Grandfather Mountain Crayfish are apparently found only in the Linville River system above Linville Falls in Burke and Avery counties, Catawba River basin. This species is most closely related to C. sciotensis and C. angularus (Thoma 2005), but was lumped with C. bartonii by some taxonomists prior to being described (J.E. Cooper, pers. comm.). Several NCMNS specimens and NCWRC records (Marsh 1998) from the upper Linville River identified as C. bartonii and C. (C.) sp. cf. bartonii likely represent C. eeseeohensis. No surveys were conducted within the range of this species during this inventory. While population trends are unknown, the small range of this species and increasing residential development in the area warrant a high priority for conservation and monitoring. A potential threat to this population is an introduced population of O. virilis in the lower Linville River. Cambarus (Cambarus) howardi Hobbs and Hall (Chattahoochee Crayfish; Fig. 5) Distribution and status. The Chattahoochee Crayfish was formerly known only from the Chattahoochee River system in Georgia and Alabama (Hobbs 1989), but specimens from North Carolina have recently been 2010 J.W. Simmons and S.J. Fraley 91 Figure 4. Collection localities for Cambarus (Cambarus) eesseeohensis (Grandfather Mountain Crayfish), Cambarus (cf. Hiaticambarus) sp. nov.?, and Cambarus (Puncticambarus) hobbsorrum (Rocky River Crayfish) in the Western Region of North Carolina. Figure 5. Collection localities for Cambarus (Cambarus) howardi (Chattahoochee Crayfish) in the Western Region of North Carolina. 92 Southeastern Naturalist Vol. 9, Special Issue 3 identified as this or a closely related species (Clamp 1999). In North Carolina, Chattahoochee Crayfish are known from the upper Broad, Catawba, and Yadkin-Pee Dee river basins (Cooper 2005). In 2004–2005, Chattahoochee Crayfish were collected from 8 of 27 stream sites sampled in the Broad River basin, including the first record from Cleveland County, and at 13 of 35 stream sites in the Catawba River basin, including the first records from Lincoln County. Populations appear to be stable in North Carolina. This species appears to be fairly tolerant of habitat alteration, indicated by its presence in a degraded urban section of Hickory Creek in Shelby. Ecology and life-history observations. Form I males were collected in April, May, June, and November in water temperatures ranging from 13–16 °C. These individuals ranged in size from 22–29 mm TCL. Ovigerous females were collected in April in water 13 °C. Number of ova ranged from 81–90. Chattahoochee Crayfish were most often collected with C. sp. C., but it was also sympatric with C. asperimanus, C. bartonii, C. spicatus, and C. johni. Habitat observations. Chattahoochee Crayfish were collected from streams 1–9 m wide and were found under rocks in mid-stream and along the margins, beneath undercut banks, and in woody debris piles. Cambarus (Cambarus) lenati Cooper (Broad River Stream Crayfish; Fig. 6) Distribution and status. The Broad River Stream Crayfish is endemic to streams in the Broad River basin of North Carolina where it is known from 6 localities in Cleveland County, one in McDowell County, five in Polk County, and 18 in Rutherford County (Cooper 2000, 2002; NCDWQ, Raleigh, NC, unpubl. data). This species was collected at 4 of the 27 stream sites sampled in the Broad River basin during this inventory. Figure 6. Collection localities for (Cambarus) lenati (Broad River Stream Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 93 The Broad River Stream Crayfish appears to be vulnerable to excess sediment. Many streams in the Broad River basin carry heavy loads of sand and fine sediment that degrade slab rock and cobble habitats where this species was collected. Most occurrence data for this species have been obtained in recent years. Of the 32 existing collection records, 31 were made during 1995–2004; thus, population trends are not clear. This endemic species should be a priority for monitoring. Elevation of state conservation status should be considered. Future efforts should focus on assessing population trends and increasing understanding of life history, habitat requirements, and vulnerability to specific threats. Ecology and life-history observations. One form I male, 33 mm TCL, was collected in April in water 17 °C. Other form I males have been collected in mid-June, late September, and late November (Cooper 2000). The first ovigerous female specimens (n = 3) were collected in late April at a water temperature of 17 °C and ranged from 23–30 mm TCL. Number of ova ranged from 18–78. Other species collected with Broad River Stream Crayfish were C. (C.) sp. A and C. spicatus. Habitat observations. The Broad River Stream Crayfish was found only under slab rocks and cobbles in small to medium streams. At one site, where substrate was predominantly sand, this species was found only among riprap at a bridge crossing. Cambarus (Cambarus) sp. A (Fig. 7) Distribution and status. An undescribed form (or forms) presently referred to as C. (C.) sp. A (Cooper 2005, Hobbs and Peters 1977) is in many respects similar to the Chattahoochee Crayfish and is found in the Hiwassee, Little Tennessee, French Broad, Watauga, and New river basins. This crayfish is widely scattered in the New and Hiwassee river basins, but its occurrence is patchy and uncommon. In the New River basin, it was not detected at any of 27 localities sampled by NCWRC in 1996–1997 (McGrath 1998); however, that study relied on field identification, few voucher specimens were retained, Figure 7. Collection localities for Cambarus (Cambarus) sp. A in the Western Region of North Carolina. 94 Southeastern Naturalist Vol. 9, Special Issue 3 and this form could have been overlooked. During this inventory, it was collected at 2 of 13 stream sites surveyed in the New River basin. These are the first records of this crayfish from the South Fork New River system. In the Hiwassee River basin, it was collected at 6 of the 16 stream sites surveyed during this inventory. Enigmatic specimens similar to C. (C.) sp. A found in the Little Tennessee, French Broad, and Watauga river basins are included here, but the taxonomic relationship between these and the other populations is presently unclear (J.E. Cooper, pers. comm.). We collected no specimens identifiable as this form from the French Broad River basin during this inventory. In the Little Tennessee River basin, we collected specimens identifiable as this form from 2 localities in the Cheoah River system. Conservation status is unclear pending further taxonomic clarification. Ecology and life-history observations. Form I males were collected in May, October, and November and ranged in size from 23–25 TCL. This crayfish was found with C. hiwasseensis, C. latimanus, O. sp. cf. spinosus, C. chasmodactylus, C. robustus, and O. cristavarius, as well as streamcollected C. dubius. Habitat observations. In the New River basin, this crayfish was collected from beneath rocks and undercut banks in streams ranging from 1–4 m wide. In the Hiwassee River basin, it was found under rocks in mid-channel and beneath undercut banks. It was most often collected in small to mediumsized streams 1–15 m wide, but was also collected from the Hiwassee River (approximately 40 m wide). Cambarus (Depressicambarus) latimanus (LeConte) (Variable Crayfish; Fig. 8) Distribution and status. The Variable Crayfish is found primarily along the Atlantic and Gulf Coastal Plain and Piedmont Plateau from North Carolina southward to Florida, and westward to the Coosa Basin in Alabama (Hobbs 1989). In the Tennessee River drainage, this species is known only from the headwaters of the Hiwassee River system in North Carolina, Georgia (Hobbs 1981), and Tennessee (Williams et al. 2007). The Variable Crayfish is known from the Hiwassee River basin in North Carolina from 2 collections, one of which was made during this inventory. Both of these collections were from an unnamed tributary to Wolf Creek in extreme southwestern North Carolina, which flows into Georgia and joins the Toccoa River. It was first collected in May 1982, and again 22 years later. This population in the North Carolina mountains appears to be stable at present. Taxonomy of this population is not entirely clear and could be revised (J.E. Cooper, pers. comm.). Habitat observations. This species was collected from a shallow stream 1–2 m wide, where it was sympatric with C (C.) sp. A. Cambarus (Depressicambarus) reduncus Hobbs (Sickle Crayfish; Fig. 9) Distribution and status. The Sickle Crayfish is relatively narrowly distributed in the Piedmont of North and South Carolina (Hobbs 1989). In North Carolina, it is known to occur almost exclusively in the Piedmont 2010 J.W. Simmons and S.J. Fraley 95 Figure 8. Collection localities for Cambarus (Depressicambarus) latimanus (Variable Crayfish) and Cambarus (Puncticambarus) brimleyorum (Valley River Crayfish) in the Western Region of North Carolina. Figure 9. Collection localities for Cambarus (Depressicambarus) reduncus (Sickle Crayfish) and Cambarus (Jugicambarus) dubius (Upland Burrowing Crayfish) in the Western Region of North Carolina. 96 Southeastern Naturalist Vol. 9, Special Issue 3 Plateau, with one population known from the Sandhills (Cooper 2002). Results of this inventory suggest that this species is relatively common within its small range in the Region. Population trends are difficult to assess from available data, but its presence in long-established residential areas suggest that it is tolerant of urban development at some level. We collected this primary burrowing species at 10 localities: 6 in the Catawba River basin, 3 in the Broad River basin, and 1 outside the Region in the Yadkin-Pee Dee River basin. Previously, this species was unknown from the Broad River basin in North Carolina, and our collections there presently represent the western extent of its known range in the state. Additional sampling in the lower Broad River basin in North Carolina is needed to better understand the distribution of this species. Color and life-history observations. Color was variable, ranging from brownish-green to forest green to bluish-green. Specimens from northwest Gaston County (Cherryville) and in central Lincoln County (Lincolnton) were predominantly blue tinted. Those from the Cherryville locality were bluish-green dorsally with chelae that were light blue ventrally. Specimens from the Lincolnton site were brownish-blue dorsally, and the dorsal surface of the chelae were blue, while the ventral surfaces were brownish-orange. One recently molted female from this site was light blue. Specimens from northern Mecklenburg County (Davidson) were forest green dorsally, orange ventro-laterally, and with chelae that had bright orange lateral margins. Specimens from all other localities were uniformly brownish-green. No form I males or ovigerous females were collected. Young-of-year (YOY) were collected from burrows at 2 localities in late July 2005. Total carapace length of YOY ranged from 6.8–7.4 mm. Habitat and burrow observations. This species was frequently found in urbanized areas such as residential neighborhoods where burrows were commonly located on the margins of small first-order streams and seeps. It was also found up to 50 m from open water in mowed lawns and wooded areas. Burrows were typically highly branched with 3 to 5 surface entrances. Many burrows contained a large central “cavern” to which several surface entrances connected. Chimney structures ranged from a mud pile less than 6 cm in height to an elaborate chimney >20 cm in height. Burrow depth varied with depth to the water table. Those located along a stream margin were usually shallow (less than 0.3 m to the water table), but some located on high stream banks were deeper. Burrows that were located farther away from streams were usually much deeper (≥1 m). Soil type where burrows were found was most often hard packed grey or red clay. Cambarus (Hiaticambarus) chasmodactylus James (New River Crayfish; Fig. 10) Distribution and status. The New River Crayfish is found only in the New-Kanawha River basin in northwestern North Carolina, southwestern Virginia, and eastern West Virginia (Hobbs 1989). It occurs throughout the New River basin in North Carolina in Watauga, Ashe, and Alleghany 2010 J.W. Simmons and S.J. Fraley 97 counties. McGrath (1998) found this species at 16 of 27 sites sampled, where it was common to abundant. During this inventory, sites were sampled in areas not previously surveyed, where it was collected from 9 of 13 sites. While this species appears to be stable, its restricted range in North Carolina warrants a place on the North Carolina Natural Heritage Program (NCNHP) watch list. Ecology and life-history observations. Form I males were collected in October in water temperatures ranging from 11–17 °C. These individuals ranged in size from 34–45 mm TCL. This species was commonly found with C. bartonii, C. robustus, and O. cristavarius. Habitat observations. This species was found in streams ranging from 1–14 m wide and was collected from beneath slab rock at all 9 localities. Cambarus (Hiaticambarus) longirostris Faxon (Longnose Crayfish; Fig. 11) Distribution and status. The Longnose Crayfish inhabits Tennessee River tributaries in the Blue Ridge and Valley and Ridge physiographic provinces in Virginia, Tennessee, North Carolina, and Georgia (Hobbs 1989). In North Carolina, this species is common in French Broad River tributaries in Madison County and in the Nolichucky River system in Yancey, Mitchell, and Avery counties. It is also known from one collection from the West Fork French Broad River in Transylvania County. One juvenile individual was identified in 1987 as C. (H.) sp. cf. longirostris from the Little Tennessee River in Macon County (NCMNS, unpubl. data). Figure 10. Collection localities for Cambarus (Hiaticambarus) chasmodactylus (New River Crayfish) in the Western Region of North Carolina. 98 Southeastern Naturalist Vol. 9, Special Issue 3 During this inventory, this species was collected at 2 localities in the South Toe River system in Yancey County in the vicinity of previous collections. Populations of the Longnose Crayfish in North Carolina appear to be presently stable but limited in range. Ecology and life-history observations. Form I males were collected in late May and September in water temperatures ranging from 11–18 °C. These individuals ranged in size from 18–28 mm TCL. Cambarus (cf. Hiaticambarus) sp. nov.? (Fig. 4) Distribution and status. Crayfish with characters that appear to be consistent with the subgenus Hiaticambarus may represent a new species and have recently been collected from Lincoln County (J.E. Cooper, pers. comm.). No other crayfish assigned to the subgenus Hiaticambarus are known from the Catawba River basin. This potentially new species should be a high priority for monitoring. Additional sampling should be conducted in this portion of the Catawba River basin to obtain more material for taxonomic evaluation and to assess the status of this crayfish. An extensive amount of sampling has been conducted in the Catawba River basin (see section on C. (P.) sp. C), and this crayfish has remained undetected until recently. Current data indicates that this animal may have an extremely limited distribution. Based on the one site where we collected this form, it appears to tolerate stream-channel alteration and high sediment loads. Figure 11. Collection localities for Cambarus (Hiaticambarus) longirostris (Longnose Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 99 Ecology and life-history observations. Two form I males were collected in early November and ranged in size from 24–34 mm TCL. This crayfish was found with C. (P.) sp. C. Habitat observations. This species was collected from Muddy Creek in the South Fork Catawba River system in southern Lincoln County. The stream was 2 m wide and channelized. Substrate was dominated by sand and gravel. Cambarus (Jugicambarus) asperimanus Faxon (Mitten Crayfish; Fig. 12) Distribution and status. The Mitten Crayfish is found primarily in streams of the Blue Ridge physiographic province in North Carolina, South Carolina, Georgia, and Tennessee (Hobbs 1989). It is widely distributed in western North Carolina, where it is known from the Little Tennessee River basin east to the Yadkin Pee-Dee River basin. Previous surveys recorded it from primarily smaller, headwater streams in every river basin in the Region except the Hiwassee. Known occurrences are concentrated along the Eastern Continental Divide, especially in headwaters of the Savannah River basin, and headwater streams along both sides of the Catawba-Broad divide. During this inventory, Mitten Crayfish were collected from 2 of 12 stream sites sampled in the Little Tennessee River basin (Tuckasegee River system, Jackson County), 3 of 46 stream sites sampled in the French Broad River basin, one of 35 sites sampled in the Catawba River basin, and 6 of 27 stream sites sampled in the Broad River basin. During this inventory, Mitten Crayfish were not collected from any stream sites in the Watauga or New river basins and are only known from single localities in each basin collected in 1984 Figure 12. Collection localities for Cambarus (Jugicambarus) asperimanus (Mitten Crayfish) in the Western Region of North Carolina. 100 Southeastern Naturalist Vol. 9, Special Issue 3 (NCMNS, unpubl. data). No surveys were conducted in streams where they are known to occur in the Savannah River basin. Many of the headwater streams along the Eastern Continental Divide where Mitten Crayfish are relatively common are within the boundaries of the Nantahala and Pisgah national forests and are relatively safe from habitat degradation. Potential long-term impacts from factors such as loss of native trees to exotic insect pests or atmospheric (e.g., acid deposition) and climate changes are difficult to predict, but may be detrimental. Mitten Crayfish populations appear to be relatively stable at present. Color, ecology, and life-history observations. Mitten Crayfish encountered ranged from deep rusty to bright red. Form I males were collected during June, August, and November in water temperatures ranging from 13–16 °C. These individuals ranged in size from 27–29 mm TCL. Mitten Crayfish were found sympatric with C. howardi, C. (P.) sp. C, C. robustus, and C. bartonii. Habitat observations. Mitten Crayfish were found in spring seeps and headwater streams under rocks and in shallow burrows. Throughout its range, this species seems to prefer small headwater streams of moderate to high gradient at higher elevations. Cambarus (Jugicambarus) carolinus (Erichson) (Red Burrowing Crayfish; Fig. 13) Distribution and status. The Red Burrowing Crayfish is reported from northeastern South Carolina, western North Carolina, and eastern Tennessee (Cooper and Braswell 1995, Dewees 1972, Williams et al. 2007). In North Carolina, it is known from 14 localities in the Little Tennessee River basin Figure 13. Collection localities for Cambarus (Jugicambarus) carolinus (Red Burrowing Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 101 in Clay, Graham, Jackson, Macon, and Swain counties (Cooper and Braswell 1995; NCWRC, unpubl. data) and a single locality in the Hiwassee River basin in Cherokee County (Cooper et al. 1998). Eight of the 14 presently known localities in the Little Tennessee River basin were discovered during this inventory. While the known distribution of this species is patchy, suitable habitats appear to be common throughout its range in the Region. Additional surveys would likely find more populations. Six additional populations were discovered in Swain and Graham counties, where only 2 records existed prior to this inventory. More undetected populations may exist within the boundaries of Great Smoky Mountains National Park (GSMNP) and Nantahala National Forest, where the likelihood of long-term habitat degradation is relatively low. Some populations on private lands may have been extirpated by, and existing populations may be vulnerable to, removal or degradation of riparian vegetation. Buffer regulations on designated trout streams offer some habitat protection. Overall status appears stable presently, but its limited range and potential threats to habitat on private land warrants its place on the NCNHP watch list. Color and life-history observations. All specimens collected were bright red. Form I males were collected in mid-October. An ovigerous female, 27 mm TCL, was collected in late June. This female possessed 51 ova. Form I males have been collected in North Carolina in April and May and one other ovigerous female was collected in late April (NCMNS, unpubl. data). Habitat and burrow observations. While this species was always collected from burrows, the situation and form of burrows varied. Most burrows were shallow (≤10 cm) in seeps and other damp areas immediately adjacent to small streams. Those burrows were more or less horizontal, often just under rocks and decaying logs that lay on the surface, and branched to multiple surface entrances. A few burrows were located up to 3 m from surface water and were relatively deep with several branches and well developed chimneys. Cambarus (Jugicambarus) dubius Faxon (Upland Burrowing Crayfish; Fig. 9) Distribution and status. The Upland Burrowing Crayfish is a primary burrower found in the Appalachian Mountains from Maryland, Pennsylvania, West Virginia, eastern Kentucky, southwestern Virginia, eastern Tennessee, and western North Carolina (Dewees 1972, Hobbs 1989, Taylor et al. 2007). In North Carolina, it is relatively widely distributed in the French Broad, Watauga, New, upper Catawba, Broad, and upper Yadkin-Pee Dee river basins. During this inventory, this species was collected at 35 sites, including localities in 7 counties in the French Broad, Broad, and Catawba River basins where it was previously unknown. This species is commonly encountered within its range in the Region, often with many burrows and individuals found at a site. Residential development is increasing in much of the species’ range; but as is true of the Sickle Crayfish, its persistence 102 Southeastern Naturalist Vol. 9, Special Issue 3 appears to be compatible with some level of development. Presently, this species appears stable. Some crayfish taxonomists consider this a species complex in need of revision (J.E. Cooper, pers. comm.). Color and life-history observations. We observed 2 color forms: red and dark blue, which were clumped geographically. With one exception, specimens from the southwestern portion of its range in the Pigeon and French Broad river systems (Buncombe, Haywood, and Henderson counties) were uniformly red. One population sampled in eastern Buncombe County in the French Broad River system (tributary to the Swannanoa River) was blue. All other specimens from the Broad, Catawba, Watauga, New, and Yadkin-Pee Dee river basins, and from the Nolichucky River system in the French Broad River basin, were blue. Form I males were collected in June, July, September, and October and ranged in size from 29–36 mm TCL. Ovigerous females were collected in mid-May and early June. The female collected during May was 34 mm TCL and possessed 49 ova. Water temperature in the burrow was 14 °C. Multiple YOY (TCL 6.0–8.6 mm) were collected from several burrows between late May and early August. Habitat and burrow observations. Burrows were found along banks of small streams, wetland seeps, and up to 50 m from surface water bodies in yards and damp wooded areas. Burrow depth varied from approximately 15 cm to 1 m. Deeper burrows were located farther from surface water bodies. Chimneys were usually present at the burrow entrance and ranged from a large mud pile to a more developed chimney approximately 15 cm in height. Burrows were usually highly branched, with as many as 5 surface entrances. On one occasion, 2 adult females, one of which was ovigerous, were collected from the same burrow. All other sampling efforts yielded only one adult per occupied burrow. This species can occur in rather large colonies. In 2005, a McDowell County landowner collected 29 specimens while excavating a small area with power equipment where 14 specimens were collected in previous months. Cambarus (Jugicambarus) nodosus Bouchard and Hobbs (Knotty Burrowing Crayfish; Fig. 14) Distribution and status. The Knotty Burrowing Crayfish is known only from the headwaters of the Savannah, Chattahoochee, and Hiwassee river systems in Georgia, South Carolina, Tennessee, and North Carolina (Hobbs 1989, Williams et al. 2007). In North Carolina, this species has been found at only 5 localities, three of which are clumped closely in southwestern Cherokee County. The other 2 localities were discovered during this inventory in southeastern Cherokee County and in Clay County, east of Chatuge Reservoir. This species is deemed “significantly rare” in North Carolina due to its limited distribution. The known distribution remains small, and population trends are not clear, but the discovery of 2 additional populations suggests that more may be present in the southwestern Hiwassee 2010 J.W. Simmons and S.J. Fraley 103 River basin. Given its narrow distribution and few known occurrences, this species is potentially vulnerable to extirpation from the state and should be a priority for monitoring. Color and life-history observations. Specimens varied in color from uniformly bright blue-green to dull bluish-green with yellow-orange on the tips and ventral surface of chelae. No form I males or ovigerous females were collected during this inventory; however, 8 YOY (7.4–9.6 mm TCL) were collected in mid-July. Habitat and burrow observations. This species burrows in stream banks, bogs, seeps, and intermittent stream channels, but can occasionally be found under surface rocks in such areas, and even in quieter sections of streams (Bouchard and Hobbs 1976, Cooper and Braswell 1995, Hobbs 1981). At the Clay County site, the species was found in burrows in a short spring seep that drained to a 2-m-wide first-order stream approximately 30 m away. The seep was covered in dense wetland vegetation; some areas contained shallow standing water, but most of the flow was subterranean. The surrounding area was a mowed lawn. Burrows lacked distinct chimneys but were surrounded by piles of mud. At the time of our survey, the water table was near the surface, and the burrows continued vertically for approximately 0.5 m below the water level. Many were highly branched, with up to 5 surface openings. No burrows were evident more than a few meters from the vegetated seep, and none were seen along the first-order stream into which the seep drained. At the southeastern Cherokee County locale, specimens were associated with vegetation Figure 14. Cambarus (Jugicambarus) nodosus (Knotty Burrowing Crayfish) and Cambarus (Tubericambarus) acanthura (Thornytail Crayfish) in the Western Region of North Carolina. 104 Southeastern Naturalist Vol. 9, Special Issue 3 and undercut banks in a small, mud-bottomed stream less than 0.5 m wide, and were syntopic with C. hiwasseensis (Hiwassee Crayfish). Several burrow entrances were observed along the stream bank, with none more than a few meters from the stream. No specimens were found in the burrows, several of which were similar to burrows at the Clay County site, while others were shallow, horizontal passages with short vertical branches. Cambarus (Jugicambarus) tuckasegee Cooper and Schofield (Tuckasegee Stream Crayfish; Fig. 15) Distribution and status. The Tuckasegee Stream Crayfish is endemic to the Tuckasegee River system of the Little Tennessee River basin in Jackson and Swain counties (Cooper and Schofield 2002). This species is restricted to a relatively small area where it occurs within the boundaries of the GSMNP, the Eastern Band Cherokee Indian Reservation (EBCIR), and on private lands (Oconoluftee River system, Deep and Cooper creeks). During 2004–2005, this species was collected at 5 of the 12 stream sites sampled in the Tuckasegee River system, which represent 5 of the 8 localities known for this species. Collections upstream from previously known occurrences produced only C. bartonii. While no population trend information is available at present, the relative protection of much of the Tuckasegee Stream Figure 15. Collection localities for Cambarus (Jugicambarus) tuckasegee (Tuckasegee Stream Crayfish), Cambarus (Puncticambarus) georgiae (Little Tennessee River Crayfish), and Orconectes (Procericambarus) sp. nov.? (cf. spinosus) (“Cheoah Crayfish”) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 105 Crayfish’s range within the GSMNP should provide long-term population viability. Streams in the EBCIR and on other private lands may be vulnerable to habitat degradation; however, this species’ tolerance to common impacts (e.g., sedimentation, eutrophication) is unknown. Given the very restricted range of the Tuckasegee Stream Crayfish, its status as “significantly rare” in North Carolina is definitely warranted, and a higher designation should be considered if future data indicate declines. This species should be a priority for monitoring. Ecology and life-history observations. At 4 of the 5 localities where Tuckasegee Stream Crayfish were collected in this inventory, C. bartonii was also present. Habitat observations. Most individuals were found along the margins of relatively high-gradient streams with substrates dominated by boulder and cobble. Cambarus (Puncticambarus) brimleyorum Cooper (Valley River Crayfish; Fig. 8) Distribution and status. The Valley River Crayfish is a recently described species endemic to the Hiwassee River basin, where it is known from the Valley River system in Cherokee County and from tributaries of the upper Hiwassee River (Fires and Tusquitee creek systems) in Clay County (Cooper 2006b). Population trends are difficult to determine from the predominately recent collection data presently available. While this species’ vulnerability to common perturbations is unknown, residential development is increasing in the Valley River watershed and throughout the Hiwassee River basin. Populations are potentially vulnerable to habitat degradation. Given the very restricted range of the Valley River Crayfish, its status as “significantly rare” in North Carolina is definitely warranted and a higher designation should be considered if future data indicate declines. This species should be a high priority for monitoring. Future efforts should focus on assessing population trends and improving our understanding of its life history and habitat requirements. Ecology and life-history observations. An ovigerous female was collected from the Valley River in late May. Habitat observations. Valley River Crayfish were found in small to medium-sized streams with substrate dominated by cobble. Cambarus (Puncticambarus) chaugaensis Prins and Hobbs (Chauga Crayfish; Fig. 16) Distribution and status. The Chauga Crayfish is endemic to the upper Savannah River basin in Georgia, South Carolina, and North Carolina (Cooper et al. 1998). No sampling was done within the known range of this species during this inventory. Fullerton (2002) reported it from 10 of 28 sites surveyed in the Savannah River basin, but many specimens were not identified with confidence and were catalogued as C. (P.) cf. chaugaensis (J.E. Cooper, pers. comm.). We collected similar confusing specimens 106 Southeastern Naturalist Vol. 9, Special Issue 3 from 2 localities in the Savannah River basin, Clay County, identified as Cambarus (C.) sp?, which are included on the Chauga Crayfish range map (Fig. 16). Much of this species’ range in North Carolina is within the Nantahala National Forest, where potential threats may be fewer. This species should be a priority for monitoring due to its limited distribution and lack of recent survey data. Cambarus (Puncticambarus) georgiae Hobbs (Little Tennessee River Crayfish; Fig.15) Distribution and status. The Little Tennessee River Crayfish is found only in the upper Little Tennessee River basin in Macon, Jackson, and Swain counties, NC, and Rabun County, GA (Cooper 2002). It is apparently restricted to the Little Tennessee and Tuckasegee River systems upstream from Fontana Reservoir, but is absent from the Nantahala and Oconaluftee River systems. The Little Tennessee River Crayfish prefers streams of medium to low gradient and seems to be tolerant of at least moderate levels of wastewater pollution and sedimentation (McLarney 1993). This species was collected from only 1 of 13 sites within its range sampled during this inventory, but few of these sites were at localities where the species had been collected in the past. This species should be a priority for future monitoring due to lack of recent survey data throughout much of its range in North Carolina. Ecology and life-history observations. This species was collected with C. bartonii in Alarka Creek. It was also collected with C. reburrus in Alarka and Connelly creeks (Cooper 2002). Figure 16. Collection localities in the Western Region of North Carolina for Cambarus (Puncticambarus) chaugaensis (Chauga Crayfish), and specimens identifiable only as Cambarus (Cambarus) sp. in the Savannah River basin in Clay County. 2010 J.W. Simmons and S.J. Fraley 107 Cambarus (Puncticambarus) hiwasseensis Hobbs (Hiwassee Crayfish; Fig. 17) Distribution and status. The Hiwassee Crayfish is endemic to the upper Hiwassee River basin in Georgia; Cherokee and Clay counties, NC, and one locality in Polk County, TN (Hobbs 1981, Williams et al. 2007). During this inventory, this species was collected at 7 of the 20 sites sampled in Cherokee and Clay counties. The majority of known localities in North Carolina were discovered since 1994, which include localities in close proximity to older records. In the smaller headwater tributaries, it is apparently replaced by C. parrishi (Hiwassee Headwater Crayfish). Recent data indicate that the Hiwassee Crayfish is currently stable in North Carolina; however, residential development is increasing throughout much of its range. Due to its narrow distribution, this species should be a priority for monitoring. Ecology and life-history observations. Two form I males were collected from the Hiwassee River in late November, and 11 YOY were collected during mid-July. In North Carolina, Hiwassee Crayfish have been found with C. bartonii, C. (C.) sp. A., and stream-collected C. nodosus. The first known syntopic occurrence with C. parrishi was found in Hothouse Branch in the Shooting Creek system in Clay County during this inventory. Habitat observations. Most collections of this species were made in small streams 1–4 m wide, but one collection was made in the Hiwassee River (approximately 40 m wide). A single specimen was collected from a shallow burrow alongside Grape Creek, Cherokee County. This site is impounded by Hiwassee Reservoir at full pool. This species was collected Figure 17. Collection localities for Cambarus (Puncticambarus) hiwasseensis (Hiwassee Crayfish) in the Western Region of North Carolina. 108 Southeastern Naturalist Vol. 9, Special Issue 3 beneath undercut banks from moderate- to low-gradient streams 0.5–2.0 m wide at 3 sites sampled in mid-July 2005. Cambarus (Puncticambarus) hobbsorum Cooper (Rocky River Crayfish; Fig. 4) Distribution and status. The Rocky River Crayfish is 1 of 3 recently described species in North Carolina that were previously included under the name Cambarus acuminatus (Cooper 2006a). This species is currently known from the Cape Fear, Yadkin-Pee Dee, Catawba, Neuse, Tar-Pamlico, and Roanoke river basins in North Carolina and from a few localities in the Catawba River basin in South Carolina (Cooper 2001; Taylor et al. 2007; J.E. Cooper, pers. comm.). Few collections of this species have been made in the Catawba River basin in North Carolina, which represents the northwestern extent of its range, but it was not collected during this inventory. Although this species does not appear to be common in the Region, it has a fairly wide distribution in the eastern portion of North Carolina, where it is considered to be currently stable (Taylor et al. 2007). Cambarus (Puncticambarus) johni Cooper (Carolina Foothills Crayfish; Fig. 18) Distribution and status. The Carolina Foothills Crayfish is a recently described species endemic to the upper Broad, Catawba, and Yadkin-Pee Figure 18. Collection localities for Cambarus (Puncticambarus) johni (Carolina Foothills Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 109 Dee river basins in North Carolina (Cooper 2006a). It was identified from 3 localities in the Catawba River basin during this inventory. Carolina Foothills Crayfish were collected by NCDWQ at several sites in the Broad River basin in 2005. Based on available data from the past 10 years, the Catawba and Broad River basin populations appear to be stable. Due to its narrow distribution, this species should be a priority for monitoring. Ecology and life-history observations. Carolina Foothills Crayfish were found with C. howardi and C. (P.) sp. C. Habitat observations. Carolina Foothills Crayfish were collected from streams 3–4 m wide with predominately sand substrate. It was found beneath undercut banks and in leaf packs and root wads. Cambarus (Puncticambarus) parrishi Hobbs (Hiwassee Headwater Crayfish; Fig. 19) Distribution and status. The aptly named Hiwassee Headwater Crayfish is endemic to the upper Hiwassee River basin in Towns County, GA and Clay County, NC (Cooper and Braswell 1995, Hobbs 1981). This species occurs in rocky headwater streams. The majority of the known localities in North Carolina were documented from 1994–2004. While population trends are not clear, it may be vulnerable to increasing development in the area. This species should be a high priority for monitoring due to its very narrow distribution. Ecology and life-history observations. The first known syntopic occurrence with C. hiwasseensis (Hiwassee Crayfish), which occupies a much wider range in the Hiwassee River basin, was found in Hothouse Branch in the Shooting Creek system in Clay County. Hiwassee Headwater Crayfish was also found with C. bartonii. Figure 19. Collection localities for Cambarus (Puncticambarus) parrishi (Hiwassee Headwater Crayfish) in the Western Region of North Carolina. 110 Southeastern Naturalist Vol. 9, Special Issue 3 Habitat observations. The only other collection of Hiwassee Headwater Crayfish made during this inventory was in Johnson Creek in Clay County, where it was associated with woody debris in both swift water and eddies. This shallow, high-gradient stream was 3–4 m wide with substrate dominated by cobbles and boulders. Cambarus (Puncticambarus) reburrus Prins (French Broad Crayfish; Fig. 20) Distribution and status. The French Broad Crayfish is endemic to North Carolina and is known from the upper French Broad, Savannah, and Little Tennessee river basins (Cooper 2002, Cooper and Braswell 1995). During this inventory, French Broad Crayfish were collected at only 3 of 15 stream sites that we sampled and at none of 23 sites sampled by cooperators within its known range in the French Broad River basin. In the Little Tennessee River basin, this species is known only from the upper Tuckasegee River system, Jackson County, from 4 localities surveyed during 1999–2003. No French Broad Crayfish were collected at 12 sites that we sampled and at 6 sites sampled by NCDWQ in the upper Tuckasegee River system during 2004–2005. It is also known from only 4 localities in the upper Savannah River basin in Jackson and Transylvania counties, two of which were from 2001. No surveys were made within the range of the French Broad Crayfish in the Savannah River basin during this inventory. Figure 20. Collection localities for Cambarus (Puncticambarus) reburrus (French Broad Crayfish) and Cambarus (Puncticambarus) sp. nov. in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 111 The French Broad Crayfish may be declining. Several of the localities sampled in 2004– 2005 that were considered within its putative range were not in specific streams that were previously known to support it, and may illustrate the patchy nature of this species’ distribution. However, a number of sites where it was not detected were at or near known historical occurrences. Much of the range of the French Broad Crayfish is on private land where human population has increased substantially in recent years. This species should be a high priority for monitoring. Future efforts should focus on assessing population trends and improving our understanding of its life history, habitat requirements, and vulnerability to specific threats. Elevation of state conservation status should be considered. Ecology and life-history observations. The French Broad Crayfish was collected with C. bartonii in Avery Creek, Buncombe County. Cambarus (Puncticambarus) robustus Girard (Big Water Crayfish; Fig. 21) Distribution and status. The Big Water Crayfish is found from Ontario and Quebec, west to Illinois and Michigan, and south to Tennessee, Virginia, and North Carolina (Hobbs 1989, Taylor et al. 2007). In North Carolina, it is known from the New, Watauga, and French Broad river basins. During this inventory, this species was collected at the majority of sites surveyed within those river basins. Oddly, Big Water Crayfish was not reported by McGrath (1998) from any of the 27 sites that they surveyed in the New River basin, but it was collected from 12 of the 14 stream sites sampled there during this inventory. While it may not have been collected from some sites reported in Figure 21. Collection localities for and Cambarus (Puncticambarus) robustus (Big Water Crayfish) in the Western Region of North Carolina. 112 Southeastern Naturalist Vol. 9, Special Issue 3 1998, it is also possible that it was overlooked in preliminary field identifi- cations at some sites where no voucher specimens were retained. Big Water Crayfish populations in North Carolina appear to be stable at present. Ecology and life-history observations. Form I males were collected during September and October in water temperatures ranging from 14–20 °C. These individuals ranged in size from 36–46 mm TCL. Young-of-year as small as 7 mm TCL were collected in mid-October. In the French Broad River basin, Big Water Crayfish were occasionally collected with C. bartonii, C. asperimanus, and C. longirostris. In the New River basin, it was commonly collected with C. bartonii, C. chasmodactylus, O. cristavarius. It was not collected with any other species in the Watauga River basin. Habitat observations. This species appears to be fairly general in its habitat requirements. It was collected from small headwater streams approximately 1 m wide to rivers ≥14 m in width, where it was found under slab rock and cobble in pools, mid-stream in flowing water, and along stream margins. Juveniles were commonly collected from beneath undercut banks or in submerged woody debris. Cambarus (Puncticambarus) spicatus Hobbs (Broad River Spiny Crayfish; Fig. 22) Distribution and status. The Broad River Spiny Crayfish is endemic to the Broad River basin in South Carolina and North Carolina (Cooper et al. 1998). In North Carolina, it is known from Cleveland, Polk, and Rutherford counties (Cooper 2002, Cooper and Braswell 1995). The majority of known occurrences are in the First Broad River system in northeast Rutherford and Cleveland counties. During this inventory, Broad River Spiny Crayfish were collected at 4 of the 27 stream sites sampled in the Broad River basin. It was represented by a single specimen in all but one Figure 22. Collection localities for and Cambarus (Puncticambarus) spicatus (Broad River Spiny Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 113 of those collections. This apparent low abundance was also true for many recent collections from both North and South Carolina (NCDWQ, unpubl. data; J. Price, South Carolina Division of Natural Resources, Eastover, SC, pers. comm.). While trends are not clear, this narrowly distributed crayfish may be declining. Many streams within in its range in North Carolina carry heavy sediment loads that can degrade habitat for this species. Broad River Spiny Crayfish may also be threatened by the non-native O. rusticus (Rusty Crayfish) which has been collected within its putative range. This species should be a high priority for monitoring. Future efforts should focus on assessing population trends and improving our understanding of its life history, habitat requirements, and vulnerability to specific threats. Elevation of conservation status should be considered. Ecology and life-history observations. A form I male, 38 mm TCL, was collected during early November in a water temperature of 15 °C. Youngof- year, 7.4 mm TCL, were also collected in early November. Broad River Spiny Crayfish were most commonly collected with C. (P.) sp. C, but also with C. howardi and C. lenati. Habitat observations. Broad River Spiny Crayfish were collected from beneath undercut banks, woody debris piles, and from cobble along the stream margin; similar to habitat reported by Hobbs (1956) and Eversole (1995). Cambarus (Puncticambarus) sp. C (C. acuminatus complex) (Fig. 23) Distribution and status. Cambarus acuminatus Faxon (s.l.) is widespread from Maryland to South Carolina (Hobbs 1989). There are many variant populations and it is generally considered to be a complex of similar, Figure 23. Collection localities for Cambarus (Puncticambarus) sp. C, (C. (P.) acuminatus complex) in the Western Region of North Carolina. 114 Southeastern Naturalist Vol. 9, Special Issue 3 undescribed species (Cooper 2001, Hobbs 1989, Taylor et al. 2007). North Carolina populations of this complex are referred to as C. (P.) sp. C (Cooper and Braswell 1995, Hobbs and Peters 1977). Three recently described North Carolina species, C. hobbsorum, C. hystricosus, and C. johni, were the result of an ongoing analysis of this complex (Cooper 2006a). Populations referred to as C. (P.) sp. C in this account represent another form in this complex and are presently under investigation by J.E. Cooper. This form occurs in most river basins in the Piedmont Plateau and Coastal Plain of North Carolina. During this inventory, it was collected from 20 of the 26 stream sites sampled in the Broad River basin and 32 of the 35 stream sites sampled in the Catawba River basin. Better taxonomic resolution may allow more relevant conservation status assignment; however, the form or forms native to the Region appear to be widespread, abundant, and stable at present. Ecology and life-history observations. Form I males were collected March–May and September–November in water temperatures ranging from 13–21 °C. These individuals ranged in size from 31–44 mm TCL. A single ovigerous female, 32 mm TCL, was collected in mid-April in a water temperature of 13 °C. This female possessed 161 ova. Young-of-year, <10 mm TCL, were collected April–July and in November. This species is syntopic with most stream dwelling crayfishes found within its range. Habitat observations. This species was found in streams 1–75 m wide and was most often collected from beneath undercut banks, root wads, or woody debris. Cambarus (Puncticambarus) sp. nov. (Fig. 20) Distribution and status. This form, similar in appearance to C. robustus (Big Water Crayfish), is known from the Pigeon River system in the French Broad River basin and from Hazel Creek in the Little Tennessee River basin (Fontana Reservoir). It is believed to be an undescribed species (J.E. Cooper, NCMNS, and R.F. Thoma, Midwest Biodiversity Institute, Columbus, OH, pers. comm.). During this inventory, this crayfish was collected at 3 of the 17 sites sampled in the Pigeon River system, Haywood County, and in Hazel Creek, GSMNP, Swain County. Most other tributaries along the north shore of Fontana Reservoir within GSMNP were surveyed during 2004–2005, either by NCWRC or others, and this species was not encountered (T.W. Savidge, The Catena Group, Inc., Hillsborough, NC, pers. comm.), nor was it found at any other site in the Little Tennessee River basin. This crayfish appears to occupy a very small and disjunct range; however, there may be as yet undetected populations in the Tuckasegee River system (Little Tennessee Basin) within the EBCIR and GSMNP. Most records of this species are from 1999–2004, and it has only recently been recognized as a new species. Population trends are not clear. While the majority of known populations are within the boundaries of the GSMNP and Pisgah National Forest, where substantial habitat degradation is less likely, residential development is increasing rapidly on surrounding private lands in Haywood County. This species should be a priority for monitoring. Better taxonomic 2010 J.W. Simmons and S.J. Fraley 115 resolution, longer term time-series data, and a better understanding of its distribution, especially in the Little Tennessee River basin, are needed to assess the conservation status of this species. Ecology and life-history observations. This crayfish was collected with C. bartonii. One large specimen was feeding on a salamander when collected. Habitat observations. This crayfish was collected in the Pigeon River system from under large slab boulders and cobble in streams 2–3 m wide with moderate to high gradient and little sediment deposition. Cambarus (Tubericambarus) acanthura Hobbs, 1981 (Thornytail Crayfish, Fig. 14) Distribution and status. The Thornytail Crayfish is found from north-central Alabama and southeastern Tennessee, through north-central Georgia to the Hiwassee River basin in North Carolina (Hobbs 1981, Williams et al. 2007). In North Carolina, it is known from a few localities in floodplains near the confluence of the Nottely and Hiwassee rivers (Hiwassee Reservoir) in Cherokee County (Cooper and Braswell 1995). We collected this species at 3 additional sites within the same general area. These populations are at the northeastern extreme of the species’ known distribution. Population trends are not clear, but its restricted range and apparent low densities make it potentially vulnerable to extirpation from North Carolina. Color and life-history observations. Specimens were predominantly grey-brown, often with pale to bright orange on the chelae and sides of the carapace. No form I males, ovigerous females, or YOY were collected in our May and July samples. Habitat and burrow observations. This species was dug from shallow streamside burrows at 2 sites in the floodplain of Hiwassee River along small streams that are impounded by Hiwassee Reservoir at full pool. A third site was located along a private reservoir, where specimens were dug from burrows in a wet area of a mowed lawn. One juvenile female was found under a rock along the reservoir margin. Burrows were in clay soil, and were located up to 5 m from the waters edge. Approximately 20 burrow entrances, surrounded by small mud piles, were seen at this site. Orconectes (Procericambarus) cristavarius Taylor (Spiny Stream Crayfish; Fig. 24) Distribution and status. The Spiny Stream Crayfish is found in the middle Ohio River drainage from the Cumberland River system in Kentucky and Tennessee east to the New River basin in North Carolina and Virginia (Taylor 2000). In North Carolina, it occurs throughout most of the New River basin and has apparently been introduced into the Yadkin-Pee Dee River basin (Cooper 2005). McGrath (1998) found Spiny Stream Crayfish to be common to abundant at all 27 sites sampled in the New River basin. During this inventory, it was collected at 7 of the 13 stream sites surveyed in the New River basin and it has been collected from many localities during 116 Southeastern Naturalist Vol. 9, Special Issue 3 1996–2005 in the vicinity of earlier collections. This species appears to be currently stable in North Carolina. Ecology and life-history observations. Spiny Stream Crayfish were commonly sympatric with C. robustus, C. chasmodactylus, and C. bartonii. Form I males were collected during mid-October in water temperatures ranging from 11–17 °C. These individuals ranged in size from 20–33 mm TCL. Juveniles, 15–18 mm TCL, were also collected in mid-October. Habitat observations. Streams where Spiny Stream Crayfish were collected ranged from 2–14 m in width. Adults were usually found under large slab rocks or cobble both in midstream and along the stream margin. Juveniles were often collected from beneath undercut banks. Orconectes (Procericambarus) sp. nov.? (cf. O. spinosus) (“Cheoah” Crayfish; Fig. 15) Distribution and status. This form, similar to Orconectes spinosus (Coosa River Spiny Crayfish), is known from 4 streams in the Cheoah River system in the Little Tennessee River basin, Graham County, upstream from Santeetlah Reservoir. As presently understood, the distribution of the Coosa River Spiny Crayfish is limited to the Coosa River drainage in Alabama, Georgia, and Tennessee (Taylor 2000). Preliminary study suggests that this form may represent an undescribed species (Cooper 2004b; C.A. Taylor, Illinois Natural History Survey, Champaign, IL, pers. comm.). During 2003–2005, surveys at 17 localities along the entire length of the Cheoah River and its Figure 24. Collection localities for Orconectes (Procericambarus) cristavarius (Spiny Stream Crayfish) in the Western Region of North Carolina. 2010 J.W. Simmons and S.J. Fraley 117 tributaries downstream from Santeetlah Reservoir did not detect “Cheoah” Crayfish. One new occurrence was found upstream from Santeetlah Reservoir in Snowbird Creek. If described, this crayfish species would be one of the rarest in North Carolina. Loss of any presently occupied habitat would make this species very vulnerable due to its already limited distribution. Due to its relatively recent recognition as a putative species and few historical records, population trends are not clear. All known occurrences are on private or EBCIR lands. Elevation of conservation status should be considered. This species should be a high priority for monitoring. Ecology and life-history observations. “Cheoah” Crayfish were found with Cambarus (C.) sp. A. Habitat observations. “Cheoah” Crayfish were collected from medium to large creeks among cobbles and small boulders near the mid-channel. Orconectes (Gremicambarus) virilis Hagen (Virile Crayfish; Fig. 25) Distribution and status. The native range of the Virile Crayfish extends from Hudson Bay southeastward through New England and west through the upper Ohio River drainage and the Missouri and upper Mississippi river basins to western Montana, south to Oklahoma and northern Arkansas Figure 25. Collection localities for Orconectes (Gremicambarus) virilis (Virile Crayfish) in the Western Region of North Carolina (circles represent sites sampled for stream-dwelling crayfish in the Catawba River basin during 2004–2005 where this species was not encountered). 118 Southeastern Naturalist Vol. 9, Special Issue 3 (Pflieger 1996). It has been widely introduced outside of its native range. Presently, introductions of this species in North Carolina are known only in the Catawba River basin. It is known from Long and Mauney creeks in Gaston County, and in the Linville River and White Creek, tributaries to Lake James in Burke County (Cooper and Armstrong 2007). Virile Crayfish in North Carolina were first discovered in Long Creek, a tributary to the South Fork Catawba River, in July 1990 and were recently collected from this tributary in July 2004 (Cooper and Armstrong 2007). No Virile Crayfish were found in recent surveys elsewhere in the South Fork Catawba system. This species was first detected in the Linville River in 1993, a few kilometers upstream from Lake James (Cooper et al. 1998, Marsh 1998). It was collected in 2005 from the Linville River in the vicinity of earlier collections, and a previously unknown population was found in White Creek, a nearby tributary to Lake James. Virile Crayfish were not found at any other site surveyed in the area. Invasion upstream in the Linville River could threaten the endemic Grandfather Mountain Crayfish which occurs upstream from Linville Falls. Pflieger (1996) reported that this species is more abundant in waters without strong base flows in Missouri. Further upstream range expansion in the Linville Gorge may be hampered by high stream gradient and swift water. Monitoring for further expansion of this invasive species should be a priority. Measures to prevent new introductions and further expansion should be considered. Ecology and life-history observations. Virile Crayfish were collected with C. (P.) sp. C. A form I male and a female were collected in copulation while snorkeling in late March in the Linville River. An ovigerous female, 27 mm TCL, was also collected. This individual possessed 103 ova. Orconectes (Procericambarus) rusticus (Girard) (Rusty Crayfish; Fig. 26) Distribution and status. The Rusty Crayfish is native to tributaries of the Ohio River in southwestern Ohio, central Kentucky, eastern and central Indiana, and the western Lake Erie drainage in southeastern Michigan and northwestern Ohio (Taylor 2000). Recently it has spread, likely through baitbucket introductions, as far north as Maine and Ontario, south to Tennessee, and west to New Mexico (Hobbs and Jass 1988). When introduced, negative impacts include destruction of aquatic vegetation, direct competition with native crayfishes, possible hybridization, and predation on other aquatic species (Hobbs et al. 1989, Lodge et al. 2000). Known introductions of the Rusty Crayfish in North Carolina are limited to 3 sites on the Broad River in Rutherford County, and Sawmill Creek in the Little Tennessee River basin, Swain County (Cooper 2005, Cooper and Armstrong 2007, Fullerton and Watson 2001). The Broad River population was discovered in September 1999, and the most recent collections were made in 2002 (Cooper and Armstrong 2007). This population was known from just downstream of Lake Lure to about 10 km downstream. Rusty Crayfish were not encountered during 2004–2005 surveys in several tributaries in that 2010 J.W. Simmons and S.J. Fraley 119 general area. Native endemic species C. lenati and C. spicatus are potentially threatened by further invasion of this species. Cambarus spicatus and Rusty Crayfish are reported from the same reach of the Broad River. Monitoring for expansion of Rusty Crayfish and status of native species in the Broad River should be a high priority. Rusty Crayfish were first collected in Sawmill Creek, a Little Tennessee River tributary, in 2001. In 2005, surveys at this site and elsewhere in the surrounding area failed to detect Rusty Crayfish. While not conclusive, these results suggest that Rusty Crayfish have not spread in detectable numbers and may not have established a nascent population. The Rusty Crayfish would potentially threaten Little Tennessee River basin endemics such as C. tuckasegee, C. georgiae, and the undescribed C. (P.) sp. nov. Further monitoring of Sawmill Creek and surrounding tributaries to the Little Tennessee River and Fontana Reservoir should be a priority. Procambarus (Ortmannicus) acutus (Girard) (White River Crawfish; Fig. 27) Distribution and status. The native range of the White River Crawfish extends along the Atlantic Coastal Plain and Piedmont Plateau from Maine to Georgia, along the Gulf Coastal Plain from the Florida panhandle to central Mexico, in the Mississippi Alluvial Plain to Minnesota, and in the southern Great Lakes from Minnesota to Ohio (Hobbs 1989, Pflieger 1996). Figure 26. Known localities for Orconectes (Procericambarus) rusticus (Rusty Crayfish), and Procambarus (Scapulicambarus) clarkii (Red Swamp Crayfish) introductions in the Western Region of North Carolina (Circles represent 2004–2005 sampling locations for stream-dwelling crayfish in the Broad and Little Tennessee River basins where these species were not encountered). 120 Southeastern Naturalist Vol. 9, Special Issue 3 In North Carolina, it is native to all river basins in the Coastal Plain and eastern Piedmont Plateau (Cooper 2005). In the Region, it is presently known from presumed introductions outside its native range in 3 river basins (Watauga, French Broad, and Broad), as well as from a small portion of its native range in the lower Catawba River basin. Within its native range in the Region, this species appears to be stable. Two records exist for White River Crawfish in the Watauga River basin from Price Lake, an artificial impoundment of Boone Fork, Watauga County: the first in November 2001 and the second during this inventory in June 2004. Thirteen sites were surveyed downstream from Price Lake in Boone Fork, the Watauga River, and other tributaries, as well as one tributary to Price Lake (NCDWQ, unpubl. data). No expansion of this population was documented. White River Crawfish were first collected from the Pigeon River, French Broad River basin, Haywood County in October 2001 (Fullerton 2002). In 2005, this species was collected from the vicinity of the initial occurrence locale and approximately 7.5 km downstream. None were collected in any other surveys in the Pigeon River system. In the North Pacolet River system, Broad River basin, Polk County, four juveniles tentatively assigned to this species were collected from Hughes Creek in April 2002 (J.E. Cooper, pers. comm.). In 2005, this locale and 3 Figure 27. Collection localities for Procambarus (Ortmannicus) acutus (White River Crayfish) in the Western Region of North Carolina. Occurrences in the Watauga, French Broad, and Broad River basins represent introduced populations (Small circles in these basins in the vicinity of known occurrences represent 2004–2005 sampling locations for stream-dwelling crayfish where this species was not encountered). 2010 J.W. Simmons and S.J. Fraley 121 adjacent streams were surveyed. No White River Crawfish were detected; however, the endemic C. spicatus was collected at 2 of these sites. Pflieger (1996) reported that most occurrences of this species in Missouri were in standing or slow-flowing waters, with none reported from the open channels of rivers. Because many streams in the Region where this species has been and is likely to be introduced are relatively high gradient and swift flowing, the invasive potential of this species may be limited. Native species adapted to these conditions may be resistant to displacement by White River Crawfish. Ecology and life-history observations. One form I male was collected in late June from Price Lake. Five form I males, 31–36 mm TCL, were collected during late July within its native range in 24 °C water. Two YOY, 8 mm TCL, were also collected at this site in late July. Within its native range in the Region, White River Crawfish were found sympatric with C. (P.) sp. C. It was collected with C. bartonii in the Pigeon River. Habitat observations. Within its native range, White River Crawfish were found in a slow-flowing stream approximately 2 m wide with substrate dominated by sand and gravel. Many individuals were observed moving about the sandy bottom, not immediately associated with cover. It occupies a small, cool-water impoundment (Price Lake) in the Watauga River system, and was collected from pools and eddies in the Pigeon River. Procambarus (Scapulicambarus) clarkii (Girard) (Red Swamp Crawfish; Fig. 26) Distribution and status. Red Swamp Crawfish are native to the Gulf Coastal Plain from western Florida to northern Mexico, in the Rio Grande River Basin to New Mexico, and north in the Mississippi Alluvial Plain to the Mississippi River and Ohio River confluence (Hobbs 1989, Pflieger 1996). Red Swamp Crawfish are widely utilized in aquaculture and have been introduced extensively outside its native range in the United States and elsewhere (Hobbs 1989). It has been introduced and is naturalized in all North Carolina river basins in the Piedmont Plateau and Coastal Plain, where it has rapidly increased in distribution and abundance (Cooper and Armstrong 2007, Cooper et al. 1998, Fullerton and Watson 2001). In the Region, this species was known from one locality: a pond in northwestern Rutherford County in the Cove Creek system, Broad River basin. Fullerton and Watson (2001) reported that it was introduced by the landowner circa 1993. After this study was completed, Red Swamp Crawfish were collected from multiple localities in Mountain Island Reservoir, an impoundment of the Catawba River in Gaston and Mecklenburg counties (G.E. Vaughn, Duke Energy, Inc., Charlotte, NC, pers. comm.). Most Red Swamp Crawfish introductions have had negative consequences for native species (Hobbs et al. 1989). Native endemic species C. lenati and C. spicatus, could be threatened if this species invades their range. In 2005, two sites were surveyed in the Cove Creek system within a 5-km radius of the Rutherford County Pond. No Red Swamp Crawfish were 122 Southeastern Naturalist Vol. 9, Special Issue 3 collected at these or other sites surveyed in the Region. In future efforts, the Cove Creek system should be a monitoring priority for potential spread of this invasive species. Contact with the landowner of the Rutherford County pond is recommended to determine the status of that population. Summary and Conclusions This inventory was successful in completing a majority of the distribution gaps for species occurring in western North Carolina. Prior to this effort, distribution data gaps, combined with lack of analysis of available historic data, hindered our knowledge of the overall status and distribution of crayfish in the Region. A compilation of historic records and new records obtained during this inventory into a GIS database provides a baseline for population trend assessments and direction for future monitoring efforts. This database allowed us to identify spatial and temporal gaps in distributional data for all species known from the Region. During this inventory, 28 species of crayfish were collected from 199 stream and 58 burrow localities, which provided additional information to help close these gaps, as well as relative abundance and demographic information for many species. Some observations of life-history traits, such as reproductive condition, fecundity, and habitat use, were also reported and represent a significant contribution to our knowledge of the biology of crayfishes in the Region. Although this inventory greatly advanced overall understanding of crayfish distributions in the Region, taxonomic clarity is needed for many species before a conservation status can be determined. When we began this inventory in 2004, 8 potentially undescribed species were known from the Region. Since then, 3 of these have been described. During this inventory, we consulted with crayfish taxonomists to determine current needs for specimens of undescribed taxa which we collected and provided to the best of our ability. Voucher specimens from a majority of our collections were deposited at the North Carolina State Museum of Natural Sciences and are available for future study. Taylor et al. (2007) listed endemism, introduction of non-indigenous species, and habitat loss, degredation, or alteration as major factors that could imperil crayfish populations; crayfish in the Region are affected by all of these. In the Region, 14 of the 31 species (26 described and 5 putative undescribed species), are endemic or “near endemic” and are priorities for monitoring due to their limited distribution (Table 1). Ten endemic or “near endemic” species occur in the Catawba, Broad, and Little Tennessee river basins where O. virilis (Catawba River basin), P. clarkii (Catawba and Broad River basins), and O. rusticus (Broad and Little Tennessee River basins) have been introduced. In addition, these species with narrow geographic ranges may be vulnerable to habitat alterations that are often synonymous with human population growth. North Carolina’s human population grew by approximately one million people between 2000 and 2007; 35% of this growth occurred within counties in the Region (USCB 2008). 2010 J.W. Simmons and S.J. Fraley 123 We identified 15 native species as priorities for future monitoring, 7 of which were identified as high priorities. Three non-native species, O. rusticus, O. virilis, and P. clarkii, were also identified as priorities for monitoring the expansion of established populations and status of native sympatric species. Native species designated as priorities for monitoring have a restricted distribution, face potential near-term threats of habitat loss, degredation, or alteration from anthropogenic activities, or recent data is not suitable to adequately assess their conservation status within the Region. Native species designated as high priorities for monitoring are very narrowly distributed, are known from only a small number of populations throughout their range, face potential near-term threats of habitat loss, degredation, or alteration from anthropogenic activities, or recent data indicates decline. Presently available data indicate that C. reburrus (French Broad Crayfish) and C. spicatus (Broad River Crayfish) have become undetectable at several known localities and may be in decline. Population trends for 14 species are presently undetermined, and 15 species are presumed stable based on presently available data. Future research needs should include obtaining material needed for taxonomic resolution of currently undescribed taxa, monitoring the spread of non-native species and the status of native sympatric species, monitoring species that have a small native range and are threatened by present or foreseeable habitat disturbance, monitoring species that may be declining based on data compiled during this effort, and obtaining more information on specific habitat requirements and the tolerance of individual species to physical and chemical changes to their habitats. Of particular importance in the Region are factors associated with global climate change and deposition of atmospheric pollutants that may affect rare and endemic species found at high elevations, and land-use changes occurring in rapidly developing areas. Priorities for crayfish monitoring will be re-evaluated on a 5-year basis and will be outlined in the periodically revised NCWRC Wildlife Action Plan. Data obtained during future monitoring will be added to the crayfish GIS and NCWRC databases and will continue to be provided to the NCWRC Non-game Wildlife Advisory Committee, the North Carolina Natural Heritage Program, and the US Fish and Wildlife Service to aid in updating the legal conservation status of rare crayfishes in western North Carolina. Acknowledgments This project was supported by a federal State Wildlife Grant, North Carolina Natural Heritage Trust Fund, and other state matching-fund sources. Collections from the Great Smoky Mountains National Park were made under Scientific Research and Collecting Permit number GRSM-2005-SCI-0018. Specimens from the Blue Ridge Parkway (BRP) were provided by E.M. Scott (Tennessee Valley Authority, permitted by the BRP). Other specimens were provided by M.C. Campen, J.A. Coombs, W. Taylor, and L. Williams. Additional species occurrence data were provided by J.E. Cooper, S.A. Cooper, B.H. Tracy, and R.F. Thoma. Special thanks to J.E. Cooper for his invaluable guidance and for verifying voucher specimens, and to our dedicated 124 Southeastern Naturalist Vol. 9, Special Issue 3 field technicians, J.A. Hartsell and D.L. Bell, without whom this work could not have been completed. We also appreciate the contributions of L.H. Lawson, who served briefly as a technician in the early phase of this project and whose work with S. Buback and the Southern Appalachian Biodiversity Project provided additional data. Special thanks to the citizens of western North Carolina who informed us of burrowing crayfish localities and who allowed us to collect on their property. We thank R.S. Butler, J.E. Cooper, R.B. Nichols, S.A. Welsh, C.J. Wood, and 2 anonymous reviewers for their editorial review and contributions to the quality of this manuscript. 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 Bouchard, R.W., and H.H. Hobbs, Jr. 1976. A new subgenus and two new species of crayfishes of the genus Cambarus (Decapoda: Cambaridae) from the southeastern United States. Smithsonian Contributions to Zoology 224:1–15. Butler, R.S. 2002. Crayfishes of the southern Appalachian ecosystem, with emphasis on the imperiled fauna. US Fish and Wildlife Service, Asheville, NC. Clamp, J.C. (Compiler). 1999. A report on the conservation status of North Carolina’s freshwater and terrestrial crustacean fauna. North Carolina Wildlife Resources Commission, Raleigh, NC. 100 pp. Cooper, J.E. 2000. A new species of crayfish of the genus Cambarus, subgenus Cambarus (Decapoda: Cambaridae), from the Broad River basin of North Carolina. Journal of the Elisha Mitchell Scientific Society 116(1):1–12. Cooper, J.E. 2001. Cambarus (Puncticambarus) hobbsorum, a new crayfish (Decapoda: Cambaridae) from North Carolina. Proceedings of the Biological Society of Washington 114(1):152–161. Cooper, J.E. 2002. North Carolina crayfishes (Decapoda: Cambaridae): Notes on distribution, taxonomy, life history, and habitat. Journal of the North Carolina Academy of Science 118(3):167–180. Cooper, J.E. 2004a. Guide to crayfishes of the Hiwassee River basin, North Carolina. North Carolina State Museum of Natural Sciences, Raleigh, NC. Cooper, J.E. 2004b. Guide to crayfishes of the Little Tennessee River basin, North Carolina. North Carolina State Museum of Natural Sciences, Raleigh, NC. Cooper, J.E. 2005. Crayfishes occurring in North Carolina. North Carolina State Museum of Natural Sciences, Raleigh, NC. Cooper, J.E. 2006a. A new species of crayfish of the genus Cambarus Erichson, 1846 (Decapoda: Cambaridae) from the eastern Blue Ridge foothills and western Piedmont Plateau of North Carolina. Proceedings of the Biological Society of Washington 119(1):67–80. Cooper, J.E. 2006b. A new crayfish of the genus Cambarus Erichson, 1846, subgenus Puncticambarus Hobbs, 1969 (Decapoda: Cambaridae), from the Hiwassee River basin of North Carolina. Proceedings of the Biological Society of Washington 119(1):81–90. Cooper, J.E., and S.A. Armstrong. 2007. Locality records and other data for invasive crayfishes (Decapoda: Cambaridae) in North Carolina. Journal of the North Carolina Academy of Science 123(1):1–13. Cooper, J.E., and A.L. Braswell. 1995. Observations on North Carolina crayfishes (Decapoda: Cambaridae). Brimleyana 22:87–132. 2010 J.W. Simmons and S.J. Fraley 125 Cooper, J.E., and K.A. Schofield. 2002. Cambarus (Jugicambarus) tuckasegee, a new species of crayfish (Decapoda: Cambaridae) from the Little Tennessee River basin, North Carolina. Proceedings of the Biological Society of Washington 115(2):371–381. Cooper, J.E., A.L. Braswell, and C. McGrath. 1998. Noteworthy distributional records for crayfishes (Decapoda: Cambaridae) in North Carolina. Journal of the Elisha Mitchell Scientific Society 114(1):1–10. Dewees, J.P. 1972. Geographic variation in the primary burrowing crayfish, Cambarus dubius Faxon and Cambarus carolinus (Erichson) (Decapoda: Astacidae) in Tennessee with notes on ecology and life history. M.Sc. Thesis. University of Tennessee, Knoxville, TN. 75 pp. Eversole, A.G. 1995. Distribution of three rare crayfish species in South Carolina, USA. Freshwater Crayfish 8:113–120. Fullerton, A.H. 2002. Status inventory of uncommon crayfishes in North Carolina, Final Report 2001: Savannah, French Broad, and Lumber (Waccamaw) river basins. North Carolina Wildlife Resources Commission, Nongame Project Report, Raleigh, NC. Fullerton, A.H., and B.T. Watson. 2001. New distributional records for two nonindigenous and one native crayfish in North Carolina. Journal of the Elisha Mitchell Scientific Society 117(1):66–70. Hobbs, H.H., Jr. 1956. A new crayfish of the Extraneus Section of the genus Cambarus with a key to the species of the Section (Decapoda, Astacidae). Proceedings of the Biological Society of Washington 69:115–122. Hobbs, H.H., Jr. 1972. Crayfishes (Astacidae) of North and Middle America. US Environmental Protection Agency, Washington, DC. 173 pp. Hobbs, H.H., Jr. 1981. The crayfishes of Georgia. Smithsonian Contributions to Zoology 318:1–549. Hobbs, H.H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, Parastacidae). Smithsonian Contributions to Zoology 480:1–236. Hobbs, H.H., Jr., and D.J. Peters. 1977. The entocytherid ostracods of North Carolina. Smithsonian Contributions to Zoology 247:1–73. Hobbs, H.H., III. 1993. Trophic relationships of North American freshwater crayfishes and shrimps. Milwaukee Public Museum Contributions in Biology and Geology 85:1–110. Hobbs, H.H., III, and J.P. Jass. 1988. The crayfishes and shrimp of Wisconsin. Milwaukee Public Museum, Special Publications in Biology and Geology No. 5, Milwaukee, WI. 177 pp. Hobbs, H.H., III, J.P. Jass, and J.V. Huner. 1989. A review of global crayfish introductions with particular emphasis on two North American species (Decapoda, Cambaridae). Crustaceana 56(3):299–316. LeGrand, H.E., S.E. McRae, S.P. Hall, and J.T. Finnegan (Compilers). 2008. Natural Heritage Program list of the rare animal species of North Carolina. North Carolina Natural Heritage Program, NC Department of Environment and Natural Resources, Raleigh, NC. 119 pp. Lodge, D.M., C.A. Taylor, D.M. Holdich, and J. Skurdal. 2000. Nonindigenous crayfishes threaten North American freshwater biodiversity: Lessons from Europe. Fisheries 25(8):7–20. 126 Southeastern Naturalist Vol. 9, Special Issue 3 Marsh, S.S. 1998. Linville River sub-basin aquatic inventory. North Carolina Wildlife Resources Commission, Nongame Project Report, Raleigh, NC. McGrath, C. 1993. Terrestrial snails, crayfish, and aquatic snails collected in the Nolichucky River basin in North Carolina. North Carolina Wildlife Resources Commission, Nongame Project Report, Raleigh, NC. McGrath, C. 1997. Monitoring and surveying for Little-wing Pearlymussel, Appalachian Elktoe, Parrish Crayfish and Hiwassee Crayfish. North Carolina Wildlife Resources Commission, Annual Performance Report, Nongame and Endangered Wildlife Program, Raleigh, NC. McGrath, C. 1998. New River basin aquatic inventory. North Carolina Wildlife Resources Commission, Nongame Project Report, Raleigh, NC. McLarney, W.O. 1993. Status of the crayfish Cambarus georgiae in the upper Little Tennessee watershed. Final report to North Carolina Wildlife Resources Commission, Raleigh, NC. North Carolina Wildlife Resources Commission (NCWRC). 2005a. North Carolina Wildlife Action Plan. Raleigh, NC. NCWRC. 2005b. 2005–2006 North Carolina inland fishing, hunting, and trapping regulations digest. Raleigh, NC. Pflieger, W.L. 1996. The crayfishes of Missouri. Missouri Department of Conservation, Jefferson City, MO. 152 pp. Taylor, C.A. 2000. Systematic studies of the Orconectes juvenilis complex (Decapoda: Cambaridae), with descriptions of two new species. Journal of Crustacean Biology 20(1):132–152. Taylor, C.A., G.A. Schuster, J.E. Cooper, R.J. DiStefano, A.G. Eversole, P. Hamr, H.H. Hobbs III, H.W. Robison, 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(8):372–389. Thoma, R.F. 2005. Cambarus (Cambarus) eeseeohensis (Decapoda: Cambaridae), a new species of crayfish from the Linville River of North Carolina and its bearing on understanding the evolution of the subgenus Cambarus. Proceedings of the Biological Society of Washington 118(4):794–802. US Census Bureau (USCB). 2008. Annual estimates of the population for counties of North Carolina: April 1, 2000 to July 1, 2007. Available online at http://quickfacts. census.gov/qfd/states/37000lk.html. Accessed April 2009. Williams, A.B., L.G. Abele, D.L. Felder, H.H. Hobbs, Jr., R.B. Manning, P.A. McLaughlin, and I. Perez Farfante. 1989. Common and scientific names of aquatic invertebrates from the United States: Decapod Crustaceans. American Fisheries Society, Special Publication 17, Bethesda, MD. 77 pp. Williams, C.E., R.D. Bivens, and B.D. Carter. 2007. Annotated list of the crayfishes of Tennessee. Tennessee Wildlife Resources Agency, Morristown, TN.