Conservation, Biology, and Natural History of Crayfishes from the Southern US 2010 Southeastern Naturalist 9(Special Issue 3):155–164 The Extirpation of Orconectes limosus (Spinycheek Crayfish) Populations in West Virginia Casey D. Swecker1,2,*, Thomas G. Jones1, Keith Donahue II1, Derek Mckinney1, and Geoffrey D. Smith1 Abstract - Introductions of nonnative crayfish species have resulted in the global decline of native crayfish populations, including those in North America. Historically, the North American range of Orconectes limosus (Spinycheek Crayfish) extended from Maine, southward into northern Virginia, including West Virginia’s eastern panhandle. A 1988–1989 survey of the eastern panhandle of West Virginia resulted in the capture of only 14 O. limosus, but an abundance of the nonnative Orconectes virilis (Virile Crayfish). These data along with additional unpublished accounts of declines of O. limosus populations prompted our survey of eastern West Virginia. In 2005 and 2006, crayfishes were collected from streams within the West Virginia range of the O. limosus, including historic capture locations. Our collection of 600 individuals comprised crayfishes from 3 species: O. virilis, Orconectes obscurus (Allegheny Crayfish), and Cambarus bartonii bartonii (Common Crayfish). The nonnative O. virilis was present at 26 of the 30 sites, whereas O. limosus was absent from all collections. Our results may indicate extirpation of some or all populations of O. limosus in eastern West Virginia, but absence data may also reflect a low detection probability of individuals from small populations. Competition between the nonnative O. virilis and O. limosus have been reported elsewhere, and likely explains the extirpation of populations of O. limosus in West Virginia. Introduction The decline of Orconectes limosus (Rafinesque) (Spinycheek Crayfish) in West Virginia has concerned researchers, conservationists, and agencies over the past decade. The historical range of O. limosus extended from Maine southward into northern Virginia, including Morgan, Berkeley, and Jefferson counties of eastern West Virginia (Fig. 1; Hobbs 1989). Orconectes limosus was first recorded from West Virginia in Cherry Run, a tributary of the Potomac River in Morgan County (Ortmann 1906). Jezerinac et al. (1995) examined and reported a record of O. limosus from Back Creek collected in 1975 by G.E. Lewis, West Virginia Division of Natural Resources. Also, Jezerinac et al. (1995) collected O. limosus in low numbers at 3 sites from Opequon Creek and 1 site from Back Creek during surveys of 1988 and 1989 (Fig. 2). During the 1988–1989 surveys, Jezerinac et al. (1995) also collected the nonnative Orconectes virilis Hagen (Virile Crayfish) from Opequon and Back creeks. Orconectes virilis was first reported in West Virginia in 1Department of Integrated Science and Technology, Marshall University, Huntington, WV 25755. 2Current address - Environmental Solutions and Innovations, 781 Neeb Road, Cincinnati, OH 45233. *Corresponding author - cswecker@environmentalsi.com. 156 Southeastern Naturalist Vol. 9, Special Issue 3 1970 in the New River Gorge National River, Summers County (Edmundson 1974). At that time, the species was considered to be an introduction (Jezerinac et al. 1995) because the range of O. virilis extended from Saskatchewan to Ontario, Canada, and from Montana and Utah to Arkansas, New York, and Maine (Hobbs 1989). As of 1990, the documented range of O. virilis in West Virginia was limited to Berkeley and Jefferson counties in eastern West Virginia and the New (Kanawha) River drainage in Monroe, Summers, Fayette, and Kanawha counties in southern West Virginia (Jezerinac et al. 1995). Figure 1. North American distributions of Orconectes limosus (Spinycheek Crayfish) and Orconectes obscurus (Allegheny Crayfish) from Hobbs (1989). 2010 C.D. Swecker, T.G. Jones, K. Donahue II, D. Mckinney, and G.D. Smith 157 The displacement of native crayfishes by nonnative crayfishes has been widely studied (Capelli 1982, Capelli and Munjal 1982, Holdich 1988, Kilian et al. 2009, Lodge et al. 2000, Schwartz et al. 1963, Taylor et al. 1996). Meredith and Schwartz (1960) documented the presence of the nonnative O. virilis at 5 sites in the Patapsco River near Woodstock, MD. By 1963, this population had expanded to occupy the entire Patapsco River drainage, Figure 2. The 2005–2006 study locations and historic collection locations of Orconectes limosus (Spinycheek Crayfish) from West Virginia by Ortmann (1906), Jezerinac et al. (1995), and G. Lewis (West Virginia Division of Natural Resources, Elkins, WV unpubl. data). 158 Southeastern Naturalist Vol. 9, Special Issue 3 displacing 2 native species: Cambarus bartonii bartonii (Fabricius) (Common Crayfish) and O. limosus (Schwartz et al. 1963). The further expansion of O. virilis into other Maryland watersheds has been followed by a concurrent decline in native crayfishes (Kilian et al. 2010). Orconectes virilis have been documented in California (Eng and Daniels 1982), Canada (McAlpin et al. 2007), Mexico (Hamr 2002), and recently in the United Kingdom (Ahern et al. 2008). We speculate that O. limosus has been displaced by O. virilis in West Virginia. Also O. virilis may impact populations of C. b. bartonii and O. obscurus (Hagen) (Allegheny Crayfish) in this region. Orconectes virilis is expected to homogenize the species composition of the crayfish community in eastern West Virginia. Similar shifts were supported by data from Schwartz et al. (1963) and Kilian et al. (2010). Our objectives were to determine the current status of O. limosus in West Virginia, document the range expansion of nonnative O. virilis, and document shifts in species composition of crayfish communities throughout eastern West Virginia. Methods We focused our survey on Back Creek and Opequon Creek, two streams with historic records of O. limosus (Jezerinac et al. 1995); however, study sites from 18 streams were sampled. We used 3 sampling strategies to document composition of crayfish communities in tributaries of the Potomac River in eastern West Virginia. Our first strategy involved sampling random and historic stream sites by hand collection (wading and snorkel assisted) and seining. In this initial effort, crayfishes were collected at 27 sites from 18 streams (Fig. 2). Secondly, we used a canoe to sample areas of Back Creek with limited public access during 2 float trips. We wanted to avoid public access areas, and sample areas where bait-bucket introductions would be less likely. Float trip 1 was approximately 8.7 km from the bridge on State Highway 45 east of Glengary to the bridge on County Highway 18 east of Shanghai. Float trip 2 was approximately 8 km from the bridge on County Highway 6 to the bridge on State Highway 9 just west of Hedgesville (Fig. 2). Crayfishes were sampled by hand, seine, and dip net at multiple intervals within all available habitat types during the float trips. Snorkeling and SCUBA also assisted in the collections of deep pools. For the third strategy, we electrofished and seined 3 sites on Back Creek. At each site, three 10-m reaches represented 3 habitat types: riffle, run, and glide. Pools were not sampled due to electrofishing limitations in deep water. At the downstream end of each reach, a seine (1.2 m x 6 m) was staked to the river bottom perpendicular to stream flow. Two methods were used to collect crayfish at each site. First, a kick method disturbed substrate and moved rocks from the upstream end of the reach to the seine at the downstream end. Second, a backpack electrofishing unit was used to sample each reach. All specimens were wet-weighed using a digital balance (Ohaus Portable Scout Pro Model-SP602) to the nearest 0.01 gram, and the total carapace 2010 C.D. Swecker, T.G. Jones, K. Donahue II, D. Mckinney, and G.D. Smith 159 length was recorded using digital calipers (Mitutoyo Absolute Digimatic Model-500-172-20) to the nearest 0.01 mm for each crayfish collected. Results We collected a total of 600 crayfish during this study representing 3 species: O. virilis, O. obscurus, and C. b. bartonii. No individuals of O. limosus were collected during our study. Orconectes virilis represented 71.5% (n = 429) of our total sample, whereas O. obscurus (19.3%, n = 116) and C. b. bartonii (9.2%, n = 55) were collected less often. For the first sampling strategy, 358 crayfishes were collected from 27 sites in 18 streams within Morgan, Jefferson, and Berkeley counties of eastern West Virginia. Orconectes virilis comprised most of the total sample (83%, 297 individuals; Table 1). Sample sizes were small for C. b. bartonii Table 1. Study site and abundance data from crayfish surveys in eastern West Virginia. UTM UTM O. O. C. b. Site Stream Zone East North virilis obscurus bartonii 1 Tributary to Back Creek 18 241248 4386910 20 0 1 2 Tomahawk Run 17 754797 4378544 0 0 6 3 Middle Fork Sleepy Creek 17 738459 4375853 5 0 0 4 Sleepy Creek 17 741180 4384676 2 0 0 5 Opequon Creek 18 248816 4371911 36 0 0 6 Rockmarsh Run 18 254846 4369470 0 0 1 7 Rattlesnake Run 18 261144 4364892 6 0 1 8 Elk Run 18 258732 4357695 0 0 6 9 Unnamed Stream 18 258781 4355312 2 0 0 10 Shenandoah River 18 257042 4348164 2 1 0 11 Evitts Run 18 254903 4348502 0 0 6 12 Bullskin Run 18 253388 4344630 3 0 6 13 Opequon Creek 18 242357 4356541 4 0 0 14 Middle Creek 18 238837 4362586 5 0 1 15 Opequon Creek 18 246795 4363032 12 0 0 16 Opequon Creek 18 247031 4367817 30 0 0 17 Harlan Run 18 246358 4385936 3 0 2 18 Sleepy Creek 17 749363 4394970 7 4 1 19 Furnace Run 17 774816 4346411 18 0 2 20 Tuscarora Creek 18 241142 4372568 8 0 2 21 Back Creek 18 234485 4373183 6 0 0 22 Harlan Run 18 246130 4382978 11 0 3 23 Back Creek 17 757433 4386523 26 2 0 24 Back Creek 17 748099 4369121 29 0 1 25 Opequon Creek 18 241551 4355498 28 0 2 26 Opequon Creek 18 244407 4360269 13 0 1 27 Cherry Run 17 751866 4387383 21 0 12 D1 Back Creek 17 756225 4382741 4 5 0 D2 Back Creek 17 754786 4377906 28 5 1 D3 Back Creek 17 754024 4377270 41 58 0 F1 Back Creek 17 745846 4363295 27 31 0 F2 Back Creek 17 754940 4378168 32 10 0 Total 429 116 55 160 Southeastern Naturalist Vol. 9, Special Issue 3 (15%, n = 54) and O. obscurus (2%, n = 7). Headwater streams (less than 1.2 m wide) typical of C. b. bartonii habitat were included in the assessment, given the possibility for smaller streams to serve as refugia for O. limosus. Eight of these smaller high-gradient streams contained C. b. bartonii and O. virilis, while O. obscurus was recorded from only 3 streams (Back Creek, Sleepy Creek, and Shenandoah River; Table 1). During the two canoe trips, we collected a total of 100 crayfish: 58 individuals during float trip 1, and 42 during float trip 2 (Table 1). Sampling during the first float trip produced 27 O. virilis and 31 O. obscurus. During the second float trip, we collected 32 O. virilis and 10 O. obscurus. After pooling samples from the 2 float trips, O. virilis was the dominant species (59% of total sample) followed by O. obscurus (41%). Kick seining and electrofishing at 3 sites on Back Creek produced 142 crayfishes. The first site yielded a low sample size (4 O. virilis and 5 O. obscurus), whereas higher numbers were collected at the second site (28 O. virilis, 5 O. obscurus, and 1 C. b. bartonii) and the third site (41 O. virilis and 58 O. obscurus). A pooled sample of the 3 sites yielded 73 O. virilis, 68 O. obscurus, and 1 C. b. bartonii. The total weight of O. virilis was 519.7 g from a sample of 73 individuals (54 form II males and 19 females), where 68 individuals of O. obscurus weighed 146.4 g (1 form I male, 38 form II males, and 29 females). Discussion Several studies have reported declines or extirpation of populations of O. limosus in West Virginia, Maryland, Pennsylvania, and New York following introduction of O. virilis (Bouchard et al. 2007; Crocker 1957, 1979; Jezerinac et al. 1995; Kilian et al. 2010; Schwartz et al.1963). Our findings in eastern West Virginia are consistent with these previous studies, and document an increase in locality records of O. virilis and no recent locality records for O. limosus. Possibly, O. virilis may also negatively impact other crayfish species in eastern West Virginia, such as O. obscurus and C. b. bartonii. Our sample numbers for O. obscurus and C. b. bartonii were low relative to that of O. virilis. Based on previous studies in the New River, WV, O. virilis was believed to inhabit larger streams and be absent from small streams, such as those used by C. b. bartonii in our study area of eastern West Virginia (Jezerinac 1995). During another study, however, O. virilis were collected in small high-gradient tributaries of the New River within the New River Gorge National River, WV (C. Swecker, unpubl. data). Similarly, we found O. virilis in small streams of eastern West Virginia, and laboratory studies are needed to experimentally examine competitive interactions between O. virilis and C. b. bartonii. Nonnative crayfish species may negatively impact native crayfish species through competitive effects linked to body and chela size, or from 2010 C.D. Swecker, T.G. Jones, K. Donahue II, D. Mckinney, and G.D. Smith 161 hybridization. Orconectes virilis attains a larger body and chela size compared to O. limosus and O. obscurus, and larger crayfish often have a competitive advantage over smaller individuals (Bywater et al. 2008, Garvey and Stein 1993,). Also, because larger crayfish species are less vulnerable to fish predation (DiDonato and Lodge 1993, Keller and Moore 2000), the nonnative O. virilis may have an additional advantage over smaller crayfish species. We have not documented hybridization of O. virilis with another crayfish species in the upper Potomac River drainage, but we did observe an O. virilis amplexing an O. obscurus. We identified hybrids between O. virilis and O. obscurus from a stream outside of our study area in the upper Potomac River drainage, MD (C.D. Swecker, unpubl. data). It is possible that causes other than the nonnative O. virilis are responsible for the reduction or extirpation of populations of O. limosus in eastern West Virginia. Interestingly, O. limosus is tolerant of habitat degradation, siltation, low oxygen, and pollutants (Lindqvist and Huner 1999). Orconectes limosus is among the most widely distributed nonnative crayfish species in Europe (Bearn and Petruskey 2006, Souty-Grosset et al. 2006). Since its introduction to Europe in 1890, it has spread to many countries because of its high fecundity and fast development (Kozak et al. 2006). Orconectes limosus use detritus, sediment-laden substrate, clay burrows, soft benthic habitat, and silty or turbid waters (Hamr 2002, Jezerinac 1995). In Europe, O. limosus were reported from calm, deep waters such as ponds and lakes, which may be organically rich and polluted. We sampled areas of high sedimentation and leaf packs, but only O. virilis were present in these habitats. Areas with clay substrates were also sampled; one female O. virilis was excavated at 0.5 m depth from a burrow in a clay bank. Orconectes limosus is tolerant to habitat degradation and pollution (Lindqvist and Huner 1999) and has persisted through eradication efforts after introduction in several countries (Holdich and Black 2007), suggesting that habitat degradation and pollution may not explain the absence of O. limosus in West Virginia. In addition to the nonnative O. virilis, it is possible that O. obscurus and O. limosus are not native to the upper Potomac River (Bouchard et al. 2007, Ortmann 1906). Ortmann (1906) suggested that O. limosus accessed the Potomac highlands via the Chesapeake and Ohio Canal. This species may have dispersed (on its own) west of the Piedmont through the low-gradient, silt-laden habitats within the canal system. These habitats are of the same type as those associated with the invasion of O. limosus in Europe (Hamr 2002). Ortmann (1906) also suggested that the Allegheny crayfish was not native to the Potomac River and was likely introduced by anglers in Wills Creek near Cumberland, MD. Astacologists and zoogeographers may never reach a consensus on the native distribution range of O. limosus; however, a determination of nonnative status for O. limosus would remove conservation concerns for this species in eastern West Virginia. 162 Southeastern Naturalist Vol. 9, Special Issue 3 An alternative explanation for reduction and extirpation of populations of O. limosus can be hypothesized if one considers O. obscurus as nonnative. If O. obscurus was introduced to the upper Potomac River drainage before O. virilis, then O. obscurus may have reduced or extirpated populations of O. limosus. Possibly O. obscurus reduced population sizes of O. limosus, and the subsequent introduction of O. virilis yielded further impact on O. limosus population. This hypothesis was also supported by Bouchard et al. (2007). Although typically not as large or as aggressive as O. virilis, O. obscurus may have competed with and reduced populations sizes of O. limosus until the introduction of O. virilis overwhelmed them. It is uncertain if this scenario is accurate, and more research is needed to confirm the interaction of O. obscurus and O. limosus from West Virginia. This possibility does not downplay the threat that O. virilis is having on native West Virginia crayfishes; however, it adds an additional topic that is often overlooked. When species are transferred from one watershed to another, the negative impact potential is still great. Our study supports the reduction or extirpation of populations of O. limosus in West Virginia. Other studies have also speculated or documented declines or extirpation of populations of O. limosus following introduction of O. virilis (Bouchard et al. 2007; Crocker 1957, 1979; Jezerinac et al. 1995; Kilian et al. 2010; Schwartz et al.1963). We did not sample many deep-water areas; hence, further surveys should focus on deep pools of the Potomac River, Back Creek, or Opequon Creek. Our study did not document direct evidence for competitive exclusion, but we speculate that competition with the nonnative O. virilis is a likely cause of the decline or extirpation of O. limosus in West Virginia. Acknowledgments This study was funded and conducted in agreement with the West Virginia Department of Natural Resources 2005 DNR Non Game Grant # 60218 and Marshall University Research Corporation # 204167. Special thanks to Dr. Mike Little and the Department of Integrated Science and Technology at Marshall University for use of GIS software and laboratory access. Stuart Welsh provided publication support. 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 Ahern, D., J. England, and A. Ellis. 2008. The Virile Crayfish, Orconectes virilis (Hagen, 1870) (Crustacea: Decapoda: Cambaridae), identified in the UK. Aquatic Invasions 3:102–104. Bouchard, R.W., D.A. Lieb, R.F. Carline, T.R. Nuttal, C.B. Wengert, and J.R. Wallace. 2007. 101 years of change (1906–2007). The distribution of the crayfishes of Pennsylvania. Part I: Eastern Pennsylvania. Academy of Natural Sciences of Philadelphia. Report Number 07–11, Philadelphia, PA. 39 pp. 2010 C.D. Swecker, T.G. Jones, K. Donahue II, D. Mckinney, and G.D. Smith 163 Capelli, G.M. 1982. Displacement of northern Wisconsin crayfish by Orconectes rusticus (Girard). Limnology and Oceanography 27:741–745. Capelli, G.M., and B.L. Munjal. 1982. Aggressive interactions and resource competition in relation to species displacement among crayfish of the genus Orconectes. Journal of Crustacean Biology 2:486–492. Crocker, D.W. 1957. The crayfishes of New York State (Decapoda, Astacidae). New York State Museum Bulletin 355:1–97. Crocker, D.W. 1979. The crayfishes of New England. Proceedings of the Biological Society of Washington 92:255–252. DiDonato, G.T., and D.M. Lodge. 1993. Species replacements among Orconectes crayfishes in northern Wisconsin lakes: The role of predation by fish. Canadian Journal of Fisheries and Aquatic Sciences 50:1484–1488. Edmundson, J.P., Jr. 1974. Food habits, age and growth of Flathead Catfish Pylodictis olivaris in Bluestone Reservoir, West Virginia. M.Sc. Thesis. West Virginia University, Morgantown, WV. 87 pp. Eng, L.L., and R.A. Daniels. 1982. Life history, distribution, and status of Pacifastacus fortis (Decapoda: Astacidae). California Fish and Game 68:197–212. Hamr, P. 2002. Orconectes. Pp. 585–608, In D.M. Holdich (Ed.). Biology of Freshwater Crayfish. Blackwell Science, Oxford, UK. 702 pp. Hobbs, H.H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology 480. Smithsonian Institute Press, Washington, DC. Holdich, D.M. 1988. The dangers of introducing alien animals, with particular reference to crayfish. Freshwater Crayfish 7:15–30. Holdich, D.M., and J. Black. 2007. The Spiny-cheek Crayfish, Orconectes limosus (Rafinesque, 1817) [Crustacea: Decapoda: Cambaridae], digs into the UK. Aquatic Invasions 2:1–16. Jezerinac, R.F., G.W. Stocker, and D.C. Tarter. 1995. The crayfishes (Decapoda:Cambaridae) of West Virginia. Bulletin of the Ohio Biological Survey, New Series 10(1):1–193. Keller, T.A., and P.A. Moore. 2000. Context-specific behavior: Crayfish size influences crayfish-fish interactions. Journal of the North American Benthological Society19:344–351. Kilian, J.V., A.J. Becker, S.A. Stranko, M. Ashton, R.J. Klauda, J. Gerber, and M. Hurd. 2010. The status and distribution of Maryland crayfishes. Southeastern Naturalist 9(Special Issue 3):11–32. Kilian, J.V., J. Frentress, R.J. Klauda, A.J. Becker, and S.A. Stranko. 2009. The invasion of Procambarus clarkii (Decapoda: Cambaridae) into Maryland streams following its introduction in outdoor aquaculture ponds. Northeastern Naturalist 16(4):655–633. Kozak P., M. Buric, and Z. Duris. 2006. The fecundity, time of egg development and juvenile production in Spiny-cheek (Orconectes limosus) under controlled conditions. Bulletin Français de la Peche et de la Pisciculture 380–381:1171–1182. Lodge, D.M., C.A. Taylor, D.M. Holdich, and J. Skudal. 2000. Nonindigenous crayfishes threaten North American freshwater biodiversity: Lessons from Europe. Fisheries 25(8):7–20. Lindqvist, O.V., and J.V. Huner. 1999. Life-history characteristics of crayfish: What makes some of them good colonizers? Pp. 23–30, In F. Gherardi and D.M. Holdich (Eds.). Crayfish in Europe as Alien Species. How to Make the Best of a Bad Situation? A. A. Balkema, Rotterdam, The Netherlands. 733 pp. 164 Southeastern Naturalist Vol. 9, Special Issue 3 McAlpin, D.F., A.H.E. McAlpin, and A. Madden. 2007. Occurrence of the potentially invasive crayfish, Orconectes virilis (Decapoda, Cambaridae) in eastern New Brunswick, Canada. Crustaceana 80:509–511. Meredith, W.G., and F.J. Schwartz. 1960. Maryland Crayfishes. Maryland Department of Resources and Education, Education Series 60, Solomons, MD. Ortmann, A.E. 1906. The crawfishes of the state of Pennsylvania. Memoirs of the Carnegie Museum 2:343–533. Petrusek, A., L. Filipová, Z. Duriš, I. Horká, P. Kozák, T. Policar, M. Stambergová, and Z. Kučera. 2006. Distribution of the invasive Spiny Cheek Crayfish (Orconectes limosus) in the Czech Republic. Past and present. Bulletin Francaisde la Peche et de la Pisciculture 380–381:903–917. Schwartz. F.J., R. Rubelmann, and J. Allison. 1963. Ecological population expansion of the introduced crayfish, Orconectes virilis. Ohio Journal of Science 63:266–273. Souty-Grosset C., D.M. Holdich, P.Y. Noel, J.D. Reynolds, and P. Haffner (Eds.). 2006. Atlas of Crayfish in Europe. Patrimoines Naturels 64. Museum National d’Histoire Naturelle, Paris, France. Taylor, C.A., M.L., Warren, Jr., J.F. Fitzpatrick, Jr., H.H. Hobbs III, R.F. Jezerinac, W.L. Pflieger, and H.W. Robison. 1996. Conservation status of crayfishes of the United States and Canada. Fisheries 21:25–38.