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Rediscovery of Cambarus diogenes (Devil Crayfish) in Pennsylvania
Mael G. Glon, Andrew J. Mularo, David A. Lieb, and Zachary J. Loughman

Northeastern Naturalist, Volume 25, Issue 3 (2018): 355–361

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Northeastern Naturalist Vol. 25, No. 3 M.G. Glon, A.J. Mularo, D.A. Lieb, and Z.J. Loughman 2018 355 2018 NORTHEASTERN NATURALIST 25(3):355–361 Rediscovery of Cambarus diogenes (Devil Crayfish) in Pennsylvania Mael G. Glon1,*, Andrew J. Mularo1, David A. Lieb2, and Zachary J. Loughman3 Abstract - North American crayfish biodiversity is in need of conservation attention, which requires monitoring to assess the status of potentially imperiled species. Pennsylvania has a long history of crayfish studies, dating back to the early 1900s and continuing today. We sampled 37 new and historical sites in southeastern Pennsylvania in an effort to locate remnant populations of Cambarus diogenes (Devil Crayfish), an ecologically important burrowing crayfish. While we did not find Devil Crayfish at any historical sites, we discovered a single population in Neshaminy State Park, indicating a decline in this species’ range in Pennsylvania since the last survey, circa 1905. We attribute the statewide decline of Devil Crayfish to an inherently limited amount of suitable habitat, urbanization, and biological invasions, and recommend that steps be taken to protect this sp ecies. Introduction North American freshwater biodiversity has decreased at an alarming rate since the beginning of the 20th century (Burkhead 2012, Ricciardi and Rasmussen 1999). Among North American freshwater organisms facing the greatest threats are crayfishes, of which 32% (127 of 391) are considered endangered or vulnerable by the American Fisheries Society Endangered Species Crayfish Subcommittee (C.A. Taylor, Illinois Natural History Survey, Champaign, IL, pers. comm.; Taylor et al. 2007). Recent research has elucidated major threats to freshwater crayfish biodiversity, including but not limited to climate change, habitat loss/alteration, and biological invasions (Bland 2017, Richman et al. 2015). However, very few North American crayfish species have been the focus of the types of detailed studies necessary to inform conservation assessments, leading to an underrepresentation of crayfishes on the federal Endangered Species list (Moore et al. 2013, Richman et al. 2015). Pennsylvania has a long history of crayfish research, starting with the work of naturalist and Carnegie Museum of Natural History Curator Arnold E. Ortmann in the early 1900s (Ortmann 1905, 1906). More recently, substantial resources have been devoted to crayfish surveys aimed at quantifying trends in the ranges of Pennsylvania crayfishes over the last century (Lieb et al. 2008, 2011a, 2011b; Loughman et al. 2017, 2018). These surveys have led to recommendations for management 1The Ohio State University, Department of Evolution, Ecology, and Organismal Biology, 318 West 12th Avenue, Columbus, OH 43210. 2Western Pennsylvania Conservancy and Pennsylvania Fish and Boat Commission, 595 East Rolling Ridge Drive, Bellefonte, PA 16823. 3West Liberty University, Department of Natural Sciences and Mathematics, PO Box 295, West Liberty, WV 26074. *Corresponding author - glon.1@osu.edu. Manuscript Editor: Stuart Welsh Northeastern Naturalist 356 M.G. Glon, A.J. Mularo, D.A. Lieb, and Z.J. Loughman 2018 Vol. 25, No. 3 actions, as well as 2 new state species records (Cambarus acuminatus Faxon [Acuminate Crayfish] [Lieb et al. 2008]; Creaserinus fodiens [Cottle] [Digger Crayfish] [Loughman et al. 2018]), highlighting the need for continued biodiversity surveys even in populated areas. However, except for Loughman et al. (2017, 2018), recent crayfish research in Pennsylvania has focused on stream-dwelling species, whereas the status of burrowing crayfishes remains lar gely unknown. Cambarus diogenes Girard (Devil Crayfish), a primary burrower (Hobbs 1969), is one of the rarest crayfish species in Pennsylvania. The Devil Crayfish, as currently described, is the most widespread North American crayfish, found in 32 US states and parts of Ontario, Canada (Taylor et al. 2007). However, this taxon has long been thought to be a species complex composed of multiple undescribed species (e.g., Faxon 1885, Hobbs 1989, Jezerinac 1993), and M. Glon has undertaken a systematic revision of this species complex. Although the Devil Crayfish’s range is extensive between the Appalachian and Rocky Mountains of North America, its distribution east of the Appalachian Mountains is limited to the Atlantic Coastal Plain physiographic region (Jezerinac 1993). As such, any populations of Devil Crayfish in southeastern Pennsylvania may represent the northern-most extent of this species’ east-coast range and therefore harbor important genetic variation (Hardie and Hutchings 2010). In his 2 seminal publications on the crayfish diversity of Pennsylvania, Ortmann (1905, 1906) documented numerous populations of Devil Crayfish from eastern and western Pennsylvania but emphasized morphological variation between these disparate populations. Jezerinac (1993), subsequently investigated this morphological variation and described all populations of Devil Crayfish in western Pennsylvania (as well as those in much of West Virginia, Ohio, Kentucky, and Tennessee) as a new species: Cambarus thomai Jezerinac (Little Brown Mudbug). This taxonomic split left only 4 known records of Devil Crayfish in Pennsylvania, all of which were prior to 1905, and none of which were specific beyond the name of a town: Penns Manor in Bucks County, Essington in Philadelphia County, and Ridley Park and Marcus Hook in Delaware County. Field-site Description and Methods From 11 to 14 October 2017, we sampled 37 sites in the Atlantic Coastal Plain in southeastern Pennsylvania in an attempt to locate populations of Devil Crayfish. Our primary objective was to sample suitable habitat near Ortmann’s (1906) 4 historical sites to survey changes to this species’ distribution, but we supplemented our sampling at these historical sites with new sites that appeared to contain suitable habitat for Devil Crayfish. At each site, we spent ~1 h (2 person-hours) searching for any signs of burrowing crayfishes (i.e., mud chimneys or burrow openings in the vicinity of streams or wetlands). Except for 1 site (see below), we excavated every burrow we encountered either by hand or with the help of a small pickaxe to capture the crayfish inside. We identified, preserved in 70% ethanol, and vouchered all captured crayfishes at Ohio State University’s Museum of Biological Diversity crayfish collection (OSUMC 10440–10447). Northeastern Naturalist Vol. 25, No. 3 M.G. Glon, A.J. Mularo, D.A. Lieb, and Z.J. Loughman 2018 357 Results Over the course of our sampling, we captured burrowing crayfishes at 11 of our 37 sites (29.7%; Fig. 1). We found a single crayfish burrow with a chimney at a 12th site but could not locate a crayfish inside of it. However, this burrow did not appear to have been excavated by a primary burrowing crayfish because it led directly into the stream and lacked any perceptible tunnels or chambers. We found crayfish burrows near only 1 of Ortmann’s historical sites (Ridley Park; Fig. 1), all of which contained non-native Procambarus clarkii (Girard) (Red Swamp Crayfish), which is a secondary burrowing crayfish (Hobbs 1974). We also collected Procambarus acutus (Girard) (White River Crayfish) and Cambarus bartonii (Fabricius) (Appalachian Brook Crayfish) during our sampling, both of which are native to Pennsylvania and are also secondary burrowers (Hobbs 1974). We discovered a single population of Devil Crayfish in Neshaminy State Park (Fig. 1). The substrate at the site where we located this population of Devil Crayfish consisted primarily of fine-grained clay soil, which is the preferred soil type of this species (Grow 1982, Helms et al. 2013). Vegetation in surrounding areas was quite dense and consisted primarily of deciduous trees and shrubs. A more thorough survey of the area will be necessary to quantify the size of this Devil Crayfish colony, but we observed several dozen active burrows. To minimize disturbance Figure 1. Map of southeastern Pennsylvania depicting sampling locations where we found crayfish burrows (shapes representing different species), where we did not find burrows (black dots), approximate historical locations (black dashed circles) and the Atlantic coastal plain (grey dotted and dashed line) . Northeastern Naturalist 358 M.G. Glon, A.J. Mularo, D.A. Lieb, and Z.J. Loughman 2018 Vol. 25, No. 3 to this area, we only excavated 5 burrows, and in each case captured a single adult Devil Crayfish (3 form I males, 2 females; carapace length = 32.2 ± 2.3 mm; Fig. 2). We did not capture any other species at this site, however, we cannot rule out the possibility that other species were present in burrows that we did not excavate. Given that we collected Red Swamp Crayfish at several nearby sites and the large potential impacts of non-native crayfishes on native crayfishes (Lodge et al. 2012, Twardochleb et al. 2013), assessing whether this species is present at this site and acting accordingly (e.g., eradication if it is present, prevention if it is not present) should be a conservation priority moving forward. Discussion Our inability to locate crayfish burrows at more than 2/3 of our sites does not preclude the presence of Devil Crayfish there. Possible limitations of our sampling efforts include the fact that burrowing-crayfish chimneys are typically more conspicuous in spring before groundcover emerges than in fall when the ground is covered in leaf litter. It is also possible that we overlooked a small number of Figure 2. Form I male Cambarus diogenes (Devil Crayfish) collected in Neshaminy State Park, PA. Northeastern Naturalist Vol. 25, No. 3 M.G. Glon, A.J. Mularo, D.A. Lieb, and Z.J. Loughman 2018 359 burrows during our search due to the large amount of habitat that we sampled. However, we relied on our substantial experience sampling for Devil Crayfish in other states to locate habitat that seemed suitable for this species, and numerous sites appeared to contain ideal Devil Crayfish habitat yet were entirely devoid of burrows. Therefore, although we cannot rule out the possibility that we overlooked 1 or more Devil Crayfish burrows, we feel confident that the Devil Crayfish was rare, if not entirely absent, from all but the single site in N eshaminy State Park. We strongly recommend that the results of our survey serve as a warning that the Devil Crayfish may be on the brink of extirpation from the state of Pennsylvania. Our findings corroborate those of Bouchard et al. (2007) and Lieb et al. (2011a), who found that crayfish biodiversity in southeastern Pennsylvania has declined due to urbanization and biological invasions by non-native crayfishes. In the case of the Devil Crayfish, this situation is exacerbated by the relatively small portion of southeastern Pennsylvania containing suitable habitat (i.e., the floodplain of the Delaware River). The extirpation of the Devil Crayfish in Pennsylvania could mean losing important genetic variation for this species (Hughes et al. 2008), as well as the disappearance of the critical ecosystem engineering performed by primary burrowing crayfishes (e.g., Glon and Thoma 2017, Pintor and Soluk 2006, Richardson 1983). We suggest that additional sampling be conducted in southeastern Pennsylvania in the Delaware River Floodplain and in parts of the neighboring Piedmont region in an attempt to locate any additional populations of Devil Crayfish. We also recommend that the Devil Crayfish population in Neshaminy State Park be thoroughly surveyed and managed. Acknowledgments We thank the Wild Resources Conservation Fund Grant for providing funding that made this project possible. 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