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Home , Alabama literature, Barbicambarus cornutus, Big Sandy crayfish, Bouchardina, Cambarellus, Cambarellus diminutus

2011. Proceedings of the Academy of Science 120(1-2):71–95

CONSERVATION STATUS OF NORTH AMERICAN FRESHWATER (: ) FROM THE SOUTHERN

Thomas P. Simon: Department of Biology, Indiana State University, 600 Chestnut Street, Room 271, Terre Haute, IN 47809, USA

ABSTRACT. A list is provided of all (family Cambaridae) in the southern United States, which includes common names, global , an alternative review of the conservation status based on the IUCN red list criteria, and state distribution. This list includes 357 native crayfishes, of which 12 (3.4%) are critically endangered, 37 (10.4%) are endangered, 126 (35.3%) are vulnerable, 181 (50.7%) are lower risk, and 1 (0.3%) is not evaluated. The leading factors causing imperilment are restricted ranges caused by anthropogenic impacts from changes in land use, contaminants, invasion by non-indigenous , and habitat fragmentation. In order to conserve and manage diversity of native crayfish, consistency is needed in determining conservation status and more complete distribution and life history information are needed for about 60% of species. Keywords: imperiled species, global vulnerability, International Union for Conservation of Nature, biodiversity,

INTRODUCTION assessments used the best professional judg- North American aquatic biodiversity has ment of taxonomic group experts to determine been disproportionately affected by anthropo- imperilment. genic influences. A major focus is to fill gaps in Crayfishes are closely related to marine research on crayfish ecology and present new (Crandall et al. 2000) and are members information on imperilment of aquatic species. of the order Decapoda, which includes crabs, This need has benefited from the focus pro- lobsters, and . Crayfishes are included duced by the Natural Heritage Global (G) in three families, and native inhabitants occur compilation of conservation status ranks (Mas- in freshwater ecosystems on every continent ter 1991). As a result of the Global status ranks, except Africa and Antarctica. Two families, other groups have increased their emphasis on and Cambaridae, are native to North assessing other freshwater faunal diversity. The America and include about 408 species and American Fisheries Society (AFS) Endangered subspecies, representing about 77% of the Species Committee evaluated the conservation global species diversity in status of North America’s freshwater fish (Taylor 2002). The majority of the North fauna (Deacon et al. 1979; Williams et al. American fauna (99%) is assigned to the family 1989; Warren et al. 2000), freshwater mussels Cambaridae with over two-thirds of its species (Williams et al. 1993), and North American endemic to the southeastern United States. freshwater crayfishes (Taylor et al. 1996, 2007). Crayfishes occur in every seasonally wet and Taylor et al. (2007) assessed the conservation terrestrial habitat including a variety of aquatic status and threats to native crayfishes in the habitats. They possess unique life-history traits United States and Canada using the best adapted for these habitats, including alteration information available, provided updated state/ of reproductive form and burrowing abilities provincial distributions, updated the list of that allow colonization (Hobbs 1981; Welch & references on the biology, conservation, and Eversole 2006). Our purpose for this research is distribution of crayfishes in the United States to report on the conservation status of the and Canada, and assigned standardized com- North American crayfish fauna distributed mon names. All of the AFS conservation in the southeastern United States using the International Union for Conservation of Na- Correspondence: [email protected] ture and Natural Resources (IUCN) red list

71 72 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE criteria. The purpose is to report the status and the wild in the immediate future, as defined by effects that might cause future declines in any of the criteria (A to E) as described in relative abundance and loss of species richness Appendix 1. ENDANGERED (EN) – A taxon (Huner 1994; Taylor et al. 1996; Holdich 2002). is EN when it is not CR, but is facing a very high risk of EW in the near future, as defined METHODS by any of the criteria (A to E) as described in Our review of the conservation status of Appendix 1. VULNERABLE (VU) – A taxon crayfishes includes all species and subspecies is VU when it is not CR or EN but is facing a from the southern United States. We do not high risk of in the wild in the recognize ornatus following Taylor medium-term future, as defined by any of the (1997), we continue to recognize criteria (A to E) as described in Appendix 1. ferrugineus until the work by Robison and LOWER RISK (LR) – A taxon is LR when it Crandall is published in peer-reviewed literature, has been evaluated and does not satisfy the and carinirostris is recognized criteria for any of the categories CR, EN, or as C. carinirostris Hay (Rock Crawfish) following VU. Taxa included in the LR category can be Thoma & Jezerinac (1999). Additional southern separated into three subcategories: (1) Conser- taxa included in our work were described vation Dependent (cd) – Taxa which are the subsequent to Taylor et al. (1996, 2007). focus of a continuing taxon-specific or habitat- Conservation status.—The current assess- specific conservation program targeted to- ment of southern crayfish includes several wards the taxon in question, the cessation of rating systems that are used by the Heritage which would result in the taxon qualifying for Database and the International Union for one of the threatened categories above within a Conservation of Nature and Natural Resources period of five years; (2) Near Threatened (nt) – (IUCN) red list criteria. Ranking categories Taxa which do not qualify for cd, but which for conservation status follow Master (1991) are close to qualifying for VU; and (3) Least and are defined as follows: G1 5 critically Concern (lc) – Taxa which do not qualify for imperiled, G2 5 imperiled, G3 5 vulnerable cd or nt. DATA DEFICIENT (DD) – A taxon to extirpation or extinction, G4 5 apparently is DD when there is inadequate information to secure, G5 5 demonstrably widespread, abun- make a direct, or indirect, assessment of its dant, and secure, GH 5 possibly extinct, known risk of extinction based on its distribution and/ only from historical collections, T 5 threatened, or population status. A taxon in this category and GX 5 presumed extinct. The conservation may be well studied, and its biology well ranks for each taxon follow the system devel- known, but appropriate data on abundance oped by The Nature Conservancy/ NatureServe and/or distribution are lacking. Data Deficient and the Network of Natural Heritage Programs is therefore not a category of threat or Lower (Master 1991, www.natureserve.org/explorer/ Risk. Listing of taxa in this category indicates ranking.htm). The International Union for that more information is required and ac- Conservation of Nature and Natural Resources knowledges the possibility that future research (IUCN) includes the following six conservation will show that threatened classification is categories: EXTINCT (EX) – when there is no appropriate. It is important to make positive reasonable doubt that the last individual has use of whatever data are available. In many died. EXTINCT IN THE WILD (EW) – when it cases great care should be exercised in choos- is known only to survive in cultivation, in ing between DD and threatened status. If the captivity or as a naturalized population (or range of a taxon is suspected to be relatively populations) well outside the past range. A circumscribed, and if a considerable period of taxon is presumed EW when exhaustive surveys time has elapsed since the last record of the in known and/or expected habitat, at appropri- taxon, threatened status may well be justified. ate times (diurnal, seasonal, annual), through- NOT EVALUATED (NE) – A taxon is NE out its historic range have failed to record an when it is has not yet been assessed against the individual. Surveys should be over a time frame criteria (Appendix 1). appropriate to the taxon’s life cycle and life List of taxa.—The list of crayfish species and form (Appendix 1). CRITICALLY ENDAN- subspecies is arranged alphabetically by GERED (CR) – A taxon is CR when it is and by species and subspecies within genus facing an extremely high risk of extinction in (Table 1). Following scientific names, common SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 73 names are based primarily on those provided Hobbs 1990; Johnson & Johnson 2008), by two internet sites; the Crayfish Tree of Life – District of Columbia (Loughman crayfish.byu.edu, and the Global Crayfish and 2009), and West (Jezerinac et al. 1995; browser – iz.carnegiemnh. Loughman et al. 2009; Jones et al. 2010; org/crayfish/NewAstacidea/index.aspwww. Loughman & Welsh 2010). Each state is crayfish.byu.edu. However, we also used com- designated by the two-digit postal code. mon names as reported in state lists, taxonomic Parentheses around states indicate known or descriptions, McLaughlin et al. (2005), and suspected introductions. Taylor et al. (2007). In determining conservation status and RESULTS AND DISCUSSION distribution, a variety of sources were used The list of southern crayfishes includes 357 including state and federal endangered species taxa (Table 1). This list includes 357 native lists, government agency reports and websites, crayfishes, of which 12 (3.4%) are critically research publications, and books. Global Her- endangered, 37 (10.4%) are endangered, 126 itage Rank criteria were based on numerals G1 (35.3%) are vulnerable, 181 (50.7%) are lower through G5 and correspond to those defined risk, and 1 (0.3%) was not evaluated. Taxo- in the Methods. Global Heritage ranks imme- nomic efforts since Hobbs’ (1989b) checklist of diately follow common name. Following the North American crayfishes have resulted in the Global Heritage ranks is the conservation description of many new crayfish species in the status based on the IUCN conservation for- United States. New descriptions have averaged mula. Finally, the distribution of each taxon slightly less than two new species per year. This is indicated by an alphabetical listing of USA demonstrates the need to train taxonomists to states within the Southern Division of the further study biodiversity that continues to be American Fisheries Society where it occurs. undiscovered in North America. The number Distribution.—The distribution reflected for of imperiled crayfishes (49.0%) ranks interme- each species is based only on its range within diate between the high levels observed in fishes the Southern United States. This geographic and freshwater mussels, almost 33% and 72%, state inclusion in based on the American respectively (Williams et al. 1989; Williams Fisheries Society Southern Division member- et al. 1993; Warren & Burr 1994; Warren et al. ship (Fig. 1). Distribution information is based 2000). Our assessment is similar to the findings on reports for (Bouchard 1976; of Taylor et al. (2007), who assessed 48% of Harris 1990; McGregor et al. 1999; Ratcliffe crayfish as imperiled. This assessment confirms & DeVries 2004; Schuster & Taylor 2004; the assertion that North American diversity in Heath et al. 2010), (Williams 1954; aquatic systems is in poorer condition than that Bouchard & Robison 1980; Hobbs & Robison in terrestrial systems (Master 1991; Master 1988; Hobbs 1989a), (Hobbs 1942; et al. 2000). The benefit of this assessment is Deyrup & Franz 1994; Franz & Franz 1990; the validation of the assessment of conservation Hobbs & Hobbs 1991), (Hobbs 1981; status by Taylor et al. (2007) and the stabiliza- Skelton 2010), (Rhoades 1944; Burr tion of the assessment process by removing best & Hobbs 1984; Taylor & Schuster 2004), professional judgment from the analysis. Tay- (Penn 1950, 1952, 1956, 1959; Penn & Marlow 1959; Walls & Black 1991; Walls lor et al. (1996) used as part of the American and Shively 2003; Walls 2009), Fisheries Society’s review process the recogni- (Meredith & Schwartz 1959, 1960; Kilian et tion of either the actual or potential imperil- al. 2010; Loughman 2010), (Adams ment of crayfishes between governmental agen- 2008), North Carolina (Cooper & Braswell cies charged with protecting natural resources 1995; Cooper et al. 1998; Cooper 2002; and non-profit conservation organizations as a LeGrand et al. 2006; Simmons and Fraley rationale for listing. Only four crayfish species 2010), (Creaser & Ortenburger 1933; (Pacifastacus fortis, , Cam- Dunlap 1951; Reimer 1969; Jones et al. 2005; barus zophonastes, and shoupi) Taylor et al. 2004), (Hobbs receive protection under the Federal Endan- et al. 1976; Eversole 1995; Eversole & Jones gered Species Act of 1973 (ESA) and 66 species 2004), (Bouchard 1972), received varying levels of protection at the state (Penn & Hobbs 1958; Albaugh & Black 1973; level, which is lower than the 197 species listed 74 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Checklist of crayfishes of the Southeastern United States including common name, Global Heritage rank, International Union for Conservation of Nature and Natural Resources (IUCN) red list, and distribution.

Species Global Rank Red List Distribution cornutus (Faxon) G4 LR lc KY, TN (Bottle Brush Crayfish) Barbicambarus simmonsi Taylor G3 VU D2 TN and Shuster (Tennessee Bottle Brush Crayfish) robisoni Hobbs G2 VU B1+2c AR (Bayou Bodcau Crayfish) blacki Hobbs G1 EN B1+2c FL (Cypress Crayfish) Hobbs G3 VU B1+2c AL, MS (Least Crayfish) Cambarellus lesliei Fitzpatrick G3 VU B1+2c AL, MS and Laning (Angular Dwarf Crayfish) Cambarellus ninae Hobbs G3 VU B1+2c TX (Aransas Dwarf Crawfish) Hobbs (Swamp G5 LR lc AR, KY, LA, MS, Dwarf Crayfish) OK, TN, TX Cambarellus schmitti Hobbs G3 LR lc FL (Fontal Dwarf Crawfish) (Faxon) G5 LR lc AL, AR, KY, LA, (Cajun Dwarf Crayfish) MS, TN, TX Cambarellus texanus Albaugh G3G4 LR lc TX and Black (Brazos Dwarf Crawfish) Cambarus acanthura Hobbs G4G5 LR lc AL, GA, NC, TN (Thornytail Crayfish) Cambarus aculabrum Hobbs and G1 EN B1+2c AR Brown (Benton County Cave Crayfish) Cambarus acuminatus Faxon G4 LR nt MD, NC, SC, VA (Acuminate Crayfish) Cambarus aldermanorum Cooper G3 VU B1, D1 SC and Price (Carolina needlenose crayfish) Cambarus angularis Hobbs and G3 LR lc TN, VA Bouchard (Angled Crayfish) Faxon G4 LR lc GA, NC, SC, TN (Mitten Crayfish) Cambarus bartonii bartonii G5T5 LR lc AL, GA, KY, MD, (Fabricius) (Common NC, SC, TN, VA, Crayfish) WV Cambarus bartonii cavatus Hay G5T5 LR lc AL, KY, WV (Appalachian Brook Crayfish) Cambarus batchi Schuster G3 VU B1+2c, D1 KY (Bluegrass Crayfish) Hobbs G2 VU B1+2c, D1 KY, TN (Big South Fork Crayfish) Cambarus brachydactylus Hobbs G4 LR lc TN (Shortfinger Crayfish) Cambarus brimleyorum Cooper G3 VU B1+2c NC (Valley River Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 75

Table 1.—Continued. Species Global Rank Red List Distribution Cambarus buntingi Bouchard G4 LR nt KY, TN, VA (Bunting Crayfish) Cambarus carinirostris Hay G5 LR lc MD, VA, WV (Rock Crawfish) Cambarus carolinus (Erichson) G4 LR lc NC, SC, TN (Red Burrowing Crayfish) Hobbs and G3 VU B1+2c NC Perkins (Greensboro Burrowing Crayfish) Cambarus causeyi Reimer G2 VU A1,B1+2c AR (Boston Mountains Crayfish) James G4 LR lc NC, VA, WV (New River Crayfish) Prins and G2 VU B1+2c, D1 GA, NC, SC Hobbs (Chauga Crayfish) Cambarus clivosus Taylor, G2 VU B1+2c, D1 TN Soucek, and Organ (Short Mountain Crayfish) Cambarus conasaugaensis Hobbs G3 VU B1+2c GA, TN and Hobbs (Mountain Crayfish) Cambarus coosae Hobbs (Coosa G5 LR lc AL, GA, TN Crayfish) Cambarus coosawattae Hobbs G1 VU B1+2c, D1 GA (Coosawattae Crayfish) Cambarus cracens Bouchard and G1 EN B1+2c AL Hobbs (Slenderclaw Crayfish) Cambarus crinipes Bouchard G3 LR lc TN (Bouchard’s Crayfish) Hobbs G2G3 VU B1+2c FL, GA (Dougherty Plain Cave Crayfish) Cambarus cumberlandensis Hobbs G5 LR lc KY, TN and Bouchard (Cumberland Crayfish) Hobbs G1 VU B1+2c, D1 GA, TN (Conasauga Blue Burrower Crayfish) Cooper G3 LR lc NC (Carolina Ladle Crayfish) Cambarus deweesae Bouchard G4 EN B1+2c KY, TN and Etnier (Valley Flame Crayfish) Girard G5 LR lc AL, AR, FL, GA, (Devil Crawfish) KY, LA, MD, MS, NC, OK, SC, TN, TX, VA Cambarus distans Rhoades G5 LR lc AL, GA, KY, TN (Boxclaw Crawfish) Cambarus doughertyensis Cooper G1G2 EN B1+2c GA and Skelton (Dougherty Burrowing Crayfish) Cambarus dubius Faxon (Upland G5 LR lc KY, MD, NC, TN, Burrowing Crayfish) VA, WV Cambarus eeseeohensis Thoma G1 EN B1+2c NC (Linville River Crayfish) Cambarus elkensis Jezerinac and G2 VU B1+2c WV Stocker (Elk River Crayfish) 76 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Cambarus englishi Hobbs and G3 VU B1+2c AL, GA Hall (Tallapoosa Crayfish) Cambarus erythrodactylus Simon G5 LR lc AL, GA, MS and Morris (Warpaint Mudbug) Hagen G2 VU B1+2c GA, TN (Chickamauga Crayfish) Hobbs G3 VU B1+2c GA (Etowah Crayfish) Cambarus friaufi Hobbs (Hairy G4 LR lc KY, TN Crayfish) Cambarus gentryi Hobbs (Linear G4 LR lc TN Cobalt Crayfish) Hobbs (Little G2 VU B1+2c, D1 GA, NC Tennessee Crayfish) Cambarus girardianus Faxon G5 LR lc AL, GA, MS, TN (Tanback Crayfish) Cambarus graysoni Faxon G5 LR lc AL, KY, TN (Twospot Crayfish) Cambarus halli Hobbs G3G4 LR lc AL, GA (Slackwater Crayfish) Cambarus hamulatus (Cope) G3G4 LR lc AL, TN (Prickly Cave Crayfish) Hobbs ( G1 EN B1+2c GA Blue Burrower) Hobbs G3G4 VU B1+2c GA, NC, TN (Hiwassee Crayfish) Cambarus hobbsorum Cooper G3G4 LR lc NC, SC (Rocky River Crayfish) Hobbs and G3 VU B1+2c AL, GA, NC Hall (Chattahoochee Crayfish) Cambarus hubbsi Creaser G5 LR lc AR (Hubb’s Crayfish) Cambarus hystricosus Cooper G2 VU B1+2c NC and Cooper (Sandhills Spiny Crayfish) Cambarus jezerinaci Thoma G3 LR nt TN, VA (Powell River Crayfish) Cambarus johni Cooper (Carolina G3 VU B1+2C NC Foothills Crayfish) Cambarus jonesi Hobbs and Barr G2 LR lc AL (Alabama Cave Crayfish) Cambarus laconensis Buhay and G1 VU B1+2C AL Crandall (Lacon Exit Cave Crayfish) Cambarus latimanus (LeConte) G5 LR lc AL, FL, GA, NC, (Variable Crayfish) SC, TN Cambarus lenati Cooper (Broad G2 VU B1+2c NC River Crayfish) Cambarus longirostris Faxon G5 LR lc AL, GA, NC, (SC), (Longnose Crayfish) TN, VA Cambarus longulus Girard G5 LR lc NC, VA, WV (Atlantic Slope Crayfish) Cambarus ludovicianus Faxon G5 LR lc AL, AR, KY, LA, (Painted Devil Crayfish) MS, TN, TX SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 77

Table 1.—Continued. Species Global Rank Red List Distribution Cambarus manningi Hobbs G4 LR lc AL, GA, TN (Greensaddle Crayfish) Cambarus miltus Fitzpatrick G2 VU B1+2c AL, FL (Rusty Gravedigger) Cambarus monongalensis G5 LR lc VA, WV Ortmann (Blue Crawfish) Cambarus nerterius Hobbs G2 VU B1+2c, D1 WV (Greenbrier Cave Crayfish) Cambarus nodosus Bouchard and G4 LR lc GA, NC, SC, TN Hobbs (Knotty Burrowing Crayfish) Hobbs and G1 VU B1+2c TN Shoup (Obey Crayfish) Cambarus obstipus Hall (Sloped G4 VU B1+2c AL Crayfish) Cambarus ortmanni Williamson G5 LR lc KY (Ortmann’s Mudbug) Hobbs G1 VU B1+2c, D1 GA, NC (Hiwassee Headwater Crayfish) Cambarus parvoculus Hobbs and G5 LR lc AL, GA, KY, TN, Shoup (Mountain Midget VA Crayfish) Cambarus pecki Hobbs (Phantom G1G2 VU B1+2c AL Cave Crayfish) Cambarus polychromatus Thoma, G5 LR lc AL, KY, TN Jezerinac, and Simon (Paintedhand Mudbug) Hobbs G2 VU B1+2c, D1 TN (Pristine Crayfish) Cambarus pyronotus Bouchard G2 EN B1+2c FL (Fireback Crayfish) Prins (French G3 VU B1+2c NC Broad Crayfish) Cambarus reduncus Hobbs G4G5 LR lc NC, SC (Sickle Crayfish) Cambarus reflexus Hobbs (Pine G4 LR lc GA, SC Savannah Crayfish) Girard (Big G5 LR lc KY, NC, TN, VA, Water Crayfish) WV Cambarus rusticiformis Rhoades G5 LR lc (AL), KY, TN (Depression Crayfish) Cambarus sciotensis Rhoades G5 LR lc KY, VA, WV (Teays River Crayfish) Cambarus scotti Hobbs G3 VU B1+2c AL, GA (Chatooga Crayfish) Cambarus setosus Faxon G3 LR lc AR, OK (Bristly Cave Crayfish) Cambarus smilax Loughman, G4 LR lc WV Simon, and Welch (Greenbrier Crayfish) Cambarus sp. 1 (Emory River G1? NE TN Crayfish) Cambarus speleocoopi Buhay and G1 VU B1+2C AL Crandall (Sweet Home Alabama Crayfish) 78 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Cambarus speciosus Hobbs G2 VU B1+2c, D1 GA (Beautiful Crayfish) Cambarus sphenoides Hobbs G4 LR lc KY, TN (Triangleclaw Crayfish) Hobbs G3 VU B1+2c NC, SC (Broad River Spiny Crayfish) Cambarus striatus Hay (Hay G5 LR lc AL, FL, GA, KY, Crayfish) MS, SC, TN Cambarus strigosus Hobbs (Lean G2 EN B1+2c GA Crayfish) Cambarus subterraneus Hobbs G1 EN B1+2c OK ( County Cave Crayfish) Cambarus tartarus Hobbs and G1 CR B1+2c OK Cooper (Oklahoma Cave Crayfish) Cambarus tenebrosus Hay G5 LR lc AL, GA, KY, TN (Cavespring Crayfish) Cambarus thomai Jezerinac G5 LR lc KY, TN, WV (Little Brown Mudbug) Cambarus trojanensis Simon and G5 LR lc AL, FL, MS Morris (Spiny Collared Mudbug) Cambarus truncatus Hobbs G2 EN B1+2c GA (Oconee Burrowing Crayfish) Cambarus tuckasegee Cooper G1G2 VU B1+2c NC and Schofield (Tuckasegee Stream Crayfish) Hobbs and G2 VU B1+2c AL, GA Hall (Blackbarred Crayfish) Cambarus veitchorum Cooper and G1 EN A1, B1+2c AL Cooper (White Spring Cave Crayfish) Faxon G2G3 VU B1+2c KY, VA, WV () Cambarus williami Bouchard and G2 EN B1+2c TN Bouchard (Brawleys Fork Crayfish) Hobbs G1 CR B1+2c AR and Bedinger (Hell Creek Cave Crayfish) carlsoni Hobbs G2G3 VU B1+2c SC (Mimic Crayfish) Distocambarus crockeri Hobbs G3 VU B1+2c, D1 SC and Carlson (Piedmont Prairie Burrowing Crayfish) Distocambarus devexus (Hobbs) G1 VU B1+2c, D1 GA (Broad River Burrowing Crayfish) Distocambarus hunteri Fitzpatrick G1 EN B1+2c SC and Eversole (Saluda Burrowing Crayfish) Distocambarus youngineri Hobbs G1 VU B1+2c, D1 SC and Carlson (Newberry Burrowing Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 79

Table 1.—Continued. Species Global Rank Red List Distribution burrisi Fitzpatrick G3 VU B1+2c AL, MS (Burrowing Bog Crayfish) Fallicambarus byersi (Hobbs) G4 LR lc AL, FL, MS (Lavender Burrowing Crayfish) Fallicambarus caesius Hobbs G4 LR lc AR (Timberlands Burrowing Crayfish) Hobbs G2 VU B1+2c AL, MS (Speckled Burrowing Crayfish) Fallicambarus devastator Hobbs G3 VU B1+2c TX and Whiteman (Texas Prairie Crayfish) Fallicambarus dissitus (Penn) G4 VU B1+2c AR, LA (Pine Hills Digger Crayfish) Fallicambarus fodiens (Cottle) G5 LR lc AL, AR, FL, GA, (Digger Crayfish) KY, LA, MD, MS, NC, OK, SC, TN, TX, VA, WV Fallicambarus gilpini Hobbs and G1 VU B1+2c, D1 AR Robison (Jefferson County Crayfish) Fitzpatrick G1 VU B1+2c, D1 MS (Camp Shelby Burrowing Crayfish) Fallicambarus harpi Hobbs and G1 EN B1+2c AR Robison (Ouachita Burrowing Crayfish) Fallicambarus hortoni Hobbs and G1 EN B1+2c TN Fitzpatrick (Hatchie Burrowing Crayfish) Fallicambarus houstonensis G3 VU B1+2c, D1 TX Johnson (Houston Burrowing Crayfish) Fallicambarus jeanae Hobbs G2 EN B1+2c AR (Daisy Burowing Crayfish) Fallicambarus kountzeae Johnson G3 VU B1+2c, D1 TX (Big Thicket Burrowing Crayfish) Fallicambarus macneesei (Black) G3 VU B1+2c, D1 LA, TX (Old Prairie Digger Crayfish) Fallicambarus oryktes (Penn and G4 EN B1+2c, B2 AL, LA, MS Marlow) (Flatwoods Digger Crayfish) Fallicambarus petilicarpus Hobbs G1 CR B1+2c AR and Robison (Slenderwrist Burrowing Crayfish) (Reimer) G1G2 VU B1+2c AR (Saline Burrowing Crayfish) beyeri (Penn) (Sabine G4 LR lc LA, TX Fencing Crayfish) Faxonella blairi Hayes and G2 LR lc AR, OK Reimer (Blair’s Fencing Crayfish) Faxonella clypeata (Hay) G5 LR lc AL, AR, FL, GA, (Ditch Fencing Crayfish) LA, MS, OK, SC, TX 80 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Walls G2 VU B1+2c, D1 LA (Ouachita Fencing Crayfish) attenuatus Black (Pearl G2 VU B1+2c, D1 MS Riverlet Crayfish) (Hobbs) G3 VU B1+2c MS (Crested Rivulet Crayfish) G3 VU B1+2c MS Fitzpatrick and Payne (Oktibbeha Riverlet Crayfish) Fitzpatrick G2 VU B1+2c MS (Tombigbee Riverlet Crayfish) Hobbseus prominens (Hobbs) G4G5 LR lc AL, MS (Prominence Riverlet Crayfish) Hobbseus valleculus (Fitzpatrick) G1 VU B1+2c MS (Choctaw Riverlet Crayfish) Hobbseus yalobushensis G3 VU B1+2c, D1 MS Fitzpatrick and Busack (Yalobusha Riverlet Crayfish) Orconectes acares Fitzpatrick G4 LR lc AR, OK (Redspotted Stream Crayfish) Orconectes alabamensis (Faxon) G5 LR lc AL, MS, TN (Alabama Crayfish) Orconectes australis australis G5T4 LR lc AL, TN (Rhoades) (Southern Cave Crayfish) Orconectes australis packardi G2 EN B1+2c KY Rhoades (Appalachian Cave Crayfish) Orconectes barrenensis Rhoades G4 LR lc KY, TN (Barren River Crayfish) Orconectes barri Buhay and G2 VU B1+2C KY, TN Crandall ( Cave Crayfish) Orconectes bisectus Rhoades G1 VU B1+2c, D1 KY (Crittenden Crayfish) Orconectes blacki Walls G2 VU B1+2c, D1 LA (Calcasieu Crayfish) Orconectes burri Taylor and G1 EN B1+2c, D1 KY, TN Sabaj (Burr Crayfish) Orconectes carolinensis Cooper G3 LR lc NC and Cooper (North Carolina Crayfish) Orconectes causeyi Jester G4 LR lc OK, TX (Western Plains Crayfish) Orconectes chickasawae Cooper G5 LR lc AL, MS and Hobbs (Chickasaw Crayfish) Orconectes compressus (Faxon) G5 LR lc AL, KY, MS, TN (Slender Crayfish) Orconectes cooperi Cooper and G1 VU B1+2c, D1 AL, TN Hobbs (Flint River Crayfish) Orconectes cristavarius Taylor G5 LR lc KY, NC, TN, VA, (Spiny Stream Crayfish) WV Orconectes deanae Reimer and G4 VU B1+2c, D1 OK Jester (Conchas Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 81

Table 1.—Continued. Species Global Rank Red List Distribution Orconectes difficilis (Faxon) G3 LR lc AR, OK (Painted Crayfish) Orconectes durelli Bouchard and G5 LR lc AL, KY, TN Bouchard (Saddle Crayfish) (Faxon) G5 LR lc AL, GA, TN, VA (Reticulate Crayfish) Orconectes etnieri Bouchard G4 LR lc MS, TN and Bouchard (Etnier’s Crayfish) Orconectes eupunctus Williams G2 EN B1+2c, D2 AR, MO (Coldwater Crayfish) Orconectes forceps (Faxon) G5 LR lc AL, GA, TN, VA (Surgeon Crayfish) Orconectes hartfieldi Fitzpatrick G2 VU B1+2c MS and Suttkus (Yazoo Crayfish) Orconectes hathawayi Penn G3 VU B1+2c LA (Teche Painted Crayfish) Orconectes hobbsi Penn G4 LR lc LA, MS (Pontchartrain Painted Crayfish) Orconectes holti Cooper and G3 VU B1+2c AL Hobbs (Bimaculate Crayfish) Orconectes immunis (Hagen) G5 LR lc AL, KY, TN (Calico Crayfish) Orconectes incomptus Hobbs and G1 VU B1+2c, D1 TN Barr (Tennessee Cave Crayfish) Orconectes inermis inermis Cope G5T4 LR lc KY (Northern Cave Crayfish) Orconectes jeffersoni Rhoades G1 VU B1+2c, D1 KY (Louisville Crayfish) Orconectes jonesi Fitzpatrick G3 VU B1+2c AL, MS (Sucarnoochee River Crayfish) Orconectes juvenilis (Hagen) G4 LR lc KY (Kentucky River Crayfish) Orconectes kentuckiensis Rhoades G4 VU B1+2c KY (Kentucky Crayfish) Orconectes lancifer (Hagen) G5 LR lc AL, AR, KY, LA, ( Crayfish) MS, OK, TN, TX Orconectes leptogonopodus Hobbs G4 LR lc AR, OK (Little River Creek Crayfish) (Rafinesque) G5 VU A1, B1 MD, WV (Spinycheek Crayfish) Orconectes longidigitus (Faxon) G4 LR lc AR (Longpincered Crayfish) Orconectes luteus (Creaser) G5 LR lc AR (Golden Crayfish) Orconectes macrus Williams G4 LR lc AR, OK (Neosho Midget Crayfish) Walls G2 VU B1+2c LA, TX (Kisatchie Painted Crayfish) Orconectes marchandi Hobbs G2 VU B1+2c, D1 AR (Sharp River Crayfish) Orconectes margorectus Taylor G2 EN B1+2c KY (Livingston Crayfish) 82 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Orconectes meeki brevis G4T3 EN B1+2c AR, OK Williams (Meek’s Short Pointed Crayfish) Orconectes meeki meeki (Faxon) G4T4 LR lc AR (Meek’s Crayfish) Orconectes menae (Creaser) G3 VU B1+2c AR, OK (Mena Crayfish) Orconectes mirus (Ortmann) G4 LR lc AL, TN (Wonderful Crayfish) Orconectes mississippiensis G3 VU B1+2c MS (Faxon) (Mississippi Crayfish) Orconectes nais (Faxon) G5 LR lc AR, OK, TX (Water Nymph Crayfish) Orconectes nana Williams G3 VU B1+2c AR, OK (Midget Crayfish) Orconectes neglectus G5T3 VU B1+2c AR chaenodactylus Williams (Gaped Ringed Crayfish) Orconectes neglectus neglectus G5T5 LR lc AR, OK (Faxon) (Ringed Crayfish) (Hagen) G5 LR lc MD, VA, WV (Allegheny Crayfish) Orconectes ozarkae Williams G5 LR lc AR (Ozark Crayfish) Orconectes pagei Taylor and G4 LR lc TN Sabaj (Mottled Crayfish) Orconectes palmeri creolanus G5T4 LR lc (GA), LA, MS (Creaser) (Creole Painted Crayfish) Orconectes palmeri longimanus G5T5 LR lc AR, OK, TX (Faxon) (Western Painted Crayfish) Orconectes palmeri palmeri G5T5 LR lc AR, KY, MS, TN (Faxon) (Gray-speckled Crayfish) Orconectes pardalotus Wetzel, G1 VU B1+2C KY Poly, Fetzner (Leopard Crayfish) Orconectes pellucidus (Tellkampf) G4 LR lc KY, TN (Mammoth Cave Crayfish) Orconectes perfectus Walls G4G5 LR lc AL, MS (Complete Crayfish) Orconectes placidus (Hagen) G5 LR lc AL, KY, TN (Placid Crayfish) Orconectes punctimanus (Creaser) G4G5 LR lc AR (Spothand Crayfish) Orconectes putnami (Faxon) G5 LR lc AL, KY, TN (Phallic Crayfish) Orconectes rafinesquei Rhoades G3 VU B1+2c, D1 KY (Rafinesque Crayfish) Orconectes rhoadesi Hobbs G4 LR lc TN (Fishhook Crayfish) Orconectes ronaldi Taylor G3 LR lc KY (Mud River Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 83

Table 1.—Continued. Species Global Rank Red List Distribution Orconectes rusticus (Girard) G5 LR lc KY, (NC), (TN), () (VA), (WV) G4G5 LR lc WV erismophorous Hobbs and Fitzpatrick Orconectes sanbornii sanbornii G4G5T4 LR lc KY, WV (Faxon) (Sanborn’s Crayfish) Bouchard G1 CR B1+2c OK and Bouchard (Kiamichi Crayfish) Orconectes sheltae Cooper and G1 EN B1+2c, D1 AL Cooper (Shelta Cave Crayfish) Hobbs G1G2 CR B1+2c TN (Nashville Crayfish) Orconectes spinosus (Bundy) G4 LR lc AL, GA, TN (Coosa River Spiny Crayfish) Orconectes taylori Schuster G2 VU B1+2C TN (Crescent Crayfish) Orconectes tricuspis Rhoades G4 LR lc KY (Rhode Crayfish) (Faxon) G5 LR lc AL, MS, TN (Powerful Crayfish) Orconectes virginiensis Hobbs G3 LR lc NC, VA (Chowanoke Crayfish) Orconectes virilis Hagen (Virile G5 LR lc (AL), AR, (MD), Crayfish) MS, (NC), OK, (TN), TX, (VA), (WV) Orconectes williamsi Fitzpatrick G3G4 LR lc AR (Williams Crayfish) Orconectes wrighti Hobbs G2 VU B1+2c, D1 MS, TN (Hardin Crayfish) Procambarus ablusus Penn G4 LR lc MS, TN (Hatchie River Crayfish) Procambarus acherontis G1 EN B1+2c FL (Lonnberg) (Orlando Cave Crayfish) Procambarus acutissimus (Girard) G5 LR lc AL, GA, MS (Sharpnose Crayfish) (Girard) G5 LR lc AL, AR, DC, FL, (White River Crawfish) GA, KY, LA, MD, MS, NC, OK, SC, TN, TX, VA, WV Procambarus advena (LeConte) G3 LR lc GA (Vidalia Crayfish) Procambarus alleni (Faxon) G4 LR lc FL (Everglades Crayfish) Procambarus ancylus Hobbs G4G5 LR lc NC, SC (Coastal Plain Crayfish) Procambarus angustatus GX CR B1+2c GA (LeConte) (Sandhills Crayfish) Procambarus apalachicolae Hobbs G2 VU B1+2c FL (Coastal Flatwoods Crayfish) Procambarus attiguus Hobbs and G1G2 CR B1+2c FL Franz (Silver Glen Springs Crayfish) 84 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Procambarus barbatus (Faxon) G5 LR lc GA, SC (Wandering Crayfish) Procambarus barbiger Fitzpatrick G2 VU B1+2c MS (Jackson Prairie Crayfish) Procambarus bivittatus Hobbs G5 LR lc AL, FL, LA, MS (Ribbon Crayfish) Procambarus blandingii (Harlan) G4 LR lc NC, SC (Santee Crayfish) Procambarus braswelli Cooper G3 VU B1+2c NC, SC (Waccamaw Crayfish) Procambarus brazoriensis Albaugh G1 VU B1+2c, D1 TX (Brazoria Crayfish) Procambarus capillatus Hobbs G3 VU B1+2c AL, FL (Capillaceous Crayfish) Procambarus caritus Hobbs G4 LR lc GA (Poor Crayfish) Procambarus ceruleus G1G3 EN B1+2c TX Fitzpatrick and Wicksten (Blueclaw Chimney Crayfish) Procambarus chacei Hobbs G4 LR lc GA, SC (Cedar Creek Crayfish) (Girard) G5 LR lc AL, AR, FL, (GA), (Red Swamp Crawfish) KY, LA, (MD), MS, (NC), OK, (SC), TN, TX, (VA) Procambarus clemmeri Hobbs G5 LR lc AL, LA, MS (Cockscomb Crayfish) Procambarus cometes Fitzpatrick G1 VU B1+2c, D1 MS (Mississippi Flatwoods Crayfish) Procambarus connus Fitzpatrick GH VU B1+2c, D1 MS (Carrollton Crayfish) Procambarus curdi Reimer G5 LR lc AR, OK, TX (Red River Burrowing Crayfish) Hobbs G1 CR B1+2c FL and Franz (Bigcheek Cave Crayfish) Procambarus dupratzii Penn G5 LR lc AR, OK, LA, TX (Southwestern Creek Crayfish) Procambarus echinatus Hobbs G3 VU B1+2c SC (Edisto Crayfish) Hobbs G1 EN B1+2c FL (Panama City Crayfish) Procambarus elegans Hobbs G4 LR lc AR, LA, MS (Elegant Creek Crayfish) Procambarus enoplosternum G4G5 LR lc GA, SC Hobbs (Black Mottled Crayfish) Procambarus epicyrtus Hobbs G3 VU B1+2c GA (Humpback Crayfish) Relyea G1G2 EN B1+2c FL and Sutton (Santa Fe Cave Crayfish) Procambarus escambiensis Hobbs G2 VU B1+2c, D1 AL, FL (Escambia Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 85

Table 1.—Continued. Species Global Rank Red List Distribution Procambarus evermanni (Faxon) G4 LR lc AL, FL, MS (Panhandle Crayfish) (Hagen) G5 LR lc FL, GA (Slough Crayfish) Procambarus ferrugineus Hobbs G1 EN B1+2c AR and Robison (Lonoke Crayfish) Hobbs G2 VU B1+2c MS (Spinytail Crayfish) Procambarus franzi Hobbs and G1G2 EN B1+2c FL Lee (Orange Lake Cave Crayfish) Procambarus geminus Hobbs G3G4 LR lc AR, LA (Twin Crawfish) Procambarus geodytes Hobbs G4 LR lc FL (Muddiver Crayfish) Procambarus gibbus Hobbs G3 VU B1+2c GA (Muckalee Crayfish) Procambarus gracilis (Bundy) G5 LR lc AR, OK, TX (Prairie Crayfish) Procambarus hagenianus G4G5T4 LR lc AL, MS hagenianus (Faxon) (Southeastern Prairie Crayfish) Procambarus hagenianus vesticeps G4G5T3 VU B1+2c MS Fitzpatrick (Egyptian Crayfish) Procambarus hayi (Faxon) G5 LR lc AL, MS, TN (Straightedge Crayfish) Procambarus hinei (Ortmann) G5 LR lc LA, TX (Marsh Crayfish) Procambarus hirsutus Hobbs G4 LR lc SC (Shaggy Crayfish) Hobbs and G1 EN B1+2c FL Means (Big Blue Springs Crayfish) Procambarus howellae Hobbs G5 LR lc GA (Ornate Crayfish) Procambarus hubbelli (Hobbs) G4 LR lc AL, FL (Jackknife Crayfish) Procambarus hybus Hobbs and G5 LR lc AL, MS Walton (Smoothnose Crayfish) Procambarus incilis Penn (Cut G4 LR lc TX Crayfish) Procambarus jaculus Hobbs and G4 LR lc LA, MS Walton (Javelin Crayfish) Procambarus kensleyi Hobbs G4 LR lc LA, TX (Free State Chimney Crayfish) Procambarus kilbyi (Hobbs) G4 LR lc FL (Hachet Crayfish) Black G2 VU B1+2c AL, MS (Lagniappe Crayfish) Procambarus latipleurum Hobbs G2 VU B1+2c, D1 FL (Wingtail Crayfish) Procambarus lecontei (Hagen) G3G4 VU B2+3a AL, MS (Mobile Crayfish) 86 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Procambarus leitheuseri Franz G1G2 VU B1+2c, D1 FL and Hobbs (Coastal Lowland Cave Crayfish) Procambarus leonensis Hobbs G4 LR lc FL (Blacknose Crayfish) Procambarus lepidodactylus G4 LR lc SC Hobbs (Pee Dee Lotic Crayfish) Procambarus lewisi Hobbs and G4 VU B2+3a AL Walton (Spur Crayfish) Procambarus liberorum G4 LR lc AR, OK Fitzpatrick (Osage Burrowing Crayfish) Procambarus litosternum Hobbs G4 LR lc GA (Blackwater Crayfish) Procambarus lophotus Hobbs G5 LR lc AL, GA, TN and Walton (Mane Crayfish) Procambarus lucifugus alachua G2G3T2T3 VU B2+3a, D2 FL (Hobbs) (Alachua Light Fleeing Crayfish) Procambarus lucifugus lucifugus G2G3T2 EN B1+2c FL (Hobbs) (Florida Cave Crayfish) Procambarus lunzi (Hobbs) G4 LR lc GA, SC (Hummock Crayfish) Procambarus lylei Fitzpatrick G2 VU B1+2c MS and Hobbs (Shutispear Crayfish) Procambarus machardyi Walls G1G2 LR lc LA (Caddo Chimney Crawfish) Procambarus mancus Hobbs G4 EN B1+2c MS and Walton (Lame Crayfish) Procambarus marthae Hobbs G3 VU B1+2c AL (Crisscross Crayfish) Procambarus medialis Hobbs G3 VU B1+2c NC (Tar River Crayfish) Hobbs G1 EN B1+2c FL ( Cave Crayfish) Procambarus morrisi Hobbs and G1 CR B1+2ae, C1+2b FL Franz (Putnum County Cave Crayfish) Procambarus natchitochae Penn G5 LR lc AR, LA, TX (Red River Crayfish) Procambarus nechesae Hobbs G2 VU B1+2c TX (Neches Crayfish) Hobbs G1G2 VU B1+2c, D1 TX (Blackbelted Crayfish) Procambarus nueces Hobbs and G1 CR B1+2c TX Hobbs (Nueces Crayfish) Procambarus okaloosae Hobbs G4 LR lc AL, FL (Okaloosa Crayfish) Hobbs and G1 VU B1+2c, D1 FL Means (Woodville Karst Cave Crayfish) Procambarus ouachitae Penn G5 LR lc AR, MS (Ouachita River Crayfish) SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 87

Table 1.—Continued. Species Global Rank Red List Distribution Procambarus paeninsulanus G5 LR lc AL, FL, GA (Faxon) (Peninsula Crayfish) Procambarus pallidus (Hobbs) G2G3 VU B1+2c, D1 FL (Pallid Cave Crayfish) Procambarus parasimulans Hobbs G4 LR lc AR and Robison (Bismark Burrowing Crayfish) Procambarus pearsei (Creaser) G4 LR lc NC, SC (Sandhills Crayfish) Procambarus penni Hobbs G3 VU B1+2c AL, LA, MS (Pearl Blackwater Crayfish) Procambarus pentastylus Walls G3 LR lc LA and Black (Calcasieu Creek Crayfish) Procambarus petersi Hobbs G3 VU B1+2c GA (Ogeechee Crayfish) (Hobbs) G2 VU B1+2c FL (Spotted Royal Crayfish) Procambarus planirostris Penn G4 LR lc LA, MS (Flatnose Crayfish) Procambarus plumimanus Hobbs G4 VU B1+2c NC and Walton (Croatan Crayfish) Procambarus pogum Fitzpatrick G1 VU B1+2c, D1 MS (Bearded Red Crayfish) Procambarus pubescens (Faxon) G4G5 LR lc GA, SC (Brushnose Crayfish) Procambarus pubischelae deficiens G5T3 LR lc GA Hobbs (Hookless Crayfish) Procambarus pubischelae G5T5 LR lc FL, GA pubishelae Hobbs (Brushpalm Crayfish) Procambarus pycnogonopodus G4G5 LR lc FL Hobbs (Stud Crayfish) Hobbs G4 LR lc FL, GA (Christmas Tree Crayfish) Procambarus raneyi Hobbs G4 LR lc GA, SC (Disjunct Crayfish) Procambarus rathbunae (Hobbs) G2 VU B1+2c FL (Comb Claw Crayfish) Procambarus regalis Hobbs and G2G3 VU B1+2c AR Robison (Regal Burrowing Crayfish) Hobbs G1 VU B1+2c, D1 AR (Irons Fork Burrowing Crayfish) Procambarus rogersi campestris G4T3 VU B1+2c, D1 FL Hobbs (Field Crayfish) Procambarus rogersi expletus G4T1 EN B1+2c FL Hobbs and Hart (Perfect Crayfish) Procambarus rogersi G4T2T3 VU B1+2c FL ochlocknensis Hobbs (Ochlockonee Crayfish) Procambarus rogersi rogersi G4T1T2 EN B1+2c FL (Hobbs) (Seepage Crayfish) 88 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

Table 1.—Continued. Species Global Rank Red List Distribution Procambarus seminolae Hobbs G5 LR lc FL, GA (Seminole Crayfish) Procambarus shermani Hobbs G4 LR lc AL, FL, LA, MS (Gulf Crayfish) Procambarus simulans (Faxon) G5 LR lc AR, LA, OK, TX (Southern Plains Crayfish) Procambarus spiculifer (LeConte) G5 LR lc AL, FL, GA, TN, SC (White Tubercled Crayfish) Procambarus steigmani Hobbs G1G2 CR B1+2c TX (Parkhill Prairie Crayfish) Procambarus suttkusi Hobbs G3G4 LR lc AL, FL (Choctawhatchee Crayfish) Procambarus talpoides Hobbs G5 LR lc FL, GA (Mole Crayfish) Hobbs G3 VU B1+2c AR, OK (Ouachita Mountain Crayfish) Procambarus texanus Hobbs G1 CR B1+2c TX (Bastrop Crayfish) Procambarus troglodytes G5 LR lc GA, SC (LeConte) (Eastern Red Swamp Crayfish) Procambarus truculentus Hobbs G3 LR lc GA (Bog Crayfish) Procambarus tulanei Penn (Giant G5 LR lc AR, LA Bearded Crayfish) Procambarus verrucosus Hobbs G4 LR lc AL, GA (Grainy Crayfish) Procambarus versutus (Hagen) G5 LR lc AL, FL, GA (Sly Crayfish) Procambarus viaeviridis (Faxon) G5 LR lc AL, AR, KY, LA, (Vernal Crayfish) MS, TN Procambarus vioscai paynei G5T4 LR lc AL, MS, TN Fitzpatrick (Payne’s Creek Crayfish) Procambarus vioscai vioscai G5T4 LR lc AR, LA Penn (Percy’s Creek Crayfish) Procambarus youngi Hobbs G2 EN B1+2c FL (Florida Longbeak Crayfish) Hobbs G5 LR lc AL, LA, (MD), MS, and Hobbs (Southern White TX, VA, (WV) River Crawfish) maclanei Hobbs G2 VU B1+2c FL (Spider Cave Crayfish) Troglocambarus sp. 1 (Orlando G1 VU B1+2c, D1 FL Spider Cave Crayfish) by Master (1991) as species in need of con- overexploitation (Allan & Flecker 1993; Richter servation attention. et al. 1997; Wilcove et al. 2000). The imperilment The causes of loss of aquatic species and of some crayfish taxa is due to limited natural population declines have been attributed to three range (e.g., one locality or one drainage system). major categories: (1) habitat fragmentation due The lack of recent distributional information is to loss, degradation, or alteration of habitat; (2) problematic; however, significant progress has chemical pollution from contaminants and been made mostly as a result of the publication disturbance from anthropogenic use; and (3) of numerous state field guides. Life history, introduction of nonindigenous organisms and ecological, and current distribution information SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 89

to conservation of biodiversity of native crayfish (Lodge et al. 2000; Clavero & Garcı´a-Berthou 2005). In North America, crayfishes are transported easily and can be inadvertently introduced into aquatic habitats when they are discarded as unused bait (Simon 2002). Such bait-bucket introductions have led to dramatic range extensions of several species, most notably Orconectes rusticus (Rusty Crayfish) and Procambarus clarkii (Red Swamp Crawfish). The Rusty Crayfish is native to the Greater Miami River in and Indiana, northern Kentucky, and Figure 1.—Distribution of states included in the the Maumee River drainage in Indiana, conservation status assessment of southeastern cray- northwestern Ohio, and extreme southeastern fishes. State inclusion is based on membership in the . The species has been introduced American Fisheries Society Southern Division. States across the upper midwestern United States in gray are included in the conservation status, while and Canada (Page 1985; Lodge et al. 2000) by those in white are excluded. expanding its range and displacing native crayfishes (Taylor & Redmer 1996). Possible are still lacking for about 60% of the North displacement mechanisms include faster indi- American fauna (Flinders & Magoulick 2005). vidual growth rates (Hill et al. 1993), differ- For crayfishes, land use change can affect habitat ential susceptibility to fish predation (DiDo- resources (Burskey & Simon 2010) by degrading nato & Lodge 1993), and hybridization (Perry habitat and homogenization of substrate surfac- et al. 2001). Imperiled crayfishes also have es. Loss of such habitat components through been affected by nonindigenous species. The dredging and channelization can drastically federally endangered Pacifastacus fortis has affect crayfish populations by making them been displaced over large portions of its native more susceptible to predation. Crayfish depend range by the nonindigenous P. leniusculus on instream cover including gravel and boulder (Erman et al. 1993). substrates, woody debris, and vegetation for Additional risk from non-indigenous cray- refuge from predators (Stein 1977; Burskey & fish is due to the escape of pets from the Simon 2010). These factors increase fish preda- aquarium and aquaculture trade (Simon tion by impounding lotic habitat, which affects 2002); however, increased prevention requires relative abundance of crayfish by increasing bait regulations that reduce nonindigenous concentrations of major predators on crayfish, crayfish spread (Holdrich 1999). States have such as black basses and sunfish, and altering either banned the sale of crayfish as bait or both the physical and chemical structure of created a list of prohibited species, restricting streams (Williams et al. 1993). the sale by bait dealers of nonindigenous Crayfish are among the most sensitive species including transport across state lines. aquatic organisms and exhibit specific differ- While no known North American cases of ences in tolerance to contaminants (Simon & overexploitation of crayfish have been docu- Morris 2009). Crayfish show extreme sensitivity mented, commercial harvest has the potential when exposed to pesticides and metals (Mayer of being a serious contributing factor when & Ellersieck 1986; Jarvinen & Ankley 1999; harvesting from wild populations. Species Besser et al. 2006) and data show significant most vulnerable to over-harvesting are those variability among genera, species, and life stages with small native ranges, long life spans prior (Berrill et al. 1985; Peake et al. 2004; Wigginton to adult maturation, and low reproductive & Birge 2007; Simon & Morris 2009). These potential. Gulf Coast populations are perhaps observations suggest that crayfish may be the most vulnerable to over-harvesting if not important indicators of habitat degradation managed properly because of exposure to due to pollutants (Burskey & Simon 2010). hurricanes. However, the majority of states The introduction of nonindigenous organ- that possess a highly diverse crayfish fauna isms may be among the most serious threats with high levels of endemism lack any 90 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE protective measures or adequate funding genic influences are needed to protect crayfish structures to monitor the status of their re- biodiversity in North America. spective state faunas. In conclusion, more than half of the southern ACKNOWLEDGEMENTS crayfish species are stable and slightly less are The opinions expressed in this paper are in need of critical management intervention to those of the author and do not reflect those of protect remaining populations. More effort is the federal or state government. Determination needed to document distribution and habitat of imperilment does not constitute endorsement needs. Human encroachment causing habitat for listing using the ESA. Although this fragmentation, followed by contaminants and manuscript may have been funded wholly or introductions of nonindigenous species, are in part by the Federal government, no endorse- among the most frequent reasons for imperil- ment by any agency is implied. Reference to ment. Wise land use development, protection of trade names does not imply endorsement of riparian corridors, and control of anthropo- commercial products by the US government.

APPENDIX 1. THE INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL RESOURCES (IUCN) RED LIST CRITERIA ASSESSMENT PROCESS FOR CONSERVATION STATUS DETERMINATIONS. CRITICALLY ENDANGERED (CR)

A) A taxon is Critically Endangered when it is facing d) number of locations or subpopulations an extremely high risk of extinction in the wild in the e) number of mature individuals immediate future, as defined by any of the following criteria (A to E): 3) Extreme fluctuations in any of the following: 1) An observed, estimated, inferred or suspected a) extent of occurrence b) area of occupancy reduction of at least 80% over the last 10 years or three generations, whichever is the longer, based c) number of locations or subpopulations on (and specifying) any of the following: d) number of mature individuals C) Population estimated to number less than 250 a) direct observation mature individuals and either: b) an index of abundance appropriate for the taxon c) a decline in area of occupancy, extent of 1) An estimated continuing decline of at least 25% occurrence and/or quality of habitat within three years or one generation, whichever is d) actual or potential levels of exploitation longer or e) the effects of introduced taxa, hybridization, 2) A continuing decline, observed, projected, or pathogens, pollutants, competitors or parasites. inferred, in numbers of mature individuals and population structure in the form of either: 2) A reduction of at least 80%, projected or suspected to be met within the next 10 years or a) severely fragmented (i.e. no subpopulation esti- three generations, whichever is the longer, based mated to contain more than 50 mature individuals) on (and specifying) any of (b), (c), (d) or (e) above. b) all individuals are in a single subpopulation B) Extent of occurrence estimated to be less than D) Population estimated to number less than 50 100 km2 or area of occupancy estimated to be less mature individuals. than 10 km2, and estimates indicating any two of the following: E) Quantitative analysis showing the probability of extinction in the wild is at least 50% within 10 years 1) Severely fragmented or known to exist at only a or three generations, whichever is the longer. single location. 2) Continuing decline, observed, inferred or ENDANGERED (EN) projected, in any of the following: A taxon is Endangered when it is not Critically a) extent of occurrence Endangered but is facing a very high risk of b) area of occupancy extinction in the wild in the near future, as defined c) area, extent and/or quality of habitat by any of the following criteria (A to E): SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 91

A) Population reduction in the form of either of the E) Quantitative analysis showing the probability of following: extinction in the wild is at least 20% within 20 years or five generations, whichever is the longer. 1) An observed, estimated, inferred or suspected reduction of at least 50% over the last 10 years or VULNERABLE (VU) three generations, whichever is the longer, based on (and specifying) any of the following: A taxon is Vulnerable when it is not Critically Endangered or Endangered but is facing a high risk a) direct observation of extinction in the wild in the medium-term future, b) an index of abundance appropriate for the as defined by any of the following criteria (A to E): taxon c) a decline in area of occupancy, extent of A) Population reduction in the form of either of the occurrence and/or quality of habitat following: d) actual or potential levels of exploitation 1) An observed, estimated, inferred or suspected e) the effects of introduced taxa, hybridisation, reduction of at least 20% over the last 10 years or pathogens, pollutants, competitors or parasites. three generations, whichever is the longer, based on (and specifying) any of the following: 2) A reduction of at least 50%, projected or suspected to be met within the next 10 years or a) direct observation three generations, whichever is the longer, based on b) an index of abundance appropriate for the taxon (and specifying) any of (b), (c), (d), or (e) above. c) a decline in area of occupancy, extent of occurrence and/or quality of habitat B) Extent of occurrence estimated to be less than d) actual or potential levels of exploitation 5000 km2 or area of occupancy estimated to be less 2 e) the effects of introduced taxa, hybridisation, than 500 km , and estimates indicating any two of pathogens, pollutants, competitors or parasites. the following: 2) A reduction of at least 20%, projected or 1) Severely fragmented or known to exist at no suspected to be met within the next ten years or more than five locations. three generations, whichever is the longer, based 2) Continuing decline, inferred, observed or on (and specifying) any of (b), (c), (d) or (e) above. projected, in any of the following: B) Extent of occurrence estimated to be less than a) extent of occurrence 20,000 km2 or area of occupancy estimated to be less b) area of occupancy than 2000 km2, and estimates indicating any two of c) area, extent and/or quality of habitat the following: d) number of locations or subpopulations 1) Severely fragmented or known to exist at no e) number of mature individuals more than ten locations. 3) Extreme fluctuations in any of the following: 2) Continuing decline, inferred, observed or projected, in any of the following: a) extent of occurrence a) extent of occurrence b) area of occupancy b) area of occupancy c) number of locations or subpopulations c) area, extent and/or quality of habitat d) number of mature individuals d) number of locations or subpopulations C) Population estimated to number less than 2500 e) number of mature individuals mature individuals and either: 3) Extreme fluctuations in any of the following: 1) An estimated continuing decline of at least 20% a) extent of occurrence within five years or two generations, whichever is b) area of occupancy longer, or c) number of locations or subpopulations 2) A continuing decline, observed, projected, or d) number of mature individuals inferred, in numbers of mature individuals and population structure in the form of either: C) Population estimated to number less than 10,000 mature individuals and either: a) severely fragmented (i.e. no subpopulation estimated to contain more than 250 mature 1) An estimated continuing decline of at least 10% individuals) within 10 years or three generations, whichever is b) all individuals are in a single subpopulation. longer, or 2) A continuing decline, observed, projected, or D) Population estimated to number less than 250 inferred, in numbers of mature individuals and mature individuals. population structure in the form of either: 92 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

a) severely fragmented (i.e. no subpopulation Burr, B.M. & H.H. Hobbs, Jr. 1984. Additions to the estimated to contain more than 1000 mature crayfish fauna of Kentucky, with new locality individuals) records for Cambarellus shufeldtii. Transactions of b) all individuals are in a single subpopulation the Kentucky Academy of Science 45:14–18. Burskey, J.L. & T.P. Simon. 2010. Reach- and D) Population very small or restricted in the form of watershed-scale associations of crayfish within an either of the following: area of varying agricultural impact in west-central 1) Population estimated to number less than 1000 Indiana. Southeastern Naturalist 9(Special Issue mature individuals. 3):199–216. 2) Population is characterized by an acute Clavero, M. & E. Garcı´a-Berthou. 2005. Invasive restriction in its area of occupancy (typically less species are a leading cause of . than 100 km2) or in the number of locations Trends in Ecology and Evolution 20:110. (typically less than five). Such a taxon would thus Cooper, J.E. 2002. North Carolina crayfishes (Dec- be prone to the effects of human activities (or apoda: Cambaridae): notes on distribution, tax- onomy, life history, and habitat. Journal of the stochastic events whose impact is increased by North Carolina Academy of Science 118:167–180. human activities) within a very short period of Cooper, J.E. & A.L. Braswell. 1995. Observations on time in an unforeseeable future, and is thus North Carolina crayfishes (Cambaridae). Brim- capable of becoming Critically Endangered or leyana 22:87–132. even Extinct in a very short period. Cooper, J.E., A.L. Braswell & C. McGrath. 1998. E) Quantitative analysis showing the probability of Noteworthy distributional records for crayfishes extinction in the wild is at least 10% within 100 years. (Cambaridae) in North Carolina. Journal of the Elisha Mitchell Scientific Society 114(1):1–10. LITERATURE CITED Crandall, K.A., D.J. Harris & J.W. Fetzner, Jr. 2000. Adams, S.B. 2008. Mississippi Crayfish. Available at: The monophyletic origin of freshwater crayfish http://maps.fs.fed.us/crayfish/crayfish_general_info. estimated from nuclear and mitochondrial DNA jsp. (Accessed: March 2009). sequences. Proceedings of the Royal Society of Albaugh, D.W. & J.B. Black. 1973. A new craw- London B. 267:1679–1686. fish of the genus Cambarellus from Texas, with Creaser, E.P. & A.I. Ortenburger. 1933. The decapod new Texas distributional records for the genus of Oklahoma. Publications of the (Decapoda, Astacidae). Southwestern Naturalist University of Oklahoma Biological Survey 5:14–47. 1:177–185. Deacon, J.E., G.K. Kobetich, J.D. Williams, S. Allan, J.D. & A.S. Flecker. 1993. Biodiversity Contreras, other members of the AFS Endangered conservation in running waters. Bioscience 4: Species Committee. 1979. Fishes of North Amer- 32–43. ica endangered, threatened, or of special concern. Fisheries 4(2):29–44. Berrill, M., L. Hollett, A. Margosian & J. Hudson. Deyrup, M., & R. Franz (Eds.). 1994. Rare and 1985. Variation in tolerance to low environ- Endangered Biota of Florida, Vol. IV. Inverte- mental pH by the crayfish Orconectes rusticus, brates. Gainesville (FL): University Press of O. propinquus and Cambarus robustus. Canadian Florida. Journal of Zoology 63:2586–2589. DiDonato, G.T. & D.M. Lodge. 1993. Species Besser, J.M., W.G. Brumbaugh, T.W. May & C.J. replacements among Orconectes crayfishes in Schmitt. 2006. Biomonitoring of lead, zinc, and northern lakes: the role of predation cadmium in streams draining lead-mining and by fish. Canadian Journal of Fisheries and non-mining areas, southeast , USA. Aquatic Sciences 50:1484–1488. Environmental Monitoring and Assessment 129: Dunlap, P.M. Jr. 1951. Taxonomic characteristics of 227–241. the decapod crustaceans of the subfamily Cam- Bouchard, R.W. 1972. A contribution to the barinae in Oklahoma with descriptions of two new knowledge of Tennessee crayfish. Ph.D. disserta- species and two keys to species. M.S. Thesis, tion, University of Tennessee, Knoxville. Oklahoma Agricultural and Mechanical College, Bouchard, R.W. 1976. Crayfishes and shrimps. Stillwater. Pp. 13–20. In Endangered and Threatened Plants Erman, D.C., T. Light & C. Myrick. 1993. Survey of and of Alabama (H. Boschung, Ed.), the status of the Shasta crayfish (Pacifastacus Bulletin, Alabama Museum of Natural History 2. fortis) in northeastern California (1991 study Bouchard, R.W. & H.W. Robison. 1980. An year). Final report to California of Fish and inventory of the decapod crustaceans (crayfishes Game, Sacramento. 56 pp. and shrimps) of Arkansas with a discussion of Eversole, A.G. 1995. Distribution of three rare their habitats. Arkansas Academy of Science crayfish species in South Carolina. Freshwater Proceedings 34:22–30. Crayfish 8:113–120. SIMON—CONSERVATION STATUS OF FRESHWATER CRAYFISH 93

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