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An Annotated Checklist and Preliminary Designation of Drainage Distributions of the Crayfishes of Alabama
Guenter A. Schuster, Christopher A. Taylor, and John Johansen

Southeastern Naturalist, Volume 7, Number 3 (2008): 493–504

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2008 SOUTHEASTERN NATURALIST 7(3):493–504 An Annotated Checklist and Preliminary Designation of Drainage Distributions of the Crayfishes of Alabama Guenter A. Schuster1,*, Christopher A. Taylor2, and John Johansen3 Abstract - As a first step toward elucidating the current status of Alabama’s crayfish fauna, museums known to have significant crayfish collections were queried for their holdings from that state. A total of 4649 records of Alabama crayfishes were obtained from seven museums. Three-hundred thirty of the records did not have suitable information for geo-referencing. The largest holdings were in the National Museum of Natural History (2544 records). Specimen identifications were verified, and all records were geo-referenced. Geographic distribution strongly favored upland drainages in the northern and central portions of the state. The Tennessee River drainage was the most collected drainage (1018 records, 23.6% of records). A total of 85 species of crayfishes are reported for the state of Alabama; only Orconectes virilis (Virile Crayfish) is deemed to be non-native. Even with the extensive collection of crayfish records in museums, there is a need for crayfish inventory work in Alabama. A total of 3107 records (76.3%) were collected prior to 1987, and 1379 (33.8%) were collected prior to 1970. In addition, there is a paucity of records from the coastal drainages of SE Alabama. There is also an underrepresentation of burrowing crayfishes, especially those classified as either primary or secondary burrowers. Only 212 (4.9%) of all records are of burrowing species. Lastly, a limited amount of fieldwork in Alabama has documented the presence of a species previously unreported for the state. Introduction As pointed out by Schuster and Taylor (2004) and others (Boschung and Mayden 2004, Crandall 1997, Lydeard and Mayden 1995), the aquatic fauna of Alabama is extremely diverse, and the total number of taxa may exceed that of any other state or province in North America. Recent distributional and ecological information has been compiled for major aquatic groups such as fishes (Boschung and Mayden 2004, Mettee et al. 1996), caddisfl ies (Harris et al. 1991), and unionid mussels (Williams et al., in press). However, such information for crayfishes is lacking. Alabama has a relatively mild, stable climate, was spared from recent glacial advances, and has a diverse geology and topology ranging from high-gradient Appalachian streams to coastal fl oodplains, factors that have undoubtedly contributed to the State’s high biodiversity. 1Department of Biological Sciences, Eastern Kentucky University, 521 Lancaster Avenue, Richmond, KY 40475. 2Division of Biodiversity and Ecological Entomology, Illinois Natural History Survey, 1816 South Oak, Champaign, IL 61820. 3Tulane University Museum of Natural History, 3705 Main Street, Building A-3, New Orleans, LA 70075. *Corresponding author - Guenter.Schuster@eku.edu. 494 Southeastern Naturalist Vol.7, No. 3 Ortmann (1905, 1931) and Hobbs (1969) speculated that the Cumberland Plateau is the center of radiation for the crayfish genera Cambarus and Orconectes, both of which are major components (combined total of >50%) of the Alabama crayfish fauna. Whether or not that speculation is correct, the Cumberland Plateau is the second largest physiographic province in the state (Boschung and Mayden 2004), and its hydrology (containing both the Tennessee and Black Warrior river drainages) and highly dissected topology strongly infl uence aquatic faunal components for most of the northern section of the state. Certainly another important aspect of the evolution of aquatic organisms in Alabama is the diversity of river systems in the state. Three major basins drain Alabama: the Ohio River, Mobile Bay, and the Apalachicola River. In addition, there are four relatively isolated Coastal Plain drainages: Pascagoula River, Perdido Bay, Pensacola Bay, and Choctawhatchee Bay (Boschung and Mayden 2004). The long-term isolation (until the recent connection of the Tombigbee and Tennessee river systems) of the major basins and the coastal drainages, undoubtedly resulted in divergent evolution among populations leading to speciation. To address the deficiency in baseline distributional information for Alabama crayfishes, a study was undertaken with three main objectives. The first was to determine the extent of Alabama crayfish holdings in various museums in the United States. Secondly, museums with significant holdings were visited and, where appropriate, specimens were identified or had their identifications confirmed or corrected. Invalid taxonomic names were also corrected. Thirdly, all museum specimens with adequate locality information were geo-referenced and compiled into a single database. In addition to museum work, a limited amount of field collecting was conducted between 2005 and 2007 using standard field techniques. Methods Museums known to have significant crayfish collections were queried for their holdings of Alabama crayfishes. Electronic records from those museums containing more than 50 records were then integrated into a single Microsoft Excel database. A record is defined as a collection of one species of crayfish from a single location. With two exceptions, museums with more than 50 Alabama records that had not been previously examined by the authors were then visted. A concentrated effort was made to identify previously unidentified Alabama specimens and confirm identifications of specimens that did not fit known distributions. Once confirmed, all records in the Microsoft Excel database were geo-referenced using GEOLocate (V 2.13, Tulane University), and entered into a Microsoft Access database in a decimal degree format. All geo-referenced records were then visually confirmed on county road or topographic maps or electronic maps in the software package TopoUSA (DeLorme). 2008 G.A. Schuster, C.A. Taylor, and J. Johansen 495 Institutional acronyms are as follows: Auburn University of Natural History (Auburn); University of Alabama Museum of Natural History (UA); Eastern Kentucky University Branley A. Branson Museum of Zoology (EKU); Museum of Comparative Zoology at Harvard University (MCZ); Illinois Natural History Survey Crustacean Collection (INHS); National Museum of Natural History, Smithsonian Institution (USNM); North Carolina State Museum of Natural History (NCSM); Tulane University Museum of Natural History (TU). Figure 1. Distribution of known crayfish records for Alabama (n = 4319). 496 Southeastern Naturalist Vol.7, No. 3 Results A total of 4649 records of Alabama crayfishes were obtained from seven museums and recent fieldwork (Fig. 1). Of these, 330 (7.1%) records did not have adequate information for geo-referencing. In addition, 249 records did not contain date of collection. The largest holdings of records were found at USNM with 2544 records (54.8%), TU with 804 (17.3%), and UA with 487 (10.5%). The temporal distribution of Alabama crayfish collections was very uneven (Fig. 2). Sixty-three percent of all records were collected during the 1960s and 1970s, with 17.1% of collections made during the 1990s. Only 9.0% of collections were made prior to 1960. The geographical distribution of Alabama records strongly favored upland drainages in the northern and central portions of the state. The Tennessee River drainage was the most represented drainage having 1018 records (23.6%) (Fig. 3). The Tombigbee (428 records), Black Warrior (539), Cahaba (217), Coosa (462), Tallapoosa (480), and Alabama (308) drainages combined accounted for 56.3% of all records. The coastal drainages of southeastern Alabama, including the Choctawhatchee, Conecuh, Escatawpa, Perdido, and Yellow, accounted for only 9.3% of all records (Fig. 4). In addition to museum work, recent fieldwork in Alabama has resulted in the addition of one new species. Orconectes durelli (Saddle Crayfish) was collected from Second Creek, Lauderdale County, on October 22, 2005. The compilation and examination of museum records and fieldwork revealed 85 described crayfish taxa known to occur in Alabama (Appendix 1). Of these, only O. virilis is deemed to be non-native. Taxa are presented in alphabetical order by subfamily, genus and species with common name, known range by drainage basin, and habitat category for each species. Figure 2. Number of geo-referenced Alabama Crayfish records by decade of collections (n = 4070). Number does not include records with incomplete locality data or missing date of collection. 2008 G.A. Schuster, C.A. Taylor, and J. Johansen 497 Figure 3. Number of geo-referenced Alabama Crayfish records by major river drainage (AL = Alabama, BW = Black Warrior, CH = Chattahoochee, CO = Coosa, MO = Mobile, TA = Tallapoosa, TN = Tennessee, and TO = Tombigbee) (n = 4319). Figure 4. Number of geo-referenced Alabama Crayfish records by minor river drainage (BW = Blackwater, CA = Cahaba, CP = Chipola, CT = Choctawhatchee, CO = Conecuh, ES = Escatawpa, PE = Perdido, and YE = Yellow) (n = 4319). 498 Southeastern Naturalist Vol.7, No. 3 All crayfish living in Alabama belong to the family Cambaridae, which contains two subfamilies, Cambarellinae and Cambarinae. Representatives of both subfamilies reside in Alabama. Cambarellinae has a single genus, Cambarellus, of which three species are known from Alabama. The remaining 82 species belong to Cambarinae. Six genera of Cambarinae are present in Alabama: Cambarus, Fallicambarus, Faxonella, Hobbseus, Orconectes, and Procambarus. Ten subgenera and 27 species of Cambarus occur in Alabama, making it one of the largest crayfish genera in the state. All five species of Fallicambarus in Alabama belong to the subgenus Creaserinus. Faxonella is represented by a single species, as is the genus Hobbseus. Orconectes, also one of the largest crayfish genera in Alabama, is represented in the state by seven subgenera and 19 species. Procambarus is the most diverse genus in Alabama, with six subgenera and 29 species found within its boundaries. Discussion History of astacology in Alabama Until 1870, the crayfishes of Alabama went almost unnoticed. Hagen (1870) provided the first checklist of crayfishes from Alabama and remarked that “nearly the whole state remains unexplored.” His list included 5 species: Cambarus advena (= Procambarus hagenianus hagenianus [Southeastern Prairie Crayfish]), C. immunis (= Orconectes immunis (Hagen) [Calico Crayfish]), C. acutus (= P. acutus [White River Crayfish]), C. lecontei (= P. lecontei [Mobile Crayfish]), and C. versutus (= P. versutus [Sly Crayfish]). Two of these species, P. lecontei and P. versutus, were the first Alabama crayfishes described from Alabama specimens. The specimens of both were collected by Louis Agassiz on a trip to Mobile in the early 1850s. A century after Hagen’s work, Bouchard (1976) provided the first attempt at a comprehensive checklist of Alabama crayfishes. He listed 58 species for the state and estimated that approximately 75 will be known when the fauna has been completely surveyed. In addition, he was the first to attempt to determine the conservation status for the crayfishes of Alabama. He listed 14 species in need of protection. One was listed as threatened, while the remaining 13 were listed as species of special concern. Hobbs’ (1989) illustrated checklist of American crayfishes listed 72 species and subspecies from Alabama, with two species being listed as questionable. One of these, Fallicambarus hedgpethi, has been synonymyzed with Fallicambarus fodiens (Digger Crayfish), while the other, Procambarus bivittatus (Ribbon Crayfish), is now considered to be a member of the state’s fauna. In a conservation review of North American crayfish, Taylor et al. (1996) listed 78 taxa as occurring in Alabama. Crandall (1997) erroneously cited Taylor et al. (1996) in reporting 76 taxa as occurring in Alabama. Most recently, Taylor et al. (2007) presented an update on the conservation of North American crayfish. They listed 86 taxa, including the subspecies Cambarus 2008 G.A. Schuster, C.A. Taylor, and J. Johansen 499 bartonii bartonii (Farbricius), in their list for Alabama, which we exclude here. The marked increase in the number of taxa reported from Alabama between 1976 and 2007 was largely due to taxonomic revisionary work and new species descriptions. Current status The records reported herein do not reflect an up-to-date assessment of the Alabama crayfish fauna. With a total of 3107 (76.3.0%) records collected prior to 1987 and 1379 (33.8%) predating 1970, the ability to determine current population trends and distributions is severely hampered. More alarming is the under-representation of burrowing crayfishes (Hobbs 1981, 1989). Of the 4649 records, only 212 (4.9%) are of species that spend a significant portion of their lives in subterranean burrows. This under-sampling prevents an assessment of both historical and current population levels and distributions. Our preliminary examinations reveal that at least one manuscript name may be applicable to a previously described species (Orconectes jonesi [Sucarnoochie River Crayfish]); however, others may represent undescribed species. There remains a great need for additional taxonomic evaluation of museum collections and field collecting. One noteworthy example is the presence of Procambarus zonangulus (Southern White River Crayfish) in Alabama. Procambarus zonangulus is a member of the P. acutus species group and has been considered by some to be the form present in most of the Gulf Coastal Plain, including Alabama (Huner 2002). We did not find specimens assignable to P. zonangulus during our visits to museums; however, we follow Huner (2002) in including it in our species list. Finally, in addition to specimen-based taxonomic uncertainties, recent molecular work has suggested the presence of at least two undescribed taxa in the genus Cambarus from karst regions of northern Alabama (Buhay et al. 2007). Future molecular analyses will surely reveal the presence of unique species in other regions of the state. Future efforts in Alabama should include: 1) taxonomic revisions of numerous crayfish groups and descriptions of new taxa; 2) additional fieldwork in under-sampled drainages, especially in coastal drainages of southeastern Alabama; 3) additional fieldwork in regions where the last known collections predate 1987; and 4) statewide collections of primary and secondary burrowing species. Acknowledgments We gratefully acknowledge Jim McHugh and Steve Rider of the Alabama Division of Wildlife and Freshwater Fisheries and the State Wildlife Grants program for their financial support of crayfish work in Alabama. We also acknowledge Hank Bark and Nelson Rios of Tulane University Museum of Natural History for their assistance with the curation of the TU Collection and the administration of the grant funds. We also thank Karen Reed and Rafael Lemaitre (UNSM), Phil Harris and Bernard Kuhadja (UA), and Michael Gangloff (Auburn) for access to the 500 Southeastern Naturalist Vol.7, No. 3 respective collections under their care and assistance during our visits. Thanks also to John Cooper (NCSM) and Ardis Johnston (MCZ) for providing collection records. We would also like to thank Courtney Graydon, Emily Hartfield, and Dan Jones for their help in the field. Finally, we thank the following for providing specimens: Paul Freeman (The Nature Conservancy), Jim Godwin (Alabama Heritage Program), Courtney Graydon (Alabama State Lands), Dennis DeVries and Emily Hartfield (Auburn), Stuart McGregor and Pat O’Neil (Geologic Survey of Alabama), and Steve Rider (Alabama Department of Wildlife and Freshwater Fisheries). Literature Cited Boschung, H.T., and R.L. Mayden. 2004. Fishes of Alabama. Smithsonian Institution, Washington, DC. Bouchard, R.W. 1976. Crayfishes and Shrimps. Pp.13–21, In H. Boschung (Ed.). Endangered and Threatened Plants and Animals of Alabama. Bulletin of the Alabama Museum of Natural History 2. Buhay, J.E., G. Moni, N. Mann, and K.A. Crandall. 2007. Molecular taxonomy in the dark: Evolutionary history, phylogeography, and diversity of cave crayfish in the subgenus Aviticambarus, genus Cambarus. Molecular Phylogenetics and Evolution 42:435–448. Crandall, K.A. 1997. The crayfish component to an endangered aquatic ecosystem of the southeastern United States. Freshwater Crayfish 11:83–86. Hagen, H.A. 1870. Monograph of the North American Astacidae. Illustrated Catalogue of the Museum of Comparative Zoology No. 3. 109 pages. Harris, S.C., P.E. O’Neil, and P.K. Lago. 1991. Caddisfl ies of Alabama. Geologic Survey of Alabama Bulletin 142:1–442. Hobbs, H.H., Jr. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pp. 92–178, In P.C. Holt, R.L. Hoffman, and C.W. Hart, Jr. (Eds.). The Distributional History of the Biota of the Southern Appalachians, Part I: Invertebrates. Virginia Polytechnic Institute, Research Division, Blacksburg, VA. Monograph 1. Hobbs, H.H., Jr. 1981. The crayfishes of Georgia. Smithsonian Contributions to Zoology 318:1–549. Hobbs, H.H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology 480:1–236. Huner, J.V. 2002. Procambarus. Pp. 541–584, In D.M. Holdich (Ed.). Biology of Freshwater Crayfish, Blackwell Science, Oxford, UK. Lydeard, C., and R.L. Mayden. 1995. A diverse and endangered aquatic ecosystem of the southeast United States. Conservation Biology 9(4):800–805. Mettee, M.F., Jr., P.E. O’Neil, and J.E. Pierson. 1996. Fishes of Alabama and the Mobile Basin. Oxmore House, Inc., Birmingham, AL. Ortmann, A.E. 1905. The mutual affinities of the species of the Genus Cambarus, and their dispersal over the United States. Proceedings of the American Philosophical Society 44:435–442. Ortmann, A.E. 1931. Crawfishes of the southern Appalachians and the Cumberland Plateau. Annals of the Carnegie Museum 20:61–160. Schuster, G.A., and C.A. Taylor. 2004. Report on the crayfishes of Alabama: Literature review museum database review, and species list with abbreviated annotations and proposed conservation statuses. Illinois Natural History Survey, Center for Biodiversity Technical Report 2004 (12). 47 pp. 2008 G.A. Schuster, C.A. Taylor, and J. Johansen 501 Taylor, C.A., M.L. Warren, Jr., J.F. Fitzpatrick, Jr., H.H. Hobbs III, R.F. Jezerinac, W.L. Pfl ieger, and H.W. Robison. 1996. Conservation status of crayfishes of the United States and Canada. Fisheries 21:25–38. Taylor, C.A., G.A. Schuster, J.E. Cooper, R.J. DiStefano, A.G. Eversole, P. Hamr, H.H. Hobbs III, H.W. Robison, C.E. Skelton, and R.R. Thoma. 2007. A reassessment of the conservation status of crayfishes of the United States and Canada after 10+ years of increased awareness. Fisheries 32:372–389. Williams, J.D., A.E. Bogan, and J.T. Garner. In press. Freshwater Mussels of Alabama and the Mobile Basin in Georgia, Mississippi, and Tennessee. University of Alabama Press, Tuscaloosa, AL. 502 Southeastern Naturalist Vol.7, No. 3 Appendix 1. List of Alabama crayfish species alphabetically by subfamily, Genus and species with common names, habitat (C = caves, Le = lentic, Lo = lotic, PB = primary burrower, and SB = secondary burrower), and known range by drainage basin (AL = Alabama; BW = Black Warrior; CA = Cahaba; CH = Chattahoochee; CO = Coosa; CT = Choctawhatchee; ES = Escambia: Blackwater, Conecuh, and Yellow; MO = Mobile Bay: Mobile, Tensaw, and direct tributaries; PA = Pascagoula: Escatawpa; PE = Perdido; TA = Tallapoosa; TN = Tennessee; and TO = Tombigbee); UN=Uncertain range or species under taxonomic revision. Species Common name Habitats Drainages Cambarellus diminutus Hobbs Least Crayfish Le MO, PA C. lesliei Fitzpatrick & Laning Angular Dwarf Crawfish Le AL ,MO, TO C. shufeldtii (Faxon) Cajun Dwarf Crayfish Le MO Cambarus acanthura Hobbs Thornytail Crayfish PB AL, BW, CA, CH, CO, ES, PA, PE, TO, TN C. bartonii cavatus Hay Appalachian Brook Crayfish Lo CH,CO C. coosae Hobbs Coosa Crayfish Lo CA,CO C. cracens Bouchard & Hobbs Slenderclaw Crayfish Lo TN C. diogenes Girard Devil Crayfish PB,SB AL, BW, CO, CT, ES, MO, TA, TN, TO C. distans Rhoades Boxclaw Crayfish Lo TN C. englishi Hobbs & Hall Tallapoosa Crayfish Lo TA C. girardianus Faxon Tanback Crayfish Lo CA, CO, TN C. graysoni Faxon Twospot Crayfish Lo TN C. halli Hobbs Slackwater Crayfish Lo TA C. hamulatus (Cope) Prickly Cave Crayfish C BW, TN C. howardi Hobbs & Hall Chattahoochee Crayfish Lo CH C. jonesi Hobbs & Barr Alabama Cave Crayfish C TN C. latimanus (Le Conte) Variable Crayfish Lo AL, BW, CA, CH, CO, CT, ES, TA, TN, TO C. longirostris Faxon Longnose Crayfish Lo CO, TN C. ludovicianus Faxon Painted Devil Crayfish PB MO C. manningi Hobbs Greensaddle Crayfish Lo CO C. miltus Fitzpatrick Rusty Grave Digger PB MO, PE C. obstipus Hall Sloped Crayfish Lo BW C. parvoculus Hobbs & Shoup Mountain Midget Crayfish Lo TN C. polychromatus Thoma et al. Paintedh& Mudbug PB, SB CT C. rusticiformis Rhoades Depression Crayfish Lo TN C. scotti Hobbs Chattooga River Crayfish Lo CO C. striatus Hay Ambiguous Crayfish Lo, PB, SB AL, BW, CA, CH, CO, CT, ES, TA, TO, TN C. tenebrosus Hay Cavespring Crayfish C, Lo TN C. unestami Hobbs & Hall Blackbarred Crayfish Lo TN C. veitchorum Cooper & Cooper White Spring Cave Crayfish C TN Fallicambarus burrisi Fitzpatrick Burrowing Bog Crayfish PB MO, PA F. byersi (Hobbs) Lavender Burrowing Crayfish PB ES, PA, PE F. danielae Hobbs Speckled Burrowing Crayfish PB MO, PA 2008 G.A. Schuster, C.A. Taylor, and J. Johansen 503 Species Common name Habitats Drainages F. fodiens (Cottle) Digger Crayfish PB, SB AL, BW, CA, CH, ES, MO, PE, TA, TN, TO F. oryktes (Penn & Marlow) Flatwoods Digger PB MO Faxonella clypeata (Hay) Ditch Fencing Crayfish Le AL, CH, CT, ES, MO, PA, TA, TO Hobbseus prominens (Hobbs) Prominence Riverlet Crayfish Le AL, BW, CA, TO Orconectes alabamensis (Faxon) Alabama Crayfish Lo TN O. australis australis (Rhoades) Southern Cave Crayfish C TN O. chickasawae Cooper & Hobbs Chickasaw Crayfish Lo UN O. compressus (Faxon) Slender Crayfish Lo TN O. cooperi Cooper & Hobbs Flint River Crayfish Lo TN O. durelli Bouchard & Bouchard Saddle Crayfish Lo TN O. erichsonianus (Faxon) Reticulate Crayfish Lo BW, CA, CO, TA, TN O. forceps (Faxon) Surgeon Crayfish Lo TN O. holti Cooper & Hobbs Bimaculate Crayfish Lo UN O. jonesi Fitzpatrick Sucarnoochee River Crayfish Lo UN O. lancifer (Hagen) Shrimp Crayfish Le AL O. mirus (Ortmann) Wonderful Crayfish Lo TN O. perfectus Walls Complete Crayfish Lo AL, BW, TA, TO O. placidus (Hagen) Bigclaw Crayfish Lo TN O. putnami (Faxon) Phallic Crayfish Lo TN O. sheltae Cooper & Cooper Shelta Cave Crayfish C TN O. spinosus (Bundy) Coosa River Spiny Crayfish Lo BW, TN O. validus (Faxon) Powerful Crayfish Lo BW, TN, TO O. virilis Hagen Virile Crayfish Lo BW, CA, CO, TA, TN Procambarus acutissimus (Girard) Sharpnose Crayfish Le, Lo AL, BW, CA, CO, CT, TA, TN, TO P. acutus (Girard) White River Crawfish Le, Lo AL, BW, CO, CT, ES, MO, PA, TN, TO P. bivittatus Hobbs Ribbon Crayfish Le, Lo AL, CT, MO, TO P. capillatus Hobbs Capillaceous Crayfish Le, SB ES P. clarkii (Girard) Red Swamp Crawfish Le, Lo AL, BW, CA, CO, MO, PA, TA, TN, TO P. clemmeri Hobbs Cockscomb Crayfish Lo PA, TO P. escambiensis Hobbs Escambia Crayfish Le, Lo, SB ES, PE P. evermanni (Faxon) Panhandle Crayfish Lo AL, MO, PE P. hagenianus hagenianus (Faxon) Southeastern Prairie Crayfish PB TO P. hayi (Faxon) Straightedge Crayfish Le, Lo TO P. hubbelli (Hobbs) Jackknife Crayfish Le, SB ES P. hybus Hobbs & Walton Smoothnose Crayfish Le, SB BW, CA, TO P. lagniappe Black Lagniappe Crayfish Lo TO 504 Southeastern Naturalist Vol.7, No. 3 Species Common name Habitats Drainages Procambarus lecontei (Hagen) Mobile Crayfish Lo MO, PA, TO P. lewisi Hobbs & Walton Spur Crayfish Le, Lo AL, CH, ES, TA P. lophotus Hobbs & Walton Mane Crayfish Le, Lo AL, BW, CO, CT, ES, TA, TN, TO P. marthae Hobbs Crisscross Crayfish Le, Lo AL, BW, CA P. okaloosae Hobbs Okaloosa Crayfish Le, Lo ES P. paeninsulanus (Faxon) Peninsula Crayfish Le, Lo CH P. pecki Hobbs Phantom Cave Crayfish C TN P. penni Hobbs Pearl Blackwater Crayfish Lo AL, PA, TO P. shermani Hobbs Gulf Crayfish Le, Lo AL, MO, PA, TO P. spiculifer (Le Conte) White Tubercled Crayfish Lo AL, BW, CA, CH, CO, CT, ES, MO, PA, PE, TA, TO P. suttkusi Hobbs Choctawhatchee Crayfish Lo CT P. verrucosus Hobbs Grainy Crayfish Lo AL, CH, ES, TA P. versutus (Hagen) Sly Crayfish Lo AL, BW, CA, CH, CO, CT, ES, MO, PA, PE, TA, TO P. viaeviridis (Faxon) Vernal Crayfish Le, Lo BW, TN P. vioscai paynei Fitzpatrick Payne’s Creek Crayfish Lo BW, TO P. zonangulus Hobbs & Hobbs Southern White River Crawfish Le, Lo UN