Zootaxa 3717 (2): 101–157 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3717.2.1 http://zoobank.org/urn:lsid:zoobank.org:pub:C8CDC011-974C-48B4-9E03-88F570EEDE13 Crayfishes (Decapoda : Cambaridae) of Oklahoma: identification, distributions, and natural history

REID L. MOREHOUSE1 & MICHAEL TOBLER Department of Zoology, Oklahoma State University, 501 Life Sciences West, Stillwater, OK 74078, USA 1Corresponding author. E-mail: [email protected]

Table of contents

Abstract ...... 102 Introduction ...... 102 Oklahoma and its crayfishes ...... 103 Material and methods ...... 103 Results ...... 108 Dichotomous Key for Oklahoma Crayfish ...... 108 Species accounts ...... 121 Cambarellus puer Hobbs, 1945: Swamp Dwarf Crayfish ...... 121 Cambarus diogenes Girard, 1852: Devil Crawfish ...... 122 Cambarus ludovicianus Faxon, 1884: Painted Devil Crayfish ...... 123 Cambarus subterraneus Hobbs, 1993 : Delaware County Cave Crayfish ...... 123 Cambarus tartarus Hobbs and Cooper, 1972: Oklahoma Cave Crayfish ...... 124 Fallicambarus fodiens (Cottle, 1863): Digger Crayfish ...... 124 Faxonella blairi Hayes and Reimer, 1977: Blair's Fencing Crayfish ...... 125 Faxonella clypeata (Hay, 1899): Ditch Fencing Crayfish ...... 126 Orconectes causeyi Jester, 1967: Western Plains Crayfish ...... 126 Orconectes deanae Reimer and Jester, 1975: Conchas Crayfish ...... 127 Orconectes difficilis (Faxon, 1898): Painted Crayfish ...... 128 Orconectes lancifer (Hagen, 1870): Shrimp Crayfish ...... 128 Orconectes leptogonopodus Hobbs, 1948: Little River Creek Crayfish ...... 129 Orconectes macrus Williams, 1952: Neosho Midget Crayfish ...... 129 Orconectes menae Creasar, 1933: Mena Crayfish ...... 130 Orconectes meeki brevis Williams, 152: Meek's Short Pointed Crayfish ...... 131 Orconectes nais (Faxon, 1885): Water Nymph Crayfish ...... 131 Orconectes nana Williams, 1952: Midget Crayfish ...... 132 Orconectes neglectus neglectus (Faxon, 1885): Ringed Crayfish ...... 132 Orconectes palmeri longimanus (Faxon, 1898): Western Painted Crayfish ...... 133 Orconectes saxatilis Bouchard and Bouchard, 1976: Kiamichi Crayfish ...... 133 Orconectes virilis Hagen, 1870: Virile Crayfish ...... 134 Procambarus acutus (Girard, 1852): White River Crawfish ...... 135 Procambarus clarkii (Girard, 1852): Red Swamp Crawfish ...... 135 Procambarus curdi Reimer, 1975: Red River Burrowing Crayfish ...... 136 Procambarus dupratzi Penn, 1953: Southwestern Creek Crayfish ...... 137 Procambarus gracilis (Bundy, 1876): Prairie Crayfish ...... 137 Procambarus liberorum Fitzpatrick, 1978: Osage Burrowing Crayfish ...... 138 Procambarus simulans (Faxon, 1884): Southern Plains Crayfish ...... 139 Procambarus tenuis Hobbs, 1950: Ouachita Mountain Crayfish ...... 139 Discussion ...... 140 Acknowledgements ...... 141 References ...... 142 Glossary ...... 157

Accepted by J. Goy: 21 Aug. 2013; published: 27 Sept. 2013 101 Abstract

We furnish an updated crayfish species list for the state of Oklahoma (United States of America), including an updated and illustrated dichotomous key. In addition, we include species accounts that summarize general characteristics, life col- oration, similar species, distribution and habitat, life history, and syntopic species. Current and potential distributions were analyzed using ecological niche models to provide a critical resource for the identification of areas with conservation pri- orities and potential susceptibility to invasive species. Currently, Oklahoma harbors 30 species of crayfish, two of which were recently discovered. Eastern Oklahoma has the highest species diversity, as this area represents the western distribu- tion extent for several species. The work herein provides baseline data for future work on crayfish biology and conserva- tion in Oklahoma and surrounding states.

Key words: Cambaridae, Crayfish, Crustacean, Dichotomous Key, Ecological Niche Modeling

Introduction

Crayfish are freshwater crustaceans of the families Astacidae, Cambaridae, and Parastacidae, and are native to every continent except for Antarctica and Africa (Hobbs, 1988). They inhabit a wide variety of aquatic habitats including streams, rivers, lakes, reservoirs, swamps, roadside ditches, wet pastures, and fields (Bouchard, 1978). In many habitats, crayfish play important roles in ecosystem functioning and can comprise the majority of invertebrate biomass (Momot et al., 1978; Rabeni et al., 1995). They are typically considered omnivores, exploiting a diversity of food sources from particulate organic matter to aquatic vertebrates and fish, and occupy an intermediate trophic position between other invertebrate consumers and fish (Schofield et al., 2001; Taylor & Soucek, 2010; Whitledge & Rabeni, 1997). Furthermore, crayfish have a substantial economic impact in the southern United States, particularly in Louisiana, where nearly $45 million worth of crayfish have been exported annually since the early 1980's (Walls, 2009). Over 75% of the known crayfish species diversity (roughly 406 species and subspecies) occur in North America (Taylor, 2002). Two families of crayfish inhabit North America, Astacidae with 4 species is restricted to the Pacific Northwest (Larson & Olden, 2011), and Cambaridae distributed east of the Rocky Mountains, with two- thirds of its species endemic to the southeastern United States (Taylor et al., 2007). Cambarids are classified into two subfamilies, Cambarellinae including a single genus (Cambarellus) and Cambarinae, which contains ten genera. Three of the ten Cambarinae genera (Cambarus, Orconectes, and Procambarus) encompass 85% of all crayfish species and subspecies known in North America; Procambarus alone includes 163 described species and represents the most species-rich genus worldwide (Taylor, 2002; Taylor et al., 2007). The species level diversity of crayfishes is relatively well documented, but new species are continuously being described (e.g., Schuster, 2008; Taylor & Soucek, 2010; Taylor et al., 2006). In addition, a host of studies over the past decades have started to elucidate the phylogenetic relationships among North American taxa (e.g., Crandall, 2006; Fetzner & Crandall, 2002). Nonetheless, relatively little information is available about the ecology and life history of many species, even though there is a growing interest in crayfish biology, particularly because of their increasing need for conservation. A recent review indicated that nearly 50% of all crayfish north of Mexico are imperiled (Taylor et al., 2007). Crayfish are particularly threatened because of loss or degradation of suitable habitats and the introduction of nonindigenous species, which is often exacerbated by narrow distributional ranges and high levels of endemism (Larson & Olden, 2011; Taylor et al., 2007; Welsh et al., 2010). While habitat loss and degradation have caused many detrimental impacts to crayfish, the introduction of nonindigenous species is likely the biggest threat (Gherardi, 2006; Holdich et al., 2009; Lodge et al., 2012; Lodge et al., 2000; Morehouse & Tobler, 2013). For example, nonindigenous crayfish introductions have caused drastic changes to ecosystems both through competitive exclusion of native congeners and physical habitat alterations (Morehouse & Tobler, 2013; Olden et al., 2009; Olden et al., 2006). In addition, nonindigenous crayfish species have been documented to negatively impact primary production and affect macroinvertebrate community structure in some systems (Lodge et al., 2012; Lodge et al., 1994; McCarthy et al., 2006). On the ground conservation efforts for crayfish are often hindered by the lack of basic knowledge about resident crayfish species and their ecology: (1) Crayfish can be difficult to identify, particularly for non-specialists. While species are typically delineated based on multiple morphological characters (Hobbs, 1989), the structures of

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