2001. Proceedings of the Academy of Science 110:104-110

CHECKLIST OF THE AND FRESHWATER SHRIMP () OF INDIANA

Thomas P. Simon: U.S. Fish and Wildlife Service, 620 South Walker Street, Bloomington, Indiana 47401

ABSTRACT. Crayfish and freshwater shrimp are members of the order Decapoda. All crayfish in In- diana are members of the family , while the freshwater shrimp belong to . Two genera of freshwater shrimps, each represented by a single , occur in Indiana. ka- diakensis and ohione are lowland forms. Macrobrachium ohione occurs in the drainage, while P. kadiakensis occurs statewide in wetlands and lowland areas including inland lakes. Currently, 21 crayfish taxa, including an undescribed form of Cambarus diogenes, are found in Indiana.

Another two species are considered hypothetical in occurrence. Conservation status is recommended for the Ohio shrimp Macrobrachium ohione, Indiana crayfish Orconectes indianensis, and both forms of the cave crayfish Orconectes biennis inennis and O. i. testii.

Keywords: Cambaridae, Palaemonidae, conservation, ecology

The crayfish and freshwater shrimp belong- fish is based on collections between 1990 and ing to the order Decapoda are among the larg- 2000. Collections were made at over 3000 lo- est of Indiana's aquatic . Crayfish calities statewide, made in every county of the possess five pair of periopods, the first is mod- state, but most heavily concentrated in south- ified into a large chela and dactyl (Pennak ern Indiana, where the greatest diversity of 1978; Hobbs 1989). The North American species occurs. families, crayfish belong to two Astacidae and The current list of species is intended to Cambaridae with all members east of the Mis- provide a record of the extant and those ex- sissippi River belong to the family Cambari- tirpated from the fauna of Indiana over the last dae (Hobbs 1974a). The freshwater shrimps two centuries (Table 1 ). This list includes new are represented by two genera in a single fam- information and taxonomic changes. This ef- ily, Palaemonidae. fort is the first step of the Indiana Biological The species of Cambaridae in North Amer- Survey to compile a listing of all known spe- ica represent a large family of over 300 de- cies of biota in Indiana. The taxonomic and scribed and numerous undescribed species in nomenclature sources include Hobbs (1989), two subfamilies (Hobbs 1989). The family Williams et al. (1989), Page (1985), Jezerinac also occurs in Japan, Korea, and the Amur et al. (1995) and Pflieger (1996). Many cray- basin of eastern Asia (Hobbs 1974). The Cam- fish species do not possess common names. barellinae include only the Cambarel- Species without accepted common names fol- lus, while the Cambarinae include ten genera (Hobbs 1974, 1977; Hobbs & Carlson 1983). lowing Williams et al. (1989) are highlighted The Palaemonidae is represented in North in brackets to signify those proposed in this America by 68 described species in 16 genera study. These names were based on those from (Williams et al. 1989). The family is world- adjacent state lists in (Pflieger 1996) wide in distribution. and Ohio (Thoma & Jezerinac 2000a). Sub- The purpose of this paper is to list the spe- genera are indicated for all crayfish species. cies of crayfish and freshwater shrimp known Crandall & Fitzpatrick (1996) and Fetzner to occur in Indiana and describe the range, (1993) have completed recent molecular stud- relative abundance, and recommended conser- ies on the phylogeny of Orconectes subgen- vation status. era, which suggests that these relationships METHODS may be doubtful. Use of the Orconectes phy- Distribution and range.—This present logeny relationships presented in this paper survey of Indiana freshwater shrimp and cray- follows Fitzpatrick (1987).

104 SIMON—CRAYFISH AND SHRIMP OF INDIANA 05

Historical collections of Indiana crayfish watt T & J generator capable of 300 V output and freshwater shrimps curated at the Ohio and usually 6—8 amps. All specimens ob- State University Museum (OSM), Illinois Nat- served were captured and a portion was re- ural History Survey (INHS), National Muse- tained for later identification in the laboratory. um of Natural History (NMNH), and Univer- Sites were sampled so that a minimum dis- sity of Michigan Museum of Zoology tance of 15 times the stream width was sam- (UMMZ) were compiled. Historical and re- pled. Each surveyed site consisted of a mini- cent records were compiled to formulate the mum distance of 50 m and a maximum current distribution of Indiana crayfish. distance of 500 m in streams and rivers. Lake The following codes were established to fa- and great river habitats were surveyed for 500 cilitate documentation of species in the State m of littoral shoreline habitat. All available

(Table 1). The general range in Indiana is in- habitats were sampled at each location includ- dicated as statewide (I), north (N), south (S), ing riffle, run, pool, various instream cover west (W), east (E), and various combinations types (e.g., woody debris, slab bedrock crev- of these regions. Relative abundance refers to ices, boulders, aquatic macrophytes), and be- the general population levels of the species neath undercut banks. All specimens were within the State. These are based on the abun- placed into a live well and retained until the dance and number of localities where the spe- end of the collection zone. cies has been found. Depending on local and Burrowing species of crayfish were more seasonal conditions, population levels will difficult to obtain. Two collection procedures vary. Relative abundance is listed as four cat- were attempted. For prairie crayfish that re- egories: abundant (A) designations are species mained in burrows, a modified toilet plunger that are easily found at a site in a particular was used to force the crayfish from the bur- region, common (C) are species that are reg- row. An aliquot of water was poured into the ularly found at a site (generally greater than burrow until full, then suction was established 25 individuals) in an area but perhaps not at at the entrance so that a good seal was estab- all locations, occasional (O) designations are lished. Plunging the burrow caused the exit for species that are found at either fewer than holes to become noticeable, and after several five locations or represent less than five spec- attempts the exits were examined to determine imens at a site, and rare (R) designations are if crayfish were present. Attempts to excavate for species that occur at less than five loca- the burrow by digging was foolishly attempt- tions or are represented by a single individual ed next; however, if this failed the identity of at a site. Extirpated species (Ex) are listed the crayfish was based on the external mor- along with the estimated date of disappear- phology of the burrow. Large multiple cham- ance. The State of Indiana does not have a bered chimney-type burrows was assumed to formal conservation listing for crayfish or be Cambarus diogenes, while piled burrows freshwater shrimp. The recommended conser- were assumed to be Procambarus gracilis. vation status has no formal or legal impetus; Single chambered chimney-type burrows were instead it refers to information based on our considered Fallicambarus fodiens. More than data. Federally endangered species are denot- likely I underestimated the distribution of ed as federally endangered (FE), federally prairie species since many of our collections threatened (FT), and federal candidates for were not completed during times when they Federal listing as FC; however, the candidate would have been more vulnerable to our col- listing has no formal status. State endangered lection methods (December-May). species will be represented by three codes: en- RESULTS DISCUSSION dangered (StE), threatened (StT), and special AND concern in need of further study (SC). An ad- Crayfish systematies.—The cambarid ditional designation, exotic (X) is included for crayfish differ from the Astacidae of western accidentally or deliberately released species. North America and Europe by the presence of Collection methodology.—Open water hooks on the ischia of the second through crayfish and freshwater shrimp were sampled fourth pereiopods and dimorphic cycling in by seining, dipnetting, or electrofishing all males (Hobbs 1974). Astacids do not exhibit representative habitats at a locality. Electro- cyclic dimorphism nor ischial hooks. Cam- fishing included the use of a pulsed DC 1850- barid crayfish have a variety of ornate termi- 106 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE nal elements on the gonopod, while astacid Momot 1967, 1978; Aiken 1969; Weagle & crayfish have simple cylindrical distal ele- Ozburn 1972; Momot & Gowing 1977). ments. The cambarid females have an annulus Life history strategies of Indiana cray- ventralis that is lacking in astacid females. fish.—The life history strategies occurring During the reproductive season, males devel- among Indiana crayfish include cave-dwell- op one or more terminal elements on the gon- ing, non-burrowing and burrowing species. opods (first pleopods) that are corneous. These Cave-dwelling species are adapted for spend- males are referred to as form I. ing their entire existence in caves. Non-bur- Historical studies of Indiana crayfish and rowing crayfish carry out their entire life his- freshwater shrimp.—Most historical infor- tory in surface waters. Burrowing species mation published on Indiana crayfish and spend variable amounts of time in surface wa- freshwater shrimp can be found in Cope ters and periodically leave their burrows to (1872), Packard (1873), Bundy (1877), Hay mate (Simon et al. 2000). Adults of these spe- (1891, 1893, 1896), Williamson (1907), Ev- cies remain above ground only during periods ermann & Clark (1920), Eberly (1955), Hobbs of late winter to spring flooding (January- (1989), Page (1994), and Page & Mottesi May). Sexually mature P. gracilis leave their (1995). Cope (1872) published on the cave burrows on warm rainy nights but are other- fauna of Wyandotte Cave, while Packard wise seldom found out of their burrows. Adult (1873) published on cave crayfish throughout C. diogenes leave their burrows on the edge the State. Bundy (1877) was the first to pub- of stream banks more often than the other bur- lish on crayfish distributions outside of caves rowing species and may forage as well as in Indiana, focusing on species in northern In- mate. Roaming occurs most frequently during diana. Hay (1891) published on the crusta- the mating season and when females are car- ceans of Indiana, including information on rying eggs or young. freshwater shrimp, and followed with a paper Distribution and conservation status.— on the observations of blind cave crayfish in- The State of Indiana possesses 21 crayfish cluding the description of Cambarus inermis taxa representing 19 species and possibily an- testii (Hay 1896). Hay (1896) first published other two undescribed species (R. Thoma, an annotated species list that included infor- pers. commun.). Page & Mottesi (1995) indi- mation on the of crayfish species cated that 18 crayfish species occurred in In- occurring in Indiana. Williamson (1907) pub- diana; however, their list did not include the lished information on the crayfish of Wells recently-elevated O. juvenilis nor Cambarus County and described a new species (Cam- robustus. Hobbs (1989) tentatively listed O. barus ortmanni). Evermann & Clark (1920) putnami from Indiana. Page & Mottesi (1995) listed the species occurring in Lake Maxink- found O. putnami to be widespread in south- uckee. Eberly (1955) summarized distribu- eastern portions of the State as did I. Taylor tions of five species and included new distri- (1997) synonymized C laevis, C. ornatus, and bution records. Page (1994) conducted a study C tenebrosus east of the as to determine the conservation status of Or- C tenebrosus based on meristic and morpho- conectes indianensis that included a listing of metric characters. This present paper still the species of crayfish found in Indiana (Page maintains the separate listing of the species & Mottesi 1995); and he indicated that 17 spe- for Indiana while further work is being con- cies and an additional undescribed species are ducted to support this revision. known to occur in Indiana. An additional two species are considered Life history and distribution information for hypothetical for their occurrence in the State. Indiana crayfish include C. diogenes (Grow The possibility that C thomai exists in south- 1981), C fodiens (Bovbjerg 1952), C. tene- eastern Indiana is based on the species distri- brosus (Prins 1968), O. immunis (Tack 1941), bution in southwestern Ohio (Jezerinac 1993; O. propinquus (Van Deventer 1937; Bovbjerg Thoma & Jezerinac 2000b). Finally, O. stan- 1952), O. rusticus (Langlois 1935; Busch nardi occurs as an endemic of the Little Wa- 1940; Prins 1968), O. sloani (Rhoades 1962; bash River in Illinois. The possibility of the Jezerinac 1986; St. John 1988) and O. virilis species occurring in direct tributaries to the (Bovbjerg 1953, 1970; Hazlett et al. 1974; Wabash River near the Little Wabash River Threinen 1958; Caldwell & Bovbjerg 1969; needs further survey. SIMON—CRAYFISH AND SHRIMP OF INDIANA [07

Table 1. —Checklist of the crayfish and freshwater shrimp (Decapoda) of Indiana following Hobbs (1974) including distribution, relative abundance, and recommended conservation status. Common names follow Williams et al. (1988), with the exception of names in brackets, which are suggested additions based on Pflieger (1996) and Thoma & Jerezinac (2000a). Subgenera of Orconectes follow Fitzpatrick (1987). Statewide (I), north (N), south (S), west (W), east (E), and various combinations of these regions. Relative abundance: R = rare; C = common; O = occasional. Conservation status: StE = state endan- gered; FC = candidate for federal listing; SC = candidate for special concern; NI* = nonindigenous. and introduction has occurred in a portion of the species range in Indiana.

Recommended Relative conservation Taxa Range abundance status

Order Decapoda Family Palaemonidae (freshwater shrimp) Macrobrachium ohione (Smith), Ohio shrimp R StE Palaemonetes kadiakensis Rathbun, Mississippi grass shrimp C Family Cambaridae (crayfish) Genus Procambarus Subgenus Girardiella P. gracilis (Bundy), prairie crayfish w o Subgenus Ortmannicus

P. acutus (Girard), White River crayfish I () Subgenus Scapulicambarus P. clarkii (Girard), red swamp crayfish NW, SW R NI : Genus Orconectes Subgenus Crockerinus

O. propinquus (Girard), northern clearwater crayfish 1 C Subgenus O. indianensis (Hay), [Indiana crayfish] SW R FC, SC Subgenus Gremicambarus

O. immunis (Hagen) [papershell crayfish] 1 C

O. virilis (Hagen), northern crayfish 1 C Subgenus Orconectes O. inermis inermis Cope [Indiana cave crayfish] s R StE O. inermis testii Cope [Hoosier cave crayfish] s R StE Subgenus Procericambarus O. putnami (Faxon) [Putnam's crayfish] SE () O. juvenilis (Faxon) [miniature crayfish] SE R

O. rusticus (Girard), rusty crayfish I C NI* Subgenus Rhoadesius O. sloanii (Bundy) [Sloan's crayfish] SE R Genus Fallicambarus Subgenus Creaserinus F. fodiens (Cottle) [digger crayfish] () Genus Cambarus Subgenus Cambarus C. bartoni cavatus (Fabricius) [Ohio crawfish] SE R C. ortmanni Williamson [Ortmann's mudbug] N R Subgenus Erebicambarus C. laevis Faxon [karst crayfish] S C C. tenebrosus Hay [cavespring crayfish] s c Subgenus Lacunicambarus

C. diogenes Girard, devil crayfish 1 R Subgenus Tubericambarus C. cf diogenes new species [Painted hand mudbug] s R Subgenus Puncticambarus C. robitstus Girard [bigwater crayfish] NE O Hypothetical in occurrence in Indiana Subgenus Tubericambarus Cambarus thomai (Jezerinac) [little brown mudbug] SE Subgenus Crockerinus O. stannardi Page [Little Wabash River crayfish] SW 108 PROCEEDINGS OF THE INDIANA ACADEMY OF SCIENCE

The rusty crayfish O. rusticus and red verely reduced over the species' former range. swamp crayfish Procambarus clarki have The species, reaching 100 mm carapace length been found in the Lake Michigan drainage (CL), once occurred throughout the Ohio Riv- (Simon, unpubl. data). The rusty crayfish is a er and lowland tributaries. During our study native species of southeastern Indiana that has we did not collect a single specimen and con- been widely distributed through bait bucket sider the species so rare as to warrant "State release by anglers. The species is native to the Endangered" status. It may already be extir- Whitewater River drainage and should be con- pated from Indiana's portion of the Ohio Riv- sidered non-indigenous outside of that water- er. However, Hobbs & Massmann (1952) sug- shed. The red swamp crayfish was collected gested that the species occurs in deeper during 2000 from Lake Michigan. Page & waters, which requires trapping to collect. The Mottesi (1995) indicated that this species was species rarity may only be a reflection of the among the rarest crayfish of Indiana. The red collection methods attempted or could be ex- coloration of the species and the recent mar- tirpated as a result of the navigation lock and keting of them as freshwater lobster in the dam system on the Ohio River prevented mi- aquarium trade perhaps have aided in the gration. spread of the species into the West Branch of ACKNOWLEDGMENTS the Grand Calumet River. Although the State of Indiana does not rec- I appreciate the assistance of numerous col- ognize any conservation status for crusta- leagues that assisted in the field collection of ceans, I recommend that three rare species be specimens especially Brant Fisher, Steve Wen- designated as either State Endangered or State te, Andy Ellis, Doug Campbell, Erich Emery Special Concern. Page (1994) surveyed for and Greg Nottingham. Additional specimens the Indiana crayfish O. indianensis (Hay), a were donated by Tony Branam, Ronda Dufour Federal candidate species, over the historic and Steve Newhouse. Conversations with Lar- range of the species in Illinois and Indiana. ry Page and Chris Taylor (Illinois Natural His- The species was collected from the Patoka tory Survey), Roger Thoma (Ohio State Uni- River and at additional locations in several versity), and Horton Hobbs, III (Wittenberg watersheds in southwestern Indiana. The wa- College) greatly helped nurture and stimulate tersheds where this species occurs are prone my interest in . Roger Thoma, to severe land use disturbance from oil and Ohio State University, reviewed a draft of this gas explorations, acid mine drainage and coal manuscript. The U.S. Fish and Wildlife Ser- mining. The species has been severely re- vice, Division of Contaminants supported this duced in its former range in Illinois, but Page study through grant number 31440-1261- did not recommend the species for listing 3N29. since it occurred at many of the historic sites LITERATURE CITED where it had been collected in Indiana. Due to Aiken, D.E. 1969. Ovarian maturation and egg lay- the species primary distribution in areas se- ing in the crayfish Orconectes virilis: Influence verely impaired by anthropogenic disturbance of temperature and photoperiod. Canadian Jour- in the Patoka and Pigeon River drainage and nal of Zoology 47:931-935. southwestern Indiana, it is recommended that Bovbjerg, R.V. 1952. Comparative ecology and the species be considered "State Special Con- physiology of the crayfish Orconectes propin- cern" until it can be determined whether the quus and Cambarus fodiens. Physiological Zo- species range is being threatened. ology 25:34-56. Orconectes biennis inermis and O. biennis Bovbjerg, R.V. 1953. Dominance order in the cray- testii are cave-dwelling crayfish found in fish Orconectes virilis (Hagen). Physiological 26:173-178. southern Indiana. The two taxa are rare with Zoology Bovbjerg, R.V. 1970. Ecological isolation and O. inermis testii being restricted to Monroe competitive exclusion in two crayfish {Orconec- County, while O. inermis inermis being broad- tes virilis and Orconectes iinmunis). Ecology 51: er ranging. two taxa are consid- The seldom 225-236. ered abundant. It is that both recommend Bundy, W.F. 1877. On the Cambari of northern In- forms be considered "State Endangered" be- diana. Proceedings of the Academy of Natural cause of the fragile nature of karst ecosystems. History of Philadelphia 29:171-174. The Ohio shrimp (M. ohione) has been se- Busch, K.H.D. 1940. Embryology of the crayfish, SIMON—CRAYFISH AND SHRIMP OF INDIANA 09

Cambarus rusticus Girard. Doctoral dissertation. Hobbs, H.H., Jr. & PH. Carlson. 1983. Distocam- The Ohio State University, Columbus. barus (Decapoda: Cambaridae) elevated to ge- Caldwell, M.J. & R.V. Bovbjerg. 1969. Natural neric rank, with an account of D. crockeri, new history of the two crayfish of northwestern Iowa, species, from South Carolina. Proceedings of the Orconectes virilis and O. immunis. Proceedings Biological Society of Washington 96:420-428.

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fish and Shrimp Atlas. Ohio Biological Survey Manuscript received 29 May 2001 , revised 25 Au- Miscellaneous Contribution 7. 28 pp. gust 2001.