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Distribution patterns of freshwater shrimp and crayfish (Decapoda: Cambaridae) in the Patoka River basin of Indiana.

ABSTRACT. Eleven species of crayfish and freshwater shrimp were collected during a study of the Patoka River drainage of Indiana between 2000-2002. The Mississippi grass shrimp Palaemonetes kadiakensis was the only shrimp species collected from wetland ponds and stream areas adjacent to vernal ponds. Several rare species were encountered, including Orconectes (Faxonius) indianensis and O. (Orconectes) inermis inermis. Orconectes (Orconectes) inermis inermis was collected from a single cave location near Valeene. Orconectes (Faxonius) indianensis was found throughout the watershed, but was most common in the upper third of the watershed within the Hoosier National Forest. Procambarus (Ortmannicus) acutus and two Erebicambarus species members, Cambarus laevis and C. tenebrosus, were collected from small streams emanating from caves and karst springs. Three primary burrowing crayfish forms (Fallicambarus (Creaserinus) fodiens, Cambarus (Tubericambarus) sp. A and Cambarus (Lacunicambarus) sp. A.) were more common throughout the watershed, while Cambarus (L.) sp. A was collected only from the glaciated portions of the watershed.

Keywords: Cambarus, Orconectes, Fallicambarus, Procambarus, Palaemonetes, Cambaridae, Palaemonidae

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The crayfish family Cambaridae represents a large group of over 300 described and undescribed species in two subfamilies (Hobbs 1989). Simon (2001) provided a checklist of Indiana species that documented the occurrence of two freshwater shrimp and 22 crayfish species within the political boundaries of Indiana. Despite the study of decapod crustaceans within Indiana for the last 130 years (Cope 1872; Packard 1873; Bundy 1877), limited surveys have documented the faunas of complete watersheds. The most studied Indiana faunas have been the crayfish of northern Indiana (Bundy 1877; Williamson 1907), Lake Maxinkuckee (Evermann & Clark 1920), and cave faunas (Cope 1872; Packard 1873; Hay 1893; Lewis 1983; Lewis et al. 2002, 2003).

The Patoka River drainage is a large tributary of the lower Wabash River that flows into the Ohio River and possesses an interesting crayfish fauna because of the east to west direction of flow and the crossing of several physiographic provinces (Schneider 1966). Virtually nothing is known of the decapod fauna of the Patoka River drainage; however, Simon et al. (1995) collected fish throughout the watershed and reported on anthropogenic changes between 1888 and 2001 (Simon et al. 2003). No previous distribution studies of the crayfish and freshwater shrimp have been conducted in the Patoka River watershed. The purpose of the current study was to document the shrimp and crayfish fauna of a large watershed in southern Indiana.

METHODS

Study area.--The Patoka River is a narrowly-confined tributary of the lower Wabash River that originates near Valeene, Orange County, Indiana (Simon et al. 1995). We sampled at 125 lentic and lotic sites throughout the entire watershed (Fig. 1). The Patoka River flows west for 162 miles (260 km) across the Crawford Upland and the Wabash Lowland physiographic units (Schneider 1966), draining 862 square miles (1388 [km.sup.2]) in Gibson, Pike, Dubois, and Orange Counties. Several large public land holdings include the Patoka River unit of the Hoosier National Forest, Patoka Lake and the U.S. Army Corps of Engineers holdings, and the U.S. Fish and Wildlife Service's Patoka River National Wildlife Refuge.

[FIGURE 1 OMITTED]

Collection methodology.--Open-water crayfish and freshwater shrimp were sampled by seining, dipnetting, or electrofishing of all representative habitats at a locality (Simon 2001). Electrofishing included the use of a pulsed DC Smith-Root generator backpack electrofishing unit capable of 300 V output and usually 3-6 amps. All specimens observed were captured, and a portion was retained for later identification in the laboratory. Sites were sampled so that a minimum distance of 15 times the stream width was sampled. Each surveyed site consisted of a minimum distance of 50 m and a maximum distance of 500 m in large-to-moderate sized rivers. Oxbow pond and lake littoral shoreline habitat was surveyed for 500 m. All available habitats were sampled at each location including riffle, run, pool, various instream cover types (e.g., woody debris, slab bedrock crevices, boulders, aquatic macrophytes), and beneath undercut banks. All specimens were placed in a live well and retained until the end of the collecting zone.

Burrowing crayfish were collected using two approaches. Prairie crayfish that remained in burrows were collected using a modified toilet plunger to force the crayfish from the burrow (Simon 2001). Water was poured into the burrow until full, then suction was applied at the entrance so that a good seal was established. Plunging the burrow caused the exit holes to be exposed, and after several attempts the exit holes were examined for the presence of the resident crayfish. If the plunger dial not reveal the crayfish, then a spade was used to excavate the burrow. Depths that were excavated in the Patoka River drainage ranged between 0.6-2 m.

Specimens retained for laboratory identification were identified using Page (1985), Pflieger (1996), and Thoma & Jezerinac (2000). All specimens were deposited in the crustacean collection of the Indiana Biological Survey, Bloomington.

RESULTS AND DISCUSSION

Distribution of decapods.--The Patoka River drainage includes 10 species of crayfish and a single freshwater shrimp (Tables 1, 2). Only a single species of freshwater shrimp was collected from the watershed (Fig. 2). The Mississippi grass shrimp Palaemonetes kadiakensis was found primarily in the mainstem Patoka River; however, the species was also found in small tributaries adjacent to vernal ponds and wetlands. This species is usually more common in pond, marsh, and slough habitats than in streams. Female specimens collected during April were ovigerous and were pigmented with lime-green bars along their sides.

Ten Cambaridae species were collected from the Patoka River drainage including members of each of the four Indiana crayfish genera (Tables 1, 2). Procambarus acutus was the only Procambarus species collected (Fig. 4). The species was most common in small headwater streams that possessed overhanging grasses and dense root wads along the stream margins. No ovigerous females were collected during this study.

Two Erebicambarus species, Cambarus laevis and C. tenebrosus were collected from karst areas of the watershed (Tables 1, 2; Fig. 4). Taylor (1997) has considered these species as synonymous; however, we follow Simon (2001) in separating them until further study can verify Taylor's hypothesis. Cambarus laevis was collected from two sites in the upper portion of the watershed (Fig. 4). One site was inside Spring Spring cave and the other from a breakdown area where groundwater was emanating from a newly forming cave. The two species were found beneath large boulder rocks in springs outflows and near the entrances of caves. Within the Patoka River drainage the two species were not sympatric; however, elsewhere the two species are commonly collected together at the same site (Simon unpubl. data).

Three primary burrowing species were collected from the watershed (Fig. 3, 4). Two Cambarus species and Fallicambarus (Creaserinus) fodiens were collected from roadside ditches, farm fields, and wetland habitats (Tables 1, 2). The widest-ranging burrowing species was Cambarus (Tubericambarus) sp. A (Fig. 3). This species was mentioned by Jezerinac (1993) as ranging throughout southern Indiana, Ohio, and Illinois. This species was the only crayfish at many of the sites affected by acid mine drainage, oil brine, and coal mining. Fallicambarus (Creaserinus) fodiens was the second most abundant and widest distributed burrowing species (Fig. 4). We collected F. (C.) fodiens and Cambarus (T.) sp. A together and Cambarus (T.) sp. A and Cambarus (Lacunicambarus) sp. A at several of the same sites, but all three burrowing species are not sympatric at the same site.

Four secondary burrowing species of the genus Orconectes were collected during this study (Tables 1, 2). The most abundant and widely distributed was O. (Gremicambarus) immunis, which was collected from among woody debris, overhanging vegetation, and debris piles (Fig. 2). Orconectes (Crockerinus) propinquus was collected from four sites in the lower Patoka River (Fig. 4). Surprisingly, Orconectes (Faxonius) indianensis was much more common than previously considered (Thoma & Simon unpubl, data), primarily being collected from the upper third of the watershed (Fig. 2). The troglobitic crayfish O. inermis inermis was found in a single cave in the extreme headwaters of the Patoka River. Thoma et al. (unpubl. data) has reviewed the distribution of records for the northern cave crayfish and found that the species was not as broadly distributed in the Patoka River drainage as originally considered (Fig. 3).

Several sites were dry or were intermittent during the sampling period. They include Flat Creek tributary (site 104), Hog Branch (site 109), unnamed tributary at SR 364 bridge (site 110), Wheeler Creek (site 10), the old Wabash and Erie Canal (site 111), and Lick Creek (site 112). All of these streams also lacked water during the sampling conducted in 1993 and 2000 (Simon et al. 1995).

No crayfish were collected from Rough Creek (sites 9, 22, 23), Wheeler Creek (site 10), Durham ditch (site 12), South Fork Patoka River (site 19), Rough Creek tributary (site 21), Robinson Creek (site 25), and Sugar Creek (site 106). These sites were affected by acid mine drainage, oil brine, or coal mining (Simon et al. 1995, 2003). No crayfish were collected from any of the lake of oxbow sites sampled in the watershed. The lakes and oxbows sampled included Patoka Lake (sites 113-118), Huntingburg Lake (sites 119, 120), Beaver Dam Lake (site 121), New Oakland City Lake (sites 122, 123), Big Bottom wetland (site 124), and Tucker Lake (site 125).

Gradient Patterns Affecting Distribution.--The distribution of crayfish and shrimp follows several landscape scales including Ecoregions, Sub-ecoregions, and Natural Divisions. Omernik and Gallant (1988) defined Ecoregions for the midwest states, Woods et al. (1996) defined Sub-ecoregions for Indiana and Ohio, while Natural Divisions were defined by Homoya et al. (1985). Both Ecoregion frameworks are based on climate, land use, physiography, potential natural vegetation at different scales, while Natural Divisions include zoogeographic patterns. The Patoka River contains two Ecoregions, the Interior River Lowland to the west and the Interior Plateau to the east, while several Sub-ecoregions are recognized within the Interior River Lowland. Homoya divided the Patoka River into six subdivisions. These subdivisions from west to east include the Southern Bottom Lands, Southwestern Lowlands-glaciated, Southwestern Plainville, Southwestern Lowlands-driftless, and Shawnee Hill-Crawford Upland.

Species distributions that were exclusively contained within the Interior River Lowland Ecoregion included Procambarus (Ortmannicus) acutus, Cambarus (Lacunicambarus) sp. A, Fallicambarus (Creaserinus) fodiens, and O. (Crockerinus) propinquus (Figs. 3, 4). Species distributions that are contained within the Interior Plateau Ecoregion include Cambarus (Erebicambarus) tenebrosus and O. (Orconectes) inermis inermis. No pattern in ecoregion distribution was observed for Palaemonetes kadiakensis, Cambarus (Tubericambarus) sp. A, O. (Gremicambarus) immunis, and O. (Faxonius) indianensis.

Cambarus (L.) sp. A (Fig. 3) and Palaemonetes kadiakensis (Fig. 2) occur along the Southwestern Lowlands-glaciated subdivision portions of the watershed, while Fallicambarus (C.)fodiens is limited to the Southwestern Lowlands-driftless subdivision (Fig. 4). Cambarus (E.) tenebrosus and O. inerrnis inermis were limited to the Shawnee Hill-Crawford Upland subdivision (Fig. 3).

Patterns in zoogeographic distributions of Indiana crayfish species are largely determined by the last glacial advance (Simon unpubl. data). The Wisconsian glacier only covered the western-most portion of the lower Patoka River. Because glacial refugia were established and more southern species were able to remain in these areas, watersheds such as the Patoka River are more faunistically diverse than northern watersheds that were completely obliterated (Simon 2001). Thus, southern portions of Indiana that were not affected by the last glacial advance will require further collection efforts to document the biodiversity of the crayfish and shrimp fauna.
Table 1.--List of collection information and corresponding spatial
coordinates (decimal degree) for sites sampled in the Patoka River
drainage between 2001 and 2002. Abbreviations include, CR = County
Road, SR = State Road, mi = miles, Twp = Township, u/s = upstream,
d/s = downstream.

 Site
number County Site

 1 Gibson Hull ditch, CR 350 N bridge, 3 mi E East Mt.
 Cannel, White River Twp

 2 Tripplet ditch, CR 400 N bridge, 2.5 mi E Patoka,
 Washington Twp

 3 Lost Creek, CR 50 N bridge, 2.5 mi N Francisco,
 Center Twp

 4 Tributary E Branch Keg Creek, CR 50 N bridge, 2.5
 mi E Francisco

 5 W Fork Keg Creek, SR 64 bridge, 2 mi E Francisco,
 Columbia Twp

 6 Turkey Creek, CR 1275 E bridge, 0.75 mi E Oakland
 City, Columbia Twp

 7 Pike Hat Creek, CR 50 E bridge, 1 mi N Coe, Monroe Twp

 8 Tributary South Fork Patoka River, CF 900 S
 bridge, 1 mi S Coe, Monroe Twp

 9 Rough Creek, CR 1000 S bridge, 0.5 mi E Spurgeon,
 Monroe Twp

 10 Wheeler Creek, CR 775 S bridge, 1 mi E Coe, Monroe
 Twp

 11 Tributary Turkey Creek, unnamed CR, 2 mi SW
 Arthur, Monroe Twp

 12 Durham ditch, CR 800 S bridge, 0.25 mi NW
 Scottsburg, Monroe Twp

 13 South Fork Patoka River, near CR 200 E bridge, 2
 mi W Stendal, Monroe Twp

 14 Rock Creek, CR 1000 E bridge, 1.5 mi E Pikesville,
 Lockhart Twp

 15 South Fork Patoka River, CR 1200 S bridge, 2.5 mi
 E Spurgeon, Monroe Twp

 16 Cup Creek, SR 257 bridge, 0.3 mi SW Pikeville,
 Lockhart Twp

 17 Farm field adjacent Cup Creek mouth, 0.5 mi NW
 Pikeville, Lockhart Twp

 18 Barren ditch, unnamed CR bridge, 2 mi SW Winslow,
 Patoka Twp

 19 South Fork Patoka River, CR 875 S bridge, 1.2 mi
 E Coe, Monroe Twp

 20 South Fork Patoka River, CR 300 E bridge, 1.5 mi
 SE Coe, Monroe Twp

 21 Tributary Rough Creek, CR 200 E bridge, 1.5 mi SSE
 Coe, Monroe Twp

 22 Rough Creek, CR 200 E bridge, 1.75 mi SSE Coe,
 Monroe Twp

 23 Rough Creek, CR 200 E bridge, 2.5 mi S Coe, Monroe
 Twp

 24 Stone Coe Creek, SR 64 bridge, 0.25 mi N Winslow,
 Patoka Twp

 25 Robinson Creek, CR 475 E bridge, 1.2 mi NE
 Winslow, Patoka Twp

 26 Bruster Branch, CR 200 S bridge, 0.75 mi NE
 Winslow, Patoka Twp

 27 Patoka River, SR 61 bridge, Winslow, Patoka Twp

 28 Tributary Patoka River, SR 364 bridge, 0.75 mi S
 Winslow, Patoka Twp

 29 Mill Creek, u/s SR 364 bridge, 0.8 mi SE Winslow,
 Patoka Twp

 30 Mill Creek, d/s CR 450 E bridge, 0.8 mi SE
 Winslow, Patoka Twp

 31 Tributary Patoka River, SR 364 bridge, 1.5 mi SE
 Winslow, Marion Twp

 32 Tributary Patoka River, Forest Road, 2.0 mi E
 Winslow, Patoka Twp

 33 Tributary Patoka River, CR 450 E bridge, 2 mi E
 Winslow, Marion Twp

 34 Dubois E Branch Ell Creek, CR 585 W bridge, 2.0 on WSW
 Huntingburg, Patoka Twp

 35 Ell Creek, CR 400 S bridge, 2.0 mi NW Huntingburg,
 Patoka Twp

 36 Flat Creek, CR 250 S bridge, 4 on SSE Otwell,
 Madison Twp

 37 Flat Creek, SR 257 bridge, 2.5 mi S Otwell, Marion
 Twp

 38 Flat Creek. CR 300 W bridge, 1.8 mi SSW Otwell,
 Marion Twp

 39 Little Flat Creek, CR 300 W bridge, 2 mi SE
 Orwell, Madison Twp

 40 Unnamed tributary, SR 56 bridge, Ireland, Madison
 Twp

 41 Mill Creek tributary, SR 231 bridge bypass, 1.5 on
 S Haysville, Harbison Twp

 42 Mill Creek, CR 600 N bridge, 1.5 mi SW Haysville,
 Boone Twp

 43 Flat Creek, CR 230 S bridge, 0.5 mi E
 Mathersville, Jackson Twp

 44 Indian Creek, SR 64 bridge

 45 Hall Creek tributary, Meridian Road

 46 Short Creek, Sunset Road, 1.5 mi SE Huntingburg,
 Ferdinand Twp

 47 Indian Creek, SR 162 bridge

 48 Patoka River, SR 162 bridge

 49 Davis Creek, Cuzco Road bridge, Crystal, Columbia
 Twp

 50 Cane Creek tributary, CR 650 S bridge, 1 mi E
 Cuzco, Columbia Twp

 51 Cane Creek, CR 500 S bridge, 4.5 mi S French Lick,
 Jackson Twp

 52 Cane Creek, Cuzco Road bridge, S Cuzco, Columbia
 Twp

 53 Patoka River, Dubois-Cuzco Road bridge, 0.8 mi SW
 Cuzco, Columbia Twp

 54 Patoka River tributary, Jasper-Dubois Road bridge,
 0.5 mi S Dubois, Marion Twp

 55 Beaver Dam Lake Creek, CR 175 E bridge, 4.5 mi NW
 Jasper, Marion Twp

 56 Flat Creek, CR 450 S bridge, 0.5 on W St. Anthony,
 Jackson Twp

 57 Flat Creek, CR 600 E bridge. 1 mi ESE St. Anthony,
 Jackson Twp

 58 Grassy Fork, Santine Road bridge, 1.25 an NNE St.
 Anthony, Jackson Twp

 59 Unnamed tributary Patoka Lake, 0.25 on SW
 Wickliffe, Patoka Twp

 60 Hall Creek, Santine Road bridge, 1.25 mi SW
 Celestine, Marion Twp

 61 Hall Creek, Schnellville-Birdseye Road bridge,
 0.25 mi E Schnellville, Jefferson Twp

 62 Unnamed tributary Patoka Lake, 1.2 mi N Birdseye,
 Patoka Twp

 63 Crawford Unnamed tributary Patoka Lake, 1 mi NW Taswell,
 Patoka Twp

 64 Unnamed tributary Patoka Lake, 0.8 mi N Taswell,
 Patoka Twp

 65 Dog Creek, CR 33 bridge, 1.2 mi NE English,
 Sterling Twp

 66 Orange Unnamed tributary Patoka Lake, CR 650 W bridge, 1
 mi SW Greenbrier, Hoosier National Forest, Jackson
 Twp

 67 Young's Creek, CR 600 S bridge, Youngs Creek,
 Hoosier National Forest, Greenfield Twp

 68 Patoka River, SR 37 bridge, 1.5 on S Pine Valley,
 Southeast Twp

 69 Dubois/ Unnamed tributary Crane Creek, Cuzco-Norton Road
 Orange bridge, 3.25 mi SW French Lick, Columbia/Jackson
 Twp

 70 Orange Unnamed tributary Patoka River, 0.2 mi E Bacon,
 Hoosier National Forest, Southeast Twp

 71 Patoka River, CR 375 E bridge, 0.1 mi S Valeene,
 Southeast Twp

 72 Patoka River unnamed tributary, CR 825S bridge,
 Valeene, Southeast Twp

 73 Patoka River, CR 500 E bridge, 2 mi SSE
 Chambersburg, Stampers Creek Twp

 74 Spring Spring Cave, CR 475 path into State Forest,
 Stampers Creek Twp

 75 Gibson South Fork Patoka River, SR 57 bridge, 2 mi N
 Oakland City, Columbia Twp

 76 Patoka River, CR 1200 E bridge, 2.25 mi N Oakland
 City, Columbia/Logan Twp

 77 Patoka River, SR 57 bridge, 2.25 mi N Oakland
 City, Columbia Twp

 78 Patoka River, CR 1050 E bridge, 2 mi SE Oatsville,
 Columbia Twp

 79 Houchins ditch, CR 450 E bridge, 1 mi SW Wheeling,
 Washington Twp

 80 Patoka River, Old SR 41/CR 75 W bridge, Patoka,
 White River Twp

 81 Patoka River, US 41 bridge, access off Service
 Road, 1 mi W Patoka, White River Twp

 82 Patoka River, CR 890 W bridge, 1 mi NNE East Mt
 Carmel, White River Twp

 83 Patoka River, CR 850 E, 2 mi E East Mt. Carmel,
 White River Twp

 84 Patoka River, 350 m E CR 450 N, 2 mi SW Cuzco,
 Columbia Twp

 85 Yellow Creek, N CR 400 N, 1.2 mi E Wheeling,
 Washington Twp

 86 Cup Creek, CR 625 N, ca 0.25 mi W HWY 257, 0.5 mi
 W Pikeville, Lockhart Twp

 87 Dubois Hall Creek, S Celestine Road, 1.25 mi S Celestine,
 Hall Twp

 88 Ell Creek unnamed tributary, 200 m N SR 64 bridge,
 2 mi W Huntingburg, Patoka Twp

 89 Orange Patoka River, 0.5 mi E CR 150 W bridge

 90 Pike South Fork Patoka River, CR 300 E bridge, 2.25 mi
 SSE Coe, Monroe Twp

 91 Dubois Patoka River. CR 300 W, 5.5 mi S Jasper, Patoka
 Twp

 92 Short Creek, 200 m W SR 231, 4.25 mi S Jasper,
 Bainbridge Twp

 93 Hall Creek tributary, Hall Creek Road, 0.75 mi W
 SR 162, 1.6 mi SE Jasper, Bainbridge Twp

 94 Pike Flat Creek, 0.25 mi W Flat Creek Road

 95 Dubois Hunley Creek, 0.5 mi N SR 64, 3 mi W Huntingburg,
 Patoka Twp

 96 Gibson Patoka River, 0.5 mi N CR 400 N, 3 mi W Wheeling,
 Washington Twp

 97 Dubois Davis Creek, 200 m S SR 56, 2 mi N Cuzco, Columbia
 Twp

 98 Pike Cup Creek, 0.5 mi S SR 64, 1.5 mi S Pikeville,
 Lockhart Twp

 99 Dubois Flat Creek, 200 m S SR 64, 0.75 mi W St. Anthony,
 Jackson Twp

 100 Pike Bruster Branch, CR 200 N, 2 mi NE Winslow, Patoka
 Twp

 101 Gibson Patoka River, CR 350 N, 0..5 mi S Patoka, White
 River Twp
 102 Dubois Dillon Creek, 0.25 mi N Dubois-Cuzco Road at CR
 850 E, 1.5 mi SW Cuzco, Columbia Twp

 103 Gibson Indian Creek, CR 150 N bridge, 2.5 mi NE
 Princeton, Patoka Twp

 104 Pike Flat Creek tributary, CR 125 S bridge, 1 mi S
 Glezen, Washington Twp

 105 Flat Creek, CR 125 S bridge

 106 Sugar Creek

 107 Patoka River, Survant

 108 Patoka River, SR 257 bridge

 109 Hog Branch, Sugar Ridge State Park, 0.75 mi SW
 Survant, Marion Twp

 110 Unnamed tributary, SR 364 bridge, 1.5 mi SSW
 Winslow, Patoka Twp

 111 Gibson Wabash and Erie Canal, CR 50 N bridge, 2 mi N
 Francisco, Center Twp

 112 Pike Lick Creek, CR 300 S, 0.5 mi W Survant, Marion Twp

 113 Orange Patoka Lake

 114 Patoka Lake

 115 Patoka Lake

 116 Patoka Lake

 117 Dubois Patoka Lake

 118 Patoka Lake

 119 Huntingburg Lake

 120 Huntingburg Lake

 121 Beaver Dam Lake

 122 Pike New Oakland City Lake

 123 New Oakland City Lake

 124 Dubois Big Bottom wetland

 125 Orange Tucker Lake

 Site
number County Latitude (N) Longitude (W)

 1 Gibson 38.40711[degrees] 87.68587[degrees]
 2 38.41376[degrees] 87.50745[degrees]
 3 38.36307[degrees] 87.44113[degrees]
 4 38.36307[degrees] 87.39940[degrees]
 5 38.32655[degrees] 87.39950[degrees]
 6 38.33827[degrees] 87.33295[degrees]
 7 Pike 38.32513[degrees] 87.25730[degrees]
 8 38.28945[degrees] 87.26575[degrees]
 9 38.24602[degrees] 87.25209[degrees]
 10 38.30800[degrees] 87.28200[degrees]
 11 38.32667[degrees] 87.22233[degrees]
 12 38.30383[degrees] 87.22233[degrees]
 13 38.25217[degrees] 87.19350[degrees]
 14 38.31683[degrees] 87.08567[degrees]
 15 38.24600[degrees] 87.18967[degrees]
 16 38.31667[degrees] 87.12050[degrees]
 17 38.33100[degrees] 87.12500[degrees]
 18 38.34083[degrees] 87.22517[degrees]
 19 38.29300[degrees] 87.29400[degrees]
 20 38.28717[degrees] 87.21933[degrees]
 21 38.28150[degrees] 87.24233[degrees]
 22 38.28000[degrees] 87.24250[degrees]
 23 38.26470[degrees] 87.25000[degrees]
 24 38.39450[degrees] 87.22017[degrees]
 25 38.41950[degrees] 87.18817[degrees]
 26 38.39367[degrees] 87.18733[degrees]
 27 38.37550[degrees] 87.21950[degrees]
 28 38.35533[degrees] 87.23333[degrees]
 29 38.35550[degrees] 87.19333[degrees]
 30 38.36933[degrees] 87.18733[degrees]
 31 38.35583[degrees] 87.16683[degrees]
 32 38.36717[degrees] 87.16100[degrees]
 33 38.38033[degrees] 87.15550[degrees]
 34 Dubois 38.29433[degrees] 87.01183[degrees]
 35 38.31950[degrees] 86.99400[degrees]
 36 38.38583[degrees] 87.05850[degrees]
 37 38.40917[degrees] 87.09183[degrees]
 38 38.42350[degrees] 87.11433[degrees]
 39 38.42250[degrees] 87.06867[degrees]
 40 38.41500[degrees] 87.00417[degrees]
 41 38.46367[degrees] 86.92467[degrees]
 42 38.46717[degrees] 86.94717[degrees]
 43 38.34817[degrees] 86.88100[degrees]
 44 38.29933[degrees] 86.91533[degrees]
 45 38.35450[degrees] 86.90983[degrees]
 46 38.27433[degrees] 86.93217[degrees]
 47 38.28083[degrees] 86.86583[degrees]
 48 38.38750[degrees] 86.92800[degrees]
 49 38.48783[degrees] 86.75633[degrees]
 50 38.47233[degrees] 86.69633[degrees]
 51 38.47900[degrees] 86.65883[degrees]
 52 38.47050[degrees] 86.72200[degrees]
 53 38.45933[degrees] 86.74917[degrees]
 54 38.43550[degrees] 86.80417[degrees]
 55 38.42333[degrees] 86.87083[degrees]
 56 38.31433[degrees] 86.84383[degrees]
 57 38.30067[degrees] 86.80183[degrees]
 58 38.34400[degrees] 86.81217[degrees]
 59 38.35850[degrees] 86.65067[degrees]
 60 38.36666[degrees] 86.79733[degrees]
 61 38.33950[degrees] 86.74567[degrees]
 62 38.34633[degrees] 86.67833[degrees]
 63 Crawford 38.34967[degrees] 86.59366[degrees]
 64 38.34833[degrees] 86.54133[degrees]
 65 38.36617[degrees] 86.44283[degrees]
 66 Orange 38.44650[degrees] 86.57801[degrees]
 67 38.51583[degrees] 86.51567[degrees]
 68 38.44033[degrees] 86.45650[degrees]
 69 Dubois/ 38.49300[degrees] 86.68217[degrees]
 Orange
 70 Orange 38.40883[degrees] 86.42100[degrees]
 71 38.43517[degrees] 86.39667[degrees]
 72 38.43417[degrees] 86397500[degrees]
 73 38.48433[degrees] 86.37367[degrees]
 74 (see Indiana Cave Atlas)
 75 Gibson 38.37733[degrees] 87.33617[degrees]
 76 38.38317[degrees] 87.33283[degrees]
 77 38.38283[degrees] 87.33816[degrees]
 78 38.37800[degrees] 87.37067[degrees]
 79 38.40333[degrees] 87.46467[degrees]
 80 38.40233[degrees] 87.58633[degrees]
 81 38.39867[degrees] 87.59900[degrees]
 82 38.39700[degrees] 87.73000[degrees]
 83 38.37719[degrees] 87.38542[degrees]
 84 38.44116[degrees] 86.28818[degrees]
 85 38.41468[degrees] 87.45415[degrees]
 86 38.32717[degrees] 87.12247[degrees]
 87 Dubois 38.36598[degrees] 86.78764[degrees]
 88 38.30034[degrees] 86.98246[degrees]
 89 Orange 38.44104[degrees] 86.47832[degrees]
 90 Pike 38.27296[degrees] 87.20605[degrees]
 91 Dubois 38.33564[degrees] 86.96543[degrees]
 92 38.26234[degrees] 86.96615[degrees]
 93 38.36204[degrees] 86.87996[degrees]
 94 Pike 38.42045[degrees] 87.11727[degrees]
 95 Dubois 38.30687[degrees] 86.92697[degrees]
 96 Gibson 38.42058[degrees] 87.49355[degrees]
 97 Dubois 38.49864[degrees] 86.73064[degrees]
 98 Pike 38.30055[degrees] 87.11840[degrees]
 99 Dubois 38.30016[degrees] 86.80841[degrees]
 100 Pike 38.39312[degrees] 87.18423[degrees]
 101 Gibson 38.40180[degrees] 87.54927[degrees]
 102 Dubois 38.46348[degrees] 86.74559[degrees]
 103 Gibson 38.37806[degrees] 87.53528[degrees]
 104 Pike 38.40222[degrees] 87.30167[degrees]
 105 38.40183[degrees] 87.30067[degrees]
 106 38.39867[degrees] 87.27900[degrees]
 107 38.37250[degrees] 87.15567[degrees]
 108 38.32900[degrees] 87.11533[degrees]
 109 38.36722[degrees] 87.16083[degrees]
 110 38.35528[degrees] 87.23417[degrees]
 111 Gibson 38.36361[degrees] 87.44083[degrees]
 112 Pike 38.37528[degrees] 87.16444[degrees]
 113 Orange 38.29488[degrees] 86.60517[degrees]
 114 38.40171[degrees] 86.61315[degrees]
 115 38.41473[degrees] 86.67350[degrees]
 116 38.43208[degrees] 86.64675[degrees]
 117 Dubois 38.37540[degrees] 86.68725[degrees]
 118 38.37040[degrees] 86.61808[degrees]
 119 38.29488[degrees] 86.98229[degrees]
 120 38.29087[degrees] 86.97665[degrees]
 121 38.39849[degrees] 86.84378[degrees]
 122 Pike 38.31928[degrees] 87.32203[degrees]
 123 38.31757[degrees] 87.32003[degrees]
 124 Dubois 38.39538[degrees] 86.91330[degrees]
 125 Orange 38.48425[degrees] 86.56184[degrees]

Table 2.--Freshwater shrimp and crayfish species collected from the
Patoka River drainage. The numbers represent sites listed in Table 1,
while numbers in parentheses refer to the number of individuals
collected. The sequence of species folllows Simon (2001).

 Species Sites where species was collected

Family Palaemonidae (freshwater
 shrimp) 3(3), 36(1), 44(110), 46(11), 48(88),
 Palaemonetes kadiakensis 75(1), 76(9), 77(7), 78(18), 79(2),
 Rathbun, Mississippi grass 83(4), 84(1), 91(2), 95(2), 96(30),
 shrimp 101(2), 102(1), 107(11), 108(2)
Family Cambaridae (crayfish)
 Procambarus (Ortmannicus) 4(5), 44(3), 46(1), 47(1)
 acutus (Girard), White
 River crayfish
 Orconectes (Crockerinus) 41(2), 42(2), 81(2), 82(4)
 propinquus (Girard), north-
 ern clearwater crayfish
 O. (Faxonius) indianensis 16(2), 37(1), 43(11), 47(41), 48(38),
 (Hay), Indiana crayfish 49(53), 50(21), 51(106), 52(20),
 53(11), 54(2), 56(2), 57(26),
 58(11), 59(16), 60(3), 61(12),
 62(12), 63(36), 64(54), 66(9),
 67(30), 68(65), 69(63), 70(1),
 71(79), 72(2), 73(67), 77(1), 78(6),
 79(2), 80(1), 82(5), 86(1), 89(5),
 91(1), 93(1), 94(5), 97(8), 98(2),
 99(4), 101(8), 102(8)
 O. (Gremicambarus) immunis 6(4), 7(1), 14(5), 20(1), 24(2),
 (Hagan), papershell 28(15), 29(2), 31(5), 32(1), 33(3),
 crayfish 34(5), 35(5), 37(1), 39(18), 40(14),
 41(33), 42(11), 43(15), 45(29),
 46(173), 47(58), 49(15), 50(37),
 52(4), 55(41), 56(1), 57(28),
 58(11), 60(12), 61(1), 62(5), 80(2),
 81(1), 82(1), 84(1), 85(14), 86(3),
 87(7), 90(2), 92(8), 93(5), 96(2),
 97(5), 98(1), 99(3), 102(1), 107(1),
 108(8)
 O. (Orconectes) inermis 74(2)
 inermis Cope, Indiana cave
 crayfish
 Fallicambarus (Creaserinus) 5(1), 8(1), 15(1), 17(2), 20(2),
 fodiens (Cottle), digger 24(1), 29(1), 38(4)
 crayfish
 Cambarus (Erebicambarus) 67(5), 74(3)
 laevis Faxon, karst
 crayfish
 C. (E.) tenebrosus Hay, 31(3), 32(3), 42(1), 49(1), 51(6),
 cavespring crayfish 52(1), 53(6), 59(8), 62(1), 63(2),
 64(3), 66(12), 68(1), 69(14), 70(3),
 71(1), 72(1), 73(3)
 C. (Lacunicambarus) sp. A, 96(2), 100(1)
 great plains mudbug
 C. (Tubericambarus) sp. A, 1(3), 2(1), 3(1), 6(3), 7(1), 8(1),
 painted-hand mudbug 11(5), 13(3), 14(1), 15(3), 16(2),
 18(2), 20(2), 26(4), 27(5), 29(5),
 30(1), 31(2), 32(1), 35(1), 39(2),
 40(3), 41(1), 45(8), 46(6), 47(1),
 48(1), 49(2), 51(2), 53(6), 54(1),
 55(13), 56(1), 57(3), 58(11), 59(2),
 60(12), 62(3), 63(1), 67(1), 68(2),
 69(7), 70(2), 71(1), 73(5), 75(2),
 76(1), 77(4), 78(5), 79(7), 80(l),
 82(2), 88(1), 90(1), 93(2), 100(1),
 103(1), 105(6), 107(3), 108(1)


ACKNOWLEDGMENTS

We appreciate the assistance of several colleagues that assisted in the field collection of specimens especially, Foster Purrington, Ohio State University; Anthony Brannam, Natural Resource Conservation Service; Stacy Sobat, Indiana Department of Environmental Management; Charles Morris, Troy State University; and Thomas Simon IV and Cameron Simon. The U.S. Fish and Wddlife Service supported this study through grant number 31440-1261-3N29.

LITERATURE CITED

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Cope, E.D. 1872. Report on the Wyandotte Cave and its fauna. Third and Fourth Annual Report to the Geological Survey of Indiana. Pp. 157-182.

Evermann, B.W. & H.W. Clark. 1920. Lake Maxinkuckee, a physical and biological survey. Indiana Department of Conservation Publication 7. 620 pp.

Hay, W.P. 1893. Observations on the blind crayfishes of Indiana, with a description of a new subspecies; Cambarus pellucidus testii. Proceedings of the United States Natural History Museum XVI:283-286.

Hobbs, H.H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology 480. 236 pp.

Homoya, M.A., D.B. Abrell, J.R. Aldrich & T.W. Post. 1985. The natural regions of Indiana. Proceedings of the Indiana Academy of Science 94: 245-268.

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Lewis, J.J. 1983. The obligatory subterranean invertebrates of glaciated southeastern Indiana. National Speleolugical Society Bulletin 45:34-40.

Lewis, J.J., R. Burns & S. Rafail. 2002. The subterranean fauna of the Hoosier National Forest. U.S. Department of Agriculture, Forest Service. 115 pp.

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Omernik, J.M. & A.L. Gallant. 1988. Ecoregions of the Midwestern United States. EPA 600/3-88/ 037. U.S. Environmental Protection Agency. Corvallis, Oregon.

Packard, A.S. 1873. On the cave fauna of Indiana. Fifth Annual Report, Peabody Academy of Sciences, Salem. Pp. 93-97.

Page, L.M. 1985. The crayfishes and shrimps (Decapoda) of Illinois. Illinois Natural History Survey Bulletin 33:335-448.

Pflieger, W.L. 1996. The crayfishes of Missouri. Missouri Department of Conservation, Jefferson City.

Schneider, A.F. 1966. Physiography. Pp. 40-56, In Natural Features of Indiana. Indiana Academy of Science, Indianapolis, Indiana.

Simon, T.P. 2001. Checklist of the crayfish and freshwater shrimp (Decapoda) of Indiana. Proceedings of the Indiana Academy of Science 110:104-110.

Simon, T.P., S.A. Sobiech, T.H. Cervone & N.E. Morales. 1995. Historical and present distribution of fishes in the Patoka River basin: Pike, Gibson, and Dubois Counties, Indiana. Proceedings of the Indiana Academy of Science 104: 193-206.

Simon, T.P., R.L. Dufour & B.E. Fisher. 2003. Changes in the biological integrity of fish communities in the Patoka River drainage as a result of anthropogenic disturbance from 1888 to 2001. Pp. 245-257, In Changes in the Biology of Large Rivers. Historical Changes in Large River Fish Assemblages of North America. R.M. Hughes, J. Rinne, & R. Calmusso, (eds.). American Fisheries Society, Bethesda, Maryland.

Taylor, C.A. 1997. Taxonomic status of members of the subgenus Erebicambarus, genus Cambarus (Decapoda: Cambaridae), east of the Mississippi River. Journal of Crustacean Biology 17: 352-360.

Thoma, R.F. & R.F. Jezerinac. 2000. Ohio Crayfish and Shrimp Atlas. Ohio Biological Survey Miscellaneous Contribution 7. 28 pp.

Williamson, E.B. 1907. Notes on the crayfish of Wells County, Indiana, with the description of a new species. Pp. 749-763, In Annual Report to the Department of Geology and Natural Resources of Indiana.

Woods, A.J., J.M. Omernik, C.S. Brockman, T.D. Gerber, W.D. Hosteter & S.H. Azevedo. 1996. Ecoregions of Indiana and Ohio. Map Circular, U.S. Environmental Protection Agency, Corvallis, Oregon.

Manuscript received 24 March 2003, revised 12 September 2003.

Thomas P. Simon: U.S. Fish and Wildlife Service, 620 South Walker Street, Bloomington, Indiana 47403-2121 USA

Roger F. Thoma: The Ohio State University, Museum of Biological Diversity, Crustacean Range, 1351 Kinnear Drive, Columbus, Ohio 43212-1192 USA
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