U.S. FISH AND WILDLIFE SERVICE SPECIES ASSESSMENT AND LISTING PRIORITY ASSIGNMENT FORM
SCIENTIFIC NAME: Etheostoma maculatum Kirtland
COMMON NAME: spotted darter
LEAD REGION: Region 3
INFORMATION CURRENT AS OF: July 25, 2011
STATUS/ACTION
We determined that threats to spotted darter are not of sufficient imminence, intensity, or magnitude that would cause substantial losses of population distribution or viability. Therefore, it was not elevated to Candidate status.
ANIMAL/PLANT GROUP AND FAMILY: Fishes, Percidae
HISTORICAL STATES/TERRITORIES/COUNTRIES OF OCCURRENCE: New York, Pennsylvania, Ohio, Indiana, Kentucky, West Virginia
CURRENT STATES/COUNTIES/TERRITORIES/COUNTRIES OF OCCURRENCE: New York, Pennsylvania, Ohio, Indiana, Kentucky, West Virginia
LAND OWNERSHIP: Spotted darters are found in perennial streams. Ownership of the bottoms of streams varies; however, perennial streams are typically regulated as state and/or federal jurisdictional waters. Land use decisions throughout entire watersheds can affect fish populations within individual streams. Land ownership within each watershed occupied by spotted darters is variable and comprised of a mix of private, federal, state and other publicly owned land. There is no comprehensive database describing land ownership within the range of the spotted darter. However, the U.S. Forest Service maintains a database of ownership of forested land by county. Some available information for land ownership within the range of the spotted darter follows.
New York: No federally owned land occurs in the only county in the range of the spotted darter (Chautauqua County) (United States Forest Service 2008). Approximately 4.3 percent of the forested land in Chautauqua County is state-owned (U.S. Forest Service 2008).
Pennsylvania: Approximately 15.1 percent of forested areas in counties within the range of the spotted darter are in federal ownership (Forest Service land) and 6.3 percent is in state ownership (U.S. Forest Service 2008). Ownership of land adjacent to French Creek in Pennsylvania is almost exclusively private (Mayasich et al. 2004, p. 17).
Ohio: Approximately 0.7 percent of forested land in counties in the current range of the spotted darter is federally owned (agency owner unspecified) and 7.9 percent is state-owned (U.S. Forest Service 2008). In the Big Darby Creek watershed, an area targeted for conservation, the three 1 main land-owning conservation entities (Franklin County Metroparks, The Nature Conservancy (TNC), Ohio Department of Natural Resources (ODNR)) own approximately 4,047 ha (10,000 ac) of land (approximately 2.8 percent of the watershed) (Sasson 2008, pers. comm.).
Indiana: Approximately 7,821 hectares (ha) (19,325 acres (ac)) (4.1 percent) of the 190,202-ha (470,000-ac) Blue River watershed are in conservation ownership, including the Harrison/Crawford State Forest Complex along the lower section of the Blue River (Hauswald 2008, pers. comm.). Several federal and state properties are located in the East Fork White River watershed, including the Hoosier National Forest and Martin State Forest (Mayasich et al. 2003, p. 17).
Kentucky: With the exception of Mammoth Cave National Park, ownership of riparian areas adjacent to the Green and Barren rivers in the range of the spotted darter is almost entirely private (Mayasich et al. 2004, p. 17). Approximately 2.0 percent of forested areas in counties within the range of the spotted darter is federally owned (Department of Defense: 1.3 percent; Unspecified: 0.7 percent) (U.S. Forest Service 2008). The state owns no forested land (U.S. Forest Service 2008).
West Virginia: Approximately 1 percent of forested land in counties within the range of the spotted darter is in unspecified federal ownership and 3.1 percent is state-owned (U.S. Forest Service 2008). West Virginia streambeds are owned by the state (Mayasich et al. 2004, p. 17).
LEAD REGION CONTACT: Laura Ragan, 612-713-5157, [email protected]
LEAD FIELD OFFICE CONTACT: Jeromy Applegate, 614-416-8993 ext. 21, [email protected]
BIOLOGICAL INFORMATION
Species Description
The spotted darter is a member of the Perch family (Percidae), a group characterized by the presence of a dorsal fin separated into two parts, one spiny and the other soft (Kuehne and Barbour 1983, p. 1). Darters are smaller and more slender than other percids. Most darters, including those in the genus Etheostoma, have a vestigial swim bladder, which decreases buoyancy, allowing them to remain near the bottom with little effort (Evans and Page 2003, p.64).
Etheostoma is the largest and most diverse of the darter genera (Kuehne and Barbour 1983, p. 66). Species of Etheostoma differ from Percina spp. in possessing scales on the midline of their belly that are similar in shape and size to the scales on their flanks (Stauffer et al. 1995, p. 294). Species of the darter genera Crystallaria and Ammocrypta have more elongate bodies than Etheostoma spp. and have a naked midline on their belly (Bailey et al. 1954, p.140).
Distinguishing morphological characteristics of the spotted darter include: laterally compressed body, subequal jaws, sharp snout, short pectoral fins, an absent/weak suborbital bar, and a rounded posterior edge of the caudal fin (Zorach and Raney 1967, p. 300). They often exceed 60 2 millimeters (mm) (2.36 inches (in)) standard length (Kuehne and Barbour 1983, p. 116). The opercle and belly are scaled, the cheek is slightly scaled to unscaled, and the nape and breast are unscaled (Page 1983, p. 100). Lateral line counts are usually 56 to 65 scales, and vertebrae number 37 to 39 (Kuehne and Barbour 1983, p. 117). Spotted darters are sexually dimorphic. Males have black-edged red spots on the body and a bluish-green breast that intensifies in color at spawning time. Females have dark spots on the body that are larger and more diffuse than the males (Keuhne and Barbour 1983, p. 116).
Spotted darters superficially resemble bluebreast darters (E. camurum), but the two can be distinguished by the latter having a black margin on its soft dorsal, caudal, and anal fins (Stauffer et al. 1995, p. 304). Small spotted darters can resemble Tippecanoe darters (E. tippecanoe), but Tippecanoe darters have an incomplete lateral line (Stauffer et al. 1995, p. 304).
Taxonomy
The spotted darter was described as Etheostoma maculata by Kirtland (1841, pp. 276-277). Jordan and Eigenmann (1885, p. 71) emended the species epithet to maculatum to conform to the neuter gender of Etheostoma. The spotted darter was subsequently listed under the genera Etheostoma, Nothonotus, and Poecilichthys by various workers through the early 1950s. Bailey et al. (1954, pp. 139-141), and Bailey and Gosline (1955, pp. 6, 10) reduced the number of darter genera to three (Ammocrypta, Etheostoma, and Percina), placing the spotted darter in the subgenus Nothonotus. Three subspecies were subsequently recognized by Zorach and Raney (1967, p. 297): the spotted darter (Etheostoma maculatum maculatum) Kirtland in the Ohio River system including the Wabash and Green river systems, bloodfin darter (E. m. sanguifluum ) (Cope) in the upper Cumberland River system below Cumberland Falls, and wounded darter (E. m. vulneratum) (Cope) in the upper Tennessee River system. These subspecies have since been elevated to distinct species within the genus Etheostoma, subgenus Nothonotus: E. maculatum (spotted darter), E. sanguifluum (bloodfin darter) and E. vulneratum (wounded darter) by Etnier and Williams (1989, p. 987).
Preliminary results from genetics research suggest that all known populations of E. maculatum are appropriately described as a common species (Keck and Near 2008, pers. comm.; Porter 2008, pers. comm.). However, ongoing morphological assessments of the Elk River population may reveal characteristics that distinguish it as distinct from other E. maculatum populations (Welsh 2008, pers. comm.). Until and unless a morphological and genetics analysis of broader scope is completed and can demonstrate distinct differences, we will consider all populations of E. maculatum as a single species, and a valid taxon for consideration as a candidate.
Habitat/Life History
Darters typically exploit riffle habitat in streams, in part because shallow riffles contain few predators capable of eating darters (Kuehne and Barbour 1983, p. 1). Substrate heterogeneity in riffles creates “velocity shelters,” areas of low velocity used by darters (Harding et al. 1998, p. 995). Interstices of coarse riffle substrate also trap organic debris, which is processed by bacteria and fungi, creating an energy source for aquatic invertebrates, the preferred food type of most darters (Kuehne and Barbour 1983, p. 1). Darters are typically more selective feeders than other benthic stream fishes (Hansen et al. 1986, p. 68).
3
Spotted darters are habitat specialists that take advantage of their extremely laterally compressed body to live under and among large, heterogeneous, unembedded substrates in riffles and glides (Raney and Lachner 1939, pp. 157-159; Burr and Warren 1986, p. 306; Bowers et al. 1992, p. 19; Osier and Welsh 2007, p. 457; Kessler and Thorp 1993, p. 1090; Kessler et al. 1995, p. 368). They are associated with deeper water and larger rocks than similar species (Raney and Lachner 1939, p. 158; Kessler and Thorp 1993, pp. 1087-1089; Osier and Welsh 2007, p. 456). They typically do not tolerate silt or embedded substrates (Kessler and Thorp 1993, p. 1090; Osier and Welsh 2007, p. 457). In the Blue River, Indiana and the lower Allegheny River and upper Ohio River in Pennsylvania, spotted darters are found in the tailwaters of some dams, where higher velocities help maintain these conditions (Baker et al. 1985, pg. 605; Fisher 2008, pers. comm.; Spears 2008, pers. comm.). Slight winter movement into deeper areas within and adjacent to riffles has been observed (Raney and Lachner 1939, p. 158; Albin 2008, pers. comm.).
Spotted darters typically spawn in May and June (Raney and Lachner 1939, p. 160; Weddle and Kessler 2008, p. 21; Ruble et al. 2008, Appendix 2). Raney and Lachner (1939, p. 159) found that spawning sites were spaced at least 120 centimeters (cm) (47.24 in) apart in the head of a riffle in water 15-60 cm (5.9-23.62 in) deep. Up to 350 adhesive pale yellow 2 mm (0.079 in) diameter eggs were deposited in tight wedge-shaped masses on the undersides of 90-275 cm (35.43-108.27 in) diameter flat rocks (Raney and Lachner 1939, p. 161). Weddle and Kessler (2008, p. 22) found that egg clump dimensions averaged 20 mm (0.79 in) long by 13 mm (0.51 in) wide and were deposited under rocks averaging 24.7 cm (9.72 in) long and 18.2 cm (7.17 in) wide. Observations of up to five distinct egg size classes in females indicate that spotted darters spawn multiple times in a single season (Raney and Lachner 1939, p. 162; Weddle and Kessler 2008, p. 24). Male spotted darters guard the eggs while remaining mostly under or adjacent to the nest rock (Raney and Lachner 1939, p. 162). First spawning activity is reported to occur at two years for both males and females; males spawn through year four and females through year five (Raney and Lachner 1939, p. 164).
The species’ extremely pointed snout makes them well-adapted for picking macroinvertebrate prey from underneath rocks (Kessler et al. 1995, p. 368). Macroinvertebrates, especially larval insects, comprise a large portion of their diet. Larval midges (Diptera, family Chironomidae), stoneflies (Plecoptera), caddisflies (Trichoptera), mayflies (Ephemeroptera), and beetles (Coleoptera), as well as adult water mites (Hydracarina) are important food items (Raney and Lachner 193, p. 162; Hansen 1983, Appendix B; Kessler 1994, p. 29). Spotted darter eggs have been found in the stomachs of spotted darter adults (Raney and Lachner 1939, p. 162).
Historical Range/Distribution
The spotted darter historically occurred in the Ohio River drainage in New York, Pennsylvania, Ohio, Indiana, Kentucky and West Virginia (Table 1). In addition to the historical streams of record in Table 1, spotted darters probably occurred in other streams in the Ohio River basin with suitable habitat. Raney and Lachner (1939, p. 158) speculated that its presence had likely been overlooked by many collectors who had not thoroughly worked deeper riffles. In addition, small benthic fishes are difficult to collect in deeper water (Ohio Environmental Protection Agency (OEPA) 1988, p. 4-10). Troutman (1981, p. 670) noted that there may be considerable variation in the numbers of spotted darters in individual populations from one year to another, 4 although he did not discuss a cause for this phenomenon. These factors may help explain why spotted darters went undetected in the Elk, Blue, East Fork White, lower Allegheny, and Ohio Rivers until after 1975. Considering that many larger parent streams in the Ohio River Basin were extensively impounded and polluted beginning in the 1800’s, degrading or eliminating spotted darter habitat (Ortmann 1909, pp. 90-110; U.S. Army Corps of Engineers (USACE) 1981; Trautman 1981, pp. 17-24), it is reasonable to believe that the species also inhabited some of these parent streams historically but were extirpated prior to detection.
Current Range/Distribution and Population Estimates/Status
Range-wide status assessments in the literature indicate that spotted darters are localized and uncommon (Kuehne and Barbour 1983, p. 117; Page 1983, p. 100; Page and Burr 1991, p. 305). Although there is no range-wide systematic sampling to monitor distribution and status, a number of river-wide surveys have been conducted in some basins in some years. Available information on distribution and status in each major system is presented below and summarized in Table 1 and Figure 1.
Ohio River, Pennsylvania. Spotted darters were first discovered in the Ohio River in 2007 when 5 individuals were collected using benthic trawling collection methods below Dashields Locks and Dam, Allegheny County, Pennsylvania (Criswell 2008, pers. comm.). This discovery may represent a recent range expansion or the initial detection of an existing population established here for a number of years.
Allegheny River System, Pennsylvania and New York. Spotted darters are extant and considered uncommon in the middle Allegheny River mainstem between Franklin, Pennsylvania (Venango County) and Kinzua Dam (Warren County), however, there is little definitive population information available (Mayasich et al. 2004, p. 9). One 1983 record occurs for Sandy Creek, an Allegheny River tributary in Venango County, Pennsylvania (Fischer 2008, pers. comm.). They were first found in the lower Allegheny River below Locks and Dam No. 3 in 2007 using benthic trawling collection methods (Criswell 2008, pers. comm.; Spears 2008, pers. comm.). Similar to the Ohio River discovery, this discovery may represent a range expansion in this river, or reflect recent efforts to sample benthic fishes here.
Spotted darters are extant in the French Creek system in New York and Pennsylvania. Bowers et al. (1992, pp. 9-13) found spotted darters at 7 of 8 sites sampled in 1991-1992 in the New York portion of French Creek with population estimates that ranged from 0.03 to 0.33 individuals per square meter (m2). They concluded that the abundance and age structure indicated a healthy, self-supporting population (Bower et al 1992, pp. 18, 22). Subsequent sampling of 28 other sites in 1994 and 1995 did not reveal any additional spotted darter locations in New York portions of French Creek (Mayasich et al. 2004, p. 9) and limited additional sampling in 2010 detected spotted darter at only 1 location, near the Pennsylvania border (Carlson 2010, p. 1). Current estimates of spotted darter population trends in the French Creek system in New York are unknown (Mayasich et al. 2004, p. 9). In Pennsylvania, Smith and Crabtree (2005, pp. 149, 173) found spotted darters in “very low numbers” at 12 of 31 sites sampled in French Creek. The spotted darter population in the Pennsylvania portion of French Creek has generally been stable over the past decade, but has possibly declined some in Erie County (Mayasich et al. 2004, p. 9). A possible decline in Erie County is supported by Smith and Crabtree (2005, p. 149) who found 5 spotted darters at only 1 of 8 sites sampled in Erie County. In West Branch French Creek, spotted darters were discovered in 1985 in Erie County, Pennsylvania (Fischer 2008, pers. comm.) and additional individuals were captured in Chautauqua County, New York, in 1992 (Mayasich et al. 2004, p. 6). They are presumed extant in West Branch French Creek, but the current trends are unknown.
Beaver River System, Pennsylvania and Ohio. Spotted darters appear to be extirpated from the Beaver River system (Shenango River, Yellow Creek and Mahoning River) (Trautman 1981, p. 670; Mayasich et al. 2004, p. 6; Mishne 2008, pers. comm.).
Muskingum River System, Ohio. Spotted darters were last collected in the Walhonding River in 1999, despite subsequent collection efforts in that stream (Mayasich et al. 2004, p. 7; Kibbey 2008, pers. comm.). They may be declining or vulnerable in the Walhonding River.
Spotted darters are extant in the Kokosing River, a tributary to the Walhonding River. The Kokosing River population, although apparently restricted to an approximately 5-km reach of the river (Mishne 2008, pers. comm.; Applegate 2008, p. 1), appears to be healthy, as evidenced by large numbers and robust adults (Kibbey 2008, pers. comm.).
Scioto River System, Ohio. The spotted darter is extant in the middle portion of the Scioto River system in the mainstem and some tributaries, including Big and Little Darby Creeks, Walnut Creek, and Paint Creek. One individual was discovered in 1998 in the Scioto River mainstem near Big Darby Creek and another found in 2002 in the Scioto River near Big Walnut Creek (Mishne 2008, pers. comm.). It has recently expanded its range upstream in Big Darby Creek and Little Darby Creek (Yoder 2006, pers. comm.; Crail 2008, pers. comm.; Kibbey 2008, pers. comm.; Mishne 2008, pers. comm.). Six individuals were found for the first time in two locations in Walnut Creek in 2005 (Mishne 2008, pers. comm.), although limited subsequent sampling in Walnut Creek failed to produce additional specimens (Kibbey 2008, pers. comm.). Three individuals were captured in Paint Creek in 2003 (Crail 2008, pers. comm.) and one individual in 2006 (Mishne 2008, pers. comm.).
Although recent discoveries suggest that spotted darter range is expanding in a portion of the Scioto River basin, they have likely been extirpated from other streams in this system, including the Olentangy River, Big Walnut Creek, and Deer Creek (Mayasich et al. 2004, p. 6; Applegate 2008, pers. comm.; Kibbey 2008, pers. comm.).
Blue River, Indiana. Spotted darters were first discovered in the Blue River, a direct tributary of the Ohio River, in 1976 (Baker et al. 1985, p. 603). Sampling in the Blue River between 1999 and 2006 produced densities of 1-2 individuals per m2, with no appreciable change over time (Hauswald 2008, pers. comm.). They are considered abundant in areas of suitable habitat (Fisher 2008, pers. comm.).
Wabash River System, Indiana. Spotted darters were first recorded in the East Fork White River in 1997 (Mayasich et al. 2004, p. 8) and are present in the mainstem of the East Fork White River in Lawrence, Martin, Dubois, and Daviess Counties (Fisher 2008, pers. comm.). They are abundant in areas of suitable habitat in the East Fork White River (Fisher 2008, pers. comm.). 6
The Tippecanoe River population may be extirpated or extremely reduced (Fisher 2008, pers. comm.). Despite substantial effort to document spotted darter’s continued existence there, none have been verified since 1985 when four individuals were collected in Pulaski County (Carney et al. 1993, p. 209). Spotted darters are presumed extirpated from Deer Creek where they have not been found since 1888 (Fisher 2008, pers. comm.).
Green River System, Kentucky. The spotted darter is extant throughout approximately 161 kilometers (km) (100 miles (mi)) of the Green River mainstem from Locks and Dam No. 6 upstream past Green River Lake (Cicerello 2003, p. 9; Evans 2008, pers. comm.). As many as 25 individual spotted darters were collected in 1998 from the Green River near Munfordville, Kentucky (Cicerello 2003, p. 9). In 2002, a total of 46 adult and young-of-year specimens were collected or observed at 11 of 30 sampling sites in the Green River between Munfordville and Greensburg (Cicerello 2003, p. 9). Extensive searches below Locks and Dam No. 5 and Locks and Dam No. 6 on the Green River have not detected spotted darters (Evans 2008, pers. comm.).
Several tributaries in the upper Green River system, including Little Barren River, South Fork Little Barren River, Russell Creek, Big Pittman Creek, and Meadow Creek also harbor extant populations. Up to 10 individuals were collected at each of five sites in the lower 39 km (24 mi) of the Little Barren River and as many as 74 individuals were collected per site on the lower 32 k (20 mi) of Russell Creek (Cicerello 2003, p. 9; Evans 2008, pers. comm.). One spotted darter was collected in Big Pittman Creek in 1999, the first record in that stream since 1962. Spotted darters were discovered in Meadow Creek in 2001 when three individuals were found (Cicerello 2003, p. 9; Evans 2008, pers. comm.).
Cicerello (2003, p. 10) concluded that the spotted darter was relatively common and evenly distributed in the Green River system above Locks and Dam No. 6; however, he noted that it has not been collected from the Green River mainstem downstream of Mammoth Cave National Park, despite the presence of suitable habitat.
A total of nine individuals have been captured from the Barren River system (tributary of the Green River downstream of Locks and Dam No. 5) since 1990. Five individuals were collected in 1990 from the mainstem Barren River below Locks and Dam No. 1, one individual was collected in 2001 from the Gasper River, and three individuals were collected in 2001 from West Fork Drakes Creek (Cicerello 2003, p.8; Evans 2008, pers. comm.). Cicerello (2003, p. 8) speculated that the Gasper River population is limited to a rather small area around the collection site (in Warren County), and noted that several subsequent samplings of West Fork Drakes Creek failed to produce spotted darters. He concluded that the spotted darter inhabits a fraction of its historical range in the Barren River basin, as it is currently restricted to West Fork Drakes Creek, lower Gasper River, and the mainstem Barren River downstream of Locks and Dam No. 1. It is extirpated from Line Creek and Trammel Fork, where it has not been found since 1961 and 1970, respectively (Cicerello 2003, pp. 23-24; Zorach and Raney 1967, pp. 299-300).
Kentucky River System, Kentucky. The most recent record from this system is a single individual collected in the North Fork Kentucky River in 1981 (Cicerello 1984, p. 158). Subsequent sampling efforts have failed to capture additional specimens (Evans 2008, pers. comm.). Sampling efforts in Greasy Creek, a headwater tributary of the North Fork Kentucky 7 River, have also failed to produce additional spotted darters since they were discovered there in 1960 (Evans 2008, pers. comm.). Spotted darters are likely extirpated from the Kentucky River system.
Licking River System, Kentucky. Spotted darters have not been found in the Licking River system since before 1892 and appear to be extirpated from this system (Evans 2008, pers. comm.).
Kanawha River System, West Virginia. Spotted darters were first discovered in the Elk River in 1978 immediately above Sutton Lake (Cincotta et al. 1986, pp. 115-116). They were first collected downstream of Sutton Lake in 1991 (Mayasich et al. 2004, p. 6; Osier 2005, p. 75). They are extant in the middle portion of the Elk River below Sutton Lake, and may be expanding their range there (Osier 2005, p. 75, Cincotta 2008, pers. comm.; Welsh 2008, pers. comm.). A single individual was also collected in the lower Elk River in 2009, suggesting further range expansion downstream (Welsh 2011, pers. comm.). No individuals have been retaken at the initial collection site above the reservoir, despite several attempts in the 1990s (Mayasich et al. 2004, p. 6).
Distribution Summary
The spotted darter is considered extant in the mainstem Ohio River (PA) and in the Allegheny (NY, PA), Muskingum (OH), Scioto (OH), Blue (IN), Wabash (IN), Green (KY), and Kanawha (WV) river systems. Of the 37 known streams that historically supported or currently support spotted darters, the species is likely extant in 24, likely extirpated in 12, and potentially extirpated in 1. Of the 24 streams that currently support spotted darters, populations are likely stable or expanding in 9 and declining or vulnerable in 4. Recent trends are unknown in the remaining 11 streams with extant populations. Fourteen of the 24 extant populations were discovered after 1975, and 9 of these 14 were discovered after 1990. Given the recent discoveries of new populations of spotted darters, and considering the potential difficulties in collecting them (see Historical Range/Distribution section), it is reasonable to believe that they may also be present, but have gone unrecorded, in other streams within the river systems discussed herein.
THREATS
Anthropogenic threats to aquatic life in the Ohio River basin were recognized a century ago by Ortmann (1909, pp. 90-110). More recently, Jelks et al. (2008, p. 377) reported that 700 fish taxa in continental North America are imperiled, in large part from negative anthropogenic impacts. This represents a 92 percent increase since 1989 and a 179 percent increase since 1979 (Jelks et al. 2008, pp. 381, 382). Although a portion of this increase is attributable to finer scales of taxonomy and revised interpretation of species concepts, 33 percent of previously recognized taxa moved to a more severe imperilment category between 1989 and 2008 and 16 are now extinct (Jelks et al. 2008, p. 382). Forty-four percent of the 191 described North American percid species are considered imperiled (Jelks et al. 2008, p. 379). The spotted darter was categorized by the authors as threatened, a more severe level of imperilment than its special concern status in 1989. No details were given regarding this change in status.
8 A. The present or threatened destruction, modification, or curtailment of its habitat or range.
Habitat degradation is the leading cause of imperilment of North American fishes (92 percent of imperiled taxa) (Jelks et al. 2008, p. 381). Due to their geographically restricted distribution, freshwater fishes are highly susceptible to extirpation from localized habitat degradation through impoundment, siltation, and stream flow modification (Warren et al. 2000, p. 8). Prior research indicates that many darter species, including the spotted darter, are sensitive to anthropogenic disturbance (OEPA 1988, pp. 4-10, B-10). The spotted darter’s association with large, clean substrates suggests that the species may be especially sensitive to changes in chemicophysical stream characteristics. The isolated and small apparent size of some remaining populations (e.g., Barren River System, KY, and Walhonding River, OH) makes those populations more susceptible to local extirpation from habitat perturbation.
Sedimentation. Sedimentation is the addition of fine soil particles (e.g., sands, silts, clays) to streams. It is a significant stressor leading to stream impairment in the United States (U.S. Environmental Protection Agency (USEPA) 1998, p. 32; 2000, p. 61; 2002, p. 13; 2007, p. 13; 2009, p. 11). Agriculture, the leading source of stream impairment (USEPA 1998, p. 32; 2000, p. 62; 2002, p.14; 2007, p. 10; 2009, p. 12), is a major contributor of sediment to streams (Waters 1995, pp. 17-23), especially where activities such as row-crops and livestock grazing encroach into riparian areas (Waters 1995, p. 17; Smith and Crabtree 2005, p. 174). Other common sources of sedimentation include road construction, urban development, and resource extraction and transport (e.g., coal, oil, gas, timber), especially in riparian zones (Waters 1995, pp. 23-51). Sediment can enter streams via overland flow, which is often exacerbated when a dirt road provides a conduit (Reed and Carpenter 2002, p. 575), or through stream bank erosion. Stream bank erosion is 30 times more prevalent in streams lacking riparian vegetation and is further compounded where livestock have unrestricted access to streams (Naiman and Decamps 1997, p. 630).
Sedimentation modifies aquatic habitats by reducing light penetration, altering heat radiation, increasing turbidity, and covering the stream bottom, filling interstitial spaces and reducing depth (Ellis 1936, p. 41; Waters 1995, pp. 67-69). These changes disrupt reproductive behavior and alter food resources utilized by stream fishes (Berkman and Rabeni 1987, pp. 291-292). Berkman and Rabeni (1987, p. 292) found that increased siltation in Missouri streams significantly reduced the number of benthic insectivores, including darter species. Powell (2003, p. 35) concluded that substrate embeddedness resulting from the cumulative effects of sedimentation was the greatest single factor impacting stream fish communities in areas of high crop-land density. Smith (1968, p. 37) concluded that siltation was a principal cause for extirpation of 2 fish species and range reduction of 14 others in Illinois between 1905 and 1971.
Spotted darters require relatively deep runs and riffles (22 cm (8.7 in) on average) with cobble and boulder substrates of high complexity and abundant interstitial space for refugia, foraging, courtship, and egg laying, incubation, and guarding (Raney and Lachner 1939, pp. 157-159; Kessler and Thorp 1993, p. 1090; Kessler et al. 1995, p. 368; Osier and Welsh 2007, p. 457). Sedimentation and resulting substrate embeddedness greatly reduce the complexity and abundance of interstitial spaces (Waters 1995, p. 118) and consequently, the amount and complexity of important microhabitat (Kessler and Thorp 1993, p. 1090; Osier and Welsh 2007, p. 457-458). Reduction in refugia makes darters more susceptible to predation. In addition, they 9 must expend greater amounts of energy maintaining their position if velocity refugia are limited. Because spotted darters lay their eggs on the undersides of rocks, reproduction can be impacted by any reductions in interstitial space (Powell 2003, p. 34; Bowers et al. 1992, p. 1) or interstitial flow rates (which leads to reduced dissolved oxygen (D.O.), affecting egg survival) (Waters 1995, p. 86). A reduction in interstitial spaces also reduces the diversity and abundance of aquatic invertebrate prey (Cobb and Flannagan 1990, pp. 35-37; Powell 2003, pp. 34-35) and elevated turbidity reduces foraging efficiency of site-feeders, such as the spotted darter (Waters 1995, p. 83). Darters are typically more selective feeders than other benthic stream fishes (Hansen et al. 1986, p. 68), and therefore, would reasonably be expected to be less resistant to reduction in diversity of aquatic invertebrates than other benthivores, which could more easily switch prey type.
In French Creek, New York, Bowers et al. (1992, p. 22) listed increased turbidity and siltation from poor agricultural and silvicultural practices as a primary threat to French Creek spotted darters. They also hypothesized that spotted darters in New York may be more susceptible to environmental perturbations because New York is at the edge of their range (p. 21), but concluded that French Creek had a healthy, self-supporting population (pp. 18, 22). The New York Department of Environmental Conservation (NYDEC) (2007, p. 5) listed agriculture as a significant threat to the Allegheny watershed, in part from poor management practices including unrestricted livestock access to streams and intensively cultivated crop lands with little riparian buffer. However, no lotic portions of the French Creek system were listed as impaired (NYDEC 2007, p. 99). In addition, Crabtree (2008, pers. comm.) reports that much of the French Creek watershed is forested and sedimentation by isolated sources is largely attenuated by the time it reaches the mainstem.
Natural gas drilling, especially from recently exploited Marcellus Shale reserves, represents a potential expanding source of sediment in the Allegheny River watershed. Marcellus Shale drilling requires large amounts of support necessary for the creation of permeable flow paths for natural gas movement using hydraulic fracturing techniques, including moving large amount of vehicles, equipment, and supplies, often into remote areas (Soeder & Kappel 2009, pp. 2, 4). These activities may cause increased erosion and sediment overload, which could pose a risk to small watersheds (Kargbo et al. 2010, p. 5681). The overall effect to spotted darters downstream in the Allegheny River mainstem, however, is unclear.
In the Scioto River system, Ohio, siltation is a documented threat to fish in the Big Darby Creek watershed (OEPA 2004, p. A.29). The majority (63 percent) of the watershed is agricultural, while the eastern portion is “under immediate or imminent development pressure from the Columbus metropolitan area” (OEPA 2004, p. A.10). The Ohio EPA documented sedimentation impacts to fish communities, resulting from a lack of adequate erosion control and stormwater management during road and suburban construction in portions of the headwaters of the Big Darby Creek system (OEPA 2006, pp. A.64, A.84). The report also noted threats to the system that included removal of riparian vegetation and free access of cattle to streams (OEPA 2004, p. A.72). Despite the documented sedimentation threat, spotted darters are expanding their range in the Big Darby Creek system. It is possible that sedimentation in headwaters may be somewhat attenuated downstream in the mainstem of the Big and Little Darby Creeks. Trautman (1981, p. 31) also noted a “noticeable decrease in the amount of silt deposited” in a section of Big Darby Creek inhabited with historical and current spotted darter populations over the 40-year period 10 preceding 1980. It is possible, however, that further range expansion may be impeded in areas of the watershed where sedimentation is prevalent, such as the rapidly developing eastern portion near Columbus.
The Blue River watershed, Indiana, is composed of 50 percent agriculture (27 percent pasture and 23 percent row crop) (Hauswald 2008, pers. comm.). Row crop agriculture is viewed as a threat to the Blue River, especially in the upper watershed, where lack of riparian buffer is still common (Hauswald 2008, pers. comm.). In lower portions of the watershed, some discrete areas are subject to direct access by cattle (Hauswald 2008, pers. comm.). Similar to the Big Darby Creek system, however, the effects of sedimentation have not resulted in recent population declines in the Blue River.
The Tippecanoe River watershed, Indiana, is intensively farmed with a high density of row crops in riparian areas (Watts 2008, pers. comm.; McWilliams-Munson and Simon 2001, p. 2.14). Portions of the Tippecanoe River watershed are impaired due to sedimentation from agricultural land uses (Commonwealth Biomonitoring 2003, p. 35), which likely contributed to the spotted darter’s decline and possible extirpation from the Tippecanoe River (Simon 2008, pers. comm.). Sedimentation also contributed to the extirpation of spotted darters from Deer Creek (Simon 2008, pers. comm.). Land use in the East Fork White River watershed, Indiana, is primarily agricultural in the northern part and varies (e.g., forestry, livestock, oil and gas production) in the southern portion (Dufour 2000, p. 2). Any adverse effects of sedimentation on East Fork White River spotted darters from these land uses are not evident, as their populations are considered relatively stable in this stream.
Livestock have access to streams in portions of the upper Green River watershed, Kentucky, where approximately 40 percent of land-use is agricultural (Kentucky Department of Environmental Protection (KDEP) 2008, p. 8). In 2002, 11,219 farms in counties partially or entirely within the upper Green River watershed produced 578,486 cattle (KDEP 2008, pp. 9- 12). Oil and gas extraction in the Green River watershed has increased dramatically with the demand for domestic energy supplies (Kessler 2008, pers. comm.). In the counties within the range of the spotted darter, the number of oil and gas permits issued between January 1, 2006 and September 29, 2008 (1164 permits) was greater than the total number of permits issued in the decade prior to 1996 (1,012 permits) (Kentucky Geological Survey 2008a). Sedimentation from these sources may be contributing to the Barren River system population declines, where the species has been extirpated from two historical streams of record and reduced in range and abundance in the remaining three streams. .
Coal mining is prevalent in the upper Kentucky River watershed (Kentucky Geological Survey 2008b), including in the North Fork basin where it has been a chronic contributor of high sediment loads (Bradfield and Porter 1990, p. 12). Sedimentation from mining and other sources likely contributed to the apparent extirpation of spotted darters from this system.
Coal mining is also a contributor of sediment to the largely forested (90 percent) middle Elk River watershed, West Virginia (Strager 2008, p. 13). As of 2003, there were 510.5 ha (1,261.5 ac) of abandoned mine land in the middle Elk River watershed and 124 mine sites that were in forfeiture of their bond for failure to properly reclaim the site (Strager 2008, p.18). Overall, there are 4,857 ha (12,000 ac) of active mine permits, 582.9 km (362.2 mi) of haul roads, 1,304 11 ha (3,223 ac) of valley fills, and 426 permits to release pollutants (including sediment) associated with coal mining in the Elk River watershed (Strager 2008, p.18). Recent exploitation of Marcellus Shale natural gas reserves has contributed to increased oil and gas extraction in the Elk River watershed, where over 3,600 active wells are located (Taylor 2011, pers. comm.). Logging activities, which contribute sediment to streams, have also been observed at riparian sites in the Elk River watershed (West Virginia Department of Environmental Protection (WVDEP) 1997, p. 59). Similar to some other systems within the range of the species, Elk River spotted darter populations appear to be stable or expanding, and specific effects of sedimentation to spotted darters in this system are not clear.
Perhaps as important as chronic sediment release from coal mines are the large-scale mine waste (coal slurry) impoundment structures. Two major slurry impoundments are located in the middle Elk River watershed. The Big Branch facility, which is located less than 16 km (10 mi) upstream of extant populations of spotted darters, includes a 79-meter (260-foot) dam and a capacity of 6,258,023 cubic meters (m3) (221,000,000 cubic feet (ft3)) of slurry (Strager 2008, p. 22). Structural failure of these impoundments would result in a massive release of mine waste into nearby streams, smothering stream bottoms. For instance, in October 2000, a coal slurry impoundment near Inez, Kentucky breached, releasing almost 991,090 m3 (35,000,000 ft3) of slurry into the Big Sandy Creek Watershed. “The slurry left, fish, turtles, snakes and other aquatic species smothered as the slurry covered the bottoms of the streams and rivers and extended out into the adjacent floodplain” (USEPA 2001, p. 2). Over 161 km (100 mi) of stream were impacted by the spill (USEPA 2001, p. 2). If a similar failure occurs in the Elk River watershed, it could have dramatic impacts on spotted darters.
Despite the documented and wide ranging threat of sedimentation to aquatic life, especially for habitat specialists like spotted darters, some spotted darter populations, even those in areas with land uses frequently identified as contributing high sediment loads to streams (e.g., Big Darby Creek watershed: 63 percent agriculture with expanding suburban development), appear to be stable or expanding. This may be partially explained by an overall reduction in sedimentation in the last couple of decades. The U.S. Department of Agriculture (USDA) (2006, pp. 5-6) reports that soil erosion on cultivated cropland in the Ohio River watershed decreased 53 percent between 1982 and 2003, due to an increase in conservation tillage and decrease in total cultivated cropland, including a 36 percent decrease in cultivation of highly erodible lands. The decrease in cultivated land can be attributed, at least in part, to agricultural programs such as the Conservation Reserve Program (CRP), begun in 1986, and the Conservation Reserve Enhancement Program (CREP), started in 1996. Both programs provide financial incentives to farmers and ranchers to remove land from agricultural production and plant trees, grass, and other vegetation along streams. The latter program provides additional financial incentives for these activities in watersheds of local and national significance. As of 2009, over 526,091 ha (1,300,000 ac) of land were enrolled in CRP in the six states in the range of the spotted darter (USDA 2010a) and CREP programs covered the Ohio River watershed in Pennsylvania and the Scioto River, East Fork White River, and Green River watersheds (USDA 2010b). Yoder et al. (2005, pp. 8, 21) found improvements in fish communities of almost all non-wadeable rivers throughout the state of Ohio since 1990, in part because of conservation farming practices that reduce sedimentation.
12 Sedimentation from coal mining has likely decreased from historical levels, primarily due to requirements and restoration programs adopted following passage of the Surface Coal Mining and Reclamation Act of 1977 (SMCRA). The Act requires environmental performance standards for coal mining operations and bonding to ensure post-mining reclamation. In addition, the Act created the Abandoned Mine Land fund to fund abandoned mine reclamation, as many abandoned sites continue to be stream sedimentation sources. Since 1977, almost seven billion dollars of funding has been distributed to states and tribes to remediate safety and environmental hazards at abandoned mines in the United States (U.S. Office of Surface Mining 2010).
In addition, after initial large scale tree clearing and agricultural development following European settlement, some watersheds have reverted to forest to a large degree. From 1890 to 1920, for example, forests in the Allegheny Plateau in Pennsylvania, had been almost completely clear-cut (Marquis 1975, p. 11). Much of this forest (64 percent of the Allegheny River watershed (White, et al. 2005, p.423)) has now regenerated and is managed in some manner, thereby limiting the excessive erosion that had historically impacted streams in this system. In addition, special stormwater management requirements are in place in some watersheds (e.g., Big Darby Creek) in an attempt to reduce the effects of sedimentation associated with development (OEPA 2007, pp. 1-45).
Sedimentation has contributed to historical extirpations of some spotted darter populations (e.g., Deer Creek, North Fork Kentucky River) and to recent declines in parts of the range (e.g., Tippecanoe River, Barren River system). Sedimentation, however, does not appear to be causing widespread declines throughout the range of the species, as spotted darter populations appear to be stable or expanding in 9 of the 13 streams where enough information is available to determine current status. This may be due in part to the effects of environmental legislation and programs targeting erosion problems, and reforestation of some previously clear-cut watersheds, which has helped reduce sedimentation from historical levels. It is reasonable to believe that these improvements have contributed to stabilization of population size in some systems (e.g. Blue River) and range expansion in some others (e.g., middle Scioto River system). Overall, the best available information does not indicate that sedimentation will cause substantial losses of population distribution or viability in all or a significant portion of the spotted darter’s range.
Impoundment. Artificial impoundment structures (dams) greatly reduce stream flow upstream, drastically increasing sediment deposition in the impounded reaches, thereby eliminating availability of coarse substrates that spotted darters require (Baxter 1977, p. 260; Bhowmik and Adams 1989, pp. 17-18). Impoundment may also lead to thermal stratification of the water column, which can result in a hypolimnion (bottom thermal layer) with greatly reduced D.O. concentrations (Baxter 1977, p. 261; Poff and Hart 2002, p. 660). Spotted darters evolved in oxygen rich glide-riffle environments, and presumably cannot live and reproduce successfully in oxygen-scarce habitat. Elimination of oxygen-rich, fast-flowing, coarse-substrate habitat also greatly reduces the abundance and diversity of benthic invertebrate food sources (Santucci et al. 2005, pp. 986-987). Dams generally make the areas within the impoundment uninhabitable by spotted darters.
Dams also fragment stream habitat by blocking fish movement, creating a physical barrier and in some cases long reaches of unsuitable habitat through which darters can or will not migrate. 13 This blocks movement among populations and restricts recolonization from source populations (Benstead et al. 1999, pp. 662-664; Poff and Hart 2002, p. 660). As a result, recolonization may be unavailable as a buffer to counter local extirpations caused by environmental perturbations or demographic stochasticity (Lande 1988, p. 1458).
Reservoir operation that results in variable water temperature and flow-rates can preclude spotted darter recolonization below some dams (Mayasich et al. 2004, p. 13). However, it has been hypothesized that darters can persist in riffle habitats downstream from impoundments if velocity and oxygen concentrations are sufficient (Baker et al. 1985, p. 605) and spotted darters have been found below dams that do not actively regulate discharge from the impoundments (e.g., in the Blue River, IN, Allegheny River, PA, and Ohio River, PA).
Dams have been constructed throughout the range of the spotted darter. Some have led to population extirpations and many currently restrict range expansion and recolonization of unoccupied suitable habitat. Kinzua Dam on the Allegheny River in Warren County, Pennsylvania, represents the upper extent of spotted darter range on this river. The area within the dam pool is now unsuitable for spotted darters and recolonization of suitable upstream reaches is blocked by the dam. The downstream range of the spotted darter in the Allegheny River is also likely somewhat limited by navigational dams that were constructed in the 1920s and 1930s, however, the species has been discovered in the lower Allegheny River and upper Ohio River in recent years.
There are nine lowhead dams in the Walhonding River system (ODNR 2010a) that alter riffle habitat and may restrict movement of spotted darters. In the Scioto River system, six major water supply or flood control reservoirs and many low head dams (18 in Franklin County alone), restrict migration of spotted darters from the middle Scioto system to other parts of the watershed with suitable habitat (ODNR 2010a).
A dam at Milltown, Indiana divides the Blue River spotted darters into two populations, likely limiting upstream and downstream movements (Fisher 2008, pers. comm.). In the East Fork White River, Williams Dam (Lawrence County) represents the upstream extent of spotted darters in this stream and prevents range expansion upstream (Fisher 2008, pers. comm.).
Reservoirs created by Barren and Green River dams in Kentucky almost certainly destroyed spotted darter habitat (Cicerello 2003, p. 7). Construction of Barren River Reservoir in 1964 caused extirpation of spotted darter in the Barren River upstream of the dam (Cicerello 2003, p. 7) and remaining populations in these watersheds are now isolated and unable to recolonize formerly occupied habitat blocked by the impoundments (Mayasich et al. 2004, p. 13).
Sutton Lake on the main channel of the Elk River, West Virginia may have played a role in the spotted darter’s extirpation immediately above the impounded reach (Mayasich et al. 2004, p. 12). Suitable habitat is present above the reservoir (WVDEP 1997, p. 67), however, Sutton Lake Dam blocks any recolonization of these areas by middle Elk River populations.
Removal of some existing dams is occurring throughout the range of the spotted darter. American Rivers (2009) reports that approximately 800 dams have been removed across the United States, 358 since 1999. Thirty-five dams were removed or planned to be removed in 14 2008 and 2009 in streams throughout Pennsylvania (American Rivers 2008; 2009). Seven lowhead dams have been removed in the Scioto River system since 1989 (ODNR 2010b). Following the 1989 removal of one of these on Little Darby Creek, spotted darter range expanded upstream of the location of the former dam. Although dam removal is occurring within the range of the spotted darter, dam construction continues as well. The USACE (2010) reports that 2,290 dams have been constructed in the United States since 2000, 146 in states within the range of spotted darter. There are a total of 9,001 dams in states within the range of the spotted darter. It is not known how many of these dams are on perennial streams with the potential to support spotted darters.
Stream impoundment has contributed to historical range reduction of spotted darters (e.g., in the Green and Barren Rivers, in the Upper Elk River) through direct habitat loss. In addition, dams currently impede full range expansion of spotted darters by restricting darter migration to formerly occupied habitats (e.g., Elk River, East Fork White River, Scioto River system). However, given the apparent stability and documented and potential range expansion of spotted darters in several stream systems, the best available information does not indicate that stream impoundment will cause substantial losses of population distribution or viability in all or a significant portion of the spotted darter’s range.
Other habitat modifications. Channelization is any combination of widening, straightening, and deepening streams, and often includes removal of riparian vegetation (Brooker 1985, p. 63). It is usually conducted for drainage improvements and flood control, but also as part of stream relocation during road construction, mining, or residential development. The effects of channelization and riparian vegetation removal on stream fish communities are well documented. Etnier (1972, pp. 373-375) found some fish species, including darters, declined or disappeared from a Tennessee stream following channelization and attributed the changes to a loss of macroinvertebrates caused by substrate instability and habitat homogeneity. Ebert and Filipek (1988, p. 29) documented smaller substrates and decreased biodiversity in channelized reaches of a third-order Ozark stream. Sensitive darters like the spotted darter are usually one of the first species to disappear from streams as biodiversity declines (OEPA 1988, pp. 4-10, B-10). Removal of riparian vegetation, which frequently accompanies channelization, increases siltation, reduces cover for aquatic species, increases water temperature and reduces input of high-quality coarse particulate organic matter (CPOM) (Sabeter et al 2000, p. 614; Naiman and Decamps 1997, p. 632). Decreases in riparian vegetation can also increase algal production (Sabeter et al. 2000, p. 614) (see Chemical Pollutants section below for implications of increased algae).
Channelization typically results in the loss of stream length, which contributes to flashier hydrographs (higher peak flows during rain events and lower base flows during dry periods) (Brooker 1985, p. 63). These effects are often exacerbated in watersheds with a high rate of wetland loss and a high density of impervious surfaces (e.g., parking lots, roads, roofs) (Paul and Meyer 2001, p. 335; Booth and Jackson 1997, p. 1080; USEPA 1997, p. 2). As little as 10 percent impervious cover in a watershed can modify hydrographs, resulting in erosion, channel instability and widening, substrate alteration, in-stream and riparian habitat loss, and loss of fish populations (Booth and Jackson 1997, p. 1084). Low base flows usually favor tolerant species, to the exclusion of intolerant forms (including spotted darter) (Rankin and Armitage 2004, pp. 26-27). Smith (1971, pp. 10-11) attributed the range reduction of 12 species of fish in Illinois, 15 including darters, to low base flows during some years.
Although channelization frequently occurs in headwater streams too small for spotted darters, impacts to headwater streams can result in indirect impacts to the species. Meyer et al. (2007, p. 99) stated, “The cumulative impact of degraded headwaters contributes to the loss of ecological integrity in ecosystems downstream.” Rankin and Armitage (2004, p. 34) suggest that headwater streams are biodiversity “cold spots,” in that they may be rather low in biodiversity, but they control ecological functions that strongly influence downstream biodiversity. Headwater streams play an important role in processing CPOM into forms usable by downstream biota. For example, leaf-shredding insects commonly dominate the macroinvertebrate fauna in intact headwater streams in the eastern U.S., and the fine particulate organic matter (FPOM) that shredders generate is exported as seston (suspended particles) to support food webs downstream (Vannote et al. 1980, p. 132). Diverse forms of organic matter are critical in supporting an abundant and varied insect food source for spotted darters. Headwater streams can also export the insects themselves to larger streams where they are sources of food for fish (Wipfli and Gregovich 2002, p. 966). Cuffney et al. (1990, p. 292) demonstrated that seston export to downstream ecosystems declined 67 percent following removal of aquatic insects from a headwater stream. Channelization and riparian vegetation removal degrade or eliminate CPOM sources and macroinvertebrate habitat in headwater streams (Brooker 1985, p. 64).
Channelization and riparian vegetation removal occur in parts of the spotted darter’s range. Bowers et al. (1992, p. 22) listed stream channel alterations as one of two main anthropogenic threats to spotted darters in French Creek, New York. In the Big Darby Creek, Ohio, a large 1997 flood prompted some landowners to dredge and bulldoze streambed sediments to re- establish damaged dikes, causing impact to local aquatic communities in the vicinity of spotted darter populations (OEPA 2004, p. A.58). Ohio EPA also reports recent direct channelization of portions of upper Little Darby Creek and suspects that as these impacts accumulate they will result in the eventual partial or non-attainment of the current aquatic life uses (OEPA 2004, p. A.59). Watts (2008, pers. comm.) reports that headwater streams in the Tippecanoe River are extensively channelized to improve agricultural drainage and that row-crops extend to the edge of the stream throughout much of the watershed. Commonwealth Biomonitoring (2003, p. 18) found habitat at some sites in the Indian Creek watershed, a tributary to the Tippecanoe River, Indiana “hampered by a paucity of stable bottom substrate and instream cover, by a lack of any riparian buffer zone, and by channelization.” Kessler (2008, pers. comm.) expressed concern about the long term health of spotted darters in the Green River watershed because of increased stormwater runoff (altered hydrograph) resulting from development and increased gas and oil production in the region. Heavy channelization and dredging has also been documented on some tributaries to the Elk River (WVDEP 1997, p.59).
Other habitat modifications such as channelization and altered hydrographs can adversely affect spotted darters and have likely contributed to population declines in some stream systems (e.g., Tippecanoe River), and may be slowing range expansion in others (e.g., Big Darby Creek). However, given the apparent stability and documented and potential range expansion of spotted darters in several stream systems, the best available information does not indicate that these threats will cause substantial losses of population distribution or viability in all or a significant portion of the spotted darter’s range.
16 Chemical Pollutants. Chemical pollution can cause direct mortality to sensitive species and, at sub-lethal levels, can increase vulnerability to other threats (Maitland 1995, p. 260). Major sources of aquatic pollutants include domestic wastes, agricultural runoff, and industrial discharges, all of which have been identified as threats to spotted darters. Reash and Berra (1987, p. 117) found that polluted streams in Ohio contained a simplified fish community, characterized by the absence of pollution intolerant species, including the fantail darter. Polluted sites generally supported habitat and diet generalists that were able to tolerate degraded environments.
Untreated and poorly treated municipal wastewater (sewage), and livestock waste, especially where livestock have unrestricted stream access, are common sources of chemical pollution. Sewage and livestock waste contain chemical contaminants that include ammonia, pathogenic bacteria, nutrients (e.g., phosphorous and nitrogen), and organic matter that increases Biological Oxygen Demand (BOD) (Cooper 1993, p. 405). Biological Oxygen Demand is a measure of the oxygen consumed through aerobic respiration of microorganisms that break down organic matter in the sewage or livestock waste. Nutrients and BOD may have the greatest impact on spotted darters because they cause decreases in D.O. Elevated levels of nutrients lead to excess algal growth. Nocturnal respiration of live algae and decomposition of dead algae consumes oxygen (Cooper 1993, p. 405). Low D.O. may be particularly harmful to spotted darters, which live and lay eggs under bottom substrates, where D.O. can be significantly lower than surface water (Whitman and Clark 1982, p. 653). Adequate oxygen is an important aspect of egg development (Sowden and Power 1985, p. 808). Excessive algal growth can also lead to physical alterations of habitat when it covers bottom substrates (Cooper 1993, p. 405).
Although sewage treatment has improved dramatically since the 1972 amendments to the Federal Water Pollution Control Act (Clean Water Act), some sewage treatment plants continue to experience maintenance and operation problems that lead to poorly treated sewage (OEPA 2006, p. 2-67). In addition, untreated domestic sewage (straight piping) (KDEP 2008, p. 8) and poorly operating septic systems (Hauswald 2008, pers. comm.) are still present in some watersheds in the range of spotted darters. In the Allegheny watershed, agricultural runoff from silage leachate, manure or milk-house wastewater, and improper manure application on fields has been noted as a source of nutrients (NYDEC 2007, p. 5). Rankin and Armitage (2004, p. 31), however, indicate that background nutrient levels in French Creek are much lower than typical effects levels that they have observed in other streams, suggesting that effects to spotted darters in French Creek may be minimal. Crabtree (2008, pers. comm.) also reports that fairly low overall agricultural land use in the Allegheny/French Creek watershed limits most nutrient impacts to localized stream segments. In portions of the Big Darby Creek watershed, pollution from poorly treated or untreated sewage was cited as a factor in water quality impairment (OEPA 2006, pp. 2- 68, 2-70). Yoder et al. (2006, p. 8), however, conclude that the expansion of spotted darter in the middle Scioto River system, including Big Darby Creek, is attributable mainly to chemical water quality improvements required by the Clean Water Act. In the Tippecanoe River drainage, excessive nutrients from agricultural sources have caused stream impairment (Commonwealth Biomonitoring 2003, p. 36). In the Blue River drainage, Hauswald (2008, pers. comm.) reports that failing septic systems are impacting water quality, and new residential development is anticipated to contribute additional impacts. In the Green River drainage, straight pipes discharge untreated domestic sewage into some streams while some others are impaired because of pathogens from sewage treatment plants (KDEP 2008, p. 8). Livestock access to streams in 17 portions of the Blue and Green River drainages systems may also contribute to nutrient enrichment (Hauswald 2008, pers. comm.; KDEP 2008, p. 8).
Unpermitted releases of contaminants (i.e., spills) have the potential to impact fish species, including the spotted darter. For instance, OEPA (2004, p. A.45) documented 22 major spills in the Big Darby Creek watershed between 1979 and 2002, resulting in 147,587 fish killed. The largest spill was a 194,000-gallon sewage release. The second largest was a mixture of fermenting grain and molasses (~24,000 gal). The substantial BOD from the material resulted in anoxic and hypoxic conditions for several miles downstream for approximately one week, killing an estimated 24,000 fish and other aquatic organisms. Half of all spills, which caused 96 percent of the total fish kill, were directly attributable to agricultural sources.
Oil and gas extraction can contribute chemical pollution to streams. Production of commercial quantities of natural gas, especially from the Marcellus Shale reserves, requires large volumes of water for proper bit cooling, rock cutting removal, and creation of permeable flow paths for natural gas movement using hydraulic fracturing techniques (Soeder & Kappel, 2009, pp. 2, 4). Estimates place water usage for Marcellus wells at between 500,000 to several million gallons of water (Harper, 2008, pp. 11-12). Water used during natural gas drilling must be recovered and disposed (Soeder & Kappel 2009, p. 4). The waste water may contain contaminants (e.g., chemical additives, shale materials, including brines, heavy metals, radionuclides) and must be treated prior to disposal (Soeder & Kappel 2009, pp. 4-5). Salts and other dissolved solids in brines, however, are not usually removed successfully by wastewater treatment, and reports of high salinity in some Appalachian rivers have been linked to the disposal of Marcellus Shale brines (Renner 2009a, p. 6120). Inadequately treated Marcellus Shale wastewater has been implicated in the large Prymnesium parvum (commonly referred to as golden algae) bloom on Dunkard Creek, a Monongahela River tributary (Renner 2009b, p. 9046). The bloom caused the death of thousands of fish, mussels, and salamanders in a 48-km (30-mi) section of the creek (Renner 2009b, p. 9046).
In the range of the spotted darter, oil and gas extraction is prevalent and expanding in the Allegheny River watershed in Pennsylvania, due primarily to the recent exploitation of Marcellus Shale natural gas reserves. Bradfield and Porter (1990, p. 4) report that oil and gas operations are prevalent in the upper Kentucky River watershed, and that many streams support only pollution-tolerant organisms. In the Green River watershed, oil and gas operations have increased significantly over the last several years (see Sedimentation section for further details). However, Cicerello (2003, p. 9) stated that the collection of a single spotted darter in the Green River tributary Big Pittman Creek in 1999, the first since 1962, indicated that the fish fauna is recovering from past brine pollution there, which may indicate improved waste water management practices. Extraction from the Marcellus Shale reserves has also contributed to an increase in oil and gas wells in the middle Elk River watershed, where there are over 3,600 active oil and gas wells (Taylor 2011, pers. comm.).
Coal mining can also contribute to stream pollution, primarily through acid mine drainage (AMD), which is created from the formation of sulfuric acid in the oxidation of iron-sulfide minerals such as pyrite (Sams and Beer 2000, pp. 3-5). Acid mine drainage may react with compounds in adjacent rock and mine spoil to produce high concentrations of aluminum, manganese, zinc, and other constituents. Precipitated iron and aluminum oxides can be toxic to 18 benthic invertebrates and fish. In addition, high acid levels can be acutely and chronically toxic to aquatic life (Erichsen Jones 1962, pp. 258-259). The properties of water coming from mines and effects of AMD at a watershed scale depend greatly on underlying geology. Implementation of SMCRA (see Sedimentation section), and development of practices in the mid 1980’s to reduce AMD chemical reactions, have resulted in overall improvement in water quality of completed coal mines.
The Northern Appalachian Coal Field covers much of western Pennsylvania and extends into the range of the spotted darter in the upper Allegheny River watershed and southern portion of French Creek watershed in Pennsylvania (Anderson et al. 2000, p. 20). Crabtree (2008, pers. comm.), however, reports that active and historical coal mining is mostly limited to watersheds downstream of the majority of extant spotted darter populations in the French Creek/Allegheny River watershed, and that mining activities do not pose a current threat to these populations. However, ongoing acid mine drainage is a problem in the Allegheny River watershed downstream of the current spotted darter range, and may be limiting expansion of spotted darter range in portions of the Allegheny River system (Crabtree 2008, pers. comm.).
In the upper Kentucky River watershed where the spotted darter is extirpated, historical and current coal mining activities, including unreclaimed abandoned mines, are prevalent (see Sedimentation section for further details) (Kentucky Geological Survey 2008b). Bradfield and Porter (1990, p. 12) noted that the effects of mining were evident in North Fork Kentucky River from Whitesburg to Jackson, but that some small tributary streams were relatively unaffected.
Coal mining is widespread in the Elk River watershed (see Sedimentation section). The middle Elk River watershed also contains 426 NPDES permits to release pollutants associated with coal mining. Currently, the quality of the Elk River and its tributaries are being negatively affected by acidic drainage from mines that were abandoned before modern mining laws (USEPA 2001, p. 1-1). Portions of the Elk River mainstem (including those within the range of the spotted darter) and some tributaries, many a short distance upstream of extant spotted darter populations, have been impaired because of metals (including iron, aluminum, lead, and zinc) and low pH caused by mining (Strager 2008, p.36; USEPA 2001 p. 1-1). Several tributaries of Buffalo Creek, which enters the Elk River near extant spotted darter populations, are affected by mine drainage (pH and metals violations) and have impaired benthic communities (WVDEP 1997, pp. 55-57).
Industrial effluent is or has in the past been a threat to spotted darters. Troutman (1981, p. 27), for example, cites pollution from steel mills on the Mahoning River, beginning in 1925, as the cause for extirpation of spotted darters there. Simon (2008, pers. comm.) cited metal toxicity in sediments as a threat to spotted darters in the Tippecanoe River, and polychlorinated biphenols (PCBs) in sediment as a threat to the East Fork White River spotted darters, although he noted that PCB inputs have been curtailed and some existing PCB contaminated sites have been remediated.
Chemical pollutants have contributed to the extirpation of some spotted darter populations (e.g., Mahoning River, North Fork Kentucky River) and may be restricting the full range expansion of some others (e.g., Allegheny River system). It is not certain how chemical pollutants are affecting extant populations, however, given the apparent stability and documented and potential 19 range expansion of spotted darters in several stream systems, the best available information does not indicate that chemical pollutants will cause substantial losses of population distribution or viability in all or a significant portion of the spotted darter’s range.
Summary of Factor A
Although few quantitative data have been gathered directly linking the effects of sedimentation, impoundment, chemical water quality, and other habitat modifications on spotted darter declines, the best available information strongly suggests that these factors resulted in historical extirpations of some populations (e.g., Mahoning River, Deer Creek, North Fork Kentucky River). The data also suggest that these factors have contributed to recent declines in parts of the range, such as the Tippecanoe River, where the species is potentially extirpated, and the Barren River system, where spotted darters have been extirpated from two historical streams of record and reduced in range and abundance in the remaining three streams. Sedimentation, impoundment, chemical water quality, and other habitat modifications, however, have not been linked to recent widespread declines throughout the range of the species. It is likely that the effects of environmental legislation such as the Clean Water Act and SMCRA, and conservation programs including CRP and CREP have contributed to improvements in chemical water quality and habitat quality in many stream systems with remaining extant populations of the species. It is reasonable to believe that these improvements have contributed to stabilization of population size in some systems (e.g. Blue River) and range expansion in some others (e.g., middle Scioto River system). In addition, the relatively intact (i.e., heavily forested) composition of some watersheds helps ameliorate the effects of activities that degrade stream quality (e.g., in the Allegheny River watershed). Overall, the best available information does not indicate that the present or threatened destruction, modification, or curtailment of the spotted darter’s habitat or range will cause substantial losses of population distribution or viability in all or a significant portion of its range.
B. Overutilization for commercial, recreational, scientific, or educational purposes.
Overutilization of spotted darters for commercial, recreational, scientific, or educational purposes is not known to be a significant threat to spotted darters. Kessler (2008, pers. comm.), however, suggested the need to coordinate collection among researchers, especially in heavily sampled locations, to avoid the potential for over-collection.
C. Disease or predation.
There is no significant known threat from disease or predation. Some natural predation by piscivorous fish and wildlife occurs (Page 1983, p. 172). Commonly reported parasites of darters include metacercarial trematodes (black-spot disease) flukes, nematodes, leeches, spiny- headed worms, and copepods (Page 1983, p. 173).
D. The inadequacy of existing regulatory mechanisms.
Clean Water Act. Significant improvements in chemical water quality have occurred nationwide following the 1972 revisions to the Clean Water Act, primarily due to improvements in wastewater treatment (Andreen 2004, p. 538). However, streams throughout the U.S. remain 20 impaired. The most recent (2004) National Water Quality Report to Congress reports that 44 percent of assessed streams were impaired or did not meet their designated uses (USEPA 2009, p. 1). Non-point sources of pollution (e.g., sedimentation) and habitat degradation continue to be identified as significant water quality stressors (USEPA 2002, p. 13; 2007, p. 9; 2009, p. 11). In addition, evolving case law and lack of clear definition regarding what bodies of water are under Clean Water Act jurisdiction (e.g., Solid Waste Agency of Northern Cook County (SWANCC) v. U.S. Army Corps of Engineers, 531 U.S. 159 (2001); Rapanos v. United States, 547 U.S. 715 (2006)), may lead to inadequate stream protection, particularly in the headwaters.
State Endangered Species Laws. Except for West Virginia, all states within the range of the spotted darter have legislation that provides protections for rare animal species. The spotted darter is on the State list of protected species in New York, Pennsylvania, and Ohio. Of these three, only the New York law extends protection beyond prohibiting the possession, sale, transportation, or killing of listed species. The New York law also prohibits any alteration of occupied habitat that is likely to negatively affect one or more essential behaviors of such species (6 NYCRR, Part 182). Except for in New York, State threatened and endangered species laws do not address the primary threat to spotted darters: the present or threatened destruction, modification, or curtailment of its habitat or range.
In summary, existing regulatory mechanisms, including the Clean Water Act and State endangered species laws provide some protection to spotted darters, but do not mitigate all threats, especially those emanating from habitat degradation and nonpoint source pollution. However, these threats, discussed under Factor A, do not appear to be causing substantial losses of population distribution or viability in all or a significant portion of the spotted darter’s range
E. Other natural or manmade factors affecting its continued existence.
Genetic Variation. Some spotted darter populations appear to be small and isolated. Individuals in small populations are more likely to suffer from decreased fitness (i.e., ability to produce viable offspring) as inbreeding among close relatives occurs and results in greater expression of deleterious recessive genes (Allendorf and Luikart 2007, pp. 306, 315). Genetic drift (i.e., random change in gene frequencies) is also more likely to result in reduced genetic diversity in small populations, which may cause loss of genes that could allow the population to adapt to environmental change. These factors can increase the likelihood of extirpation (Allendorf and Luikart 2007, p. 355). The specific effects of genetic isolation on population dynamics in extant spotted darter populations, however, are not clear.
Climate Change. Climate change is expected to result in rising average temperatures throughout the range of the spotted darter and altered precipitation patterns, likely resulting in elevated stream temperature regimes and lower summer base-flows (Karl et al. 2009, pp. 107, 111-112, 117-118). Higher stream temperatures may result in reduced reproductive success and low base flows favor more tolerant stream fishes. Migration of spotted darters as an adaptation to climate changes is unlikely, due to their limited mobility, restriction to defined stream systems, and extensive impoundment throughout the Ohio River basin. According to the NatureServe Climate Change Vulnerability Index, release 2.01, spotted darters are considered moderately vulnerable to climate change, which means their abundance and/or range extent are likely to decrease by 2050 (Applegate 2010). Specific impacts to spotted darters resulting from 21 climate change are not clear.
In summary, both limited genetic variation and the effects of climate change are potential threats to spotted darter. However, the available information is not adequate to determine specific impacts to the species, or to identify either as a significant threat affecting its viability.
Summary and Finding
Threats to stream fishes are widespread throughout much of the spotted darter’s range. Because it is a habitat specialist that requires large unembedded substrate in riffle and glide habitat, the spotted darter can be particularly susceptible to threats that result in destruction, modification, or curtailment of its habitat or range. Although it is a wide ranging species, historical records indicate that it likely was never common, probably because of its specialized habitat requirements. Threats including sedimentation, impoundment, chemical pollutants, and other habitat modifications likely contributed to extirpation of spotted darters in parts of its range, and to population declines in some others. However, improvements in chemical water quality and habitat quality, in part because of environmental legislation and programs, have also likely contributed to the recent stabilization and/or range expansion in several stream systems.
Of the 24 streams with extant populations, enough available data exists to allow an assessment of current population trends in 13 streams. Of these, 9 have either stable or expanding populations, while 4 are declining or vulnerable. In addition, the discovery of 9 populations since 1990 and the documented difficulty in detecting the species suggest that they may also be present in other streams but have gone undetected. Further range expansion in some stream systems is reasonable to predict given current trends, but full range expansion to areas of suitable habitat is restricted by the presence of dams throughout every stream system within the range of the species.
In summary, the available evidence indicates that spotted darters are a wide ranging but uncommon species with relatively stable populations in much of its current range. Our review of the best available scientific and commercial information pertaining to the five factor threat analysis does not indicate that there are threats of sufficient imminence, intensity, or magnitude that would cause substantial losses of population distribution or viability of spotted darters. Therefore, we do not believe that the spotted darter is in danger of extinction (endangered), nor is it likely to become endangered in the foreseeable future (threatened) throughout all or a significant portion of its range. Accordingly, we have decided not to elevate spotted darter (Etheostoma maculatum) to Candidate status under the Federal Endangered Species Act of 1973 (as amended).
CONSERVATION MEASURES PLANNED OR IMPLEMENTED
No known conservation measures are planned or implemented specifically for the spotted darter. However, several general conservation efforts to improve water quality and aquatic habitat are planned or are being implemented for watersheds in the range of the spotted darter. The following are some known conservation measures within the range of the spotted darter.
Allegheny River System. The Nature Conservancy maintains an office on French Creek to direct 22 outreach and conservation actions. In addition, conservation measures are often implemented for development projects affecting spotted darter habitat in Pennsylvania, through the State Environmental Review system and Pennsylvania Natural Diversity Inventory (Fischer 2011, pers. comm.).
Scioto River System. Franklin County Metroparks, TNC, and ODNR own approximately 4,047 ha (10,000 ac) of land in the Big Darby Creek Watershed (Sasson 2008, pers. comm.), which is managed primarily for habitat conservation and stream quality improvements. The Franklin County Metroparks and TNC are actively acquiring additional properties (Morrow 2008, pers. comm.). In addition, recent and ongoing implementation of the Big Darby Total Maximum Daily Load recommendations, strict stormwater discharge rules, and the Big Darby Accord (multijurisdictional development plan emphasizing watershed protection) should help reduce future threats (Sasson 2008, pers. comm.).
Blue River. Hauswald (2008, pers. comm.) reports that The Nature Conservancy owns or manages approximately 405 ha (1000 ac) of the 190,202-ha (470,000-ac) Blue River watershed. They have planted approximately 221 ha (545 ac) of trees along 24 km (15 mi) of riparian buffer. They are also working with the city of Salem on upgrades to its wastewater treatment plant and the Washington County Soil and Water Conservation District to develop a watershed plan. Private forest management is also being implemented to minimize disturbance.
Green River. A joint project involving TNC, Southern Illinois University, and Eastern Kentucky University is modifying water releases from Green River Lake Dam on a trial basis to determine if they will benefit the downstream aquatic community (Mayasich et al. 2004, p. 17, 18).
RECOMMENDED CONSERVATION MEASURES
Regular evaluation of extant range and population status. Reintroduction in its historical range where suitable habitat is present. Augmentation of extant populations where appropriate. Riparian protection where existing land uses affect habitat. Dam removal, where feasible. Minimization of watershed-scale cumulative stressors such as channelization and sedimentation. Improved communication among researchers to prevent over-collection. Broad genetics research on a large number of individuals to assess phylogenetic structure and quantify degree of gene flow and movement between populations.
COORDINATION WITH STATES
All states within the range of the spotted darter provided information and comments that contributed to the candidate assessment.
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34 Table 1. Summary of historical (< 1985) records of the spotted darter and current distribution and hypothesized status. Current status designations are based on number of locations recorded, number of individuals collected, and date of last collection relative to amount of known sampling effort in each stream, as well as professional opinions of researchers and natural resource managers working in the range of the species.
State System River Counties of Historical Record (Year) Current Status Extant Ohio Ohio None (discovered 2007) a (possible range expansion) Extant PA Allegheny Erie (1975) b; Crawford (1966) b; Venango (1968) b (possible range expansion) Extant Sandy Creek Venango (1983) b (recent trends unknown) Allegheny NY: Chautauqua (1937, 1951) c Extant French Creek PA: Erie (1935, 1938) c; Crawford (1935, 1959) c; (recent trends unknown) NY/PA Mercer (1935, 1938) c; Venango (1935) c NY: None (discovered 1992) d Extant W. Branch French Creek PA: None (discovered 1985) b (recent trends unknown) PA Shenango River Mercer (1934-1936) c Extirpated Beaver Mahoning River Mahoning (≤ 1841) e Extirpated Yellow Creek Mahoning (1859) c Extirpated Walhonding Coshocton (1960, 1961 and 1963) f Declining/Vulnerable Muskingum Kokosing Coshocton (1963) f Extant Scioto None (discovered 1998) g Extant in Small Portion Olentangy River Franklin (1958 and 1960) f Extirpated OH Big Walnut Creek Franklin (1897, 1958-1960, 1962) c Extirpated Walnut Creek None (discovered 2005) g Extant Scioto Pickaway (1930, 1943, 1947, 1948, 1956, 1957, Big Darby Creek Expanding Range 1960, 1962, 1973-1976) f Little Darby Creek None (discovered 2006) g Expanding Range Deer Creek Ross (1956) f Extirpated Paint Creek None (discovered 2006) g Extant IN Blue Blue River Crawford, Harrison (1976-1983) h Stable
35 Tippecanoe River Fulton (1888 i, 1890 c, 1899 c) Possibly Extirpated IN Wabash Deer Creek Carroll (1888) i Extirpated E. Fork White River None (discovered 1997) j Stable Extant upstream of Locks and Green River Green (1963) c Dam 6, Stable Little Barren River Green (1956) c Extant South Fork Little Barren None (discovered 2006) k Extant River Russel Creek None (discovered 1991) k Stable Big Pittman Creek Green (1962) l Extant Green l Meadow Creek None (discovered 2001) Extant KY Extirpated upstream of L&D #1, Barren River Allen (1959) c; Monroe (1961) c Declining/Vulnerable below L&D #1 Gasper River Warren (1956 c, 1982 k) Declining/Vulnerable W. Fork Drakes Creek Simpson (1970) l Declining/Vulnerable Line Creek Monroe (1961) c Extirpated Trammel Fork Allen (1970) l Extirpated N. Fork Kentucky River Breathitt (1981) m Extirpated Kentucky Greasy Creek Leslie (1960) k Extirpated Licking S. Fork Licking River Harrison (<1892) k Extirpated Extant WV Kanawha Elk River Webster (1978) n (possible range expansion in middle Elk River)
(a) Criswell 2008, pers. comm.; (b) Fischer 2008, pers. comm.; (c) Zorach and Raney 1967, pp. 299-300; (d) Mayasich et al. 2004, p. 6; (e) Kirtland 1841, pp. 276-277 (Type locality); (f) Kibbey 2008, pers. comm.; (g) Mishne 2008, pers. comm.; (h) Baker et al. 1985, p. 603; (i) Jordan 1890; (j) Fisher 2008, pers. comm.; (k) Evans 2008, pers. comm.; (l) Cicerello 2003, pp. 23-24; (m) Cicerello 1984, p. 158; (n) Cincotta et al. 1986, p. 115-116
36 37