JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA AND TENNESSEE, USA: A FEATURED COLLECTION1 Carl E. Zipper, Braven Beaty, Gregory C. Johnson, Jess W. Jones, Jennifer Lynn Krstolic, Brett J.K. Ostby, William J. Wolfe, and Patricia Donovan2 ABSTRACT: The Clinch River of southwestern Virginia and northeastern Tennessee is arguably the most important river for freshwater mussel conservation in the United States. This featured collection presents investigations of mussel population status and habitat quality in the Clinch River. Analyses of historic water- and sediment-quality data suggest that water column ammonia and water column and sediment metals, including Cu and Zn, may have contributed historically to declining densities and extirpations of mussels in the river’s Virginia sections. These studies also reveal increasing temporal trends for dissolved solids concentrations throughout much of the river’s extent. Current mussel abundance patterns do not correspond spatially with physical habitat quality, but they do correspond with specific conductance, dissolved major ions, and water column metals, suggesting these and/or associated constituents as factors contributing to mussel declines. Mussels are sensitive to metals. Native mussels and hatchery-raised mussels held in cages in situ accumulated metals in their body tissues in river sections where mussels are declining. Organic compound and bed-sediment contaminant analyses did not reveal spatial correspondences with mussel status metrics, although potentially toxic levels were found. Collectively, these studies identify major ions and metals as water- and sediment-quality concerns for mussel conservation in the Clinch River. (KEY TERMS: aquatic ecology; freshwater mussels; major ions; metals; water quality; water resources management.) Zipper, Carl E., Braven Beaty, Gregory C. Johnson, Jess W. Jones, Jennifer Lynn Krstolic, Brett J.K. Ostby, William J. Wolfe, and Patricia Donovan, 2014. Freshwater Mussel Population Status and Habitat Quality in the Clinch River, Virginia and Tennessee, USA: A Featured Collection. Journal of the American Water Resources Association (JAWRA) 1-13. DOI: 10.1111/jawr.12220 important freshwater systems of North America and is one of the most important rivers for mussel diversity and conservation in the world. Here, we introduce a featured collection of articles describing investigations of freshwater mussel population status BACKGROUND The Clinch River of southwestern Virginia and eastern Tennessee is among the most ecologically 1 Paper No. JAWRA-13-0110-P of the Journal of the American Water Resources Association (JAWRA). Received May 7, 2013; accepted April 3, 2014. © 2014 American Water Resources Association. This article is a U.S. Government work and is in the public domain in the USA. Discussions are open until six months from print publication. 2 Professor (Zipper) and Geospatial Laboratory Specialist (Donovan), Department of Crop and Soil Environmental Sciences, Research Specialist (Ostby), Department of Fish and Wildlife Conservation, 416 Smyth Hall, Virginia Tech, Blacksburg, Virginia 24061; Aquatic Biologist (Beaty), Clinch Valley Program, The Nature Conservancy, Abingdon, Virginia 24210; Hydrologist (Johnson), Tennessee Water Science Center, U.S. Geological Survey, Knoxville, Tennessee 37921; Restoration Biologist (Jones), U.S. Fish and Wildlife Service and Department of Fish and Wildlife Conservation, Virginia Tech, Blacksburg, Virginia 24061; Geographer (Krstolic), Virginia Water Science Center, U.S. Geological Survey, Richmond, Virginia 23228; and Supervisory Hydrologist (Wolfe), Tennessee Water Science Center, U.S. Geological Survey, Nashville, Tennessee 37211 (E-Mail/Zipper: [email protected]). JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION 1 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, DONOVAN cal Services Strategic Plan (2010-2014) (USFWS, 2012). The first mussel surveys in the Clinch River were conducted in the late 1800s and early 1900s by Adams, Walker, and Ortmann, and were compiled by Ortmann (1918). These surveys identified a diverse mussel fauna that was already experiencing declines due to river system stresses. Ortmann (1918) expressed concern that sedimentation and toxins from logging, tanneries, and other land uses threatened the mussel fauna. Qualitative assessments in the 1970s continued to show a rich mussel community in the Clinch River (Stansbery, 1973; Bates and Dennis, 1978). Quantitative riverine mussel surveys have been conducted regularly at approximately fiveyear intervals since 1979. Subsequent analyses have revealed continuing population declines throughout most of the Clinch River in Virginia but generally stable populations in Tennessee (Jones et al., 2014). Mussel biodiversity conservation challenges in the Clinch River are complicated by the complex and multistage life history of mussels that generally requires a parasitic intermediate stage on fish before an offspring can survive on its own (McMahon, 1991). Freshwater mussels spawn via males releasing sperm into the water and females collecting sperm during siphoning, requiring adequate population densities for gamete exchange. Freshwater mussels begin life as fertilized eggs that develop into an embryonic form called glochidia within the female’s gills. Once mature, these glochidia must be released from the female mussel, attach to a host fish, and encyst into the fish tissue. There is a substantial degree of taxonomic specificity between mussel and host fish species. The developing mussels disperse as encysted glochidia attached to their hosts. As the host fish moves up or down river, fully metamorphosed juvenile mussels excyst from the host fish and are deposited to the river bottom in the habitat of its host fish. This is when the juvenile mussels begin self-sustaining life. This obligate parasitic life history and limited ability to select and disperse to suitable habitats complicates mussel conservation. Once a species becomes extirpated or depressed within a reach, recovery is hampered by this complex life history and lack of adult mobility (Neves et al., 1997; Strayer et al., 2004). Reduced mussel population densities may lower fertilization rates due to decreased probability of successful gamete encounters during spawning. The mussel fauna also depends on a suitable composition and density of fish assemblages to ensure recruitment success (Neves et al., 1997; Strayer, 2004). Similarly, for the fish fauna to remain viable and diverse, suitable numbers and types of food items (mostly aquatic insects) must be present. The aquatic and habitat quality in the Clinch River in response to a pattern of decline in some reaches of the river but not others. This introductory article provides general background for the Clinch River system, introduces the featured collection, and summarizes findings of the collected articles. Biodiversity Value and Challenges The Clinch River and its tributaries in the largely free-flowing system upstream of Norris Lake, Tennessee, support 133 species of fish (Jenkins and Burkhead, 1994) and at least 46 extant species of mussels, including 20 species that are federally endangered (Neves et al., 1997; Jones et al., 2014). Fifty-six mussel species, 18.5% of the United States (U.S.) and Canada freshwater total, have been reported from this section of the Clinch River (Stansbery, 1973; Jones et al., 2014). Of these, 10 species that are believed to be extinct or extirpated (Jones et al., 2014), 29 are globally imperiled (Master et al., 1998), and 31 are described as “at-risk” by state Natural Heritage programs (Tennessee Natural Heritage Program, 2009; VDGIF, 2013). Fish species richness in the Clinch River system is also important for biodiversity conservation. A total of 116 fish species are native to the Clinch River system, representing 52% of the fishes in the entire Tennessee River watershed, including 17 endemic species (Jenkins and Burkhead, 1994; Warren et al., 1997). The 87 native fishes in the Virginia portion of the Clinch River basin comprise 44% of Virginia’s total native fish species (Jenkins and Burkhead, 1994; Warren et al., 1997). Local species richness can be high, as many as 95 species at a single shoal, based on repeated sampling over several years (David A. Etnier, University of Tennessee, 2003, personal communication to B. Beaty), presenting strategic conservation opportunities. While the fishes are not as imperiled as mussels, five species are federally endangered or threatened and 14 are “at-risk” (Natureserve, 2012). For several of the rarer fishes, the Clinch River populations are important for rangewide conservation of the species. The Clinch River’s mussel and fish fauna rank among the most important in North America based on imperilment (Master et al., 1998). The extent of imperilment, especially for its mussel species, also makes the Clinch River an important conservation focal area. Using a rarity- and richness-weighted metric for analysis and comparison of all U.S. areas, Chaplin et al. (2000) identified the Clinch River as among the top U.S. conservation priorities. The U.S. Fish and Wildlife Service (FWS) designated this watershed as a priority area in the Virginia EcologiJAWRA AND 2 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA insects in the riffle habitats typically occupied by mussels and lithophilic spawning fish are also sensitive to environmental disturbances and contaminants (Roy et al., 2003). Robust mussel communities provide ecological benefits for epibenthic flora and fauna supporting fish, leading to a feedback loop among mussels and fish (Vaughn et al., 2008). We define a high-quality mussel assemblage or fauna as one composed of most mussel species expected to occur in the reach based on historical surveys and exhibiting evidence of recruitment for many to all species present; while a low-quality mussel assemblage exhibits substantially reduced species richness and abundance compared to historical data and/or little evidence of recruitment in extant species; we use these terms following the Johnson et al. (2014) and Ostby et al. (2014) contributions to this collection. Furthermore, we use the term “impacted” to describe river reaches occupied by low-quality assemblages, given historic accounts of dense and diverse mussel assemblages occurring throughout the Clinch River’s Virginia extent (Ortmann, 1918; Stansbery, 1973; Ahlstedt, 1986), and the term “integrity” to describe an assemblage or community composition that appears as relatively unimpacted by human activities. The Clinch River flows southwest from its source in Virginia’s Tazewell County through Russell, Wise, and Scott counties, Virginia, and Hancock and Claiborne counties, Tennessee, to Norris Lake, an impoundment north of Knoxville, Tennessee. The basin upstream of Norris Lake is the Clinch River’s primary biodiversity refuge. The papers in this collection focus on reaches upstream of Norris Lake in Tennessee and Virginia (Figure 1). The hydrologic units (Seaber et al., 2007) containing these river segments and their contributing watershed areas (study area, Figure 1) occupy 3,721 km2. Major land uses are forest (66%) and agriculture (>20%), primarily hay crops and livestock pasture (Table 1). Agriculture, primarily cattle grazing, is the principal managed land use throughout the watershed’s Ridge and Valley areas. Developed land uses, including developed open space, cover about 7.3% of the basin and are most concentrated in lower-elevation stream valleys. The area’s human population, as determined through analysis of tract-level data (U.S. Census, 2010), is approximately 105,000, with an average density of approximately 28 people per km2. The Clinch River watershed contains lands of both the Ridge and Valley and Appalachian Plateaus physiographic provinces (Figure 1) (Fenneman, 1938). The OF THE AMERICAN WATER RESOURCES ASSOCIATION TENNESSEE, USA: A FEATURED COLLECTION Ridge and Valley is defined by a folded and faulted geologic sequence composed of carbonates, sandstones, siltstones, and shales of Cambrian to Devonian age (roughly 541-360 million years) with local inclusions of younger, more flat-lying Mississippianand Pennsylvanian-aged (roughly 360-300 million years) limestones, sandstones, and other sedimentary strata (Figures 1 and 2). The province is characterized by parallel ridges, commonly capped with sandstones, and fertile valleys with soils derived from carbonates which commonly include karst features. The Appalachian Plateaus have formed from sandstones, shales, and other sedimentary strata of Mississippian and Pennsylvanian age that are generally flat-lying but locally dissected by streams. These strata contain bituminous coal seams. Coal mining, including surface mines, underground mines, coal processing and transportation, is a major industry in this portion of the watershed. Coal-bed methane production, including well-pad development and road and pipeline construction, has been expanding in the region as well. Because of terrain, much of this region’s residential and urban development has occurred in the valleys close to tributaries and the main river. Several population centers occur within the watershed of the Clinch River and its tributaries, including Richlands, Cedar Bluff, Cleveland, and St. Paul, along the Clinch River’s banks; Wise, Norton, and Coeburn along the Clinch’s largest tributary, the Guest River; Castlewood and Lebanon, to the mainstem’s southeast; Tazewell, near the headwaters; and Sneedville, in Tennessee. All of these and some smaller towns utilize sewage treatment plants that discharge treated effluent into the river or its tributaries. A number of industrial facilities also occur along the river’s length, including a coal-fired electric power plant located at Carbo, Virginia, at the Clinch River’s confluence with Dumps Creek since the 1950s, and a second coal-fired power plant recently constructed in a tributary drainage near St. Paul, Virginia. The Clinch River watershed’s southwestern (downstream) segment has a lower population density and fewer developed areas than the northeastern (upstream) segment; and it has less agriculture and more forest than the northeastern segment’s Ridge and Valley areas (Table 1). This area also contains the most dense and diverse mussel assemblages. Watershed Features JOURNAL AND Ecosystem Stressors Concerns for water contamination effects on the Clinch River’s biodiversity, first voiced by Ortmann (1918), remain current today. Although the Clinch River’s fauna remain diverse, it has been in decline 3 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, AND DONOVAN LEGEND Watershed boundary Clinch River KY Major tributaries Population centers > 3000 Appalachian plateaus VA < 3000 Ridge and valley Geologic cross section (see Figure 2) St. Paul Coeburn Separates northeast, southwest (see Table 1) Wise Guest River Lick Creek Cedar Bluff Richlands Dumps Creek Tazewell Norton Little River Claypool Dungannon Stock Creek Lebanon Cleveland Castlewood Pendleton Island Sneedville WV Indian Creek Hill Copper Creek Swan Island NC Clinchport TN Horton Ford Limits of Jones et. al. (2014) mussel assemblage designations H Diverse, no significant perturbations S Stable, higher densities than other VA sections D In decline, little evidence of recruitment L Low density, but with recruitment Limits (approx.) of upstream reach with low-quality fauna from past impacts (Price et. al., 2014) High-quality and Low-quality mussel assemblages, as sampled and classified by Johnson et al. (2014), Ostby et al. (2014). D S H L VA S D TN FIGURE 1. The Upper Clinch River Watershed, as Defined by 12-Digit Hydrologic Units Containing Mainstem Monitoring Locations Addressed by the Featured Collection and Upstream Tributary Areas. Above: Physiographic, hydrographic, and cultural features within the study area. Physiography designations as ecoregions defined by Omernik (1987) and U.S. EPA (2012a), Level III. Below: Mussel assemblage status designations, as defined and documented by Featured Collection articles. The impounded waters of Norris Lake begin approximately 20 km southeast of the furthest downstream sampling point in the lower figure. to wastewater treatment plant effluents have also been noted (Goudreau et al., 1993). Other sources of stress within the system may include point-source discharges from wastewater treatment and industrial facilities, tributary streambank and channel erosion, domestic livestock activity within stream channels, atmospheric deposition of pollutants such as nitrates, sulfates, and mercury, and discharges from active and abandoned coal facilities. Collectively, these and perhaps other stressors have contributed to the declines of the mussel fauna in Virginia reaches of the Clinch River and have adversely affected other faunal groups including fishes. Surface-water and bed-sediment quality are critical environmental variables that likely affect the health and status of mussel communities. Juvenile mussels throughout much of its Virginia extent (Burkhead and Jenkins, 1991; Neves, 1991; Neves et al., 1997; Jones et al., 2014). Since Ortmann’s time, numerous human activities in the watershed have delivered sediment and contaminants to the river, potentially altering stream temperatures and creating other stresses to the aquatic fauna. A risk assessment found that agricultural and urban land uses as well as proximity to mining activities and transportation corridors were inversely related to a fish index of biotic integrity and mussel species diversity (Diamond and Serveiss, 2001; USEPA, 2002). This study also found that habitat quality measures and acute, episodic stressors such as chemical spills were also strongly related to fishand mussel population metrics. Historic impacts due JAWRA 4 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA AND TENNESSEE, USA: A FEATURED COLLECTION TABLE 1. Area, Population Density, and Land Use within the Clinch River Watershed Areas Represented by Figure 1 as Determined from Analysis of National Land Cover Database 2006 Data (MRLC, 2006) and U.S. Census (2010). Appalachian Plateaus Area (km2) Population density (people/km2) Land use (percent of total) Water, wetlands Forested Shrub/scrub Agricultural (pasture, hay, and crops) and grasslands/herbaceous1 Developed: open Developed: low, medium, and high density Barren Ridge and Valley, Northeast 1,116 33 1,857 32 Ridge and Valley, Southwest 748 12 Total 3,721 28 0.2 78.8 0.1 10.9 0.2 54.4 0.6 37.9 0.9 73.0 0.6 19.1 0.3 65.5 0.4 26.0 4.7 4.2 4.0 2.7 4.9 1.3 4.4 2.9 1.1 0.2 0.2 0.5 1 Review of aerial photography reveals that extensive areas mapped as grassland/herbaceous within the Ridge and Valley are in agricultural use for pasture and hay. 600m Pennsylvanian quartzarenite Mississippian 300m Mid-Cambrian Mississippian Mid-Cambrian Ordovician Devonian Lower Cambrian Lower Cambrian Upper Cambrian sandstone siltstone shale limestone/dolomite FIGURE 2. Geologic Cross Section for the Surface Trace Represented in Figure 1, Oriented Northwest (left) to Southeast (right). Predominant characters of geologic units are represented but rock types within most geologic units are mixed. Geologic periods of origin for geologic units are represented along the horizontal axis. Pennsylvanian and Mississippian strata form Appalachian Plateaus physiography and also include coal seams. Bold lines separating geologic units represent faults. Vertical axis is elevation above sea level and is scaled at 29 the horizontal. Figure has been redrawn from Evans and Troensegaard (1991). example, is a leading cause of water-quality impairment of streams in the U.S. (USEPA, 2012b); however, it remained unclear whether physical habitat disturbance is correlated with the observed mussel declines. Natural physical habitat features, such as bed stability and channel geomorphology, have been linked to mussel density, diversity, and survival (Vannote and Minshall, 1982; Church, 1997; Brim Box and Mossa, 1999; Krstolic, 2001; Diamond et al., 2002; Strayer, 2004; Strayer et al., 2004; Gangloff and Feminella, 2007; Baldigo et al., 2008; Osterling et al., 2010). Excessive sedimentation has been shown to harm mussel populations through degrading habitat, clogging gills, and impairing filter-feeding efficiency (Ellis, 1936; Aldridge et al., 1987; Neves et al., 1997). Sedimentation and toxicity are related issues because many contaminants are preferentially bound to organic or mineral sediments (Adams et al., 1992), which may preferentially expose juvenile mussels. live burrowed in the sediment and are directly influenced by the quality of sediments and interstitial waters (Yeager et al., 1994; Cope et al., 2008). As filter-feeding organisms, mature mussels process large volumes of water and suspended and deposited particles to extract sustenance. Freshwater mussels are known to be sensitive to a variety of water and sediment contaminants, especially at early-life stages, including ammonia (Augspurger et al., 2003; Mummert et al., 2003; Wang et al., 2007), sodium chloride (Gillis, 2011), and other mineral salts (Kunz et al., 2013), chlorine (Goudreau et al., 1993), metals including Cd, Cu, Ni, Pb, and Zn (Havlik and Marking, 1987; Jacobson et al., 1993, 1997; Naimo, 1995; Keller et al., 2007; Cope et al., 2008; Wang et al., 2010), and K (Keller et al., 2007). At the outset of this research, much remained unknown about contaminant and habitat influences on mussels in the Clinch River. Sedimentation, for JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION 5 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, DONOVAN Jones et al. (2014) conducted and interpreted mussel population surveys at 18 locations from 2004 to 2009, and described current status of mussel populations within various river reaches. They reported dramatic differences in mussel-assemblage status indicators among their study sites. The mussel assemblages in the Clinch River’s Tennessee section, upstream of Norris Lake, are thriving. Mussel populations in Virginia vary in status, but none exhibited the density and richness observed in the Tennessee reaches. These authors noted particular concern with the river reach containing Pendleton Island, in Virginia, where dramatic declines in density have been recorded. They designated the Clinch River section extending from the confluence with Lick Creek, just upstream of the town of St. Paul, past Pendleton Island to the Clinch River’s confluence with Stock Creek (see Figure 1) as harboring mussel fauna that are “in decline with little evidence of recruitment.” Price et al. (2014) analyzed historic water- and sediment-quality data collected by government agencies from the Clinch River over a period extending from 1964 through 2010. They found that water column ammonia, water column metals (including Cu), and sediment metals (including Pb and Zn) have historically occurred at levels exceeding thresholds established by environmental quality guidelines (MacDonald et al., 2000; VDEQ, 2010; USEPA, 2013) and may have negatively impacted mussels. Most of these exceedances were recorded prior to 1979 when formal mussel surveys were initiated and they occurred throughout the river’s extent, although most frequently in the river’s upper reaches (upstream of Dumps Creek). The analysis also revealed that water column occurrences of ammonia and metals at potentially toxic concentrations have declined in frequency and have not been recorded during the past decade. Price et al. (2014) also noted that dissolved solids exhibited rising trends over the study period throughout most of the Clinch River’s extent. The Price et al. (2014) analysis revealed no significant differences in dissolved solids concentrations between the impacted reach that includes Pendleton Island, where concentrations over the 2000-2010 period averaged 208 mg/l, and the Clinch River’s Tennessee reach where mussels are thriving. The sensitivity of those comparisons was hampered by variations of sampling frequencies and locations and of measurement methods over the study period. Ostby et al. (2014) sought to determine if the observed variations in mussel assemblages throughout the Clinch River reflect differences in physical habitat quality. This study was conducted in response to work in the southern Appalachians and elsewhere that suggests mussel assemblages are influenced by Elevated specific conductance (SC), an indicator of water’s dissolved solids content, and also a concern in the Clinch River, has been shown to be strongly correlated with benthic macroinvertebrate community structure change in headwater and other low order Appalachian streams (Pond et al., 2008; USEPA, 2011). The toxicity of dissolved solids to freshwater organisms is dependent upon ionic composition (Mount et al., 1997). Dissolved solids in coal mining influenced Appalachian streams, such as those that occur in the Clinch River watershed, are often dominated by the anions SO42 and HCO3 and the cations Ca2+ and Mg2+ (Pond et al., 2008) as is the Clinch River itself (Johnson et al., 2014). Although toxicity of some pesticide formulations to freshwater mussels have been studied (Bringolf et al., 2007a; Keller et al., 2007), effects of many other organic constituents, such as polycyclic aromatic hydrocarbons (PAHs) and other nonpesticide persistent organics have not been well studied. Trace metals, pesticides, and other organic contaminants occur in the water column and in sediments (Hampson et al., 2000; USEPA, 2004; Wang et al., 2013). Given the changes experienced by mussel fauna in the Clinch River, we see investigation of water, sediment, and physical habitat quality as essential to an improved understanding of how human activities are influencing mussels in the river, and development of effective conservation measures. This collection of studies addresses these issues in the Clinch River with the intent of providing a foundation for conservation of the imperiled mussel assemblages and further research. OVERVIEW OF FEATURED COLLECTION This featured collection contains studies of water, sediment, and physical habitat quality and mussel population status conducted in the free-flowing section of the Clinch River upstream of Norris Lake. These studies focus on spatial associations of potential stressors with indicators of mussel health and diversity, with the goal of resolving uncertainties concerning which stressors are most influential. The studies were conducted using field-derived data; as such, they do not present direct causal relationships between mussel declines and specific environmental stressors. However, they have been conducted with the intent that their findings may contribute to identification of environmental factors responsible for the mussel declines that have been observed in the Virginia sections of the Clinch River. JAWRA AND 6 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA stream channel geomorphology. These authors measured channel geomorphology, assessed lotic habitats, and characterized mussel assemblages at 10 locations, seeking relationships among key habitat variables. They also compared habitat quality metrics and associated physical habitat variables measured in the Clinch River to those in other streams throughout the upper Tennessee River basin that are known to harbor mussels. While the habitat characteristics varied among rivers and among sites, the sites in the Clinch River supporting both high- and low-quality mussel assemblages were more similar to each other than to occupied habitats in other rivers. Their findings provided little evidence to suggest that geomorphology and physical habitat are associated with the variations in mussel assemblage conditions in the Clinch River. None of the habitat metrics evaluated by Ostby et al. (2014) were consistently correlated with patterns of mussel decline. However, they and other authors (Church, 1997; Krstolic, 2001) have found evidence that geomorphology may have played a more substantial role in structuring assemblages historically. These findings suggest Clinch River sites and others in the upper Tennessee River basin have the potential to support greater richness than they do currently. Johnson et al. (2014) evaluated water and sediment chemistry at many of the same sites studied by Ostby et al. (2014), with the goal of identifying environmental factors that differed between sites in the Clinch River’s impacted section and those supporting stable or recovering populations. Environmental variables studied included SC, major ions, selected metals and other trace elements, and organic constituents (PAHs, and aliphatic compounds) in water and sediments. Turbidity, SC, and water column concentrations of several constituents, including Cl, Ca, F, K, Fe, Na, Mg, Se, and SO42 , were significantly greater at a site in the impacted reach near Pendleton Island, compared to a site in a reach supporting high-quality mussel assemblages. Evaluating water quality at 15 sites along the mainstem Clinch River, they observed that the spatial distributions of elevated dissolved ion (Ca, Na, Cl, and Fl) and metal (total Fe and Mn) concentrations correlated with spatial patterns of mussel decline. They also found bedsediment concentrations of several elements varied in regular patterns over the river’s extent. Several trace elements and metals in bed sediments tended to increase moving from upstream to downstream (e.g., Al, As, Cr, Ni), while several other elements exhibited the opposite pattern (e.g., C, Ca, Mg, Na, P), but neither pattern correlated with the spatial distribution of declining or stable mussel populations. Considering findings by Wang et al. (2013), Johnson et al. (2014) determined that all sediment JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION AND TENNESSEE, USA: A FEATURED COLLECTION samples had combined metals PAH contents at levels potentially toxic to freshwater mussels. Caged hatchery-raised mussels exposed to ambient waters and native mussels harvested in impacted reaches accumulated higher tissue concentrations for a number of metals (e.g., Cd, Co, Cu, Fe, Pb, Ni, Th, V) than similarly treated mussels placed in reaches harboring high-quality assemblages, suggesting that water column exposure is more important to metals uptake in mussels than bed sediment. The Johnson et al. (2014) findings indicate that dissolved major ion concentrations vary among different segments of the mainstem river, and suggest that inflows to the mainstem from tributaries that drain mined areas increase dissolved constituent concentrations while water influxes from tributaries that drain land without mines cause dilution and lower concentrations. Johnson et al. (2014) observed that the spatial distributions of elevated dissolved ion and metal concentrations in the mainstem correspond with impacted reaches. Johnson et al. (2014) found that water column metals (measured as total forms) and major ions consistently correlated with patterns of mussel decline, while organic compounds, water column nutrients, and bed-sediment metals and organic compounds did not. COLLECTIVE INTERPRETATION It is now clear, based on quantitative data, that the mussel fauna in the Clinch River has declined dramatically in a 68-km reach from St. Paul downstream to Clinchport, Virginia (Jones et al., 2014), and that other faunal impacts have been noted elsewhere in the Clinch River’s Virginia sections. At Pendleton Island where 46 species have been recorded, density has declined from 24 m 2 in 1979 to 0.66 m 2 in 2009 (Ahlstedt et al., 2005; Jones et al., 2014). Such a precipitous decline at the best documented site in the river — a greater than 96% decline in density over a 30-year period — represents a catastrophic loss in mussel abundance; essentially it is a collapse of the fauna at what was formerly among the most species-rich sites in the country (Jones et al., 2014). Because of the high number of mussel and fish species listed as federally endangered that occur or occurred at this site and in this reach of the river, federal and state natural resource agencies are concerned about long-term viability of the fauna and are seeking to understand the causes of the decline. The main cause appears to be pollution emanating from various local sources and over different time periods. Natural ecological factors, such as peri7 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, DONOVAN found 12 additional observations in the data record where ammonia concentrations were >50% of the CCC, but most of these occurred in the 1970s and the most recent was recorded in 1983. Hence, the Price et al. (2014) findings support the claim that frequencies of elevated ammonia concentrations in the water column have been declining with time. As source data for Johnson et al. (2014), Krstolic et al. (2013) documented ammonia concentrations below detection limits of 0.02 mg/l for all but 1 of 32 base-flow samples from 2009 to 2011 and <0.06 mg/l for 20 storm samples regardless of sampling location. These ammonia values were well below the USEPA CCC intended to protect freshwater mussels (USEPA, 2013). Although mean total N and median total P from state-agency data evaluated by Price et al. (2014) were above the USEPA’s ecoregional reference levels (USEPA, 2000), they found no indication that the recent levels cause environmental stress to mussels. Throughout most of the Clinch River’s extent over the study period, total N and total P levels were found to be either stable or declining (Price et al., 2014). odic floods and droughts, reduced availability of food and fish hosts, are not considered the suspected drivers of mussel population declines. The temporal pattern of the decline occurring within the section of the river that includes Pendleton Island is downward and unidirectional. Ostby et al. (2014) found no differences in more than 50 physical habitat variables among high-quality and impacted river reaches, concluding that measured physical habitat differences do not explain the observed patterns in mussel assemblages. Natural ecological drivers would cause populations to fluctuate randomly and similarly throughout the entire length of the river in Tennessee and Virginia. Mussel populations in Tennessee, downstream of the impacted reaches in Virginia, have maintained much higher density levels and do not exhibit a downward pattern of decline, while those further upstream in Virginia are exhibiting some recovery potential (Ahlstedt et al., 2005; Jones et al., 2014). Thus, ecological factors do not offer plausible scientific explanations for understanding such sharp mussel population declines in the Clinch River section that includes Pendleton Island. In contrast, contaminants affecting water and habitat quality offer a plausible and logical explanation for the observed declines. High levels of ammonia, metals, dissolved solids, and organic compounds have been documented in the water, sediment, and tissues of native mussels and other bivalves in the river over the last 30 years, while severe pollution discharges and contaminant spills have also been documented (as reviewed by Price et al., 2014). Hence, this featured collection of articles documents past and current mussel population status and scientific evidence concerning the role of historical and contemporary pollution in the river as a likely driver of the faunal losses occurring in the Clinch River’s Virginia sections. Major Ions One water-quality measure that has deteriorated in the Clinch River is dissolved solids, an indicator of major ion concentrations. Using data records extending from the late 1960s through 2010, Price et al. (2014) found that dissolved solids increased over time throughout the river’s extent, and, hence, exhibited a negative association with the pattern of mussel decline that has been observed at Pendleton Island. A complementary pattern was observed by Johnson et al. (2014) through continuous measurements of SC. That study found that SC, frequently used as a surrogate for dissolved solids, was lower at Cleveland and Horton Ford (high-quality mussel assemblage) than at Dungannon (low-quality mussel assemblage). Together, these two studies provide evidence that dissolved solids concentrations in the Clinch River are associated, negatively, with the ecological condition of mussel assemblages. Interpretation of these results to infer potential effects of major ions in the Clinch River, however, is not straightforward. Freshwater mussel sensitivity to major ions has been most studied in laboratory settings for NaCl solutions in reconstituted waters of varying hardness. Such studies have found acute toxicities for NaCl to vary widely among freshwater mussel species (Valenti, 2004; Bringolf et al., 2007b; Gillis, 2011), among life stages (Bringolf et al., 2007b; Blakeslee et al., 2013), and with water hardness (Gillis, 2011). Gillis (2011) also found acute toxicity to native mussel glochidia at lower NaCl concentrations in laboratory reconstituted Ammonia, Nutrients Water quality of free-flowing streams within many human-altered landscapes of the U.S. has passed through a pattern of substantial degradation occurring before passage of federal environmental laws. Price et al. (2014) analyzed historic datasets to determine water-quality change. They found that most measured parameters improved from the 1960s to 2010, particularly in the most recent decade. Total P concentrations and frequencies of elevated concentrations for metals and ammonia generally declined over the analysis period. Ammonia levels exceeded USEPA’s criterion chronic concentration (CCC) (USEPA, 2013) three times in the data record, most recently in 1990. The Price et al. (2014) analysis also JAWRA AND 8 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA Metals Studies investigating effects of other contaminants on Clinch River mussels began in the 1980s, as scientists postulated that chronic toxic contaminant OF THE AMERICAN WATER RESOURCES ASSOCIATION TENNESSEE, USA: A FEATURED COLLECTION exposures were partially responsible for local or reach-scale declines. Metals, both dissolved and sediment-borne, have received the greatest attention (Belanger et al., 1986, 1990; Jacobson et al., 1993, 1997; Farris et al., 1994; Cherry et al., 2001; Wang et al., 2013). Metals, particularly Cu and Zn originating from an electric power plant’s wastewater discharge, were suspected of preventing recovery of the mussel fauna downstream of the coal-fired electric power plant located on the Clinch River’s bank at the Dumps Creek confluence following a pair of chemical spills in 1967 and 1970 (Crossman et al., 1973). Cherry et al. (1991) summarized these events, highlighting that Cu was detrimental to mussel growth and physiological activity at water concentrations in the 15-20 lg/l range, with acutely toxic effects noted at 19.4 lg/l. Additional research reported Cu-acute toxicity levels (median effective concentrations, EC50s) as low as 17-36 lg/l for native freshwater mussels (Jacobson et al., 1993; Gillis et al., 2010). Analysis of historical water-quality data suggests that water column metals concentrations have declined in recent decades (Price et al., 2014). Metals and associated trace elements in the Clinch River system since 2000 demonstrate water column dissolved concentrations for As, Cd, Cr, Cu, Pb, Ni, Se, and Zn as low, relative to USEPA’s CCC (Price et al., 2014). Johnson et al. (2014) also found concentrations of the same metals below the CCC from 2009 to 2011. However, Se concentrations during base flow and storm events in whole water and dissolved phase were significantly higher at an impacted site, relative to a site harboring high-quality mussel assemblages. They also found dissolved (0.45-lm filtered) Fe and Mn in the water column were correlated with mussel population status along the mainstem Clinch River. Cherry et al. (2001) reported concentrations of Al, Fe, and Se in Clinch River interstitial waters that exceeded acute toxicity values for sensitive freshwater test organisms, with the highest levels measured in the Clinch River reach with low-quality mussel assemblages. Cherry et al. (2001) also reported that the Cu and Pb tissue concentrations of in situ exposed Corbicula fluminea correlated with declines in mussel species richness at known mussel-assemblage locations. A similar pattern was detected by Johnson et al. (2014), who found higher tissue concentrations of several metals (including, Cd, Cu, Fe, Pb, K, and Th) in caged mussels and all measured metals in native mussels from sites with lower mussel-assemblage quality compared to Horton Ford in the downstream segment supporting high-quality assemblages. The caged mussels were removed from the bed-sediment interface and exposed only to the water column. Sediment concentrations at all of the Clinch River sites are at potentially stressful levels, with probable waters than in natural river waters, suggesting that the full range of water chemistry is influential as demonstrated for laboratory test organisms by Mount et al. (1997). The above studies with NaCl cannot be translated to infer toxic dissolved solids levels in the Clinch River where predominant major ions are SO42 , HCO3 , Ca2+, and Mg2+, not Na+ and Cl . The only study characterizing toxicity to mussels of solutions with SO42 , HCO3 , Ca2+, and Mg2+ as predominant major ions (Kunz et al., 2013) found toxicities to juvenile Lampsilis siliquoidea at SC values of 504 and 565 lS/cm for 28-day exposures to two reconstituted waters. Although the Kunz et al. (2013) study tested a suite of major ions similar to that found in the Clinch River for mussel toxicity, the tested concentrations differed from mean base-flow values in the Clinch River (Krstolic et al., 2013), especially for SO42 and Mg2+. Continuous monitoring of the Clinch River documents SC values occurring at the levels reported as causing toxicity by Kunz et al. (2013). Between March 2009 and October 2011, at Dungannon within the Clinch River’s impacted reach, 0.5% of the in situ SC measurements exceeded 500 lS/cm, and the maximum recorded value was 553 lS/cm (Krstolic et al., 2013); while SC in the Guest River, a major mined tributary of the Clinch River upstream of Dungannon, exceeded 500 lS/cm ~50% of the time (Johnson et al., 2014). However, no SC measurements exceeding 500 lS/cm were recorded at sites with high-quality mussel assemblages during this period (maximum recorded SC values were 470 lS/cm at Horton Ford, Tennessee, and 479 lS/cm at Cleveland, Virginia; Krstolic et al., 2013). During low-flow years, even higher SCs occurred in the Clinch River mainstem. For example, between October 2007 and March 2009, SC exceeded 500 lS/cm at a high-quality mussel assemblage site (Horton Ford, Tennessee) 8% of the time (U.S. Geological Survey, National Water Information System. Accessed by G.C. Johnson 16 December 2013, http://waterdata.usgs.gov/nwis); and, thus, it can be inferred that 500 lS/cm exceedances in the impacted reach near Dungannon, closer to the Guest River’s high-SC water influx, were more frequent. Gaining greater understanding of major-ion concentrations, extremes, and effects, including potential chronic effects of persistent exposures to concentrations that are elevated above natural background, is essential to improved understanding of causes for mussel declines in the Clinch River. JOURNAL AND 9 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, DONOVAN tent with origination from coal (Krstolic et al., 2013). However, the current pattern of PAH distributions among Clinch River bed sediments did not correlate with mussel-assemblage quality (Johnson et al., 2014), and concentrations at Pendleton Island were lower than reported by Hampson et al. (2000). For 2010 and 2011, the fraction of bed sediments composed of coal particles did not correlate with the mussel-assemblage quality, metals, or PAH concentrations (Krstolic et al., 2013; Johnson et al., 2014). effects concentration quotients (PECQs, which characterize Cd, Cu, Ni, Pb, and Zn in sediments) above 0.2 threshold found by Wang et al. (2013) to cause growth impairments (Johnson et al., 2014; Price et al., 2014). Although none of the metals concentrations in the bed sediment corresponded to the spatial distribution of the declining mussel populations, Ca, Mg, Na, P, and Zn were negatively correlated with multiple mussel health metrics, such as total density, total recruitment, number of threatened or endangered species, and number of globally listed species (Johnson et al., 2014). This current collection of evidence indicates that metals likely have played a role in mussel declines, but the evidence is not conclusive. Metal concentrations in the water column are low relative to water quality guidelines (USEPA, 2010), but Fe and Mn in the water column were significantly higher in the impacted reaches than in reaches harboring highquality assemblages (Johnson et al., 2014). Elevated bed-sediment concentrations were found at levels that have been shown to impair mussel growth (Wang et al., 2013) throughout the Clinch River, indicating that the bed sediment and associated interstitial water could be affecting mussel populations. Mussel tissue concentrations from two separate tests show that mussels are bioaccumulating metals, with higher concentrations occurring in the impacted reaches. While the caged mussels were exposed only to the water column, and for native mussels tests the sediment metals were higher but mussel tissue metals were lower in the reach supporting high-quality assemblages, it appears that the water column exposure is the dominant route of exposure to metals. Bed-sediment metals are possibly a greater concern for juvenile mussels due to their feeding and burrowing behaviors within the sediments (Yeager et al., 1994). Past Impacts: Spills and Releases While episodic spills and release events have occurred and have been influential in upstream segments above St. Paul (Price et al., 2014), there is no evidence that these events have been influential as causes for recent declines in the impacted segment downstream of the Guest River confluence that includes Pendleton Island (Jones et al., 2014). CONCLUSION The precipitous decline in mussel assemblages documented in the Clinch River’s Virginia sections raises serious conservation concerns. This fauna is a stronghold for 20 extant federally endangered species; and, for most of these species, it is the best or only remaining broodstock for propagation and source of individuals for relocation. The findings of Price et al. (2014) and Johnson et al. (2014) suggest that primary current concerns are chronic rather than acute stresses, as no contaminant levels known to be acutely toxic, based on prior research, are evident. Price et al. (2014) has also found that measured water quality is better today than ≥2 decades ago in most respects. The exceptions are the major ions that predominate in Clinch River water chemistry. Dissolved solid concentrations are increasing and correlate temporally while elevated SC correlates spatially with patterns of mussel decline, but potential effects of the major ions present are not well understood. Sediment metals concentrations are at levels indicating their potential role as chronic stressors throughout the river, and a number of water column metal concentrations correlated negatively to mussel indices; however, sediment metals concentrations did not correlate spatially with mussel assemblage quality. Water column metals also appear as problematic given mussels’ known sensitivities and the bioaccumulation observed in impacted sections. Further studies to evaluate multi-stressor Organics Coal mining is a major landscape activity in parts of the Clinch River watershed (Figure 1). Hampson et al. (2000) reported elevated and potentially toxic PAH levels in Clinch River bed sediment at Pendleton Island, in the impacted reach. Organic contaminants, particularly PAHs, were investigated as possible mussel stressors in the Clinch River reach extending from Artrip, Virginia (5 km upstream from Cleveland) to Swan Island, Tennessee, an approximately 165-km distance (Johnson et al., 2014). The lightweight organic fraction of the bed sediment, which predominantly consisted of coal particles, were found to comprise 2.2-6.5% of the Clinch River sediments, and PAHs found in the sediments were consisJAWRA AND 10 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA relationships with mussel physiological and life history response are needed to better understand causal relationships. The information reported by these studies can aid resource management, including enhanced environmental monitoring, mussel conservation efforts, and future research for the Clinch River. Adams, W.J., R.A. Kimerle, and J.W. Barnett, Jr., 1992. Sediment Quality and Aquatic Life. Environmental Science and Technology 26:1865-1875. Ahlstedt, S.A., 1986. Cumberlandian Mollusk Conservation Program. Activity 1: Mussel Distribution Surveys. Final Report, Tennessee Valley Authority, Office of Natural Resources and Economic Development, Knoxville, Tennessee, 125 pp. Ahlstedt, S.A., M.T. Fagg, R.S. Butler, and J.F. Connell, 2005. Long-Term Trend Information for Freshwater Mussel Populations at Twelve Fixed-Station Monitoring Sites in the Clinch and Powell Rivers of Eastern Tennessee and Southwestern Virginia 1979-2004. Final Report for U.S. Fish and Wildlife Service, Cookeville, Tennessee, 38 pp. Aldridge, D.W., B.S. Payne, and A.C. Miller, 1987. The Effects of Intermittent Exposure to Suspended Solids and Turbulence on Three Species of Freshwater Mussels. Environmental Pollution 45:17-20. Augspurger, T., A. Keller, M. Black, W.G. Cope, and F. Dwyer, 2003. Water Quality Guidance for Protection of Freshwater Mussels (Unionidae) from Ammonia Exposure. Environmental Toxicology and Chemistry 22:2569-2575. Baldigo, B.P., A.G. Ernst, G.E. Schuler, and C.D. Apse, 2008. Relations of Environmental Factors with Mussel-Species Richness in the Neversink River, New York, U.S. Geological Survey Open File Report 2007-1283, 8 pp. Bates, J.M. and S.D. Dennis, 1978. The Mussel Fauna of the Clinch River, Tennessee and Virginia. Sterkiana 69-70:3-23. Belanger, S.E., J.L. Farris, D.S. Cherry, and J. Cairns, Jr., 1986. Growth of Asiatic Clams (Corbicula sp.) during and after LongTerm Zinc Exposure in Field-Located and Laboratory Artificial Streams. Archives of Environmental Contamination and Toxicology 15:427-434. Belanger, S.E., J.L. Farris, D.S. Cherry, and J. Cairns, Jr., 1990. Validation of Corbicula fluminea Growth Reductions Induced by Copper in Artificial Streams and River Systems. Canadian Journal of Fisheries and Aquatic Sciences 47:904-914. Blakeslee, C.J., H.S. Galbraith, L.S. Robertson, and B. St. John White, 2013. The Effects of Salinity Exposure on Multiple Life Stages of a Common Freshwater Mussel, Elliptio complanata. Environmental Toxicology and Chemistry 32:28492854. Brim Box, J. and J. Mossa, 1999. Sediment, Land Use, and Freshwater Mussels: Prospects and Problems. Journal of the North American Benthological Society 18:99-117. Bringolf, R.B., W.G. Cope, M.C. Barnhart, S. Mosher, P.R. Lazaro, and D. Shea, 2007a. Acute and Chronic Toxicity of Pesticide Formulations (Atrazine, Chlorpyrifos, and Permethrin) to Glochidia and Juveniles of Lampsilis siliquoidea. Environmental Toxicology and Chemistry 2:2101-2107. Bringolf, R.B., W.G. Cope, C.B. Eads, P.R. Lazaro, M.C. Barnhart, and D. Shea, 2007b. Acute and Chronic Toxicity of TechnicalGrade Pesticides to Glochidia and Juveniles of Freshwater Mussels (Unionidae). Environmental Toxicology and Chemistry 26:2086-2093. OF THE AMERICAN WATER RESOURCES ASSOCIATION TENNESSEE, USA: A FEATURED COLLECTION Burkhead, N. and R. Jenkins, 1991. Fishes. In: Virginia’s Endangered Species: Proceedings of a Symposium, K. Terwilliger (Editor). McDonald and Woodward Publishing Co., Blacksburg, Virginia, pp. 321-410. Chaplin, S., R. Gerrard, H. Watson, L. Master, and S. Flack, 2000. The Geography of Imperilment: Targeting Conservation Toward Critical Biodiversity Areas. In: Precious Heritage: The Status of Biodiversity in the United States, B. Stein, L. Kutner, and J. Adams (Editors). Oxford University Press, New York City, New York, pp. 159-200. Cherry, D.S., J.L. Farris, and R.J. Neves, 1991. Laboratory and Field Ecotoxicological Studies at the Clinch River Plant, Virginia. University Center for Environmental Studies, Virginia Polytechnic Institute & State University, Blacksburg, Virginia, 228 pp. Cherry, D.S., D.J. Soucek, and T.S. Schmidt, 2001. Metal Accumulation in Sediment, Asian Clams (Corbicula fluminea), and Periphyton at Selected Freshwater Mussel Preserves in the Clinch and Powell Rivers, Virginia. Report to The Nature Conservancy, Abingdon, Virginia. Church, G.W., 1997. Macrohabitat Factors Affecting Distribution Patterns of Freshwater Mussels in the Clinch River (Virginia, Tennessee). Ph.D. Dissertation, Virginia Polytechnic Institute and State University, Blacksburg, Virginia. Cope, W.G., R.B. Bringolf, D.B. Buchwalter, T.J. Newton, C.G. Ingersoll, N. Wang, T. Augspurger, F.J. Dwyer, M.C. Barnhart, R.J. Neves, and E. Hammer, 2008. Differential Exposure, Duration, and Sensitivity of Unionoidean Bivalve Life Stages to Environmental Contaminants. Journal of the North American Benthological Society 27:451-462. Crossman, J.S., J. Cairns, Jr., and R.L. Kaesler, 1973. Aquatic Invertebrate Recovery in the Clinch River Following Hazardous Spills and Floods. Virginia Water Resources Research Center Bulletin 63. Virginia Polytechnic Institute and State University, Blacksburg, Virginia, 56 pp. Diamond, J.M., D.W. Bressler, and V.B. Serveiss, 2002. Assessing Relationships Between Human Land Uses and the Decline of Native Mussels, Fish, and Macroinvertebrates in the Clinch and Powell River Watershed, U.S.A. Environmental Toxicology and Chemistry 21:1147-1155. Diamond, J.M. and V.B. Serveiss, 2001. Identifying Sources of Stress to Native Aquatic Fauna Using a Watershed Ecological Risk Assessment Framework. Environmental Science and Technology 35:4711-4718. Ellis, M.M., 1936. Erosion Silt as a Factor in Aquatic Environments. Ecology 17:29-42. Evans, N.H. and K.W. Troensegaard, 1991. Geology of the St. Paul and Carbo Quadrangles, Virginia. Virginia Division of Mineral Resources, Charlottesville, Virginia. Publication 106. Farris, J.L., J.L. Grudzien, S.E. Belanger, D.S. Cherry, and J. Cairns, Jr., 1994. Molluscan Cellulolytic Activity Responses to Zinc Exposure in Laboratory and Field Stream Comparisons. Hydrobiologia 287:161-178. Fenneman, N.M., 1938. Physiography of Eastern United States. McGraw Hill, New York City, New York. Gangloff, M.M. and J.W. Feminella, 2007. Stream Channel Geomorphology Influences Mussel Abundance in Southern Appalachian Streams, U.S.A. Freshwater Biology 52:64-74. Gillis, P.L., 2011. Assessing the Toxicity of Sodium Chloride to the Glochidia of Freshwater Mussels: Implications for Salinization of Surface Waters. Environmental Pollution 159:17021708. Gillis, P.L., J.C. McGeer, G.L. Mackie, M.P. Wilkie, and J.D. Ackerman, 2010. The Effect of Natural Dissolved Organic Carbon on the Acute Toxicity of Copper to Larval Freshwater Mussels (Glochidia). Environmental Toxicology and Chemistry 29:25192528. LITERATURE CITED JOURNAL AND 11 JAWRA ZIPPER, BEATY, JOHNSON, JONES, KRSTOLIC, OSTBY, WOLFE, DONOVAN Mount, D.R., D.D. Gulley, J.R. Hockett, T.D. Garrison, and J.M. Evans, 1997. Statistical Models to Predict the Toxicity of Major Ions to Ceriodaphnia dubia, Daphnia magna and Pimephales promelas (Fathead Minnows). Environmental Toxicology and Chemistry 16:2009-2019. MRLC (Multi-Resolution Land Characteristics Consortium, 2006. National Land Cover Database 2006. http://www.mrlc.gov/ nlcd06_data.php, accessed April 2012. Mummert, A.K., R. Neves, T. Newcomb, and D. Cherry, 2003. Sensitivity of Juvenile Freshwater Mussels (Lampsilis fasciola, Villosa iris) to Total and Un-Ionized Ammonia. Environmental Toxicology and Chemistry 22:2545-2553. Naimo, T.J., 1995. A Review of the Effects of Heavy Metals on Freshwater Mussels. Ecotoxicology 4:341-362. Natureserve, 2012. NatureServe Explorer: An Online Encyclopedia of Life, www.natureserve.org, accessed March 2012. Neves, R., 1991. Mollusks. In: Virginia’s Endangered Species: Proceedings of a Symposium, K. Terwilliger (Editor). McDonald and Woodward Publishing, Blacksburg, Virginia, pp. 251-320. Neves, R., A. Bogan, J. Williams, S. Ahlstedt, and P. Hartfield, 1997. Status of Aquatic Mollusks in the Southeastern United States: A Downward Spiral of Diversity. In: Aquatic Fauna in Peril: The Southeastern Perspective, G. Benz and D. Collins (Editors). Lenz Design and Communications, Decatur, Georgia, pp. 43-85, ISBN-13: 978-0965484107. Omernik, J.M., 1987. Ecoregions of the Conterminous United States. Map (Scale 1:7,500,000). Annals of the Association of American Geographers 77:118-125. Ortmann, A.E., 1918. The Nayads (Freshwater Mussels) of the Upper Tennessee Drainage with Notes on Synonymy and Distribution. Proceedings of the American Philosophical Society 57:521-626. Ostby, B.J.K., J.L. Krstolic, and G.C. Johnson, 2014. Reach-Scale Comparison of Habitat and Mollusk Assemblages for Select Sites in the Clinch River with Regional Context. Journal of the American Water Resources Association, doi: 10.1111/jawr.12218. Osterling, M.E., B.L. Arvidsson, and L.A. Greenberg, 2010. Habitat Degradation and the Decline of the Threatened Mussel Margaritifera margaritifera: Influence of Turbidity and Sedimentation on the Mussel and Its Host. Journal of Applied Ecology 47:759-768. Pond, G.J., M.E. Passmore, F.A. Borsuk, L. Reynolds, and C.J. Rose, 2008. Downstream Effects of Mountaintop Coal Mining: Comparing Biological Conditions Using Family- and GenusLevel Macroinvertebrate Bioassessment Tools. Journal of the North American Benthological Society 27:717-737. Price, J.E., C.E. Zipper, J.W. Jones, and C.T. Franck, 2014. Water and Sediment Quality in the Clinch River, Virginia and Tennessee, USA, over Nearly Five Decades. Journal of the American Water Resources Association, doi: 10.1111/jawr.12219. Roy, A., A. Rosemond, D. Leigh, M. Paul, and J. Wallace, 2003. Habitat-Specific Responses of Stream Insects to Land Cover Disturbance: Biological Consequences and Monitoring Implications. Journal of the North American Benthological Society 22:292307. Seaber, P.R., F.P. Kapinos, and G.L. Knapp, 2007. Hydrologic Unit Maps. U.S. Geological Survey, Water Supply Paper 2294. Stansbery, D., 1973. A Preliminary Report on the Naiad Fauna of the Clinch River in the Southern Appalachian Mountains of Virginia and Tennessee (Mollusca: Bivalvia: Unionoida). Bulletin of the American Malacological Union for 1972, pp. 20-22. Strayer, D.L., 2004. Freshwater Mussel Ecology: A Multifactor Approach to Distribution and Abundance. University of California Press, Berkeley, California, 204 pp. Strayer, D.L., J.A. Downing, W.R. Haag, T.L. King, J.B. Layzer, T.J. Newton, and S. Jerrine, 2004. Changing Perspectives on Pearly Mussels, North America’s Most Imperiled Animals. BioScience 54:429-439. Goudreau, S.E., R.J. Neves, and R.J. Sheehan, 1993. Effects of Wastewater Treatment Plant Effluents on Freshwater Mollusks in the Upper Clinch River, Virginia, U.S.A. Hydrobiologia 252:211-230. Hampson, P.S., M.W. Treece, Jr., G.C. Johnson, S.A. Ahlstedt, and J.F. Connell, 2000. Water Quality in the Upper Tennessee River Basin, Tennessee, North Carolina, Virginia, and Georgia, 1994-98. U.S. Geological Survey, Water Resources Division, National Water Quality Assessment Program. Circular 1205: 1-32. Havlik, M.E. and L. Marking, 1987. Effects of Contaminants on Naiad Mollusks (Unionidae): A Review. Resource Publication 164. U.S. Fish and Wildlife Service, Washington, D.C. Jacobson, P.J., J. Farris, D. Cherry, and R. Neves, 1993. Juvenile Freshwater Mussel (Bivalvia: Unionidae) Responses to Acute Toxicity Testing with Copper. Environmental Toxicology and Chemistry 12:879-833. Jacobson, P.J., R. Neves, D. Cherry, and J. Farris, 1997. Sensitivity of Glochidial Stages of Freshwater Mussels (Bivalvia: Unionidae) to Copper. Environmental Toxicology and Chemistry 16:2384-2392. Jenkins, R. and N. Burkhead, 1994. Freshwater Fishes of Virginia. American Fisheries Society, Bethesda, Maryland, ISBN-13: 978-0913235874. Johnson, G.C., J.L. Krstolic, and B.J.K. Ostby, 2014. Influences of Water and Sediment Quality and Hydrologic Processes on Mussels in the Clinch River. Journal of the American Water Resources Association, doi: 10.1111/jawr.12221. Jones, J., S. Ahlstedt, B. Ostby, B. Beaty, M. Pinder, N. Eckert, R. Butler, D. Hubbs, C. Walker, S. Hanlon, J. Schmerfeld, and R. Neves, 2014. Clinch River Freshwater Mussels Upstream of Norris Reservoir, Tennessee and Virginia: A Quantitative Assessment from 2004 to 2009. Journal of the American Water Resources Association, doi: 10.1111/jawr.12222. Keller, A., M. Lydy, and S. Ruessler, 2007. Unionid Mussel Sensitivity to Environmental Contaminants. In: Freshwater Bivalve Ecotoxicology, J.L. Farris and J.H. Van Hassel (Editors). CRC Press, Boca Raton, Florida, pp. 151-167, ISBN-13: 978-1420 042849. Krstolic, J.L., 2001. A Comparative Study of Freshwater Mussel Habitat along the Clinch River, Tennessee and Virginia, USA: Hydraulic Factors and Their Influences. M.S. Thesis, University of Tennessee, Knoxville, 99 pp. Krstolic, J.L., G.C. Johnson, and B.J.K. Ostby, 2013. Water Quality, Sediment Characteristics, Aquatic Habitat, Geomorphology, and Mussel Population Status of the Clinch River, Virginia and Tennessee, 2009-2011. U.S. Geological Survey Data Series 802, including data tables, http://pubs.usgs.gov/ds/0802/, accessed February 20, 2014. Kunz, J.L., J.M. Conley, D.B. Buchwalter, T.J. Norberg-King, N.E. Kemble, N. Wang, and C.G. Ingersoll, 2013. Use of Reconstituted Waters to Evaluate Effects of Elevated Major Ions Associated with Mountaintop Coal Mining on Freshwater Invertebrates. Environmental Toxicology & Chemistry 32:28262835. MacDonald, D.D., C.G. Ingersoll, and T.A. Berger, 2000. Development and Evaluation of Consensus-Based Sediment Quality Guidelines for Freshwater Ecosystems. Archives for Environmental Contamination and Toxicology 39:20-31. Master, L., S. Flack, and B. Stein (Editors), 1998. Rivers of Life: Critical Watersheds for Protecting Freshwater Biodiversity. The Nature Conservancy, Arlington, Virginia, 71 pp., ISBN: 1-886765-09-X, http://www.natureserve.org/library/riversoflife.pdf, accessed May 7, 2013. McMahon, R., 1991. Mollusca Bivalvia. Ecology and Classification of North American Freshwater Invertebrates. Academic Press, New York. ISBN-13: 978-0126906455. JAWRA AND 12 JOURNAL OF THE AMERICAN WATER RESOURCES ASSOCIATION FRESHWATER MUSSEL POPULATION STATUS AND HABITAT QUALITY IN THE CLINCH RIVER, VIRGINIA OF THE AMERICAN WATER RESOURCES ASSOCIATION TENNESSEE, USA: A FEATURED COLLECTION Wang, N., C.G. Ingersoll, C.D. Ivey, D.K. Hardesty, T.W. May, T. Augspurger, A.D. Roberts, E. van Genderen, and M.C. Barnhart, 2010. Sensitivity of Early Life Stages of Freshwater Mussels (Unionidae) to Acute and Chronic Toxicity of Lead, Cadmium, and Zinc in Water. Environmental Toxicology and Chemistry 29:2053-2063. Wang, N., C.G. Ingersoll, J.L. Kunz, W.G. Brumbaugh, C.M. Kane, R.B. Evans, S. Alexander, C. Walker, and S. Bakaletz, 2013. Toxicity of Sediments Potentially Contaminated by Coal Mining and Natural Gas Extraction to Unionid Mussels and Commonly Tested Benthic Invertebrates. Environmental Toxicology and Chemistry 32:207-221. Warren, M.L., P.L. Angermeier, B.M. Burr, and W.R. Haag, 1997. Decline of a Diverse Fish Fauna: Patterns of Imperilment and Protection in the Southeastern United States. In: Aquatic Fauna in Peril: The Southeastern Perspective, G. Benz and D. Collins (Editors). Lenz Design and Communications, Decatur, Georgia, pp. 105-164. ISBN-13: 978-0965484107. Yeager, M.M., D.S. Cherry, and R.J. Neves, 1994. Feeding and Burrowing Behaviors of Juvenile Rainbow Mussels, Villosa iris (Bivalvia: Unionidae). Journal of the North American Benthological Society 13:217-222. Tennessee Natural Heritage Program, 2009. Rare Animals List. Tennessee Department of Environment & Conservation, Nashville, Tennessee, 61 pp. U.S. Census (U.S. Bureau of the Census), 2010. United States Census 2010. Maps and Data, Geographic Data. http://www.census. gov/geo/maps-data/index.html, accessed June 2012. USEPA (U.S. Environmental Protection Agency), 2000. Ambient Water Quality Criteria Recommendations Information Supporting the Development of State and Tribal Nutrient Criteria Rivers and Streams in Nutrient Ecoregion XI. EPA 822-B-00020. USEPA (U.S. Environmental Protection Agency), 2002. Clinch and Powell Valley Watershed Ecological Risk Assessment. National Center for Environmental Assessment, Washington, D.C. EPA/ 600/R-01/050, 145 pp. USEPA (U.S. Environmental Protection Agency), 2004. The Incidence and Severity of Sediment Contamination in Surface Waters of the United States. National Sediment Quality Survey (Second Edition) EPA-823-R-04-007. Office of Science and Technology, Standards and Health Protection Division, http:// water.epa.gov/polwaste/sediments/cs/upload/nsqs2ed-complete-2.pdf, accessed November 2004. USEPA (U.S. Environmental Protection Agency), 2010. National Recommended Water Quality Criteria. http://water.epa.gov/ scitech/swguidance/waterquality/standards/current/index.cfm, accessed March 2011. USEPA (U.S. Environmental Protection Agency), 2011. A FieldBased Aquatic Life Benchmark for Conductivity in Central Appalachian Streams. Office of Research and Development, National Center for Environmental Assessment, Washington, DC. EPA/600/R-10/023. USEPA (U.S. Environmental Protection Agency), 2012a. Ecoregions of North America. Western Ecology Division. http://www.epa. gov/wed/pages/ecoregions/na_eco.htm, accessed June 2012. USEPA (U.S. Environmental Protection Agency), 2012b. National Water Quality Inventory Report to Congress. http://water.epa. gov/lawsregs/guidance/cwa/305b/index.cfm, accessed March 2013. USEPA (U.S. Environmental Protection Agency), 2013. Aquatic Life Ambient Water Quality Criteria for Ammonia-Freshwater 2013. Office of Water. EPA 822-R-13-001. USFWS (U.S. Fish and Wildlife Service), 2012. Virginia Ecological Services Strategic Plan 2010-2014. Northeast Region, Virginia Ecological Services, 206 pp. http://www.fws.gov/northeast/virgin iafield/strategic_plan.html, accessed April 2013. Valenti, T.W., 2004. Approaches for Assessing Toxicity of Selected Contaminants to Freshwater Mussels (Bivalvia: Unionidae). M.S. Thesis, Virginia Tech, Blacksburg, Virginia. Vannote, R.L. and G.W. Minshall, 1982. Fluvial Processes and Local Lithology Controlling Abundance, Structure, and Composition of Mussel Beds. Proceedings of the National Academy of Sciences USA 79:4103-4107. Vaughn, C.C., S.J. Nichols, and D.E. Spooner, 2008. Community and Foodweb Ecology of Freshwater Mussels. Journal of the North American Benthological Society 27:409-423. VDEQ (Virginia Department of Environmental Quality), 2010. 2010 Water Quality Assessment Guidance Manual. http://www. deq.state.va.us/wqs/documents/WQS_eff_1FEB2010.pdf, accessed August 2013. VDGIF (Virginia Department of Game and Inland Fisheries), 2013. Virginia Fish and Wildlife Information Service. https://fwis web1.dgif.virginia.gov/fwis/, accessed April 2013. Wang, N., C.G. Ingersoll, D.K. Hardesty, C.D. Ivey, J.L. Kunz, T.W. May, F.J. Dwyer, A.D. Roberts, T. Augspurger, C.M. Kane, R.J. Neves, and M.C. Barnhart, 2007. Acute Toxicity of Copper, Ammonia, and Chlorine to Glochidia and Juveniles of Freshwater Mussels (Unionidae). Environmental Toxicology and Chemistry 26:2036-2047. JOURNAL AND 13 JAWRA
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