Southeastern Naturalist 347 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 22001144 SOUTHEASTERN NATURALIST 1V3o(2l.) :1334,7 N–3o6. 62 Freshwater Mussels (Unionidae) in the Paint Rock River (Jackson, Madison, and Marshall Counties), Alabama Todd B. Fobian1,*, Michael L. Buntin1, Jesse T. Holifield1, Thomas A. Tarpley1, Jeffrey T. Garner2, and Paul D. Johnson1 Abstract - The Paint Rock River (PRR) drainage in northern Alabama and southern Tennessee historically supported 58 freshwater mussel species. This study semi-quantitatively examined the mussel assemblage at 42 sites in the Paint Rock mainstem and 5 sites in Estill Fork, a headwater tributary. A total of 1825 live mussels were collected over 78.9 personhours, with an overall catch per unit effort (CPUE) of 23.1 mussels/person-hour. Forty-one species were collected live and/or fresh dead, including federally protected Epioblasma triquetra (Snuffbox), Fusconaia cor (Shiny Pigtoe), Lampsilis abrupta (Pink Mucket), Lampsilis virescens (Alabama Lampmussel), Pleuronaia dolabelloides (Tennessee Pigtoe), Quadrula cylindrica cylindrica (Rabbitsfoot), and Toxolasma cylindrellus (Pale Lilliput). The river system continues to support a high diversity of mussels (48 species collected in the past 25 years). The survey also identified several sites in the basin suitable for the reintroduction of extirpated species. Introduction The Paint Rock River (PRR) drainage in northern Alabama and southern Tennessee is identified as a global hotspot for aquatic biodiversity, historically supporting 58 freshwater mussel and 98 fish species (Boschung and Mayden 2004, Williams et al. 2008). Surveys over the past 25 years indicate the continued presence of at least 48 mussel species (Godwin 2002, Williams et al. 2008), including Cumberlandian Basin endemics, whose range is limited to the upper and middle Tennessee and Cumberland River watersheds. Currently, 10 of 58 documented mussel species appear extirpated from the drainage, including four federally protected and two extinct species. Eight federally protected species are currently extant (USFWS 1976, 2011; Williams et al. 2008). This survey is part of a multifaceted effort by the Alabama Aquatic Biodiversity Center (AABC; Alabama Department of Conservation and Natural Resources [ADCNR]), Marion, AL, to restore the PRR mussel fauna. Due to the high species richness and improved habitat quality over the past two decades as a result of the ADCNR Landowner Incentive Program and efforts by The Nature Conservancy (TNC), the river is identified as a primary location for Cumberlandian mussel restoration efforts (CRMRC 2010). Mussel recovery through reintroduction, translocation, and augmentation is a viable restoration strategy identified by the USFWS and NMFS (2000). 1Alabama Division of Wildlife and Freshwater Fisheries, 2200 Highway 175, Marion, AL 36756, (334) 683-5000. 2Alabama Division of Wildlife and Freshwater Fisheries, 350 County Road 275, Florence, AL 35633.*Corresponding author - todd.fobian@dcnr.alabama.gov. Manuscript Editor: Arthur Bogan Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 348 Prior to commencement of species-restoration activities, a thorough assessment of the current mussel population was necessary. Previous mussel surveys of the mainstem river were completed in 1980 (Ahlstedt 1986) and 1991 (Ahlstedt 1995), and headwaters tributaries (Estill Fork, Larkin Fork, and Hurricane Creek) were last surveyed in 1995 and 2002 (Godwin 2002; McGregor and Shelton 1995). Mussel species historically inhabiting the river system can be inferred based on the following sources: Ortmann (1925), a summary of multiple collector’s material prior to 1925; Johnson et al. (2008), a catalog of curator Herbert D. Athearn’s mollusk collection that were acquired between the years of 1957–1969; Isom and Yokley (1973), a mussel-fauna report between1965–1967; and Williams et al. (2008), a comprehensive review of US museum material. The present study updates the freshwater mussel inventory of the PRR and recommends priority restoration actions to improve mussel populations within the drainage. Fieldsite Description The PRR is located in Jackson, Madison, and Marshall counties in northeast Alabama (Fig. 1). The river flows southwest along the southern edge of the Cumberland Plateau for 97 river km (rkm; 60 river miles [rm]), confluencing with the Tennessee River at TRM 343.2. The lower 21 rkm (13 rm) is impounded by Wheeler Reservoir. The river drainage area encompasses 1186 km2 (458 mi2) and includes three major headwater tributaries: Larkin Fork, Estill Fork, and Hurricane Creek. Estill Fork and Hurricane Creek originate in Franklin County, TN, and flow south where they join to form the PRR. The upper headwaters of the river system generally have narrow floodplains adjacent to forested mountains. These areas have a low residential density, and land use is primarily large farms and commercial timberlands. Lower reaches of the watershed consist of a wider river valley floodplain with a narrow riparian zone usually surrounded by pasture and row crops (Ahlstedt 1995). The PRR drainage was severely affected in past decades by small impoundments, stream channelization, erosion, and agricultural runoff. These habitat influences have led to the possible extirpation of 10 mussel and 8 fish species from the river within the past 75 years (Boschung and Mayden 2004, Mirarchi 2004, Williams et al. 2008). A major detrimental impact on habitat occurred with the channelization and removal of snags and riverbank timber in the upper drainage and the lower reaches of Larkin and Estill forks and Hurricane Creek by the US Army Corps of Engineers during the 1960s (Ahlstedt 1995). This direct headwater habitat manipulation was probably a large contributor to freshwater mussel loss in the drainage. Wheeler Dam was completed by the Tennessee Valley Authority (TVA) in 1936, resulting in loss of most of the mussel fauna and riverine habitat in the lower 21 km of PRR (Ahlstedt 1995). Loss of habitat and fragmentation created by impoundments is the leading cause of decline and extinction of North American mollusk species (Vaughn and Taylor 1999). Continuing threats to the watershed include siltation and erosion from poor farming practices along with commercial and residential development (Godwin Southeastern Naturalist 349 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 Figure 1. Paint Rock River drainage mollusk survey sites in Jackson, Madison, and Marshall counties, AL. Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 350 2002). A heavy layer of silt can suffocate entire mussel beds and has contributed to extirpation of mussels in several rivers (Anderson et al. 1991). Specific activities that increase siltation and erosion include clearing of riparian vegetation, cattle access/grazing, timber clear cutting, head cutting, gravel mining, in-stream ATV traffic, and runoff from poor farming and construction practices (Vaughn and Taylor 1999). Nonpoint source agricultural runoff and chemical spills are also a threat. There appears to be direct and inadvertent impacts from some of the widely used pesticides, herbicides, and fertilizers, but many of the potential effects are still not fully understood (Cope et al. 2008). Uncertainties remain about the relationship of laboratory data to actual contaminant exposure routes for various mussel species and life stages. Additional research is needed to understand these complex environmental interactions so that the risks of exposures can be properly assessed and managed (Cope et al. 2008, Milam et al. 2005). Methods Free-flowing sections of the river were float surveyed by kayak (RM 14–60) and motor boat (RM 5–14) in June, July, and August 2008, and an additional 10 rkm (6 rm) of Estill Fork, was kayaked and surveyed in June 2008. We used US Geological Survey (7.5-minute) topographical maps and a hand held global positioning system (GPS) (Magellan Meridian Platinum unit, WAAS enabled) for navigation and sample-site location. The GPS coordinates were verified and plotted on topographical maps using Maptech Terrain Navigator GIS software. We based sampling-site selection on three criteria: 1) presence and abundance of live mussels and fresh dead shell, 2) stream-bank and channel stability, and 3) maintenance of uniform spacing of sites throughout the study area. Searches focused on shoal areas consisting of riffles, runs, and shallow pools. The total amount of time spent sampling was recorded as catch per unit effort (CPUE), a semi-quantitative method used for determining the number of mussels collected per person-hour (ph). This technique is reliable for estimating total species richness and locating rare species, the primary goal of this study (Obermeyer 1998). Each site was sampled by a three-person crew. Collection methods included snorkeling, visual searching, and hand grubbing. We collected all mussels encountered within a sampling reach approximately 50–100 m long. Collected mussels were placed in net bags and taken to shore where they were identified and counted. Taxonomy followed that of Williams et al. (2008). Mussel species considered abundant were counted in situ and left undisturbed in the substrate. We determined maximum total length by measuring the distance from the anterior to posterior end of the shell margins parallel to the hinge line to the nearest millimeter (mm) using vernier calipers. Collecting time varied among sites and continued until no additional species were present and all suitable habitats adequately examined. Live individuals were returned to location of capture. Dead shells were gathered along the shore, streambed, and feeding stations (middens) left by Ondatra zibethica (L.) (Muskrat), and were classified as fresh Southeastern Naturalist 351 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 dead (shiny nacre, intact hinge ligament, decomposing tissue inside shell), weathered dead (dull nacre, relatively intact periostracum), and relic (chalky nacre, flaky or absent periostracum). Shell materials from different sites were segregated, labeled, and curated at the North Carolina Museum of Natural Science (NCMNS) in Raleigh, NC. Recorded site descriptions include date and general habitat information. We took site photographs and described the distribution of mussel beds. These details are not included in this manuscript, but are on file at the Alabama Aquatic Biodiversity Center (AABC). Results During this study, we assessed a total of 46 sites in the PRR system, including 5 in Estill Fork and 41 in the mainstem (Tables 1, 2). The mainstem covered 88.5 rkm (55 rm; from PRR mile [RM] 5–60; Fig. 1). River discharge was 302 cfs and 196 cfs, respectively, during the first two sampling trips in June 2008. However, flows ranged from 12–24 cfs for the remainder of the sampling trips in July and August (USGS gage station # 03574500 Woodville, AL). In all, 1825 live mussels were collected during 78.9 ph (mean = 1.7 ph/site, range = 0.5–5.3 ph/site). Overall CPUE was 23.1 live mussels/ph. Forty-one species were found live and/or fresh dead, and all species occurred in the mainstem except Medionidus conradicus (Lea) (Cumberland Moccasinshell) and Toxolasma cylindrellus (Lea) (Pale Lilliput), which were found only in Estill Fork. The most abundant species encountered was Amblema plicata (Say) (Threeridge), which comprised 30.0% of the cumulative total, followed by Quadrula cylindrica cylindrica (Say) (Rabbitsfoot; 12.0%), Quadrula verrucosa (Rafinesque) (Pistolgrip; 8.2%), Lampsilis ovata (Say) (Pocketbook; 7.9%), Potamilus alatus (Say) (Pink Heelsplitter; 7.2%), and Cyclonaias tuberculata (Rafinesque) (Purple Wartyback; 6.0%). The remaining species were all below 4.0% in relative abundance. The most widely distributed mussel was Villosa vanuxemensis (Lea) (Mountain Creekshell), which was found at 63.8% of the sites sampled, followed by A. plicata at 61.7%, Lampsilis fasciola (Rafinesque) (Wavyrayed Lampmussel) at 61.7%, L. ovata at 57.4%, Villosa iris (Lea) (Rainbow) at 55.3%, and Pleuronaia barnesiana (Lea) (Tennessee Pigtoe) at 48.9%. The remaining species were detected at fewer than 45.0% of the sites. Table 1. Estill Fork river mile/kilometer (RM/KM), locality, and latitude/longitude coordinates for sites sampled during summer 2008 in Jackson County, AL. RM/KM Locality Coordinates EF 5.6/9.0 Downstream of Burks Creek confluence N34º 57.759', W86º 09.241' EF 3.6/5.8 Upstream of Reid Hollow tributaries N34º 56.332', W86º 09.355' EF 2.8/4.8 Downstream of Houston Hollow confluence N34º 55.793', W86º 09.562' EF 2.2/3.5 Northeast of Estill Fork N34º 55.327', W86º 09.533' EF 1.0/1.6 Upstream of Freedom Bridge N34º 55.330', W86º 09.528' Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 352 The following species were represented by single live individuals: Ellipsaria lineolata (Rafinesque) (Butterfly), Elliptio dilatata (Rafinesque) (Spike), Fusconaia ebena (Lea) (Ebonyshell), M. conradicus, Pleurobema cordatum (Rafinesque) (Ohio Pigtoe), Pleurobema rubrum (Rafinesque) (Pyramid Pigtoe), Pyganodon grandis (Say) (Giant Floater), and T. cylindrellus. Other relatively rare species, represented by five or fewer individuals, included: Actinonaias ligamentina (Lamarck) Table 2. Paint Rock river mile/kilometer (RM/RKM), locality, and latitude/longitude coordinates for sites sampled during summer 2008 in Jackson, Madison, and Marshall Counties Alabama. RM/RKM Locality Coordinates 60.0/96.6 Hurricane Creek confluence N34º 53.883', W86º 10.237' 59.8/96.2 East of Bostik Hill Church N34º 53.753', W86º 10.315' 59.6/95.9 Upstream of private bridge N34º 53.545', W86º 10.400' 59.4/95.6 Downstream of private bridge N34º 53.397', W86º 10.483' 59.2/95.3 Downstream of Henshaw Cove N34º 53.293', W86º 10.497' 59.1/95.1 East of Henshaw Cemetery N34º 52.198', W86º 10.674' 59.0/95.0 Southeast of Henshaw Cemetery N34º 53.069', W86º 10.898' 58.3/93.8 West of Bouldin Cemetery N34º 52.739', W86º 11.277' 58.0/93.3 Upstream of Robertson Cove confluence N34º 52.655', W86º 11.341' 57.5/92.5 West of Swaim N34º 52.162', W86º 11.536' 56.7/91.2 Upstream of Fowler Cove tributaries N34º 51.598', W86º 11.935' 56.2/90.4 Downstream of Fowler Cove tributaries N34º 51.603', W86º 12.203' 55.3/89.0 Downstream of Larkin Fork confluence N34º 51.674', W86º 12.649' 54.7/88.0 Northeast of Church Hill Cemetery N34º 51.510', W86º 13.001' 53.9/86.7 Downstream of County Road 142 bridge N34º 51.017', W86º 13.302' 53.3/85.8 West of Princeton N34º 50.657', W86º 13.601' 52.3/84.2 Upstream of Cowen Hollow confluence N34º 51.031', W86º 13.952' 50.7/81.6 Upstream of gravel ford N34º 49.225', W86º 14.615' 50.3/81.0 Upstream of Graham Cove confluence N34º 49.093', W86º 14.339' 48.4/77.9 Downstream of County Road 12 bridge N34º 47.826', W86º 14.418' 48.2/77.6 Downstream of County Road 12 bridge N34º 47.330', W86º 14.396' 46.7/75.2 South of Hollytree N34º 46.762', W86º 14.950' 44.4/71.5 Upstream of Frazier Cove confluence N34º 45.434', W86º 14.246' 43.6/70.2 Downstream of County Road 20 bridge N34º 44.898', W86º 13.906' 42.0/67.6 Downstream of Flippo Ford N34º 44.224', W86º 14.198' 38.5/62.0 North of Gilliam Spring N34º 43.303', W86º 16.709' 36.5/58.7 Downstream of heavy equipment ford N34º 42.861', W86º 18.122' 34.3/55.2 Downstream of Big Lake bridge N34º 41.868', W86º 18.604' 33.7/54.2 Downstream of Hales Cove confluence N34º 41.414', W86º 18.658' 33.3/53.6 Upstream of gravel ford N34º 41.245', W86º 18.610' 30.0/48.3 Downstream of Whitaker Preserve N34º 38.978', W86º 19.107' 29.2/47.0 Upstream of Webb Hollow confluence N34º 39.318', W86º 19.512' 27.6/44.4 West of Splitrock Mountain N34º 38.266', W86º 18.514' 25.2/40.6 Downstream of Stillhouse Branch N34º 36.652', W86º 19.331' 21.2/34.1 Upstream of Butler ’s Mill bridge N34º 34.880', W86º 18.080' 20.6/33.2 Downstream of Butler's Mill bridge N34º 34.711', W86º 18.368' 19.0/30.6 Bend upstream of Tremble Creek N34º 34.396', W86º 19.904' 18.2/29.3 Bend downstream of Tremble Creek N34º 33.886', W86º 19.631' 16.7/26.9 Near Butlers Mill Road bend N34º 33.545', W86º 18.475' 15.5/24.9 Downstream of Fish Trap Ford N34º 32.665', W86º 18.743' 13.1/21.1 Downstream of Buck Ford N34º 31.090', W86º 19.915' Southeastern Naturalist 353 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 (Mucket), Lampsilis teres (Rafinesque) (Yellow Sandshell), Lampsilis virescens (Lea) (Alabama Lampmussel), Quadrula pustulosa (Lea) (Pimpleback), Truncilla donaciformis (Lea) (Fawnsfoot), and Truncilla truncata (Rafinesque) (Deertoe). Two live A. ligamentina were found at Paint Rock RM 43.6. Prior to this survey, this species was last reported live in Alabama from Wheeler Reservoir in the early 1990s with no indication of abundance (Bowen et al. 1994), and subsequent status assessments of freshwater mussels in this Tennessee River reach indicate that if present it is very rare (Garner and McGregor 2001). Obovaria subrotunda (Rafinesque) (Round Hickorynut), occurred at 10% of the sites sampled, but was only collected fresh dead; no live specimens were found. Another rare species, Lampsilis abrupta (Say) (Pink Mucket), was not found during this survey, but a fresh-dead individual was collected during the survey period (Paul Freeman, TNC, Birmingham, AL, pers. comm.). A lotic-habitat specialist, Utterbackia imbecillis (Say) (Paper Pondshell), was collected fresh dead from Cole Spring Branch, a PRR tributary, on 25 November 2007 (Paul Freeman, pers. comm.). Three species were represented by weathered dead and/or relic shells only: Fusconaia cuneolus (Lea) (Finerayed Pigtoe), Ligumia recta (Lamarck) (Black Sandshell), and Quadrula quadrula (Rafinesque) (Mapleleaf). Seven sites had mussel species richness ≥19 live or fresh dead, and 12 sites had a CPUE of ≥19 live mussels per ph. (Appendix 1). Sites with both high species richness and CPUE included RM 48.2, RM 43.6, RM 34.3, RM 33.3, and RM 19.0. Based on our survey results, these five localities were considered the best mussel habitat in the river. We observed evidence of recent recruitment for 26 species as indicated by the presence of subadult/juvenile age classes (1–5 annuli, ≤50 mm shell length for most medium-to-large species). Populations of abundant mussels, A. plicata, Q.c. cylindrica, C. tuberculata, and Pleuronaia dolabelloides (Lea) (Slabside Pearlymussel), appeared healthy and comprised of several age classes, though most individuals were large adults. Interestingly, subadult Fusconaia subrotunda (Lea) (Longsolid) and Pleurobema oviforme (Conrad) (Tennessee Clubshell) outnumbered older individuals of those species. Species of Conservation Concern Of the twelve federally protected mussel species historically present in the PRR system, we encountered only seven live or fresh dead: Epioblasma triquetra (Rafinesque) (Snuffbox), Fusconaia cor (Conrad) (Shiny Pigtoe), L. abrupta (1 fresh dead), L. virescens, P. dolabelloides, Q.c. cylindrical, and T. cylindrellus. Only weathered dead and relic shells of F. cuneolus were collected. All federally protected species and candidates encountered are considered priority 1 (species of highest conservation concern in Alabama; Mirarchi 2004). In the Cumberlandian Region Mollusk Recovery Plan, F. cor, F. cuneolus, L. virescens, and T. cylindrellus are tier 1 species (highest priority); P. dolabelloides is tier 2; and E. triquetra, L. abrupta, and Q.c. cylindrica are tier 3 (CRMRC 2010). Federally protected species historically found in the PRR but not collected during this survey are Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 354 Epioblasma capsaeformis (Lea) (Oyster Mussel), Epioblasma florentina (Lea) (Yellow Blossom), Pleurobema plenum (Lea) (Rough Pigtoe), and Villosa trabalis (Conrad) (Cumberland Bean). Epioblasma triquetra This survey collected 21 live individuals from seven sites and fresh dead from eight sites (RM 46.7–13.1). The sizes ranged from 9–52 mm in length, and 15 sexually mature individuals yielded a 3:2 male-to-female ratio. Epioblasma triquetra appears to be recruiting and viable, and its presence in PRR is considered the best of only two known Alabama populations. Fusconaia cor This survey collected seven live F. cor from three sites (RM 43.6, RM 33.3, and between RM 34.3–33.7), and fresh dead from two sites (RM 48.2 and RM 46.7). Weathered dead shells also occurred at six sites (RM 25.2–RM 59.05). The size range of live individuals was 21–80 mm in length. RM 43.6, at which five of the seven live specimens were found, had a CPUE of 44.5 mussels/ph and a species richness of 20, making it one of the best sites evaluated. The presence of a single live sub-adult and multiple fresh dead juveniles suggests that F. cor is reproducing in PRR. Fusconaia cuneolus This survey did not yield any live Fusconaia cuneolus, but did find weathered dead shells at RM 20.6 and relic shells at four sites. The identification of relic F. cuneolus is difficult since they are very similar to those of F. cor, and the most discerning differences between the two species (periostracum color and texture), are generally absent. Fusconaia cuneolus has not been collected live in the river since 1991 (Ahlstedt 1995), when two old individuals were found at RM 13.3 and RM 16.0. Lampsilis abrupta This survey did not find any Lampsilis abrupta, but a single fresh-dead shell was reported from RM 34.0 during the survey period (Paul Freeman, pers. comm.). No additional individuals were found despite a concerted effort at RM 34.3. That site was one of the best sampled, with a CPUE of 64.3 mussels/ph and species richness of 23. Lampsilis virescens This survey found live Lampsilis virescens at three sites (RM 50.3, RM 48.2, and Estill Fork mile 3.6), fresh dead at two (RM 53.9 and RM 52.3) weathered dead shells at RM 50.7, and relic shells at RM 53.3. This mussel occurred at 10.6% of sites surveyed. Extant populations were believed restricted to Estill Fork, but this survey extended the known range downstream into the river (RM 48) where it is rare. The size range of live individuals was 53–89 mm in length. Pleuronaia dolabelloides This survey collected a total of 66 live P. dolabelloides at 13 sites and fresh dead from four (RM 59.05–13.1). The cumulative relative abundance of P. dolabelloides Southeastern Naturalist 355 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 was 3.62%, and the size range of live individuals was 11–78 mm in length. Several size classes are indicative of recent recruitment and healthy populations. Quadrula c.cylindrica This survey collected a total of 218 live Q.c.cylindrica from 18 sites and fresh dead at one site (RM 56.7–RM 13.1). The size range of live individuals was 21–117 mm in shell length. Several size classes suggest recent recruitment and healthy populations. Toxolasma cylindrellus This survey found one live gravid female at Estill Fork mile 5.6. A weathered dead shell was collected from Estill Fork mile 2.2, and one relic valve was collected from Estill Fork mile 1.0. The presence of a gravid female is a positive sign of reproduction potential, but recruitment viability is unknown. Discussion and Summary There appears to have been little interest in the PRR mussel fauna prior to the latter half of the 20th century, since there were no systematic surveys of the system up to that point. Ortmann (1925) summarized early mussel records from the middle and lower Tennessee River based on material collected by other individuals. Herbert D. Athearn, a collector of southeastern aquatic mollusks visited several PRR system sites from 1957–1969 and documented 39 mussel species, which he curated privately (Johnson et al. 2008, Williams et al. 2008). His personal collection was donated in 2007 to NCMNS where it was reviewed and is being databased (Johnson et al. 2008, Williams et al. 2008). The first published account of PRR mussels was Isom and Yokley (1973), who made several visits to the river from 1965–1967 and reported 30 species from 6 sites. Subsequent surveys were more intensive, but results were variable. The TVA surveyed the PRR in 1980 (Ahlstedt 1986) and again in 1991 (Ahlstedt 1995), reporting 25 and 37 species, respectively. Godwin (2002) reported only 19 species from a survey that was concentrated in the headwaters. Williams et al. (2008) included a comprehensive review of museum material from US institutions that house significant collections of mussel mat erial. Parmalee and Bogan (1998) report 102 species of mussels known to occur in the Tennessee River system and its tributaries; 58 species, approximately 57%, are known from the PRR drainage (Ahlstedt 1995, Isom and Yokley 1973, Mc- Gregor and Shelton 1995, Ortmann 1925). Based on the 41 species collected live and/or fresh dead during this survey, the PRR system ranks among those with the highest remaining mussel diversity in North America. For comparison, the Duck and Clinch rivers in Tennessee are both nationally recognized as significant conservation resources for their diverse mussel faunas (Hubbs et al. 2011, Jones and Neves 2011). The Duck River historically supported 75 mussel species and harbors 56 extant species (Ahlstedt et al. 2004). The Clinch River presumably supports >40 mussel species (Ahlstedt 1991). The current PRR mussel assemblage includes at least eight federally protected species (E. triquetra, F. cor, Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 356 L. abrupta, L. virescens, P. dolabelloides, Q.c. cylindrical, and T. cylindrellus, plus the potentially extant F. cuneolus). The mussel fauna is globally significant, including the last remaining population of T. cylindrellus and one of two remaining populations of L. virescens. The PRR mussel fauna remains diverse and relatively healthy, but some species are healthier than others. Recent recruitment and the presence of smaller size classes suggest healthy populations observed for several species, including federally protected E. triquetra, F. cor, and P. dolabelloides. Two species, P. oviforme and F. subrotunda, have been eliminated from much of their former ranges and are among species considered in the CRMRC (2010). Both mussels were more commonly encountered as sub-adults rather than older individuals. Two species previously thought extirpated from the drainage, A. ligamentina and L. abrupta, were collected in the 2008 survey, and Lasmigona complanata (Barnes) (White Heelsplitter) and U. imbecillus were subsequently found live by the authors. Mussel species reported from the drainage within the last 25 years that were not found during this survey are presented in Appendix 2. However, among those listed are Alasmidonta viridis (Rafinesque) (Slippershell Mussel) and Lasmigona holstonia (Lea) (Tennessee Heelsplitter); both species occur almost exclusively in small headwater streams that were not covered in this survey. Weathered dead shells of F. cuneolus found suggest that it may still occur in the PRR at very low densities. Toxolasma parvum (Barnes) (Lilliput) is a widespread species that typically occurs in lotic habitats, which are not prevalent in the river. A non-dated T. parvum record near RM 30 (Williams et al. 2008) suggests it may still be extant despite not being found. Alasmidonta marginata (Say) (Elktoe) was not collected, but a shell was found in 2004 in a Muskrat midden near Whitaker Nature Preserve at RM 30 (Doug Shelton, private consultant, Mobile, AL, pers. comm.), which indicates its continued persistence in the river. Two species more commonly found in big rivers, L. recta and Quadrula metanevra (Rafinesque) (Monkeyface), are likely peripheral to their primary distributions in larger rivers. Their density in PRR was probably always low, and they might still persist at very low densities. Similarly, for all species reported from the PRR in the last 25 years but not encountered during this survey, the possibility exists that they are still extant but rare. There have been some losses to the historically documented mussel fauna. Two species are now considered extinct: Epioblasma biemarginata (Lea) (Angled Riffleshell) and Epioblasma lenior (Lea) (Narrow Catspaw). Three species, P. plenum, Pleurobema sintoxia (Rafinesque) (Round Pigtoe), and Q. quadrula, are primarily big-river species, and the PRR was likely peripheral to their primary distributions. Potamilus ohiensis (Rafinesque) (Pink Papershell) is also a big-river species that has expanded its range into Alabama since impoundment of the Tennessee River increased the prevalence of its preferred lotic habitats (Williams et al. 2008). The only report of PRR P. ohiensis is Williams et al. (2008), and no date was provided for the record. However, Ortmann (1925) reported P. ohiensis from adjacent Flint River, indicating that the species occurred at least in the PRR vicinity prior to Tennessee River impoundment. Other historically reported species include Actinonaias Southeastern Naturalist 357 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 pectorosa (Conrad) (Pheasantshell), E. capsaeformis, E. florentina, and V. trabalis (Appendix 2). All Epioblasma spp. disappeared from the PRR except E. triquetra, and current numbers of that species appear to have increased (Ahlstedt 1995, Isom and Yokley 1973). In summary, the PRR drainage remains home to a diverse, globally significant freshwater mussel assemblage, despite the loss of major faunal components due to habitat alteration and poor land-use practices. However, past habitat perturbations have diminished over time, and for over a decade, the Alabama TNC and the ADCNR Landowner Incentive Program have worked to accelerate land-use improvements. Also, great strides have been made in techniques for captive propagation of mussels, and ADCNR has made a large investment in facilities dedicated to that purpose at AABC. With these recent developments taken into account, pro - tection of the remaining mussel assemblage appears promising, and restoration of its extirpated fauna is a real possibility. Acknowledgments Thanks to Doug Fears and Paul Freeman of The Nature Conservancy for help with this study. They provided river-access information, housing, local knowledge and shell material during the survey efforts. Additional expertise in shell identification was provided by Steven Ahlstedt, Mike Gangloff, and Art Bogan. Funding was provided by Section 6 of the Endangered Species Act and the Alabama Division of Wildlife and Freshwater Fisheries. Literature Cited Ahlstedt, S.A. 1986. Cumberlandian Mollusk Conservation Program. Activity 1: Mussel distribution surveys. Tennessee Valley Authority, Services and Field Operations, Norris, TN. 125 pp. Ahlstedt, S.A. 1991. Twentieth-century changes in the freshwater mussel fauna of the Clinch River (Tennessee and Virginia). Walkerana 5:73–122. Ahlstedt, S.A. 1995. Status survey for federally listed endangered freshwater mussel species in the Paint Rock River system, Northeastern Alabama, USA. Walkerana 8:63–80. Ahlstedt, S.A., P.D. Johnson, J.R. Powell, R.S. Butler, M.T. Fagg, D.W. Hubbs, S.F. Novak, and S.R. Palmer. 2004. Historical and current examination of freshwater mussels (Bivalvia: Margaritiferidae, Unionidae) in the Duck River basin Tennessee. Final Report: Tennessee Wildlife Resources Agency, Nashville, TN. Contract No. FA-02-14725-00. 213 pp. Anderson, R.M., J.B. Layzer, and M.E. Gordon. 1991. Recent catastrophic decline of mussels (Bivalvia: Unionidae) in the Little South Fork Cumberland River, Kentucky. Brimleyana 17:1–8 Boschung, H., and R. Mayden. 2004. Fishes of Alabama. Smithsonian Books, Mobile, AL. 736 pp. Bowen, Z.H., S.P. Malvestuto, W.D. Davies, and J.H. Crance. 1994. Evaluation of the mussel fishery in Wheeler Reservoir, Tennessee River. Journal of Freshwater Ecology 9(4):313–319. 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(2):451–462. Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 358 Cumberlandian Region Mollusk Restoration Committee (CRMRC). 2010. Plan for the population restoration and conservation of freshwater mollusks of the Cumberlandian Region. V + 145 pp. Garner, J.T., and S.W. McGregor. 2001. Current status of freshwater mussels (Unionidae, Margaritiferidae) in the Muscle Shoals area of the Tennessee River in Alabama (Muscle Shoals revisited again). American Malacological Bulletin 16(1–2):155–170. Godwin, J.C. 2002. Monitoring of federally listed and rare mussels in the Paint Rock River. Alabama Heritage Program, The Nature Conservancy, Birmingham, AL. 80 pp. Hubbs, D.W., S. Chance, L. Colley, and R.S. Butler. 2011. 2010 Duck River quantitative mussel survey. Final Report: Tennessee Wildlife Resources Agency Fisheries Division, Nashville, TN. Report 11-04. 47 pp. Isom, B.G., and P.H. Yokley. 1973. The mussels of the Flint and Paint Rock River systems of the southwest slope of the Cumberland Plateau in North Alabama 1965 and 1967. American Midland Naturalist 89:442–447. Johnson, P.D., S.F. Novak, and A.E. Bogan. 2008. An Electronic Database of the Museum of Fluviatile Mollusks. A Final Report to United States Fish and Wildlife Service, Cookeville, TN. Jones, J.W., and R.J. Neves. 2011. Influence of life-history on demographic responses of three freshwater mussel species (Bivalvia: Unionidae) in the Clinch River, USA. Aquatic Conservation: Marine and Freshwater Ecosystems 21:57–73. McGregor, S.W., and D.N. Shelton. 1995. A qualitative assessment of the Unionid fauna of the headwaters of the Paint Rock and Flint Rivers of North Alabama and adjacent areas of Tennessee. Report to the Alabama Department of Conservation and Natural Resources, Montgomery, AL. 65 pp. Milam, C.D., J.L. Farris, F.J. Dwyer, and D.K. Hardesty. 2005. Acute toxicity of six freshwater mussel species (glochidia) to six chemicals: Implications for Daphnids and Utterbackia imbecillis as surrogates for protection of freshwater mussels (Unionidae). Archives of Environmental Contamination and Toxicology 48(2):166–173. Mirarchi, R.E. (Ed.). 2004. Alabama Wildlife: Volume 2: Imperiled Aquatic Wildlife. University of Alabama Press, Tuscaloosa, AL. 256 pp. Obermeyer, B.K. 1998. A comparison of quadrats versus timed snorkel searches for assessing freshwater mussels. American Midland Naturalist 139(2):331–339. Ortmann, A.E. 1925. The Naiad-fauna of the Tennessee River system below Walden Gorge. American Midland Naturalist 9(8):321–372. Parmalee, P.W., and A.E Bogan. 1998. The Freshwater Mussels of Tennessee. University of Tennessee Press, Knoxville, TN. 328 pp. United States Fish and Wildlife Service (USFWS). 1976. Endangered and threatened wildlife and plants; endangered status for 159 taxa of animals. Department of the Interior Federal Register. 41:24062–24067. USFWS. 2011. Endangered and threatened wildlife and plants; review of native species that are candidates for listing as endangered or threatened; annual notice of findings of resubmitted petitions; annual description of progress on listing actions; proposed rule. Department of the Interior Federal Register. 76:66370–66439. USFWS and National Marine Fisheries Service (NMFS). 2000. Policy regarding controlled propagation of species listed under the Endangered Species Act. Department of Interior Federal Register. 65:56916–56922. Southeastern Naturalist 359 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 Vaughn, C.C., and C.M.Taylor. 1999. Impoundments and the decline of freshwater mussels: A case study of an extinction gradient. Conservation Biology 13:912–920. Williams, J.D., A.E. Bogan, and J.T. Garner. 2008. Freshwater Mussels of Alabama and the Mobile Basin in Georgia, Mississippi, and Tennessee. The University of Alabama Press, Tuscaloosa, AL. 908 pp. Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 360 Appendix 1. Summary of PRR system mussel collections. Includes species collected at each site (denoted by Paint Rock meanstem and Estill Fork river mile), search time, CPUE, and species richness. * = federally protected species, FD = fresh-dead shell, WD = weathered-dead shell, and R = relic shell. Paint Rock River mainstem (RM) Species 60.0 59.8 59.6 59.4 59.2 59.1 59.0 58.3 58.0 57.5 56.7 56.2 55.3 54.7 53.9 Actinonaias ligamentina Amblema plicata R R R R R 4 4 FD 57 2 7 Cyclonaias tuberculata Ellipsaria lineolata Elliptio crassidens (Lamarck) 1 R Elliptio dilatata Epioblasma triquetra* Fusconaia cor* WD WD Fusconaia ebena Fusconaia subrotunda 4 WD R 3 2 R Lampsilis abrupta* Lampsilis fasciola WD 1 1 FD WD R 1 Lampsilis ovata 1 R 2 5 1 9 R Lampsilis teres Lampsilis virescens* R FD Lasmigona costata (Rafinesque) 1 FD 1 R 2 R R Leptodea fragilis (Rafinesque) Medionidus conradicus Megalonaias nervosa (Rafinesque) R Obliquaria reflexa Rafinesque Obovaria subrotunda R R Pleurobema cordatum Pleurobema oviforme 1 FD Pleurobema rubrum Pleuronaia barnesiana R R R WD 2 1 FD 1 2 FD 2 R 2 Pleuronaia dolabelloides WD 1 R R R 1 2 R 1 Potamilus alatus 1 R 1 Ptychobranchus fasciolaris (Rafinesque) 1 WD R 2 1 R 3 Pyganodon grandis Southeastern Naturalist 361 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 Paint Rock River mainstem (RM) Species 60.0 59.8 59.6 59.4 59.2 59.1 59.0 58.3 58.0 57.5 56.7 56.2 55.3 54.7 53.9 Quadrula c.cylindrica* R R R 2 6 R Quadrula pustulosa Quadrula verrucosa 1 R R 2 Toxolasma cylindrellus* Toxolasma lividum Rafinesque 2 WD FD FD Truncilla donaciformis Truncilla truncata Villosa iris 1 FD FD 2 FD R Villosa taeniata (Conrad) WD 1 9 WD WD 1 WD 4 FD Villosa vanuxemensis 1 WD WD 2 1 WD FD WD FD FD FD FD 1 Search time (ph) 1.0 1.0 1.0 0.5 2.0 1.75 0.75 1.25 1.0 3.0 3.0 2.25 2.25 0.5 3.0 Total number of mussels 1 0 2 0 16 9 1 1 1 10 22 1 88 2 17 CPUE (mussels/hour) 1 0 0.5 0 8.0 5.1 1.3 0.8 1 3.3 7.3 0.4 39.1 4 5.6 Total number of species (L, FD) 1 0 2 0 6 5 1 4 2 9 12 5 12 1 10 Paint Rock River mainstem (RM) Species 53.3 52.3 50.7 50.3 48.4 48.2 46.7 44.4 43.6 42.0 38.5 36.5 34.3 33.7 33.3 Actinonaias ligamentina 2 Amblema plicata 5 10 29 6 2 50 L 12 74 34 3 5 85 18 125 Cyclonaias tuberculata 2 1 1 WD 5 2 1 2 2 2 22 Ellipsaria lineolata Elliptio crassidens R FD WD 2 1 1 Elliptio dilatata Epioblasma triquetra* FD 1 1 3 4 3 Fusconaia cor* FD FD WD 5 WD WD 1 Fusconaia ebena 1 Fusconaia subrotunda WD FD 1 WD 1 7 WD 1 3 1 11 Lampsilis abrupta* FD Lampsilis fasciola 2 1 2 1 FD 1 L 2 9 2 FD 2 2 WD 2 Lampsilis ovata 4 1 7 11 2 10 L 14 8 7 13 9 7 13 Lampsilis teres 1 Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 362 Paint Rock River mainstem (RM) Species 53.3 52.3 50.7 50.3 48.4 48.2 46.7 44.4 43.6 42.0 38.5 36.5 34.3 33.7 33.3 Lampsilis virescens* FD WD 1 2 Lasmigona costata 2 2 7 1 10 FD 11 10 1 1 5 2 13 Leptodea fragilis 2 FD 1 4 3 FD 2 Medionidus conradicus Megalonaias nervosa 1 1 3 4 Obliquaria reflexa 2 1 2 FD 2 Obovaria subrotunda FD FD FD Pleurobema cordatum WD R Pleurobema oviforme R FD WD 1 19 1 28 Pleurobema rubrum Pleuronaia barnesiana 2 2 2 1 1 L FD 2 2 Pleuronaia dolabelloides WD 3 1 R FD WD 4 WD 4 15 4 14 Potamilus alatus 1 R 2 2 4 3 R FD 15 13 2 15 19 15 32 Ptychobranchus fasciolaris 2 2 2 1 WD FD L FD 6 FD FD 1 WD FD Pyganodon grandis 1 Quadrula c.cylindrica* 3 R 13 1 12 7 8 11 4 14 3 119 Quadrula pustulosa R 1 Quadrula verrucosa 1 R 20 2 4 100 L 5 2 12 Toxolasma cylindrellus* Toxolasma lividum FD 1 FD FD 1 FD 1 6 Truncilla donaciformis FD 2 Truncilla truncata FD 1 FD 1 Villosa iris 2 2 2 1 FD FD FD 4 WD FD Villosa taeniata WD FD 2 Villosa vanuxemensis WD FD FD 1 2 FD FD FD 3 WD FD FD 1 WD Search time (ph) 1.5 1.3 2.7 2.25 1.5 2.25 0.75 3.0 4.0 1.25 0.6 1.0 3.0 1.0 3.0 Total number of mussels 24 22 87 37 17 196 N/A 17 178 78 27 52 193 61 410 CPUE (mussels/hour) 16 16.9 32.2 16.4 11.3 87.1 N/A 5.7 44.5 62.4 45 52 64.3 61 137 Total number of species (L, FD) 12 10 16 12 12 22 11 13 19 9 12 12 22 16 21 Southeastern Naturalist 363 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 Paint Rock River mainstem (RM) Estill Fork (RM) Species 30.0 29.2 27.6 25.2 21.2 20.6 19.0 18.2 16.7 15.5 13.1 5.6 3.6 2.8 2.2 1.0 Totals Actinonaias ligamentina R 2 Amblema plicata 2 1 1 2 6 1 1 R FD R R 2 R 548 Cyclonaias tuberculata 5 1 8 2 43 10 1 WD L 110 Ellipsaria lineolata 1 R 1 Elliptio crassidens FD FD 2 4 R FD 11 Elliptio dilatata R R R 1 R 1 Epioblasma triquetra* FD FD FD 2 7 FD FD FD FD 21 Fusconaia cor* WD 6 Fusconaia ebena 1 Fusconaia subrotunda WD WD 1 5 6 FD 1 FD FD 47 Lampsilis abrupta* R FD Lampsilis fasciola 1 FD FD 1 FD FD 6 FD FD 1 FD 38 Lampsilis ovata 4 FD 7 1 5 2 3 L 144 Lampsilis teres R FD FD R 1 2 FD 4 Lampsilis virescens* 1 4 Lasmigona costata FD R R FD FD 69 Leptodea fragilis 2 2 FD FD FD FD 16 Medionidus conradicus 1 1 Megalonaias nervosa 3 2 1 1 WD FD 16 Obliquaria reflexa WD FD FD 1 FD 8 Obovaria subrotunda WD WD FD FD Pleurobema cordatum 1 R R R FD 1 Pleurobema oviforme WD FD 1 9 4 FD 1 L 65 Pleurobema rubrum WD R 1 1 Pleuronaia barnesiana WD 3 FD FD 5 2 1 WD 31 Pleuronaia dolabelloides R 3 15 FD FD FD L 66 Potamilus alatus 3 1 2 1 FD L 132 Ptychobranchus fasciolaris FD FD FD R 21 Pyganodon grandis 1 Quadrula c.cylindrica* 3 2 4 6 FD L 218 Quadrula pustulosa WD WD FD WD 1 FD FD L 2 Quadrula verrucosa R FD R 149 Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 364 Paint Rock River mainstem (RM) Estill Fork (RM) Species 30.0 29.2 27.6 25.2 21.2 20.6 19.0 18.2 16.7 15.5 13.1 5.6 3.6 2.8 2.2 1.0 Totals Toxolasma cylindrellus* 1 WD R 1 Toxolasma lividum FD FD FD 2 WD 2 FD 2 1 L WD 1 19 Truncilla donaciformis 2 Truncilla truncata 1 3 Villosa iris FD FD 1 FD FD 1 FD FD FD FD WD 4 4 WD 24 Villosa taeniata 1 18 Villosa vanuxemensis 1 2 FD FD FD FD WD FD 4 FD WD 19 Search time (ph) 1.25 1.0 0.75 1.0 3.0 3.0 5.3 1.0 1.25 1.5 1.0 1.5 1 0.75 1.0 1.25 Total number of mussels 22 2 2 5 38 21 108 22 6 6 N/A 12 5 2 7 1 1825 CPUE (mussels/hour) 17.6 2 2.7 5 12.7 7 20.3 22 4.8 4 N/A 8 5 2.7 7 0.8 Total # of species (L, FD) 10 7 5 5 22 16 22 17 13 9 23 5 2 1 4 1 Southeastern Naturalist 365 T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 Appendix 2. Historical species documentation and PRR mussel-survey comparisons. W = Williams et al. (2008) , O = Ortmann (1925, At = Athearn (1957–1969) collections from North Carolina State Museum of Natural Sciences (Johnson et al. 2008), I & Y = Isom and Yokley (1973), Ah 86 = Ahlstedt (1986), Ah 95 = Ahlstedt (1995), G = Godwin (2002), and PS = present survey (2008). *Not found during this survey but collected by P. Freeman (TNC) during survey period. **Not found during this survey but subsequently collected by the authors and P. Freeman. Species W O At I & Y Ah 86 Ah 95 G PS Actinonaias ligamentina X 2 Actinonaias pectorosa X Alasmidonta marginata X X Alasmidonta viridis X X 2 1 Amblema plicata X X X X 27 182 79 548 Cyclonaias tuberculata X X X 99 110 Ellipsaria lineolata X X X 4 1 Elliptio crassidens X X X X 10 11 Elliptio dilatata X X X X 1 1 Epioblasma biemarginata X X Epioblasma capsaeformis X X X X 1 Epioblasma florentina X Epioblasma lenior X X Epioblasma triquetra X X X 12 21 Fusconaia cor X X X X 16 30 4 6 Fusconaia cuneolus X X X X 1 2 Fusconaia ebena X X 1 Fusconaia subrotunda X X 10 47 Lampsilis abrupta X FD* Lampsilis fasciola X X X X 8 26 1 38 Lampsilis ovata X X X X 24 60 12 144 Lampsilis teres X X 4 4 Lampsilis virescens X X X X 8 1 4 Lasmigona complanata X X** Lasmigona costata X X X X 12 37 7 69 Lasmigona holstonia X 1 50 Leptodea fragilis X X X X 8 16 Ligumia recta X X X 1 Medionidus conradicus X X X 6 2 3 1 Megalonaias nervosa X X 1 47 16 Obliquaria reflexa X X X 2 37 8 Obovaria subrotunda X X X X 7 9 FD Pleurobema cordatum X X X 8 1 Pleurobema oviforme X X X X 2 1 65 Pleurobema plenum X X Pleurobema rubrum X 1 Pleurobema sintoxia X Pleuronaia barnesiana X X X X 8 1 48 31 Pleuronaia dolabelloides X X X X 14 309 4 50 Potamilus alatus X X X X 20 103 6 132 Potamilus ohiensis X Ptychobranchus fasciolaris X X X X 12 29 1 21 Pyganodon grandis X X X 1 1 Quadrula c.cylindrica X X X X 2 40 1 218 Quadrula metanevra X X 4 Southeastern Naturalist T.B. Fobian, M.L. Buntin, J.T. Holifield, T.A. Tarpley, J.T. Garner, and P.D. Johnson 2014 Vol. 13, No. 2 366 Species W O At I & Y Ah 86 Ah 95 G PS Quadrula pustulosa X X X X 1 14 2 Quadrula quadrula X Quadrula verrucosa X X X X 8 8 149 Toxolasma cylindrellus X X 3 1 5 1 Toxolasma lividum X X X X 3 82 2 19 Toxolasma parvum X Truncilla donaciformis X X X X 2 Truncilla truncate X X X X 3 3 Utterbackia imbecillis X X FD* Villosa iris X X X X 14 109 80 24 Villosa taeniata X X X X 24 10 18 Villosa trabalis X X Villosa vanuxemensis X X X X 4 81 17 19 Total species 57 39 39 30 25 37 19 41