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2019 SOUTHEASTERN NATURALIST 18(2):346–372
Freshwater Mussels (Bivalvia: Margaritiferidae and
Unionidae) of the Buffalo River Drainage, Tennessee
Matthew P. Reed1,*, Gerald R. Dinkins2, and Steven A. Ahlstedt3
Abstract - The Buffalo River in Tennessee historically contained a rich diversity of freshwater
mussels. Sampling efforts in the 1980s documented declines in most of the main channel.
Recent collection data indicated recovery in the upper and lower reaches of the river. The objective
of this study was to update the current status, distribution, and species composition of
the mussel fauna in the main channel and major tributaries through qualitative sampling, and
document community structure using quantitative sampling at the most diverse location in the
main channel for use in future monitoring efforts. In the qualitative sampling portion of this
study, timed searches established catch-per-unit effort (CPUE) at intervals of ~5 river miles
in the main channel and tributaries. We recorded a total of 36 species at 62 sites, including 3
federally protected species: Margaritifera monodonta (Spectaclecase), Pleuronaia dolabelloides
(Slabside Pearlymussel), and Theliderma cylindrica (Rabbitsfoot). An additional 3
species being considered for the federal endangered species list were also found extant in the
main channel: Obovaria subrotunda (Round Hickorynut), Pleuronaia barnesiana (Tennessee
Pigtoe), and Toxolasma lividum (Purple Lilliput). Multiple sites in the upper and lower mainstem
were suitable for reintroduction of species. These findings should be considered in future
management and conservation efforts.
Introduction and Historical Review
Tennessee is one of the most biologically rich inland states and has the highest
number of freshwater fish species (Boschung and Mayden 2004, Etnier and Starnes
1993) and the second highest number of freshwater mussels among US states (Parmalee
and Bogan 1998, Williams et al. 2008). Parmalee and Bogan (1998) reported
129 mussel taxa historically known to occur in Tennessee, and subsequent taxonomic
changes and recent surveys have raised this number to 139 (G.R. Dinkins,
unpubl. data). To date, Tennessee has lost roughly 26 taxa to extinction or statewide
extirpation, lowering the total number of extant mussel species to 113, of which 3
have uncertain status (G.R. Dinkins, unpubl. data).
The Buffalo River joins the Duck River ~15.5 river miles upstream of the confluence
with the Tennessee River, and is the largest tributary to the Duck River.
The Duck River has long been recognized as exceptionally diverse, and is home to
roughly 151 species of fish, 66 species of mussels, and 22 species of aquatic snails
(Ahlstedt et al. 2017, Etnier and Starnes 1993, Hubbs et al. 2011). In contrast, the
Buffalo River has received little attention except for occasional surveys at a few
isolated locations on the main channel.
1Tennessee Department of Transportation, 7512 Volkswagen Drive, Chattanoooga, TN
37416. 2McClung Museum of Natural History and Culture, University of Tennessee, 1327
Circle Park Drive, Knoxville, TN 37996. 3PO Box 460, Norris, TN 37828. *Corresponding
author - matthew.reed@tn.gov.
Manuscript Editor: Paul M. Stewart
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Only a few published surveys have documented the historical mussel fauna
of the Buffalo River. Arnold Ortmann visited the Buffalo River in 1922 during a
mussel study of the Duck River drainage (Ortmann 1924) and sampled near Riverside
(Buffalo River Mile [BRM] 80.5). He noted a substantial mussel fauna, and
reported 20 species. In 1931, Calvin Goodrich and Henry van der Schalie sampled
BRM 80.5, BRM 41.0, BRM 31.6, and BRM 19.1. Their results, published over
40 y after the survey, added 13 species to Ortmann’s list (van der Schalie 1973).
Goodrich and van der Schalie’s most speciose site was BRM 19.1 (near Lobelville),
where they reported 29 species. Van der Schalie (1973:49) noted that “the Duck and
Buffalo rivers had a surprisingly rich mussel fauna, both in the numbers of species
and in individuals”. Based on the number of mussels found at BRM 41.0, BRM
31.6, and BRM 19.1, Van der Schalie (1973:49) concluded the Duck and Buffalo
Rivers “have some of the finest shoals in the world but, as previously indicated, the
mussels have now been depleted drastically”.
The decline of the mussel fauna in the Buffalo River was also noted by Isom and
Yokley (1968). In their field notes accompanying speciments deposited in the Museum
of Biological Diversity at Ohio State Universtiy (OSUM), they wrote, “The
Buffalo is a clear, cool, rapidly flowing small river with no evident pollution. In spite
of limestone bedrock, cobbles, and gravel, the mollusks were not abundant.” Isom
and Yokley found no live mussels or dead shells when they visited BRM 80.5, but
they found 16 species at BRM 74.8 and added several new species to the historical
record. David Stansbery visited the Buffalo River in 1972 and made this entry in his
fieldnotes accompanying speciments deposited in OSUM: “Water low, clear and cool
… Naiades [freshwater mussels] all but absent, a single Toxolasma lividus lividus
[= Toxolasma lividum Rafinesque, 1831] was taken alive along with a few dead shells
of other species.” In 1980, biologists from the Tennessee Valley Authority (TVA)
sampled 24 sites on the Buffalo River as part of the Cumberlandian Mollusk Conservation
Program and Columbia Dam project on the Duck River (Ahlstedt 1991). They
found only 3 or fewer live species at any site, and most sites yielded none. Initially,
more sites were to be surveyed, but the study was terminated because of the paucity
of mussels encountered. These surveys did not investigate the reason(s) for the decline
of the mussel fauna in the Buffalo River, but Fitz (1973) and Mast and Turk
(1999) suggested the decline was associated with degraded water quality caused by
anthropogenic activities.
In 2002, a survey of the Duck River drainage included 5 sites in the Buffalo
River previously surveyed by earlier investigators (Ahlstedt et al. 2017). Of the 9
species found across the 5 sites, none represented a new species for the river. Those
authors observed severe substrate destabilization and silt accumulation, and mussels
were extremely rare.
Despite the lack of a drainage-wide survey, based on these reports, the Buffalo
River was known to have contained 43 species, including 7 that are now federally
endangered: Epioblasma ahlstedti (Duck River Dartersnapper), E. aureola
(Golden Riffleshell), Hemistena lata (Cracking Pearlymussel), Pleuronaia dolabelloides
(Slabside Pearlymussel), Ptychobranchus subtentus (Fluted Kidneyshell),
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Theliderma cylindrica (Rabbitsfoot), and Toxolasma cylindrellus (Pale Lilliput)
(Appendix 1). In 2012, we recognized the need for a thorough survey of the Buffalo
River when biologists from the TVA brought several fresh dead mussels collected
during a fish survey in the lower reach to the McClung Museum of Natural History
and Culture at the University of Tennessee. In this sample of shells, we identified 2
species not previously reported from the Buffalo River: Margaritifera monodonta
(Spectaclecase [endangered]) and Ellipsaria lineolata (Butterfly). The sample also
included Eurynia dilatata (Spike), which had not been seen in the Buffalo River
since 1922 (Ortmann 1924), Cyclonaias pustulosa (Pimpleback), and Pleuronaia
dolabelloides, not seen since 1931 (van der Schalie 1973).
Study Area
The Buffalo River is located in western Middle Tennessee in the Western
Highland Rim of the Interior Low Plateau physiographic province (Mast and Turk
1999) and is one of the state’s longest free-flowing rivers (201 km) (Fig. 1). Rising
in Lawrence County, the river flows west through Lewis and Wayne counties,
then turns north to flow through Perry and Humphreys counties, draining ~1227
km2. The upper Buffalo River, in Lawrence County, is designated a “State Scenic
River” under the Tennessee State Scenic Rivers Act. The Western Highland Rim is
highly dissected by perennial streams that create irregular topography consisting
of peneplain ridgelines and flat-bottomed drainages, often separated by steep-sided
ridges. The Western subdivision of the Highland Rim province encompasses the
entire Buffalo River, and varies in elevation from 157 m to 305 m. Mississippian
limestone, chert, sandstone, and shale compose primary deposits, with sinkholes,
caves, and karst topography occurring throughout the region (Smalley 1981).
Mean temperatures vary from 8.8 °C in January to 31.6 °C in July. Mean annual
precipitation is 142 cm, with peak precipitation occurring in May and December.
May is generally the wettest month and averages 15.5 cm of precipitation. The driest
months are August through October. Mast and Turk (1999) and the Tennessee
Department of Environment and Conservation (TDEC 2005) described land use in
the Buffalo River watershed as primarily forest (77%) and agriculture (18%). Over
90% of the watershed is private land, and urban development constitutes less than
1%. Despite its rural setting, the Buffalo River drainage experienced considerable
development from the mid-1960s to mid-1980s.
Methods
Museum records
We collected records from several museums to supplement the list of species in
past survey reports: Carnegie Museum of Natural History (CM), McClung Museum
of Natural History and Culture at the University of Tennessee (UTMM), North
Carolina State Museum of Natural Sciences (NCMNS), Museum of Biological Diversity
at The Ohio State University (OSUM), and University of Michigan Museum
of Zoology (UMMZ).
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Qualitative sampling
We sampled 47 sites in the main channel Buffalo River and 15 sites in tributary
streams between October 2012 and December 2015 (Table 1, Fig. 1). We selected
sampling locations surveyed in previous investigations and new sites that we considered
to possess suitable habitat. Each locality was recorded in decimal degrees
Figure 1. Sampling localities for mussels in the Buffalo River drainage in Lawrence, Lewis,
Wayne, Perry, and Humphreys counties, TN.
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Table 1. Location of mussel survey sites in the Buffalo River system, TN. [Table continued on following
page.]
Site/location Latitude (°N) Longitude (°W) River-mile Date
1 Buffalo River 35.9957330 87.8407500 0.2 11 October 2012
2 Buffalo River 35.9894400 87.8555600 1.9 16 November 2012
3 Buffalo River 35.9850500 87.8603100 2.2 16 November 2012
4 Buffalo River 35.9784300 87.8626100 2.8 16 November 2012
5 Buffalo River 35.9746800 87.8627500 3.2 16 November 2012
6 Buffalo River 35.9722400 87.8577100 3.5 17 November 2012
7 Buffalo River 35.9657000 87.8541400 4.0 17 November 2012
8 Buffalo River 35.9567100 87.8524000 4.6 17 November 2012
9 Buffalo River 35.9517200 87.8522000 4.9 17 November 2012
10 Buffalo River 35.9316590 87.8501960 6.8 11 October 2013
11 Buffalo River 35.9195000 87.8461000 8.0 11 October 2013
12 Buffalo River 35.8766670 87.8303500 12.0 11 October 2013
13 Buffalo River 35.8576170 87.8135670 13.8 30 August 2013
14 Buffalo River 35.8279670 87.8086830 16.3 30 August 2013
15 Buffalo River 35.8121830 87.7964500 17.7 30 August 2013
16 Buffalo River 35.8126170 87.7759000 19.4 30 August 2013
17 Buffalo River 35.7867330 87.7766670 22.4 29 August 2013
18 Buffalo River 35.7620830 87.7727500 25.8 29 August 2013
19 Buffalo River 35.7049100 87.7937700 32.0 10 August 2013
20 Buffalo River 35.6880300 87.8026200 34.3 10 Aug 2013
21 Buffalo River 35.6154300 87.8320300 41.4 9 August 2013
22 Buffalo River 35.6040700 87.8387900 42.8 9 August 2013
23 Buffalo River 35.5827400 87.8385200 45.6 9 August 2013
24 Buffalo River 35.5432900 87.8201000 52.1 9 August 2013
25 Buffalo River 35.5235500 87.8419670 55.1 18 December 2012
26 Buffalo River 35.5206330 87.8435670 55.6 18 December 2012
27 Buffalo River 35.5125267 87.8395990 56.0 18 December 2012
28 Buffalo River 35.5050500 87.8266330 57.4 18 December 2012
29 Buffalo River 35.5004000 87.8322330 58.0 18 December 2012
30 Buffalo River 35.4890835 87.8342990 59.0 18 December 2012
31 Buffalo River 35.4657500 87.8476000 62.7 21 June 2013
32 Buffalo River 35.4734830 87.8148670 67.4 21 June 2013
33 Buffalo River 35.4556670 87.8029500 70.6 21 June 2013
34 Buffalo River 35.4551000 87.7737670 73.5 25 May 2013
35 Buffalo River 35.4343500 87.7210000 78.7 25 May 2013
36 Buffalo River 35.4360170 87.6990500 80.5 25 May 2013
37 Buffalo River 35.4427170 87.6451170 87.0 24 May 2013
38 Buffalo River 35.4521500 87.6030170 90.6 24 May 2013
39 Buffalo River 35.4497700 87.5690900 93.6 17 July 2013
40 Buffalo River 35.4625800 87.5352000 98.2 17 July 2013
41 Buffalo River 35.4681200 87.5012900 100.6 17 July 2013
42 Buffalo River 35.4645100 87.4786700 102.9 16 July 2013
43 Buffalo River 35.4654800 87.4652900 105.3 17 July 2013
44 Buffalo River 35.4367000 87.4232300 111.3 17 July 2013
45 Buffalo River 35.3971400 87.3877100 116.7 18 July 2013
46 Buffalo River 35.3939800 87.3111500 121.0 18 July 2013
47 Buffalo River 35.3911100 87.2994600 122.0 18 July 2013
48 Cane Creek 35.7048170 87.7554670 3.4 11 December 2012
49 Cane Creek 35.7086670 87.7601500 2.4 11 December 2012
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using a hand-held global positioning system unit (Garmin GPSMAP 76Cx unit,
WAAS enabled) and is reported in river miles because this is the increment used in
US Geological Survey 7.5 minute topographic maps.
Mussels encountered at each sampling location were removed from the substrate
and held in a submerged mesh bag until they were sorted and identified. Fresh-dead
(shiny nacre, hinge ligament intact) and relic shells (weathered periostracum, faded
or chalky nacre) were identified, counted, recorded on a field data sheet, and taken
to UTMM where they were cleaned, verified, and catalogued. Live mussels were
measured (anterior to posterior in millimeters) using dial calipers, photographed,
and returned to the substrate. Each site was surveyed by at least 2 experienced biologists
for 1 h or more. Shorelines and adjacent submerged habitats were searched
for shells deposited by Ondatra zibethicus (L.) (Muskrat) or high water.
Quantitative sampling
Based on the results of the qualitative survey, we chose the most diverse and
abundant mussel community for quantitative sampling in November 2013 (shoal
at BRM 3.2). We established parallel transects in an area measuring 67 m long
and 35 m wide (2345 m2). Transects were oriented perpendicular to the shoreline,
spaced at 6-m intervals, and spanned the wetted channel. A total of 100 quadrats
(0.25 m2) evenly spaced 4 m apart along the transects was sampled using skin diving
gear and SCUBA, and each quadrat was excavated to an attempted minimum
depth of 15 cm. Water depth varied from a few centimeters along the right descending
bank to 2 m in the thalweg, located near the left descending bank.
Analyses
We calculated species richness, relative abundance, and CPUE (determined as
number of observed live and fresh dead mussels/snorkel time per observer x number
of observers) for each site. Numerous studies involving freshwater mussels
have documented the potential bias of selecting for larger individuals in qualitative
versus quantitative sampling methods (Hornbach and Deneka 1996, Miller and
Table 1, continued.
Site/location Latitude (°N) Longitude (°W) River mile Date
50 Cane Creek 35.7077819 87.7575284 2.6 11 December 2012
51 Cane Creek 35.6992830 87.7475330 4.1 11 December 2012
52 Brush Creek 35.6751170 87.8015000 0.1 12 December 2012
53 Coon Creek 35.6349000 87.8045330 1.3 12 December 2012
54 Coon Creek 35.6375500 87.8088670 0.9 12 December 2012
55 Coon Creek 35.6377000 87.8101801 0.7 12 December 2012
56 Lower Opossum Creek 35.4878336 87.8325710 0.1 18 December 2012
57 Hurricane Creek 35.5721100 87.8019500 0.9 10 August 2013
58 Green River 35.4299440 87.7750700 2.3 19 December 2015
59 Green River 35.3431100 87.7592800 10.6 19 December 2015
60 Fortyeight Creek 35.4040640 87.6667760 4.4 19 December 2015
61 Little Buffalo River 35.4143080 87.5105710 4.8 19 December 2015
62 Little Buffalo River 35.3261060 87.4866630 15.3 19 December 2015
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Payne 1993, Miller et al. 1993, Obermeyer 1998). To test for this potential bias,
we conducted a 2-tailed Kolmogorov–Smirnov (K–S) nonparametric test (Smirnov
1948) to examine size-class variance in qualitative versus quantitative sampling
data for the most abundant species, Pleuronaia dolabelloides. We hypothesized
qualitative size-class data would vary significantly from quantitative size-class
data. We conducted our analyses in XLSTAT statistical software. Taxonomy follows
Williams et al. (2017).
Results
Museum records
The search for Buffalo River specimens at 5 museums yielded a total of 32
species, including 4 new records from 16 collecting localities (Appendix 2):
Epioblasma turgidula (Turgid Blossom), Megalonaias nervosa (Giant Washboard),
Ligumia subrostrata (Pondmussel), and Utterbackiana suborbiculata
(Flat Floater). Epioblasma turgidula (2 males and 1 female) and Megalonaias
nervosa were collected by H. Athearn in 1966 at the Gilmore Bridge near Lobelville
(BRM 22.6) (J. Smith, NCMNS, Raleigh, NC, pers. comm.). Athearn also
collected M. nervosa at the US Route 412 bridge near Linden (BRM 41.0). In
July 1994, S. Ahlstedt found Ligumia subrostrata (Pondmussel) in an overflow
channel of the Buffalo River at Mill Bridge (BRM 24.0). There are only a few
records of this species from Tennessee, but it is found throughout the Mississippi
River basin (Parmalee and Bogan 1998). Ligumia subrostrata typically occurs
in areas with little to no current (Williams et al. 2008), and it seems to be particularly
adaptable to newly created ponds or channels, where it can be abundant.
Its presence in an overflow channel of the Buffalo River, while well upstream of
its previously known range in the Tennessee River drainage, is consistent with
its habitat preference. Utterbackiana suborbiculata (Flat Floater) was found at
Beardstown (BRM 32.6) in 1973. A large, single valve of this species was found
in the Buffalo River and is cataloged at UTMM. Utterbackiana suborbiculata appears
to be expanding its range upstream in both the Tennessee and Cumberland
river drainages (Bates 1962, Parmalee and Bogan 1998) and it is now found in the
Tennessee River as far upstream as Knoxville and in the Cumberland River as far
upstream as Hartsville (G. Dinkins, pers. observ.). Williams et al. (2008) speculated
U. suborbiculata has spread into the Tennessee River due to conditions
associated with impoundment of the formerly free-flowing river. While the Buffalo
River is completely unimpounded, Kentucky Lake impounds the lower Duck
River upstream to its confluence with the Buffalo River.
Qualitative results
We found a total of 33 species in the qualitative survey; 2 additional species
were found after the qualitative survey was completed and were added to the
qualitative survey results (Appendix 3). We observed live or fresh dead mussels at
30 of the 47 main channel sites. By far, the most diverse sites were BRM 3.2 and
BRM 3.5. We found no live mussels at the 5 most upstream sites (BRM 105.3 to
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BRM 122.0). Mussel diversity varied in the main channel, and although mussels
were sparse in the middle reaches compared to historical sampling efforts, we detected
significant increases in the upper and lower Buffalo River. We documented
Lampsilis fasciola (Wavyrayed Lampmussel) and Villosa taeniata (Painted
Creekshell) at 18 and 21 sites, respectively; these were the most abundant and
widespread species. Species richness and CPUE were correlated throughout the
river. Sites in the middle portion of the river (BRM 30.0 to BRM 98.0) supported
few live mussels. Species richness was similarly low in this stretch of the river.
Tributaries to the Buffalo River (Sites 48 to 60, 62) were devoid of mussels except
for a single relic Strophitus undulatus (Creeper) found in the Little Buffalo
River at Site 61. We did not observe Strophitus undulatus in the main channel of
the Buffalo River. The apparent lack of mussels in Buffalo River tributaries mirrors
recent findings from sampling in tributaries of the Duck River (K. Irwin,
Tennessee Tech University, Cookeville, TN, unpubl. data). For this reason, tributary
sampling results are not included in Appendix 3.
Length–frequency data indicated recent recruitment for V. taeniata (20–89 mm),
L. fasciola (56–86 mm), Pleuronaia barnesiana (Tennessee Pigtoe) (27–59 mm),
and P. dolabelloides (36–70 mm). We observed multiple size-classes of Potamilus
alatus (Pink Heelsplitter) and T. cylindrica, but there were insufficient numbers to
infer recent recruitment. During quantitative sampling at BRM 3.2, a large, submerged
muskrat midden was located just outside of the study area. After a brief
assessment, it became evident this midden contained a large number of fresh dead
shells; these numbers are omitted from the study results.
Quantitative results
Quadrat excavations yielded 178 live individuals representing 20 species and an
average of 1.8 mussels per 0.25-m2 quadrat (Table 2). We collected 1 live Actinonaias
ligamentina (Mucket) in the quadrat excavations, which was the only individual
of the species found in our study. Two federally protected species, P. dolabelloides
and T. cylindrica, comprised nearly 30% of the live mussels in the samples;
of these, P. dolabelloides was the most abundant (26.4%). There were multiple
size-classes (11–65 mm) of this species, which indicated recent recruitment. Other
species with multiple size-classes included Cyclonaias pustulosa, Cyclonaias tuberculata
(Purple Wartyback), and Eurynia dilatata. Evidence of recent recruitment
was inconclusive for Lampsilis fasciola, P. barnesiana, T. cylindrica, Tritogonia
verrucosa (Pistolgrip), and Truncilla truncata (Deertoe) due to insufficient number
of individuals.
There was no significant difference in size-classes of P. dolabelloides in qualitative
versus quantitative data (P = 0.093), suggesting qualitative sampling methods
were not biased toward detecting larger individuals.
Species of conservation concern
Margaritifera monodonta (Spectaclecase).
Margaritifera monodonta was once widespread in the Mississippi, Ohio, and
Missouri River basins from Minnesota and western Pennsylvania south to the Gulf
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of Mexico (Parmalee and Bogan 1998). It was listed as federally endangered in
2012 (USFWS 2012). Prior to this study, it was not known to occur in the Buffalo
River. Two live individuals were found in 1998 in the Tennessee River at RM 170,
approximately 60 river-miles downstream of the mouth of the Duck River (Hubbs
and Jones 2000), and in the early 2000s, a single individual was observed in the
Duck River (USFWS 2012). To our knowledge, no additional individuals of this
species have been found in the Duck River, and it appears to be equally rare in the
Buffalo River. The periostracum of the specimen brought to us by the TVA fishsampling
crew was fairly weathered but not overly so, and the nacre was lustrous.
We cannot precisely estimate how much time had elapsed since its death, but we
have found shells of M. monodonta from other locations in the Tennessee River
drainage that appeared weathered soon after death; consequently, it is likely the
specimen found by the TVA was alive within 1 y of its shell being found, and from
this we infer that there is an extant but extremely small population remaining in the
Buffalo River.
Epioblasma ahlstedti (Duck River Dartersnapper).
Epioblasma ahlstedti historically occurred in the Duck and Buffalo rivers,
as well as the Tennessee River at Muscle Shoals and Shoal Creek, a large direct
tributary to the Tennessee River (Jones 2004). Currently, it is restricted to ~48.3
river km in the Duck River, from the Old Columbia Dam upstream to Lillard Mill
Table 2. Summary of quantitative sampling at BRM 3.2. Number = number observed among 100
quadrats. % Occurrence = percent of occurrence across all quadrat samples. Number of quadrats =
number of quadrats occupied by species. Size (min–max if more than 1 observed) is presented for
live individuals only.
Species Number % occurrence # of quadrats Size (mm)
Actinonaias ligamentina 1 0.6 1 49
Actinonaias pectorosa 1 0.6 1 14
Cyclonaias pustulosa 15 8.4 10 20–61
Cyclonaias tuberculata 46 26.0 25 18–99
Ellipsaria lineolata 3 1.7 2 22–54
Elliptio crassidens 3 1.7 3 111–141
Eurynia dilatata 20 11.2 15 15–85
Lampsilis fasciola 6 3.3 5 47–82
Lampsilis ovata 2 1.1 2 16–74
Lasmigona costata 2 1.1 2 82–149
Megalonaias nervosa 1 0.6 1 88
Obliquaria reflexa 1 0.6 1 29
Pleuronaia barnesiana 5 2.8 5 18–59
Pleuronaia dolabelloides 47 26.4 28 11–65
Potamilus alatus 2 1.1 2 109–113
Theliderma cylindrica 6 3.3 5 64–94
Toxolasma lividum 1 0.6 1 16
Tritogonia verrucosa 6 3.3 4 64–89
Truncilla truncata 9 5.0 8 18–37
Villosa taeniata 1 0.6 1 84
Total 178
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(Jones and Neves 2010, Parmalee and Bogan 1998). Its historical occurrence in
the Buffalo River is based on 1 specimen found in the Buffalo River in 1931 (van
der Schalie 1973) near the Humphreys/Perry County border (approximate BRM
19.0). We did not find this species in this survey. Epioblasma ahlstedti was listed
as federally endangered in 1997 (as Epioblasma capsaeformis [Oyster Mussel];
USFWS 1997).
Epioblasma aureola (Golden Riffleshell).
Epioblasma aureola presumably occurred in numerous tributary streams of the
middle and upper Tennessee River system downstream to the Duck and Buffalo rivers,
but is now thought to occur only in Indian Creek, a tributary to the upper Clinch
River in southwestern Virginia (Jones and Neves 2010). Its historical occurrence in
the Buffalo River is based on 3 specimens found in the Buffalo River in 1931 (van
der Schalie 1973) near the Humphreys/Perry County border (approximate BRM
19.0). We did not find this species in this survey. Epioblasma aureola was listed as
federally endangered in 1977 (as Epioblasma florentina walkeri [Wilson and Clark]
[Tan Riffleshell]; USFWS 1977).
Epioblasma turgidula (Turgid Blossom).
Epioblasma turgidula appears to have been endemic to the Tennessee and Cumberland
River drainages in Alabama and Tennessee, where it occurred in numerous
large tributary rivers and streams (Parmalee and Bogan 1998, Williams et al. 2008).
H. Athearn collected 3 individuals of this species (presumably alive) in September
1966 in the Buffalo River at Gilmore Bridge (BRM 22.6) (J. Smith, pers. comm.).
Epioblasma turgidula is presumed extinct and was last seen alive in 1972 in the
upper Duck River (Stansbery 1976). Johnson (1978) included records for the species
from the St. Francis River in Arkansas, but its taxonomic status outside of
the Tennessee and Cumberland systems is uncertain and yet unresolved (J. Harris,
Arkansas State University, Jonesboro, AR, pers. comm.). It was listed as federally
endangered in 1976 (USFWS 1976).
Hemistena lata (Cracking Pearlymussel).
Hemistena lata is restricted to the Ohio River drainage, including the Tennessee
and Cumberland rivers (Watters et al. 2009). There are 2 records from the Buffalo
River: 1 individual was collected at BRM 19.1 in 1931 (van der Schalie 1973) and
an unknown number were collected at BRM 74.8 in 1965 (Isom and Yokley 1968).
We did not find H. lata in this survey. Hemistena lata was listed as federally endangered
in 1989 (USFWS 1989).
Obovaria subrotunda (Round Hickorynut).
Obovaria subrotunda is known from parts of the Great Lakes Basin and is
widespread in the Tennessee, Cumberland, and Ohio River drainages (Haag and
Cicerello 2016, Parmalee and Bogan 1998, Watters et al. 2009, Williams et al.
2008). Obovaria subrotunda was first reported from the Buffalo River in 1922 (Ortmann
1922), and it was subsequently collected in 1931 and 1965 (Isom and Yokley
1968, van der Schalie 1973). We found 1 live individual at BRM 3.2 during a timed
search but did not find it in the quantitative sampling at this same location. The
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USFWS is currently conducting a status assessment to determine if O. subrotunda
warrants federal protection under the Endangered Species Act (ESA).
Pleuronaia barnesiana (Tennessee Pigtoe).
Pleuronaia barnesiana occurs in medium-sized creeks to large rivers in the
Tennessee River drainage (Parmalee and Bogan 1998, Williams et al. 2008). Pleuronaia
barnesiana was relatively widespread in the Buffalo River. We found live
or dead shells at 13 locations from just upstream of the Duck River confluence to
BRM 102.9. In the quantitative survey at BRM 3.2, P. barnesiana comprised ~3%
of the live mussels found in the quadrats and several age classes were evident. The
USFWS is currently conducting a status assessment to determine if P. barnesiana
warrants federal protection under the ESA.
Pleuronaia dolabelloides (Slabside Pearlymussel).
Pleuronaia dolabelloides is endemic to the Tennessee and Cumberland River
drainages and was listed as federally endangered in 2013 (USFWS 2013a). This
species was one of the most abundant and wide-ranging species we found in the
Buffalo River. Individuals collected from quadrats at BRM 3.2 exhibited a healthy
size-class distribution (11–65 mm), with strong evidence of recent recruitment.
Currently, healthy communities of P. dolabelloides persist in the upper and lower
reaches of the Buffalo River (e.g., BRM 102.9 and BRM 3.2). Clinal variation in
shell morphology for this species is consistent with observations made by Ortmann
(1924) and Van der Schalie (1973); individuals collected upstream (e.g., BRM
102.9) were notably compressed compared to individuals at downstream sites (e.g.,
BRM 3.2).
Pleurobema oviforme (Tennessee Clubshell).
Pleurobema oviforme historically occurred in small to large rivers in the Tennessee
and Cumberland River drainages, but it has disappeared from much of its
historical range and is in danger of disappearing from Kentucky (Haag and Cicerello
2016). We observed relic shells of P. oviforme at 2 sites during this study—
BRM 3.2 and BRM 3.5. The USFWS is currently conducting a status assessment to
determine if P. oviforme warrants federal protection under the ESA.
Ptychobranchus subtentus (Fluted Kidneyshell).
Ptychobranchus subtentus is endemic to the Tennessee and Cumberland River
drainages (Parmalee and Bogan 1998). It appears to be extirpated from Alabama
(Garner et al. 2004), and populations have declined dramatically in Kentucky
(Haag and Cicerello 2016). We did not find P. subtentus in this survey, and it has not
been seen in the Buffalo River since 1922 at BRM 90.4 (Ortmann 1924). Ptychobranchus
subtentus was listed as federally endangered in 2013 (USFWS 2013a).
Theliderma cylindrica (Rabbitsfoot).
Theliderma cylindrica is endemic to the Tennessee River drainage and was
listed as threatened in 2013 (USFWS 2013b). The Buffalo River was not included
as critical habitat for T. cylindrica, and a recent status update assumed it was extirpated
from the river (USFWS 2015). During our qualitative search, T. cylindrica
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was found to be mostly restricted to the lower reach of the Buffalo River. We
documented live individuals at BRM 2.2, BRM 3.2, and BRM 17.7. Quantitative
sampling at BRM 3.2 produced 6 T. cylindrica from 5 quadrat samples.
Toxolasma cylindrellus (Pale Lilliput).
Toxolasma cylindrellus is endemic to the middle and lower Tennessee River
drainage in Tennessee and Alabama (Parmalee and Bogan 1998, Williams et al.
2008). It was thought to have been extirpated from the Duck River, but was rediscovered
in 2015 in Lick Creek, a tributary to the Duck River in Maury County (D.
Hubbs, Tennessee Wildlife Resource Agency, Camden, TN, pers. comm.). Prior to
this discovery, the only known population occurred in a tributary of the Paint Rock
River in Alabama and Tennessee. Toxolasma cylindrellus was last seen in the Buffalo
River in 1922 at BRM 80.5 (Ortmann 1924), and we did not find the species
in this survey. Toxolasma cylindrellus was listed as federally endangered in 1976
(USFWS 1976).
Toxolasma lividum (Purple Lilliput).
Toxolasma lividum occurs in the Ohio River drainage, including the Cumberland
and Tennessee rivers, the Arkansas River drainage, and in the Lake Erie drainage
(Parmalee and Bogan 1998, Watters et al. 2009). In the Buffalo River, it was rare
and restricted to lower reaches where it occurred along the margins of backwater
areas and shallow pools in soft substrate. We found a single individual during
quantitative sampling at BRM 3.2. The taxonomic status of populations in the Cumberland
River drainage are under review (D. Campbell, Gardner-Webb University,
Boiling Springs, NC, pers. comm.), and the USFWS is currently conducting a status
assessment to determine if T. lividum warrants federal protection under the ESA.
The following species were not detected in our survey and may be extirpated
from the Buffalo River. The date of last occurrence is given in brackets following
the common name: Alasmidonta marginata (Say) (Elktoe) [1965], Alasmidonta
viridis (Rafinesque) (Slippershell Mussel) [1922], Lasmigona complanata (Barnes)
(White Heelsplitter) [1931], Ligumia subrostrata (Say) (Pondmussel) [1994],
Medionidus conradicus (Lea) (Cumberland Moccasinshell) [1968], Pleurobema
sintoxia (Rafinesque) (Round Pigtoe) [1931], Ptychobranchus fasciolaris (Rafinesque)
(Kidneyshell) [1965], Utterbackia imbecillis (Say) (Paper Pondshell)
[1931], and Utterbackia suborbiculata [1973].
Discussion and Recommendations
The Buffalo River is a rare example of a large unimpounded stream. Our review
of published literature, unpublished reports, and museum records along with our
survey results indicate that the river once contained 51 freshwater mussel species
(Appendix 4). Most of the river’s mussel fauna was nearly extirpated in the
years following the early surveys by Ortmann, which was documented in subsequent
surveys by Goodrich and van der Schalie, Isom and Yokley, and Ahlstedt.
As a result, management decisions and conservation efforts aimed at recovering
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2019 Vol. 18, No. 2
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imperiled species in the Tennessee River system have largely overlooked the Buffalo
River’s potential. The present survey recorded 36 species in the main channel
of the river and an additional species found a few months before fieldwork began
(Margaritifera monodonta). Of these, 29 species were represented by live or freshdead
material, and 8 species were represented by relic shells only. There were 14
species present in the historical record that we did not detect.
We rarely encountered the degree of species richness or abundance noted in
previous studies. In our qualitative survey, 22 species were represented by 10 or
fewer live or fresh dead individuals and 7 species were represented by relic shells
only. A substantial reach of the Buffalo River had few or no live mussels (BRM
32.0 to 98.2; BRM 105.3 to BRM 122.0). The middle reach from BRM 32.0 to
BRM 58.0 contained only L. fasciola and V. taeniata. The paucity of mussels in
the middle reach is comparable to the survey data from 1980 (Ahlstedt 1991). Of
the 30 species still present in the river, at least 8 species that historically occurred
in the upper reaches of the river are now restricted to the lower reaches: Actinonaias
ligamentina, Actinonaias pectorosa (Pheasantshell), Lampsilis cardium (Plain
Pocketbook), Lampsilis ovata (Pocketbook), Lasmigona costata (Flutedshell),
Leptodea fragilis (Fragile Papershell), Obovaria subrotunda, and T. verrucosa.
We observed obvious signs of stream-bank instability and significant erosion
and siltation, likely due to clearing of riparian vegetation adjacent to agricultural
operations, in various stretches of the river. In general, most of these reaches produced
low CPUE values. Another factor that may be inhibiting recovery of the
mussel communities is elevated levels of toxic metals. Denton (2007) reported
high levels of mercury in the tissue of Micropterus dolomieu (Lacepède) (Smallmouth
Bass) at BRM 17.7, which prompted TDEC to issue a fish-consumption
advisory for the lower Buffalo River extending from BRM 31.6 to the confluence
with the Duck River. Naimo (1995) reported that elevated levels of mercury can
alter growth, filtration efficiency, enzyme activity, and behavior in freshwater
mussels. Gravel dredging and unrestricted livestock access have also destabilized
many of the tributaries. We found no mussels in the tributaries except for a single,
relic shell of Strophitus undulatus in the Little Buffalo River at mile 4.8. Despite
reaches with unstable habitat and heavy sedimentation, the Buffalo River and its
tributaries have excellent fish diversity, (C. Saylor, TVA, Knoxville, TN, pers.
comm.), and support multiple federally listed fishes including Etheostoma boschungi
(Wall and Williams) (Slackwater Darter) and Erimonax monachus (Cope)
(Spotfin Chub) (Etnier and Starnes 1993).
Analysis of spatial distributions demonstrated several findings of immediate
conservation importance. Between BRM 3.2 to BRM 4.9, we found 18 species
that occurred nowhere else in the Buffalo River, and a number of species that are
apparently restricted to its lower reaches (e.g., Lampsilis cardium, Lampsilis teres
[Yellow Sandshell], Obliquaria reflexa [Threehorn Wartyback], Pleurobema cordatum
[Ohio Pigtoe], Theliderma cylindrica, Cyclonaias pustulosa, and Quadrula
quadrula [Mapleleaf]) have not been reported in the Buffalo River since 1931 (van
der Schalie 1973). We documented 5 new records for the drainage in this reach
(Margaritifera monodonta, Arcidens confragosus [Rock Pocketbook], Fusconaia
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subrotunda, [Longsolid], Ligumia recta, [Black Sandshell], and Theliderma
metanevra [Monkeyface]). With the exception of F. subrotunda, all were reported
in the lower Duck River by Schilling and Williams (2002).
In addition, the area extending from BRM 100.6 to BRM 102.9 should be
re-examined and monitored in the future; BRM 102.9 had populations of both
P. barnesiana and P. dolabelloides. We observed evidence of recent recruitment for
both species. Habitat at sites in this upper reach appeared excellent, with minimal
evidence of disturbance. We found no mussels above BRM 102.9, further increasing
the importance of preserving this isolated reach.
The continued monitoring and conservation of mussel assemblages identified in
this study is vital to future management of mussel resources in the Buffalo River.
Fish hosts of threatened and endangered mussels should also be monitored. Quadrat
excavations conducted at BRM 3.2 provide a baseline for future quantitative mussel
monitoring. Because it is a tributary to the Duck River, the Buffalo River has
the potential, with the proper attention and management, to regain much of its lost
mussel biodiversity. Sedimentation and channel instability appear to be the primary
factors impacting mussel habitat in the river. We strongly recommend that landowners
in the Buffalo River drainage be encouraged to implement best management
practices. Better management of riparian zones and limiting cattle movements near
main-channel mussel habitats could play a pivotal role in mitigating sedimentation
and habitat degradation in the Buffalo River.
Acknowledgments
We thank Larry Wilson for providing financial support for the fieldwork, and Michael
McKinney and Jess Jones for assistance with design of the quantitative study. Thanks to
Todd Amacker, Robert Eldridge, Meredith Hayes, Kristin Irwin, Drew Mallinak, Josh Peterson,
Jackson Sibley, Ashley Slater, and Dan Walker for providing assistance in the field, and
to Dinkins Biological Consulting, LLC, the University of Tennessee (UT) Fisheries Laboratory,
and the UT Outdoor Program for providing equipment. Thanks to Chuck Howard and
the TVA for additional support and insight, and to Susan Lanier, Jeanette Jones, and Craig
Phillips for mapping assistance. Museum records were provided by Tim Pearce at Carnegie
Museum, and Jamie Smith and Art Bogan at North Carolina State Museum of Natural
Sciences. Thanks especially to 3 anonymous reviewers for suggestions that improved this
manuscript significantly. Finally, we gratefully acknowledge the Mayberry family for allowing
us access to their farm on the lower reach of the Buffalo River on multiple occasions,
and for their keen interest in preserving the natural history of the Buffalo River.
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Appendix 1. List of mussel species documented in the Buffalo River, by survey year. 1922
= Ortmann (1924), 1931 = Van der Schalie (1973), 1965 = Isom and Yokley (1968), 1980 =
Ahlstedt (1991), 2002 = Ahlstedt et al. (2017), and 2011 = TVA field crew (unpubl. data).
Present
Species 1922 1931 1965 1980 2002 2011 study
Margaritiferidae
Margaritifera monodonta X
Unionidae
Actinonaias ligamentina X X X X
Actinonaias pectorosa X X X X X
Alasmidonta marginata X X
Alasmidonta viridis X
Amblema plicata X X X X
Arcidens confragosus X
Cyclonaias pustulosa X X X
Cyclonaias tuberculata X X X X X
Ellipsaria lineolata X X
Elliptio crassidens X X X X X
Eurynia dilatata X X X
Epioblasma ahlstedti X
Epioblasma aureola X
Fusconaia subrotunda X
Hemistena lata X X
Lampsilis cardium X X X
Lampsilis fasciola X X X X
Lampsilis ovata X X X X X
Lampsilis teres X X X
Lasmigona complanata X
Lasmigona costata X X X X X
Leptodea fragilis X X X
Ligumia recta X
Megalonaias nervosa X
Obliquaria reflexa X X
Obovaria subrotunda X X X X
Pleurobema cordatum X X X
Pleurobema oviforme X X X X
Pleurobema sintoxia X
Pleuronaia barnesiana X X X X
Pleuronaia dolabelloides X X X X
Potamilus alatus X X X X
Ptychobranchus fasciolaris X
Ptychobranchus subtentus X
Pyganodon grandis X X X X
Quadrula quadrula X X
Strophitus undulatus X X X
Theliderma cylindrica X X
Theliderma metanevra X
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Present
Species 1922 1931 1965 1980 2002 2011 study
Toxolasma cylindrellus X
Toxolasma lividum X X X
Tritogonia verrucosa X X X X
Truncilla truncata X X X
Utterbackia imbecillis X
Villosa iris X X X X X X
Villosa taeniata X X X X
Villosa vanuxemensis X X X X X X
Number of Sites Sampled 1 4 2 24 5 1 62
Total (48 species) 20 31 16 7 8 17 36
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Appendix 2. Mussel species from the Buffalo River and cataloged at Carnegie Museum of
Natural History (CM), University of Tennessee, McClung Museum of Natural History and
Culture (UTMM), North Carolina State Museum of Natural Sciences (NCMNS), Museum
of Biological Diversity at The Ohio State University (OSUM), and University of Michigan
Museum of Zoology (UMMZ).
Species CM UTMM NCMNS OSUM UMMZ
Actinonaias pectorosa X X X X
Alasmidonta marginata X X X
Alasmidonta viridis X
Amblema plicata X X X
Cyclonaias tuberculata X X X X
Elliptio crassidens X X
Epioblasma ahlstedti X
Epioblasma aureola X
Epioblasma turgidula X
Hemistena lata X
Lampsilis cardium X
Lampsilis fasciola X X X
Lampsilis ovata X X
Lasmigona costata X X
Ligumia subrostrata X
Megalonaias nervosa X
Obovaria subrotunda X X X
Pleurobema oviforme X X
Pleuronaia barnesiana X
Pleuronaia dolabelloides X X X
Potamilus alatus X
Ptychobranchus subtentus X
Pyganodon grandis X X
Theliderma cylindrica X
Strophitus undulatus X
Toxolasma cylindrellus X
Toxolasma lividum X X
Utterbackia imbecillis X
Utterbackiana suborbiculata X
Villosa iris X X
Villosa taeniata X X X
Villosa vanuxemensis X X X
Number of species 15 9 19 9 6
Number of locations from which 1 6 2 5 4
specimens originated
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Appendix 3. Summary of mussels found in the Buffalo River during qualitative collections
in 2012–2013, and an opportunistic collection made in 2016 at Buffalo River Mile (BRM)
3.2. Number of live and fresh dead mussels combined; number of relic shells in parentheses
and shells collected in 2016 in brackets. Catch-per-unit effort (CPUE) is based on combined
number of live and fresh dead found in qualitative sampling only and does not include opportunistic
collection. Percent collected (% coll.) is calculated using the combined number
of live mussels and dead shells. Species richness is the combined number of live mussels
and dead shells. AFederally protected species. BNew drainage record
Site 1 2 3 4 5 6 7 8 9 10 11
BRM 0.2 1.9 2.2 2.8 3.2 3.5 4.0 4.6 4.9 6.8 8.0
Species
Actinonaias pectorosa (1)
Amblema plicata 2 2 (1) (1)
Arcidens confragosusB [1]
Cyclonaias pustulosa 4 (41) 1
Cyclonaias tuberculata 1 3 (17) 2(2) 2(1) 1 (1)
Ellipsaria lineolataB 3 (1)
Elliptio crassidens 2
Eurynia dilatata 2 (1) 2(2) (14) (5) 1(1) 1(1) 1(2)
Fusconaia subrotundaB (2)
Lampsilis cardium 1(1)
Lampsilis fasciola 2 1(1) (1) 2(1) (3) 1(2) 1(2) 2(1) 1
Lampsilis ovata 1 (2) 1
Lampsilis teres [1]
Lasmigona costata (1) (1) 1 (1) 1
Leptodea fragilis 1 1 (1)
Ligumia rectaB (1)
Obliquaria reflexa (1) 2
Obovaria subrotunda 1
Pleurobema cordatum (1)
Pleurobema oviforme (1) (13)
Pleuronaia barnesiana 6 2 1(2) (20) (4) (1) 1
Pleuronaia dolabelloidesA 6 1 2(3) (58) 1(3) (6) 1
Potamilus alatus 1 2(3) 1(1) (1)
Pyganodon grandis (1)
Quadrula quadrula 1(1)
Theliderma cylindricaA 2 6 (1)
Theliderma metanevraB [1]
Toxolasma lividum 2 1(1)
Tritogonia verrucosa 4 1(1)
Truncilla truncata 1(4) (10)
Villosa iris (1) (1)
Villosa taeniata 1(1)
Villosa vanuxemensis (1) (1)
Effort (person hours) 1.5 1 1 1 1.5 1.5 1 1 1 1 1
Total live and fresh dead 27 1 3 3 34 7 5 6 7 5 0
Total relic 1 1 2 1 21 284 17 16 6 5 1
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Site 1 2 3 4 5 6 7 8 9 10 11
BRM 0.2 1.9 2.2 2.8 3.2 3.5 4.0 4.6 4.9 6.8 8.0
CPUE 18 1 3 3 20 4 5 6 7 5 0
Species richness 9 2 3 3 23 18 7 10 9 7 1
Site 12 13 14 15 16 17 18 19 20 21 22
BRM 12.0 13.8 16.3 17.7 19.4 22.4 25.8 32.0 34.3 41.4 42.8
Species
Actinonaias pectorosa
Amblema plicata (1)
Arcidens confragosusB
Cyclonaias pustulosa
Cyclonaias tuberculata (1) (1)
Ellipsaria lineolataB
Elliptio crassidens
Eurynia dilatata (1) 1
Fusconaia subrotundaB
Lampsilis cardium
Lampsilis fasciola 2(2) 4(1) 2 2 1
Lampsilis ovata 1
Lampsilis teres
Lasmigona costata 1(1) 1 1
Leptodea fragilis 1(1) 1 1
Ligumia rectaB
Obliquaria reflexa
Obovaria subrotunda
Pleurobema cordatum
Pleurobema oviforme
Pleuronaia barnesiana (2) 1
Pleuronaia dolabelloidesA (1) (2) (3) 1 1
Potamilus alatus 1 2
Pyganodon grandis
Quadrula quadrula
Theliderma cylindrica 1
Theliderma metanevraB
Toxolasma lividum
Tritogonia verrucosa 1 1
Truncilla truncata
Villosa iris
Villosa taeniata 1 1 2 1 2(2)
Villosa vanuxemensis
Effort (person hours) 1 1 1 1 1 1 1 1 1 1 1
Total live and fresh dead 1 2 5 9 7 6 3 0 0 0 2
Total relic 1 6 5 1 2 2 0 0 0 0 2
CPUE 1 2 5 9 7 6 3 0 0 0 2
Species richness 2 3 3 9 6 7 3 0 0 0 1
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Site 23 24 25 26 27 28 29 30 31 32 33
BRM 45.6 52.1 55.1 55.6 56.0 57.4 58.0 59.0 62.7 67.4 70.6
Species
Actinonaias pectorosa
Amblema plicata
Arcidens confragosusB
Cyclonaias pustulosa
Cyclonaias tuberculata (1) (1)
Ellipsaria lineolataB
Elliptio crassidens (1)
Eurynia dilatata
Fusconaia subrotundaB
Lampsilis cardium
Lampsilis fasciola 1 (1) 1(2)
Lampsilis ovata
Lampsilis teres
Lasmigona costata
Leptodea fragilis
Ligumia rectaB
Obliquaria reflexa
Obovaria subrotunda
Pleurobema cordatum
Pleurobema oviforme
Pleuronaia barnesiana (1) 2
Pleuronaia dolabelloidesA
Potamilus alatus 1
Pyganodon grandis
Quadrula quadrula
Theliderma cylindricaA
Theliderma metanevraB
Toxolasma lividum
Tritogonia verrucosa
Truncilla truncata
Villosa iris (1)
Villosa taeniata (1) 1 1 3(7) (4) (2) 1 (1) 2
Villosa vanuxemensis 1
Effort (person hours) 1 1 1 1 1 1 1 1 1 1 1
Total live and fresh dead 0 0 2 1 3 1 0 2 3 0 2
Total relic 0 1 0 3 8 7 2 0 1 1 0
CPUE 0 0 2 1 3 1 0 2 3 0 2
Species richness 0 1 2 4 2 3 1 2 3 1 1
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Site 34 35 36 37 38 39 40 41 42 43
BRM 73.5 78.7 80.5 87.0 90.6 93.6 98.2 100.6 102.9 105.3
Species
Actinonaias pectorosa
Amblema plicata
Arcidens confragosusB
Cyclonaias pustulosa
Cyclonaias tuberculata
Ellipsaria lineolataB
Elliptio crassidens (1)
Eurynia dilatata
Fusconaia subrotundaB
Lampsilis cardium
Lampsilis fasciola 1 1
Lampsilis ovata
Lampsilis teres
Lasmigona costata
Leptodea fragilis
Ligumia rectaB
Obliquaria reflexa
Obovaria subrotunda
Pleurobema cordatum
Pleurobema oviforme
Pleuronaia barnesiana (9) 3(5) 5(3)
Pleuronaia dolabelloides1 2 5
Potamilus alatus
Pyganodon grandis
Quadrula quadrula
Theliderma cylindricaA
Theliderma metanevraB
Toxolasma lividum
Tritogonia verrucosa
Truncilla truncata
Villosa iris
Villosa taeniata (3) 3 2 (8) (4) 7(42) 12(13) (5)
Villosa vanuxemensis 2 1
Effort (person hours) 1 1 1 1 1 1 1 1 1 1
Total live and fresh dead 0 3 0 2 0 1 0 14 24 0
Total relic 3 1 0 0 0 17 4 47 16 5
CPUE 0 3 0 2 0 1 0 14 24 0
Species richness 1 2 0 1 0 3 1 4 5 1
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Site 44 45 46 47 Site Distrib. %
BRM 111.3 116.7 121.0 122.0 Totals freq. (BRM) coll.
Species
Actinonaias pectorosa (1) 1 3.2 0.2
Amblema plicata 4(3) 5 0.2–22.4 1.1
Arcidens confragosusB 1 1 3.2 0.2
Cyclonaias pustulosa 5(41) 3 3.2–4.9 7.9
Cyclonaias tuberculata 9(25) 10 0.2–57.4 5.9
Ellipsaria lineolataB 3(1) 2 0.2–3.2 0.7
Elliptio crassidens 2(2) 3 0.2–78.7 0.7
Eurynia dilatata 8(27) 10 0.2–22.4 6.0
Fusconaia subrotundaB (2) 1 4.6 0.2
Lampsilis cardium 1(1) 1 0.2 0.4
Lampsilis fasciola 25(17) 19 0.2–102.9 7.3
Lampsilis ovata 3(2) 4 3.2–17.7 0.9
Lampsilis teres 1 1 3.2 0.2
Lasmigona costata 4(4) 8 1.9–25.8 1.6
Leptodea fragilis 5(2) 6 3.5–19.4 1.1
Ligumia rectaB (1) 1 4.6 0.2
Obliquaria reflexa 2(1) 2 3.2–3.5 0.5
Obovaria subrotunda 1 1 3.2 0.2
Pleurobema cordatum (1) 1 3.5 0.2
Pleurobema oviforme (14) 2 3.2–3.5 2.4
Pleuronaia barnesiana 21(47) 14 0.2–102.9 11.2
Pleuronaia dolabelloidesA 20(76) 14 0.3–102.9 16.6
Potamilus alatus 8(5) 7 1.9–59.0 2.2
Pyganodon grandis (1) 1 3.2 0.2
Quadrula quadrula 1(1) 1 3.5 0.4
Theliderma cylindricaA 9(1) 4 2.2–17.7 1.7
Theliderma metanevraB 1 1 3.2 0.2
Toxolasma lividum 3(1) 2 3.2–3.5 0.7
Tritogonia verrucosa 7(1) 4 0.2–22.4 1.4
Truncilla truncata 1(14) 2 3.2–3.5 2.6
Villosa iris (3) 3 4.6–62.7 0.7
Villosa taeniata 40(93) 23 4.9–102.9 22.9
Villosa vanuxemensis 4(2) 5 3.2–102.9 1.0
Effort (person hours) 1 1 1 1
Total live and fresh dead 0 0 0 0 189
Total relic 0 0 0 0 390
CPUE 0 0 0 0
Species richness 0 0 0 0 33
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Appendix 4. Conservation status of mussels of the Buffalo River, TN. Federal status according
to USFWS (2016). Species with Unknown status were either undetected or represented
by relic shells only. E = Endangered, T = Threatened, X = Extinct.
Status
Buffalo
Scientific name Common name USFWS River
Margaritiferidae
Margaritifera monodonta (Say) Spectaclecase E Extant
Unionidae
Actinonaias ligamentina (Lamarck) Mucket Extant
Actinonaias pectorosa (Conrad) Pheasantshell Extant
Alasmidonta marginata Say Elktoe Unknown
Alasmidonta viridis (Rafinesque) Slippershell Unknown
Amblema plicata (Say) Threeridge Extant
Arcidens confragosus (Say) Rock Pocketbook Extant
Cyclonaias pustulosa (Lea) Pimpleback Extant
Cyclonaias tuberculata (Rafinesque) Purple Wartyback Extant
Ellipsaria lineolata (Rafinesque) Butterfly Extant
Elliptio crassidens (Lamarck) Elephantear Extant
Epioblasma ahlstedti Jones and Neves Duck River Dartersnapper E Unknown
Epioblasma aureola Jones and Neves Golden Riffleshell E Unknown
Epioblasma turgidula (Lea) Turgid Blossom E,X Unknown
Eurynia dilatata Rafinesque Spike Extant
Fusconaia subrotunda (Lea) Longsolid Unknown
Hemistena lata (Rafinesque) Cracking Pearlymussel E Unknown
Lampsilis cardium Rafinesque Plain Pocketbook Extant
Lampsilis fasciola Rafinesque Wavyrayed Lampmussel Extant
Lampsilis ovata (Say) Pocketbook Extant
Lampsilis teres (Rafinesque) Yellow Sandshell Extant
Lasmigona complanata (Barnes) White Heelsplitter Unknown
Lasmigona costata (Rafinesque) Flutedshell Extant
Leptodea fragilis (Rafinesque) Fragile Papershell Extant
Ligumia recta (Lamarck) Black Sandshell Unknown
Ligumia subrostrata (Say) Pondmussel Unknown
Megalonaias nervosa (Rafinesque) Washboard Extant
Obliquaria reflexa Rafinesque Threehorn Wartyback Extant
Obovaria subrotunda (Rafinesque) Round Hickorynut Extant
Pleurobema cordatum (Rafinesque) Ohio Pigtoe Unknown
Pleurobema oviforme (Conrad) Tennessee Clubshell Unknown
Pleurobema sintoxia (Rafinesque) Round Pigtoe Unknown
Pleuronaia barnesiana (Lea) Tennessee Pigtoe Extant
Pleuronaia dolabelloides (Lea) Slabside Pearlymussel E Extant
Potamilus alatus (Say) Pink Heelsplitter Extant
Ptychobranchus fasciolaris (Rafinesque) Kidneyshell Unknown
Ptychobranchus subtentus (Say) Fluted Kidneyshell E Unknown
Pyganodon grandis (Say) Giant Floater Unknown
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Status
Buffalo
Scientific name Common name USFWS River
Quadrula quadrula (Rafinesque) Mapleleaf Extant
Strophitus undulatus (Say) Creeper Unknown
Theliderma cylindrica (Say) Rabbitsfoot T Extant
Theliderma metanevra (Rafinesque) Monkeyface Extant
Toxolasma cylindrellus (Lea) Pale Lilliput E Unknown
Toxolasma lividum Rafinesque Purple Lilliput Extant
Tritogonia verrucosa (Rafinesque) Pistolgrip Extant
Truncilla truncata Rafinesque Deertoe Extant
Utterbackia imbecillis (Say) Paper Pondshell Unknown
Utterbackiana suborbiculata (Say) Flat Floater Unknown
Villosa iris (Lea) Rainbow Unknown
Villosa taeniata (Conrad) Painted Creekshell Extant
Villosa vanuxemensis (Lea) Mountain Creekshell Extant
Total (51 species)