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Transactions of the Illinois State Academy of Science received 10/26/07
(2008), Volume 101, #1&2, pp. 113-124 accepted 4/30/08
Freshwater Mussel (Bivalvia: Unionidae)
Survey of the Galena River Basin,
Wisconsin and Illinois
Jeremy S. Tiemann
Illinois Natural History Survey
Division of Biodiversity and Ecological Entomology
1816 South Oak Street
Champaign, Illinois 61820
Correspondence: jtiemann@inhs.uiuc.edu
ABSTRACT
I investigated the freshwater mussel assemblage of the Galena River basin in Wisconsin
and Illinois in 2005-2006 to determine distribution and structure of the fauna. I collected
27 live individuals representing 5 species during 47 person-hours of sampling at 28 sites;
I also found valves of an additional 20 species, including 5 species listed at the state level
in Wisconsin and/or Illinois. Freshwater mussels were evident at 18, or 64%, of the sites.
Regression analysis indicated an increase in extant species richness (r2 = 0.21, F0.05(2), 26 =
6.97, P = 0.02) and historic species richness (r2 = 0.58, F0.05(2), 26 = 40.18, P < 0.0001)
from upstream to downstream. Expanding drainage areas usually offer decreased gradi-
ents, more habitat complexities, and higher host fish diversity. A t-test concluded extant
and historic species richness differed significantly (t0.05(1), 17 = 3.56, P = 0.003) suggesting
that the assemblage has been reduced.
Key Words: bivalve, Mollusca, unionids, Wisconsin Driftless Division
INTRODUCTION
The Wisconsin Driftless Division (herein after Driftless area), a region of nearly 35,000
km2, was surrounded but never covered by late Pleistocene glacial ice (Page et al. 1992;
Knox 2001; Knox 2006). Today, the area has rolling topography characterized by steep-
sided (up to150 m/km vertical relief) limestone/sandstone valleys, forested ridges, and
streams that have spring-fed headwaters. Situated between the tallgrass prairie to the west
and the deciduous forest to the east, the Driftless area historically was dominated by tall-
grass prairie, oak savanna, southern oak forest, and southern mesic forest. Today, the
region has agricultural fields (e.g., row crops or grazing pastures) that have riparian areas
composed of either grassy or woody buffer strips. About 40 species of freshwater mussels
(Bivalvia: Unionidae) historically inhabited the Driftless area (unpublished data from the
Illinois Natural History Survey [INHS] Mollusk Collection, Champaign). It appears that
5 species (13%) are extirpated from the area, and include fluted shell Lasmigona costata,
114
elephantear Elliptio crassidens, ebonyshell Fusconaia ebena, sheepnose Plethobasus
cyphyus, and mucket Actinonaias ligamentina.
The Galena River, one of the streams in the Driftless area, drains nearly 525 km2. The
stream originates in Grant and Lafayette counties, Wisconsin, and flows south-southwest
through Jo Daviess County, Illinois, until reaching the Mississippi River (Figure 1). The
Galena River basin contains a unique fish assemblage, including Ozark minnow Notropis
nubilus and longnose dace Rhinichthys cataractae, which, in Wisconsin and Illinois, are
found only in a few basins (INHS Fish Collection, Champaign, data). Monitoring aquatic
assemblages is vital for natural resource agencies to accurately assess their statuses (e.g.,
rare species) and provide baseline data to evaluate the effects of human activities. How-
ever, not all aquatic assemblages have been adequately sampled. One such example is the
freshwater mussel fauna. I sampled freshwater mussels in the Galena River basin to
determine distribution and structure of the assemblage. Prior to this study, no comprehen-
sive survey on the freshwater mussel fauna of the Galena River basin had been con-
ducted. Data collected will allow future comparisons for monitoring the assemblage and
provide information on which to base management goals for the basin.
METHODS
I collected freshwater mussels at 28 sites in the Galena River basin (Figure 1; Appendix
1) during August 2005 and August 2006. Four sites were selected because historical data
existed for them (INHS Mollusk Collection data) and the remaining 24 sites were
selected to provide coverage of streams throughout the entire watershed. Of the four his-
torical locations, Site 19 was surveyed in 1993 and 1994 (unknown amount of effort) and
Site 22 was sampled for 30 minutes in 2002, whereas incidental encounters were discov-
ered at Site 18 in 2000 and Site 21 in 1941. I collected live freshwater mussels and valves
of dead specimens by hand-grabbing for 1-2 person-hours at each site depending upon
stream condition and amount of success (Tiemann 2006). For example, collecting ceased
at sites where substrates were predominantly clay/silt and no specimens were found
within the first hour of sampling. Hand-grabbing is a cost-effective, semi-quantitative
method effective for obtaining information on relative abundance and species richness
but can be biased to an unknown extent (Strayer et al. 1997; Vaughn et al. 1997). Sam-
pling occurred while wading in all available habitats but primarily was concentrated in
areas that appeared likely to support freshwater mussels, such as non-silted areas (Tie-
mann 2006). No effort was made to sample ponds/lakes or wetlands in the basin. Below
average water levels during summer 2005 allowed sampling in the channelized, lower
portions of the basin. I obtained Global Positioning System coordinates at each site using
a Garmin GPS 12 XL (Garmin International, Romsey, Hampshire, United Kingdom). I
classified shell material as live, fresh dead (shiny nacre), or relict (chalky nacre) based on
condition of the best specimen found (Sietman et al. 2004). Identification and taxonomy
followed Cummings and Mayer (1992). All live individuals were counted and returned
upon completion of a site.
Extant species richness, historical species richness, and relative abundance were calcu-
lated for each site (Tiemann et al. 2005). I figured extant species richness as the number
of species represented by live or fresh dead shell, historical species richness as the total
number of species found including museum records located at the INHS Mollusk Collec-
115
tion, and relative abundance as catch-per-unit-effort (CPUE). I used simple regression
analysis to test if extant species richness, historical species richness, and CPUE increased
from upstream to downstream in the basin. Drainage area was determined by digitizing
topographic maps. At those sites where specimens were found, I applied a t-test to deter-
mine if extant species richness was significantly lower than historic species richness.
Statistical analyses was preformed with SAS Version 8 (SAS Institute Inc., Cary, North
Carolina) and considered significant at P 0.05.
RESULTS AND DISCUSSION
I collected a total of 27 live individuals representing 5 species in 47 person-hours in the
Galena River basin; I also found valves of an additional 20 species, including 5 species
listed at the state level in Wisconsin and/or Illinois (Table 1, Appendix 2). Freshwater
mussels were evident at 18, or 64%, of the 28 sites (Appendix 1). CPUE in the Galena
River basin ranged from 0-3 individuals per hour per site (Appendix 2) and did not
increase significantly (r2 = 0.06, F0.05(2), 26 = 1.53, P = 0.23) from upstream to downstream
(Figure 2a). Giant floater Pyganodon grandis was the most abundant and widely distrib-
uted species (12 individuals from 6 sites) followed by white heelsplitter Lasmigona com-
planata (5 individuals from 5 sites), plain pocketbook Lampsilis cardium (5 individuals
from 3 sites), creek heelsplitter Lasmigona compressa (3 individuals from 2 sites), and
lilliput Toxolasma parvus (2 individuals from 2 sites) (Appendix 2). Excluding L. com-
pressa, the species found live are widespread and common throughout streams in the
Midwest; L. compressa occasionally is found in small streams in the region (Cummings
and Mayer 1992). Most of the other specimens found were relict valves (Table 1, Appen-
dix 2). Based on historical records, some species (e.g., slippershell mussel Alasmidonta
viridis, T. parvus, and ellipse Venustaconcha ellipsiformis) were found throughout the
basin, whereas other species (e.g., pistolgrip Tritogonia verrucosa, pink heelsplitter
Potamilus alatus, and threehorn wartyback Obliquaria reflexa) probably were not widely
distributed (e.g., found only in the lower portions). The extant species in the basin were
found only in small, isolated populations, which could hinder reproduction and recoloni-
zation efforts.
No live threatened or endangered species were collected; however, 5 species listed at the
state level in Wisconsin and/or Illinois were found only as valves. Alasmidonta viridis
(Wisconsin state threatened and Illinois state threatened), E. dilatata (Illinois state threat-
ened), and V. ellipsiformis (Wisconsin state threatened) were found throughout the basin
(Appendix 2). These 3 species were once widely distributed in the Midwest but are now
are sporadic in their distributions (Cummings and Mayer 1992). The other 2 species,
wartyback Quadrula nodulata and T. verrucosa, both listed as state threatened in Wis-
consin, were found only in the downstream areas of the mainstem (Appendix 2). Both of
these species are found in larger rivers or in the downstream sections of medium-sized
streams (Cummings and Mayer 1992).
The number of extant species in the Galena River basin ranged from 0-3 species per site,
whereas the number of historic species varied from 0-11 species (Appendix 1; Appendix
2). The differences in species richness between historic and extant ranged from 0-11 spe-
cies. Extant species richness and historic species richness differed significantly (t0.05(1), 17
= 3.56, P = 0.003) suggesting that the freshwater mussel assemblage has declined. There
116
was a linear increase in extant species richness (r2 = 0.21, F0.05(2), 26 = 6.97, P = 0.02) and
historic species richness (r2 = 0.58, F0.05(2), 26 = 40.18, P < 0.0001) from upstream to
downstream (Figure 2b). This positive relationship between species richness and drainage
area has been shown for freshwater mussels (Watters 1992) and fishes (Edds 1993).
Expanding drainage areas offer decreased gradients, more habitat complexities, and
higher fish diversity to serve as glochidia hosts (Vannote et al. 1980; Watters 1992). No
evidence of freshwater mussels was found in the middle portions of the basin and likely
is the result of sub-optimal habitat (e.g., silt-laden cobble).
The temporal decline in species richness in the Galena River basin (80%) is substantially
greater than other basins in the Driftless area region. The Apple River basin (Wisconsin
and Illinois) has a 16% (26 extant species out of 31 historic species) reduction in historic
species richness (Anderson and Sietman 2004), whereas the upper Iowa and Turkey river
basins (Iowa) together have a 23% (10 extant species out of 13 historic species) reduction
(Eckblad et al. 2002), and the Menominee, Little Menominee, and Sinsinawa river basins
(Wisconsin and Illinois) have a 22% (7 extant species out of 9 historic species), 36% (9
extant species out of 14 historic species), and 33% (8 extant species out of 12 historic
species) reduction, respectively (Jeremy Tiemann, Illinois Natural History Survey,
unpubl. data). The Lost Mound Unit of the Upper Mississippi River National Wildlife
and Fish Refuge, an area of the Mississippi River that lies on the southern edge of the
Driftless area, has a 30% (26 extant species out of 37 historic species) reduction in his-
toric species richness (Sietman et al. 2004). In the Apple, Menominee, Little Menominee,
and Sinsinawa river basins, the majority (> 80%) of live individuals and species richness
were found within the lower quarter of their respective basins (Anderson and Sietman
2004; Jeremy Tiemann, Illinois Natural History Survey, unpubl. data), whereas in the
upper Iowa and Turkey river basins, the majority (> 85%) of live individuals and species
richness were found in the headwaters (Eckblad et al. 2002). These distribution patterns
were attributed to the complexity and amount of available habitat at a given site. A simi-
lar pattern of downstream distribution was seen in the Galena River for historic species
richness but not extant species richness or CPUE, perhaps because the lower portion of
the river has been dredged and now offers unsuitable habitat. The amount of effort per
site (1-2 person-hours) in the Galena River basin was the same as in the Menominee, Lit-
tle Menominee, and Sinsinawa river basins (Jeremy Tiemann, Illinois Natural History
Survey, unpubl. data), but was greater than the 25 minutes per site in the upper Iowa and
Turkey river basins (Eckblad et al. 2002); the amount of effort in the Apple River basin
varied from 20-540 minutes per site (Anderson and Sietman 2004).
Habitat appears to be the limiting factor for freshwater mussels in the Driftless area.
Mathiak (1979) suggested that the Driftless area lacks freshwater mussels because of
poor habitat. Since the early 19th century, post-settlement modifications of the natural
land cover have altered all aspects of the hydrologic landscape in the Galena River basin,
including floods, erosion, transportation and deposition of sediments, and the morphol-
ogy of stream channels and the associated floodplains (Magilligan 1985; Knox 2001;
Knox 2006). Very little sand, gravel, or pebble existed in the Galena River basin. The
majority of the habitat in these areas was silt-laden cobble / boulder with patches of bare
bedrock or clay hardpan. Even though freshwater mussels can colonize bedrock, it is sub-
optimal habitat (Sietman et al. 1999). Silt, the number one pollutant of streams in the
Driftless area (Page et al. 1992), has been shown to decrease species richness of macroin-
117
vertebrates (Weigel 2003) and fishes (Wang et al. 1997). Several anthropogenic distur-
bances that cause siltation, including dredging, mining, unrestricted livestock access in
streams, and cutting of riparian areas, have occurred in the Galena River basin (Magilli-
gan 1985; Knox 2001; Knox 2006). These types of disturbances, along with organic pol-
lution (e.g., effluents from sewage treatment plants), have been shown to alter stream
habitat and change freshwater mussel assemblages (e.g., Aldridge 2000; Hoke 1997).
Unless mitigated, these disturbances will continue to threaten the existing assemblage and
might prevent the expansion / recolonization of future species.
I recognize the limitations of this study (e.g., limited number of sites and limited sam-
pling effort). Additional surveys could be done to further explore the freshwater mussel
assemblage in the Galena River basin by adding more sites (e.g., pools) and incorporating
other methods (e.g., quadrat searches). Nevertheless, the reduction in freshwater mussel
species richness in the Galena River basin is a cause of concern. The apparent 80% loss
of a taxonomic group in a basin could result in the loss of valuable genetic diversity.
Imlay (1973) suggested that the Driftless area be protected as a possible “seed area” for
the redistribution of species. An active, positive partnership between natural resource
agencies and landowners to promote sound management practices should aid in the pres-
ervation and restoration of this unique area.
ACKNOWLEDGMENTS
R. Szafoni of the Illinois Department of Natural Resources (IL-DNR) provided the idea
for this project. Funding was provided by the IL-DNR Illinois Wildlife Preservation Fund
(grant #06-34W) and the Illinois Department of Transportation. The Wisconsin Depart-
ment of Natural Resources issued a Wisconsin collecting permit. F. Hutto (INHS)
assisted in map creation. E. Anderson (IL-DNR) and B. Sietman (Minnesota Department
of Natural Resources) shared their Apple River data. K. Cummings (INHS), G. Levin
(INHS), R. Szafoni (IL-DNR), D. Thomas (INHS), B Tiemann, and C. Warrick (INHS)
offered constructive comments.
LITERATURE CITED
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sels (Bivalvia: Unionidae). Biological Conservation 95:247-257.
Anderson, E.A., and B.E. Sietman. 2004. Unionid mussel fauna of the Apple River above Hanover,
Illinois. Technical Report to the Illinois Department of Natural Resources, Springfield, Illinois.
23 pp + appendix.
Cummings, K.S., and C.A. Mayer. 1992. Field guide to freshwater mussels of the Midwest. Manual
5. Illinois Natural History Survey, Champaign, Illinois. 194 pp.
Eckblad, J., B. Ostby, and K. Tenneson. 2002. The unionid mussels of the upper Iowa and Turkey
river watersheds. Journal of the Iowa Academy of Science 109:43-49.
Edds, D.R. 1993. Fish assemblage and environmental correlates in Nepal’s Gandaki River. Copeia
1993:48-60.
Hoke, E. 1997. The unionid mollusks of the upper Kansas basin of northwestern Kansas and
southwestern Nebraska. Transactions of the Nebraska Academy of Science 24:35-62.
Imlay, M.J. 1973. A case for protecting the Driftless area of the upper Midwest. Malacological
Review 6:64-65.
Knox, J.C. 2001. Agricultural influence on landscape sensitivity in the Upper Mississippi River
Valley. Catena 42:193-224.
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Knox, J.C. 2006. Floodplain sedimentation in the Upper Mississippi Valley: Natural versus human
accelerated. Geomorphology 79:286-310.
Mathiak, H.A. 1979. A river survey of the unionid mussels of Wisconsin 1973-1977. Sand Shell
Press. Horicon, Wisconsin. 75 pp.
Magilligan, F.J. 1985. Historical floodplain sedimentation in the Galena River basin, Wisconsin
and Illinois. Annals of the Association of American Geographers 75:583–594.
Page, L. M., K. S. Cummings, C. A. Mayer, S. L. Post, and M. E. Retzer. 1992. Biologically sig-
nificant Illinois streams - an evaluation of the streams of Illinois based on aquatic biodiversity.
Center for Biodiversity Technical Report 1992(1), Illinois Natural History Survey, Champaign,
Illinois. vi + 479 pp + appendices.
Sietman, B.E., M.A. Furman, and F.A. Pursell. 1999. Colonization of bedrock by freshwater mus-
sels (Bivalvia: Unionidae). American Midland Naturalist 141:209-211.
Sietman, B.E., E.A. Anderson, R. Nÿboer, and F.R. Hutto. 2004. Native freshwater mussels
(Bivaliva: Unionidae) and infestation by zebra mussels at the Lost Mound Unit of the Upper
Mississippi River National Wildlife and Fish Refuge. Transactions of the Illinois State Academy
of Science 97:234-254.
Strayer, D.L., S. Claypool, and S.J. Sprague. 1997. Assessing unionid population with quadrats and
timed searches. p. 163-169. In K.S. Cummings, A.C. Buchanan, C.A. Mayer, and T.J. Naimo
(eds.). Conservation and Management of Freshwater Mussels II: Initiatives for the Future. Pro-
ceedings of a Upper Mississippi River Conservation Committee Symposium, October 1995, St.
Louis, Missouri. Upper Mississippi River Conservation Committee, Rock Island, Illinois. 293 p.
Tiemann, J.S., R.E. Szafoni, and K. Roman. Freshwater mussel (Bivalvia: Unionidae) survey of the
Kyte River, Ogle County, Illinois. Transactions of the Illinois State Academy of Science 98:159-
169.
Tiemann, J.S. 2006. Freshwater mussel (Bivalvia: Unionidae) survey of the Wakarusa River basin,
Kansas. Transactions of the Kansas Academy of Science 109:221-230.
Vannote, R.L., G.W. Minshall, K.W. Cummins, J.R. Sedell, and C.E. Cushing. 1980. The river
continuum concept. Canadian Journal of Fisheries and Aquatic Sciences 37:130-137.
Vaughn, C.C., C.M. Taylor, and K.J. Eberhard. 1997. A comparison of the effectiveness of timed
searches vs. quadrat sampling in mussel surveys. p. 157-162. In K.S. Cummings, A.C.
Buchanan, C.A. Mayer, and T.J. Naimo (eds.). Conservation and Management of Freshwater
Mussels II: Initiatives for the Future. Proceedings of a Upper Mississippi River Conservation
Committee Symposium, October 1995, St. Louis, Missouri. Upper Mississippi River Conserva-
tion Committee, Rock Island, Illinois. 293 p.
Wang, L. J. Lyons, P. Kanehl, and R. Gatti. 1997. Influences of watershed land use on habitat
quality and biotic integrity in Wisconsin streams. Fisheries 22(6):6-12.
Watters, G.T. 1992. Unionids, fishes, and the species-area curve. Journal of Biogeography 19:481-
490.
Weigel, B.M. 2003. Development of stream macroinvertebrate models that predict watershed and
local stressors in Wisconsin. Journal of North American Benthological Society 22:123-142.
119
Table 1. Freshwater mussels collected during the 2005-2006 survey of the Galena River
basin, Wisconsin and Illinois. See appendices for site-specific data. Numbers
within a given species row represent the number of individuals that species was
collected alive, D indicates those species collected only as fresh dead specimens,
R signifies those species collected only as relict specimens, and * indicates those
species not found during survey but an INHS Mollusk Collection record exits
for the basin. WI-ST = Wisconsin state-threatened and IL-ST = Illinois state-threat-
ened.
Sub-family
Scientific name
Common name
Status
Anodontinae
Alasmidonta viridis WI-ST, IL-ST
Slippershell mussel
R
Anodontoides ferussacianus
Cylindrical papershell
D
Lasmigona complanata
White heelsplitter
5
Lasmigona compressa
Creek heelsplitter
3
Lasmigona costata
Flutedshell
R
Pyganodon grandis
Giant floater
12
Strophitus undulatus
Creeper
R
Utterbackia imbecillis
Paper pondshell
D
Ambleminae
Amblema plicata
Threeridge
R
Elliptio dilatata IL-ST
Spike
R
Fusconaia flava
Wabash pigtoe
R
Quadrula nodulata WI-ST
Wartyback
R
Quadrula pustulosa
Pimpleback
R
Quadrula quadrula
Mapleleaf
R
Tritogonia verrucosa WI-ST
Pistolgrip
R
Lampsilinae
Lampsilis cardium
Plain pocketbook
5
Lampsilis siliquoidea
Fatmucket
*
Leptodea fragilis
Fragile papershell
D
Obliquaria reflexa
Threehorn wartyback
D
Potamilus alatus
Pink heelsplitter
R
Potamilus ohiensis
Pink papershell
R
Toxolasma parvus
Lilliput
2
Truncilla donaciformis
Fawnsfoot
R
Truncilla truncata
Deertoe
R
Venustaconcha ellipsiformis WI-ST
Ellipse
R
120
Figure 1. Map of the Galena River basin and sampling sites, Wisconsin and Illinois.
121
Figure 2. (a) Freshwater mussel CPUE (r2 = 0.06, F0.05(2), 26 = 1.53, P = 0.23) and (b)
extant species richness (r2 = 0.21, F0.0 5(2), 26 = 6.97, P = 0.02) (left y-axis; cir-
cles with solid line) and historic species richness (r2 = 0. 58, F0.0 5(2), 26 = 40.18,
P < 0.0001) (right y-axis; triangles with dashed line) compared drainage area in
the Galena River basin, Wisconsin and Illinois.
(a)
(b)
122
Appendix 1
Collecting locations for the 2005-2006 freshwater mussel survey of the Galena River
basin, Wisconsin and Illinois. FWM is freshwater mussel material collected at that site. L
is number of species collected alive and V is number of species collected only as valves.
Site
State: County
Stream
Latitude, Longitude
FWM
01
WI: Lafayette
Galena River
42.6673, -90.4134
02
Galena River
42.6405, -90.3969
03
Galena River
42.6308, -90.3621
L(2)
04
Galena River
40.6092, -90.3602
05
Galena River
42.5714, -90.3639
L(2), V(3)
06
Galena River
42.5529, -90.3537
L(2), V(5)
07
Galena River
42.5428, -90.3578
L(3), V(3)
08
Galena River
42.5161, -90.3931
09
Pats Creek
42.6528, -90.3836
L(1)
10
Madden Branch
42.6311, -90.3552
11
Shullburg Branch
42.5659, -90.3277
V(1)
12
Ellis Branch
42.5473, -90.3395
V(1)
13
Kelsey Branch
42.5108, -90.3575
14
Coon Branch
42.5136, -90.3781
V(1)
15
Bull Branch
42.5146, -90.3965
L(1)
16
Scrabble Branch
42.5136, -90.3978
17
IL: Jo Daviess
Galena River
42.4947, -90.3947
18
Galena River
42.4768, -90.4066
V(3)
19
Galena River
42.4511, -90.3879
L(3), V(8)
20
Galena River
42.4285, -90.4017
L(3), V(3)
21
Galena River
42.4163, -90.4237
V(6)
22
Galena River
42.4012, -90.4366
V(6)
23
Galena River
42.3757, -90.4455
V(11)
24
East Fork Galena
River
42.4901, -90.2991
25
East Fork Galena
River
42.4748, -90.3187
L(1)
26
East Fork Galena
River
42.4665, -90.3483
V(4)
27
East Fork Galena
River
42.4536, -90.3779
V(2)
28
Hughlett Branch
42.4367, -90.4237
123
Appendix 2
Site-specific data for only those locations where freshwater mussels were collected during the 2005-2006 freshwater mussel survey of the
Galena River basin, Wisconsin and Illinois. Site No. is the site number, which is referenced in Appendix 1. Numbers within a given species
row represent the number of individuals that species was collected alive at that site, D indicates those species collected only as fresh dead
specimens, R signifies those species collected only as relict specimens, and * indicates those species represented by INHS Mollusk Collection
records. Abundance is the total number of live unionids, extant species richness is the number of species represented by live or fresh dead shell
material, and historical species richness is the total number of species found (including museum records). Effort is in person-hours. Note: sites
01, 02, 04, 08, 17, and 24 were sampled for two person-hours each and sites 10, 13, 16, and 28 were sampled for one person-hour each, but no
evidence of freshwater mussels was found.
Site No
Species
03
05
06
07
09
11
12
14
15
18
19
20
21
22
23
25
26
27
Anodontinae
Alasmidonta viridis
R
R
*
*
R
Anodontoides ferussacianus
R
R
Lasmigona complanata
1
1
1
R
1
1
R
Lasmigona compressa
1
2
Lasmigona costata
R
R
Pyganodon grandis
2
2
3
R
2
2
1
Strophitus undulatus
R
*
Utterbackia imbecillis
D
Ambleminae
Amblema plicata
R
R
R
R
R
Elliptio dilatata
R
R
R
*
R
Fusconaia flava
R
*
R
R
Quadrula nodulata
R
Quadrula pustulosa
R
124
Site No
Species
03
05
06
07
09
11
12
14
15
18
19
20
21
22
23
25
26
27
Quadrula quadrula
R
Tritogonia verrucosa
R
R
R
Lampsilinae
Lampsilis cardium
R
1
R
R
3
1
R
R
R
Lampsilis siliquoidea
*
Leptodea fragilis
D
Obliquaria reflexa
D
Potamilus alatus
R
Potamilus ohiensis
R
Toxolasma parvus
R
R
1
1
*
Truncilla donaciformis
R
Truncilla truncata
*
R
R
R
Venustaconcha ellipsiformis
R
R
R
R
R
Abundance
3
3
2
6
1
0
0
0
1
0
6
4
0
0
0
1
0
0
Extant species richness
2
2
2
3
1
0
0
0
1
0
3
3
0
1
2
1
0
0
Historical species richness
2
5
7
6
1
1
1
1
1
3
11
6
8
6
11
1
4
2
Effort
2
2
2
2
1
1
1
1
1
2
2
2
2
2
2
2
2
2
Catch-per-unit-effort
1.5
1.5
1
2
1
0
0
0
1
0
3
2
0
0
0
0.5
0
0
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We propose that the Concept provides a framework for integrating predictable and observable biological features of lotic systems. Implications of the concept in the areas of structure, function, and stability of riverine ecosystems are discussed. -from Authors