ABSTRACT. Seventy-five species of freshwater mussels (Order
Unionoida) have historically inhabited the Wabash River drainage of
Indiana. Nine of these species have always been restricted to Wabash
River tributaries and never maintained reproducing populations in the
mainstem Wabash River. Of the 66 remaining species, 18 are currently
considered extirpated from the entire drainage and 18 maintain
reproducing populations only in Wabash River tributaries. Currently, 30
species maintain reproducing populations in the mainstem Wabash River,
which represents a 55% reduction in its freshwater mussel fauna. To
date, the entire Wabash River drainage of Indiana has seen a 24%
reduction in its freshwater mussel fauna.
Keywords: Freshwater mussels, Wabash River
**********
The freshwater mussel (Order Unionoida) fauna of the Wabash River
drainage has been well documented historically. Stein (1881) attempted
the first complete list of the 'molluscous fauna of Indiana,'
and referenced many species as inhabiting the Wabash River and its
tributaries. Call (1894, 1896, 1897, 1900), Blatchley & Daniels
(1903), Daniels (1903, 1915), and Goodrich & van der Schalie (1944)
continued to add to the knowledge of Indiana's mollusca fauna and
provided invaluable information on those species found in the Wabash
River drainage.
Three important Wabash River studies were completed during the
1960s and 1970s. Meyer (1968) and Krumholz et al. (1970) studied the
commercially valuable species of the Wabash and White rivers. Clark
(1976) inventoried mussels from the lower Wabash River.
Between 1987 and 1991, Cummings et al. (1992) sampled 100 sites in
the Wabash River drainage, including 53 sites on the mainstem Wabash
River. Several of the lower Wabash River sites sampled by Cummings et
al. (1992) were re-sampled in 1996 by Frankland (1996). Ball &
Schoenung (1996) and EnviroScience (2006) intensively sampled freshwater
mussels at several locations in the upper mainstem Wabash River. Page et
al. (1992) and Cummings & Mayer (1997) provide information on the
status of freshwater mussels in the Wabash River drainage of Illinois.
Many of the larger tributaries of the Wabash River have also had
recent survey work completed (from upstream to downstream): Salamonie
River (Ecological Specialists, Inc. 1995), Mississinewa River
(Ecological Specialists, Inc. 1995), Eel River (upper Wabash River)
(Henschen 1987), Tippecanoe River (Cummings & Berlocher 1990;
Cummings et al. 1992; Ecological Specialists, Inc. 1993, 1998; Ball
& Schoenung 1996; Commonwealth Biomonitoring 2005; EnviroScience
2006), Middle Fork Wildcat (Henschen 1990), small streams of Tippecanoe
County (Myers-Kinzie et al. 2001), Jordan Creek (Szafoni et al. 2000),
Sugar Creek (middle Wabash River tributary) (Lewis 1991), Brouilletts
Creek (Tiemann 2005), East Fork White River drainage (Cummings et al.
1992; Ball & Schoenung 1996; Harmon 1998; Clarke et al. 1999;
EnviroScience 2006), West Fork White River drainage (Cummings et al.
1992; Henschen 1993, 1995; L. Bowley, Muncie Bureau of Water Quality
pers. comm.), and Patoka River (Ecological Specialists, Inc. 2001).
Figure 1 illustrates the recent collections cited here, which include
samples from 1987-2004.
[FIGURE 1 OMITTED]
In addition to this information, the Wildlife Diversity Section,
Division of Fish and Wildlife, Indiana Department of Natural Resources,
collected freshwater mussel information from nearly 900 sites within the
Wabash River drainage between 1995-2006 (Fig. 2). This information,
along with the previously cited surveys, was used to determine the
current status of freshwater mussel species within the Wabash River
drainage of Indiana.
[FIGURE 2 OMITTED]
METHODS
Several different collecting methods were used in the previously
cited surveys: the individual reports should be reviewed to determine
the specific methods used by those researchers.
An informal sampling design (Strayer & Smith 2003) was utilized
for most of our surveys within the Wabash River drainage. Sampling
locations were chosen to provide information from watersheds where few
or no previous freshwater mussel surveys had been completed. Locations
were waded and visually (if possible) and physically searched for live
freshwater mussels and dead shell material. At locations where
visibility was limited, the stream bottom was searched with hands or
shoed-feet.
Live freshwater mussels were identified onsite and returned;
representative dead shell material was retained from most locations.
Species lists indicating the best condition of shell material (live,
fresh dead, weathered dead, or subfossil) encountered were prepared for
all sampled locations. All shell material retained from our sampling
efforts is currently vouchered at the Atterbury Fish and Wildlife Area,
Edinburgh, Indiana.
Besides the specific freshwater mussel sampling described,
additional information was obtained incidentally while completing survey
work for fishes. Notes were made on live freshwater mussels and shell
material encountered, although no formal surveys were completed.
RESULTS AND DISCUSSION
For the purpose of this paper, freshwater mussel species are
considered extirpated if they no longer maintain a reproducing
population. It is possible remnant live individuals of some of the
species considered extirpated may still be found. However, because their
populations have reached levels that no longer support reproduction,
they are deemed functionally extirpated. Viable populations refer only
to those aggregations of freshwater mussels that are reproducing, as
evidenced by the presence of live, young individuals.
Of the 75 species of freshwater mussels historically known from the
Wabash River drainage of Indiana, 30 still maintain populations in both
the mainstem Wabash River and its tributaries, 18 are extirpated from
the mainstem, but maintain populations in some portion of the drainage,
and 18 are now extirpated from the entire drainage (Tables 1, 2). The
nine remaining species are small-stream species or species of unique
habitats and probably never had populations in the mainstem Wabash
River; they maintain populations in tributaries of the Wabash River
drainage within Indiana.
In describing the distribution of some of the freshwater mussel
species in the following discussion, the 'upper Wabash River'
is used to describe the area upstream from where Sugar Creek enters the
Wabash River in Parke and Vermillion counties. The 'lower Wabash
River' refers to the area downstream from Sugar Creek to its
confluence with the Ohio River.
Live species of mainstem Wabash River and its tributaries.--Thirty
species of freshwater mussels still maintain populations in both the
mainstem Wabash River and its tributaries (Table 1). The most widely
distributed of these species can be found throughout the entire
drainage, in a variety of stream sizes. These species include:
threeridge (Amblema plicata), Wabash pigtoe (Fusconaia flava), plain
pocketbook (Lampsilis cardium), fatmucket (Lampsilis siliquoidea), white
heelsplitter (Lasmigona complanata), giant floater (Pyganodon grandis),
and paper pondshell (Utterbackia imbecillis). Although common throughout
the drainage, these species tend to be less common in the tributaries
and mainstem of the extreme lowest section of the Wabash River.
Several species are primarily restricted to the mainstem Wabash
River and the lower parts of its largest tributaries. These species
include: yellow sandshell (Lampsilis teres), fragile papershell
(Leptodea fragilis), three-horn wartyback (Obliquaria reflexa),
hickorynut (Obovaria olivaria), pink heelsplitter (Potamilus alatus),
pink papershell (Potamilus ohiensis), monkeyface (Quadrula metanevra),
pimpleback (Quadrula pustulosa pustulosa), mapleleaf (Quadrula
quadrula), pistolgrip (Tritogonia verrucosa), fawnsfoot (Truncilla
donaciformis), and deertoe (Truncilla truncata). These species are also
the most common inhabitants of the tributaries of the lower Wabash
River, where most of the other live species are not found.
Some species are restricted to the tributaries and mainstem of the
upper Wabash River and the upper watersheds of its other larger
tributaries (East Fork and West Fork White rivers). These species
include: mucket (Actinonaias ligamentina), elktoe (Alasmidonta
marginata), wavyrayed lampmussel (Lampsilis fasciola), flutedshell
(Lasmigona costata), round pigtoe (Pleurobema sintoxia), and creeper
(Strophitus undulatus).
Black sandshell (Ligumia recta) is restricted to the lower sections
of the larger tributaries and mainstem of the upper Wabash River. Purple
wartyback (Cyclonaias tuberculata) and pocketbook (Lampsilis ovata) have
similar distributions but are also found in the lower mainstem East Fork
White River.
Fat pocketbook (Potamilus capax) and wartyback (Quadrula nodulata)
have the most restricted ranges of these live species. Both are
concentrated in the extreme lower section of the mainstem Wabash River.
They do ascend some tributaries of the lower mainstem; however, their
reproductive success in these areas is somewhat questionable.
Live tributary species extirpated from mainstem Wabash
River.--Eighteen freshwater mussel species that once maintained
populations throughout the Wabash River drainage are now restricted to
populations that are viable only in the tributaries (Table 1). For many
of these species, live individuals can be found in the mainstem Wabash
River; however, they no longer constitute reproducing populations.
Many species that are now restricted to the tributaries of the
Wabash River have been gone from the mainstem for a long time. These
species include: butterfly (Ellipsaria lineolata), spike (Elliptio
dilatata), snuffbox (Epioblasma triquetra), round hickorynut (Obovaria
subrotunda), clubshell (Pleurobema clava), Ohio pigtoe (Pleurobema
cordatum), salamander mussel (Simpsonaias ambigua), purple lilliput
(Toxolasma lividus), rayed bean (Villosa fabalis), and rainbow (Villosa
iris). Many of these species are now rare in the tributaries, and most
have incurred a substantial reduction in their historic distribution.
Live individuals of the following species can occasionally be found
in the mainstem Wabash River; however, all are functionally extirpated
and restricted to the tributaries: rock pocketbook (Arcidens
confragosus), fanshell (Cyprogenia stegaria), elephantear (Elliptio
crassidens), ebonyshell (Fusconaia ebena), washboard (Megalonaias
nervosa), sheepnose (Plethobasus cyphyus), kidneyshell (Ptychobranchus
fasciolaris), and rabbitsfoot (Quadrula cylindrica cylindrica).
Of these species, rock pocketbook, elephant-ear, ebonyshell, and
washboard are the most likely to be collected live. Though these species
may still maintain small, isolated, reproducing populations somewhere in
the mainstem Wabash River, their continued presence is uncertain at
best. We are considering them extirpated until further collections are
made that can repudiate this claim.
Fanshell, sheepnose, kidneyshell, and rabbits-foot are much rarer
in the mainstem Wabash River than the previously mentioned species. All
have been found sporadically in the mainstem in recent years (Cummings
et al. 1992; Ball & Schoenung 1996; EnviroScience 2006). Occasional
live individuals of these species could be entering the Wabash River
from the lower Tippecanoe River where populations of these species
remain.
Tributary species.--Nine species are found live in tributaries of
the Wabash River drainage (Table 1) and likely never maintained
populations in the mainstem Wabash River in Indiana. These species
include: slippershell mussel (Alasmidonta viridis), flat floater
(Anodonta suborbiculata), cylindrical papershell (Anodontoides
ferussacianus), creek heelsplitter (Lasmigona compressa), pondmussel
(Ligumia subrostrata), lilliput (Toxolasma parvus), Texas lilliput
(Toxolasma texasiensis), pondhorn (Uniomerus tetralasmus), and little
spectaclecase (Villosa lienosa). Shell material of these species is
occasionally found in the mainstem Wabash River, likely washing in from
tributaries.
Slippershell mussel, cylindrical papershell, creek heelsplitter,
and lilliput are small-stream species found in the smaller watersheds
throughout the Wabash River drainage (except the southwest portion).
These species are likely found live in the very upper mainstem Wabash
River in Ohio.
Flat floater, pondhorn and Texas lilliput are primarily restricted
to the southwest portion of the Wabash River drainage, where they
inhabit the ditches, oxbows, and other similar habitats of the region.
Pondmussel can be found in this portion of the drainage as well but is
also an inhabitant of the natural lakes of the upper Wabash River
drainage.
Little spectaclecase is a small- to medium-sized stream species. It
has a rather sporadic distribution within the Wabash River drainage. It
inhabits tributaries of the middle Wabash, East Fork White, and West
Fork White rivers.
Extirpated species.--Eighteen freshwater mussel species are now
considered extirpated from the entire Wabash River drainage (Table 1).
All 18 species are also extirpated from the entire state of Indiana.
These species include: spectaclecase (Cumberlandia monodonta), leafshell
(Epioblasma flexuosa), white cat-spaw (Epioblasma obliquata perobliqua),
round combshell (Epioblasma personata), Tennessee riffleshell
(Epioblasma propinqua), Wabash riffleshell (Epioblasma sampsonii),
northern riffleshell (Epioblasma torulosa rangiana), tubercled blossom
(Epioblasma torulosa torulosa), longsolid (Fusconaia subrotunda),
cracking pearlymussel (Hemistena lata), pink mucket (Lampsilis abrupta),
scale-shell (Leptodea leptodon), ring pink (Obovaria retusa), white
wartyback (Plethobasus cicatricosus), orangefoot pimpleback (Plethobasus
cooperianus), rough pigtoe (Pleurobema plenum), pyramid pigtoe
(Pleurobema rubrum), and winged mapleleaf (Quadrula fragosa). Most of
these species have been lost from the fauna for many years, and several
are even extinct.
A single live rough pigtoe was collected from the East Fork White
River in 1992 (Ball & Schoenung 1996). A single live longsolid was
collected from Sugar Creek in 1991 (Lewis 1991). These represent the
last known live collections of either species from Indiana waters. If
these species are live in the Wabash River drainage they are extremely
rare and are no longer reproducing.
Federally-endangered species.--Three federally-endangered
freshwater mussel species maintain populations in the Wabash River
drainage of Indiana. Clubshell is found throughout the Tippecanoe River
in the upper Wabash River drainage. Fanshell is found in the lower
Tippecanoe River (below Lake Freeman) and the lower East Fork White
River. The fat pocketbook survives in the extreme lowest section of the
mainstem Wabash River, where it is one of the most common live
freshwater mussels found.
ACKNOWLEDGMENTS
Mussel collections by the Wildlife Diversity Section, IDNR, were
funded through State Wildlife Grant T-3. Sampling was aided by a series
of dedicated naturalist aides, including Ted V. Briggs, Kathy M. Eddy,
Kevin L. Rowe, Amelia R. Barnett, Kara L. Bieker, Melissa R. Jones, and
Sarah A. Bales.
Manuscript received 7 September 2006, revised 18 October 2006.
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Table 1.-Current status of freshwater mussels
(Order Unionoida) in the Wabash River drainage of
Indiana. Scientific and common names follow Turgeon
et al. (1998). L = reproducing populations of
species still found in mainstem Wabash River and
its tributaries; XT = reproducing populations of
species historically found in mainstem Wabash River
but now restricted to its tributaries; T = reproducing
populations of species always restricted to
Wabash River tributaries; X = species extirpated
from entire Wabash River drainage.
Species Status
Family Margaritiferidae
Cumberlandia monodonta (spectaclecase) X
Family Unionidae
Actinonaias ligamentina (mucket) L
Alasmidonta marginata (elktoe) L
Alasmidonta viridis (slippershell mussel) T
Amblema plicata (threeridge) L
Anodonta suborbiculata (flat floater) T
Andontoides ferussacianus (cylindrical
papershell) T
Arcidens confragosus (rock pocketbook) XT
Cyclonaias tuberculata (purple warty-
back) L
Cyprogenia stegaria (fanshell) XT
Ellipsaria lineolata (butterfly) XT
Elliptio crassidens (elephantear) XT
Elliptio dilatata (spike) XT
Epioblasma fiexuosa (leafshell) X
Epioblasma obliquata perobliqua (white
catspaw) X
Epioblasma personata (round combshell) X
Epioblasma propinqua (Tennessee rif-
fleshell) X
Epioblasma sampsonii (Wabash riffle-
shell) X
Epioblasma torulosa rangiana (northern
riffleshell) X
Epioblasma torulosa torulosa (tubercled
blossom) X
Epioblasma triquetra (snuffbox) XT
Fusconaia ebena (ebonyshell) XT
Fusconaia flava (Wabash pigtoe) L
Fusconaia subrotunda (longsolid) X
Hemistena lata (cracking pearlymussel) X
Lampsilis abrupta (pink mucket) X
Lampsilis cardium (plain pocketbook) L
Lampsilis fasciola (wavyrayed lampmus-
sel) L
Lampsilis ovata (pocketbook) L
Lampsilis siliquoidea (fatmucket) L
Lampsilis teres (yellow sandshell) L
Lasmigona complanata (white heelsplit-
ter) L
Lasmi gona costata (flutedshell) T
Lasmigona compressa (creek heelsplitter) L
Leptodea fragilis (fragile papershell) L
Leptodea leptodon (scaleshell) X
Ligumia recta (black sandshell) L
Ligumia subrostrata (pondmussel) T
Megalonaias nervosa (washboard) XT
Obliquaria reflexes (threehorn wartyback) L
Obovaria olivaria (hickorynut) L
Obovaria retusa (ring pink) X
Obovaria subrotunda (round hickorynut) XT
Plethobasus cicatricosus (white warty-
back) X
Plethobasus cooperianus (orangefoot
pimpleback) X
Plethobasus cyphyus (sheepnose) XT
Pleurobema clava (clubshell) XT
Pleurobema cordatum (Ohio pigtoe) XT
Pleurobema plenum (rough pigtoe) X
Pleurobema rubrum (pyramid pigtoe) X
Pleurobema sintoxia (round pigtoe) L
Potamilus alatus (pink heelsplitter) L
Potamilus capax (fat pocketbook) L
Potamilus ohiensis (pink papershell) L
Ptychobranchus fasciolaris (kidneyshell) XT
Pyganodon grandis (giant floater) L
Quadrula cylindrica cylindrica (rabbits-
foot) XT
Quadrula fragosa (winged mapleleaf) X
Quadrula metanevra (monkeyface) L
Quadrula nodulata (wartyback) L
Quadrula pustulosa pustulosa (pimple-
back) L
Quadrula quadrula (mapleleaf) L
Simpsonaias ambigua (salamander mus-
sel) XT
Strophitus undulatus (creeper) L
Toxolasma lividus (purple lilliput) XT
Toxolasma parvus (lilliput) T
Toxolasma texasiensis (Texas lilliput) T
Tritogonia verrucosa (pistolgrip) L
Truncilla donaciformis (fawnsfoot) L
Truncilla truncata (deertoe) L
Uniomerus tetralasmus (pondhorn) T
Utterbackia imbecillis (paper pondshell) L
Villosa fabalis (rayed bean) XT
Villosa iris (rainbow) XT
Villosa lienosa (little spectaclecase) T
Table 2.-Summary of status for freshwater mus-sels in the Wabash
River drainage of Indiana.
Status Number of
species
Reproducting populations still found in
the mainstem Wabash River and its
tributaries 30
Reproducing populations historically
found in the mainstem Wabash River
but now restricted to its tributaries 18
Reproducing populations always restricted
to Wabash River tributaries 9
Extirpated from the entire Wabash River
drainage 18
Total species: 75