Endangered and Threatened Wildlife and Plants; Endangered Status for the Sheepnose and Spectaclecase Mussels, 3392-3420 [2011-469]
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FOR FURTHER INFORMATION CONTACT:
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R3–ES–2010–0050; MO
92210–0–0008–B2]
RIN 1018–AV93
Endangered and Threatened Wildlife
and Plants; Endangered Status for the
Sheepnose and Spectaclecase
Mussels
Fish and Wildlife Service,
Interior.
ACTION: Proposed rule.
AGENCY:
We, the U.S. Fish and
Wildlife Service (Service), propose to
list two freshwater mussels, the
spectaclecase mussel (Cumberlandia
monodonta) and sheepnose
(Plethobasus cyphyus) as endangered
under the Endangered Species Act of
1973, as amended (Act). If we finalize
this rule as proposed, it would extend
the Act’s protections to these species
throughout their ranges, including
sheepnose in Alabama, Illinois, Indiana,
Iowa, Kentucky, Minnesota, Mississippi,
Missouri, Ohio, Pennsylvania,
Tennessee, Virginia, West Virginia, and
Wisconsin, and spectaclecase in
Alabama, Arkansas, Illinois, Indiana,
Iowa, Kentucky, Kansas, Minnesota,
Missouri, Nebraska, Ohio, Tennessee,
Virginia, West Virginia, and Wisconsin.
We determined that critical habitat for
these species is prudent, but not
determinable at this time. The Service
seeks data and comments from the
public on this proposed listing rule.
DATES: We will consider comments and
information we receive from all
interested parties by March 21, 2011.
We must receive requests for public
hearings, in writing, at the address
shown in the FOR FURTHER INFORMATION
CONTACT section by March 7, 2011.
ADDRESSES: You may submit comments
by one of the following methods:
• Federal eRulemaking Portal: https://
www.regulations.gov. Follow the
instructions for submitting comments
on docket number FWS–R3–ES–2010–
0050.
• U.S. mail or hand-delivery: Public
Comments Processing, Attn: FWS–R3–
2010–0050; Division of Policy and
Directives Management; U.S. Fish and
Wildlife Service; 4401 North Fairfax
Drive, Suite 222; Arlington, VA 22203.
We will post all comments on https://
www.regulations.gov. This generally
means that we will post any personal
information you provide us (see Public
Comments section below for more
information).
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SUMMARY:
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Richard Nelson, Field Supervisor, at the
U.S. Fish and Wildlife Service, Rock
Island, Illinois Ecological Services Field
Office, 1511 47th Avenue, Moline, IL
61265 (telephone 309–757–5800).
SUPPLEMENTARY INFORMATION:
Public Comments
Our intent is to use the best available
commercial and scientific data as the
foundation for all endangered and
threatened species classification
decisions. We request comments or
suggestions from other concerned
governmental agencies, the scientific
community, industry, or any other
interested party concerning this
proposed rule to list the spectaclecase
and sheepnose mussels as endangered.
We particularly seek comments
concerning:
(1) Biological, commercial trade, or
other relevant data concerning any
threats (or lack thereof) to the species
and regulations that may be addressing
those threats.
(2) Additional information concerning
the ranges, distributions, and
population sizes of the species,
including the locations of any
additional populations of these species.
(3) Any additional information on the
biological or ecological requirements of
these species.
(4) Current or planned activities in the
areas occupied by these species and
possible impacts of these activities on
the species and their habitats.
(5) Potential effects of climate change
on these species and their habitats.
(6) The reasons why areas should or
should not be designated as critical
habitat as provided by section 4 of the
Act (16 U.S.C. 1531 et seq.), including
whether the benefits of designation
would outweigh threats to the species
that designation could cause (e.g.,
exacerbation of existing threats, such as
overcollection), such that the
designation of critical habitat is
prudent.
(7) Specific information on:
• What areas contain physical and
biological features essential for the
conservation of these species;
• What areas are essential to the
conservation of these species and
• Special management considerations
or protection that proposed critical
habitat may require.
Please note that submissions merely
stating support for or opposition to the
action under consideration without
providing supporting information,
although noted, will not be considered
in making a determination, as section
4(b)(1)(A) of the Act directs that
determinations as to whether any
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species is an endangered or threatened
species must be made ‘‘solely on the
basis of the best scientific and
commercial data available.’’
You may submit your comments and
materials concerning this proposed rule
by one of the methods listed in the
ADDRESSES section. We will not accept
comments sent by e-mail or fax or to an
address not listed in the ADDRESSES
section. Comments must be submitted to
https://www.regulations.gov before 11:59
(Eastern Time) on the date specified in
the DATES section. We will not consider
hand-delivered comments that we do
not receive, or mailed comments that
are not postmarked, by the date
specified in the DATES section.
We will post your entire comment—
including your personal identifying
information—on https://www.
regulations.gov. If you provide personal
identifying information in your
comment, you may request at the top of
your document that we withhold this
information from public review.
However, we cannot guarantee that we
will be able to do so.
Comments and materials we receive,
as well as supporting documentation we
used in preparing this proposed rule,
will be available for public inspection
on https://www.regulations.gov, or by
appointment, during normal business
hours at the Rock Island, Illinois
Ecological Services Field Office (see the
FOR FURTHER INFORMATION CONTACT
section).
Public Hearing
The Act provides for one or more
public hearings on this proposal, if
requested. Requests must be received by
March 7, 2011. Such requests must be
made in writing and be addressed to the
Field Supervisor at the address
provided in the FOR FURTHER
INFORMATION CONTACT section. We will
schedule public hearings on this
proposal, if any are requested, and
announce the dates, times, and places of
those hearings, as well as how to obtain
reasonable accommodations, in the
Federal Register and local newspapers
at least 15 days before the hearing.
Persons needing reasonable
accommodations to attend and
participate in a public hearing should
contact the Rock Island, Illinois
Ecological Services Field Office by
telephone at 309–757–5800, as soon as
possible. To allow sufficient time to
process requests, please call no later
than one week before the hearing date.
Information regarding this proposed
rule is available in alternative formats
upon request.
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Background
Species Descriptions
The spectaclecase (Cumberlandia
monodonta) is a member of the mussel
family Margaritiferidae and was
originally described as Unio monodonta
Say, 1829. The type locality is the Falls
of the Ohio (on the Ohio River in the
vicinity of Louisville, Kentucky, and
adjacent Indiana), and the Wabash River
(probably the lower portion in Illinois
and Indiana) (Parmalee and Bogan 1998,
p. 49). Parmalee and Bogan (1998, p. 49)
summarized the synonymy of the
spectaclecase. The species has been
placed in the genera Unio, Margaritana,
Alasmidonta, Margarita, Margaron, and
Margaritifera at various times in history.
Ortmann (1912, p. 13) placed it in the
monotypic (a taxonomic group with
only one biological type) genus
Cumberlandia in the family
Margaritiferidae. Currently recognized
synonymy includes Unio soleniformis
(Lea). Smith (2001, p. 43) reassigned the
spectaclecase to the Holarctic genus
Margaritinopsis based on shell and gill
characters. However, the Service will
defer to the Committee on Scientific and
Vernacular Names of Mollusks of the
Council of Systematic Malacologists,
American Malacological Union
(Turgeon et al. 1998), on whether the
genus Margaritinopsis is accepted as
valid for the spectaclecase. Until an
official decision is made, the Service
will use the commonly accepted
Cumberlandia for the genus of this
species. Spectaclecase is the accepted
common name for Cumberlandia
monodonta (Turgeon et al. 1998, p. 32).
The spectaclecase is a large mussel
that reaches at least 9.25 inches (23.5
centimeters (cm)) in length (Havlik
1994, p. 19). The shape of the shell is
greatly elongated, sometimes arcuate
(curved), and moderately inflated, with
the valves being solid and moderately
thick, especially in older individuals
(Parmalee & Bogan 1998, p. 49). Both
anterior and posterior ends of the shell
are rounded with a shallow depression
near the center of shell (Baird 2000, p.
6; Parmalee & Bogan 1998, p. 49). The
anterior end is higher than the posterior
end (Baird 2000, p. 6). The posterior
ridge is low and broadly rounded
(Parmalee & Bogan 1998, p. 50). Yearone specimens have heavy ridges
running parallel with the growth arrests,
which are shell lines that indicate
slower periods of growth, thought to be
laid down annually (Baird 2000, p. 6).
The periostracum (external shell
surface) is somewhat smooth, rayless,
and light yellow, greenish-tan, or brown
in young specimens, becoming rough
and dark brown to black in old shells
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(Parmalee & Bogan 1998, p. 50). The
shell commonly will crack posteriorly
when dried (Oesch 1984, p. 31).
Internally, the single pseudocardinal
tooth (a triangular tooth-like structure
along the hinge line of the internal
portion of the shell) is simple and peglike in the right valve, fitting into a
depression in the left (Parmalee & Bogan
1998, p. 50). The lateral teeth are
straight and single in the right valve,
and double in the left valve but become
fused with age into an indistinct raised
hinge line (Parmalee & Bogan 1998, p.
50). The soft anatomy was described by
Williams et al. (2008, pp. 497–498). The
color of the nacre (interior covering of
the shell) is white, occasionally granular
and pitted, mostly iridescent in young
specimens, but becoming iridescent
posteriorly in older shells (Parmalee &
Bogan 1998, p. 50). There are no
differences between the sexes in the
shells of this species (Baird 2000, p. 19).
Key characters for distinguishing the
spectaclecase from other mussels are its
large size, elongate shape, arcuate
ventral margin, dark coloration,
roughened periostracum, poorly
developed teeth, and white nacre
(Oesch 1984, pp. 31–32). No other North
American mussel species has this suite
of characters.
The sheepnose (Plethobasus cyphyus)
is a member of the mussel family
Unionidae and was originally described
as Obliquaria cyphya Rafinesque, 1820.
The type locality is the Falls of the Ohio
(Parmalee & Bogan 1998, p. 175) on the
Ohio River in the vicinity of Louisville,
Kentucky, and adjacent Indiana.
Parmalee and Bogan (1998, p. 175)
summarized the synonymy of the
species. Over the years, the name of this
species has been variably spelled
cyphya, scyphius, cyphius, cyphia,
cyphyum, and ultimately cyphyus. Over
the years the species has been placed in
the genera Obliquaria, Unio,
Pleurobema, Margarita, and Margaron.
It was ultimately placed in the genus
Plethobasus by Ortmann (1919, pp. 65–
66) where it remains today (Turgeon et
al. 1998, p. 35). The Service recognizes
Unio aesopus and U. compertus as
synonyms of Plethobasus cyphyus.
Sheepnose is the accepted common
name for Plethobasus cyphyus as
established by the Committee on
Scientific and Vernacular Names of
Mollusks of the Council of Systematic
Malacologists, American Malacological
Union (Turgeon et al. 1998, p. 35). The
Service also recognizes ‘‘bullhead’’ and
‘‘clear profit’’ as older common names
for the sheepnose.
Key characters useful for
distinguishing the sheepnose from other
mussels are its color, the occurrence of
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central tubercles, and its general shape.
Oesch (1984, p. 120) and Parmalee and
Bogan (1998, p. 176), describe the
sheepnose as a medium-sized mussel
that reaches nearly 5 inches (13 cm) in
length. The shell is elongate ovate in
shape, moderately inflated, and with
thick, solid valves. The anterior end of
the shell is rounded, but the posterior
end is somewhat bluntly pointed to
truncate. The dorsal margin of the shell
is nearly straight, while the ventral
margin is uniformly rounded or slightly
convex. The posterior ridge is gently
rounded, becoming flattened ventrally
and somewhat biangular. There is a row
of large, broad tubercular swellings on
the center of the shell extending from
the beak to the ventral margin. A broad,
shallow sulcus (depression on furrow
on the outside surface of shell) lies
between the posterior ridge and central
row. Beaks are elevated, high, and
placed near the anterior margin.
Juvenile beak sculpture consists of a few
concentric ridges at the tip of the beaks.
The periostracum is generally smooth,
shiny, rayless, and light yellow to a dull
yellowish brown. Concentric ridges
resulting from growth arrests are usually
darker.
Oesch (1984, p. 120) describes the
internal anatomy of the sheepnose as
the left valve having two heavy, erect,
roughened, somewhat triangular, and
divergent pseudocardinal teeth. The
right valve has a large, triangular,
roughened pseudocardinal tooth. The
lateral teeth are heavy, long, slightly
curved, and serrated. The beak cavity is
shallow to moderately deep. The soft
anatomy was described by Williams et
al. (2008, p. 94). The color of the nacre
is generally white, but may be pinkish
to cream-colored and iridescent
posteriorly. There are no differences
between the sexes in the shells of this
species. The shell of the sheepnose is
extremely hard and was given the name
‘‘clear profit’’ by early commercial
shellers, being too hard to cut into
buttons (Wilson & Clark 1914, p. 57).
The species also preserves well in
archaeological material (Morrison 1942,
p. 357).
Life History
The general biology of the
spectaclecase and sheepnose are similar
to other bivalve mollusks belonging to
the families Margaritiferidae and
Unionidae, order Unioniformes or
Unionoida. Adult mussels suspensionfeed, spending their entire lives
partially or completely buried within
the substrate (Murray and Leonard 1962,
p. 27). Adults feed on algae, bacteria,
detritus, microscopic animals, and
dissolved organic material (Christian et
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al. 2004, pp. 108–109; Nichols & Garling
2000, p. 873; Silverman et al. 1997, p.
1859; Strayer et al. 2004, pp. 430–431).
Recent evidence suggests that adult
mussels may also deposit feed on
particles in the sediment (Raikow &
Hamilton 2001, p. 520). For their first
several months, juvenile mussels
employ foot (pedal) feeding, consuming
bacteria, algae, and detritus (Yeager et
al. 1994, p. 221).
As a group, mussel longevity varies
tremendously with some species living
only about 4 years (Haag & Rypel 2010,
p. 5) but possibly up to 100 to 200 years
in other species (Ziuganov et al. 2000,
p. 102). However, the vast majority of
species live a few decades (Haag &
Rypel 2010, pp. 4–6). Baird (2000, pp.
54, 59, 67) aged 278 specimens of the
spectaclecase in Missouri by sectioning
the hinge ligament, as most
margaritiferids are aged. The maximum
age determined was 56 years, but he
surmised that some large individuals
may have been older. A very large
specimen (9.25 inches (23.5 cm)) from
the St. Croix River, Minnesota and
Wisconsin, was estimated (based on
external growth ring counts) to be
approximately 70 years old (Havlik
1994, p. 19). Sheepnose longevity has
been reported as being nearly 30 years
(Watters et al. 2009, p. 221). Thick
shelled mussels from large rivers, like
sheepnose, are thought to live longer
than other species (Stansbery 1961, p.
16).
Mussels tend to grow relatively
rapidly for the first few years, and then
slow appreciably at sexual maturity,
when energy presumably is being
diverted from growth to reproductive
activities (Baird 2000, pp. 66–67). In
spectaclecase, the biggest change in
growth rate appears to occur at 10 to15
years of age, which suggests that
significant reproductive investment
does not occur until they reach 10 years
of age (Baird 2000, pp. 66–67).
Margaritiferids and unionids have an
unusual mode of reproduction. With
very few exceptions, their life cycle
includes a brief, obligatory parasitic
stage on a host organism, typically fish.
Eggs develop into microscopic larvae
(glochidia) within special gill chambers
of the female. The female expels the
mature glochidia, which must attach to
an appropriate host species (generally a
fish) to complete development. Host
specificity varies among margaritiferids
and unionids. Some species appear to
use a single host, while others can
transform on several host species.
Following successful infestation,
glochidia encyst (enclose in a cyst-like
structure), remain attached to the host
for several weeks, and then drop off as
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newly transformed juveniles. For further
information on the life history of
freshwater mussels, see Williams et al.
2008.
Mussel biologists know relatively
little about the specific life-history
requirements of the spectaclecase and
sheepnose. Most mussels, including the
spectaclecase and sheepnose, have
separate sexes. Age at sexual maturity of
the spectaclecase was estimated to be 4
to 5 years for males and 5 to 7 years for
females, with sex ratios approximating
50:50 (Baird 2000, p. 24). The
spectaclecase life cycle includes a
parasitic phase; however, despite
extensive investigation, the host species
is not yet known. The spectaclecase is
thought to release glochidia from early
April to late May in the Meramec and
Gasconade Rivers, Missouri (Baird 2000,
p. 26). Gordon and Smith (1990, p. 409)
reported the species as producing two
broods, one in spring or early summer
and the other in the fall, also based on
Meramec River specimens. In the
Meramec and Gasconade Rivers,
however, Baird (2000, pp. 26–27) found
no evidence of two spawns in a given
year.
Age at sexual maturity for the
sheepnose is unknown, but given its
estimated longevity, probably occurs
after a few years. The sheepnose is
thought to be a short-term brooder, with
egg fertilization taking place in early
summer (Parmalee & Bogan 1998, p.
177; Williams et al. 1998, p. 498), and
glochidial release presumably occurring
later in the summer. Hermaphroditism
occurs in many mussel species (van der
Schalie 1966, p. 77), but is not known
for the sheepnose. If hermaphroditism
does occur in the sheepnose, it may
explain the occurrence of small, but
persistent populations over long periods
of time.
Glochidia of spectaclecase and
sheepnose are released in conglutinates
(gelatinous structures containing
numerous glochidia and analogous to
cold capsules). Spectaclecase glochidia
lack hooks (teeth-like structures that
presumably function to pierce through
skin tissue of the host) and are the
smallest glochidia known of any North
American freshwater mussel; they
measure approximately 0.0024 inches
(0.06 mm) in both length and height
(Baird 2000, p. 22). Tens to hundreds of
thousands of glochidia may occur in
each conglutinate. Based on eight
Missouri spectaclecase specimens, the
number of conglutinates released per
female varied from 53 to 88, with a
mean of 64.5 (Baird 2000, p. 23). Total
fecundity (reproductive potential,
including glochidia and ova) in Baird’s
(2000, p. 27) Missouri study varied from
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1.93 to 9.57 million per female. In
mussels, fecundity is related positively
to body size and inversely related to
glochidia size (Bauer 1994, pp. 940–
941). The reproductive potential of the
spectaclecase is therefore phenomenal.
However, the fact that extant
populations are generally skewed
towards larger adults strongly indicates
that survival rates to the adult stage
must be extraordinarily low.
Researchers in Wisconsin observed
female spectaclecase under boulders in
the St. Croix River simultaneously
releasing their conglutinates (Heath
2008, pers. comm.). The spectaclecase
conglutinates are entrained along a
transparent, sticky mucous strand up to
several feet in length (Lee & Hove 1997,
p. 9). Baird (2000, p. 29) observed the
release of loose glochidia and small
fragments of conglutinates. Based on his
observations, he hypothesized that
conglutinates sometimes contain mostly
immature glochidia, and that
conglutinates containing mostly
immature glochidia may be aborted
when disturbed.
Sheepnose conglutinates are narrow
and lanceolate in outline, solid and red
or pink in color, and discharged in
unbroken form (Oesch 1984, pp. 118–
119). Discharge of sheepnose
conglutinates have been observed in late
July (Ortmann 1911, p. 306) and August
(Williams et al. 2008, p. 498). Ortmann
(1911, p. 306) described them as being
pink and ‘‘lying behind the posterior
end of the shell, which were greedily
devoured by a number of minnows.’’
Sheepnose glochidia are semicircular in
outline, with the ventral margin
obliquely rounded, hinge line long, and
medium in size. The length (0.009
inches (0.23 mm)) is slightly greater
than the height (0.008 inches (0.20 mm))
(Oesch 1984, p. 119). Several hundred
glochidia probably occur in each
conglutinate. Judging from the size of
the glochidia, total fecundity (including
glochidia and ova) per female sheepnose
is probably in the tens of thousands.
Like many freshwater mussels, the
complex life histories of the
spectaclecase and sheepnose have many
vulnerable components that may
prevent successful reproduction or
recruitment of juveniles into existing
populations. Glochidia must come into
contact with a specific host species for
their survival to be ensured. Without the
proper host, the glochidia will perish.
The host(s) for the spectaclecase is
unknown, although over 60 species of
fish, amphibians, and crayfish have
been tested in the lab during host
suitability studies (Baird 2000, pp. 23–
24; Henley & Neves 2006, p. 3; Hove et
al. 2009b, pp. 22–23; Hove et al. 1998,
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pp. 13–14; Hove et al. 2008, p. 4;
Knudsen & Hove 1997, p. 2; Lee & Hove
1997, pp. 9–10). Two of 690 wildcollected fish checked by Baird (2000, p.
24) had spectaclecase glochidia attached
to their gills; these fish were the bigeye
chub (Hybopsis amblops) and pealip
redhorse (Moxostoma pisolabrum).
However, these fish are not confirmed
as hosts, because the encysted glochidia
had not grown measurably and
glochidial transformation was not
observed (Baird 2000, p. 24).
Spectaclecase populations are
oftentimes highly aggregated (see
Habitat) with many apparently evenaged individuals, suggesting that
glochidia may excyst simultaneously
from a host (Gordon & Layzer 1989, p.
19). Additional host work is underway
to test the wild-collected fish species
that were found with encysted
spectaclecase glochidia (pealip redhorse
and bigeye chub), as well as to test
additional species of fish and other
aquatic organisms for suitability. Host
information is needed so that existing
populations can be artificially cultured
for potential population augmentation
and reintroduction efforts.
Little is known regarding host fish of
the sheepnose. Until recently the only
cited host for this species came from a
1914 report that found glochidia
naturally attached to sauger (Sander
canadense) in the wild. No confirmation
of successful transformation was
recorded in this early report (Surber
1912, p. 110; Wilson 1914, pp. 338–
340). However, recent laboratory studies
at the Genoa National Fish Hatchery, the
University of Minnesota, and Ohio State
University have successfully
transformed sheepnose glochidia on
fathead minnow (Pimephales promelas),
creek chub (Semotilus atrromaculatus),
central stoneroller (Campostoma
anomalum), and brook stickleback
(Culaea inconstans) (Watters et al. 2005,
pp. 11–12; Brady 2008, pers. comm.;
Watters 2008, pers. comm.). Although
these are identified as suitable hosts in
laboratory studies, natural interactions
between the aforementioned fishes and
the sheepnose seem rare and infrequent
due to habitat preferences. Fish that
frequent medium to large rivers near
mussel beds, like the sauger, may act as
hosts in the natural environment.
Habitat
The spectaclecase generally inhabits
large rivers, and is found in
microhabitats sheltered from the main
force of current. It occurs in substrates
from mud and sand to gravel, cobble,
and boulders in relatively shallow riffles
and shoals with a slow to swift current
(Baird 2000, pp. 5–6; Buchanan 1980, p.
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13; Parmalee & Bogan 1998, p. 50).
According to Stansbery (1967, pp. 29–
30), this species is usually found in firm
mud between large rocks in quiet water
very near the interface with swift
currents. Specimens have also been
reported in tree stumps, in root masses,
and in beds of rooted vegetation (Oesch
1984, p. 33). Similar to other
margaritiferids, spectaclecase
occurrences throughout much of its
range tend to be aggregated (Gordon &
Layzer 1989, p. 19), particularly under
slab boulders or bedrock shelves (Baird
2000, p. 6; Buchanan 1980, p. 13;
Parmalee & Bogan 1998, p. 50), where
they are protected from the current. Up
to 200 specimens have been reported
from under a single large slab in the
Tennessee River at Muscle Shoals,
Alabama (Hinkley 1906, p. 54). Unlike
most species that move about to some
degree, the spectaclecase may seldom if
ever move except to burrow deeper and
may die from stranding during droughts
(Oesch 1984, p. 17).
The sheepnose is primarily a largerstream species occurring primarily in
shallow shoal habitats with moderate to
swift currents over coarse sand and
gravel (Oesch 1984, p. 121). Habitats
with sheepnose may also have mud,
cobble, and boulders. Sheepnose in
larger rivers may occur at depths
exceeding 6 m (Williams et al. 2008, p.
498).
Genetics
A recent genetic study (Monroe et al.
2007, pp. 7–13) indicates that much of
the remaining genetic variability in the
spectaclecase is represented in each of
the remaining large populations, and
that these populations do not appear to
differ significantly from one another.
Genetics studies of sheepnose are
currently under investigation; however,
no conclusions were available at the
time of publication (Roe 2010, pers.
comm.).
Species Distribution
We use the term ‘‘population’’ here in
a geographical and not genetic sense,
defining it as all individuals of the
spectaclecase or sheepnose living in one
stream. Using the term in this way
allows the status, trends, and threats to
be discussed comparatively across
streams where the species occur. In
using this term we do not imply that
their populations are currently
reproducing and recruiting or that they
are distinct genetic units. We
considered populations of the
spectaclecase and sheepnose as extant if
live or fresh-dead specimens have been
observed or collected since 1990. A
‘‘population cluster’’ refers to where two
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3395
or more adjacent stream populations of
a species occur without a barrier (for
example, a dam and impoundment)
between them.
Following are generalized sets of
criteria that were used to categorize the
relative status of populations of
spectaclecase and sheepnose. The status
of a population is considered
‘‘improving’’ if: (1) There is evidence
that habitat degradation appears
insignificant, (2) live or fresh dead
mussel abundance has improved during
post-1990 surveys, or (3) ample
evidence of recent recruitment has been
documented during post-1990 surveys.
The status of a population is considered
‘‘stable’’ if: (1) There is little evidence of
significant habitat loss or degradation,
(2) live or fresh dead mussel abundance
has been fairly consistent during post1990 surveys, or (3) evidence of
relatively recent recruitment has been
documented during post-1990 surveys.
The status of a population is considered
‘‘declining’’ if: (1) There is ample
evidence of significant habitat loss or
degradation, (2) live or fresh dead
mussel numbers have declined during
recent surveys, or (3) no evidence of
relatively recent recruitment has been
documented during recent surveys. The
status of a population is considered
‘‘extirpated’’ if: (1) All known suitable
habitat has been destroyed, or (2) no live
or fresh dead mussels of any age have
been located during recent surveys. The
status of a population is considered
‘‘unknown’’ if the available information
is inadequate to place the population in
one of the above four categories. In a
few cases, additional information not
listed above may have been used to
categorize a population.
Spectaclecase Historical Range and
Distribution
The spectaclecase occurred
historically in at least 44 streams in the
Mississippi, Ohio, and Missouri River
basins (Butler 2002a, p. 6, Heath 2008,
pers. comm.). Its distribution comprised
portions of 15 States (Alabama,
Arkansas, Illinois, Indiana, Iowa,
Kansas, Kentucky, Minnesota, Missouri,
Nebraska, Ohio, Tennessee, Virginia,
West Virginia, and Wisconsin).
Historical occurrence by stream system
(with tributaries) include the: upper
Mississippi River system (Mississippi
River (St. Croix, Chippewa, Rock, Salt,
Illinois (Des Plaines, Kankakee Rivers),
Meramec (Bourbeuse, Big Rivers),
Kaskaskia Rivers; Joachim Creek));
lower Missouri River system (Missouri
River (Platte, River Aux Vases, Osage
(Sac, Marais des Cygnes Rivers),
Gasconade (Osage Fork, Big Piney River)
Rivers)); Ohio River system (Ohio River
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(Muskingum, Kanawha, Green, Wabash
Rivers)); Cumberland River system
(Cumberland River (Big South, Caney
Fork; Stones, Red Rivers)); Tennessee
River system (Tennessee River (Holston,
Nolichucky, Little, Little Tennessee,
Clinch (Powell River), Sequatchie, Elk,
Duck Rivers)); lower Mississippi River
system (Mulberry, Ouachita Rivers).
Spectaclecase Current Range and
Distribution
Extant populations of the
spectaclecase are known from 19
streams in 11 States (Butler 2002b, p. 7).
These include the following stream
systems (with tributaries):
• Upper Mississippi River system
(Mississippi River (St. Croix, Meramec
(Bourbeuse, Big Rivers) Rivers));
• Lower Missouri River system (Sac
and Gasconade (Osage Fork, Big Piney
River) Rivers);
• Lower Ohio River system
(lowermost Ohio River (Kanawha, Green
Rivers));
• Cumberland River system
(Cumberland River);
• Tennessee River system (Tennessee
River (Nolichucky, Clinch, Duck
Rivers)); and
• Lower Mississippi River system
(Mulberry, Ouachita Rivers).
The 19 extant spectaclecase
populations occur in the following 11
States (with streams):
• Alabama (Tennessee River),
• Arkansas (Mulberry, Ouachita
Rivers),
• Illinois (Mississippi, Ohio Rivers),
• Iowa (Mississippi River),
• Kentucky (Ohio, Green Rivers),
• Minnesota (Mississippi, St. Croix
Rivers),
• Missouri (Mississippi, Meramec,
Bourbeuse, Big, Gasconade, Sac, Big
Piney Rivers; Osage Fork),
• Tennessee (Tennessee, Clinch,
Nolichucky, Duck Rivers; Caney Fork),
• Virginia (Cumberland, Clinch
Rivers),
• West Virginia (Kanawha River), and
• Wisconsin (Mississippi, St. Croix
Rivers).
Spectaclecase Population Estimates and
Status
Based on historical and current data,
the spectaclecase has declined
significantly rangewide and is now
known from only 19 of 44 streams
(Table 1), representing a 57 percent
decline. The species is presumed
extirpated from thousands of river miles
and from numerous reaches of habitat in
which it occurred historically, including
long reaches of upper Mississippi, Ohio,
Cumberland, and Tennessee Rivers and
many other streams and stream reaches.
Of the 19 extant populations, 6 are
represented by only one or two recent
specimens each and are likely declining
and some may be extirpated.
Populations in Mississippi and Clinch
Rivers have recently experienced
significant population declines. Most
surviving populations face significant
threats and with few exceptions are
highly fragmented and restricted to
short stream reaches. The spectaclecase
is considered extirpated from Indiana,
Kansas, Nebraska, and Ohio. The only
relatively strong populations remaining
are in the Meramec and Gasconade
Rivers in Missouri and in the St. Croix
River in Minnesota and Wisconsin.
TABLE 1—SPECTACLECASE STATUS IN ALL STREAMS OF HISTORICAL OR CURRENT OCCURRENCE
River Basin
Stream
Current Status
Date of
Last Live
Observation
Upper Mississippi River ...................
Mississippi River .............................
St. Croix River ................................
Chippewa River ..............................
Rock River ......................................
Salt River ........................................
Illinois River ....................................
Des Plaines River ...........................
Kankakee River ..............................
Meramec River ...............................
Bourbeuse River .............................
Big River .........................................
Kaskaskia River ..............................
Joachim Creek ................................
Missouri River .................................
Platte River .....................................
River Aux Vases .............................
Osage River ....................................
Sac River ........................................
Marais des Cygnes River ...............
Gasconade River ............................
Big Piney River ...............................
Osage Fork .....................................
Ohio River .......................................
Muskingum River ............................
Kanawha River ...............................
Green River ....................................
Wabash River .................................
Cumberland River ...........................
Big South Fork ................................
Caney Fork .....................................
Stones River ...................................
Red River ........................................
Tennessee River .............................
Holston River ..................................
Nolichucky River .............................
declining .............
stable ..................
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
stable ..................
stable ..................
stable ..................
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
declining .............
extirpated ...........
stable ..................
unknown .............
unknown .............
declining .............
extirpated ...........
unknown .............
unknown .............
extirpated ...........
unknown .............
extirpated ...........
extirpated ...........
extirpated ...........
extirpated ...........
unknown .............
extirpated ...........
unknown .............
2009 .......
2008 .......
1989 .......
∼1970 .....
1980 .......
∼1914 .....
∼1921 .....
1906 .......
2003 .......
1997 .......
2002 .......
∼1970 .....
∼1965 .....
∼1914 .....
∼1917 .....
∼1974 .....
1980 .......
2001 .......
unknown
2007 .......
2004 .......
1999 .......
1994 .......
unknown
2005 .......
2006 .......
1970 .......
2008 .......
1911 .......
1988 .......
1968 .......
1966 .......
2001 .......
1981 .......
1991 .......
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Lower Missouri River .......................
Ohio River ........................................
Cumberland River ............................
Tennessee River ..............................
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Comments
relic shell observed in 1998.
single individual observed.
relic shell observed in 1995.
two live individuals observed.
single individual observed.
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3397
TABLE 1—SPECTACLECASE STATUS IN ALL STREAMS OF HISTORICAL OR CURRENT OCCURRENCE—Continued
Stream
Current Status
Date of
Last Live
Observation
Little River .......................................
Little Tennessee River ....................
extirpated ...........
extirpated ...........
∼1911 .....
unknown
Clinch River ....................................
Powell River ....................................
Sequatchie River ............................
Elk River .........................................
Duck River ......................................
Mulberry River ................................
Ouachita River ................................
declining .............
extirpated ...........
extirpated ...........
extirpated ...........
declining .............
unknown .............
declining .............
2005 .......
∼1978 .....
∼1925 .....
unknown
1999 .......
∼1995 .....
2000 .......
River Basin
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Lower Mississippi River ...................
Based on collections made over 100
years ago, the spectaclecase was
historically widespread and locally
common in many streams rangewide.
The spectaclecase is often absent from
archaeological shell middens (Morrison
1942, p. 353) and is generally difficult
to find due to its habit of occurring
under rocks or ledges and burrowing
deep into the substrate (Parmalee 1967,
p. 25). Therefore, the chance of casually
finding the species where population
numbers are low is remote.
The spectaclecase was considered a
rare species by mussel experts as early
as 1970 (Stansbery 1970, p. 13), when
the first attempt was made to compile a
list of imperiled mussels. The
spectaclecase is considered widely
distributed but absent from many areas
where it formerly occurred (Cummings
& Mayer 1992, p. 22). The American
Malacological Union and American
Fisheries Society consider the
spectaclecase to be threatened (Williams
et al. 1993, p.10). Six of the 19 streams
(or big river reaches) considered to
harbor extant populations of the
spectaclecase are represented by one or
two recent specimens (for example,
Ohio, Kanawha, Cumberland, Duck,
Ouatchita, and Mulberry Rivers),
exemplifying the species’ imperiled
status rangewide.
In some streams, the last reported
records for the spectaclecase occurred
decades ago (for example, Rock, Des
Plaines, Kaskaskia, Platte, Wabash,
Stones, Red, and Little Rivers; River
Aux Vases; Big South Fork). Parmalee
(1967, p. 25) considered the
spectaclecase to be ‘‘rare and of local
occurrence’’ in Illinois in the 1960s, but
that it had ‘‘[a]pparently already been
extirpated from the Illinois and
Kankakee Rivers.’’ The only records
known from some streams are relic
specimens collected around 1975 (for
example, Marais des Cygnes,
Muskingum, and Elk Rivers).
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Although quantitative historical
abundance data for the spectaclecase is
rare, generalized relative abundance (the
percent abundance of a species, divided
by the total abundance of all mussel
species combined) was sometimes noted
in the historical literature and can be
inferred from museum lots. The
following is a summary of what is
known about the relative abundance
and trends of presumably extant
spectaclecase populations by stream
system.
Upper Mississippi River System
The spectaclecase was historically
known from 13 streams in the upper
Mississippi River system. Currently,
only four streams in the system are
thought to have extant spectaclecase
populations.
Mississippi River mainstem: In 1907,
Bartsch found spectaclecase at
approximately nine of the 140 sampled
sites from what are now Mississippi
River Pools (MRP) 9 to 22 (Havlik
2001b, p. 10). Grier (1922, p. 11) did not
find spectaclecase in sampled portions
of MRP 4 to 6. Van der Schalie and van
der Schalie (1950, p. 456), reporting on
studies from the upper Mississippi
River to the Missouri River mouth,
stated that no live spectaclecase were
found in their study of 254 sites during
1930–31. Havlik and Stansbery (1977, p.
12) thought the spectaclecase had
disappeared from MRP 8 by the 1920s.
Thiel (1981, p. 10) found only shell
material in MRP 11 in a survey that
spanned MRP 3 to 11 conducted during
1977 to 1980. Whitney et al. (1997, p.
12) recorded a single individual during
1994–1995 in MRP 15, for a density of
0.004 per square foot (sq. ft) (0.04 per
square meter (sq. m)). Helms (2008, p.
8) found eight live individuals and
numerous shells during a recent search
of MRP 19, representing the most recent
and numerous collection of the species
in the Mississippi River.
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Comments
relic shell observed in 1980, previous record archaeological.
relic shell observed in 1998.
single individual observed.
single individual observed.
two individuals observed.
The spectaclecase is thought to be
extant in at least four pools of the
Mississippi River mainstem, albeit in
very low numbers. Records include
MRP 15 (Quad Cities area, Illinois and
Iowa; in 1998), MRP 16 (Muscatine area,
Iowa and Illinois in 1997), MRP 19
(Burlington area, Illinois and Iowa in
2009), and MRP 22 (Quincy, Illinois and
Hannibal, Missouri, area in 1996).
Populations may still persist in MRP 9
and 10 where specimens were found in
the 1980s (Heath 2010a, pers. comm.).
Only a relic spectaclecase shell was
found in MRP 3 above the St. Croix
River confluence in 2001, and none
were found in subsequent surveys
(Kelner 2008, pers. comm.). In general,
spectaclecase population levels in the
upper Mississippi River appear to have
always been fairly small and difficult to
locate, and are now of questionable
long-term persistence.
St. Croix River: The northernmost and
one of the three most significant extant
populations of the spectaclecase occurs
in the St. Croix River, Minnesota and
Wisconsin. The population is primarily
found in the middle reaches of the river
in Chisago and Washington Counties,
Minnesota, and Polk and St. Croix
Counties, Wisconsin (river miles (RM)
17 to 118). Havlik (1994, p. 19) reported
spectaclecase in the St. Croix Wild River
State Park portion of the river
(approximately RM 62 to 65) and the
reproducing population below the St.
Croix Falls Dam at St. Croix Falls,
Wisconsin (dam located at
approximately RM 52). Additional
survey work in the lower river at Afton
State Park (approximately RM 7 to 9)
failed to find the spectaclecase (Havlik
1994, p. 19).
Hornbach (2001, p. 218) reported 68
live specimens from 4 of 16 river
reaches. Relative abundance for the
spectaclecase varied from 0.67 percent
from RM 78 to 92 (20 live spectaclecase
among 17 species collected), 0.008
percent from RM 63 to 78 (41 live, 24
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species), 0.0006 percent from RM 42 to
52 (6 live, 33 species), and 0.003 percent
from RM 40 to 42 (1 live, 21 species).
Reaches where the spectaclecase is
extant are fragmented by the pool
formed from the power dam at St. Croix
Falls.
Baird (2000, p. 70) presented a lengthfrequency histogram for the
spectaclecase in the St. Croix River
using data from an unpublished 1989
study. The 962 specimens were fairly
evenly distributed over the length scale,
indicating multiple age classes
including healthy numbers of young
spectaclecase recruiting into the
population. Baird (2000, p. 70) used
growth curves determined from his
Missouri study of the species to estimate
the ages of spectaclecase of known size
in the St. Croix River. The percentage of
newly recruited individuals (less than
or equal to 10 years of age) in the St.
Croix was 40 percent— considerably
higher than that noted from the
Gasconade (10.4 percent) and Meramec
(2.8 percent) Rivers in Missouri, two
other streams with abundant
spectaclecase populations that he
studied. The St. Croix spectaclecase
population, while among the largest
known, may also be the healthiest based
on this metric. The spectaclecase is
currently distributed from RM 17 to 118
and appears to be recruiting from RM 17
to 54 (downstream of the St. Croix Falls
Dam) (Heath 2008, pers. comm.).
The long-term health of mussel
populations in the St. Croix may be in
jeopardy, however. Hornbach et al.
(2001, pp. 12–13) determined that
juvenile mussel density had suffered a
statistically significant decline at three
of four lower St. Croix sites sampled in
the 1990s and in 2000. Zebra mussels
also threaten the spectaclecase and
other mussel populations in the lower
St. Croix River. A 2000 survey at 20
sites on the lowermost 24 miles of the
St. Croix River estimated that nearly one
percent of the mussels were infested
with zebra mussels (Kelner & Davis
2002, p. 36).
Meramec River: The Meramec River
flows into the Mississippi River
downstream of St. Louis in east-central
Missouri. Its spectaclecase population
represents one of the best remaining
rangewide. In the late 1970s, Buchanan
(1980, p. 13) reported this species from
31 sites, 19 with live individuals. Live
or fresh dead individuals occurred from
RM 17.5 to 145.7. Buchanan (1980, p. 6)
considered it to be common in the lower
108 miles (174 km) of the Meramec
River, but locally abundant from RM
17.5 to 84. In 1997, Roberts and
Bruenderman (2000, pp. 39, 44), using
similar sampling methods as Buchanan
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(1980, pp. 4–5), resurveyed the
Meramec River system and collected
spectaclecase from 23 sites, 19 of which
had live individuals. They found the
largest populations between RM 56.7
and 118.8. Among 17 sites where
spectaclecase were found during both
surveys, the species was less abundant
at nine sites and more abundant at five
sites in 1997. At three sites, only relic
shells were found during both surveys.
In the 1970s, Buchanan (1980, p. 10)
reported finding 456 live individuals
among the 17 shared sites, whereas
Roberts and Bruenderman (2000, p. 44)
recorded only 198. A reduction in
spectaclecase numbers (260 to 33) at RM
59.5 accounted for most of the overall
decrease in abundance between the
studies. Confounding the decrease in
numbers among shared survey sites,
Roberts and Bruenderman (2000, p. 44)
surveyed three sites between RM 56.7
and 118.8 that were unsampled by
Buchanan (1980, pp. 1–69) and found
500, 538, and 856 live spectaclecase.
The most specimens found at a single
site in the earlier study was 260 (RM
59.5). Currently, the population in the
Meramec River stretches over much of
the mainstem, a distance of over 100
miles (161 km) from RM 18.5 to 120.4.
The spectaclecase represented 28
percent of all mussels sampled in the
Meramec River in 1997 (Roberts &
Bruenderman 2000, p. 39). Baird (2000,
pp. 62, 68,77) extensively studied the
demographics of the Meramec River
spectaclecase population in the late
1990s. The mean estimated age of the
population was 32 years. Individuals
less than 10 years of age comprised only
2.8 percent of the Meramec population
sampled (a total of 2,983 individuals).
At the four sites he intentionally
selected for their large spectaclecase
populations, densities ranged from 0.01
to 0.12 per sq. ft (0.1 to 1.3 per sq. m)
while estimated population numbers at
these sites ranged from 933 to 22,697.
Baird (2000, p. 71) thought that
conditions for spectaclecase recruitment
in the Meramec had declined in the past
20 to 30 years, but the causes were
undetermined. The prevalence of larger
adults in the Meramec population may
be cause for concern, as it appears to
indicate a low level of recruitment in
the population.
Bourbeuse River: The Bourbeuse River
is a northern tributary of the Meramec
River joining it at RM 68. Its
spectaclecase population was sampled
in 1997 at a single site (RM 10.3), and
7 live individuals were found (Roberts
& Bruenderman 2000, p. 91). Sampling
near the mouth (RM 0.4), Buchanan
(1980, p. 16) found only relic shells. The
Bourbeuse population is probably
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dependent on the much larger Meramec
population for long-term sustainability.
Big River: Another Meramec tributary
with a population of the spectaclecase,
the Big River flows northward into the
Meramec River at RM 38. The
spectaclecase is only known from the
lower end (RM 1.3), where 14 live
specimens were found in 1997 (Roberts
& Bruenderman 2000, p. 96). At RM 0.4,
Buchanan (1980, p. 13) found only relic
shells. Similar to the Bourbeuse River
population, the population in the Big
River is probably dependent on the
much larger Meramec population for
sustainability. The Meramec River
system, including the lower Bourbeuse,
lower Big, and Meramec River
mainstems, can be considered a single
spectaclecase population cluster.
Lower Missouri River System
The spectaclecase was historically
known from 10 streams in the Missouri
River system. Currently, only four of
these streams are thought to have extant
populations.
Sac River: The Sac River is a large
tributary to the Osage River. The
spectaclecase was considered extirpated
in the 2002 status review of the species
(Butler 2002a). However, three old, live
individuals were collected at two sites
during a survey of the Sac River in 2004
(Hutson & Barnhart 2004, p. 17). The
same survey revealed ‘‘numerous’’ relic
shells from six other sites, indicating
that the spectaclecase may have been
relatively abundant at one time. Prior to
the 2004 survey, the spectaclecase had
not been collected from this river since
1978 (Bruenderman 2001, pers. comm.).
Given the age of the live individuals and
the abundance of shell material, Hutson
& Barnhart (2004, p. 17) predicted the
species would ‘‘soon be extirpated’’ from
the river.
Gasconade River: The Gasconade
River is a southern tributary of the
Missouri River in south-central Missouri
and flows into the mainstem east of
Jefferson City. When Stansbery (1970, p.
13) included this species in the first
compiled list of imperiled mussels, he
noted that ‘‘the only population of
substantial size presently known is
found in the Gasconade River.’’ In 1994,
Buchanan found over 1,000 individuals
between RM 7 and 84 (Buchanan 1994,
pp. 5, 8–13). Today, one of the three
best spectaclecase populations
remaining rangewide occurs in the
Gasconade. The spectaclecase
population occurs over approximately
200 miles (322 km) of the mainstem
from RM 4.9 upstream (Bruenderman et
al. 2001, p. 54). Baird (2000, pp. 61, 71)
studied the demographics of the
Gasconade River spectaclecase
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population in the late 1990s. Based on
his limited number of sampling sites,
this species comprised about 20 percent
of the entire mussel fauna in this
system. The mean estimated age of the
population was 25 years. Individuals
less than 10 years of age comprised 10.4
percent of the Gasconade population
sampled (n = 2,111), indicating a
significant level of recent recruitment.
Historically, Stansbery (1967, p. 29)
noted that ‘‘[t]he size of some
aggregation[s] * * * is impressive,’’ and
that ‘‘the number of individuals may
reach a density of well over a dozen per
square foot.’’ Both statements are
probably in reference to the Gasconade
River, Missouri, population, which he
had described in the text of his note.
Densities at the four sites Baird (2000,
pp. 61, 71) intentionally selected for
their large spectaclecase populations
ranged from 0.03 to 0.06 per sq. ft (0.3
to 0.6 per sq. m); estimated population
numbers at these selected sites ranged
from 2,156 to 4,766. Baird (2000, p. 71)
thought that conditions for
spectaclecase recruitment in the
Gasconade River had declined in the
past 20 to 30 years, but the causes were
undetermined.
Big Piney River: The Big Piney River,
a southern tributary of the Gasconade
River, harbors a small population of the
spectaclecase. Although overlooked
during a 1999 survey (Bruenderman et
al. 2001, pp. 14, 28), 15 individuals
were collected from the lower mainstem
(RM 24) in 2004 (Barnhart et al. 2004,
p. 5). The status of the population is
unknown, but it is probably dependent
on the much larger source population in
the Gasconade River for sustainability
(McMurray 2008, pers. comm.).
Osage Fork: The Osage Fork is a
southwestern headwater tributary of the
Gasconade River. The spectaclecase is
known from the lower portion of this
Gasconade River tributary, specifically
from RM 13.9. Sampling in the Osage
Fork in 1999 yielded 26 live individuals
from this site (Bruenderman et al. 2001,
p. 9). Relative abundance of the
spectaclecase in the Osage Fork was 3.9
percent, and catch-per-unit effort was
1.3 per person-hour. This population is
thought to be stable, but it may also be
dependent on the much larger source
population in the Gasconade River for
long-term sustainability. The Gasconade
River system, including the lower Big
Piney, lower Osage Fork, and Gasconade
mainstems, can be considered a single
population cluster.
Ohio River System
The spectaclecase’s continued
existence in the Ohio River is extremely
uncertain. Once known from five rivers,
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it has been extirpated from two, and two
of the remaining three are recently
represented by only one or two
individuals each.
Ohio River: The Ohio River is the
largest eastern tributary of the
Mississippi River, with its confluence
marking the divide between the upper
and lower portions of the Mississippi
River system. Historically, the
spectaclecase was documented from the
Ohio River from the vicinity of
Cincinnati, Ohio, to its mouth. Although
no specimens are known from the
mainstem upstream of Cincinnati,
populations are known from two
upstream tributaries, the Muskingum
and Kanawha Rivers. Nearly all
spectaclecase records from the Ohio
River were made around 1900 or before
(Schuster 1988, p. 186). The only recent
record is for a single live individual
found in an abandoned gill net near the
Illinois shore in 1994 (Cummings 2008,
pers. comm.). If a population of the
spectaclecase continues to occur in the
Ohio River, its future persistence is
extremely doubtful and continued
existence seriously threatened by the
exotic zebra mussel.
Kanawha River: The Kanawha River is
a major southern tributary of the Ohio
River that drains much of West Virginia.
The spectaclecase was not known from
this stream until 2002, when a single,
very old, live individual was discovered
near Glasgow, Kanawha County
(Zimmerman 2002, pers. comm.).
Another live individual was found in
the same vicinity in 2005, as well as two
additional weathered shells in 2006
(Clayton 2008a, pers. comm.). This site
is approximately 20 miles (32.2 km)
downstream of Kanawha Falls, below
which is the only significant mussel bed
known from the Kanawha River. It is
doubtful that a recruiting spectaclecase
population occurs in the Kanawha River
due to the small number of individuals
found and their advanced age.
Green River: The Green River is a
lower Ohio River tributary in westcentral Kentucky. The spectaclecase has
been collected sparingly in the Green
River. That it was not reported in early
collections made in the system is
indicative of the difficulty in finding
specimens (Price 1900, pp. 75–79).
Stansbery (1965, p. 13) was the first to
find it in the mid-1960s at Munfordville,
Hart County, where he reported 47
mussel species collected over a severalyear period in the early 1960s. More
recently, from 1987 to 1989, Cicerello
and Hannan (1990, p. 20) reported
single fresh dead specimens at six sites
and relic specimens from an additional
five sites in Mammoth Cave National
Park (MCNP). A single specimen was
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3399
recorded from MCNP, Edmonson
County, in 1995. Sampling conducted
from 1996 to 1998 located fresh dead
specimens at two sites above MCNP,
with a relic shell at a third site farther
upstream (Cicerello 1999, pp. 17–18). At
least one fresh dead specimen was
reported from MCNP in 2001, as well as
several live individuals in 2005 and
2006 (Layzer 2008, pers. comm.).
A small spectaclecase population
remains in the upper Green River from
below Lock and Dam 5 upstream
through MCNP, Edmonson County, into
western Hart County. Most recent
specimens have been reported from the
upstream portion of this reach, where it
is generally distributed from MCNP
upstream to western Hart County. Its
distribution is much more sporadic and
localized in the lower portion of this
reach due to the pooling effect of two
locks and dams (5 and 6). In 2001, a
concerted effort (approximately 15
person-hours) to locate rare mussels
below Lock and Dam 5 and at other sites
downstream failed to find spectaclecase
(live or shell), although a fresh dead
shell had been collected in this area in
1993 (Cicerello 2008, pers. comm.). The
occurrence of variable-sized individuals
in the 1990s indicates different year
classes but not necessarily recent
recruitment (Cicerello 2008, pers.
comm.). The long-term sustainability of
the Green River population, primarily
limited to an approximately 15-mile (24km) reach of the river, is therefore
questionable, and its status is unknown.
Cumberland River System
With few exceptions, most records of
the spectaclecase in the Cumberland
River system were made before the
1920s. It was historically known from
the mainstem and four tributaries but
appears currently to be restricted to the
lowermost Cumberland River a few
miles above its confluence with the
Ohio River.
Cumberland River mainstem: The
Cumberland River is a large southern
tributary of the lower Ohio River. The
spectaclecase was considered ‘‘not rare’’
in the Cumberland River by Hinkley and
Marsh (1885, p. 6), whereas it was found
at six sites by Wilson and Clark (1914,
pp. 17, 19) during their survey primarily
for commercial species in the
Cumberland River system. In a 1947–
1949 survey of the Kentucky portion of
the upper Cumberland River, Neel and
Allen (1964, p. 453) reported live
specimens only from one of six
mainstem sites that they sampled below
Cumberland Falls. Neel and Allen
(1964, p. 432) considered it to be
‘‘uncommon’’ in the lower Cumberland
River (where they did not sample), a
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statement possibly based on its sporadic
occurrence as reported by Wilson and
Clark (1914, pp. 17, 19). One of the last
mainstem records is that of a single live
specimen found in the cold tailwaters of
Wolf Creek Dam, Kentucky, near the
Tennessee border in 1982 (Miller et al.
1984, p. 108). This was one of only two
live mussels found during a survey of
the dewatered river reach below the
dam, the mussel community having
been eliminated from decades of cold
water releases. The most recent record
is of a single live individual found at
RM 10 below Barkley Lock and Dam in
2008 (Fortenbery 2008, p. 9). A
thorough search of the area yielded no
additional individuals.
Tennessee River System
The spectaclecase was originally
known from the Tennessee River and
nine of its stream systems. Ortmann
(1924, p. 60) reported that the
spectaclecase was ‘‘frequent… in the
upper Tennessee,’’ while acknowledging
in an earlier paper (Ortmann 1918, p.
527) that it was locally abundant in
parts of the upper Tennessee River
system, but noted that it was ‘‘generally
regarded as a rare species’’ rangewide.
Hundreds of miles of large river
habitat on the Tennessee mainstem have
been converted under nine reservoirs,
with additional dams constructed in
tributaries historically harboring this
species (for example, Clinch, Holston,
and Elk Rivers). Watters (2000, p. 262)
summarizes the tremendous loss of
mussel species from various reaches of
the Tennessee. The spectaclecase is now
known only from the Tennessee
mainstem and three of its tributaries.
Despite this fact, the Tennessee River
system continues to represent one of the
last strongholds of the spectaclecase
rangewide.
Tennessee River mainstem: The
Tennessee River is the largest tributary
of the Ohio River, draining portions of
seven states. The 53-mile (85-km)
stretch of river in northwestern Alabama
collectively referred to as the Muscle
Shoals historically harbored 69 species
of mussels, making it among the most
diverse mussel faunas ever known
(Garner & McGregor 2001, p. 155). The
historical spectaclecase population in
this reach was thought to be
phenomenal given the amount of
historical habitat that was available and
literature accounts of the period.
Hinkley (1906, p. 54), in 1904,
considered the spectaclecase ‘‘plentiful,’’
noting 200 individuals under a single
slab boulder. Twenty years later,
Ortmann (1925, p. 327) stated that ‘‘this
species must be, or have been,
abundant’’ at Muscle Shoals based on
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the ‘‘considerable number of dead
shells’’ he observed. In these quotes he
predicted the demise of the
spectaclecase. The construction of three
dams (Wilson in 1925, Wheeler in 1930,
Pickwick Landing in 1940) inundated
most of the historical habitat, leaving
only small habitat remnants (Garner &
McGregor 2001, p. 155). The largest
remnant habitat remaining is the Wilson
Dam tailwaters, a reach adjacent to and
downstream from Florence, Alabama.
With the exception of 1976–1978
when it was ‘‘collected infrequently’’
from below Wilson Dam (Gooch et al.
1979, p. 90), no collections of the
spectaclecase were reported at Muscle
Shoals from 1931 to 1995 despite
surveys conducted in 1956–1957, 1963–
1964, and 1991 (Garner & McGregor
2001, p. 156).
Elsewhere along the Tennessee
mainstem, a specimen was recently
reported from the Guntersville Dam
tailwaters in northern Alabama (Butler
2002a, p. 17). From 1997–1999, 10 live,
1 fresh dead, and 4 relic spectaclecase
were reported from three sites in this
river reach based on Ohio State
University Museum (OSUM) records.
The species is found only occasionally
in the lower Tennessee River below
Pickwick Landing Dam in southeastern
Tennessee, having been unreported in
various surveys (for example, Scruggs
1960, p. 12; van der Schalie 1939, p.
456). Yokley (1972, p. 61) considered it
rare, having only found fresh dead
specimens in his 3-year study. Hubbs
and Jones (2000, p. 28) reported two live
specimens found in 1998 at RM 170,
Hardin County. The current status of
these small populations is unknown
(Garner 2008, pers. comm.; Hubbs 2008,
pers. comm.).
Clinch River: The Clinch River is a
major tributary of the upper Tennessee
River in southwestern Virginia and
¨
northeastern Tennessee. Bopple and
Coker (1912, p. 9) noted numerous
spectaclecase shells in muskrat middens
in a portion of the Clinch that is now
inundated by Norris Reservoir. Ortmann
(1918, p. 527) reported the spectaclecase
as being locally abundant in the lower
Clinch River, again in an area mostly
flooded by Norris Reservoir. Oddly, he
failed to find this species upstream of
Claiborne County, yet, in later years,
one of the spectaclecase’s largest known
populations was identified in this reach.
The species was locally common at sites
in the upper Clinch River, according to
OSUM records from the 1960s. Ahlstedt
(1991a, p. 98) considered this species to
be relatively rare in the Clinch River
based on survey work conducted during
1978 to 1983. He recorded 78 live
specimens from 22 sites between RM
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151 and 223, for an average of 3.5 per
site. The spectaclecase population
reported by Ahlstedt (1991a, pp. 89–90)
from the lower Clinch River between
Melton Hill and Norris Dam (11
specimens from 4 sites between RM 45
and 73) was considered to be small but
stable. Once considered abundant in the
Clinch River at Speers Ferry, Scott
County, Virginia (Bates & Dennis 1978,
pp. 18–19), the species is now extremely
rare at this site (Neves 1991, p. 264).
Currently, the species is locally
common in the Tennessee River system
only in the upper Clinch River, and
populations are primarily restricted to
the Tennessee portion of that stream.
Despite low numbers (0.02 per sq. ft (0.2
per sq. m)) detected in quantitative
sampling (428; 2.7 sq. ft (0.25 sq. m)
quadrats) in 1994 (Ahlstedt & Tuberville
1997, pp. 73, 81), the upper Clinch
River in Tennessee may still yield two
to three dozen specimens under
individual slab boulders. Three
individuals were collected at RM 223.6
in Virginia in 2005, and one old
individual was collected in 2007 at RM
270.8, representing the farthest
upstream record for the species (Eckert
2008, pers. comm.). The upper Clinch
River population is considered to be
reproducing, with fairly young
individuals occasionally found, but
overall the population appears to be
declining (Ahlstedt 2008, pers. comm.).
The recent occurrence of a disjunct
population in the lower Clinch River
(separated from the upper Clinch River
population by Norris Reservoir) was
recently verified (Fraley 2008, pers.
comm.). The specimens sampled likely
recruited since the Norris Dam gates
closed in 1936 (Fraley 2008, pers.
comm.), despite the cold tailwaters that
destroyed the majority of the mussel
fauna in this once incredibly diverse
river reach.
Nolichucky River: The Nolichucky
River is a tributary of the lower French
Broad River, in the upper Tennessee
River system in North Carolina and
Tennessee. The spectaclecase
population in this river was once
sizable, judging from museum lots (for
example, 23 fresh dead, OSUM
1971:0372). Sampling at 41 Nolichucky
River sites in 1980, Ahlstedt (1991b, pp.
136–137) reported 8 live spectaclecase
from 6 sites between RM 11.4 to 31.9.
A small population of the spectaclecase
also persists in a relatively short reach
of the lower river (Ahlstedt 2008, pers.
comm.). The current status of the
Nolichucky River population is
unknown.
Duck River: The Duck River is wholly
in Tennessee and represents the farthest
downstream significant tributary of the
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Tennessee River, joining it in the
headwaters of Kentucky Reservoir. A
single spectaclecase, representing a new
drainage record, was found live in lower
Duck River, Hickman County, in 1999
(Hubbs 1999, p. 1; Powell 2008, pers.
comm.). Since then, at least two
individuals from the lower part of the
river in Humphreys County have been
documented, and several relic
specimens have been reported farther
upstream (Hubbs 2008, pers. comm.;
Powell 2008, pers. comm.). These
records cover an approximately 20-mile
(32-km) reach of river, with the live
individual reported from the lower end
of this reach. The spectaclecase is
considered extremely rare in the Duck
River, and its status is unknown.
Lower Mississippi River System
The spectaclecase was apparently
never widely distributed in the lower
Mississippi River system. Records from
only two streams are known, both from
Arkansas.
Mulberry River: The Mulberry River is
a tributary of the Arkansas River in
northwestern Arkansas. Other than the
Ouachita River records, the only other
record of the spectaclecase in the lower
Mississippi River system is a single
specimen found in the mid-1990s in the
Mulberry River. There is some
uncertainty regarding the validity of this
record, as the collectors were not
experienced malacologists, and no
specimen or photograph is available to
substantiate the record. This record is,
however, accepted as valid (Harris et al.
2009, p. 67; Harris 2010, pers. comm.).
The status of the spectaclecase in the
Mulberry River is unknown.
Ouachita River: The Ouachita River
flows into lower Red River, a major
western tributary of the lower
Mississippi River, draining portions of
Arkansas and Louisiana. This species
was first reported in this portion of its
range from the Ouachita River,
southwestern Arkansas, in the early
1900s (Wheeler 1918, p. 121).
Spectaclecase records in the Ouachita
span a three-county reach of river. Only
two live specimens were found in the
mid-1990s, both in the lower portion of
Ouachita County. A single relic shell
(paired valves) was found in
Montgomery County, at the upper end
of its Ouachita River range in 2000. The
population is considered very small and
declining (Harris et al. 2009, p. 67;
Harris 2010).
Summary of Extant Spectaclecase
Populations
The spectaclecase appears to be
declining rangewide, with the exception
of a few significant populations. Its
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occurrence in the St. Croix, Meramec,
Gasconade, and Clinch Rivers represent
the only sizable, sustainable, and
reproducing populations remaining,
although the Clinch River population
appears to be in decline. The
spectaclecase has been eliminated from
three-fifths of the total number of
streams from which it was historically
known (19 streams currently compared
to 44 streams historically). This species
has also been eliminated from long
reaches of former habitat in thousands
of miles of the Illinois, Ohio,
Cumberland, and other rivers, and from
long reaches of the Mississippi and
Tennessee Rivers. In addition, the
species is no longer known from the
States of Ohio, Indiana, Kansas, and
Nebraska. The extirpation of this species
from numerous streams and stream
reaches within its historical range
signifies that substantial population
losses have occurred.
Sheepnose Historical Range and
Distribution
Historically, the sheepnose occurred
in the Mississippi, Ohio, Cumberland,
and Tennessee River systems and their
tributaries, totaling at least 77 streams
(including 1 canal) (Butler 2002b). Its
distribution comprised portions of 14
States (Alabama, Illinois, Indiana, Iowa,
Kentucky, Minnesota, Mississippi,
Missouri, Ohio, Pennsylvania,
Tennessee, Virginia, West Virginia, and
Wisconsin). Historical occurrence by
stream system (with tributaries) include
the following:
• Upper Mississippi River system
(Mississippi River (Minnesota, St. Croix,
Chippewa (Flambeau River), Wisconsin,
Rock, Iowa, Des Moines, Illinois (Des
Plaines, Kankakee, Fox, Mackinaw,
Spoon, Sangamon (Salt Creek) Rivers;
Quiver Creek; Illinois and Michigan
Canal), Meramec (Bourbeuse, Big
Rivers), Kaskaskia, Saline, Castor,
Whitewater Rivers));
• Lower Missouri River system (Little
Sioux, Little Blue, Gasconade (Osage
Fork) Rivers);
• Ohio River system (Ohio River
(Allegheny (Hemlock Creek),
Monongahela, Beaver (Duck Creek),
Muskingum (Tuscarawas, Walhonding
(Mohican River), Otter Fork Licking
Rivers), Kanawha, Scioto, Little Miami,
Licking, Kentucky, Salt, Green (Barren
River), Wabash (Mississinewa, Eel,
Tippecanoe, Vermillion, Embarras,
White (East, West Forks White River)
Rivers) Rivers);
• Cumberland River system
(Cumberland River (Obey, Harpeth
Rivers; Caney Fork));
• Tennessee River system (Tennessee
River (Holston (North Fork Holston
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River), French Broad (Little Pigeon
River), Little Tennessee, Clinch (North
Fork Clinch, Powell Rivers), Hiwassee,
Duck Rivers)); and
• Lower Mississippi River system
(Hatchie, Black, Yazoo (Big Sunflower
River), Big Black Rivers).
Sheepnose Current Range and
Distribution
Extant populations of the sheepnose
are known from 24 rivers in all 14 States
of historical occurrence. Current
populations occur in the following
systems (with tributaries):
• Upper Mississippi River system
(Mississippi River (Chippewa
(Flambeau River), Wisconsin, Kankakee,
Meramec (Bourbeuse River) Rivers));
• Lower Missouri River system
(Osage Fork, Gasconade River); Ohio
River system (Ohio River (Allegheny,
Muskingum (Walhonding River),
Kanawha, Licking, Kentucky,
Tippecanoe, Eel, Green Rivers));
• Tennessee River system (Tennessee
River (Holston, Clinch, Duck (Powell
River) Rivers)); and
• Lower Mississippi River system
(Big Sunflower River).
The 24 extant sheepnose populations
occur in the following 14 States (with
streams):
• Alabama (Tennessee River),
• Illinois (Mississippi, Kankakee,
Ohio, Wabash Rivers),
• Indiana (Ohio, Tippecanoe, Eel
Rivers),
• Iowa (Mississippi River),
• Kentucky (Ohio, Licking, Kentucky,
Green Rivers),
• Minnesota (Mississippi River),
• Mississippi (Big Sunflower River),
• Missouri (Mississippi, Meramec,
Bourbeuse, Osage Fork Gasconade
Rivers),
• Ohio (Ohio, Muskingum Rivers),
• Pennsylvania (Allegheny River),
• Tennessee (Tennessee, Holston,
Clinch, Powell, Duck Rivers),
• Virginia (Clinch, Powell Rivers),
• West Virginia (Ohio, Kanawha
Rivers), and
• Wisconsin (Mississippi, St. Croix,
Chippewa, Flambeau, Wisconsin
Rivers).
The sheepnose was last observed from
over two dozen streams decades ago
(e.g., Minnesota, Rock, Iowa, Illinois,
Des Plaines, Fox, Mackinaw, Spoon,
Castor, Little Sioux, Little Blue,
Monongahela, Beaver, Scioto, Little
Miami, Salt, Mississenewa, Vermilion,
Embarras, White, Obey, Harpeth, North
Fork Holston, French Broad, North Fork
Clinch Rivers; Caney Fork). According
to Parmalee and Bogan (1998, p. 177)
and Neves (1991, pp. 280–281), the
sheepnose has been extirpated
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throughout much of its former range or
reduced to isolated populations. The
only records known from some streams
are archeological specimens (for
example, Little Pigeon, Big Black, Yazoo
Rivers; Saline River).
Sheepnose Population Estimates and
Status
The sheepnose has been eliminated
from two-thirds of the total number of
streams from which it was historically
known (24 streams currently occupied
compared to 77 streams historically)
(Table 2). This species has also been
eliminated from long reaches of former
habitat including thousands of miles of
the Mississippi, Wisconsin, Illinois,
Ohio, Cumberland, and Tennessee
Rivers and dozens of other streams and
stream reaches.
Based on the population designation
criteria (see Species Distribution
section, above), of the 24 sheepnose
populations that are considered extant,
11 are thought to be stable and 8 are
considered declining (Table 2). Five
other populations (Walhonding,
Gasconade, Muskingum, Osage Fork,
and Duck Rivers) are considered extant,
but the status of these populations is
unknown.
TABLE 2—SHEEPNOSE STATUS AT HISTORICAL LOCATIONS
River basin
Stream
Current status
Date of
last observation
Upper Mississippi River ...................
Mississippi River .............................
Minnesota River ..............................
St. Croix River ................................
Chippewa/Flambeau River .............
Wisconsin River ..............................
Rock River ......................................
Iowa River .......................................
Des Moines River ...........................
Illinois River ....................................
Des Plaines River ...........................
Kankakee River ..............................
Fox River ........................................
Mackinaw River ..............................
Spoon River ....................................
Sangamon River .............................
Salt Creek .......................................
Quiver Creek ...................................
Illinois and Michigan (I & M) Canal
Meramec River ...............................
Bourbeuse River .............................
Big River .........................................
Kaskaskia River ..............................
Saline River ....................................
Castor River ....................................
Whitewater River ............................
Little Sioux River .............................
Little Blue River ..............................
Gasconade River ............................
Osage Fork .....................................
Declining ............
Extirpated ...........
Extirpated ...........
Stable .................
Declining ............
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Stable .................
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Stable .................
Declining ............
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Unknown ............
Unknown ............
2008 .......
~1944 ....
1988 .......
~1994 ....
2002 .......
1926 .......
1925 .......
~1915 ....
1940 .......
~1970 ....
2007 .......
~1913 ....
~1970 ....
1929 .......
~1919 ....
Unknown
1881 .......
? .............
2002 .......
1997 .......
1978 .......
1970 .......
? .............
~1965 ....
1970s .....
1916 .......
~1915 ....
~1965 ....
1999 .......
Ohio River .......................................
Allegheny River ...............................
Hemlock Creek ...............................
Monongahela River .........................
Beaver River ...................................
Duck Creek .....................................
Muskingum River ............................
Tuscarawas River ...........................
Walhonding River ...........................
Mohican River .................................
Otter Fork Licking River ..................
Kanawha River ...............................
Scioto River ....................................
Little Miami River ............................
Licking River ...................................
Kentucky River ................................
Salt River ........................................
Green River ....................................
Barren River ....................................
Wabash River .................................
Mississinewa River .........................
Eel River .........................................
Tippecanoe River ............................
Vermillion River ...............................
Embarras River ...............................
White River .....................................
East White River .............................
Stable .................
Improving ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Unknown ............
Extirpated ...........
Unknown ............
Extirpated ...........
Extirpated ...........
Stable .................
Extirpated ...........
Extirpated ...........
Declining ............
Declining ............
Extirpated ...........
Improving ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Declining ............
Stable .................
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
2007 .......
2008 .......
Unknown
~1897 ....
~1910 ....
1930 .......
1993 .......
Unknown
1993 .......
1977 .......
1973 .......
2005 .......
1963 .......
~1953 ....
1998 .......
1996 .......
~1900 ....
2007 .......
Unknown
1988 .......
1899 .......
1997 .......
1995 .......
Unknown
1953 .......
1913 .......
1969 .......
Lower Missouri River .......................
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Ohio River ........................................
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Comments
Relic shell collected in 1999.
Relic shell collected in 1999.
Relic shell collected in 1989.
Relic shell collected in 1989.
Represented by single specimen,
presumably near extirpation.
Relic shell collected in 1991.
Relic shell collected in 1998.
Relic shell collected in 1993.
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3403
TABLE 2—SHEEPNOSE STATUS AT HISTORICAL LOCATIONS—Continued
Stream
Current status
Date of
last observation
West Fork White River ....................
Cumberland River ...........................
Obey River ......................................
Harpeth River ..................................
Caney Fork River ............................
Tennessee River .............................
Holston River ..................................
North Fork Holston River ................
French Broad River ........................
Little Pigeon River ..........................
Little Tennessee River ....................
Clinch River ....................................
North Fork Clinch River ..................
Powell River ....................................
Hiwassee ........................................
Duck River ......................................
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Stable .................
Declining ............
Extirpated ...........
Extirpated ...........
Extirpated ...........
Extirpated ...........
Stable .................
Extirpated ...........
Stable .................
Extirpated ...........
Unknown ............
1908 .......
1987 .......
1939 .......
? .............
Unknown
2004 .......
2007 .......
1913 .......
1914 .......
Unknown
Unknown
2006 .......
~1921 ....
2004 .......
Unknown
2003 .......
Hatchie River ..................................
Black River ......................................
Yazoo River ....................................
Big Sunflower River ........................
Big Black River ...............................
Extirpated ...........
Extirpated ...........
Extirpated ...........
Declining .............
Extirpated ...........
1983 .......
Unknown
Unknown
2000 .......
Unknown
River basin
Cumberland River ............................
Tennessee River ..............................
EMCDONALD on DSK2BSOYB1PROD with MISCELLANEOUS
Lower Mississippi River ...................
Historically, the sheepnose was fairly
widespread in many Mississippi River
system streams, although rarely
common. Archaeological evidence on
relative abundance indicates that it has
been an uncommon or even rare species
in many streams for centuries (Morrison
1942, p. 357; Patch 1976, pp. 44–52;
Parmalee et al. 1980, p. 101; Parmalee
et al. 1982, p. 82; Parmalee and Bogan
1986, pp. 28, 30; Parmalee and Hughes
1994, pp. 25–26), and relatively
common in only a few (Bogan 1990, p.
135).
Museum collections of this species
are almost always few in number
(Cummings 2010, pers. comm.), with the
exception of the 1960s collections from
the Clinch and Powell Rivers,
Tennessee and Virginia. Moderate
numbers of individuals were also
commonly recorded historically from
the upper Muskingum River system in
Ohio and the lower Wabash River in
Indiana and Ohio, based on museum
lots. Schuster and Williams (1989, p. 21)
reported the species as being not
common in the Ohio River, while
Cummings and Mayer (1992, p. 50)
considered it rare throughout its range.
The American Malacological Union
considers the sheepnose to be
threatened (Williams et al. 1993, p. 13).
Some known populations of the
sheepnose are represented by the
collection of a single specimen. Other
populations have seen a dramatic range
decline (for example, reduced from
several hundred river miles to a single
bed of a river system) or we have
limited recent information on
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population status. The following
summaries focus primarily on those
populations for which we have
sufficient information to make status
and trend determinations, and less on
those populations that are nearly
extirpated, have no recruitment, or are
of unknown status.
Upper Mississippi River System
Judging from the archeological record,
the sheepnose may have been common
at some sites on the Mississippi River
(Bogan 1990, p. 135) but over the past
century it has become a rare species
throughout the mainstem (Grier 1922,
pp. 13–31; van der Schalie and van der
Schalie 1950, pp. 454–457). Robust
populations may have been found in
some tributary rivers. The sheepnose
has been extirpated from eight
Mississippi River tributaries
(Minnesota, Rock, Iowa, Des Moines,
Kaskaskia, Saline, Castor, and
Whitewater Rivers) and all but one
Illinois River tributary (the Kankakee
River). Today, the sheepnose is extant
(though in low numbers) in ten
mainstem pools, and six tributary rivers
of the Upper Mississippi River System.
Mississippi River mainstem:
Sheepnose populations in the mainstem
of the Upper Mississippi River are
declining. Despite the discovery of a
juvenile in Mississippi River Pool
(MRP) 7 in 2001, recruitment is limited
at best. The mainstem population is
comprised of a few old individuals
spread across a very large geographic
range (MRP 3 through MRP 24 a
distance of over 550 river miles (880
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Relic shell collected in 2000.
Relic shell collected in 1990.
Relic shell collected in 1971.
Relic shell collected in 1975.
Record represented by single specimen.
river km)) (Thiel 1981, p. 10; Havlik and
Marking 1981, p. 32; Whitney et al.
1996, p. 17; Helms and Associates,
Ecological Specialists Inc. 2008, p. 16).
The status of this species in the
Mississippi River is highly jeopardized
(Butler 2002b, p. 7).
Pools with extant populations include
MRP 3 (last seen live or fresh dead in
2000–01), MRP 4 (2008), MRP 7 (2001),
MRP 11 (2007), MRP 14 (2006–07), MRP
15 (2005–06), MRP 16 (2003), MRP 17
(2004), MRP 20 (1992), and MRP 24
(1999). The 2001 MRP 7 record was for
a live juvenile 1.3 inches (3.3 cm) long
and estimated to be 3 years old (Davis
2008, pers. comm.).
St. Croix River: The St. Croix River
population is isolated and comprised of
old individuals with little to no
recruitment (Heath 2010b, pers. comm.).
Currently, the population is thought to
be restricted to the lowermost mainstem
below RM 1 in Washington County,
Minnesota, and Pierce County,
Wisconsin (Heath 2010b, pers. comm.).
Three live individuals were collected in
1988, during a mussel relocation project
for the U.S. Highway 10 bridge
immediately upstream of the confluence
with the Mississippi River (Heath 1989,
p. 16). Hornbach (2001, p. 218) analyzed
mussel collections throughout the St.
Croix River and found that the
sheepnose was absent in 15 of the 16
river reaches he sampled, only noting
the 1988 occurrence. One historical
occurrence is known from the vicinity
of RM 53 in 1930; however, this is the
only known record upstream of RM 1
(Heath 2010b, pers. comm.). Because
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there have been no recent collections in
the St. Croix River since 1988, this
population is most likely extirpated.
Chippewa/Flambeau River: The
sheepnose population in the Chippewa
River is extant in much of the river
system including the lower end of its
tributary, the Flambeau River. This
population is stable with documented
recruitment (Butler 2002b, p. 8). Balding
and Balding (1996, p. 5) reported 50 live
specimens sampled from 1989–1994,
but more recent collections have
expanded sites of occurrence to 20 of 67
sites in the middle and upper portions
of the Chippewa River, with a relative
abundance of 0.8 percent (Balding 2001,
pers. comm.). Balding (1992, p. 166)
found 12 live specimens and 31 dead
shells from 5 of 37 sites in the lower
river. Additional survey work extended
the number of sites where it was found
live to 10 of 45 (Balding 2001, pers.
comm.). The Flambeau River supports a
small sheepnose population below its
lowest dam and near its confluence with
the Chippewa River (lower 8 miles (13
km) of river), and is most likely
dependent on the source population in
the Chippewa River. The Chippewa
River sheepnose population is
considered one of the best known extant
populations. The Flambeau River
supports a small sheepnose population
below its lowest dam and near its
confluence with the Chippewa River
(lower 8 miles (13 km) of river), and is
most likely dependent on the source
population in the Chippewa River.
Wisconsin River: The sheepnose is
declining in the Wisconsin River.
Historical records for the sheepnose are
available throughout the lower 335
miles (539 km) of the 420-mile (676-km)
Wisconsin River (Heath 2010c, pers.
comm.). In July 2002, 20 live specimens
were found in a dense mussel bed near
Port Andrew (Seitman 2008, pers.
comm.). Currently, the sheepnose is
primarily confined to RM 133.7
downstream (a reduction of over 201
river miles (232 km)). The sheepnose
population is probably recruiting in the
river, primarily in the lower section
(below RM 82) (Heath 2010b, pers.
comm.). It is unknown if the middle
river population, from RM 93 to 133.7,
is recruiting because only three living
individuals have been found in recent
years (Heath 2010c, pers. comm.).
Kankakee River: The sheepnose once
occurred along the lower two-thirds of
the Kankakee River, an Upper Illinois
River tributary, in Indiana and Illinois
(Wilson and Clark 1912, p. 47; Lewis
and Brice 1980, p. 4). The sheepnose
has been extirpated from the
channelized portion of the Kankakee in
Indiana but persists in the Illinois
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portion of the river where it appears
stable, with evidence of recent
recruitment (Butler 2002b, p. 9).
Records since 1986 identify the
sheepnose in the Kankakee River from
the Iroquois River confluence
downstream approximately 30 river
miles (48 km) (Cummings 2010, pers.
comm.; Helms and Associates 2005, p.
3). A mussel relocation effort for a
pipeline crossing in the Kankakee River
in July 2002 found 11 sheepnose
individuals, representing 0.32 percent of
the total mussels relocated (Helms 2004,
p. D–1). Subsequent monitoring of the
site in 2004 and 2007 located four new
individuals. One individual collected in
2004 measured 1.6 inches (40 mm) and
was estimated to be a juvenile of 3 years
of age.
Meramec River: The Meramec River
flows into the Mississippi River
downstream of St. Louis and drains
east-central Missouri. The Meramec
sheepnose population is stable and
recruiting, and represents one of the
best rangewide (Butler 2002b, p. 9). Two
studies (Buchanan 1980, p. 4; Roberts &
Bruenderman 2000, p. 20) extensively
surveyed the mussel fauna of the
Meramec River. The most notable
difference in the results of these studies
was the reduced range in which
sheepnose were found. Buchanan (1980,
p. 34) found live or fresh dead
individuals from RM 4.5 to 145.7 (141.2
river miles (227.2 km)), whereas Roberts
and Bruenderman (2000, p. 20) found
live or fresh dead individuals from RM
25.6 to 91.3 (65.7 river miles (105.7
km)). The trend data from the late 1970s
to 1997 indicate that the sheepnose
declined 75.5 river miles (121.5 km) in
total range within the Meramec River.
The extent of the population in the
lower end appears to be shrinking
upriver (Butler 2002b, p. 10).
In 2002, a site associated with a
railroad crossing in St. Louis County at
RM 28 yielded 43 live specimens over
3 days of sampling, including at least
one gravid female (Roberts 2008, pers.
comm.). Collectively, these data
reinforce the level of importance of the
Meramec population for the sheepnose
rangewide. Although the existing
population has been described as stable
and recruitment has been documented
in the system (Butler 2002b, pp. 11–12),
the population has shrunk by half of its
former geographic range over the past 30
years.
Bourbeuse River: The Bourbeuse River
sheepnose population is distributed in
the downstream 90 river miles (145 km)
of the river (Buchanan 1980, p. 34), but
is considered rare. Although
recruitment has been documented in the
Bourbeuse River, the sheepnose
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population is considered declining
(Roberts and Bruenderman 2000, p. 130;
Roberts 2010, pers. comm.). In the late
1970s, Buchanan (1980, p. 10) found the
sheepnose to represent 0.1 percent of
the Bourbeuse River mussel fauna, with
10 live specimens sampled from 7 sites.
Based on data collected by Buchanan
(1980, p. 34) and additional survey work
in 1980, live or fresh-dead individuals
were found in the Bourbeuse from RM
6.5 to 90.0. Data from a resurvey of the
Bourbeuse River collected in 1997
yielded nine live sheepnose from four
sites (Roberts and Bruenderman 2000, p.
39) and fresh dead shells were located
at an additional site. Sheepnose relative
abundance was 0.4 percent. Live or
fresh dead sheepnose were found
between RM 1.4 to 66.3. This
comparison indicates a decrease in the
number of extant sites (7 to 4) and a
range contraction of 18 river miles (29
km). The sheepnose in the Meramec and
Bourbeuse Rivers represents a
population cluster.
Lower Missouri River System
Osage Fork Gasconade River: The
Lower Missouri River system
population is represented by a single
sheepnose specimen and is near
extirpation. This individual was located
in 1999 at RM 21.2 in the Osage Fork,
a tributary to the Gasconade River
(Bruenderman et al. 2001, p. 14). It is
the only known record for sheepnose in
the Gasconade River drainage for over
25 years.
Ohio River System
Historically, the sheepnose was
documented from the entire length of
the Ohio River (its type locality), and
was first collected there in the early
1800s. Ohio River sampling of 664 river
miles (1,068 km) along the northern
border of Kentucky yielded 41
sheepnose (Williams 1969, p. 58). Most
of these (29) were found in the upper
portions of river (from RM 317 to 538),
but the population extended
downstream to RM 871. Relative
abundance was 0.7 percent for the entire
reach sampled. Currently, the mainstem
Ohio River and 10 tributary streams
have extant sheepnose populations.
Ohio River mainstem: The sheepnose
is generally distributed, but rare, in
most mainstem pools of the Ohio River.
The population appears to be more
abundant in the lower section of the
river with a smaller population in the
upper Ohio River pools (McGregor 2008,
pers. comm.; Schuster and Williams
1989, p. 24; Zeto et al. 1987, p. 184).
Long term monitoring data from 1993 to
2007 at RM 176 shows the sheepnose is
usually collected each survey,
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recruitment is occurring, and the
species comprises 1.0 percent of the
mussels at the site (relative abundance)
(Morrison 2008, pers. comm.). Live
sheepnose have also been collected in
recent years at RM 725 and RM 300
(Morrison 2008, pers. comm.). The
population in the lower Ohio River
mainstem is viable with documented
recruitment, but the population overall
continues to show signs of decline
(Butler 2002b, p. 12).
Allegheny River: The Allegheny River
drains northwestern Pennsylvania and
western New York and joins the
Monongahela River at Pittsburgh to form
the Ohio River. A recruiting and
improving population of sheepnose
exists within the middle Allegheny
River (Villella 2008, pers. comm.).
Sampling efforts from 2006–2008 at 63
sites over 78 miles (125 km) of river
produced sheepnose at 18 sites. A total
of 244 individuals of 7 different age
classes were collected (Villella 2008,
pers. comm.) providing ample evidence
of recent recruitment.
Kanawha River: The Kanawha River is
a major southern tributary of the Ohio
River draining much of West Virginia
and with headwaters in Virginia and
North Carolina. The Kanawha River
harbors a small, but recruiting and
stable, population of sheepnose in
Fayette County, West Virginia (Butler
2002b, p. 14). The Kanawha population
appears to be limited to 5 river miles (8
km) immediately below Kanawha Falls
(Clayton 2008c, pers. comm.).
Sheepnose collections from this reach in
1987 resulted in a density of 0.013 per
sq. m (0.140 per sq. ft), and collections
from 2005 found a density of 0.016 per
sq. m (0.172 per sq. ft) (Clayton 2008c,
pers. comm.).
Licking River: The sheepnose is
known from the lower half of the
Licking River, a southern tributary of
the Ohio River in northeastern
Kentucky. Currently, the species is
known from roughly five sites in the
middle Licking River (McGregor 2008,
pers. comm.). There is no documented
evidence of recent recruitment, and,
therefore, the sustainability of the
population is unknown. It is possible
this population represents a population
cluster with the Ohio River.
Green River: The Green River is a
lower Ohio River tributary in westcentral Kentucky. Currently, a recruiting
and improving population remains over
an approximately 25 river mile (40 river
km) reach in the upper Green River from
the vicinity of Mammoth Cave National
Park upstream into Hart County (Butler
2002b, p. 15). An investigation of
muskrat middens from 2002 and 2003
revealed 42 sheepnose shells, with 39 of
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the 42 between 1.2 and 2.2 inches (3.0
and 5.6 cm) in length and described as
juveniles (Layzer 2008, pers. comm.).
Sampling over the past several years
(2005–2007) has documented a number
of beds experiencing recruitment
(McGregor 2008, pers. comm.).
Tippecanoe River: The Tippecanoe
River drains the central portion of
northern Indiana in the upper Wabash
River system. This population of
sheepnose is considered stable with
relatively recent recruitment (Butler
2002b, p. 17). Survey work between
1987 and 1995 documented sheepnose
at 14 sites throughout the river and
extended the known range of the species
upstream into Marshall County (Butler
2002b, p. 17). The sheepnose is now
known from 45 miles (72 km) of the
Tippecanoe River (Ecological
Specialists, Inc. 1993, pp. 80–81;
Cummings and Berlocher 1990, pp. 84,
98; Cummings 2008, pers. comm.;
Fisher 2008, pers. comm.).
Kentucky, Wabash, Eel, Muskingum,
and Walhonding Rivers: In addition to
the aforementioned populations,
sheepnose in the Ohio River system are
known from the Kentucky, Wabash, and
Eel Rivers, which are each represented
by two or fewer specimens collected in
the past 25 years. Populations of the
sheepnose in these three river systems
are considered to be declining and may
be nearing extirpation (Butler 2002b, p.
15–16). A population cluster in two
additional rivers, the Muskingum River
and its tributary, the Walhonding River,
have unknown populations. Although
Watters and Dunn (1995, p. 240)
documented recruitment in the lower
Muskingum River in the mid-1980s, the
sheepnose population in the river is
extremely small, and distribution has
been reduced to only the lower portion
of the river where six individuals were
collected in 1992 (Watters and Dunn
1995, pp. 253–254).
Cumberland River System
Historical sheepnose records in the
system are known from throughout the
mainstem downstream of Cumberland
Falls and three of its tributaries (Obey,
Harpeth, and Caney Fork Rivers).
Wilson and Clark (1914, pp. 15–19, 57)
reported the species to be generally
uncommon from 14 mainstem sites from
what is now Cumberland Reservoir,
Kentucky, downstream to Stewart
County, Tennessee, a distance of nearly
500 miles (∼805 km). Sheepnose was
last documented in the Tennessee
portion of the river during the early
1980s (Butler 2002b, p. 67).
The only recent record for the
Cumberland River is from 1987, at the
extreme lower end of the river near its
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confluence with the Ohio River, below
Barkley Dam (Butler 2002b, p. 18). This
population may be influenced by the
lower Ohio River sheepnose population
(Butler 2002b, p. 18) and represents a
population cluster. Surveys conducted
in 2007–2009 found no sheepnose
(Hubbs, 2010, pers. comm.) and so this
population may be extirpated.
Tennessee River System
The sheepnose was originally known
from the Tennessee River and 10 of its
tributary streams. Historically, Ortmann
(1925, p. 328) considered the sheepnose
to occur ‘‘sparingly’’ in the lower
Tennessee River, and to be ‘‘rare’’ in the
upper part of the system (Ortmann 1918,
p. 545). Hundreds of miles of large river
habitat on the Tennessee River
mainstem have been converted under
nine reservoirs, with additional dams
constructed in tributaries historically
harboring the sheepnose (for example,
Clinch, Holston, Little Tennessee,
Hiwassee Rivers) (Tennessee Valley
Authority 1971, p. 5). Sheepnose
populations currently persist in limited
reaches of the Tennessee River
mainstem and four tributaries.
Tennessee River mainstem: The 53mile (85-km) stretch of river in
northwestern Alabama referred to as the
Muscle Shoals, historically harbored 69
species of mussels, making it the most
diverse mussel fauna ever known
(Garner and McGregor 2001, pp. 155–
157). However, with the construction of
three dams (Wilson in 1925, Wheeler in
1930, and Pickwick Landing in 1940)
most of the historical habitat was
inundated, leaving only small, flowing
habitat remnants (Garner and McGregor
2001, p. 158).
The species is found only
occasionally in the lower Tennessee
River below Pickwick Landing Dam in
southwestern Tennessee. Scruggs (1960,
p. 11) recorded a relative abundance of
0.2 percent, while Yokley (1972, p. 64)
considered it to be ‘‘very rare’’ in this
reach (relative abundance of 0.1
percent). Yokley reported only two
specimens that were each estimated to
be 20 or more years old.
The sheepnose persists in the
tailwaters of Guntersville, Wilson,
Pickwick Landing, and Kentucky Dams
on the mainstem Tennessee River,
where it is considered uncommon
(Garner & McGregor 2001, p. 165; Gooch
et al. 1979, p. 9). These populations are
considered stable overall but with very
limited recruitment (Garner and
McGregor 2001, p. 165; McGregor 2008,
pers. comm.). The species has been
found in low numbers over the past 80
years from relic habitat in the Wilson
Dam tailwaters, a several-mile reach
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adjacent to and downstream from
Florence, Alabama (Butler 2002b, pp.
20–21).
Clinch River: The Clinch River in
southwestern Virginia and northeastern
Tennessee is one of the largest and most
significant tributaries of the upper
Tennessee River system. Based on
archeological evidence, the sheepnose
was ‘‘extremely rare’’ in the lower Clinch
River (Parmalee and Bogan 1986, p. 28).
As of 2002, the largest lots of museum
material available for the sheepnose had
been from the Clinch River and its
tributary, the Powell River (Watters
2010, pers. comm.). Individual Clinch
River museum lots collected during
1963 to 1969 include 36, 39, 70, and 82
fresh dead specimens. The sheepnose
population in the Clinch River currently
occurs over approximately 60 river
miles (96 km) from northern Scott
County, Virginia downstream into
Hancock County, Tennessee, and is
considered stable with recently
documented recruitment (Eckert 2008,
pers. comm.). Survey work between
1979 and 1994 (Ahlstedt & Tuberville
1997, p. 73) reported low densities of
0.009 to 0.018 individuals per sq. ft. (0.1
to 0.2 per sq. m). Sampling efforts in
2005 and 2006 reported densities from
two sites (RM 223.6 and 213.2) in Scott
County, Virginia, of 0.021 and 0.006
individuals per sq. m (0.226 and 0.064
per sq. ft), respectively (Eckert 2008,
pers. comm.). Relative abundance for
sheepnose at these locations was 1.5
percent and 1.0 percent, respectively.
Powell River: The largest sheepnose
collection (OSUM) known rangewide
was collected in the Powell River, the
Clinch River’s largest tributary, and
included 6 live and 141 fresh dead
specimens. Today, the sheepnose
population in the Powell River is
considered stable, and recruitment has
been documented. In 1979, Ahlstedt
(1991b, pp. 129–130) reported 45 live
specimens from 17 of 78 sites (an
average 2.6 individuals per site).
Ahlstedt and Tuberville (1997, p. 96)
conducted quantitative sampling in the
Powell between 1979 and 1994, and
found the sheepnose at densities of 0.01
to 0.08 per sq. m (0.107 and 0.861 per
sq. ft). Sampling efforts in 2004 reported
densities from two sites in Lee County,
Virginia (RM 120.3 and 117.3), of 0.012
and 0.017 individuals per sq. m (0.129
and 0.183 per sq. ft), respectively
(Eckert 2008, pers. comm.). Relative
abundance for sheepnose was 0.82
percent and 0.99 percent, respectively.
Duck River: The Duck River
population is recently represented by
the collection of single 10+ year old
animal in 2003. The sheepnose was
likely always rare in the Duck River
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and, previous to 2003, the species was
thought to be extirpated. The current
status of the population is unknown.
Holston River: In July 2002, sampling
in Holston River produced live
sheepnose at 16 of 20 sites sampled
below the Cherokee Dam. This reach
extended from Nance Ferry to Monday
Island (RM 14.6), Jefferson and Knox
Counties (Fraley 2008, pers. comm.). A
total of 206 specimens were found with
an overall relative abundance of 18.2
percent among the 18 species reported
live from this reach. The collection was
comprised of extremely old individuals
with no recently recruited individuals
being found. Although the population
appeared significant in numbers, the
lack of recruitment in this population is
indicative of a remnant population on
its way to extirpation (Butler 2002b, p.
19). In 2007, Tennessee Valley
Authority biologists located sheepnose
in the Holston River while conducting
fish surveys; however, no additional
mussel survey work has been completed
in the area since 2002 (Baxter 2010,
pers. comm.).
Lower Mississippi River System
The sheepnose was apparently never
widely distributed in the lower
Mississippi River system. The only
verified records are for Hatchie River in
Tennessee and the Delta region in
Mississippi. The only records for the
Yazoo and Big Black Rivers are from
archeological sites (Butler 2002b, p. 21).
The sheepnose population in the Big
Sunflower River, Mississippi, is the
only one remaining in the lower
Mississippi River system. Once
abundant judging from museum and
archeological records, there is now only
a small declining population in the Big
Sunflower River (Jones 2008, pers.
comm.). The population is believed to
be limited to a 12- to 15-mile (19- to 24km) reach upstream of Indianola in
Sunflower County, Mississippi.
Although no juvenile mussels have been
found in recent sampling efforts,
variably-sized individuals indicate
some, possibly very low, level of
recruitment in the population (Jones
2008, pers. comm.).
Summary of Extant Sheepnose
Populations
The sheepnose has experienced a
significant reduction in range, and many
of the extant populations are disjunct,
isolated, and appear to be declining.
The extirpation of this species from over
50 streams (more than 65 percent)
within its historical range indicates that
substantial population losses have
occurred. In the majority of streams
with extant populations, the sheepnose
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appears to be uncommon at best. Only
in the Allegheny and Green Rivers is the
species considered to be improving in
population status. Several other extant
populations are thought to exhibit some
level of stability and have experienced
relatively recent recruitment
(Chippewa/Flambeau, Meramec, Ohio,
Tippecanoe, Clinch, and Powell Rivers).
Given the compilation of current
distribution, abundance, and status
trend information, the sheepnose
appears to exhibit a high level of
imperilment.
Previous Federal Actions
We identified the spectaclecase and
sheepnose as candidate species in a
notice of review published in the
Federal Register on May 4, 2004 (69 FR
24875). The spectaclecase and
sheepnose remained candidate species
in subsequent notices, including: May
11, 2005 (70 FR 24869), September 12,
2006 (71 FR 53755), December 6, 2007
(72 FR 69033), December 10, 2008 (73
FR 75176), and November 9, 2009 (74
FR 57804). A candidate species is a
species for which we have on file
sufficient information on biological
vulnerability and threats to support
issuing a proposed rule to list the
species as endangered or threatened.
Summary of Factors Affecting the
Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
CFR part 424, set forth the procedures
for adding species to the Federal lists of
Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may determine a species to be
endangered or threatened due to one or
more of the following five factors: (A)
The present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination. Each of these factors is
discussed below.
A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range.
The decline of mussels such as the
spectaclecase and sheepnose is
primarily the result of habitat loss and
degradation (Neves 1991, pp. 252, 265).
Chief among the causes of decline are
impoundments, channelization,
chemical contaminants, mining, oil and
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gas development, and sedimentation
(Neves 1991, pp. 252, 260–261; Neves
1993, pp. 1–7; Neves et al. 1997, pp. 63–
72; Strayer et al. 2004, pp. 435–437;
Watters 2000, pp. 261–268; Williams et
al. 1993, p. 7). These threats to mussels
in general (and spectaclecase and
sheepnose where specifically known)
are individually discussed below.
Dams and Impoundments
Dams eliminate or reduce river flow
within impounded areas, trap silts and
cause sediment deposition, alter water
temperature and dissolved oxygen
levels, change downstream water flow
and quality, decrease habitat
heterogeneity, affect normal flood
patterns, and block upstream and
downstream movement of species
(Layzer et al. 1993, pp. 68–69; Neves et
al. 1997, pp. 63–64; Watters 2000, pp.
261–264). Within impounded waters,
decline of freshwater mollusks has been
attributed to sedimentation, decreased
dissolved oxygen, and alteration in
resident fish populations (Neves et al.
1997, pp. 63–64; Pringle et al. 2009, pp.
810–815; Watters 2000, pp. 261–264).
Dams significantly alter downstream
water quality and habitats (Allen &
Flecker 1993, p. 36), and negatively
affect tailwater mussel populations
(Layzer et al. 1993, p. 69; Neves et al.
1997, p. 63; Watters 2000, pp. 265–266).
Below dams, mussel declines are
associated with changes and fluctuation
in flow regime, scouring and erosion,
reduced dissolved oxygen levels and
water temperatures, and changes in
resident fish assemblages (Layzer et al.
1993, p. 69; Neves et al. 1997, pp. 63–
64; Pringle et al. 2009, pp. 810–815;
Watters 2000, pp. 265–266; Williams et
al. 1992, p. 7). The decline and
imperilment of freshwater mussels in
several tributaries within the Tennessee,
Cumberland, Mississippi, Missouri, and
Ohio River basins have been directly
attributed to construction of numerous
impoundments in those river systems
(Hanlon et al. 2009, pp. 11–12; Layzer
et al. 1993, pp. 68–69; Miller et al. 1984,
p. 109; Neves et al. 1997, pp. 63–64;
Sickel et al. 2007, pp. 71–78; Suloway
1981, pp. 237–238; Watters 2000, pp.
262–263; Watters & Flaute 2010, pp. 3–
7; Williams & Schuster 1989, pp. 7–10).
Population losses due to
impoundments have likely contributed
more to the decline and imperilment of
the spectaclecase and the sheepnose
than any other factor. Large river habitat
throughout nearly all of the range of
both species has been impounded,
leaving generally short, isolated patches
of vestigial habitat in the area below
dams. Navigational locks and dams, (for
example, on the upper Mississippi,
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Ohio, Allegheny, Muskingum,
Kentucky, Green, and Barren Rivers),
some high-wall dams (for example, on
the Wisconsin, Kaskaskia, Walhonding,
and Tippecanoe Rivers), and many lowhead dams (for example, on the St.
Croix, Chippewa, Flambeau, Wisconsin,
Kankakee, and Bourbeuse Rivers) have
contributed significantly to the loss of
sheepnose and spectaclecase habitat
(Butler 2002a, pp. 11–20 2002b, pp. 9–
25).
The majority of the Tennessee and
Cumberland River main stems and
many of their largest tributaries are now
impounded. There are 36 major dams
located in the Tennessee River system
and about 90 percent of the Cumberland
River downstream of Cumberland Falls
(RM 550 (RKM 886)) is either directly
impounded by U.S. Army Corps of
Engineers (Corps) structures or
otherwise impacted by cold tail water
released from several dams. Major Corps
impoundments on Cumberland River
tributaries (for example, Stones River
and Caney Fork) have inundated an
additional 100 miles (161 km) or more
of spectaclecase and sheepnose habitat.
Coldwater releases from Wolf Creek,
Dale Hollow (Obey River), and Center
Hill (Caney Fork) Dams continue to
degrade spectaclecase and sheepnose
habitat in the Cumberland River system.
Layzer et al. (1993, p. 68) reported that
37 of the 60 pre-impoundment mussel
species of the Caney Fork River have
been extirpated. Watters (2000, pp. 262–
263) summarizes the tremendous loss of
mussel species from various portions of
the Tennessee and Cumberland River
systems. Approximately one-third of the
historical sheepnose and spectaclecase
streams are in the Tennessee and
Cumberland River systems.
Navigational improvements on the
Ohio River began in 1830, and now
include 21 lock and dam structures
stretching from Pittsburgh,
Pennsylvania, to Olmsted, Illinois, near
its confluence with the Mississippi
River. Historically, habitat now under
navigational pools once supported up to
50 species of mussels, including the
spectaclecase and sheepnose.
Tributaries to the Ohio River, such as
the Green and Allegheny rivers, were
also altered by impoundments. A series
of six locks and dams was constructed
on the lower half of the Green River
decades ago and extend upstream to the
western boundary of Mammoth Cave
National Park (MCNP). The upper two
locks and dams destroyed spectaclecase
habitat, particularly Lock and Dam 6,
which flooded the central and western
portions of MCNP. Approximately 30
river miles (48 km) of mainstem habitat
were also eliminated with the
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construction of the Green River Dam in
1969. Locks and dams were also
constructed on the lower reaches of the
Allegheny, Kanawha, Muskingum, and
Kentucky Rivers which disrupted
historical riverine habitat for the
sheepnose.
Similarly, dams impound most of the
upper Mississippi River and many of its
tributaries. A series of 29 locks and
dams constructed since the 1930s in the
mainstem resulted in profound changes
to the nature of the river, primarily
replacing a free-flowing alluvial (flood
plain) system with a stepped gradient
(higher pool area to riffle area ratio)
river. Modifications fragmented the
mussel beds where spectaclecase and
sheepnose were found in the
Mississippi River, reduced stable
riverine habitat, and disrupted fish host
migration and habitat use.
Dams and impoundments have
fragmented and altered stream habitats
throughout the Sac River Basin in the
lower Missouri River system. Stockton
Dam impounds 39 miles (63 km) of the
upper Sac River and the Truman Dam
inundates about 8 miles (13 km) of the
lower Sac River and its tributaries
(Hutson & Barnhart 2004, p. 7). The
rarity of live spectaclecase in the Sac
River, coupled with the large number of
dead shells observed in a recent study,
suggests that this species has decreased
since the river was impounded, and that
spectaclecase may soon be extirpated
from the Sac River system (Hutson &
Barnhart 2004, p. 17).
Dam construction has a secondary
effect of fragmenting the ranges of
aquatic mollusk species, leaving relict
habitats and populations isolated by the
structures as well as by extensive areas
of deep uninhabitable, impounded
waters. These isolated populations are
unable to naturally recolonize suitable
habitat that is impacted by temporary,
but devastating events, such as severe
drought, chemical spills, or
unauthorized discharges (Cope et al.
1997, pp. 235–237; Layzer et al. 1993,
pp. 68–69; Miller & Payne 2001, pp. 14–
15; Neves et al. 1997, pp. 63–75; Pringle
et al. 2009, pp. 810–815; Watters 2000,
pp. 264–265, 268; Watters & Flaute
2010, pp. 3–7).
Sedimentation
Nonpoint source pollution from land
surface runoff originates from virtually
all land use activities and includes
sediments; fertilizer, herbicide, and
pesticide residues; animal or human
wastes; septic tank leakage and gray
water discharge; and oils and greases.
Nonpoint source pollution can cause
excess sedimentation, nutrification,
decreased dissolved oxygen
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concentration, increased acidity and
conductivity, and other changes in
water chemistry that can negatively
impact freshwater mussels. Land use
types around the sheepnose and
spectaclecase populations include
pastures, row crops, timber, and urban
and rural communities.
Excessive sediments are believed to
impact riverine mollusks requiring
clean, stable streams (Brim Box & Mosa
1999, p. 99; Ellis 1936, pp. 39–40).
Impacts resulting from sediments have
been noted for many components of
aquatic communities. For example,
sediments have been shown to affect
respiration, growth, reproductive
success, and behavior of freshwater
mussels, and to affect fish growth,
survival, and reproduction (Waters
1995, pp. 173–175). Potential sediment
sources within a watershed include
virtually all activities that disturb the
land surface, and most localities
currently occupied by the spectaclecase
and sheepnose are affected to varying
degrees by sedimentation.
Sedimentation has been implicated in
the decline of mussel populations
nationwide, and is a threat to
spectaclecase and sheepnose (Brim Box
& Mosa 1999, p. 99; Dennis 1984, p. 212;
Ellis 1936, pp. 39–40; Fraley & Ahlstedt
2000, pp. 193–194; Poole & Downing
2004, pp. 119–122; Vannote & Minshall
1982, pp. 4105–4106). Specific
biological impacts include reduced
feeding and respiratory efficiency from
clogged gills, disrupted metabolic
processes, reduced growth rates, limited
burrowing activity, physical smothering,
and disrupted host fish attractant
mechanisms (Ellis 1936, pp. 39–40;
Hartfield & Hartfield 1996, p. 373;
Marking & Bills 1979, p. 210; Vannote
& Minshall 1982, pp. 4105–4106; Waters
1995, pp. 173–175). In addition, mussels
may be indirectly affected if high
turbidity levels significantly reduce the
amount of light available for
photosynthesis and thus the production
of certain food items (Kanehl & Lyons
1992, p. 7).
Studies indicate that the primary
impacts of excess sediment on mussels
are sublethal, with detrimental effects
not immediately apparent (Brim Box &
Mosa 1999, p. 101). The physical effects
of sediment on mussels are multifold,
and include changes in suspended and
bed material load; changes in bed
sediment composition associated with
increased sediment production and runoff in the watershed; changes in the
form, position, and stability of channels;
changes in depth or the width-to-depth
ratio, which affects light penetration
and flow regime; actively aggrading
(filling) or degrading (scouring)
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channels; and changes in channel
position that may leave mussels
stranded (Brim Box & Mosa 1999, pp.
109–112; Kanehl & Lyons 1992, pp. 4–
5; Vannote & Minshall 1982, p. 4106).
The Chippewa River in Wisconsin, for
example, has a tremendous bedload
composed primarily of sand that
requires dredging to maintain barge
traffic on the mainstem Mississippi
below its confluence (Thiel 1981, p. 20).
The mussel diversity in the Mississippi
River below the confluence with the
Chippewa River has predictably
declined from historical times. Lake
Pepin, a once natural lake formed in the
upper Mississippi River upstream from
the mouth of the Chippewa River, has
become increasingly silted in over the
past century, reducing habitat for the
spectaclecase and sheepnose (Thiel
1981, p. 20).
Increased sedimentation and siltation
may explain in part why spectaclecase
and sheepnose mussels appear to be
experiencing recruitment failure in
some streams. Interstitial spaces in the
substrate provide crucial habitat for
juvenile mussels. When clogged,
interstitial flow rates and spaces are
reduced (Brim Box & Mosa 1999, p.
100), thus reducing juvenile habitat.
Furthermore, sediment may act as a
vector for delivering contaminants such
as nutrients and pesticides to streams
and juveniles may ingest contaminants
adsorbed to silt particles during normal
feeding activities. Female spectaclecase
and sheepnose produce conglutinates
that attract hosts. Such a reproductive
strategy depends on clear water during
the critical time of the year when
mussels are releasing their glochidia.
Agricultural activities are responsible
for much of the sediment (Waters 1995,
p. 170) and chemical discharge into
streams, affecting 60 percent of the
impaired river miles in the country
(EPA 2007, p. 10). Unrestricted livestock
access occurs on many streams and
potentially threatens their mussel
populations (Fraley & Ahlstedt 2000,
pp. 193–194). Grazing may reduce
infiltration rates and increase runoff;
trampling and vegetation removal
increases the probability of erosion
(Armour et al. 1991, pp. 8–10; Brim Box
& Mosa 1999, p. 103). The majority of
the remaining spectaclecase and
sheepnose populations are threatened
by some form of agricultural runoff
(nutrients, pesticides, sediment). Copper
Creek, a tributary to the Clinch River,
for example, has a drainage area that
contains approximately 41 percent
agricultural land (Hanlon et al. 2009, p.
3). Fraley and Ahlstedt (2000, p. 193)
and Hanlon et al. (2009, pp. 11–12)
attributed the decline of the Copper
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Creek mussel fauna to an increase in
cattle grazing and resultant nutrient
enrichment and loss of riparian
vegetation along the stream, among
other factors. This scenario is similar in
other parts of the extant range of the
spectaclecase and sheepnose.
Sedimentation and urban runoff may
also be threats to the sheepnose in the
Kankakee River system as the Chicago
Metro area continues to expand.
Declines in mussel diversity observed in
the Ohio River are in part due to
pollution from urban centers; in many
of these areas the loss of diversity has
not recovered from water quality
problems that began prior to dam
construction (Watters & Flaute 2010, pp.
3–7).
As the spectaclecase primarily
inhabits deep water along the outside of
bends, it may be particularly vulnerable
to siltation. The current often slackens
in this habitat, more so than in riffles
and runs where other mussel species are
typically found, and suspended
sediment settles out. Spectaclecase beds
covered with a thick layer of silt have
been observed in Missouri, often
downstream from reaches with eroding
banks (Roberts 2008, pers. comm.).
Channelization
Dredging and channelization
activities have profoundly altered
riverine habitats nationwide. Hartfield
(1993, pp. 131–139), Neves et al. (1997,
pp. 71–72), and Watters (2000, pp. 268–
269) reviewed the specific effects of
channelization on freshwater mussels.
Channelization impacts stream
physically (for example accelerated
erosion, reduced depth, decreased
habitat diversity, geomorphic
instability, and loss of riparian
vegetation) and biologically (for
example decreased fish and mussel
diversity, altered species composition
and abundance, decreased biomass, and
reduced growth rates) (Hartfield 1993,
pp. 131–139). Channel construction for
navigation increases flood heights (Belt
1975, p. 684), partly as a result of a
decrease in stream length and an
increase in gradient (Hubbard et al.
1993, p. 137 (in Hartfield 1993, p. 131)).
Flood events may thus be exacerbated,
conveying into streams large quantities
of sediment, potentially with adsorbed
contaminants. Channel maintenance
may result in profound impacts
downstream (Stansbery 1970, p. 10),
such as increases in turbidity and
sedimentation, which may smother
bottom-dwelling organisms.
Channel maintenance operations for
commercial navigation have impacted
habitat for the sheepnose and
spectaclecase in many large rivers
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rangewide. Periodic channel
maintenance may continue to adversely
affect this species in the upper
Mississippi, Ohio, Muskingum, and
Tennessee rivers. Further modifications
to the Mississippi River channel are
anticipated with the recently authorized
Navigation and Environmental
Sustainability Program (NESP) (Water
Resources Development Act of 2007
(Pub. L. 110–114)), which will consist of
construction of larger locks and other
navigation improvements downstream
of MRP 14. Continual maintenance of
the Mississippi River navigation
channel requires dredging, wing and
closing dam reconstruction and
maintenance, and bank armoring.
Dredging, maintenance, and
construction activities destabilize
instream fine sediments are likely to
adversely affect the spectaclecase and
the sheepnose. Spectaclecase tend to
inhabit relatively deep water where they
are particularly vulnerable to siltation.
The current is slower in this habitat
than in riffles and runs, and suspended
sediment settles out in greater volume.
Dredging to maintain barge traffic on the
Mississippi River below the mouth of
the Chippewa River in Wisconsin has
reduced mussel diversity due to the
increase in unstable sand substrates
(Thiel 1981, p. 20; U.S. Army Corps of
Engineers 1996, p. 1, Tab 14).
Channel maintenance dredging is also
a major concern for mussel populations.
A large amount of spoil (dredged earth
and rock) was dumped directly on a
mussel bed in the Muskingum River that
included the sheepnose in the late
1990s (Watters 2008, pers. comm.).
Thousands of mussels were killed as the
result of this single event. Watters and
Dunn (1995 p. 231) also noted that the
lower ends of two mussel beds
coincided with the mouths of Wolf and
Bear Creeks. This led them to surmise
that pollutants, such as sediment loads
or agricultural runoff, in their
watersheds may adversely impact
mussels in the mainstem Muskingum
River below the confluences of Wolf
Creek and Bear Creek.
Mussels require a stable substrate to
survive and reproduce and are
particularly susceptible to channel
instability (Neves et al. 1997, p. 23;
Parmalee & Bogan 1998). Channel and
bank degradation have led to the loss of
stable substrates in the Meramec River
Basin. Roberts and Bruenderman (2000,
pp. 7–8, 21–23) pointed to the loss of
suitable stable habitat as a major cause
of decline in mussel abundance at sites
previously surveyed in 1979.
The Tennessee River was once a
stronghold for the spectaclecase
(Ortmann 1924, p. 60; 1925, p. 327), and
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the sheepnose was originally known to
occur in the Tennessee River and 10 of
its tributaries (Ortmann 1925, p. 328).
The mainstem of the Tennessee River is
maintained as a navigational channel,
and a plan to deepen it has been
proposed (Hubbs 2008, pers. comm.).
Severe bank erosion is ongoing along
some reaches of the river below
Pickwick Landing Dam, with some sites
losing several feet of stream bank per
year (Hubbs 2008, pers. comm.).
The sheepnose population within the
Big Sunflower River is threatened by a
Corps flood control project (Jones 2008,
pers. comm.). Dredging for this project
is planned to take place downstream of
Indianaola, but head-cutting may
ultimately destabilize the substrate in
which sheepnose now exists. This
activity, coupled with other threats
potentially affecting sheepnose (see
below), may lead to extirpation of the
population within 10 years (Jones 2008,
pers. comm.).
The upper Kankakee River in Indiana
was channelized several decades ago.
The sheepnose is now considered
extirpated from the upper Kankakee,
and is restricted to the unchannelized
portion of the river in Illinois
(Cummings 2008, pers. comm.).
Mining
Instream gravel mining has been
implicated in the destruction of mussel
populations (Hartfield 1993, pp. 136–
138). Negative impacts associated with
gravel mining include stream channel
modifications (altered habitat, disrupted
flow patterns, and sediment transport),
water quality modifications (increased
turbidity, reduced light penetration, and
increased temperature),
macroinvertebrate population changes
(elimination, habitat disruption, and
increased sedimentation), and changes
in fish populations (impacts to
spawning and nursery habitat and food
web disruptions) (Kanehl & Lyons 1992,
pp. 4–10).
Heavy metal-rich drainage from coal
mining and associated sedimentation
has adversely impacted portions of the
Tennessee River system in Virginia.
Low pH commonly associated with
mine runoff can reduce glochidial
encystment (attachment) rates (Huebner
& Pynnonen 1992, pp. 2350–2353). Acid
mine runoff may thus have local
impacts on recruitment of the mussel
populations close to mines.
Coal-related toxins in the Clinch River
may explain the decline and lack of
mussel recruitment at some sites in the
Virginia portion of that stream (Ahlstedt
2008, pers. comm.). Patterns of mussel
distribution and abundances have been
found to be negatively correlated with
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proximity to coal-mining activities
(Ahlstedt & Tuberville 1997, pp. 74–75).
Known mussel toxicants, such as
polycyclic aromatic hydrocarbons,
heavy metals (for example, copper,
manganese, and zinc), and other
chemicals from coal mining and other
activities contaminate sediments in the
Clinch River (Ahlstedt & Tuberville
1997, p. 75). These chemicals are toxic
to juvenile mussels (Ahlstedt &
Tuberville 1997, p. 75). Pollutant inputs
to the Clinch River from a coal-burning
power plant in Carbo, Virginia, were
shown to increase mortality and reduce
cellulolytic activity (breaking down
cellulose) in transplanted mussels
(Farris et al. 1988, pp. 705–706). Sitespecific copper toxicity studies of
unionid glochidia in the Clinch River
showed that freshwater mussels as a
group were generally sensitive to
copper, the toxic constituent of the
power plant effluent (Cherry et al. 2002,
p. 596). All of these studies indicate that
acid mine runoff may have local
impacts on spectaclecase recruitment
and survival in this river.
Gravel-mining activities may also be a
localized threat in some streams with
extant sheepnose and spectaclecase
populations. Gravel mining causes
stream instability, increasing erosion,
turbidity, and subsequent sediment
deposition (Meador & Layher 1998, pp.
8–9). Gravel mining is common in the
Meramec River system. Between 1997
and 2008, the Missouri Department of
Natural Resources issued permits for
102 sand- and gravel-mining sites in the
Meramec River (Zeaman 2008, pers.
comm.). Although rigid guidelines
prohibited instream mining and
required streamside buffers, a court
ruling deauthorized the Corps from
regulating these habitat protective
measures. The Corps still retains
oversight for gravel mining, but many
mining operations do not fall under
Corps jurisdiction (Roberts &
Bruenderman 2000, p. 23). In the lower
Tennessee River, mining is permitted in
18 reaches for a total of 47.9 river miles
(77.1 km) between the Duck River
confluence and Pickwick Landing Dam,
a distance of over 95 miles (153 km)
(Hubbs 2008, pers. comm.). This is the
reach where mussel recruitment has
been noted for many rare species in
recent years. These activities have the
potential to impact the river’s small
sheepnose population. The Gasconade
River and its tributaries have been
subject to gravel mining and other
channel modifying practices that
accelerate channel destabilization.
These physical habitat threats combined
with poor water quality and agricultural
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may result in exposure of a relatively
immobile species to extremely elevated
concentrations that far exceed toxic
levels and any water quality standards
Oil and Gas Development
that might be in effect. Some notable
Coal, oil, and natural gas resources are spills that released large quantities of
present in some of the watersheds that
highly concentrated chemicals resulting
are known to support sheepnose,
in mortality to mussels include:
including the Allegheny River.
• Massive mussel kills on the Clinch
Exploration and extraction of these
River at Carbo, Virginia occurred from a
energy resources can result in increased power plant alkaline fly ash pond spill
siltation, a changed hydrograph, and
in 1967, and a sulfuric acid spill in 1970
altered water quality even at a distance
(Crossman et al. 1973, p. 6);
from the mine or well field. Sheepnose
• Approximately 18,000 mussels of
habitat in larger streams can be
several species, including 750
threatened by the cumulative effects of
individuals from three endangered
multiple mines and well fields (adapted mussel species, were eliminated from
from Service 2008, p. 11).
the upper Clinch River near Cedar Bluff,
Coal, oil, and gas resources are
Virginia in 1998, when an overturned
present in a number of the basins where
tanker truck released 1,600 gallons
sheepnose occur, and extraction of these
(6,056 liters) of a chemical used in
resources has increased dramatically in
rubber manufacturing (Jones et al. 2001,
recent years, particularly in
p. 20; Schmerfeld 2006, p. 12); and
Pennsylvania and West Virginia.
• An on-going release of sodium
Although oil and gas extraction
dimethyl dithiocarbamate, a chemical
generally occurs away from the river,
used to reduce and precipitate
extensive road networks are required to
hexachrome, starting in 1999 impacted
construct and maintain wells. These
road networks frequently cross or occur approximately 10 river miles (16 km) of
the Ohio River and resulted in an
near tributaries, contributing sediment
estimated loss of one million mussels,
to the receiving waterway. In addition,
including individuals from two
the construction and operation of wells
federally listed species (DeVault 2009,
may result in the discharge of brine.
pers. comm.; Clayton 2008b, pers.
Point source discharges are typically
comm.).
regulated; however, nonpoint inputs
These are not the only instances
such as silt and other contaminants may
where chemical spills have resulted in
not be sufficiently regulated,
the loss of high numbers of mussels
particularly those originating some
distance from a waterway. In 2006, more (Brown et al. 2005, p. 1457; Jones et al.
2001, p. 20; Neves 1991, p. 252;
than 3,700 permits were issued for oil
Schmerfeld 2006, pp. 12–13), but are
and gas wells by the Pennsylvania
provided as examples of the serious
Department of Environmental
threat chemical spills pose to mussel
Protection, which also issued 98
species. The sheepnose and
citations for permit violations at 54
spectaclecase are especially threatened
wells (Hopey 2007; adapted from
by chemical spills because these spills
Service 2008, p. 12).
can occur anywhere that highways with
Chemical Contaminants
tanker trucks, industries, or mines
overlap with sheepnose and
Chemical contaminants are
ubiquitous throughout the environment spectaclecase distribution.
Exposure of mussels to lower
and are considered a major threat in the
concentrations of contaminants more
decline of freshwater mussel species
likely to be found in aquatic
(Cope et al. 2008, p. 451; Richter et al.
1997, p. 1081; Strayer et al. 2004, p. 436; environments can also adversely affect
mussels and result in the decline of
Wang et al. 2007a, p. 2029). Chemicals
freshwater mussel species. Such
enter the environment through both
concentrations may not be immediately
point and nonpoint discharges
lethal, but over time, can result in
including spills, industrial sources,
mortality, reduced filtration efficiency,
municipal effluents, and agricultural
runoff. These sources contribute organic reduced growth, decreased
reproduction, changes in enzyme
compounds, heavy metals, pesticides,
activity, and behavioral changes to all
and a wide variety of newly emerging
mussel life stages. Frequently,
contaminants to the aquatic
procedures which evaluate the ‘‘safe’’
environment. As a result, water and
sediment quality can be degraded to the concentration of an environmental
contaminant (for example, national
extent that mussel populations are
water quality criteria) do not have data
adversely impacted.
for freshwater mussel species or exclude
Chemical spills can be especially
data that are available for freshwater
devastating to mussels because they
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nonpoint source pollution are serious
threats to all existing mussel fauna in
the system.
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mussels (March et al. 2007, pp. 2066–
2067, 2073).
Current research is now starting to
focus on the contaminant sensitivity of
freshwater mussel glochidia and newlyreleased juvenile mussels (Goudreau et
al. 1993, pp. 219–222; Jacobson et al.
1997, p. 2390; March et al. 2007, pp.
2068–2073; Valenti et al. 2006, pp.
2514–2517; Valenti et al. 2005, pp.
1244–1245; Wang et al. 2007c, pp.
2041–2046) and juveniles (Augspurger
et al. 2003, p. 2569; Bartsch et al. 2003,
p. 2561; March et al. 2007, pp. 2068–
2073; Mummert et al. 2003, p. 2549;
Valenti et al. 2006, pp. 2514–2517;
Valenti et al. 2005, pp. 1244–1245;
Wang et al. 2007b, pp. 2053–2055;
Wang et al. 2007c, pp. 2041–2046) to
such contaminants as ammonia, metals,
chlorine, and pesticides. The toxicity
information presented in this section
focuses on recent water-only laboratory
acute (sudden and severe exposure) and
chronic (prolonged or repeated
exposure) toxicity tests with early life
stages of freshwater mussels using the
standard testing methodology published
by the American Society for Testing and
Materials (ASTM) (American Society for
Testing and Materials. 2008. Standard
guide for conducting laboratory toxicity
tests with freshwater mussels E2455–06.
In Annual Book of ASTM Standards,
Vol 11.06. Philadelphia, PA, pp. 1442–
1493.) Use of this standard testing
method generates consistent, reliable
toxicity data with acceptable precision
and accuracy (Wang et al. 2007a, p.
2035) and was used for toxicity tests on
ammonia, copper, chlorine and select
pesticides (Augspurger et al. 2007, p.
2025; Bringolf et al. 2007b, p. 2101;
Bringolf et al. 2007c, p. 2087; Wang et
al. 2007a, p. 2029; Wang et al. 2007b, p.
2048; Wang et al. 2007c, p. 2036). Use
of these tests has documented that,
while mussels are sensitive to some
contaminants, they are not universally
sensitive to all contaminants
(Augspurger et al. 2007, pp. 2025–2026).
One chemical that is particularly toxic
to early life stages of mussels is
ammonia. Sources of ammonia include
agricultural wastes (animal feedlots and
nitrogenous fertilizers), municipal
wastewater treatment plants, and
industrial waste (Augspurger et al. 2007,
p. 2026) as well as precipitation and
natural processes (decomposition of
organic nitrogen) (Augspurger et al.
2003, p. 2569; Goudreau et al. 1993, p.
212; Hickey & Martin 1999, p. 44;
Newton 2003, p. 1243). Therefore,
ammonia is considered a limiting factor
for survival and recovery of some
mussel species due to its ubiquity in
aquatic environments and high level of
toxicity, and because the highest
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concentrations typically occur in mussel
microhabitats (Augspurger et al. 2003,
p. 2574). In addition, studies have
shown that ammonia concentrations
increase with increasing temperature
and low flow conditions (Cherry et al.
2005, p. 378; Cooper et al. 2005, p. 381),
which may be exacerbated by the effects
of climate change, and may cause
ammonia to become more problematic
for juvenile mussels. The EPAestablished ammonia water quality
criteria (EPA 1985, pp. 94–99) may not
be protective of mussels (Augspurger et
al. 2003, p. 2572; Sharpe 2005, p. 28)
under current and future climate
conditions.
Mussels are also affected by metals
(Keller & Zam 1991, p. 543), such as
cadmium, chromium, copper, mercury,
and zinc, which can negatively affect
biological processes such as growth,
filtration efficiency, enzyme activity,
valve closure, and behavior (Jacobson et
al. 1997, p. 2390; Keller & Zam 1991, p.
543; Naimo 1995, pp. 351–355; Valenti
et al. 2005, p. 1244). Metals occur in
industrial and wastewater effluents and
are often a result of atmospheric
deposition from industrial processes
and incinerators. Glochidia and juvenile
freshwater mussels have recently been
studied to determine the acute and
chronic toxicity of copper to these life
stages (Wang et al. 2007b, pp. 2048–
2056; Wang et al. 2007c, pp. 2036–
2047). The chronic values determined
for copper ranged from 8.5 to 9.8
micrograms per liter (ug/L) for survival
and from 4.6 to 8.5 ug/L for growth of
juveniles. These chronic values are
below the EPA 1996 chronic water
quality criterion of 15 ug/L (hardness
170 mg/L) for copper (Wang et al.
2007b, pp. 2052–2055). March (2007,
pp. 2066, 2073) identifies that copper
water quality criteria and modified State
water quality standards may not be
protective of mussels.
Mercury is another heavy metal that
has the potential to negatively affect
mussel populations, and it is receiving
attention due to its widespread
distribution and potential to adversely
impact the environment. Mercury has
been detected throughout aquatic
environments as a product of municipal
and industrial waste and atmospheric
deposition from coal-burning plants.
One recent study evaluated the
sensitivity of early life stages of mussels
to mercury (Valenti et al. 2005, p. 1242).
This study determined that for the
mussel species used (rainbow mussel,
Villosa iris) glochidia were more
sensitive to mercury than were juvenile
mussels, with the median lethal
concentration value of 14 ug/L
compared to 114 ug/L for the juvenile
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life stage. The chronic toxicity tests
conducted determined that juveniles
exposed to mercury greater than or
equal to 8 ug/L exhibited reduced
growth. These observed toxicity values
are below EPA’s Criteria Continuous
Concentration and Criteria Maximum
Concentration, which are 0.77 ug/L and
1.4 ug/L, respectively. Based on these
data, we believe that EPA’s water
quality standards for mercury should be
protective of juvenile mussels and
glochidia, except in cases of illegal
dumping, permit violations, or spills.
However, impacts to mussels from
mercury toxicity may be occurring in
some streams. According to the National
Summary Data reported by States to the
EPA, 3,770 monitored waters do not
meet EPA standards for mercury in the
United States. (https://iaspub.epa.gov/
waters10/attains_nation_cy.control?p_
report_type=T, accessed 6/28/2010).
Acute mercury toxicity was determined
to be the cause of extirpation of a
diverse mussel fauna for a 70-mile (112km) portion of the North Fork Holston
River (Brown et al. 2005, pp. 1455–
1457).
In addition to ammonia, agricultural
sources of chemical contaminants
include two broad categories that have
the potential to adversely impact mussel
species: Nutrients and pesticides.
Nutrients (such as nitrogen and
phosphorus) can impact streams when
their concentrations reach levels that
cannot be assimilated, a condition
known as over-enrichment. Nutrient
over-enrichment is primarily a result of
runoff from livestock farms, feedlots,
and heavily fertilized row crops
(Peterjohn & Correll 1984, p. 1471).
Over-enriched conditions are
exacerbated by low-flow conditions,
such as those experienced during
typical summer-season flows and that
might occur with greater frequency and
magnitude as a result of climate change.
Bauer (1988, p. 244) found that
excessive nitrogen concentrations can
be detrimental to the adult freshwater
pearl mussel (Margaritifera
margaritifera), as was evident by the
positive linear relationship between
mortality and nitrate concentration.
Also, a study of mussel life span and
size (Bauer 1992, p. 425) showed a
negative correlation between growth
rate and eutrophication, and longevity
was reduced as the concentration of
nitrates increased. Nutrient overenrichment can result in an increase in
primary productivity, and the
subsequent respiration depletes
dissolved oxygen levels. This may be
particularly detrimental to juvenile
mussels that inhabit the interstitial
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spaces in the substrate where lower
dissolved oxygen concentrations are
more likely than on the sediment
surface where adults tend to live
(Sparks & Strayer 1998, pp. 132–133).
Elevated concentrations of pesticide
frequently occur in streams due to
pesticide runoff, overspray application
to row crops, and lack of adequate
riparian buffers. Agricultural pesticide
applications often coincide with the
reproductive and early life stages of
mussel, and thus impacts to mussels
due to pesticides may be increased
(Bringolf et al. 2007a, p. 2094). Little is
known regarding the impact of currently
used pesticides to freshwater mussels
even though some pesticides, such as
glyphosate (Roundup), are used
globally. Recent studies tested the
toxicity of glyphosate, its formulations,
and a surfactant (MON 0818) used in
several glyphosate formulations, to early
life stages of the fatmucket (Lampsilis
siliquoidea), a native freshwater mussel
(Bringolf et al. 2007a, p. 2094). Studies
conducted with juvenile mussels and
glochidia determined that the surfactant
(MON 0818) was the most toxic of the
compounds tested and that L.
siliquoidea glochidia were the most
sensitive organism tested to date
(Bringolf et al. 2007a, p. 2094).
Roundup, technical grade glyphosate
isopropylamine salt, and
isopropylamine were also acutely toxic
to juveniles and glochidia (Bringolf et
al. 2007a, p. 2097). The impacts of other
pesticides including atrazine,
chlorpyrifos, and permethrin on
glochidia and juvenile life stages have
also recently been studied (Bringolf et
al. 2007b, p. 2101). This study
determined that chlorpyrifos was toxic
to both L. siliquoidea glochidia and
juveniles (Bringolf et al. 2007b, p. 2104).
The above results indicate the potential
toxicity of commonly applied pesticides
and the threat to mussel species as a
result of the widespread use of these
pesticides. All of these pesticides are
commonly used throughout the range of
the sheepnose and spectaclecase.
A potential, but undocumented, threat
to freshwater mussel species, including
sheepnose and spectaclecase, are
contaminants referred to as ‘‘emerging
contaminants’’ that are being detected in
aquatic ecosystems at an increasing rate.
Pharmaceuticals, hormones, and other
organic contaminants have been
detected downstream from urban areas
and livestock production (Kolpin et al.
2002, p. 1202). A large potential source
of these emerging contaminants is
wastewater being discharged through
both permitted (National Pollutant
Discharge Elimination System, or
NPDES) and non-permitted sites
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throughout the country. Permitted
discharge sites are ubiquitous in
watersheds with sheepnose and
spectaclecase populations, providing
ample opportunities for contaminants to
impact the species (for example, there
are more than 250 NPDES sites in the
Meramec River, Missouri system, which
harbors large, but declining, populations
of sheepnose and spectaclecase; Roberts
and Bruenderman 2000, p. 78).
The information presented in this
section represents some of the threats
from chemical contaminants that have
been documented both in the laboratory
and field and demonstrates that
chemical contaminants pose a
substantial threat to sheepnose and
spectaclecase. This information
indicates the potential for contaminants
from spills that are immediately lethal
to species, to chronic contaminant
exposure, which results in death,
reduced growth, or reduced
reproduction of sheepnose and
spectaclecase to contribute to declining
sheepnose and spectaclecase
populations.
Summary of Factor A
The decline of the freshwater mussels
in the eastern United States is primarily
the result of the long-lasting effects of
habitat alterations such as
impoundments, channelization,
chemical contaminants, mining, oil and
gas development, and sedimentation.
Although efforts have been made to
restore habitat in some areas, the longterm effects of large-scale and wideranging habitat modification,
destruction, and curtailment will
continue into the foreseeable future.
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B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
The spectaclecase and sheepnose are
not commercially valuable species but
may be increasingly sought by collectors
as they become rarer. Although
scientific collecting is not thought to
represent a significant threat,
unregulated collecting could adversely
affect localized spectaclecase and
sheepnose populations.
Mussel harvest is illegal in some
States (for example, Indiana and Ohio),
but regulated in others (for example,
Alabama, Kentucky, Tennessee, and
Wisconsin). These species may be
inadvertently harvested by
inexperienced commercial harvesters
unfamiliar with species identification.
Although illegal harvest of protected
mussel beds occurs (Watters and Dunn
1995, p. 225, 247–250), commercial
harvest is not known to have a
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significant impact on the spectaclecase
and sheepnose.
On the basis of this analysis, we find
that overutilization for commercial,
recreational, scientific, or educational
purposes is not now a threat to the
spectaclecase or sheepnose in any
portion of its range or likely to become
a significant threat in the foreseeable
future.
C. Disease or Predation
Little is known about diseases in
freshwater mussels (Grizzle & Brunner
2007, p. 6). However, mussel die-offs
have been documented in spectaclecase
and sheepnose streams (Neves 1986, p.
9), and some researchers believe that
disease may be a factor contributing to
the die-offs (Buchanan 1986, p. 53;
Neves 1986, p. 11). Mussel parasites
include water mites, trematodes,
oligochaetes, leeches, copepods,
bacteria, and protozoa (Grizzle &
Brunner 2007, p. 4). Generally, parasites
are not suspected of being a major
limiting factor (Oesch 1984, p. 6), but a
recent study provides contrary
evidence. Reproductive output and
physiological condition were negatively
correlated with mite and trematode
abundance, respectively (Gangloff et al.
2008, pp. 28–30). Stressors that reduce
fitness may make mussels more
susceptible to parasites (Butler 2007, p.
90). Furthermore, nonnative mussels
may carry diseases and parasites that are
potentially devastating to the native
mussel fauna, including spectaclecase
and sheepnose (Strayer 1999, p. 88).
The muskrat (Ondatra zibethicus) is
cited as the most prevalent mussel
predator (Convey et al. 1989, p. 654–
655; Hanson et al. 1989, pp. 15–16;
Kunz 1898, p. 328). Muskrat predation
may limit the recovery potential of
endangered mussels or contribute to
local extirpations of previously stressed
populations, according to Neves and
Odom (1989, p. 940), but they consider
it primarily a seasonal or localized
¨
threat. Bopple and Coker (1912, p. 9)
noted the occurrence of ‘‘large piles of
shells made by the muskrats’’ on an
island in the Clinch River, Tennessee,
composed of ‘‘about one-third’’
spectaclecase shells. Predation by
muskrats may be a seasonal and
localized threat to spectaclecase and
sheepnose populations but is probably
not a significant threat rangewide.
Some species of fish feed on mussels
(for example, common carp (Cyprinus
carpio), freshwater drum (Aplodinotus
grunniens), redear sunfish (Lepomis
microlophus)) and potentially on this
species when young. Various
invertebrates, such as flatworms, hydra,
non-biting midge larvae, dragonfly
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larvae, and crayfish, may feed on
juvenile mussels (Neves 2008, pers.
comm.). Although predation by
naturally occurring predators is a
normal aspect of the population
dynamics of a healthy mussel
population, predation may amplify
declines in small populations of this
species. In addition, the potential now
exists for the black carp
(Mylopharyngodon piceus), a molluskeating Asian fish recently introduced
into the waters of the United States
(Strayer 1999, p. 89), to eventually
disperse throughout the range of the
spectaclecase and sheepnose.
The life cycle of freshwater mussels is
intimately related to that of the
freshwater fish they use as hosts for
their parasitic glochidia. For this reason,
diseases that impact populations of
freshwater fishes also pose a significant
threat to mussels. Viral hemorrhagic
septicemia (VHS) disease has been
confirmed from much of the Great Lakes
and St. Lawrence River system. In June
2008, muskellunge (Esox masquinongy)
from Clearfork Reservoir, near
Mansfield, Ohio, tested positive for
carrying VHS virus. This is the first
known occurrence of VHS virus in the
Mississippi River basin.
The VHS virus has been implicated as
a mortality factor in fish kills
throughout the Great Lakes region. It has
been confirmed in 28 fish species, but
no identified hosts for sheepnose are on
the U.S. Department of Agriculture’s
Animal and Plant Health Inspection
Service (APHIS) list of fish species
susceptible to VHS (APHIS 2008, pp. 1–
2). Since the host for spectaclecase is
unknown, we do not know how VHS
could affect reproduction for
spectaclecase. If the VHS virus
successfully migrates out of the
Clearfork Reservoir and into the Ohio
River, it could spread rapidly and cause
fish kills throughout the Mississippi
River basin. Few spectaclecase and
sheepnose populations are currently
recruiting at sustainable levels, and fish
kills could further reduce encounters
with hosts and potentially reduce
recruitment.
In summary, disease in freshwater
mollusks is poorly known and not
currently considered a threat. Although
there is no direct evidence at this time
that predation is detrimentally affecting
the spectaclecase or sheepnose, their
small populations and limited ranges
leaves them vulnerable to threats of
predation from natural or introduced
predators. Therefore, we conclude that
predation currently represents a threat
of low magnitude, but it could
potentially become a significant future
threat to the spectaclecase and
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sheepnose due to their small population
sizes.
D. The Inadequacy of Existing
Regulatory Mechanisms
States with extant spectaclecase and
sheepnose populations prohibit the
taking of mussels for scientific purposes
without a State collecting permit.
However, enforcement of this permit
requirement is difficult.
The level of protection that
spectaclecase and sheepnose receive
from State listing varies from State to
State. The sheepnose is State-listed in
every State that keeps such a list.
Collection of sheepnose in Pennsylvania
for use as fish bait is limited to 50
individuals per day. The spectaclecase
is State-listed in 8 of the 10 States that
harbor extant populations. Only in
Missouri and Tennessee is the
spectaclecase not assigned conservation
status and West Virginia does not have
any State-specific legislation similar to
the Act.
Nonpoint source pollution is
considered a primary threat to
sheepnose and spectaclecase habitat;
however, current laws do not
adequately protect spectaclecase and
sheepnose habitat from nonpoint source
pollution, as the laws to prevent
sediment entering water ways are poorly
enforced. Best management practices for
sediment and erosion control are often
recommended or required by local
ordinances for construction projects;
however, compliance, monitoring, and
enforcement of these recommendations
are often poorly implemented.
Furthermore, there are currently no
requirements within the scope of
Federal environmental laws to
specifically consider the spectaclecase
and sheepnose during Federal activities.
Point source discharges within the
range of the spectaclecase and
sheepnose have been reduced since the
inception of the Clean Water Act (33
U.S.C. 1251 et seq.), but this may not
provide adequate protection for filter
feeding organisms that can be impacted
by extremely low levels of contaminants
(see ‘‘Chemical Contaminants’’
discussion under Factor A). There is no
specific information on the sensitivity of
the spectaclecase and sheepnose to
common industrial and municipal
pollutants, and very little information
on other freshwater mussels. Therefore,
it appears that a lack of adequate
research and data prevents existing
regulations, such as the Clean Water Act
(administered by the EPA and the
Corps), from being fully used or
effective.
The U.S. Army Corps of Engineers
retains oversight authority and requires
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a permit for gravel-mining activities that
deposit fill into streams under section
404 of the Clean Water Act.
Additionally, a Corps permit is required
under section 10 of the Rivers and
Harbors Act (33 U.S.C. 401 et seq.) for
navigable waterways including the
lower 50 miles (80 km) of the Meramec
River. However, many gravel-mining
operations do not fall under these two
categories.
Despite these existing regulatory
mechanisms, the spectaclecase and
sheepnose continue to decline due to
the effects of habitat destruction, poor
water quality, contaminants, and other
factors. We find that these regulatory
measures have been insufficient to
significantly reduce or remove the
threats to the spectaclecase and
sheepnose, and therefore that the
inadequacy of existing regulatory
mechanisms is a threat to these species
throughout all of their ranges.
Based on our analysis of the best
available information, we have no
reason to believe that the
aforementioned regulations will offer
adequate protection to the spectaclecase
and sheepnose in the foreseeable future.
3413
maximums, depending on the species
(Watters & O’Dee 2000, p. 136).
Abnormal temperature changes may
cause particular problems to mussels
whose reproductive cycles may be
linked to fish reproductive cycles (for
example,Young & Williams 1984).
E. Other Natural or Manmade Factors
Affecting Its Continued Existence
Climate Change
It is a widely accepted fact that
changes in climate are occurring
worldwide (IPCC 2007, p. 30).
Understanding the effects of climate
change on freshwater mussels is of
crucial importance, because the extreme
fragmentation of freshwater drainage
systems, coupled with the limited
ability of mussels to migrate, will make
it particularly difficult for mussels to
adjust their range in response to changes
in climate (Strayer 2008, p. 30). For
example, changes in temperature and
precipitation can increase the likelihood
of flooding or increase drought duration
and intensity, resulting in direct
impacts to freshwater mussels (Golladay
et al. 2004, p. 503; Hastie et al. 2003, pp.
40–43). Indirect effects of climate
change may include declines in host
fish stocks, sea level rise, habitat
reduction, and changes in human
activity in response to climate change
(Hastie et al. 2003, pp. 43–44).
Temperature
Natural temperature regimes can be
altered by impoundments, water
releases from dams, industrial and
municipal effluents, and changes in
riparian habitat. Critical thermal limits
for survival and normal functioning of
many freshwater mussel species are
unknown. High temperatures can
reduce dissolved oxygen concentrations
in the water, which slows growth,
reduces glycogen stores, impairs
respiration, and may inhibit
reproduction (Fuller 1974, pp. 240–
241). Low temperatures can
significantly delay or prevent
metamorphosis (Watters & O’Dee 1999,
pp. 454–455). Water temperature
increases have been documented to
shorten the period of glochidial
encystment, reduce righting speed,
increase oxygen consumption, and slow
burrowing and movement responses
(Bartsch et al. 2000, p. 237; Fuller 1974,
pp. 240–241; Schwalb & Pusch 2007,
pp. 264–265; Watters et al. 2001, p.
546). Several studies have documented
the influence of temperature on the
timing of aspects of mussel
reproduction (for example, Allen et al.
2007, p. 85; Gray et al. 2002, p. 156;
Steingraeber et al. 2007, pp. 303–309).
Peak glochidial releases are associated
with water temperature thresholds that
can be thermal minimums or thermal
Population Fragmentation and Isolation
Most of the remaining spectaclecase
and sheepnose populations are small
and isolated and thus are susceptible to
genetic drift, inbreeding depression, and
random or chance changes to the
environment, such as toxic chemical
spills (Avise and Hamrick 1996, pp.
463–466; Smith 1990, pp. 311–321;
Watters and Dunn 1995, pp. 257–258
Inbreeding depression can result in
death, decreased fertility, smaller body
size, loss of vigor, reduced fitness, and
various chromosome abnormalities
(Smith 1990, pp. 311–321). Despite any
evolutionary adaptations for rarity,
habitat loss and degradation increase a
species’ vulnerability to extinction
(Noss and Cooperrider 1994, pp. 58–62).
Numerous authors (including Noss and
Cooperrider 1994, pp. 58–62; Thomas
1994, p. 373) have indicated that the
probability of extinction increases with
decreasing habitat availability. Although
changes in the environment may cause
populations to fluctuate naturally, small
and low-density populations are more
likely to fluctuate below a minimum
viable population (the minimum or
threshold number of individuals needed
in a population to persist in a viable
state for a given interval) (Gilpin and
Soule 1986, pp. 25–33; Shaffer 1981, p.
131; Shaffer and Samson 1985, pp. 148–
150).
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These species were widespread
throughout much of the upper twothirds of the Mississippi River system,
for example, when few natural barriers
existed to prevent migration (via host
species) among suitable habitats.
Construction of dams, however,
destroyed many spectaclecase and
sheepnose populations and isolated
others. Recruitment reduction or failure
is a potential problem for many small
sheepnose populations rangewide, a
potential condition exacerbated by its
reduced range and increasingly isolated
populations. If these trends continue,
further significant declines in total
sheepnose population size and
consequent reduction in long-term
survivability may soon become
apparent.
Spectaclecase are long-lived (up to 70
years; Havlik 1994, p. 19) while
sheepnose are relatively long-lived
(approximately 30 years; Watters et al.
2009, p. 221) Therefore, it may take
decades for non-reproducing
populations of both species to become
extinct following their isolation by, for
example, the construction of a dam. The
occasional discovery of relatively young
spectaclecase in river reaches between
impoundments indicates that some
post-impoundment recruitment has
occurred. The level of recruitment in
these cases, however, appears to be
insufficient to ensure the long-term
sustainability of the spectaclecase.
Small isolated populations of
spectaclecase and sheepnose that may
now be comprised predominantly of
adult specimens could be dying out
slowly in the absence of recruitment,
even without other threats described
above. Isolated populations usually face
other threats that result in continually
decreasing patches of suitable habitat.
Genetic considerations for managing
imperiled mussels and for captive
propagation were reviewed by Neves
(1997a, p. 1422) and Jones et al. (2006,
pp. 527–535), respectively. The
likelihood is high that some populations
of the spectaclecase and sheepnose are
below the effective population size
(EPS) (Soule 1980, pp. 162–164)
necessary to adapt to environmental
change and persist in the long term.
Isolated populations eventually die out
when population size drops below the
EPS or threshold level of sustainability.
Evidence of recruitment in many
populations of these two species is
scant, making recruitment reduction or
outright failure suspect. These
populations may be experiencing the
bottleneck effect of not attaining the
effective population size. Small,
isolated, below effective population
size-threshold populations of short-
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lived species (most host fishes)
theoretically die out within a decade or
so, while below-threshold populations
of long-lived species, such as the
spectaclecase and sheepnose, might take
decades to die out even given years of
total recruitment failure. Without
historical barriers to genetic
interchange, small, isolated populations
could be slowly expiring, a
phenomenon termed the extinction debt
(Tilman et al. 1994, pp. 65–66). Even
given the totally improbable absence of
anthropogenic threats, we may lose
disjunct populations to below-threshold
effective population size. However,
evidence indicates that general
degradation continues to decrease
habitat patch size and to act insidiously
in the decline of spectaclecase and
sheepnose populations.
Spectaclecase and sheepnose mussels’
scarcity and decreased population size
makes maintaining adequate
heterogeneity problematic for resource
managers. Neves (1997b, p. 6) warned
that ‘‘[i]f we let conservation genetics
become the goal rather than the
guidelines for restoring and recovering
mussel populations, then we will be
doomed to failure with rare species.’’
Habitat alteration, not lack of genetic
variability, is the driving force of
population extirpation (Caro and
Laurenson 1994, pp. 485–486; Neves et
al. 1997, p. 60). Nevertheless, genetics
issues should be considered in
maintaining high levels of
heterozygosity during spectaclecase
recovery efforts. Treating disjunct
occurrences of this wide-ranging species
as a metapopulation would facilitate
conservation management while
increasing recovery options (for
example, translocating adults or
introducing infested hosts and
propagated juveniles) to establish and
maintain viable populations (Neves
1997b, p. 6). Due to small population
size and probable reduction of genetic
diversity within populations, efforts
should be made to maximize genetic
heterogeneity to avoid both inbreeding
(Templeton & Read 1984, p. 189) and
outbreeding depression (Avise &
Hamrick 1996, pp. 463–466) whenever
feasible in propagation and
translocation efforts (Jones et al. 2006,
p. 529).
We find that fragmentation and
isolation of small remaining populations
of the spectaclecase and sheepnose are
current and ongoing threats to both
species throughout all of their ranges
that will continue into the foreseeable
future. Further, stochastic events may
play a magnified role in population
extirpation when small, isolated
populations are involved.
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Exotic Species
Various exotic or nonnative species of
aquatic organisms are firmly established
in the range of the spectaclecase and
sheepnose. The exotic species that poses
the most significant threat to the
spectaclecase and sheepnose is the
zebra mussel (Dreissena polymorpha).
Its invasion of freshwater habitats in the
United States poses a threat to mussel
faunas in many regions, and species’
extinctions are expected as a result of its
continued spread in the eastern United
States (Ricciardi et al. 1998, p. 615).
Strayer (1999, pp. 75–80) reviewed in
detail the mechanisms in which zebra
mussels impact native mussels. The
primary means of impact is direct
fouling of the shells of live native
mussels. Zebra mussels attach in large
numbers to the shells of live native
mussels and are implicated in the loss
of entire native mussel beds. Fouling
impacts include impeding locomotion
(both laterally and vertically),
interfering with normal valve
movements, deforming valve margins,
and locally depleting food resources and
increasing waste products. Heavy
infestations of zebra mussels on native
mussels may overly stress the animals
by reducing their energy stores. They
may also reduce food concentrations to
levels too low to support reproduction,
or even survival in extreme cases.
Other ways zebra mussels may impact
spectaclecase and sheepnose is through
filtering their sperm and possibly
glochidia from the water column, thus
reducing reproductive potential. Habitat
for native mussels may also be degraded
by large deposits of zebra mussel
pseudofeces (undigested waste material
passed out of the incurrent siphon)
(Vaughan 1997, p. 11). Because
spectaclecase are found in pools and
zebra mussel veligers (larvae) attach to
hard substrates at the point at which
they settle out from the water column,
spectaclecase are particularly vulnerable
to zebra mussel invasion. The
spectaclecase’s colonial tendency could
allow for very large numbers to be
affected by a single favorable year for
zebra mussels.
Zebra mussels are established
throughout the upper Mississippi, lower
St. Croix, Ohio, and Tennessee Rivers,
overlapping much of the current range
of the spectaclecase and sheepnose. The
greatest potential for present zebra
mussel impacts to the spectaclecase and
sheepnose appears to be in the upper
Mississippi River. Kelner and Davis
(2002, p. ii) stated that zebra mussels in
the Mississippi River from Mississippi
River Pool 4 downstream are ‘‘extremely
abundant and are decimating the native
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mussel communities.’’ Huge numbers of
dead and live zebra mussels cover the
bottom of the river in some localities up
to 1 to 2 inches (2.5 to 5.1 centimeters
(cm)) deep (Havlik 2001a, p. 16), where
they have reduced significantly the
quality of the habitat with their
pseudofeces (Fraley 2008, pers. comm.).
Zebra mussels likely have reduced
spectaclecase and sheepnose
populations in these heavily infested
waters.
As zebra mussels may maintain high
densities in big rivers, large tributaries,
and below infested reservoirs,
spectaclecase and sheepnose
populations in affected areas may be
significantly impacted. For example,
zebra mussel densities in the Tennessee
River remained low until 2002, but are
now abundant enough below Wilson
Dam to be measured quantitatively
(Garner 2008, pers. comm.). In addition,
there is long-term potential for zebra
mussel invasions into other systems that
currently harbor spectaclecase and
sheepnose populations. Zebra mussels
occur in the lower St. Croix River, one
of the strongholds for spectaclecase,
although it is unclear whether they are
likely to spread much further upstream
due to the transition from lake-like
conditions to almost exclusively
riverine conditions above RM 25.
The Asian clam (Corbicula fluminea)
has spread throughout the range of the
spectaclecase and sheepnose since its
introduction in the mid-1900s. Asian
clams compete with native mussels,
especially juveniles, for food, nutrients,
and space (Leff et al. 1990, p. 415; Neves
& Widlak 1987, p. 6) and may ingest
unionid sperm, glochidia, and newly
metamorphosed juveniles of native
mussels (Strayer 1999, p. 82; Yeager et
al. 2000, p. 255). Dense Asian clam
populations actively disturb sediments
that may reduce habitat for juveniles of
native mussels (Strayer 1999, p. 82).
Asian clam densities vary widely in
the absence of native mussels or in
patches with sparse mussel
concentrations, but Asian clam density
is never high in dense mussel beds,
indicating that the clam is unable to
successfully invade small-scale habitat
patches with high unionid biomass
(Vaughn & Spooner 2006, pp. 334–335).
The invading clam therefore appears to
preferentially invade sites where
mussels are already in decline (Strayer
1999, pp. 82–83; Vaughn & Spooner
2006, pp. 332–336) and does not appear
be a causative factor in the decline of
mussels in dense beds. However, an
Asian clam population that thrives in
previously stressed, sparse mussel
populations might exacerbate unionid
imperilment through competition and
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impeding mussel population expansion
(Vaughn & Spooner 2006, pp. 335–336).
A molluscivore (mollusk eater), the
black carp (Mylopharyngodon piceus) is
a potential threat to native mussels
(Strayer 1999, p. 89); it has been
introduced into North America since the
1970s. The species has been proposed
for widespread use by aquaculturists to
control snails, the intermediate host of
a trematode (flatworm) parasite that
affects catfish in commercial culture
ponds in the southeast and lower
Midwest. Black carp are known to eat
clams (Corbicula spp.) and unionid
mussels in China, in addition to snails.
They are the largest of the Asian carp
species, reaching more than 4 ft. in
length and achieving a weight in excess
of 150 pounds (Nico & Williams 1996,
p. 6). Foraging rates for a 4-year-old fish
average 3 or 4 pounds (1.4–1.8 kg) a day,
indicating that a single individual could
consume 10 tons (9,072 kg) of native
mollusks over its lifetime (MICRA 2005,
p. 1). In 1994, 30 black carp escaped
from an aquaculture facility in Missouri
during a flood. Other escapes into the
wild by non-sterile black carp are likely
to occur.
The round goby (Neogobius
melanostomus) is another exotic fish
species released into the Great Lakes
that is well established and likely to
spread through the Mississippi River
system (Strayer 1999, pp. 87–88). This
species is an aggressive competitor of
similar sized benthic fishes (sculpins,
darters), as well as a voracious
carnivore, despite its size (less than 10
in. (25.4 cm) in length), preying on a
variety of foods, including small
mussels and fishes that could serve as
glochidial hosts (Janssen and Jude 2001,
p. 325; Strayer 1999, p. 88). Round
gobies may therefore have important
indirect effects on the spectaclecase and
sheepnose through negative effects to
their hosts.
Additional exotic species will
invariably become established in the
foreseeable future (Strayer 1999, pp. 88–
89). Added to potential direct threats,
exotic species could carry diseases and
parasites that may be devastating to the
native biota. Because of our ignorance of
mollusk diseases and parasites, ‘‘it is
imprudent to conclude that alien
diseases and parasites are unimportant’’
(Strayer 1999, p. 88).
Exotic species, such as those
described above, are an ongoing threat
to the spectaclecase and sheepnose—a
threat that is likely to increase as these
exotic species expand their occupancy
within the ranges of the spectaclecase
and sheepnose.
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Summary of Threats
The decline of the spectaclecase and
sheepnose in the eastern United States
(described by Butler 2002a, entire;
Butler 2002b, entire) is primarily the
result of habitat loss and degradation
(Neves 1991, p. 252). These losses have
been well documented since the mid19th century (Higgins 1858, p. 550).
Chief among the causes of decline are
impoundments, channelization,
chemical contaminants, mining, and
sedimentation (Neves 1991, p. 252;
Neves 1993, pp. 4–6; Neves et al. 1997,
pp. 60, 63–75; Watters 2000, pp. 262–
267; Williams et al. 1993, pp. 7–9).
These stressors have had profound
impacts on sheepnose and spectaclecase
populations and their habitat.
The majority of the remaining
populations of the spectaclecase and
sheepnose are generally small and
geographically isolated (Butler 2002a, p.
27; 2002b, p. 27). The patchy
distributional pattern of populations in
short river reaches makes them much
more susceptible to extirpation from
single catastrophic events, such as toxic
chemical spills (Watters and Dunn 1995,
p. 257). Furthermore, this level of
isolation makes natural repopulation of
any extirpated population virtually
impossible without human intervention.
In addition, the fish host of
spectaclecase is unknown; thus,
propagation to reestablish the species in
restored habitats and to maintain nonreproducing populations and focused
conservation of its fish host are
currently not possible. Although there
are ongoing attempts to alleviate some
of these threats at some locations, there
appear to be no populations without
significant threats, and many threats are
without obvious or readily available
solutions.
Recruitment reduction or failure is a
threat for many small spectaclecase and
sheepnose populations rangewide, a
condition exacerbated by reduced range
and increasingly isolated populations
(Butler 2002a, p. 28; 2002b, p. 28). If
these trends continue, further
significant declines in total
spectaclecase and sheepnose population
size and consequent reduction in longterm viability may soon become
apparent.
Various exotic species of aquatic
organisms are firmly established in the
range of the spectaclecase and
sheepnose. The exotic species that poses
the most significant threat to the
spectaclecase and sheepnose is the
zebra mussel. The invasion of the zebra
mussel poses a serious threat to mussel
faunas in many regions, and species
extinctions are expected as a result of its
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continued spread in the eastern United
States (Ricciardi et al. 1998, p. 618).
Proposed Determination
The Act defines an endangered
species as any species that is ‘‘in danger
of extinction throughout all or a
significant portion of its range’’ and a
threatened species as any species ‘‘that
is likely to become endangered
throughout all or a significant portion of
its range within the foreseeable future.’’
We find that the spectaclecase and
sheepnose are presently in danger of
extinction throughout their entire range,
based on the immediacy, severity, and
scope of the threats described under
Factors A, D, and E, above. Although
there are ongoing attempts to alleviate
some threats, there appear to be no
populations without current significant
threats, and many threats are without
obvious or readily available solutions.
These isolated species have a limited
ability to recolonize historically
occupied stream and river reaches and
are vulnerable to natural or humancaused changes in their stream and river
habitats. Their range curtailment, small
population size, and isolation make the
spectaclecase and sheepnose more
vulnerable to threats such as
sedimentation, disturbance of riparian
corridors, changes in channel
morphology, point and nonpoint source
pollutants, urbanization, and introduced
species and to stochastic events (for
example, chemical spills). Therefore, on
the basis of the best available scientific
and commercial information, we
propose listing the spectaclecase and
sheepnose as endangered in accordance
with sections 3(6) and 4(a)(1) of the Act.
Under the Act and our implementing
regulations, a species may warrant
listing if it is endangered or threatened
throughout all or a significant portion of
its range. Threats to the spectaclecase
and sheepnose occur throughout their
ranges; therefore, we assessed the status
of the species throughout their entire
ranges. The threats to the survival of the
species occur throughout the species’
ranges and are not restricted to any
particular significant portion of those
ranges. Accordingly, our assessment and
proposed determination applies to both
species throughout their entire ranges.
Available Conservation Measures
Conservation measures provided to
species listed as endangered or
threatened under the Act include
recognition, recovery actions,
requirements for Federal protection, and
prohibitions against certain practices.
Recognition through listing encourages
and results in public awareness and
conservation by Federal, State, and local
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agencies, private organizations, and
individuals. The Act encourages
cooperation with the States and requires
that recovery actions be carried out for
all listed species. The protection
required of Federal agencies and the
prohibitions against take and harm are
discussed, in part, below.
The primary purpose of the Act is the
conservation of endangered and
threatened species and the ecosystems
upon which they depend. The ultimate
goal of such conservation efforts is the
recovery of these listed species, so that
they no longer need the protective
measures of the Act. Subsection 4(f) of
the Act requires the Service to develop
and implement recovery plans for the
conservation of endangered and
threatened species, unless such a plan
will not promote the conservation of the
species. The recovery planning process
involves the identification of actions
that are necessary to halt or reverse the
species’ decline by addressing the
threats to its survival and recovery. The
goal of this process is to restore listed
species to a point where they are secure,
self-sustaining, and functioning
components of their ecosystems.
Recovery planning includes the
development of a recovery outline
shortly after a species is listed,
preparation of a draft and final recovery
plan, and revisions to the plan as
significant new information becomes
available. The recovery outline guides
the immediate implementation of urgent
recovery actions and describes the
process to be used to develop a recovery
plan. The recovery plan identifies sitespecific management actions that will
achieve recovery of the species,
measurable criteria that determine when
a species may be downlisted or delisted,
and methods for monitoring recovery
progress. Recovery plans also establish
a framework for agencies to coordinate
their recovery efforts and provide
estimates of the cost of implementing
recovery tasks. Recovery teams
(comprised of species experts, Federal
and State agencies, non-government
organizations, and stakeholders) are
often established to develop recovery
plans. When completed, the recovery
outline, draft recovery plan, and the
final recovery plan will be available on
our Web site (https://www.fws.gov/
endangered), or from our Rock Island,
Illinois, Ecological Services Field Office
(see FOR FURTHER INFORMATION CONTACT
section).
Implementation of recovery actions
generally requires the participation of a
broad range of partners, including other
Federal agencies, States, Tribal,
nongovernmental organizations,
businesses, and private landowners.
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Examples of recovery actions include
habitat restoration (e.g., restoration of
native vegetation), research, captive
propagation and reintroduction, and
outreach and education. The recovery of
many listed species cannot be
accomplished solely on Federal lands
because their range may occur primarily
or solely on non-Federal lands. To
achieve recovery of these species
requires cooperative conservation efforts
on private, State, and Tribal lands.
Listing will also require the Service to
review any actions on Federal lands and
activities under Federal jurisdiction that
may adversely affect the two species;
allow State plans to be developed under
section 6 of the Act; encourage scientific
investigations of efforts to enhance the
propagation or survival of the animals
under section 10(a)(1)(A) of the Act; and
promote habitat conservation plans on
non-Federal lands and activities under
section 10(a)(1)(B) of the Act.
Section 7(a) of the Act, as amended,
requires Federal agencies to evaluate
their actions with respect to any species
that is proposed or listed as endangered
or threatened and with respect to its
critical habitat, if any is designated.
Regulations implementing this
interagency cooperation provision of the
Act are codified at 50 CFR part 402.
Federal agencies are required to confer
with us informally on any action that is
likely to jeopardize the continued
existence of a proposed species. Section
7(a)(4) requires Federal agencies to
confer with the Service on any action
that is likely to jeopardize the continued
existence of a species proposed for
listing or result in destruction or
adverse modification of proposed
critical habitat. If a species is listed
subsequently, section 7(a)(2) requires
Federal agencies to ensure that activities
they authorize, fund, or carry out are not
likely to jeopardize the continued
existence of the species or destroy or
adversely modify its critical habitat. If a
Federal action may adversely affect a
listed species or its critical habitat, the
responsible Federal agency must enter
into formal consultation with the
Service.
Federal activities that may affect the
sheepnose and spectaclecase include,
but are not limited to, the funding of,
carrying out of, or the issuance of
permits for reservoir construction,
natural gas extraction, stream
alterations, discharges, wastewater
facility development, water withdrawal
projects, pesticide registration, mining,
and road and bridge construction.
Jeopardy Standard
Prior to and following listing and
designation of critical habitat, if prudent
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and determinable, the Service applies
an analytical framework for jeopardy
analyses that relies heavily on the
importance of core area populations to
the survival and recovery of the species.
The section 7(a)(2) analysis is focused
not only on these populations but also
on the habitat conditions necessary to
support them.
The jeopardy analysis usually
expresses the survival and recovery
needs of the species in a qualitative
fashion without making distinctions
between what is necessary for survival
and what is necessary for recovery.
Generally, if a proposed Federal action
is incompatible with the viability of the
affected core area population(s),
inclusive of associated habitat
conditions, a jeopardy finding is
considered to be warranted, because of
the relationship of each core area
population to the survival and recovery
of the species as a whole.
EMCDONALD on DSK2BSOYB1PROD with MISCELLANEOUS
Section 9 Take
Section 9(a)(2) of the Act, and its
implementing regulations found at 50
CFR 17.21, set forth a series of general
prohibitions and exceptions that apply
to all endangered wildlife. These
prohibitions, in part, make it illegal for
any person subject to the jurisdiction of
the United States to take (includes
harass, harm, pursue, hunt, shoot,
wound, kill, trap, or collect, or to
attempt any of these), import or export,
ship in interstate commerce in the
course of commercial activity, or sell or
offer for sale in interstate or foreign
commerce any listed species. It also is
illegal to knowingly possess, sell,
deliver, carry, transport, or ship any
wildlife that has been taken illegally.
Certain exceptions apply to agents of the
Service and State conservation agencies.
We may issue permits to carry out
otherwise prohibited activities
involving endangered wildlife species
under certain circumstances.
Regulations governing permits are at 50
CFR 17.22 for endangered species. Such
permits are available for scientific
purposes, to enhance the propagation or
survival of the species, or for incidental
take in connection with otherwise
lawful activities.
Our policy, as published in the
Federal Register on July 1, 1994 (59 FR
34272), is to identify, to the maximum
extent practicable, those activities that
would or would not likely constitute a
violation of section 9 of the Act. The
intent of this policy is to increase public
awareness as to the potential effects of
this final listing on future and ongoing
activities within a species’ range. We
believe that the following activities are
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unlikely to result in a violation of
section 9:
(1) Existing discharges into waters
supporting these species, provided these
activities are carried out in accordance
with existing regulations and permit
requirements (for example, activities
subject to sections 402, 404, and 405 of
the Clean Water Act and discharges
regulated under the National Pollutant
Discharge Elimination System).
(2) Actions that may affect the
spectaclecase or sheepnose and are
authorized, funded, or carried out by a
Federal agency when the action is
conducted in accordance with any
reasonable and prudent measures we
have specified in accordance with
section 7 of the Act.
(3) Development and construction
activities designed and implemented
under Federal, State, and local water
quality regulations and implemented
using approved best management
practices.
(4) Existing recreational activities,
such as swimming, wading, canoeing,
and fishing, that are in accordance with
State and local regulations, provided
that if a spectaclecase or sheepnose is
collected, it is immediately released,
unharmed.
Activities that we believe could
potentially result in take of
spectaclecase or sheepnose include but
are not limited to:
(1) Illegal collection or capture of the
species;
(2) Unlawful destruction or alteration
of the species’ occupied habitat (for
example, unpermitted instream
dredging, channelization, or discharge
of fill material);
(3) Violation of any discharge or water
withdrawal permit within the species’
occupied range; and
(4) Illegal discharge or dumping of
toxic chemicals or other pollutants into
waters supporting spectaclecase or
sheepnose.
We will review other activities not
identified above on a case-by-case basis
to determine whether they are likely to
result in a violation of section 9 of the
Act. We do not consider these lists to be
exhaustive and provide them as
information to the public.
You should direct questions regarding
whether specific activities may
constitute a future violation of section 9
to the Field Supervisor of the Service’s
Rock Island, Illinois Ecological Services
Field Office (see FOR FURTHER
INFORMATION CONTACT section). You may
request copies of the regulations
regarding listed wildlife from and
address questions about prohibitions
and permits to the U.S. Fish and
Wildlife Service, Ecological Services
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Division, Henry Whipple Federal
Building, 1 Federal Drive, Fort Snelling,
MN 55111 (Phone (612) 713–5350; Fax
(612) 713–5292).
Critical Habitat
Background
Critical habitat is defined in section 3
of the Act as:
(i) The specific areas within the
geographical area occupied by a species,
at the time it is listed in accordance
with the Act, on which are found those
physical or biological features
(I) Essential to the conservation of the
species, and
(II) That may require special
management considerations or
protection; and
(ii) Specific areas outside the
geographical area occupied by a species
at the time it is listed, upon a
determination that such areas are
essential for the conservation of the
species.
Conservation is defined in section 3 of
the Act as the use of all methods and
procedures needed to bring the species
to the point at which listing under the
Act is no longer necessary.
Critical habitat receives protection
under section 7 of the Act through the
prohibition against Federal agencies
carrying out, funding, or authorizing the
destruction or adverse modification of
critical habitat. Section 7(a)(2) requires
consultation on Federal actions that
may affect critical habitat. The
designation of critical habitat does not
affect land ownership or establish a
refuge, wilderness, reserve, preserve, or
other conservation area. Such
designation does not allow the
government or public to access private
lands. Such designation does not
require implementation of restoration,
recovery, or enhancement measures by
non-Federal landowners. Where a
landowner seeks or requests Federal
agency funding or authorization for an
action that may affect a listed species or
critical habitat, the consultation
requirements of section 7(a)(2) of the
Act would apply, but even in the event
of a destruction or adverse modification
finding, the obligation of the Federal
action agency and the applicant is not
to restore or recover the species, but to
implement reasonable and prudent
alternatives to avoid destruction or
adverse modification of critical habitat.
Prudency Determination
Section 4(a)(3) of the Act, as
amended, and implementing regulations
(50 CFR 424.12), require that, to the
maximum extent prudent and
determinable, we designate critical
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habitat at the time the species is
determined to be endangered or
threatened. Our regulations (50 CFR
424.12(a)(1)) state that the designation
of critical habitat is not prudent when
one or both of the following situations
exist: (1) The species is threatened by
taking or other human activity, and
identification of critical habitat can be
expected to increase the degree of threat
to the species, or (2) such designation of
critical habitat would not be beneficial
to the species.
There is currently no imminent threat
of take attributed to collection or
vandalism under Factor B
(overutilization for commercial,
recreational, scientific, or educational
purposes) for sheepnose and
spectaclecase and identification of
critical habitat is not expected to initiate
such a threat. In the absence of finding
that the designation of critical habitat
would increase threats to a species, if
there are any benefits to a critical
habitat designation, then a prudent
finding is warranted. The potential
benefits include: (1) Triggering
consultation under section 7(a)(2) of the
Act, in new areas for actions in which
there may be a Federal nexus where it
would not otherwise occur because the
species may not be present; (2) focusing
conservation activities on the most
essential habitat features and areas; (3)
increasing awareness of important
habitat areas among State or county
governments or private entities; and (4)
preventing inadvertent harm to the
species.
Critical habitat designation includes
the identification of the physical and
biological features of the habitat
essential to the conservation of each
species that may require special
management and protection. As such,
these designations will provide useful
information to individuals, local and
State governments, and other entities
engaged in activities or long-range
planning that may affect areas essential
to the conservation of the species.
Conservation of the spectaclecase and
sheepnose and essential features of their
habitats will require habitat
management, protection, and
restoration, which will be facilitated by
disseminating information on the
locations and the key physical and
biological features of those habitats. In
the case of spectaclecase and sheepnose,
these aspects of critical habitat
designation would potentially benefit
the conservation of the species.
Therefore, since we have determined
that the designation of critical habitat
will not likely increase the degree of
threat to these species and may provide
some measure of benefit, we find that
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designation of critical habitat is prudent
for the spectaclecase and sheepnose.
Primary Constituent Elements
In accordance with sections 3(5)(A)(i)
and 4(b)(1)(A) of the Act and regulations
at 50 CFR 424.12, in determining which
areas to propose as critical habitat, we
must consider those physical and
biological features—primary constituent
elements in the necessary and
appropriate quantity and spatial
arrangement—essential to the
conservation of the species. We must
also consider those areas essential to the
conservation of the species that are
outside the geographical area occupied
by the species. Primary constituent
elements include, but are not limited to:
(1) Space for individual and
population growth and for normal
behavior;
(2) Food, water, air, light, minerals, or
other nutritional or physiological
requirements;
(3) Cover or shelter;
(4) Sites for breeding, reproduction,
and rearing (or development) of
offspring; and
(5) Habitats that are protected from
disturbance or are representative of the
historical, geographical, and ecological
distribution of a species.
We are currently unable to identify
the primary constituent elements for
spectaclecase and sheepnose because
information on the physical and
biological features that are considered
essential to the conservation of these
species is not known at this time. The
apparent poor viability of the species’
occurrences observed in recent years
indicates that current conditions are not
sufficient to meet the basic biological
requirements of these species in many
rivers. Since spectaclecase and
sheepnose have not been observed for
decades in many of their historical
locations, and much of the habitat in
which they still persists has been
drastically altered, the optimal
conditions that would provide the
biological or ecological requisites of
these species are not known. Although
we can surmise that habitat degradation
from a variety of factors has contributed
to the decline of these species, we do
not know specifically what essential
physical or biological features of that
habitat are currently lacking for
spectaclecase and sheepnose.
Key features of the basic life history,
ecology, reproductive biology, and
habitat requirements of most mussels,
including spectaclecase and sheepnose,
are unknown. Species-specific
ecological requirements have not been
determined (for example, minimum
water flow and effects of particular
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pollutants). Population dynamics, such
as species’ interactions and community
structure, population trends, and
population size and age class structure
necessary to maintain a long-term
viability, have not been determined for
these species. Basics of reproductive
biology for these species are unknown,
such as age and size at earliest maturity,
reproductive longevity, and the level of
recruitment needed for species survival
and long-term viability. Of particular
concern to the spectaclecase is the lack
of known host(s) species essential for
glochidia survival and reproductive
success. Similarly, although recent
laboratory studies have produced
successful transformation of sheepnose
glochidia on a few fish species, many
questions remain concerning the natural
interactions between the sheepnose and
its known hosts. Because the host(s) for
spectaclecase is unknown and little is
known about the sheepnose hosts, there
is a degree of uncertainty at this time as
to which specific areas might be
essential to the conservation of these
species (for example, the host(s)’s
biological needs and population sizes
necessary to support mussel
reproduction and population viability)
and thus meet a key aspect of the
definition of critical habitat. As we are
unable to identify many physical and
biological features essential to the
conservation of spectaclecase and
sheepnose, we are unable to identify
areas that contain these features.
Therefore, although we have determined
that the designation of critical habitat is
prudent for spectaclecase and
sheepnose, because the biological and
physical requirements of these species
are not sufficiently known, we find that
critical habitat for spectaclecase and
sheepnose is not determinable at this
time.
Peer Review
In accordance with our policy,
‘‘Notice of Interagency Cooperative
Policy for Peer Review in Endangered
Species Act Activities,’’ that was
published on July 1, 1994 (59 FR
34270), we will seek the expert opinion
of at least three appropriate
independent specialists regarding this
proposed rule. The purpose of such
review is to ensure listing decisions are
based on scientifically sound data,
assumptions, and analysis. We will send
copies of this proposed rule to the peer
reviewers immediately following
publication in the Federal Register. We
will invite these peer reviewers to
comment, during the public comment
period, on the specific assumptions and
the data that are the basis for our
conclusions regarding the proposal to
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list spectaclecase and sheepnose as
endangered and our proposal regarding
critical habitat for this species.
Required Determinations
Clarity of the Rule
We are required by Executive Orders
12866 and 12988 and by the
Presidential Memorandum of June 1,
1998, to write all rules in plain
language. This means that each rule we
publish must:
(a) Be logically organized;
(b) Use the active voice to address
readers directly;
(c) Use clear language rather than
jargon;
(d) Be divided into short sections and
sentences; and
(e) Use lists and tables wherever
possible.
If you feel that we have not met these
requirements, send us comments by one
of the methods listed in the ADDRESSES
section. To better help us revise the
rule, your comments should be as
specific as possible. For example, you
should tell us the names of the sections
or paragraphs that are unclearly written,
which sections or sentences are too
long, the sections where you feel lists or
tables would be useful, etc.
Executive Order 13211
On May 18, 2001, the President issued
Executive Order 13211 on regulations
that significantly affect energy supply,
distribution, and use. Executive Order
13211 requires agencies to prepare
Statements of Energy Effects when
undertaking certain actions. This rule is
not expected to significantly affect
energy supplies, distribution, or use.
Therefore, this action is not a significant
energy action, and no Statement of
Energy Effects is required.
Paperwork Reduction Act of 1995 (44
U.S.C. 3501 et seq.)
This proposed rule does not contain
any new collections of information that
require approval by the Office of
Management and Budget (OMB) under
the Paperwork Reduction Act. The rule
would not impose new recordkeeping or
reporting requirements on State or local
governments, individuals, businesses, or
organizations. We may not conduct or
sponsor, and you are not required to
respond to, a collection of information
unless it displays a currently valid OMB
control number.
National Environmental Policy Act
We determined that we do not need
to prepare an environmental
assessment, as defined under the
authority of the National Environmental
Policy Act of 1969 (42 U.S.C. 4321 et
seq.), in connection with regulations
adopted under section 4(a) of the Act.
We published a notice outlining our
reasons for this determination in the
Federal Register on October 25, 1983
(48 FR 49244).
References Cited
A complete list of all references cited
in this rule is available on the Internet
at https://www.regulations.gov or upon
request from the Field Supervisor, Rock
Species
Vertebrate population
where
endangered or
threatened
Historical range
Common name
*
CLAMS
Scientific name
*
*
*
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*
Plethobasus cyphyus
*
*
*
U.S.A. (AL, IL, IN, IA, NA .............................
KY, MN, MS, MO,
OH, PA, TN, VA,
WV, WI).
*
Spectaclecase
*
Cumberlandia
monodonta.
*
*
*
U.S.A. (AL, AR, IL,
NA .............................
IN, IA, KS, KY, MN,
MO, NE, OH, TN,
VA, WV, WI).
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Authors
The primary authors of this proposed
rule are the staff members of the
Service’s Rock Island and Twin Cities
Field Offices (see FOR FURTHER
INFORMATION CONTACT section).
List of Subjects in 50 CFR Part 17
Endangered and threatened species,
Exports, Imports, Reporting and
recordkeeping requirements,
Transportation.
Proposed Regulation Promulgation
Accordingly, we propose to amend
part 17, subchapter B of chapter I, title
50 of the Code of Feral Regulations, as
follows:
PART 17—[AMENDED]
1. The authority citation for part 17
continues to read as follows:
Authority: 16 U.S.C. 1361–1407; 16 U.S.C.
1531–1544; 16 U.S.C. 4201–4245; Pub. L. 99–
625, 100 Stat. 3500; unless otherwise noted.
2. In § 17.11(h) add the entries for
‘‘Sheepnose’’ and ‘‘Spectaclecase’’ in
alphabetical order under CLAMS to the
List of Endangered and Threatened
Wildlife, as follows:
§ 17.11 Endangered and threatened
wildlife.
*
*
*
(h) * * *
Status
*
Critical
habitat
*
*
NA
*
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rules
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*
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NA
*
NA
*
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*
When
listed
*
*
Sheepnose .................
*
Island, Illinois Ecological Services Field
Office (see FOR FURTHER INFORMATION
CONTACT section).
*
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*
3420
*
*
Federal Register / Vol. 76, No. 12 / Wednesday, January 19, 2011 / Proposed Rules
*
*
Dated: December 16, 2010.
Rowan W. Gould,
Acting Director, U.S. Fish and Wildlife
Service.
*
[FR Doc. 2011–469 Filed 1–18–11; 8:45 am]
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Agencies
[Federal Register Volume 76, Number 12 (Wednesday, January 19, 2011)]
[Proposed Rules]
[Pages 3392-3420]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2011-469]
[[Page 3391]]
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Part IV
Department of the Interior
-----------------------------------------------------------------------
Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Endangered Status for
the Sheepnose and Spectaclecase Mussels; Proposed Rule
Federal Register / Vol. 76 , No. 12 / Wednesday, January 19, 2011 /
Proposed Rules
[[Page 3392]]
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R3-ES-2010-0050; MO 92210-0-0008-B2]
RIN 1018-AV93
Endangered and Threatened Wildlife and Plants; Endangered Status
for the Sheepnose and Spectaclecase Mussels
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Proposed rule.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service (Service), propose to
list two freshwater mussels, the spectaclecase mussel (Cumberlandia
monodonta) and sheepnose (Plethobasus cyphyus) as endangered under the
Endangered Species Act of 1973, as amended (Act). If we finalize this
rule as proposed, it would extend the Act's protections to these
species throughout their ranges, including sheepnose in Alabama,
Illinois, Indiana, Iowa, Kentucky, Minnesota, Mississippi, Missouri,
Ohio, Pennsylvania, Tennessee, Virginia, West Virginia, and Wisconsin,
and spectaclecase in Alabama, Arkansas, Illinois, Indiana, Iowa,
Kentucky, Kansas, Minnesota, Missouri, Nebraska, Ohio, Tennessee,
Virginia, West Virginia, and Wisconsin. We determined that critical
habitat for these species is prudent, but not determinable at this
time. The Service seeks data and comments from the public on this
proposed listing rule.
DATES: We will consider comments and information we receive from all
interested parties by March 21, 2011. We must receive requests for
public hearings, in writing, at the address shown in the FOR FURTHER
INFORMATION CONTACT section by March 7, 2011.
ADDRESSES: You may submit comments by one of the following methods:
Federal eRulemaking Portal: https://www.regulations.gov.
Follow the instructions for submitting comments on docket number FWS-
R3-ES-2010-0050.
U.S. mail or hand-delivery: Public Comments Processing,
Attn: FWS-R3-2010-0050; Division of Policy and Directives Management;
U.S. Fish and Wildlife Service; 4401 North Fairfax Drive, Suite 222;
Arlington, VA 22203.
We will post all comments on https://www.regulations.gov. This
generally means that we will post any personal information you provide
us (see Public Comments section below for more information).
FOR FURTHER INFORMATION CONTACT: Richard Nelson, Field Supervisor, at
the U.S. Fish and Wildlife Service, Rock Island, Illinois Ecological
Services Field Office, 1511 47th Avenue, Moline, IL 61265 (telephone
309-757-5800).
SUPPLEMENTARY INFORMATION:
Public Comments
Our intent is to use the best available commercial and scientific
data as the foundation for all endangered and threatened species
classification decisions. We request comments or suggestions from other
concerned governmental agencies, the scientific community, industry, or
any other interested party concerning this proposed rule to list the
spectaclecase and sheepnose mussels as endangered. We particularly seek
comments concerning:
(1) Biological, commercial trade, or other relevant data concerning
any threats (or lack thereof) to the species and regulations that may
be addressing those threats.
(2) Additional information concerning the ranges, distributions,
and population sizes of the species, including the locations of any
additional populations of these species.
(3) Any additional information on the biological or ecological
requirements of these species.
(4) Current or planned activities in the areas occupied by these
species and possible impacts of these activities on the species and
their habitats.
(5) Potential effects of climate change on these species and their
habitats.
(6) The reasons why areas should or should not be designated as
critical habitat as provided by section 4 of the Act (16 U.S.C. 1531 et
seq.), including whether the benefits of designation would outweigh
threats to the species that designation could cause (e.g., exacerbation
of existing threats, such as overcollection), such that the designation
of critical habitat is prudent.
(7) Specific information on:
What areas contain physical and biological features
essential for the conservation of these species;
What areas are essential to the conservation of these
species and
Special management considerations or protection that
proposed critical habitat may require.
Please note that submissions merely stating support for or
opposition to the action under consideration without providing
supporting information, although noted, will not be considered in
making a determination, as section 4(b)(1)(A) of the Act directs that
determinations as to whether any species is an endangered or threatened
species must be made ``solely on the basis of the best scientific and
commercial data available.''
You may submit your comments and materials concerning this proposed
rule by one of the methods listed in the ADDRESSES section. We will not
accept comments sent by e-mail or fax or to an address not listed in
the ADDRESSES section. Comments must be submitted to https://www.regulations.gov before 11:59 (Eastern Time) on the date specified
in the DATES section. We will not consider hand-delivered comments that
we do not receive, or mailed comments that are not postmarked, by the
date specified in the DATES section.
We will post your entire comment--including your personal
identifying information--on https://www.regulations.gov. If you provide
personal identifying information in your comment, you may request at
the top of your document that we withhold this information from public
review. However, we cannot guarantee that we will be able to do so.
Comments and materials we receive, as well as supporting
documentation we used in preparing this proposed rule, will be
available for public inspection on https://www.regulations.gov, or by
appointment, during normal business hours at the Rock Island, Illinois
Ecological Services Field Office (see the FOR FURTHER INFORMATION
CONTACT section).
Public Hearing
The Act provides for one or more public hearings on this proposal,
if requested. Requests must be received by March 7, 2011. Such requests
must be made in writing and be addressed to the Field Supervisor at the
address provided in the FOR FURTHER INFORMATION CONTACT section. We
will schedule public hearings on this proposal, if any are requested,
and announce the dates, times, and places of those hearings, as well as
how to obtain reasonable accommodations, in the Federal Register and
local newspapers at least 15 days before the hearing.
Persons needing reasonable accommodations to attend and participate
in a public hearing should contact the Rock Island, Illinois Ecological
Services Field Office by telephone at 309-757-5800, as soon as
possible. To allow sufficient time to process requests, please call no
later than one week before the hearing date. Information regarding this
proposed rule is available in alternative formats upon request.
[[Page 3393]]
Background
Species Descriptions
The spectaclecase (Cumberlandia monodonta) is a member of the
mussel family Margaritiferidae and was originally described as Unio
monodonta Say, 1829. The type locality is the Falls of the Ohio (on the
Ohio River in the vicinity of Louisville, Kentucky, and adjacent
Indiana), and the Wabash River (probably the lower portion in Illinois
and Indiana) (Parmalee and Bogan 1998, p. 49). Parmalee and Bogan
(1998, p. 49) summarized the synonymy of the spectaclecase. The species
has been placed in the genera Unio, Margaritana, Alasmidonta,
Margarita, Margaron, and Margaritifera at various times in history.
Ortmann (1912, p. 13) placed it in the monotypic (a taxonomic group
with only one biological type) genus Cumberlandia in the family
Margaritiferidae. Currently recognized synonymy includes Unio
soleniformis (Lea). Smith (2001, p. 43) reassigned the spectaclecase to
the Holarctic genus Margaritinopsis based on shell and gill characters.
However, the Service will defer to the Committee on Scientific and
Vernacular Names of Mollusks of the Council of Systematic
Malacologists, American Malacological Union (Turgeon et al. 1998), on
whether the genus Margaritinopsis is accepted as valid for the
spectaclecase. Until an official decision is made, the Service will use
the commonly accepted Cumberlandia for the genus of this species.
Spectaclecase is the accepted common name for Cumberlandia monodonta
(Turgeon et al. 1998, p. 32).
The spectaclecase is a large mussel that reaches at least 9.25
inches (23.5 centimeters (cm)) in length (Havlik 1994, p. 19). The
shape of the shell is greatly elongated, sometimes arcuate (curved),
and moderately inflated, with the valves being solid and moderately
thick, especially in older individuals (Parmalee & Bogan 1998, p. 49).
Both anterior and posterior ends of the shell are rounded with a
shallow depression near the center of shell (Baird 2000, p. 6; Parmalee
& Bogan 1998, p. 49). The anterior end is higher than the posterior end
(Baird 2000, p. 6). The posterior ridge is low and broadly rounded
(Parmalee & Bogan 1998, p. 50). Year-one specimens have heavy ridges
running parallel with the growth arrests, which are shell lines that
indicate slower periods of growth, thought to be laid down annually
(Baird 2000, p. 6). The periostracum (external shell surface) is
somewhat smooth, rayless, and light yellow, greenish-tan, or brown in
young specimens, becoming rough and dark brown to black in old shells
(Parmalee & Bogan 1998, p. 50). The shell commonly will crack
posteriorly when dried (Oesch 1984, p. 31).
Internally, the single pseudocardinal tooth (a triangular tooth-
like structure along the hinge line of the internal portion of the
shell) is simple and peg-like in the right valve, fitting into a
depression in the left (Parmalee & Bogan 1998, p. 50). The lateral
teeth are straight and single in the right valve, and double in the
left valve but become fused with age into an indistinct raised hinge
line (Parmalee & Bogan 1998, p. 50). The soft anatomy was described by
Williams et al. (2008, pp. 497-498). The color of the nacre (interior
covering of the shell) is white, occasionally granular and pitted,
mostly iridescent in young specimens, but becoming iridescent
posteriorly in older shells (Parmalee & Bogan 1998, p. 50). There are
no differences between the sexes in the shells of this species (Baird
2000, p. 19). Key characters for distinguishing the spectaclecase from
other mussels are its large size, elongate shape, arcuate ventral
margin, dark coloration, roughened periostracum, poorly developed
teeth, and white nacre (Oesch 1984, pp. 31-32). No other North American
mussel species has this suite of characters.
The sheepnose (Plethobasus cyphyus) is a member of the mussel
family Unionidae and was originally described as Obliquaria cyphya
Rafinesque, 1820. The type locality is the Falls of the Ohio (Parmalee
& Bogan 1998, p. 175) on the Ohio River in the vicinity of Louisville,
Kentucky, and adjacent Indiana. Parmalee and Bogan (1998, p. 175)
summarized the synonymy of the species. Over the years, the name of
this species has been variably spelled cyphya, scyphius, cyphius,
cyphia, cyphyum, and ultimately cyphyus. Over the years the species has
been placed in the genera Obliquaria, Unio, Pleurobema, Margarita, and
Margaron. It was ultimately placed in the genus Plethobasus by Ortmann
(1919, pp. 65-66) where it remains today (Turgeon et al. 1998, p. 35).
The Service recognizes Unio aesopus and U. compertus as synonyms of
Plethobasus cyphyus. Sheepnose is the accepted common name for
Plethobasus cyphyus as established by the Committee on Scientific and
Vernacular Names of Mollusks of the Council of Systematic
Malacologists, American Malacological Union (Turgeon et al. 1998, p.
35). The Service also recognizes ``bullhead'' and ``clear profit'' as
older common names for the sheepnose.
Key characters useful for distinguishing the sheepnose from other
mussels are its color, the occurrence of central tubercles, and its
general shape. Oesch (1984, p. 120) and Parmalee and Bogan (1998, p.
176), describe the sheepnose as a medium-sized mussel that reaches
nearly 5 inches (13 cm) in length. The shell is elongate ovate in
shape, moderately inflated, and with thick, solid valves. The anterior
end of the shell is rounded, but the posterior end is somewhat bluntly
pointed to truncate. The dorsal margin of the shell is nearly straight,
while the ventral margin is uniformly rounded or slightly convex. The
posterior ridge is gently rounded, becoming flattened ventrally and
somewhat biangular. There is a row of large, broad tubercular swellings
on the center of the shell extending from the beak to the ventral
margin. A broad, shallow sulcus (depression on furrow on the outside
surface of shell) lies between the posterior ridge and central row.
Beaks are elevated, high, and placed near the anterior margin. Juvenile
beak sculpture consists of a few concentric ridges at the tip of the
beaks. The periostracum is generally smooth, shiny, rayless, and light
yellow to a dull yellowish brown. Concentric ridges resulting from
growth arrests are usually darker.
Oesch (1984, p. 120) describes the internal anatomy of the
sheepnose as the left valve having two heavy, erect, roughened,
somewhat triangular, and divergent pseudocardinal teeth. The right
valve has a large, triangular, roughened pseudocardinal tooth. The
lateral teeth are heavy, long, slightly curved, and serrated. The beak
cavity is shallow to moderately deep. The soft anatomy was described by
Williams et al. (2008, p. 94). The color of the nacre is generally
white, but may be pinkish to cream-colored and iridescent posteriorly.
There are no differences between the sexes in the shells of this
species. The shell of the sheepnose is extremely hard and was given the
name ``clear profit'' by early commercial shellers, being too hard to
cut into buttons (Wilson & Clark 1914, p. 57). The species also
preserves well in archaeological material (Morrison 1942, p. 357).
Life History
The general biology of the spectaclecase and sheepnose are similar
to other bivalve mollusks belonging to the families Margaritiferidae
and Unionidae, order Unioniformes or Unionoida. Adult mussels
suspension-feed, spending their entire lives partially or completely
buried within the substrate (Murray and Leonard 1962, p. 27). Adults
feed on algae, bacteria, detritus, microscopic animals, and dissolved
organic material (Christian et
[[Page 3394]]
al. 2004, pp. 108-109; Nichols & Garling 2000, p. 873; Silverman et al.
1997, p. 1859; Strayer et al. 2004, pp. 430-431). Recent evidence
suggests that adult mussels may also deposit feed on particles in the
sediment (Raikow & Hamilton 2001, p. 520). For their first several
months, juvenile mussels employ foot (pedal) feeding, consuming
bacteria, algae, and detritus (Yeager et al. 1994, p. 221).
As a group, mussel longevity varies tremendously with some species
living only about 4 years (Haag & Rypel 2010, p. 5) but possibly up to
100 to 200 years in other species (Ziuganov et al. 2000, p. 102).
However, the vast majority of species live a few decades (Haag & Rypel
2010, pp. 4-6). Baird (2000, pp. 54, 59, 67) aged 278 specimens of the
spectaclecase in Missouri by sectioning the hinge ligament, as most
margaritiferids are aged. The maximum age determined was 56 years, but
he surmised that some large individuals may have been older. A very
large specimen (9.25 inches (23.5 cm)) from the St. Croix River,
Minnesota and Wisconsin, was estimated (based on external growth ring
counts) to be approximately 70 years old (Havlik 1994, p. 19).
Sheepnose longevity has been reported as being nearly 30 years (Watters
et al. 2009, p. 221). Thick shelled mussels from large rivers, like
sheepnose, are thought to live longer than other species (Stansbery
1961, p. 16).
Mussels tend to grow relatively rapidly for the first few years,
and then slow appreciably at sexual maturity, when energy presumably is
being diverted from growth to reproductive activities (Baird 2000, pp.
66-67). In spectaclecase, the biggest change in growth rate appears to
occur at 10 to15 years of age, which suggests that significant
reproductive investment does not occur until they reach 10 years of age
(Baird 2000, pp. 66-67).
Margaritiferids and unionids have an unusual mode of reproduction.
With very few exceptions, their life cycle includes a brief, obligatory
parasitic stage on a host organism, typically fish. Eggs develop into
microscopic larvae (glochidia) within special gill chambers of the
female. The female expels the mature glochidia, which must attach to an
appropriate host species (generally a fish) to complete development.
Host specificity varies among margaritiferids and unionids. Some
species appear to use a single host, while others can transform on
several host species. Following successful infestation, glochidia
encyst (enclose in a cyst-like structure), remain attached to the host
for several weeks, and then drop off as newly transformed juveniles.
For further information on the life history of freshwater mussels, see
Williams et al. 2008.
Mussel biologists know relatively little about the specific life-
history requirements of the spectaclecase and sheepnose. Most mussels,
including the spectaclecase and sheepnose, have separate sexes. Age at
sexual maturity of the spectaclecase was estimated to be 4 to 5 years
for males and 5 to 7 years for females, with sex ratios approximating
50:50 (Baird 2000, p. 24). The spectaclecase life cycle includes a
parasitic phase; however, despite extensive investigation, the host
species is not yet known. The spectaclecase is thought to release
glochidia from early April to late May in the Meramec and Gasconade
Rivers, Missouri (Baird 2000, p. 26). Gordon and Smith (1990, p. 409)
reported the species as producing two broods, one in spring or early
summer and the other in the fall, also based on Meramec River
specimens. In the Meramec and Gasconade Rivers, however, Baird (2000,
pp. 26-27) found no evidence of two spawns in a given year.
Age at sexual maturity for the sheepnose is unknown, but given its
estimated longevity, probably occurs after a few years. The sheepnose
is thought to be a short-term brooder, with egg fertilization taking
place in early summer (Parmalee & Bogan 1998, p. 177; Williams et al.
1998, p. 498), and glochidial release presumably occurring later in the
summer. Hermaphroditism occurs in many mussel species (van der Schalie
1966, p. 77), but is not known for the sheepnose. If hermaphroditism
does occur in the sheepnose, it may explain the occurrence of small,
but persistent populations over long periods of time.
Glochidia of spectaclecase and sheepnose are released in
conglutinates (gelatinous structures containing numerous glochidia and
analogous to cold capsules). Spectaclecase glochidia lack hooks (teeth-
like structures that presumably function to pierce through skin tissue
of the host) and are the smallest glochidia known of any North American
freshwater mussel; they measure approximately 0.0024 inches (0.06 mm)
in both length and height (Baird 2000, p. 22). Tens to hundreds of
thousands of glochidia may occur in each conglutinate. Based on eight
Missouri spectaclecase specimens, the number of conglutinates released
per female varied from 53 to 88, with a mean of 64.5 (Baird 2000, p.
23). Total fecundity (reproductive potential, including glochidia and
ova) in Baird's (2000, p. 27) Missouri study varied from 1.93 to 9.57
million per female. In mussels, fecundity is related positively to body
size and inversely related to glochidia size (Bauer 1994, pp. 940-941).
The reproductive potential of the spectaclecase is therefore
phenomenal. However, the fact that extant populations are generally
skewed towards larger adults strongly indicates that survival rates to
the adult stage must be extraordinarily low.
Researchers in Wisconsin observed female spectaclecase under
boulders in the St. Croix River simultaneously releasing their
conglutinates (Heath 2008, pers. comm.). The spectaclecase
conglutinates are entrained along a transparent, sticky mucous strand
up to several feet in length (Lee & Hove 1997, p. 9). Baird (2000, p.
29) observed the release of loose glochidia and small fragments of
conglutinates. Based on his observations, he hypothesized that
conglutinates sometimes contain mostly immature glochidia, and that
conglutinates containing mostly immature glochidia may be aborted when
disturbed.
Sheepnose conglutinates are narrow and lanceolate in outline, solid
and red or pink in color, and discharged in unbroken form (Oesch 1984,
pp. 118-119). Discharge of sheepnose conglutinates have been observed
in late July (Ortmann 1911, p. 306) and August (Williams et al. 2008,
p. 498). Ortmann (1911, p. 306) described them as being pink and
``lying behind the posterior end of the shell, which were greedily
devoured by a number of minnows.'' Sheepnose glochidia are semicircular
in outline, with the ventral margin obliquely rounded, hinge line long,
and medium in size. The length (0.009 inches (0.23 mm)) is slightly
greater than the height (0.008 inches (0.20 mm)) (Oesch 1984, p. 119).
Several hundred glochidia probably occur in each conglutinate. Judging
from the size of the glochidia, total fecundity (including glochidia
and ova) per female sheepnose is probably in the tens of thousands.
Like many freshwater mussels, the complex life histories of the
spectaclecase and sheepnose have many vulnerable components that may
prevent successful reproduction or recruitment of juveniles into
existing populations. Glochidia must come into contact with a specific
host species for their survival to be ensured. Without the proper host,
the glochidia will perish. The host(s) for the spectaclecase is
unknown, although over 60 species of fish, amphibians, and crayfish
have been tested in the lab during host suitability studies (Baird
2000, pp. 23-24; Henley & Neves 2006, p. 3; Hove et al. 2009b, pp. 22-
23; Hove et al. 1998,
[[Page 3395]]
pp. 13-14; Hove et al. 2008, p. 4; Knudsen & Hove 1997, p. 2; Lee &
Hove 1997, pp. 9-10). Two of 690 wild-collected fish checked by Baird
(2000, p. 24) had spectaclecase glochidia attached to their gills;
these fish were the bigeye chub (Hybopsis amblops) and pealip redhorse
(Moxostoma pisolabrum). However, these fish are not confirmed as hosts,
because the encysted glochidia had not grown measurably and glochidial
transformation was not observed (Baird 2000, p. 24). Spectaclecase
populations are oftentimes highly aggregated (see Habitat) with many
apparently even-aged individuals, suggesting that glochidia may excyst
simultaneously from a host (Gordon & Layzer 1989, p. 19). Additional
host work is underway to test the wild-collected fish species that were
found with encysted spectaclecase glochidia (pealip redhorse and bigeye
chub), as well as to test additional species of fish and other aquatic
organisms for suitability. Host information is needed so that existing
populations can be artificially cultured for potential population
augmentation and reintroduction efforts.
Little is known regarding host fish of the sheepnose. Until
recently the only cited host for this species came from a 1914 report
that found glochidia naturally attached to sauger (Sander canadense) in
the wild. No confirmation of successful transformation was recorded in
this early report (Surber 1912, p. 110; Wilson 1914, pp. 338-340).
However, recent laboratory studies at the Genoa National Fish Hatchery,
the University of Minnesota, and Ohio State University have
successfully transformed sheepnose glochidia on fathead minnow
(Pimephales promelas), creek chub (Semotilus atrromaculatus), central
stoneroller (Campostoma anomalum), and brook stickleback (Culaea
inconstans) (Watters et al. 2005, pp. 11-12; Brady 2008, pers. comm.;
Watters 2008, pers. comm.). Although these are identified as suitable
hosts in laboratory studies, natural interactions between the
aforementioned fishes and the sheepnose seem rare and infrequent due to
habitat preferences. Fish that frequent medium to large rivers near
mussel beds, like the sauger, may act as hosts in the natural
environment.
Habitat
The spectaclecase generally inhabits large rivers, and is found in
microhabitats sheltered from the main force of current. It occurs in
substrates from mud and sand to gravel, cobble, and boulders in
relatively shallow riffles and shoals with a slow to swift current
(Baird 2000, pp. 5-6; Buchanan 1980, p. 13; Parmalee & Bogan 1998, p.
50). According to Stansbery (1967, pp. 29-30), this species is usually
found in firm mud between large rocks in quiet water very near the
interface with swift currents. Specimens have also been reported in
tree stumps, in root masses, and in beds of rooted vegetation (Oesch
1984, p. 33). Similar to other margaritiferids, spectaclecase
occurrences throughout much of its range tend to be aggregated (Gordon
& Layzer 1989, p. 19), particularly under slab boulders or bedrock
shelves (Baird 2000, p. 6; Buchanan 1980, p. 13; Parmalee & Bogan 1998,
p. 50), where they are protected from the current. Up to 200 specimens
have been reported from under a single large slab in the Tennessee
River at Muscle Shoals, Alabama (Hinkley 1906, p. 54). Unlike most
species that move about to some degree, the spectaclecase may seldom if
ever move except to burrow deeper and may die from stranding during
droughts (Oesch 1984, p. 17).
The sheepnose is primarily a larger-stream species occurring
primarily in shallow shoal habitats with moderate to swift currents
over coarse sand and gravel (Oesch 1984, p. 121). Habitats with
sheepnose may also have mud, cobble, and boulders. Sheepnose in larger
rivers may occur at depths exceeding 6 m (Williams et al. 2008, p.
498).
Genetics
A recent genetic study (Monroe et al. 2007, pp. 7-13) indicates
that much of the remaining genetic variability in the spectaclecase is
represented in each of the remaining large populations, and that these
populations do not appear to differ significantly from one another.
Genetics studies of sheepnose are currently under investigation;
however, no conclusions were available at the time of publication (Roe
2010, pers. comm.).
Species Distribution
We use the term ``population'' here in a geographical and not
genetic sense, defining it as all individuals of the spectaclecase or
sheepnose living in one stream. Using the term in this way allows the
status, trends, and threats to be discussed comparatively across
streams where the species occur. In using this term we do not imply
that their populations are currently reproducing and recruiting or that
they are distinct genetic units. We considered populations of the
spectaclecase and sheepnose as extant if live or fresh-dead specimens
have been observed or collected since 1990. A ``population cluster''
refers to where two or more adjacent stream populations of a species
occur without a barrier (for example, a dam and impoundment) between
them.
Following are generalized sets of criteria that were used to
categorize the relative status of populations of spectaclecase and
sheepnose. The status of a population is considered ``improving'' if:
(1) There is evidence that habitat degradation appears insignificant,
(2) live or fresh dead mussel abundance has improved during post-1990
surveys, or (3) ample evidence of recent recruitment has been
documented during post-1990 surveys. The status of a population is
considered ``stable'' if: (1) There is little evidence of significant
habitat loss or degradation, (2) live or fresh dead mussel abundance
has been fairly consistent during post-1990 surveys, or (3) evidence of
relatively recent recruitment has been documented during post-1990
surveys. The status of a population is considered ``declining'' if: (1)
There is ample evidence of significant habitat loss or degradation, (2)
live or fresh dead mussel numbers have declined during recent surveys,
or (3) no evidence of relatively recent recruitment has been documented
during recent surveys. The status of a population is considered
``extirpated'' if: (1) All known suitable habitat has been destroyed,
or (2) no live or fresh dead mussels of any age have been located
during recent surveys. The status of a population is considered
``unknown'' if the available information is inadequate to place the
population in one of the above four categories. In a few cases,
additional information not listed above may have been used to
categorize a population.
Spectaclecase Historical Range and Distribution
The spectaclecase occurred historically in at least 44 streams in
the Mississippi, Ohio, and Missouri River basins (Butler 2002a, p. 6,
Heath 2008, pers. comm.). Its distribution comprised portions of 15
States (Alabama, Arkansas, Illinois, Indiana, Iowa, Kansas, Kentucky,
Minnesota, Missouri, Nebraska, Ohio, Tennessee, Virginia, West
Virginia, and Wisconsin). Historical occurrence by stream system (with
tributaries) include the: upper Mississippi River system (Mississippi
River (St. Croix, Chippewa, Rock, Salt, Illinois (Des Plaines, Kankakee
Rivers), Meramec (Bourbeuse, Big Rivers), Kaskaskia Rivers; Joachim
Creek)); lower Missouri River system (Missouri River (Platte, River Aux
Vases, Osage (Sac, Marais des Cygnes Rivers), Gasconade (Osage Fork,
Big Piney River) Rivers)); Ohio River system (Ohio River
[[Page 3396]]
(Muskingum, Kanawha, Green, Wabash Rivers)); Cumberland River system
(Cumberland River (Big South, Caney Fork; Stones, Red Rivers));
Tennessee River system (Tennessee River (Holston, Nolichucky, Little,
Little Tennessee, Clinch (Powell River), Sequatchie, Elk, Duck
Rivers)); lower Mississippi River system (Mulberry, Ouachita Rivers).
Spectaclecase Current Range and Distribution
Extant populations of the spectaclecase are known from 19 streams
in 11 States (Butler 2002b, p. 7). These include the following stream
systems (with tributaries):
Upper Mississippi River system (Mississippi River (St.
Croix, Meramec (Bourbeuse, Big Rivers) Rivers));
Lower Missouri River system (Sac and Gasconade (Osage
Fork, Big Piney River) Rivers);
Lower Ohio River system (lowermost Ohio River (Kanawha,
Green Rivers));
Cumberland River system (Cumberland River);
Tennessee River system (Tennessee River (Nolichucky,
Clinch, Duck Rivers)); and
Lower Mississippi River system (Mulberry, Ouachita
Rivers).
The 19 extant spectaclecase populations occur in the following 11
States (with streams):
Alabama (Tennessee River),
Arkansas (Mulberry, Ouachita Rivers),
Illinois (Mississippi, Ohio Rivers),
Iowa (Mississippi River),
Kentucky (Ohio, Green Rivers),
Minnesota (Mississippi, St. Croix Rivers),
Missouri (Mississippi, Meramec, Bourbeuse, Big, Gasconade,
Sac, Big Piney Rivers; Osage Fork),
Tennessee (Tennessee, Clinch, Nolichucky, Duck Rivers;
Caney Fork),
Virginia (Cumberland, Clinch Rivers),
West Virginia (Kanawha River), and
Wisconsin (Mississippi, St. Croix Rivers).
Spectaclecase Population Estimates and Status
Based on historical and current data, the spectaclecase has
declined significantly rangewide and is now known from only 19 of 44
streams (Table 1), representing a 57 percent decline. The species is
presumed extirpated from thousands of river miles and from numerous
reaches of habitat in which it occurred historically, including long
reaches of upper Mississippi, Ohio, Cumberland, and Tennessee Rivers
and many other streams and stream reaches. Of the 19 extant
populations, 6 are represented by only one or two recent specimens each
and are likely declining and some may be extirpated. Populations in
Mississippi and Clinch Rivers have recently experienced significant
population declines. Most surviving populations face significant
threats and with few exceptions are highly fragmented and restricted to
short stream reaches. The spectaclecase is considered extirpated from
Indiana, Kansas, Nebraska, and Ohio. The only relatively strong
populations remaining are in the Meramec and Gasconade Rivers in
Missouri and in the St. Croix River in Minnesota and Wisconsin.
Table 1--Spectaclecase Status in All Streams of Historical or Current Occurrence
----------------------------------------------------------------------------------------------------------------
Date of Last Live
River Basin Stream Current Status Observation Comments
----------------------------------------------------------------------------------------------------------------
Upper Mississippi River......... Mississippi River. declining......... 2009.............. ..................
St. Croix River... stable............ 2008.............. ..................
Chippewa River.... extirpated........ 1989.............. ..................
Rock River........ extirpated........ ~1970............. ..................
Salt River........ extirpated........ 1980.............. ..................
Illinois River.... extirpated........ ~1914............. ..................
Des Plaines River. extirpated........ ~1921............. ..................
Kankakee River.... extirpated........ 1906.............. ..................
Meramec River..... stable............ 2003.............. ..................
Bourbeuse River... stable............ 1997.............. ..................
Big River......... stable............ 2002.............. ..................
Kaskaskia River... extirpated........ ~1970............. ..................
Joachim Creek..... extirpated........ ~1965............. ..................
Lower Missouri River............ Missouri River.... extirpated........ ~1914............. ..................
Platte River...... extirpated........ ~1917............. ..................
River Aux Vases... extirpated........ ~1974............. ..................
Osage River....... extirpated........ 1980.............. ..................
Sac River......... declining......... 2001.............. ..................
Marais des Cygnes extirpated........ unknown........... relic shell
River. observed in 1998.
Gasconade River... stable............ 2007.............. ..................
Big Piney River... unknown........... 2004.............. ..................
Osage Fork........ unknown........... 1999.............. ..................
Ohio River...................... Ohio River........ declining......... 1994.............. single individual
observed.
Muskingum River... extirpated........ unknown........... relic shell
observed in 1995.
Kanawha River..... unknown........... 2005.............. two live
individuals
observed.
Green River....... unknown........... 2006.............. ..................
Wabash River...... extirpated........ 1970.............. ..................
Cumberland River................ Cumberland River.. unknown........... 2008.............. single individual
observed.
Big South Fork.... extirpated........ 1911.............. ..................
Caney Fork........ extirpated........ 1988.............. ..................
Stones River...... extirpated........ 1968.............. ..................
Red River......... extirpated........ 1966.............. ..................
Tennessee River................. Tennessee River... unknown........... 2001.............. ..................
Holston River..... extirpated........ 1981.............. ..................
Nolichucky River.. unknown........... 1991.............. ..................
[[Page 3397]]
Little River...... extirpated........ ~1911............. ..................
Little Tennessee extirpated........ unknown........... relic shell
River. observed in 1980,
previous record
archaeological.
Clinch River...... declining......... 2005.............. ..................
Powell River...... extirpated........ ~1978............. ..................
Sequatchie River.. extirpated........ ~1925............. ..................
Elk River......... extirpated........ unknown........... relic shell
observed in 1998.
Duck River........ declining......... 1999.............. single individual
observed.
Lower Mississippi River......... Mulberry River.... unknown........... ~1995............. single individual
observed.
Ouachita River.... declining......... 2000.............. two individuals
observed.
----------------------------------------------------------------------------------------------------------------
Based on collections made over 100 years ago, the spectaclecase was
historically widespread and locally common in many streams rangewide.
The spectaclecase is often absent from archaeological shell middens
(Morrison 1942, p. 353) and is generally difficult to find due to its
habit of occurring under rocks or ledges and burrowing deep into the
substrate (Parmalee 1967, p. 25). Therefore, the chance of casually
finding the species where population numbers are low is remote.
The spectaclecase was considered a rare species by mussel experts
as early as 1970 (Stansbery 1970, p. 13), when the first attempt was
made to compile a list of imperiled mussels. The spectaclecase is
considered widely distributed but absent from many areas where it
formerly occurred (Cummings & Mayer 1992, p. 22). The American
Malacological Union and American Fisheries Society consider the
spectaclecase to be threatened (Williams et al. 1993, p.10). Six of the
19 streams (or big river reaches) considered to harbor extant
populations of the spectaclecase are represented by one or two recent
specimens (for example, Ohio, Kanawha, Cumberland, Duck, Ouatchita, and
Mulberry Rivers), exemplifying the species' imperiled status rangewide.
In some streams, the last reported records for the spectaclecase
occurred decades ago (for example, Rock, Des Plaines, Kaskaskia,
Platte, Wabash, Stones, Red, and Little Rivers; River Aux Vases; Big
South Fork). Parmalee (1967, p. 25) considered the spectaclecase to be
``rare and of local occurrence'' in Illinois in the 1960s, but that it
had ``[a]pparently already been extirpated from the Illinois and
Kankakee Rivers.'' The only records known from some streams are relic
specimens collected around 1975 (for example, Marais des Cygnes,
Muskingum, and Elk Rivers).
Although quantitative historical abundance data for the
spectaclecase is rare, generalized relative abundance (the percent
abundance of a species, divided by the total abundance of all mussel
species combined) was sometimes noted in the historical literature and
can be inferred from museum lots. The following is a summary of what is
known about the relative abundance and trends of presumably extant
spectaclecase populations by stream system.
Upper Mississippi River System
The spectaclecase was historically known from 13 streams in the
upper Mississippi River system. Currently, only four streams in the
system are thought to have extant spectaclecase populations.
Mississippi River mainstem: In 1907, Bartsch found spectaclecase at
approximately nine of the 140 sampled sites from what are now
Mississippi River Pools (MRP) 9 to 22 (Havlik 2001b, p. 10). Grier
(1922, p. 11) did not find spectaclecase in sampled portions of MRP 4
to 6. Van der Schalie and van der Schalie (1950, p. 456), reporting on
studies from the upper Mississippi River to the Missouri River mouth,
stated that no live spectaclecase were found in their study of 254
sites during 1930-31. Havlik and Stansbery (1977, p. 12) thought the
spectaclecase had disappeared from MRP 8 by the 1920s. Thiel (1981, p.
10) found only shell material in MRP 11 in a survey that spanned MRP 3
to 11 conducted during 1977 to 1980. Whitney et al. (1997, p. 12)
recorded a single individual during 1994-1995 in MRP 15, for a density
of 0.004 per square foot (sq. ft) (0.04 per square meter (sq. m)).
Helms (2008, p. 8) found eight live individuals and numerous shells
during a recent search of MRP 19, representing the most recent and
numerous collection of the species in the Mississippi River.
The spectaclecase is thought to be extant in at least four pools of
the Mississippi River mainstem, albeit in very low numbers. Records
include MRP 15 (Quad Cities area, Illinois and Iowa; in 1998), MRP 16
(Muscatine area, Iowa and Illinois in 1997), MRP 19 (Burlington area,
Illinois and Iowa in 2009), and MRP 22 (Quincy, Illinois and Hannibal,
Missouri, area in 1996). Populations may still persist in MRP 9 and 10
where specimens were found in the 1980s (Heath 2010a, pers. comm.).
Only a relic spectaclecase shell was found in MRP 3 above the St. Croix
River confluence in 2001, and none were found in subsequent surveys
(Kelner 2008, pers. comm.). In general, spectaclecase population levels
in the upper Mississippi River appear to have always been fairly small
and difficult to locate, and are now of questionable long-term
persistence.
St. Croix River: The northernmost and one of the three most
significant extant populations of the spectaclecase occurs in the St.
Croix River, Minnesota and Wisconsin. The population is primarily found
in the middle reaches of the river in Chisago and Washington Counties,
Minnesota, and Polk and St. Croix Counties, Wisconsin (river miles (RM)
17 to 118). Havlik (1994, p. 19) reported spectaclecase in the St.
Croix Wild River State Park portion of the river (approximately RM 62
to 65) and the reproducing population below the St. Croix Falls Dam at
St. Croix Falls, Wisconsin (dam located at approximately RM 52).
Additional survey work in the lower river at Afton State Park
(approximately RM 7 to 9) failed to find the spectaclecase (Havlik
1994, p. 19).
Hornbach (2001, p. 218) reported 68 live specimens from 4 of 16
river reaches. Relative abundance for the spectaclecase varied from
0.67 percent from RM 78 to 92 (20 live spectaclecase among 17 species
collected), 0.008 percent from RM 63 to 78 (41 live, 24
[[Page 3398]]
species), 0.0006 percent from RM 42 to 52 (6 live, 33 species), and
0.003 percent from RM 40 to 42 (1 live, 21 species). Reaches where the
spectaclecase is extant are fragmented by the pool formed from the
power dam at St. Croix Falls.
Baird (2000, p. 70) presented a length-frequency histogram for the
spectaclecase in the St. Croix River using data from an unpublished
1989 study. The 962 specimens were fairly evenly distributed over the
length scale, indicating multiple age classes including healthy numbers
of young spectaclecase recruiting into the population. Baird (2000, p.
70) used growth curves determined from his Missouri study of the
species to estimate the ages of spectaclecase of known size in the St.
Croix River. The percentage of newly recruited individuals (less than
or equal to 10 years of age) in the St. Croix was 40 percent--
considerably higher than that noted from the Gasconade (10.4 percent)
and Meramec (2.8 percent) Rivers in Missouri, two other streams with
abundant spectaclecase populations that he studied. The St. Croix
spectaclecase population, while among the largest known, may also be
the healthiest based on this metric. The spectaclecase is currently
distributed from RM 17 to 118 and appears to be recruiting from RM 17
to 54 (downstream of the St. Croix Falls Dam) (Heath 2008, pers.
comm.).
The long-term health of mussel populations in the St. Croix may be
in jeopardy, however. Hornbach et al. (2001, pp. 12-13) determined that
juvenile mussel density had suffered a statistically significant
decline at three of four lower St. Croix sites sampled in the 1990s and
in 2000. Zebra mussels also threaten the spectaclecase and other mussel
populations in the lower St. Croix River. A 2000 survey at 20 sites on
the lowermost 24 miles of the St. Croix River estimated that nearly one
percent of the mussels were infested with zebra mussels (Kelner & Davis
2002, p. 36).
Meramec River: The Meramec River flows into the Mississippi River
downstream of St. Louis in east-central Missouri. Its spectaclecase
population represents one of the best remaining rangewide. In the late
1970s, Buchanan (1980, p. 13) reported this species from 31 sites, 19
with live individuals. Live or fresh dead individuals occurred from RM
17.5 to 145.7. Buchanan (1980, p. 6) considered it to be common in the
lower 108 miles (174 km) of the Meramec River, but locally abundant
from RM 17.5 to 84. In 1997, Roberts and Bruenderman (2000, pp. 39,
44), using similar sampling methods as Buchanan (1980, pp. 4-5),
resurveyed the Meramec River system and collected spectaclecase from 23
sites, 19 of which had live individuals. They found the largest
populations between RM 56.7 and 118.8. Among 17 sites where
spectaclecase were found during both surveys, the species was less
abundant at nine sites and more abundant at five sites in 1997. At
three sites, only relic shells were found during both surveys. In the
1970s, Buchanan (1980, p. 10) reported finding 456 live individuals
among the 17 shared sites, whereas Roberts and Bruenderman (2000, p.
44) recorded only 198. A reduction in spectaclecase numbers (260 to 33)
at RM 59.5 accounted for most of the overall decrease in abundance
between the studies. Confounding the decrease in numbers among shared
survey sites, Roberts and Bruenderman (2000, p. 44) surveyed three
sites between RM 56.7 and 118.8 that were unsampled by Buchanan (1980,
pp. 1-69) and found 500, 538, and 856 live spectaclecase. The most
specimens found at a single site in the earlier study was 260 (RM
59.5). Currently, the population in the Meramec River stretches over
much of the mainstem, a distance of over 100 miles (161 km) from RM
18.5 to 120.4.
The spectaclecase represented 28 percent of all mussels sampled in
the Meramec River in 1997 (Roberts & Bruenderman 2000, p. 39). Baird
(2000, pp. 62, 68,77) extensively studied the demographics of the
Meramec River spectaclecase population in the late 1990s. The mean
estimated age of the population was 32 years. Individuals less than 10
years of age comprised only 2.8 percent of the Meramec population
sampled (a total of 2,983 individuals). At the four sites he
intentionally selected for their large spectaclecase populations,
densities ranged from 0.01 to 0.12 per sq. ft (0.1 to 1.3 per sq. m)
while estimated population numbers at these sites ranged from 933 to
22,697. Baird (2000, p. 71) thought that conditions for spectaclecase
recruitment in the Meramec had declined in the past 20 to 30 years, but
the causes were undetermined. The prevalence of larger adults in the
Meramec population may be cause for concern, as it appears to indicate
a low level of recruitment in the population.
Bourbeuse River: The Bourbeuse River is a northern tributary of the
Meramec River joining it at RM 68. Its spectaclecase population was
sampled in 1997 at a single site (RM 10.3), and 7 live individuals were
found (Roberts & Bruenderman 2000, p. 91). Sampling near the mouth (RM
0.4), Buchanan (1980, p. 16) found only relic shells. The Bourbeuse
population is probably dependent on the much larger Meramec population
for long-term sustainability.
Big River: Another Meramec tributary with a population of the
spectaclecase, the Big River flows northward into the Meramec River at
RM 38. The spectaclecase is only known from the lower end (RM 1.3),
where 14 live specimens were found in 1997 (Roberts & Bruenderman 2000,
p. 96). At RM 0.4, Buchanan (1980, p. 13) found only relic shells.
Similar to the Bourbeuse River population, the population in the Big
River is probably dependent on the much larger Meramec population for
sustainability. The Meramec River system, including the lower
Bourbeuse, lower Big, and Meramec River mainstems, can be considered a
single spectaclecase population cluster.
Lower Missouri River System
The spectaclecase was historically known from 10 streams in the
Missouri River system. Currently, only four of these streams are
thought to have extant populations.
Sac River: The Sac River is a large tributary to the Osage River.
The spectaclecase was considered extirpated in the 2002 status review
of the species (Butler 2002a). However, three old, live individuals
were collected at two sites during a survey of the Sac River in 2004
(Hutson & Barnhart 2004, p. 17). The same survey revealed ``numerous''
relic shells from six other sites, indicating that the spectaclecase
may have been relatively abundant at one time. Prior to the 2004
survey, the spectaclecase had not been collected from this river since
1978 (Bruenderman 2001, pers. comm.). Given the age of the live
individuals and the abundance of shell material, Hutson & Barnhart
(2004, p. 17) predicted the species would ``soon be extirpated'' from
the river.
Gasconade River: The Gasconade River is a southern tributary of the
Missouri River in south-central Missouri and flows into the mainstem
east of Jefferson City. When Stansbery (1970, p. 13) included this
species in the first compiled list of imperiled mussels, he noted that
``the only population of substantial size presently known is found in
the Gasconade River.'' In 1994, Buchanan found over 1,000 individuals
between RM 7 and 84 (Buchanan 1994, pp. 5, 8-13). Today, one of the
three best spectaclecase populations remaining rangewide occurs in the
Gasconade. The spectaclecase population occurs over approximately 200
miles (322 km) of the mainstem from RM 4.9 upstream (Bruenderman et al.
2001, p. 54). Baird (2000, pp. 61, 71) studied the demographics of the
Gasconade River spectaclecase
[[Page 3399]]
population in the late 1990s. Based on his limited number of sampling
sites, this species comprised about 20 percent of the entire mussel
fauna in this system. The mean estimated age of the population was 25
years. Individuals less than 10 years of age comprised 10.4 percent of
the Gasconade population sampled (n = 2,111), indicating a significant
level of recent recruitment.
Historically, Stansbery (1967, p. 29) noted that ``[t]he size of
some aggregation[s] * * * is impressive,'' and that ``the number of
individuals may reach a density of well over a dozen per square foot.''
Both statements are probably in reference to the Gasconade River,
Missouri, population, which he had described in the text of his note.
Densities at the four sites Baird (2000, pp. 61, 71) intentionally
selected for their large spectaclecase populations ranged from 0.03 to
0.06 per sq. ft (0.3 to 0.6 per sq. m); estimated population numbers at
these selected sites ranged from 2,156 to 4,766. Baird (2000, p. 71)
thought that conditions for spectaclecase recruitment in the Gasconade
River had declined in the past 20 to 30 years, but the causes were
undetermined.
Big Piney River: The Big Piney River, a southern tributary of the
Gasconade River, harbors a small population of the spectaclecase.
Although overlooked during a 1999 survey (Bruenderman et al. 2001, pp.
14, 28), 15 individuals were collected from the lower mainstem (RM 24)
in 2004 (Barnhart et al. 2004, p. 5). The status of the population is
unknown, but it is probably dependent on the much larger source
population in the Gasconade River for sustainability (McMurray 2008,
pers. comm.).
Osage Fork: The Osage Fork is a southwestern headwater tributary of
the Gasconade River. The spectaclecase is known from the lower portion
of this Gasconade River tributary, specifically from RM 13.9. Sampling
in the Osage Fork in 1999 yielded 26 live individuals from this site
(Bruenderman et al. 2001, p. 9). Relative abundance of the
spectaclecase in the Osage Fork was 3.9 percent, and catch-per-unit
effort was 1.3 per person-hour. This population is thought to be
stable, but it may also be dependent on the much larger source
population in the Gasconade River for long-term sustainability. The
Gasconade River system, including the lower Big Piney, lower Osage
Fork, and Gasconade mainstems, can be considered a single population
cluster.
Ohio River System
The spectaclecase's continued existence in the Ohio River is
extremely uncertain. Once known from five rivers, it has been
extirpated from two, and two of the remaining three are recently
represented by only one or two individuals each.
Ohio River: The Ohio River is the largest eastern tributary of the
Mississippi River, with its confluence marking the divide between the
upper and lower portions of the Mississippi River system. Historically,
the spectaclecase was documented from the Ohio River from the vicinity
of Cincinnati, Ohio, to its mouth. Although no specimens are known from
the mainstem upstream of Cincinnati, populations are known from two
upstream tributaries, the Muskingum and Kanawha Rivers. Nearly all
spectaclecase records from the Ohio River were made around 1900 or
before (Schuster 1988, p. 186). The only recent record is for a single
live individual found in an abandoned gill net near the Illinois shore
in 1994 (Cummings 2008, pers. comm.). If a population of the
spectaclecase continues to occur in the Ohio River, its future
persistence is extremely doubtful and continued existence seriously
threatened by the exotic zebra mussel.
Kanawha River: The Kanawha River is a major southern tributary of
the Ohio River that drains much of West Virginia. The spectaclecase was
not known from this stream until 2002, when a single, very old, live
individual was discovered near Glasgow, Kanawha County (Zimmerman 2002,
pers. comm.). Another live individual was found in the same vicinity in
2005, as well as two additional weathered shells in 2006 (Clayton
2008a, pers. comm.). This site is approximately 20 miles (32.2 km)
downstream of Kanawha Falls, below which is the only significant mussel
bed known from the Kanawha River. It is doubtful that a recruiting
spectaclecase population occurs in the Kanawha River due to the small
number of individuals found and their advanced age.
Green River: The Green River is a lower Ohio River tributary in
west-central Kentucky. The spectaclecase has been collected sparingly
in the Green River. That it was not reported in early collections made
in the system is indicative of the difficulty in finding specimens
(Price 1900, pp. 75-79). Stansbery (1965, p. 13) was the first to find
it in the mid-1960s at Munfordville, Hart County, where he reported 47
mussel species collected over a several-year period in the early 1960s.
More recently, from 1987 to 1989, Cicerello and Hannan (1990, p. 20)
reported single fresh dead specimens at six sites and relic specimens
from an additional five sites in Mammoth Cave National Park (MCNP). A
single specimen was recorded from MCNP, Edmonson County, in 1995.
Sampling conducted from 1996 to 1998 located fresh dead specimens at
two sites above MCNP, with a relic shell at a third site farther
upstream (Cicerello 1999, pp. 17-18). At least one fresh dead specimen
was reported from MCNP in 2001, as well as several live individuals in
2005 and 2006 (Layzer 2008, pers. comm.).
A small spectaclecase population remains in the upper Green River
from below Lock and Dam 5 upstream through MCNP, Edmonson County, into
western Hart County. Most recent specimens have been reported from the
upstream portion of this reach, where it is generally distributed from
MCNP upstream to western Hart County. Its distribution is much more
sporadic and localized in the lower portion of this reach due to the
pooling effect of two locks and dams (5 and 6). In 2001, a concerted
effort (approximately 15 person-hours) to locate rare mussels below
Lock and Dam 5 and at other sites downstream failed to find
spectaclecase (live or shell), although a fresh dead shell had been
collected in this area in 1993 (Cicerello 2008, pers. comm.). The
occurrence of variable-sized individuals in the 1990s indicates
different year classes but not necessarily recent recruitment
(Cicerello 2008, pers. comm.). The long-term sustainability of the
Green River population, primarily limited to an approximately 15-mile
(24-km) reach of the river, is therefore questionable, and its status
is unknown.
Cumberland River System
With few exceptions, most records of the spectaclecase in the
Cumberland River system were made before the 1920s. It was historically
known from the mainstem and four tributaries but appears currently to
be restricted to the lowermost Cumberland River a few miles above its
confluence with the Ohio River.
Cumberland River mainstem: The Cumberland River is a large southern
tributary of the lower Ohio River. The spectaclecase was considered
``not rare'' in the Cumberland River by Hinkley and Marsh (1885, p. 6),
whereas it was found at six sites by Wilson and Clark (1914, pp. 17,
19) during their survey primarily for commercial species in the
Cumberland River system. In a 1947-1949 survey of the Kentucky portion
of the upper Cumberland River, Neel and Allen (1964, p. 453) reported
live specimens only from one of six mainstem sites that they sampled
below Cumberland Falls. Neel and Allen (1964, p. 432) considered it to
be ``uncommon'' in the lower Cumberland River (where they did not
sample), a
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statement possibly based on its sporadic occurrence as reported by
Wilson and Clark (1914, pp. 17, 19). One of the last mainstem records
is that of a single live specimen found in the cold tailwaters of Wolf
Creek Dam, Kentucky, near the Tennessee border in 1982 (Miller et al.
1984, p. 108). This was one of only two live mussels found during a
survey of the dewatered river reach below the dam, the mussel community
having been eliminated from decades of cold water releases. The most
recent record is of a single live individual found at RM 10 below
Barkley Lock and Dam in 2008 (Fortenbery 2008, p. 9). A thorough search
of the area yielded no additional individuals.
Tennessee River System
The spectaclecase was originally known from the Tennessee River and
nine of its stream systems. Ortmann (1924, p. 60) reported that the
spectaclecase was ``frequent[hellip] in the upper Tennessee,'' while
acknowledging in an earlier paper (Ortmann 1918, p. 527) that it was
locally abundant in parts of the upper Tennessee River system, but
noted that it was ``generally regarded as a rare species'' rangewide.
Hundreds of miles of large river habitat on the Tennessee mainstem
have been converted under nine reservoirs, with additional dams
constructed in tributaries historically harboring this species (for
example, Clinch, Holston, and Elk Rivers). Watters (2000, p. 262)
summarizes the tremendous loss of mussel species from various reaches
of the Tennessee. The spectaclecase is now known only from the
Tennessee mainstem and three of its tributaries. Despite this fact, the
Tennessee River system continues to represent one of the last
strongholds of the spectaclecase rangewide.
Tennessee River mainstem: The Tennessee River is the largest
tributary of the Ohio River, draining portions of seven states. The 53-
mile (85-km) stretch of river in northwestern Alabama collectively
referred to as the Muscle Shoals historically harbored 69 species of
mussels, making it among the most diverse mussel faunas ever known
(Garner & McGregor 2001, p. 155). The historical spectaclecase
population in this reach was thought to be phenomenal given the amount
of historical habitat that was available and literature accounts of the
period. Hinkley (1906, p. 54), in 1904, considered the spectaclecase
``plentiful,'' noting 200 individuals under a single slab boulder.
Twenty years later, Ortmann (1925, p. 327) stated that ``this species
must be, or have been, abundant'' at Muscle Shoals based on the
``considerable number of dead shells'' he observed. In these quotes he
predicted the demise of the spectaclecase. The construction of three
dams (Wilson in 1925, Wheeler in 1930, Pickwick Landing in 1940)
inundated most of the historical habitat, leaving only small habitat
remnants (Garner & McGregor 2001, p. 155). The largest remnant habitat
remaining is the Wilson Dam tailwaters, a reach adjacent to and
downstream from Florence, Alabama.
With the exception of 1976-1978 when it was ``collected
infrequently'' from below Wilson Dam (Gooch et al. 1979, p. 90), no
collections of the spectaclecase were reported at Muscle Shoals from
1931 to 1995 despite surveys conducted in 1956-1957, 1963-1964, and
1991 (Garner & McGregor 2001, p. 156).
Elsewhere along the Tennessee mainstem, a specimen was recently
reported from the Guntersville Dam tailwaters in northern Alabama
(Butler 2002a, p. 17). From 1997-1999, 10 live, 1 fresh dead, a