67552 Federal Register / Vol. 75, No. 211 / Tuesday, November 2, 2010 / Proposed Rules 

DEPARTMENT OF THE INTERIOR 

Fish and Wildlife Service 

50 CFR Part 17 

[Docket No. FWS–R3–ES–2010–0019; MO 

92210–0–0008–B2] 

RIN 1018–AV96 

Endangered and Threatened Wildlife 

and Plants; Listing the Rayed Bean 

and Snuffbox as Endangered 

AGENCY: Fish and Wildlife Service, 

Interior. 

ACTION: Proposed rule. 

SUMMARY: We, the U.S. Fish and 

Wildlife Service (Service), propose to 

list the rayed bean (Villosa fabalis) and 

snuffbox (Epioblasma triquetra) as 

endangered throughout their ranges, 

under Endangered Species Act of 1973, 

as amended (Act). This proposed rule, if 

made final, would extend the Act’s 

protection to the rayed bean and the 

snuffbox. We have determined that 

designating 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 we 

receive on or before January 3, 2011. We 

must receive requests for public 

hearings, in writing, at the address 

shown in the FOR FURTHER INFORMATION

CONTACT section on or before December 

17, 2010. 

ADDRESSES: You may submit comments 

by one of the following methods: 

• Federal eRulemaking Portal: http:// 

www.regulations.gov. Follow the 

instructions for submitting comments 

on Docket No. FWS–R3–2010–0019. 

• U.S. mail or hand-delivery: Public 

Comments Processing, Attn: FWS–R3– 

2010–0019; Division of Policy and 

Directives Management; U.S. Fish and 

Wildlife Service; 4401 N. Fairfax Drive, 

Suite 222; Arlington, VA 22203. 

We will post all comments on http:// 

www.regulations.gov. This generally 

means that we will post any personal 

information you provide us (see the 

Public Comments section below for 

more information). 

FOR FURTHER INFORMATION CONTACT: 

Angela Boyer at the U.S. Fish and 

Wildlife Service, Ohio Ecological 

Services Field Office, 4625 Morse Road, 

Suite 104, Columbus, OH 43230; 

telephone 614–416–8993, ext. 22. 

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 listing 

determinations. We therefore 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 rayed bean and 

snuffbox as endangered. We particularly 

seek comments concerning: 

(1) Survey results for the rayed bean 

or snuffbox, as well as any studies that 

may show distribution, status, 

population size, or population trends, 

including indications of recruitment; 

(2) Pertinent aspects of life history, 

ecology, and habitat use of the rayed 

bean or snuffbox; 

(3) Current and foreseeable threats 

faced by the rayed bean or snuffbox, or 

both species, in relation to the five 

factors (as defined in section 4(a)(1) of 

the Act (16 U.S.C. 1531 et seq.)); 

(4) The specific physical and 

biological features to consider, and 

specific areas that may meet the 

definition of critical habitat and that 

should or should not be considered for 

a proposed critical habitat designation 

as provided by section 4 of the Act; and 

(5) The data and studies to which this 

proposal refers. 

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 

http://www.regulations.gov before 

midnight (Eastern Time) on the date 

specified in the DATES section. Finally, 

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 http:// 

www.regulations.gov. If you provide us 

personal identifying information such as 

your street address, phone number, or email address, 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 http://www.regulations.gov, or by 

appointment, during normal business 

hours at the Ohio Ecological Services 

Field Office (see FOR FURTHER

INFORMATION CONTACT). 

Public Hearing 

The Act provides for one or more 

public hearings on this proposal, if 

requested. We must receive requests by 

the date listed in the DATES section 

above. Such requests must be made in 

writing and addressed to the Field 

Supervisor of the Ohio Ecological 

Services Field Office (see FOR FURTHER

INFORMATION CONTACT). 

Background 

Species Descriptions 

The rayed bean is a small mussel 

usually less than 1.5 inches (in) (3.8 

centimeters (cm)) in length (Cummings 

and Mayer 1992, p. 142; Parmalee and 

Bogan 1998, p. 244; West et al. 2000, p. 

248). The shell outline is elongate or 

ovate in males and elliptical in females, 

and moderately inflated in both sexes, 

but more so in females (Parmalee and 

Bogan 1998, p. 244). The valves are 

thick and solid. The anterior end is 

rounded in females and bluntly pointed 

in males (Cummings and Mayer 1992, p. 

142). Females are generally smaller than 

males (Parmalee and Bogan 1998, p. 

244). Dorsally, the shell margin is 

straight, while the ventral margin is 

straight to slightly curved (Cummings 

and Mayer 1992, p. 142). The beaks are 

slightly elevated above the hingeline 

(West et al. 2000, p. 248), with sculpture 

consisting of double loops with some 

nodules (Parmalee and Bogan 1998, p. 

244). No posterior ridge is evident. 

Surface texture is smooth and subshiny, and green, yellowish-green, or 

brown in color, with numerous wavy, 

dark-green rays of various widths 

(sometimes obscure in older, blackened 

specimens) (Cummings and Mayer 1992, 

p. 142; West et al. 2000, p. 248). 

Internally, the left valve has two 

pseudocardinal teeth (tooth-like 

structures along the hinge line of the 

internal portion of the shell) that are 

triangular, relatively heavy, and large, 

and two short, heavy lateral teeth 

(Cummings and Mayer 1992, p. 142). 

The right valve has a low, triangular 

pseudocardinal tooth, with possibly 

smaller secondary teeth anteriorly and 

posteriorly, and a short, heavy, and 

somewhat elevated lateral tooth 

(Parmalee and Bogan 1998, p. 244). The 

color of the nacre (mother-of-pearl) is 

silvery white or bluish and iridescent 

posteriorly. Key characters useful for 

distinguishing the rayed bean from 

other mussels is its small size, thick 

valves, unusually heavy teeth for a 

small mussel, and color pattern 

(Cummings and Mayer 1992, p. 142). 

The snuffbox is a small- to mediumsized mussel with males reaching up to 

2.8 in. (7.0 cm) in length (Cummings 


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and Mayer 1992, p. 162; Parmalee and 

Bogan 1998, p. 108). The maximum 

length of females is about 1.8 in (4.5 cm) 

(Parmalee and Bogan 1998, p. 108). The 

shape of the shell is somewhat 

triangular (females), oblong, or ovate 

(males) with the valves solid, thick, and 

very inflated. The beaks are located 

somewhat anterior of the middle, 

swollen, turned forward and inward, 

and extended above the hingeline 

(Cummings and Mayer 1992, p. 162). 

Beak sculpture consists of three or four 

faint, double-looped bars (Cummings 

and Mayer 1992, p. 162; Parmalee and 

Bogan 1998, p. 108). The anterior end of 

the shell is rounded, and the posterior 

end is truncated, highly so in females. 

The posterior ridge is prominent, being 

high and rounded, while the posterior 

slope is widely flattened. The posterior 

ridge and slope in females is covered 

with fine ridges and grooves, and the 

posterioventral shell edge is finely 

toothed (Cummings and Mayer 1992, p. 

162). When females are viewed from a 

dorsal or ventral perspective, the 

convergence of the two valves on the 

posterior slope is nearly straight due to 

being highly inflated. This gives the 

female snuffbox a unique broadly 

lanceolate or cordate perspective when 

viewed at the substrate and water 

column interface (Ortmann 1919, p. 329; 

van der Schalie 1932, p. 104). The 

ventral margin is slightly rounded in 

males and nearly straight in females. 

Females have recurved denticles on the 

posterior shell margin that aid in 

holding host fish (Barnhart 2008, p. 1). 

The periostracum (external shell 

surface) is generally smooth and 

yellowish or yellowish-green in young 

individuals, becoming darker with age. 

Green squarish, triangular, or chevronshaped marks cover the umbone (the 

inflated area of the shell along the 

dorsal margin) but become poorly 

delineated stripes with age. Internally, 

the left valve has two high, thin, 

triangular, emarginate pseudocardinal 

teeth (the front tooth being thinner than 

the back tooth) and two short, strong, 

slightly curved, and finely striated 

lateral teeth. The right valve has a high, 

triangular pseudocardinal tooth with a 

single short, erect, and heavy lateral 

tooth. The interdentum (a flattened area 

between the pseudocardinal and lateral 

teeth) is absent, and the beak cavity is 

wide and deep. The color of the nacre 

is white, often with a silvery luster, and 

a gray-blue or gray-green tinge in the 

beak cavity. The soft anatomy was 

described by Oesch (1984, pp. 233–234), 

and Williams et al. (2008, p. 282). Key 

characters useful for distinguishing the 

snuffbox from other species include its 

unique color pattern, shape (especially 

in females), and high degree of inflation. 

Taxonomy 

The rayed bean is a member of the 

freshwater mussel family Unionidae and 

was originally described as Unio fabalis 

by Lea in 1831. The type locality is the 

Ohio River (Parmalee and Bogan 1998, 

p. 244), probably in the vicinity of 

Cincinnati, Ohio. Over the years, the 

rayed bean has been placed in the 

genera Unio, Margarita, Margaron, 

Eurynia, Micromya, and Lemiox. It was 

ultimately placed in the genus Villosa 

by Stein (1963, p. 19), where it remains 

today (Turgeon et al. 1998, p. 33). We 

recognize Unio capillus, U. lapillus, and 

U. donacopsis as synonyms of Villosa 

fabalis. 

The snuffbox is a member of the 

freshwater mussel family Unionidae and 

was described as Truncilla triqueter 

(Rafinesque 1820, p. 300). The species 

name was later changed to triquetra 

(Simpson 1900, p. 517), from the Latin 

triquetrous meaning ‘‘having three acute 

angles,’’ a reference to the general shape 

of the female. The type locality is the 

Falls of the Ohio (Ohio River, 

Louisville, Kentucky) (Parmalee and 

Bogan 1998, p. 108). The synonymy of 

the snuffbox was summarized by 

Johnson (1978, pp. 248–249), Parmalee 

and Bogan (1998, p. 108), and Roe (no 

date, p. 3). This species has also been 

considered a member of the genera 

Unio, Dysnomia, Plagiola, Mya, 

Margarita, Margaron, and Epioblasma at 

various times since its description. The 

monotypic subgenus Truncillopsis was 

created for this species (Ortmann and 

Walker 1922, p. 65). The genus 

Epioblasma was not in common usage 

until the 1970s (Stansbery 1973, p. 22; 

Stansbery 1976, p. 48; contra Johnson 

1978, p. 248), where it currently 

remains (Turgeon et al. 1998, p. 34). 

Unio triqueter, U. triangularis, U. 

triangularis longisculus, U. triangularis 

pergibosus, U. cuneatus, and U. 

formosus are recognized as synonyms of 

E. triquetra. Tricorn pearly mussel is 

another common name for this species 

(Clarke 1981a, p. 354). 

Life History 

The general biology of the rayed bean 

and the snuffbox are similar to other 

bivalved mollusks belonging to the 

family Unionidae. Adults are 

suspension-feeders, 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 

(Silverman et al. 1997, p. 1859; Nichols 

and Garling 2000, p. 873; Christian et al. 

2004, pp. 108–109; Strayer et al. 2004, 

pp. 430–431). Recent evidence suggests 

that adult mussels may also deposit-feed 

on particles in the sediment (Raikow 

and Hamilton 2001, p. 520). For their 

first several months, juvenile mussels 

employ foot (pedal) feeding, consuming 

settled algae and detritus (Yeager et al. 

1994, p. 221). Unionids have an unusual 

mode of reproduction. Their life cycle 

includes a brief, obligatory parasitic 

stage on fish. Eggs develop into 

microscopic larvae called glochidia 

within special gill chambers of the 

female. The female expels the mature 

glochidia, which must attach to the gills 

or the fins of an appropriate fish host to 

complete development. Host fish 

specificity varies among 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) and drop off as 

newly transformed juveniles. For further 

information on freshwater mussels, see 

Gordon and Layzer (1989, pp. 1–17). 

Mussel biologists know relatively 

little about the specific life-history 

requirements of the rayed bean and the 

snuffbox. Most mussels, including the 

rayed bean and snuffbox, have separate 

sexes. The age at sexual maturity, which 

is unknown for the rayed bean and 

snuffbox, is highly variable among and 

within species (0–9 years) (Haag and 

Staton 2003, pp. 2122–2123), and may 

be sex dependent (Smith 1979, p. 382). 

Both species are thought to be long-term 

brooders; rayed bean females brood 

glochidia from May through October 

(Parmalee and Bogan 1998, p. 108; 

Ecological Specialists, Inc. (ESI) 2000, p. 

5; Woolnough 2002, p. 23), and snuffbox 

brood glochidia from September to May 

(Ortmann 1912, p. 355; 1919, p. 327). 

The only published research identifies 

the Tippecanoe darter (Etheostoma 

tippecanoe) as a host fish for the rayed 

bean (White et al. 1996, p. 191). Other 

rayed bean hosts are thought to include 

the greenside darter (E. blennioides), 

rainbow darter (E. caeruleum), mottled 

sculpin (Cottus bairdi), and largemouth 

bass (Micropterus salmoides) 

(Woolnough 2002, p. 51). Based on 

inference of closely related species, 

additional hosts may be suitable, 

including other darter and sculpin 

species (Jones 2002, pers. comm.). 

Juvenile snuffbox have successfully 

transformed on logperch (Percina 

caprodes), blackside darter (P. 

maculata), rainbow darter, Iowa darter 

(E. exile), blackspotted topminnow 

(Fundulus olivaceous), mottled sculpin, 

banded sculpin (C. carolinae), Ozark 

sculpin (C. hypselurus), largemouth 


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bass, and brook stickleback (Culaea 

inconstans) in laboratory tests (Sherman 

1994, p. 17; Yeager and Saylor 1995, p. 

3; Hillegass and Hove 1997, p. 25; 

Barnhart et al. 1998, p. 34; Hove et al. 

2000, p. 30; Sherman Mulcrone 2004, 

pp. 100–103). 

Habitat Characteristics 

The rayed bean is generally known 

from smaller, headwater creeks, but 

occurrence records exist from larger 

rivers (Cummings and Mayer 1992, p. 

142; Parmalee and Bogan 1998, pp. 244). 

They are usually found in or near shoal 

or riffle areas, and in the shallow, wavewashed areas of glacial lakes, including 

Lake Erie (West et al. 2000, p. 253). In 

Lake Erie, the species is generally 

associated with islands in the western 

portion of the lake. Preferred substrates 

typically include gravel and sand. The 

rayed bean is oftentimes found among 

vegetation (water willow (Justicia 

americana) and water milfoil 

(Myriophyllum sp.)) in and adjacent to 

riffles and shoals (Watters 1988b, p. 15; 

West et al. 2000, p. 253). Specimens are 

typically buried among the roots of the 

vegetation (Parmalee and Bogan 1998, 

pp. 245). Adults and juveniles appear to 

produce byssal threads (thin, proteinbased fibers) (Woolnough 2002, pp. 99– 

100), apparently to attach themselves to 

substrate particles. 

The snuffbox is found in small to 

medium-sized creeks to larger rivers and 

in lakes (Cummings and Mayer 1992, p. 

162; Parmalee and Bogan 1998, p. 108). 

The species occurs in swift currents of 

riffles and shoals and wave-washed 

shores of lakes over gravel and sand 

with occasional cobble and boulders. 

Individuals generally burrow deep into 

the substrate except when spawning or 

attempting to attract a host (Parmalee 

and Bogan 1998, p. 108). 

Strayer (1999a, pp. 471–472) 

demonstrated in field trials that mussels 

in streams occur chiefly in flow refuges, 

or relatively stable areas that displayed 

little movement of particles during flood 

events. Flow refuges conceivably allow 

relatively immobile mussels to remain 

in the same general location throughout 

their entire lives. He thought that 

features commonly used in the past to 

explain the spatial patchiness of 

mussels (water depth, current speed, 

sediment grain size) were poor 

predictors of where mussels actually 

occur in streams. 

Rayed Bean Historical Distribution 

The rayed bean historically occurred 

in 112 streams, lakes, and some humanmade canals in 10 States: Illinois, 

Indiana, Kentucky, Michigan, New 

York, Ohio, Pennsylvania, Tennessee, 

Virginia, and West Virginia; and 

Ontario, Canada. The mussel occurred 

in parts of the upper (Lake Michigan 

drainage) and lower Great Lakes 

systems, and throughout most of the 

Ohio and Tennessee River systems. 

During historical times, the rayed bean 

was fairly widespread and locally 

common in many Ohio River system 

streams based on collections made over 

a several-decade period. The species 

was once fairly common in the Belle, 

South Branch Thames, Detroit, Scioto, 

Wabash, and Duck Rivers; several 

tributaries in the Scioto system 

(Olentangy River, and Big Darby and 

Alum Creeks); and Tippecanoe Lake 

based on literature and museum records 

(Call 1900; Watters 1994, p. 105; West 

et al. 2000, p. 251; Badra 2002, pers. 

comm.). The rayed bean was last 

reported from some streams several 

decades ago (North Branch Clinton, 

Auglaize, Ohio, West Fork, Beaver, 

Shenango, Mahoning, Mohican, Scioto, 

Green, Barren, Salamonie, White, Big 

Blue, Tennessee, Holston, South Fork 

Holston, Nolichucky, Clinch, North 

Fork Clinch, and Powell Rivers; Wolf, 

Conewango, Oil, Crooked, Pymatuning, 

Mill, Alum, Whetstone, Deer, Lick, and 

Richland Creeks; and Buckeye, 

Tippecanoe, Winona, and Pike Lakes). 

The rayed bean population in Lake Erie 

was once considerable (Ohio State 

University Museum of Biological 

Diversity (OSUM) collections), but has 

been eliminated by the zebra mussel. 

Rayed Bean Current Distribution 

Extant populations of the rayed bean 

are known from 28 streams and 1 lake 

in six States and one Canadian 

province: Indiana (St. Joseph River 

(stream) (Fish Creek (tributary)), 

Tippecanoe River (Lake Maxinkuckee, 

Sugar Creek)), Michigan (Black River 

(Mill Creek), Pine River, Belle River, 

Clinton River), New York (Allegheny 

River (Olean Creek, Cassadaga Creek, 

French Creek)), Ohio (Swan Creek, Fish 

Creek, Blanchard River, Tymochtee 

Creek, Walhonding River, Mill Creek, 

Big Darby Creek, Scioto Brush Creek), 

(Great Miami River, Little Miami River 

(East Fork Little Miami River), 

Stillwater River), Pennsylvania 

(Allegheny River (French Creek 

(Cussewago Creek))), and West Virginia 

(Elk River); and Ontario, Canada 

(Sydenham River, Thames River). 

Rayed Bean Population Estimates and 

Status 

Based on historical and current data, 

the rayed bean has declined 

significantly rangewide and is now 

known from only 28 streams and 1 lake 

(down from 112), a 74 percent decline 

(Table 1). This species has also been 

eliminated from long reaches of former 

habitat in hundreds of miles of the 

Maumee, Ohio, Wabash, and Tennessee 

Rivers and from numerous stream 

reaches and their tributaries. In 

addition, this species is no longer 

known from the States of Illinois, 

Kentucky, Tennessee, and Virginia. The 

rayed bean was also extirpated in West 

Virginia until the 2006 reintroduction 

into the Elk River (Clayton 2007, pers. 

comm.). 

In this proposed rule, mussel shell 

collection records have been classified 

according to the condition of shell 

material. Fresh dead (FD) shells still 

have flesh attached to the valves, they 

may or may not retain a luster to their 

nacre, and their periostracum is nonpeeling, all indicating relatively recent 

death (generally less than 1 year) 

(Buchanan 1980, p. 4). Relic (R) shells 

have lost the luster to their nacre, have 

peeling or absent periostracum, may be 

brittle or worn, and likely have been 

dead more than a year (Buchanan 1980, 

pp. 4–5; Zanatta et al. 2002, p. 482). 

Generally, FD shells indicate the 

continued presence of the species at a 

site (Metcalf 1980, p. 4). The presence 

of R shells only, along with repeated 

failure to find live (L) animals or FD 

shells, likely signifies that a population 

is extirpated (Watters and Dunn 1993– 

94, pp. 253–254). Shells labeled R may 

originally have been reported by 

collectors as either weathered dead (or 

weathered dry) or subfossil. If no details 

on shell condition were provided for a 

record, the shell is simply referred to as 

dead. In this document, a population is 

considered viable if it is reproducing 

and has enough individuals to sustain 

the population at its current level for the 

foreseeable future. 


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TABLE 1—RAYED BEAN STATUS AT HISTORICAL LOCATIONS

River basin Stream Last observed 

(R = relic) Current status Comments 

Upper Great Lakes 

Sub-basin.

Pigeon River ................................ 1996 (R) Extirpated ...................

Lower Great Lakes 

Sub-basin.

Black River ................................... 2001 Unknown .................... Small and of questionable viability. 

Mill Creek ..................................... 2002 Unknown .................... Unknown. 

Pine River .................................... 2002 Declining .................... Recruiting. 

Belle River .................................... 2003 Unknown ....................

Clinton River ................................ 1992 Unknown .................... Recruiting. 

North Fork Clinton River .............. 1933 Extirpated ...................

Sydenham River (Canada) .......... 2003 Stable ......................... Recruiting. 

Thames River ............................... 2008 Unknown .................... Unknown. 

Detroit River ................................. 1983 Extirpated ...................

Rouge River ................................. <1914 Extirpated.

Huron River .................................. 1931–32 Extirpated.

Raisin River .................................. 1941 Extirpated.

Macon Creek ................................ 1976–78 (R) Extirpated.

Maumee River .............................. 1913 Extirpated.

Swan Creek ................................. 2009 Stable ......................... Recruiting. 

St. Joseph River .......................... 1998 Declining .................... Probably not recruiting. 

West Branch St. Joseph River .... 1997 (R) Extirpated.

Fish Creek .................................... 2009 Declining .................... Unknown. 

Cedar Creek ................................. 1985 Extirpated.

Feeder Canal to St. Joseph River 1988 (R) Extirpated.

Auglaize River .............................. 1964 Extirpated.

Ottawa River ................................ 1998 (R) Extirpated.

Blanchard River ........................... 2009 Unknown .................... Recruiting. 

Sandusky River ............................ 1978 Extirpated.

Tymochtee Creek ......................... 1996 Unknown .................... Unknown. 

Wolf Creek ................................... 1971 (R) Extirpated.

Lake Erie ...................................... 1977–87 Extirpated.

Ohio River system ....... Ohio River mainstem ................... <1960 Extirpated.

Allegheny River ............................ 2007 Stable ......................... Recruiting. 

Chautauqua Lake outlet ............... <1919 Extirpated.

Chautauqua Lake ......................... <1919 Extirpated.

Olean Creek ................................. 2000 Unknown .................... Recruiting. 

Cassadaga Creek ........................ 1994 Unknown .................... Recruiting. 

Conewango Creek ....................... ∼1908 Extirpated.

Oil Creek ...................................... <1970 Extirpated.

French Creek ............................... 2005 Stable ......................... Recruiting. 

Cussewago Creek ........................ 1991 Unknown.

Crooked Creek ............................. ∼1908 Extirpated.

West Fork River ........................... <1913 Extirpated.

Beaver River ................................ ∼1910 Extirpated.

Shenango River ........................... ∼1908 Extirpated.

Pymatuning Creek ....................... ∼1908 Extirpated.

Mahoning River ............................ <1921 Extirpated.

Middle Island Creek ..................... 1980 (R) Extirpated.

Muskingum River ......................... 1980 (R) Extirpated.

Tuscarawas River ........................ ? Extirpated.

Walhonding River ......................... 1991–95 Declining .................... Probably not recruiting. 

Mohican River .............................. 1969 Extirpated.

Elk River ....................................... 2008 Reintroduced in 2006.

Scioto River .................................. 1964 Extirpated.

Mill Creek ..................................... 2007 Unknown.

Alum Creek .................................. 1970 Extirpated.

Blacklick Creek ............................ ? Extirpated.

Olentangy River ........................... 1962 Extirpated.

Whetstone Creek ......................... 1961 Extirpated.

Big Walnut Creek ......................... 1961 Extirpated.

Walnut Creek ............................... 1994 (R) Extirpated.

Big Darby Creek .......................... 2008 Declining .................... Unknown. 

Little Darby Creek ........................ 1990 (R) or 1986 (R) Extirpated.

Deer Creek ................................... 1981 Extirpated.

Sugar Creek ................................. <1900 Extirpated.

Scioto Brush Creek ...................... 1987 Unknown .................... Probably not recruiting. 

Cedar Creek ................................. ? Extirpated.

Buckeye Lake .............................. ? Extirpated.

Ohio and Erie Canal .................... ? Extirpated ...................

Great Miami River ........................ 2009 Unknown .................... Unknown. 

Little Miami River ......................... 1990–91 Unknown .................... Probably not recruiting. 

East Fork Little Miami River ........ 1990–91 Unknown.


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TABLE 1—RAYED BEAN STATUS AT HISTORICAL LOCATIONS—Continued 

River basin Stream Last observed 

(R = relic) Current status Comments 

Stillwater River ............................. 1987 Unknown .................... Probably not recruiting. 

South Fork Licking River ............. 1982 (R) Extirpated.

North Fork Elkhorn Creek ............ 1982 (R) Extirpated.

Eagle Creek ................................. 1981 (R) Extirpated.

Brashears Creek .......................... 1983 (R) Extirpated.

Green River .................................. 1964 Extirpated.

Nolin River ................................... 1983 (R) Extirpated.

Barren River ................................. <1900, ? Extirpated.

Wabash River .............................. 1962 (R) Extirpated.

Salamonie River ........................... 1971 Extirpated.

Mississinewa River ...................... 1994 (R) Extirpated.

Tippecanoe River ......................... 1995 Declining .................... Possibly recruiting. 

Tippecanoe Lake ......................... <1920 Extirpated.

Winona Lake ................................ 1934 Extirpated.

Pike Lake ..................................... 1906 Extirpated.

Lake Maxinkuckee ....................... 1997 Declining .................... Unknown. 

Vermilion River ............................. 1999 (R) Extirpated.

Salt Fork Vermilion River ............. 1956–57 Extirpated.

Middle Fork Vermilion River ........ 1991 Extirpated.

North Fork Vermilion River .......... 1995 (R) Extirpated.

Embarras River ............................ 1956 Extirpated.

Sugar Creek ................................. 1998 Unknown .................... Unknown. 

White River .................................. <1903 Extirpated.

West Fork White River ................. 1989–91 (R) Extirpated.

East Fork White River .................. ? Extirpated.

Big Blue River .............................. 1944 Extirpated.

Walnut Creek ............................... 1992 (R) Extirpated.

Mill Creek ..................................... 1992 (R) Extirpated.

Fall Creek ..................................... ? Extirpated.

Sugar Creek ................................. 1950 Extirpated.

Tennessee River system.Tennessee River mainstem ......... <1939 Extirpated.

Holston River ............................... 1914–15 Extirpated.

North Fork Holston River ............. 1913 Extirpated.

South Fork Holston River ............ 1914 Extirpated.

Nolichucky River .......................... 1968 Extirpated.

Lick Creek .................................... 1967 (R) Extirpated.

First Creek ................................... ? Extirpated.

Clinch River .................................. 1965 Extirpated.

North Fork Clinch River ............... <1921 Extirpated.

Powell River ................................. 1913–15 Extirpated.

Elk River ....................................... 1965 Extirpated.

Richland Creek ............................ 1892 Extirpated.

Duck River ................................... 1982 Extirpated.

Upper Great Lakes Sub-Basin 

The rayed bean was not known from 

the upper Great Lakes sub-basin until 

1996, when relic specimens were 

documented from a tributary to the St. 

Joseph River, a tributary to Lake 

Michigan. No extant populations of the 

rayed bean are currently known from 

this system. 

Lower Great Lakes Sub-Basin 

Of the 112 water bodies from which 

the rayed bean was historically 

recorded, 27 are in the lower Great 

Lakes system. The species is thought to 

be extant in 12 streams, which are 

discussed below, but historically 

significant populations have been 

eliminated from Lake Erie and the 

Detroit River. 

Black River—A tributary of the St. 

Clair River, linking Lakes Huron and St. 

Clair, the Black River is located in 

southeastern Michigan. Hoeh and Trdan 

(1985, p. 115) surveyed 17 sites in the 

Black River system, including 12 

mainstem sites, but failed to find the 

rayed bean. The rayed bean was not 

discovered there until the summer of 

2001 when a single live (L) individual 

was found in the lower river in the Port 

Huron State Game Area (PHSGA) (Badra 

2002, pers. comm.). A survey in 2003 

failed to find any rayed bean, and two 

surveys in 2005 found only two valves 

(Badra 2008, pers. comm.). An 

additional survey was performed in 

2005 at six sites, but no rayed bean were 

found (Badra 2008, pers. comm.). The 

status of this population cannot be 

accurately assessed at this time, but 

would appear to be small and of 

questionable viability (Butler 2002, p. 

8). 

Mill Creek—Mill Creek is a tributary 

of the Black River, St. Clair County, in 

southeastern Michigan. The rayed bean 

was discovered in Mill Creek in August 

2002. Five dead specimens were found 

approximately 0.5 miles (mi) (0.8 

kilometers (km)) above its confluence 

with the Black River in the PHSGA 

(Badra 2002, pers. comm.). A Mill Creek 

site 0.25 mi (0.4 km) from the 

confluence of the Black River was 

surveyed in 2003 and 2004 with one 

rayed bean shell found during each 

survey (Badra 2008, pers. comm.). 

Similar to the population in the Black 

River, the status of this newly 

discovered population cannot be 

accurately assessed at this time. 


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Pine River—Another tributary of the 

St. Clair River, the Pine River is located 

in southeastern Michigan. The rayed 

bean was apparently not collected in the 

Pine River until 1982 when specimens 

were found at three sites (Hoeh and 

Trdan 1985, p. 116). These collections 

included 5 L individuals and 23 FD 

specimens (Badra 2002, pers. comm.). 

Hoeh and Trdan (1985, p. 116) 

considered it to be ‘‘rare,’’ semiquantitatively defined as occurring at a 

rate of less than one specimen per 

person-hour sampling effort. In 1997, 

two L individuals were found. The last 

survey in the Pine River occurred in 

2002 (Badra 2008, pers. comm.), and 

one L rayed bean was documented 

(Badra and Goforth 2003, p. 6). The 

species may have declined significantly 

since the 1980s, but is probably still 

viable in the Pine River. 

Belle River—The Belle River is a third 

tributary of the St. Clair River harboring 

an extant population of the rayed bean. 

This species was first collected from the 

Belle River in 1965, when 17 FD 

specimens were collected (OSUM 

1965:0106). The same site was revisited 

in 1978, but only one FD shell is 

represented in OSUM 1978:0013. Since 

that time, L individuals or FD 

specimens have been found in 1983 and 

1992, while only R shells were found in 

1994 (Badra 2008, pers. comm.). During 

summer 2002 sampling, single L 

specimens were found at two new sites, 

with an additional four and two FD 

specimens, respectively, also found 

from these sites (Badra 2008, pers. 

comm.). The status of the population is 

still not well known, but appears to be 

small. 

Clinton River—The rayed bean was 

first recorded from the Clinton River in 

1933 (Badra 2008, pers. comm.). The 

mussel fauna in the entire mainstem of 

the Clinton River downstream of 

Pontiac, Michigan, was apparently 

wiped out by pollution between 1933 

and 1977 (Strayer 1980, p. 147). In 1992, 

Trdan and Hoeh (1993, p. 102) found 26 

L individuals using a suction dredge 

from a bridge site slated for widening 

where Strayer (1980, p. 146) found only 

R shells. The rayed bean represented 1.2 

percent relative abundance of the 10 

species collected at the site. The 

population is probably viable but 

currently restricted to about 3 mi (4.8 

km) of stream in the western suburbs of 

Pontiac. Its long-term status appears to 

be highly precarious. 

Sydenham River—The rayed bean in 

the Sydenham River represents one of 

the largest rayed bean populations 

remaining. West et al. (2000, pp. 252– 

253) presented a highly detailed 

collection history of the rayed bean in 

the Sydenham River. The rayed bean is 

currently thought to exist in an 

approximately 75-mi (120-km) reach of 

the middle Sydenham, from the general 

vicinity of Napier, Ontario, downstream 

to Dawn Mills. The species appears to 

be most abundant in the lower half of 

this river reach. Although the range has 

remained relatively consistent over 

time, abundance data at repeatedly 

sampled sites from the 1960s to the late 

1990s indicate a general decline of the 

rayed bean. Based on the range of sizes 

and roughly equal number of specimens 

in various size classes of the L and FD 

material they gathered, West et al. 

(2000, p. 256) considered the population 

to be ‘‘healthy’’ and ‘‘reproducing’’ 

(recruiting). Data from sampling in 2001 

shows evidence of recruitment and 

variable size classes for both sexes from 

most of the sites (Woolnough 2002, p. 

50). Based on this data, the rayed bean 

population in the Sydenham River is 

doing considerably better than West et 

al. (2000, pp. 252–253) suggested. 

Woolnough and Morris (2009, p. 19) 

estimates that there are 1.5 million 

mature rayed bean in the Sydenham 

River living in the 38-mile (61-km) 

stretch between Napier Road near 

Alvinston, Ontario, and Dawn Mills, 

Ontario. 

Thames River—The Thames River 

flows west through southwestern 

Ontario. The rayed bean was historically 

known from only the south branch until 

2008, when it was discovered in the 

north branch. In July 2008, six gravid 

(full of eggs) females were collected at 

two north branch sites (Woolnough 

2008, pers. comm.). In September 2008, 

four L females and two L males were 

collected at two different north branch 

sites (Woolnough 2008, pers. comm.). 

All of these individuals were collected 

within a 4.5-mi (7.2-km) reach of the 

river (Woolnough 2008, pers. comm.). 

Woolnough and Morris (2009, p. 19) 

estimates that there are 4,300 mature 

rayed bean in the Thames River. 

Maumee River System—The Maumee 

River system, which flows into the 

western end of Lake Erie, was once a 

major center of distribution of the rayed 

bean. The species was historically 

known from eight streams in the system 

in addition to the mainstem Maumee. 

Further, an additional population was 

discovered in the system in 2005 in 

Swan Creek. 

Swan Creek—Swan Creek is a 

tributary of the lower Maumee River in 

northwestern Ohio. This population was 

discovered in 2005. Surveys conducted 

in 2006 and 2007 found that the Swan 

Creek population is limited to about 3 

river mi (5 river km) between river mile 

(RM) 18.3 and 15.3 (Grabarkiewicz 

2008, p. 11). The rayed bean was the 

fourth most abundant unionid present 

within the 2006–2008 sample area, 

reaching densities of eight individuals 

per square meter in some areas and 

comprising about 14.1 percent of the 

total mussel community (Grabarkiewicz 

2008, p. 10). The rayed bean population 

in Swan Creek is viable and, although 

limited to a short reach, may be one of 

the most robust remaining populations. 

St. Joseph River—The St. Joseph River 

is one of the two major headwater 

tributaries to the Maumee, with a 

drainage area in southeastern Michigan, 

northwestern Ohio, and northeastern 

Indiana. The mainstem flows in a 

southwesterly direction to its 

confluence with the St. Mary’s River to 

form the Maumee in Ft. Wayne, Indiana. 

The rayed bean was historically known 

from numerous sites on the river, but 

now apparently persists only at a couple 

of sites in the lower St. Joseph River in 

Allen and DeKalb Counties, Indiana 

(Watters 1988b, p. 15; 1998, Appendix 

C); a few FD specimens were found in 

both studies, but no live individuals 

were found. Grabarkiewicz and Crail 

(2008, p. 13) surveyed six sites on the 

West Branch St. Joseph River in 2007, 

but did not encounter any rayed bean. 

Fish Creek—A tributary of the St. 

Joseph River that begins in Ohio, Fish 

Creek flows west then south through 

Indiana, then eventually east into Ohio 

before joining the St. Joseph River at 

Edgerton. The rayed bean persists in 

Williams County, Ohio, and possibly 

DeKalb County, Indiana. Based on the 

appearance of 2 L individuals and FD 

shells, it inhabits the lower 10 mi (16.1 

km) or less of the stream (Watters 1988b, 

p. 18; Grabarkiewicz 2009, pers. 

comm.). Watters (1988b, p. ii) 

considered Fish Creek to be ‘‘the most 

pristine tributary of the St. Joseph 

system.’’ A major diesel fuel spill from 

a ruptured pipeline in DeKalb County in 

1993 resulted in a mussel kill in the 

lower portion of the stream (Sparks et 

al. 1999, p. 12). It is not known if the 

rayed bean was affected by the spill. 

Surveys in 2004 (at 64 qualitative sites) 

and 2005 (at 11 quantitative sites) failed 

to detect the species (Brady et al. 2004, 

p. 2; 2005, p. 3). However, 

Grabarkiewicz (2009, pers. comm.) 

reported finding two L and three FD 

rayed bean in 2005 at the County Road 

3 bridge in Ohio. In 2009, two FD rayed 

bean were found in lower Fish Creek in 

Ohio (personal observation). The 

viability and status of this population is 

uncertain (Fisher 2008, pers. comm.). 

Blanchard River—The Blanchard 

River is a tributary of the Auglaize River 

in the Maumee River system, in 

northwestern Ohio. First discovered in 


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1946, this population is one of the 

largest of the rayed bean rangewide. The 

rayed bean in the Blanchard River is 

restricted to 25–30 river mi (40–48 river 

km) in the upper portion of the stream 

in Hardin and Hancock Counties 

upstream of Findley (Hoggarth et al. 

2000, p. 22). Hoggarth et al. (2000, p. 23) 

reported the rayed bean to be the fourth 

most common species in the drainage. 

The population is considered to be 

viable. 

Tymochtee Creek—Tymochtee Creek 

is a tributary to the upper Sandusky 

River in north-central Ohio, which 

flows into the southwestern portion of 

Lake Erie. The rayed bean is known 

from three sites in a reach of stream in 

Wyandot County and was first collected 

in 1970. All collections of the rayed 

bean have been small, with not more 

than five FD shells found in any one 

collection effort. The last record is for 

1996, when a pair and three unpaired 

valves were collected. The condition of 

at least one of the valves indicated that 

the rayed bean is probably still extant in 

the stream, although no L individuals 

were observed (Athearn 2002, pers. 

comm.). The rayed bean status in 

Tymochtee Creek is therefore currently 

unknown. 

Ohio River System 

The rayed bean was historically 

known from the Ohio River in the 

vicinity of Cincinnati, Ohio, 

downstream to the Illinois portion of the 

river. It undoubtedly occurred 

elsewhere in the upper mainstem. Few 

historical records are known (mostly 

circa 1900), and no recent collections 

have been made, indicating that it 

became extirpated there decades ago. It 

was historically known from 71 streams, 

canals, and lakes in the system, 

representing roughly two-thirds of its 

total range. Ortmann (1925, p. 354) 

considered the rayed bean to be 

‘‘abundant in small streams’’ in the Ohio 

River system. Currently, only 16 streams 

and a lake are thought to have extant 

rayed bean populations in the system. 

Allegheny River System—Nine 

streams and Chautauqua Lake 

historically harbored rayed bean 

populations in the Allegheny River 

system. Currently, the rayed bean is 

found in half of these water bodies, but 

in good numbers in two streams 

(Allegheny River and French Creek) in 

this drainage. 

Allegheny River—The Allegheny 

River drains northwestern Pennsylvania 

and western New York joining the 

Monongahela River at Pittsburgh, 

Pennsylvania, to form the Ohio River. 

Ortmann (1909a, p. 179; 1919, p. 262) 

was the first to report the rayed bean 

from the Allegheny. The population 

once stretched from Cataraugus County, 

New York, to Armstrong County, 

Pennsylvania. Based on historical 

collections, it appears that the rayed 

bean is more abundant now than it was 

historically in the Allegheny River. This 

may indicate that the rayed bean 

population in the Allegheny has 

expanded in the past 100 years. Many 

streams in western Pennsylvania have 

improved water quality since Ortmann’s 

time, when he reported on the 

wholesale destruction of mussels in 

several streams (Ortmann 1909b, pp. 

11–12). It currently occurs in 

Pennsylvania downstream of Allegheny 

(Kinzua) Reservoir in Warren County to 

the pool of Lock and Dam 8 in northern 

Armstrong County, a distance of over 

100 river mi (161 river km) (Villella 

Bumgardner 2008, pers. comm.). The 

Allegheny population is viable and one 

of the most important remaining 

rangewide today. 

Olean Creek—Olean Creek is a 

tributary of the Allegheny River in 

western New York. A small population 

of the rayed bean is known from the 

lower portions of the stream. Strayer et 

al. (1991, p. 67) reported the rayed bean 

from three sites during 1987–90 

sampling, although just one L 

individual was located with R shells 

from the other two sites. Only R shells 

were found in Olean Creek in 1994, but 

three L individuals were found in 2000, 

at the proposed construction site of the 

City of Olean Water Treatment Plant 

(ESI 2000, p. 8). Collected only during 

their quantitative sampling effort, the 

rayed bean represented a relative 

abundance of 11.5 percent of the seven 

L species sampled. The rayed bean age 

distribution of these specimens also 

indicates recent recruitment into the 

population (ESI 2000, p. 9). Relic 

specimens are now known from an 8-mi 

(13-km) reach of stream, with L 

individuals known from less than 1.5 mi 

(2.4 km) of the lower creek. The Olean 

Creek population appears viable, but is 

small and tenuous (Butler 2008, pers. 

comm.) 

Cassadaga Creek—Cassadaga Creek is 

a tributary of Conewango Creek in the 

Allegheny River system, in western New 

York. A small population of the rayed 

bean is known from a single riffle (Ross 

Mills) in the lower creek north of 

Jamestown. Four L specimens were 

found in 1994 (Strayer 1995). Muskrat 

middens collected during the winter of 

2002 produced 38 FD specimens with a 

size range of 0.8–1.7 in (2.0–4.3 cm) 

(Clapsadl 2002, pers. comm.). Although 

the rayed bean is not known from other 

sites in the stream, it appears to be 

viable at this site. The highly restricted 

extent of the population combined with 

its proximity to roads and retail 

development, including a gas station 

close to the flood zone upstream, makes 

it extremely susceptible to a stochastic 

event (such as a toxic chemical spill). 

French Creek—French Creek is a 

major tributary of the middle Allegheny 

River, in western New York and 

northwestern Pennsylvania. One of the 

largest rayed bean populations known is 

found in much of the lower portions of 

the stream in four Pennsylvania 

counties (the species is not known from 

the New York portion of stream). 

Ortmann (1909a, p. 188; 1919, p. 264) 

reported the species from two counties, 

Crawford and Vanango. Not until circa 

1970 did the population become more 

thoroughly known, with museum lot 

sizes indicating sizable populations at 

several sites, particularly in the lower 

reaches of the stream. Recent collections 

indicate that population levels remain 

high with the rayed bean occurring 

throughout the mainstem (Villella 

Bumgardner 2002, pers. comm.; Smith 

and Crabtree 2005, pp. 15–17; 

Enviroscience 2006, p. 5). 

Cussewago Creek—Cussewago Creek 

is a tributary of lower French Creek, 

with its confluence at Meadville, 

Crawford County, Pennsylvania. A 

small population was reported in 1991 

from Cussewago Creek (Proch 2001, 

pers. comm.). The rayed bean is thought 

to persist in the stream, but its current 

status is unknown. 

Walhonding River—The Walhonding 

River is a tributary of the upper 

Muskingum River system, in central 

Ohio, forming the latter River at its 

confluence with the Tuscarawas River at 

Coschocton. Small numbers of rayed 

bean shells are represented in OSUM 

collections from the 1960s and 1970s. 

During 1991–93, Hoggarth (1995–96, p. 

161) discovered one L individual and 

one FD specimen at one site, while four 

R specimens were found at three other 

sites. A small rayed bean population is 

thought to remain in the Walhonding 

River; its status is unknown, but is 

deemed highly tenuous given the small 

population size. The population is 

probably nearing extirpation (Hoggarth 

2008a, pers. comm.). 

Elk River—The Elk River is a major 

181-river-mi (291-river-km) tributary in 

the lower Kanawha River system 

draining central West Virginia and 

flowing west to the Kanawaha River at 

Charleston. The rayed bean was 

extirpated in the Elk River sometime in 

the 1990s. In 2006 and 2007, 

approximately 600 adults were 

reintroduced into the Elk River above 

Clendenin. In 2008, an effort was made 

to monitor the reintroduction. A 30- 


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minute search yielded two L 

individuals, but efforts were 

discontinued due to high water and 

excessive habitat disturbance caused by 

the search effort (Clayton 2008, pers. 

comm.). The translocated adults are 

thought to persist in the stream, but it 

is unknown if this new population is 

reproducing. 

Scioto River system—The Scioto River 

system, in central and south-central 

Ohio, is a major northern tributary of 

the Ohio River. A historically large 

meta-population of the rayed bean 

occupied at least 11 streams, the Ohio 

and Erie Canal, and Buckeye Lake. 

Sizable populations were noted in at 

least the Olentangy River, and Alum 

and Big Darby Creeks, based on OSUM 

collections primarily from the 1960s. A 

series of system reservoirs mostly north 

of Columbus reduced habitat and 

contributed to the elimination of some 

populations in several streams (Alum, 

Big Walnut, and Deer Creeks; Olentangy 

and Scioto Rivers). The location of the 

Columbus Metropolitan Area in the 

heart of the watershed has also taken a 

major toll on the species. The historical 

Scioto rayed bean meta-population has 

since been decimated by anthropogenic 

factors. Currently, remnant populations 

are known only from Mill Creek, Big 

Darby Creek, and Scioto Brush Creek. 

Mill Creek—Mill Creek is a tributary 

of the Scioto River in central Ohio that 

joins the Scioto River at the 

O’Shaughnessy Reservoir northwest of 

the City of Columbus. In 2004, seven FD 

specimens were found during a survey 

in the City of Marysville (Hoggarth 

2005, p. 7). In 2007, Hoggarth (2007a, 

pp. 5–6) found two L rayed bean at the 

same site and one L individual at an 

additional site. No other information is 

available on the status of this 

population. 

Big Darby Creek—Big Darby Creek is 

one of the major tributaries draining the 

northwestern portion of the Scioto River 

system in central Ohio. A sizable rayed 

bean population was noted in Big Darby 

Creek from OSUM collections primarily 

from the 1960s. Watters (1994, p. 105) 

reported finding a few FD specimens in 

1986, but none in 1990, and indicated 

that the rayed bean was probably 

extirpated from Big Darby Creek. In 

2006, one L individual was found at the 

U.S. Highway 42 bridge replacement 

project site (Hoggarth 2006, p. 6). This 

individual was relocated to a site 

upstream out of the impact zone of the 

bridge project, and nine additional L 

individuals were subsequently found at 

the relocation site (Hoggarth 2006, p. 6). 

In 2007, three L rayed bean were found 

at the relocation site (Hoggarth 2007b, p. 

9). Hoggarth (2008b, pers. comm.) 

visited the same relocation site in 2008, 

and reported finding ‘‘numerous living 

specimens’’ of the rayed bean. The status 

of this population cannot be accurately 

assessed at this time, but would appear 

to be small and of questionable viability. 

Scioto Brush Creek— Scioto Brush 

Creek is a small western tributary of the 

lower Scioto River in Scioto County, 

south-central Ohio. Watters (1988a, p. 

45) discovered the rayed bean in this 

stream in 1987, reporting two FD and 

two R specimens from a site, and a R 

specimen from a second site among the 

20 sites he collected. This population’s 

current status is uncertain. 

Great Miami River—The Great Miami 

River is a major northern tributary of the 

Ohio River in southwestern Ohio that 

originates from Indian Lake in westcentral Ohio and flows into the Ohio 

River west of Cincinnati. The 

occurrence of the rayed bean in the 

Great Miami River was discovered in 

August 2009, during a mussel survey for 

a bridge project in Logan County, Ohio. 

Only one individual was documented, a 

male approximately 7 to 8 years of age 

(Hoggarth 2009, pers. comm.). The 

status of this newly discovered 

population is not known. 

Little Miami River—The Little Miami 

River is a northern tributary of the Ohio 

River in southwestern Ohio, flowing 

into the latter at the eastern fringe of the 

Cincinnati metropolitan area. Hoggarth 

(1992, p. 248) surveyed over 100 sites in 

the entire system. He found one L 

individual at a site in Warren County 

and possibly a subfossil shell at another 

site, although there is contradictory data 

in his paper (Butler 2002, p. 17). The 

latter site may have been the same as 

that reported for a pre-1863 record 

(Hoggarth 1992, p. 265). The rayed bean 

appears to be very rare in the Little 

Miami, having been found extant at only 

1 of 46 mainstem sites. Hoggarth (1992, 

p. 267) highlighted the ‘‘fragile nature’’ 

of the extant mussel community in the 

system, while noting that localized 

reaches of the Little Miami were 

‘‘severely impacted.’’ The species status 

in the river is uncertain, but apparently 

very tenuous and probably headed 

toward extirpation (Butler 2002, p. 17). 

East Fork Little Miami River—The 

East Fork Little Miami River is an 

eastern tributary of the lower Little 

Miami River, with its confluence at the 

eastern fringe of the Cincinnati 

metropolitan area. According to OSUM 

records, eight FD specimens were 

reported from a site in eastern Clermont 

County in 1973. Hoggarth (1992, p. 265) 

reported one L, three FD, and one R 

rayed bean from three sites in a 7-rivermi (11-river-km) stretch of the stream in 

western Clermont and adjacent Brown 

County (including the 1973 site). Harsha 

Reservoir on the East Fork destroyed 

several miles of potential stream habitat 

for the rayed bean a few miles 

downstream of the extant population. 

The status of the rayed bean in the river 

is uncertain but probably of doubtful 

persistence (Butler 2002, p. 17). 

Stillwater River—The Stillwater River 

is a western tributary of the middle 

Great Miami River in southwestern 

Ohio. The rayed bean is known from 

two specimens, one FD and one R, 

collected in 1987 at two sites spanning 

the Miami–Montgomery County line 

(OSUM records). Both sites occur in the 

footprint of Englewood Reservoir 

(constructed circa 1920), which serves 

as a retarding basin (a constructed 

empty lake used to absorb and contain 

flooding in periods of high rain) that is 

normally a free-flowing river except in 

times of flood, therefore continuing to 

provide riverine habitat that is normally 

destroyed by permanently impounded 

reservoirs. The rayed bean in the 

Stillwater River may be extant, but its 

status is currently unknown and 

considered highly imperiled. 

Tippecanoe River—The Tippecanoe 

River is a large northern tributary of the 

middle Wabash River in north-central 

Indiana. The first records for the rayed 

bean date to circa 1900 (Daniels 1903, 

p. 646). Historically, this species was 

known from numerous sites in six 

counties in the Tippecanoe River. A 

total of 12 FD specimens from 5 of 30 

sites were found when sampled in 1992. 

The rayed bean ‘‘is apparently on the 

decline’’ in the river (ESI 1993, p. 87). 

The Tippecanoe rayed bean population 

was thought to be recruiting by Fisher 

(2008, pers. comm.), but appears 

tenuous and its long-term viability is 

questionable. 

Lake Maxinkuckee—Lake 

Maxinkuckee is a glacial lake in the 

headwaters of the Tippecanoe River in 

north-central Indiana. The rayed bean 

has been known from the lake for more 

than a century (Blatchley 1901). A 1997 

OSUM record included seven FD 

specimens collected at its outlet to the 

Tippecanoe River. Fisher (2002, pers. 

comm.), who made the 1997 OSUM 

collection, noted that many native 

mussels had zebra mussels attached to 

their valves and were apparently 

contributing to their mortality. The 

status of the rayed bean in Lake 

Maxinkuckee is therefore highly 

tenuous, and its long-term persistence 

questionable. 

Sugar Creek—Sugar Creek is a 

tributary of the East Fork White River, 

in the lower Wabash River system in 

south-central Indiana. A rayed bean 

population was first reported there in 


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1930. Harmon (1992, p. 33) sampled 27 

mainstem and 16 tributary sites finding 

FD specimens at 3 mainstem sites and 

R specimens from 2 other sites. The 

sites with FD material were found in the 

lowermost 6 mi (9.7 km) of stream. The 

status and viability of this tenuous 

population is uncertain (Fisher 2008, 

pers. comm.). 

Tennessee River System 

Historically, the rayed bean was 

known from the Tennessee River and 12 

of its tributary streams. Ortmann (1924, 

p. 55) reported that the rayed bean had 

a ‘‘rather irregular distribution’’; 

however, museum lots show that it was 

fairly common in some streams (North 

Fork Clinch, Duck Rivers). The last L 

rayed bean records from the system, 

with the exception of the Duck River, 

were from the 1960s or earlier. The 

species held on in the Duck until the 

early 1980s. Recent intensive sampling 

in the Duck watershed has failed to 

locate even a R shell of the rayed bean 

(Ahlstedt et al. 2004, p. 29). Tributaries 

in this system have been extensively 

sampled over the past 25 years. It is 

highly probable that this species is 

extirpated from the entire Tennessee 

River system. 

A project was initiated in 2008 to 

reintroduce rayed bean into the Duck 

River by translocating over 1,000 adults 

from the Allegheny River system. 

Although the rayed bean was extirpated 

from the Duck River about 25 years ago, 

major improvements in water quality 

and physical habitat conditions have 

occurred in the past 15 years. In 

response to these improvements, 

recruitment of nearly all extant mussel 

species has been documented and 

suggests that reintroduction of the rayed 

bean might be successful (Anderson 

2008, pers. comm.). The reintroduction 

has not yet occurred. 

The information presented in this 

document indicates that the rayed bean 

has experienced a significant reduction 

in range and most of its populations are 

disjunct, isolated, and with few 

exceptions, appear to be declining (West 

et al. 2000, p. 251). The extirpation of 

this species from over 80 streams and 

other water bodies within its historical 

range indicates that substantial 

population losses have occurred. 

Relatively few streams are thought to 

harbor sizable viable populations 

(Sydenham, Swan, Blanchard, and 

Allegheny Rivers, and French Creek). 

Small population size and restricted 

stream reaches of current occurrence are 

a real threat to the rayed bean due to the 

negative genetic aspects associated with 

small, geographically isolated 

populations. This can be especially true 

for a species, like rayed bean, that was 

historically widespread and had 

population connectivity among 

mainstem rivers and multiple 

tributaries. The current distribution, 

abundance, and trend information 

illustrates that the rayed bean is 

imperiled. 

Snuffbox Historical Distribution 

The snuffbox historically occurred in 

208 streams and lakes in 18 States and 

1 Canadian province: Alabama, 

Arkansas, Illinois, Indiana, Iowa, 

Kansas, Kentucky, Michigan, 

Minnesota, Mississippi, Missouri, New 

York, Ohio, Pennsylvania, Tennessee, 

Virginia, West Virginia, and Wisconsin; 

and Ontario. The major watersheds of 

historical streams and lakes of 

occurrence include the upper Great 

Lakes sub-basin (Lake Michigan 

drainage), lower Great Lakes sub-basin 

(Lakes Huron, Erie, and Ontario 

drainages), upper Mississippi River subbasin, lower Missouri River system, 

Ohio River system, Cumberland River 

system, Tennessee River system, lower 

Mississippi River sub-basin, and White 

River system. 

Snuffbox Current Distribution 

Extant populations of the snuffbox are 

known from 74 streams in 14 States and 

1 Canadian province: Alabama 

(Tennessee River, Paint Rock River, and 

Elk River), Arkansas (Buffalo River, 

Spring River, and Strawberry River), 

Illinois (Kankakee River and Embarras 

River), Indiana (Pigeon River, Salamonie 

River, Tippecanoe River, Sugar Creek, 

Buck Creek, Muscatatuck River, and 

Graham Creek), Kentucky (Tygarts 

Creek, Kinniconick Creek, Licking 

River, Slate Creek, Middle Fork 

Kentucky River, Red Bird River, Red 

River, Rolling Fork Salt River, Green 

River, and Buck Creek), Michigan 

(Grand River, Maple River, Pine River, 

Belle River, Clinton River, Huron River, 

Davis Creek, South Ore Creek, and 

Portage River), Minnesota (St. Croix 

River), Missouri (Meramec River, 

Bourbeuse River, St. Francis River, and 

Black River), Ohio (Grand River, Ohio 

River, Muskingum River, Walhonding 

River, Killbuck Creek, Olentangy River, 

Big Darby Creek, Little Darby Creek, Salt 

Creek, Scioto Brush Creek, South Fork 

Scioto Brush Creek, Little Miami River, 

and Stillwater River), Pennsylvania 

(Allegheny River, French Creek, West 

Branch French Creek, Le Boeuf Creek, 

Muddy Creek, Conneaut Outlet, Little 

Mahoning Creek, Dunkard Creek, 

Shenango River, and Little Shenango 

River), Tennessee (Clinch River, Powell 

River, Elk River, and Duck River), 

Virginia (Clinch River and Powell 

River), West Virginia (Ohio River, 

Dunkard Creek, Middle Island Creek, 

North Fork Hughes River, and Elk 

River), Wisconsin (St. Croix River, Wolf 

River, Embarrass River, Little Wolf 

River, and Willow Creek), and Ontario 

(Ausable River and Sydenham River). It 

is probable that the species persists in 

some of the 134 streams or lakes where 

it is now considered extirpated (Butler 

2007, p. 16); however, if extant, these 

populations are likely to be small and 

not viable. 

Snuffbox Population Estimates and 

Status 

Based on historical and current data, 

the snuffbox has declined significantly 

rangewide and is now known from only 

74 streams (down from 208 historically), 

representing a 65 percent decline in 

occupied streams (Table 2). Since 

multiple streams may comprise a single 

snuffbox population (French Creek 

system), the actual number of extant 

populations is less than 74. Extant 

populations, with few exceptions, are 

highly fragmented and restricted to 

short reaches. Available records indicate 

that 24 of 74, or 33 percent, of streams 

considered to harbor extant populations 

of the snuffbox are represented by only 

one or two recent L or FD individuals 

(Embarrass, Little Wolf, Maple, Pigeon, 

Kankakee, Meramec, Ohio, Muskingum, 

Olentangy, Stillwater, Green, Powell, 

Duck, and Black Rivers; and Little 

Mahoning, Middle Island, Big Darby, 

Little Darby, Salt, South Fork Scioto 

Brush, Slate, and Buck (Indiana), 

Graham, Buck (Kentucky) Creeks. 

Butler (2007, pp. 70–71) categorized 

the extant populations into three groups 

based on population size, general 

distribution, evidence of recent 

recruitment, and assessment of current 

viability. Stronghold populations were 

described as having sizable populations 

generally distributed over a significant 

and more or less contiguous length of 

stream (30 or more river mi (48 or more 

river km)), with ample evidence of 

recent recruitment, and currently 

considered viable. Significant 

populations were defined as small, 

generally restricted populations with 

limited recent recruitment and viability. 

Many significant populations are 

susceptible to extirpation, but this 

category has a broad range of quality. 

The third category, marginal 

populations, are defined as those which 

are very small and highly restricted with 

no evidence of recent recruitment, of 

questionable viability, and that may be 

on the verge of extirpation in the 

immediate future. Following this 

criteria, there are 6 stronghold 

populations, 23 significant populations, 


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and 45 marginal populations of 

snuffbox. 

A population is considered extant if L 

individuals or FD specimens have been 

located since approximately 1985. A 

population is considered to be 

recruiting if there was recent (within 

approximately 10 years) evidence of 

subadults (generally, individuals less 

than or equal to 1.5 in (3.8 cm) long or 

less than or equal to 4 years). 

TABLE 2—SNUFFBOX EXTANT STREAM POPULATION SUMMARY BY STREAM OF OCCURRENCE

Stream (state) Last observed Recruiting Potential viability Population size Population trend Status category 

Wolf River (WI) ...... 2006 Yes ........................ High ....................... Large ..................... Declining ............... Stronghold. 

Embarrass River 

(WI).

1995 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Little Wolf River 

(WI).

1999 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Willow Creek (WI) .. 2001 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Grand River (MI) .... 2002 Yes ........................ High ....................... Medium ................. ? ............................ Significant. 

Maple River (MI) .... 2001 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Pine River (MI) ....... 2002 ? ............................ Low ....................... Small ..................... Stable .................... Marginal. 

Belle River (MI) ...... 2002 Yes ........................ High ....................... Small ..................... ? ............................ Significant. 

Clinton River (MI) ... 2003 Yes ........................ High ....................... Large ..................... Declining ............... Significant. 

Huron River (MI) .... 2001 ? ............................ Low ....................... Medium ................. ? ............................ Significant. 

Davis Creek (MI) .... 2005 Yes ........................ High ....................... Medium ................. ? ............................ Significant. 

South Ore Creek 

(MI).

1999 Yes ........................ High ....................... Small ..................... ? ............................ Significant. 

Portage River (MI) 1998 Yes ........................ High ....................... Medium ................. ? ............................ Significant. 

Grand River (OH) ... 2006 Yes ........................ High ....................... Medium ................. ? ............................ Significant. 

St. Croix River (MN 

and WI).

2004 Yes ........................ High ....................... Large ..................... Declining ............... Significant. 

Kankakee River (IL) 1991 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Meramec River 

(MO).

1997 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Bourbeuse River 

(MO).

2006 Yes ........................ High ....................... Large ..................... Improving .............. Stronghold. 

Ohio River (OH) ..... 2001 ? ............................ Low ....................... Small ..................... ? ............................ Marginal. 

Muskingum River 

(OH).

2005 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Walhonding River 

(OH).

1991 ? ............................ ? ............................ Small ..................... Declining ............... Significant. 

Killbuck Creek (OH) 2009 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Olentangy River 

(OH).

1989 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Big Darby Creek 

(OH).

2008 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Little Darby Creek 

(OH).

1999 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Salt Creek (OH) ..... 1987 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Scioto Brush Creek 

(OH).

1987 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

South Fork Scioto 

Brush Creek (OH).

1987 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Little Miami River 

(OH).

1991 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Stillwater River 

(OH).

1987 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Pigeon River (IN) ... 1998 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Salamonie River 

(IN).

2004 Yes ........................ Low ....................... Small ..................... ? ............................ Significant. 

Tippecanoe River 

(IN).

2003 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Embarras River (IL) 2008 Yes ........................ Low ....................... Small ..................... Declining ............... Significant. 

Sugar Creek (IN) .... 1990 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Buck Creek (IN) ..... 1990 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Muscatatuck River 

(IN).

1988 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Graham Creek (IN) 1990 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

St. Francis River 

(MO).

2006 Yes ........................ High ....................... Medium ................. Stable .................... Significant. 

Black River (MO) ... 2002 Yes ........................ Low ....................... Small ..................... ? ............................ Significant. 

Tygarts Creek (KY) 1995 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Kinniconick Creek 

(KY).

2005 ? ............................ Low ....................... Small ..................... Declining ............... Marginal. 

Licking River (KY) .. 2006 ? ............................ Low ....................... Small ..................... ? ............................ Marginal. 

Slate Creek (KY) .... 1992 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Middle Fork Kentucky River (KY).1997 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.


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TABLE 2—SNUFFBOX EXTANT STREAM POPULATION SUMMARY BY STREAM OF OCCURRENCE—Continued 

Stream (state) Last observed Recruiting Potential viability Population size Population trend Status category 

Red Bird River (KY) 1995 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Red River (KY) ....... ∼2002 ? ............................ ? ............................ Small ..................... ? ............................ Significant. 

Rolling Fork Salt 

River (KY).

∼2005 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Green River (KY) ... 1989 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Buck Creek (KY) .... 1987–90 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Clinch River (TN 

and VA).

2006 Yes ........................ High ....................... Large ..................... Stable or Declining Stronghold. 

Powell River (TN 

and VA).

2008 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

Tennessee River 

(AL).

2006 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Paint Rock River 

(AL).

2008 Yes ........................ High ....................... Large ..................... Improving .............. Stronghold. 

Elk River (TN and 

AL).

2007 Yes ........................ Low ....................... Small ..................... Stable .................... Significant. 

Duck River (TN) ..... 2001 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Buffalo River (AR) .. 2006 ? ............................ ? ............................ Small ..................... ? ............................ Marginal. 

Spring River (AR) ... 2005 ? ............................ Low ....................... Medium ................. ? ............................ Significant. 

Strawberry River 

(AR).

1997 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Allegheny River 

(PA).

2001 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

French Creek (PA) 2008 Yes ........................ High ....................... Large ..................... Stable .................... Stronghold. 

West Branch 

French Creek 

(PA).

2008 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Le Boeuf Creek 

(PA).

2006 Yes ........................ Low ....................... Small ..................... ? ............................ Marginal. 

Muddy Creek (PA) 2008 Yes ........................ Low ....................... Medium ................. ? ............................ Significant. 

Conneaut Outlet 

(PA).

1997 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Little Mahoning 

Creek (PA).

1991 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Dunkard Creek (PA 

and WV).

2009 ? ............................ ? ............................ Small ..................... Declining ............... Significant. 

Shenango River 

(PA).

2002 ? ............................ ? ............................ Small ..................... ? ............................ Marginal.

Little Shenango 

River (PA).

2002 ? ............................ ? ............................ Small ..................... ? ............................ Significant. 

Middle Island Creek 

(WV).

2001 ? ............................ ? ............................ Small ..................... Declining ............... Marginal. 

North Fork Hughes 

River (WV).

2001 ? ............................ Low ....................... Small ..................... Declining ............... Significant. 

Elk River (WV) ....... 2004 ? ............................ Low ....................... Medium ................. Improving .............. Significant. 

Ausable River (ON) 2006 Yes ........................ High ....................... Medium ................. Declining ............... Significant. 

Sydenham River 

(ON).

2002 Yes ........................ High ....................... Large ..................... ? ............................ Stronghold. 

Upper Great Lakes Sub-Basin 

The snuffbox was formerly known 

from 15 streams and lakes in the upper 

Great Lakes sub-basin. The Fox River 

system in Wisconsin, particularly its 

major tributary the Wolf River (and its 

tributaries), had a widespread and 

locally abundant population. The 

species is thought to be extant in seven 

sub-basin streams; however, all but the 

Wolf and Grand Rivers have 

populations that are considered 

marginal. 

Wolf River—The Wolf River is the 

major tributary of the Fox River draining 

a large portion of northeastern 

Wisconsin and flowing southward to 

join the Fox River at Lake Butte Des 

Morts, near Oshkosh. Snuffbox records 

are known from Shawano, Waupaca, 

and Outagamie Counties. The snuffbox 

is known from a 30-river-mi (48-riverkm) reach of the Wolf River (Butler 

2007, p. 21). It is one of the few 

stronghold populations, but appears to 

exhibit a low level of recruitment. Only 

4 of 257 individuals collected in the 

mid-1990s were less than 6 years old 

(Butler 2007, p. 21). A bridge 

replacement project on the south side of 

Shawano, scheduled to begin in 2010, 

may adversely impact the large snuffbox 

bed located just downstream (ESI 2006, 

p. 10). The zebra mussel occurs in this 

river, with a 0.7 percent infestation rate 

on unionids sampled in 2006 (ESI 2006, 

p. 6). This large population continues to 

be viable but appears to be in decline 

(Butler 2008, pers. comm.). 

Embarrass River—A western tributary 

of the lower Wolf River, the Embarrass 

River parallels the western bank of the 

Wolf River before joining it at New 

London, Wisconsin. A population of the 

snuffbox is located in the headwaters 

below a small dam at Pella, Wisconsin. 

Records exist for three L individuals 

and two dead specimens during 1987– 

1988 and a single D specimen in 1995 

(Butler 2007, p. 22). Its current status is 

unknown. 

Little Wolf River—The Little Wolf 

River is a western tributary of the lower 


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Wolf River in Waupaca County, 

Wisconsin. The snuffbox is known from 

a single L individual collected in 1988 

at RM 14 below the Mill Pond dam at 

Manawa (Butler 2007, p. 22). Five D 

specimens were found during 1999 at 

RM 2, where shells were abundant in a 

muskrat midden (Butler 2007, p. 22). 

Nothing else is known regarding this 

population. 

Willow Creek—Willow Creek flows 

eastward into Lake Poygan, a large flowthrough lake of the Wolf River system, 

in Waushara County, Wisconsin. The 

snuffbox is known from a single 

observation of two L females in 2001 

(Butler 2007, p. 22). No other 

information is available on the status of 

this population. 

Grand River—The Grand River, a 

major Lake Michigan tributary, 

represents the largest lotic (moving 

water) watershed in Michigan and is 

located in the southwestern portion of 

the State. The snuffbox is sporadically 

distributed in approximately 25 river mi 

(40 river km) of the middle Grand River, 

approximately between the confluences 

of the Flat and Maple Rivers. The 

medium-sized population appears to 

have a low level of viability, with 

recruitment noted in 1999 (Badra 2008, 

pers. comm.). 

Maple River—The Maple River is a 

northeastern tributary of the Grand 

River draining south-central Michigan. 

A single snuffbox record (one L 

individual) is known from 2001 in 

southern Gratiot County, approximately 

20 river mi (32 river km) upstream of 

the Grand River (Badra 2008, pers. 

comm.). Portions of the Maple River and 

several tributaries have been 

channelized, but the suitability of these 

channelized areas for the snuffbox is 

unknown (Badra 2010, pers. comm.). 

The current status of this small 

population is unknown. 

Pigeon River—The Pigeon River is a 

headwater tributary of the St. Joseph 

River system of Lake Michigan, flowing 

westward across northern-most Indiana, 

crossing the State border to its 

confluence in southwestern Michigan. 

One very large FD specimen was found 

in 1998, among thousands of shells in 

LaGrange County, Indiana (Butler 2007, 

p. 24). The same site was sampled in 

1996 without evidence of this species, 

and R shells were found at three of nine 

sites sampled in 2004 (Butler 2007, p. 

24). The snuffbox occupied reach 

historically covered more than 10 river 

mi (16.1 river km) in north-central 

LaGrange County. The species is very 

rare in this river, and its viability is 

unknown. 

Lower Great Lakes Sub-Basin 

Of all the water bodies from which 

the snuffbox was historically recorded, 

32 are in the lower Great Lakes subbasin, including several chains-of-lakes, 

springs, and channels in some systems 

(Clinton, Huron Rivers). Historically 

sizable populations occurred in some 

streams (Lake Erie; Belle, Clinton, 

Huron, Portage, and Niagara Rivers), but 

the species had become 

‘‘characteristically uncommon’’ by the 

1970s (Strayer 1980, p. 147). A pre-zebra 

mussel decline of unionids in Lake Erie 

was noted (Mackie et al. 1980, p. 101), 

and the snuffbox appeared extirpated 

there by the late 1960s. The Lake St. 

Clair population of snuffbox persisted 

until around 1983 (Nalepa and Gauvin 

1988, p. 414; Nalepa 1994, p. 2231; 

Nalepa et al. 1996, p. 361), which was 

the year the zebra mussel is thought to 

have invaded (Schloesser et al. 1998, p. 

70). Observations of L and FD snuffbox 

from the Detroit River were made until 

1994, but the mussel fauna has since 

been devastated by zebra mussels, and 

the snuffbox is now considered to be 

extirpated (Schloesser et al. 1998 p. 69; 

Butler 2007, p. 25). Other snuffbox 

populations in the sub-basin may also 

have suffered from zebra mussel 

invasions, but not those in the Ausable 

and Sydenham Rivers in Ontario. The 

lack of impounded area on these 

streams has likely prevented the 

introduction or the establishment of 

zebra mussels (Ausable River Recovery 

Team 2005, p. 12; Dextrase et al. 2000, 

p. 10). The snuffbox is considered 

extant in 10 streams of the lower Great 

Lakes sub-basin, including a stronghold 

population in the Sydenham River and 

sizable but reach-limited populations in 

the Clinton River and Davis Creek. A 

single FD valve was reported in 1998 

from among 24 sites sampled in the 

Thames River, but no evidence of the 

snuffbox was found at 16 Thames sites 

in 2004 (McGoldrick 2005, pers. 

comm.). Currently, the species is 

considered extant in Canada only in the 

Ausable and Sydenham Rivers (Morris 

and Burridge 2006, p. 9). Both of these 

populations are viable. 

Ausable River—The Ausable River is 

a southeastern tributary of Lake Huron, 

draining southwestern Ontario, Canada. 

A survey conducted in 2006 found that 

a sizable population of snuffbox occurs 

in the lower portion of the stream in 

over 23 river mi (37 river km) 

(McGoldrick 2007, pers. comm.). The 

size range of individuals found in the 

2006 survey indicates recent 

recruitment in the viable population 

(McGoldrick 2007, pers. comm.; Staton 

2007, pers. comm.). 

Pine River—A tributary of the St. Clair 

River, the Pine River flows south and is 

located in St. Clair County, in 

southeastern Michigan. Although 

apparently stable, the snuffbox 

population is small, very restricted in 

range, and has a low potential for 

viability (Badra 2002, pers. comm.; 

Badra and Goforth 2003, p. 23). 

Belle River—The Belle River is 

another tributary of the St. Clair River 

in St. Clair County, flowing in a 

southeasterly direction. Records for the 

snuffbox date to the early 1960s, but all 

L and FD records over the past 40 years 

have been from the same lower 

mainstem site. Historically, a sizable 

population was found in the Belle (65 

specimens, 1965). The Belle is located 

in a primarily agricultural watershed 

(Hoeh and Trdan 1985, p. 115), and is 

impacted by sedimentation and runoff. 

The population has declined to the 

point of being small, but shows 

evidence of recruitment and viability 

(Badra 2002, pers. comm.; Badra and 

Goforth 2003, p. 24; Sherman 2005, 

pers. comm.). 

Clinton River—The Clinton River is 

an eastward flowing chain-of-lakes 

tributary of Lake St. Clair in 

southeastern Michigan. The snuffbox 

population in the Clinton River is 

limited to around 10 river mi (16.2 river 

km) and lakeshore in the western 

suburbs of Pontiac primarily between 

Cass and Loon Lakes. This population 

appears to be recruiting (Sherman 

Mulcrone 2004, p. 64) and viable, 

although apparently in decline since the 

early 1990s (Badra 2002, pers. comm.; 

Butler 2007, p. 27). 

Sydenham River—The Sydenham 

River is a large, southeasterly flowing, 

eastern tributary of Lake St. Clair in 

extreme southwestern Ontario. The 

snuffbox was reported in the mid-1960s 

and early 1970s but was overlooked 

during surveys in 1985 (except D shells) 

and 1991 (Butler 2007, p. 28). During 

the 1997–99 sampling, a total of 10 L 

and FD individuals were found from 4 

of 12 sites, including the 3 1960s sites 

(Metcalfe-Smith et al. 2003, p. 41). The 

snuffbox was recorded at a rate of 0.22 

per hour of effort during 1997–98 

(Metcalfe-Smith et al. 2000, p. 728). 

More recent sampling found 57 L and 

FD individuals from 21 collection 

events (some individuals may have been 

counted multiple times) at six sites 

during 2000–02. The increase in 

numbers relative to historical 

collections may be attributed to more 

intensive sampling methods rather than 

to improving population size (MetcalfeSmith et al. 2003, p. 46), thus making 

population trend assessments difficult 

(Morris and Burridge 2006, p. 12). This 


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stronghold population is recruiting 

(Butler 2007, p. 28), viable, and is 

currently known from approximately 30 

river miles (48 km) of the middle 

Sydenham. 

Huron River—The Huron River is a 

major tributary of western Lake Erie 

draining a significant portion of 

southeastern Michigan. It is a complex 

system of flow-through chains-of-lakes 

and tributaries. The snuffbox is 

considered extant in two disjunct upper 

mainstem reaches. Individuals in the 

middle Huron River reach and in Davis 

Creek are considered a single 

population segment (Marangelo 2005a, 

pers. comm.). 

Zebra mussels invaded the Huron 

River system in the early 1990s. Zebra 

mussel densities on individual mussels 

increased from less than 1 in spring 

1995 to 245 in winter 1998 (Nichols et 

al. 2000, p. 72). Despite the increasing 

presence of zebra mussels, the Huron 

population is probably recruiting and 

viable (Butler 2007, p. 29). 

Davis Creek—Davis Creek is a chainof-lakes in the upper Huron River 

system, primarily in southeastern 

Livingston County, Michigan. The 

snuffbox appears to be limited to the 

lower 3 river mi (4.8 river km), 

comprising a single population with one 

of the extant Huron River population 

segments in this area. This viable 

population appears to be sizable and is 

experiencing recent recruitment 

(Marangelo 2005a, pers. comm.; Zanatta 

2005, pers. comm.). 

South Ore Creek—South Ore Creek is 

a northern tributary of the Huron River, 

forming a southward flowing chain-oflakes draining southeastern Livingston 

County, Michigan. The snuffbox was 

discovered in 1999, just upstream of Ore 

Lake, which is near the Huron River 

confluence (Butler 2007, p. 31). Three 

subadult snuffbox (two age 2, one age 3– 

4) were recorded. Despite the lack of 

additional information, the small 

population appears to be viable based 

on recent recruitment. 

Portage River—The Portage River is a 

chain-of-lakes in the northwestern 

portion of the Huron River system. Two 

University of Michigan Museum of 

Zoology (UMMZ) records suggest 

historical abundance (Badra 2002, pers. 

comm.). The species was reported as 

‘‘rare’’ in the lower river during 1976–78 

(Strayer 1979, p. 94). At least 22 L, 

young (age 4 and younger) individuals 

were identified in 1998 at one of three 

sites upstream of Little Portage Lake and 

Portage Lake (Butler 2007, p. 31). The 

localized population appears to be 

medium-sized and viable. 

Grand River—The Grand River is a 

99-river-mi (159-river-km) tributary of 

Lake Erie, flowing north then west to its 

confluence northeast of Cleveland, 

Ohio. Several museum snuffbox records 

date back to the 1800s. Dozens of FD 

snuffbox were found washed up on the 

banks in the vicinity of the Interstate 90 

crossing in Lake County, Ohio, 

following a major flood in 2006 (Butler 

2007, p. 32). The species is known from 

approximately 12 river mi (19.3 river 

km) downstream of Harpersfield Dam 

(Huehner et al. 2005, p. 59; Zimmerman 

2008a, pers. comm.). The sizable 

population was considered recruiting 

based on the 1995 Huehner et al. (2005, 

p. 59) survey. 

Upper Mississippi River Sub-basin 

The snuffbox was historically known 

from 17 streams in the upper 

Mississippi River sub-basin. Records 

exist for Mississippi River Pools (MRPs) 

3–4, 5a–6, and 14–16 (Kelner no date, p. 

6), with early surveys summarized by 

van der Schalie and van der Schalie 

(1950, p. 456). The species was reported 

L in the upper river in the 1920s (Grier 

1922, p. 15; Grier 1926, p. 119) but not 

from subsequent surveys (254 sites 

upstream of the Ohio River during 

1930–1931 (UMMZ, Ellis 1931, pp. 1– 

10), MRPs 5–7 and 9 in 1965 (Finke 

1966, Table 2; Thiel 1981, p. 16), MRPs 

3–11 during 1977–79 (Thiel 1981, p. 

16)) and is now extirpated from the 

mainstem of the Mississippi River 

(Havlik and Sauer 2000, p. 4). Only 4 of 

17 historical populations remain, but 

they include two of the largest 

rangewide (St. Croix and Bourbeuse 

Rivers). Three populations, including 

the St. Croix, appear to be declining. 

St. Croix River—The St. Croix River is 

a major south-flowing tributary of the 

upper Mississippi River and forms the 

border between southeastern Minnesota 

and northwestern Wisconsin. Densities 

of juvenile snuffbox declined at eight 

sites between 1992 and 2002 (Hornbach 

et al. 2003, p. 344). Snuffbox density at 

Interstate Park declined significantly 

between 1988 and 2004 (WIDNR 2004). 

A flood in 2001 may have contributed 

to these declines in mussel density, but 

post-flood recruitment was also 

surprisingly low (WIDNR 2004). The St. 

Croix snuffbox population occurs from 

the Northern States Power Dam (NSPD) 

at RM 54.2 to RM 36.8 (Heath 2005, 

pers. comm.), represents the species’ 

northernmost occurrence, and despite 

recent observed declines, remains one of 

the six stronghold populations 

rangewide. 

Kankakee River—The Kankakee River 

is a major, westward-flowing, upper 

Illinois River tributary with its 

headwaters in northwest Indiana and 

northeast Illinois. The snuffbox was 

reported over a century ago (Baker 1906, 

p. 63), but surveys in 1911 (43 sites; 

Wilson and Clark 1913, pp. 41–50), 

1978 (13 sites; Suloway 1981, p. 236), 

1975–2000 (18 samples from an 

unknown number of Will County, 

Illinois, sites; Sietman et al. 2001, p. 

279), and 1999 (4 sites, Stinson et al. 

2000, Appendix C) failed to find it. It 

was considered extirpated from the 

Kankakee by Cummings et al. (1988, p. 

16), but single FD specimens in Illinois 

(Will County in 1988, Kankakee County 

in 1991) were subsequently found. Only 

R shells have been found since 1991. 

The Kankakee River population, if 

extant, appears small, localized, and of 

doubtful viability. 

Meramec River—The Meramec River 

is a 236-mi (380-km) tributary that flows 

northeasterly into the Mississippi River 

downstream of St. Louis and drains the 

northeastern slope of the Ozark Plateaus 

in east-central Missouri. Early species 

lists failed to report the snuffbox (Grier 

1916, p. 518; Utterback 1917, p. 28). 

Buchanan (1980, p. 63) found FD 

specimens at three sites and R shells at 

two other sites sampled in 1977–78. 

Roberts and Bruenderman (2000, p. 85) 

sampled 42 sites in 1997, including 26 

of Buchanan’s (1980, p. 5) sites, and 

found FD specimens at RM 33.5, 48.8, 

and 59.8; and one L individual at RM 

39.8. The L individual (2.4 in (6.1 cm), 

approximately 6 years old) was reported 

from a reach where a die-off, perhaps 

attributable to disease, was reported in 

1978 (Buchanan 1986, p. 44). There was 

an obvious decline of mussels in the 

system based on catch-per-unit-effort 

data over the 20-year period (Roberts 

and Bruenderman 2000, p. 8). The 

Meramec snuffbox population is rare, 

sporadically distributed over 

approximately 26 river mi (41.8 river 

km), and of unknown viability. 

Bourbeuse River—The Bourbeuse 

River is a 149-mi (240-km), 

northeasterly flowing, northern tributary 

of the Meramec River, joining it at RM 

68. The snuffbox is currently distributed 

over about 60 river mi (96.6 river km) 

upstream of RM 16, plus a disjunct site 

at the mouth of the river. Although it 

was considered to have ‘‘greatly 

declined’’ by the late 1990s (Roberts and 

Bruenderman 2000, p. 15), post-2000 

sampling indicates that the population 

is recruiting, viable, and improving 

(McMurray 2006, pers. comm.). The 

Bourbeuse, one of the few stronghold 

snuffbox populations rangewide, has 

been augmented with laboratory 

propagated juveniles since 2002 

(McMurray 2006, pers. comm.). 


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Lower Missouri River System 

The snuffbox was historically known 

from four streams in this system. The 

highly disjunct occurrences suggest that 

it was more widespread historically. All 

populations in the system are 

considered extirpated (Butler 2007, p. 

36). 

Ohio River System 

Half of the water body occurrences for 

the snuffbox rangewide are known from 

the Ohio River system, which 

collectively represented the largest 

block of available habitat for this 

species. Sizable populations historically 

occurred in at least a dozen streams in 

the system. Today, only French Creek is 

considered to have a stronghold 

population, although nine others are 

also significant. Currently, the species is 

known from 40 of the 107 streams of 

historical occurrence. 

Ohio River—The Ohio River is the 

largest eastern tributary of the 

Mississippi, with its confluence 

marking the divide between the upper 

and lower portions of the latter system. 

Numerous historical records are known 

from throughout the River. Recently, 

single FD and L specimens have been 

reported from just below Belleville Lock 

and Dam, Ohio and West Virginia, in 

1995 and 2001, respectively (ESI 2002, 

p. 27). Having persisted in this highly 

modified river may indicate that the 

small population exhibits a low level of 

viability. 

Allegheny River—The 325-mi (523- 

km) Allegheny River drains 

northwestern Pennsylvania and a small 

portion of adjacent New York flowing 

south before joining the Monongahela 

River at Pittsburgh to form the Ohio 

River. Snuffbox collections are 

sporadically known since around 1900 

in Pennsylvania from Forest County 

downstream to Armstrong County. The 

snuffbox is currently known from three 

disjunct sites over a 42-river-mi (67.6- 

river-km) reach centered in Venango 

County (Butler 2007, p. 37). Its 

occurrence in the lower Allegheny River 

and lower French Creek could be 

considered a single population segment. 

The viability status of the small 

population is unknown. 

French Creek—French Creek is a 

major tributary of the middle Allegheny 

River with its headwaters in western 

New York and flowing south into 

northwestern Pennsylvania. The 

snuffbox is known from the length of 

the stream in Pennsylvania in Erie, 

Crawford, Mercer, and Venango 

Counties. Most records date since 

approximately 1970 (Dennis 1971, p. 

97). Snuffbox collections made during 

2002–2004 were summarized by Smith 

(2005, pp. 3–9). Live and FD specimens 

were found at 19 sites throughout the 

stream. The size of the L individuals 

indicated that multiple year classes 

were represented, including subadults. 

The species stretches for approximately 

80 river mi (128.7 river km) from around 

RM 10, upstream. The population 

encompasses several of its tributary 

population segments as well, making it 

relatively more secure when compared 

to most of the other stronghold 

populations that are linearly distributed 

and, thus, more susceptible to stochastic 

events (Sydenham, Bourbeuse, and 

Clinch Rivers). The French Creek 

snuffbox population is considered large 

and viable (Evans 2003a, pers. comm.; 

Zimmerman 2008c, pers. comm.), 

appears stable, and may represent the 

best stronghold population rangewide. 

West Branch French Creek—West 

Branch of French Creek follows a 

southerly course to its parent stream in 

Erie County, Pennsylvania. The only 

record for the snuffbox dates from 1993, 

but the number of specimens and shell 

condition are unknown (Evans 2003b, 

pers. comm.). Union City Lake isolates 

the upper French Creek and West 

Branch French Creek population 

segment from the main French Creek 

population. The snuffbox was not found 

at three sites sampled in 2006 (Smith 

2006, pers. comm.). Zimmerman (2008c, 

pers. comm.) documented 38 L 

individuals at a site near Wattsburg, 

Pennsylvania. This population appears 

to be small and of unknown viability. 

Le Boeuf Creek—Le Boeuf Creek is a 

small western tributary of upper French 

Creek flowing in a southerly direction 

just west of West Branch French Creek 

in Erie County. The first snuffbox 

collections in this creek were made 100 

years ago (Ortmann 1909a, p. 188). Two 

FD and 6 R shells were reported in 1988 

(Evans 2003b, pers. comm.), and 1 L, 16 

FD, and 8 R specimens were found in 

1991 (Butler 2007, p. 40). Three L 

individuals were found at a site in 2006 

(Smith 2006, pers. comm.). The 

snuffbox population has recently 

recruited and exhibits some level of 

viability, but appears to be very limited 

in extent. 

Muddy Creek—Muddy Creek is an 

eastern tributary of upper French Creek 

in Crawford County, Pennsylvania. The 

snuffbox was not discovered until the 

summer of 2003. Forty-two L 

individuals were reported from 11 of 20 

lower river sites (Morrison 2005, pers. 

comm.). Low numbers were found at 

most sites, but 18 L individuals were 

collected from a site near the mouth. 

This occurrence is considered to be part 

of the more extensive French Creek 

snuffbox population. Zimmerman 

(2008c, pers. comm.) documented one L 

female in 2008. The population is 

medium-sized, occurs along 8 river mi 

(12.9 river km) of the lower mainstem, 

and is recruiting, as recent juveniles 

were recorded (Morrison 2005, pers. 

comm.). 

Conneaut Outlet—This stream forms 

the outlet to Conneaut Lake, flowing in 

a southeasterly direction until its 

confluence with middle French Creek, 

Crawford County. The snuffbox was first 

reported by Ortmann (1909a, p. 188), 

and was rediscovered L in 1997, but 

without collection details (Butler 2007, 

p. 40). No specimens were found at a 

site sampled in 2006 (Smith 2006, pers. 

comm.). The snuffbox is considered rare 

in this stream and its viability is 

unknown. 

Little Mahoning Creek—Little 

Mahoning Creek is a tributary of 

Mahoning Creek, a lower eastern 

tributary of the Allegheny River 

northeast of Pittsburgh. The snuffbox 

was discovered in 1991, when sampling 

produced two FD and one R specimen 

at 1 of 12 sites in the system (Butler 

2007, p. 41). The lower 10 miles of Little 

Mahoning Creek is subject to periodic 

inundation by a reservoir on Mahoning 

Creek (Butler 2010, pers. comm.). 

However, the impact of this periodic 

flooding on the snuffbox is not known. 

Viability is unknown. 

Dunkard Creek—Dunkard Creek is an 

easterly flowing, western tributary of the 

middle Monongahela River, straddling 

the Pennsylvania and West Virginia 

State lines. Snuffbox records occur in 

both States from several museum 

collections from 1969–74. Small 

numbers of specimens, of 

undocumented condition, were found at 

four sites during 1993–94 sampling in 

Pennsylvania (Bogan 1993, p. 8; Evans 

2003b, pers. comm.). Eight specimens, 

of undocumented condition, were 

collected at a West Virginia site in 1997. 

On September 1, 2009, a fish kill was 

reported in Dunkard Creek due to an 

unknown cause (Clayton 2009, pers. 

comm.). The Upper Monongahela River 

Association (2009) reported that 161 

aquatic species including fish, mussels, 

and plants died along Dunkard Creek 

due to this toxic event. According to 

Clayton (2009, pers. comm.), the event 

may have killed 100 percent of the 

mussel fauna in the entire stream. The 

status of this population is not known 

at this time, but the snuffbox may now 

be extirpated from Dunkard Creek. 

Shenango River—The Shenango River 

is a large tributary in the Beaver River 

system, a northern tributary of the upper 

Ohio River in west-central 

Pennsylvania. The snuffbox was 


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reported from four sites on the 

Shenango in 1908 (Ortmann 1919, p. 

328). Six L individuals were collected 

from three sites sampled in 2001–02 

between Jamestown and New Hamburg 

(about 25 river mi (40.2 river km)). The 

upper reach is considered the best 

habitat in the Shenango River. The 

population is small and has declined, 

although some recent reproduction is 

evident (Zimmerman 2008b, pers. 

comm.). 

Little Shenango River—The Little 

Shenango River is a small tributary of 

the upper Shenango River, Mercer 

County, Pennsylvania. This population 

was not located during limited surveys 

(Dennis 1971, p. 97; Bursey 1987, p. 42), 

but a single FD museum record from 

1991 exists. The species was reported to 

be relatively abundant and reproducing 

in the lower portion in 2002 

(Zimmerman 2008b, pers. comm.). 

Viability of the small population is 

unknown. 

Middle Island Creek—Middle Island 

Creek is a small tributary of the Ohio 

River in northwestern West Virginia. 

The first snuffbox records were made at 

six sites in 1969, when the species was 

locally common in Doddridge, Tyler, 

and Pleasants Counties (Taylor and 

Spurlock 1981, p. 157). The snuffbox 

was later found at two sites in Tyler 

County in 1980, and the overall mussel 

population was considered to be 

‘‘thriving’’ (Taylor and Spurlock 1981, p. 

157). The most recent record was for a 

single L individual collected in Tyler 

County in 2001 (Zimmerman 2008b, 

pers. comm.). This snuffbox population 

has declined, is currently rare, and its 

viability is questionable (Zimmerman 

2008b, pers. comm.). 

Muskingum River—The Muskingum 

River is a large, southerly flowing, 

northern tributary of the upper Ohio 

River draining a significant portion of 

east-central Ohio. The snuffbox, which 

has a long collection history dating to 

the early 1800s, occurred along the 

entire mainstem and was locally 

abundant. Two L individuals and two 

FD shells were found in 1979, but no L 

or FD snuffbox were found in surveys 

conducted in 1979–81 (Stansbery and 

King 1983) and in 1992–93 (Watters and 

Dunn 1993–94, p. 241). A single L 

specimen was located during sampling 

for a construction project in 2005 near 

Dresden (Taylor 2006, pers. comm.). 

Viability of this population is unknown. 

Walhonding River—The Walhonding 

River is a short (23.3 river mi (37.5 river 

km)), east flowing tributary of the 

Muskingum River in central Ohio, 

forming the latter river at its confluence 

with the Tuscarawas River, and formed 

by the confluence of the Mohican and 

Kokosing Rivers. The snuffbox 

historically occurred throughout the 

river. The extant snuffbox reach (RM 

1.8–6.8) is downstream from Killbuck 

Creek. The population had apparently 

declined in range and size by the early 

1990s and possibly further since. A once 

productive site about 0.25 mi (0.40 km) 

downstream of the Killbuck Creek 

confluence yielded only a few mussels 

of very common species in 2006, but no 

snuffbox (Butler 2007, p. 44). The 

Walhonding River population is 

considered small and of unknown 

viability. 

Killbuck Creek—Killbuck Creek is a 

large tributary of the lower Walhonding 

River, flowing south from southern 

Medina County to Coshocton County 

and entering the latter at approximately 

RM 7. Live and FD snuffbox were found 

by Hoggarth (1997, p. 33) at eight sites 

from RM 15 to the mouth. Its occurrence 

has become more sporadic in the last 10 

years. In spring 2006, 4 L adults were 

found at 2 sites approximately 3 river 

mi (4.8 river km) apart, while 9 large L 

individuals and a single FD specimen 

were collected near RM 13 during fall 

2006 (Ahlstedt 2007, pers. comm.; 

Butler 2007, p. 45). A shrinking 

distribution, declining population size, 

and lack of evidence of recent 

recruitment suggest that the population 

may be losing viability and trending 

towards extirpation. 

North Fork Hughes River—The North 

Fork Hughes River is a westerly flowing 

tributary of the Hughes River in the 

lower Little Kanawha River system in 

northwestern West Virginia. The 

snuffbox was found at one of six North 

Fork sites sampled during a 1981–82 

survey of the Little Kanawha River 

system (Schmidt et al. 1983). A total of 

41 L adult individuals (23 reported as 

gravid) were reported at 5 sites located 

over a 1.5-mi (2.4-km) reach in North 

Fork State Park, Richie County, in 1993 

(Butler 2007, p. 46). At least 10 L 

individuals were found at a site in the 

park in 1997 (Butler 2007, p. 46), and 

a single FD specimen was collected at 

an additional site downstream in 2001 

(Butler 2007, p. 46). This small snuffbox 

population is declining and currently 

restricted to less than 4 river mi (6.4 

river km), but may be viable. 

Elk River—The Elk River is a major, 

181-mi (291-km) tributary in the lower 

Kanawha River system draining central 

West Virginia flowing west to the 

Kanawha at Charleston. The snuffbox 

went undetected in a 1920s survey 

(Butler 2007, p. 46). Ten L individuals 

were collected during 1991–1995, the 

smallest being about 5 years old (Butler 

2007, pp. 46–47). Collectively, 16 L 

individuals were identified at 8 sites in 

a 13-river-mi (20.9-river-km) reach in 

Kanawha County in 2002, and 4 L 

individuals were found at 4 sites in 

2004 over a 16.8-river-mi (27-river-km) 

reach further upstream (Douglas 2005, 

pers. comm.). This medium-sized 

population extends over 30 river mi 

(48.3 river km), is viable, and may have 

improved since the 1970s. 

Tygarts Creek—Tygarts Creek is a 

small, north-flowing, southern tributary 

of the Ohio River in northeastern 

Kentucky. Thirteen snuffbox were 

reported from one of five sites sampled 

in 1977 (Taylor 1980, p. 90). FD 

specimens are also known from 1981 

and 1987 (Cicerello 2003, pers. comm.). 

Nine L (Butler 2007, p. 47) and 36 FD 

specimens were found at 2 sites, 

respectively, in 1988, while 1 L and 2 

FD were reported from at least 2 sites in 

1995 (Cicerello 2003, pers. comm.). The 

overall mussel population appeared 

‘‘healthy’’ in 1977 (Taylor 1980), but the 

small snuffbox population has recently 

declined, and its viability is unknown. 

Scioto River System—The Scioto 

River system in central and southcentral Ohio is a major northern 

tributary of the upper Ohio River. The 

system was one of the most routinely 

sampled watersheds for mussels (mostly 

OSUM records), and historically 

harbored a large and thoroughly 

dispersed snuffbox population in the 

mainstem and 16 tributaries. The system 

was either exceptional for its snuffbox 

population, or it provided a general 

historical perspective of what 

researchers may have found if other 

systems had been as thoroughly 

sampled. Sizable populations were 

noted in at least the Olentangy River, 

Big Darby Creek, and Big Walnut Creek. 

Development associated with the 

Columbus metropolitan area has taken a 

major toll on the aquatic fauna. 

Pollutants from the 1800s included 

wastes from sawmills, breweries, and 

slaughterhouses (Butler 2007, p. 48). 

Only a few fish species were found in 

the Scioto River 100 years ago 

(Trautman 1981, p. 33). Currently, 90 to 

95 percent of the normal summer-fall 

flow in the river consists of wastewater 

treatment plant (WWTP) discharges 

(Yoder et al. 2005, p. 410). Museum 

records indicate that the snuffbox had 

completely disappeared from the 

mainstem by the 1970s. A series of 

reservoirs around Columbus fragmented 

habitat and eliminated or reduced 

populations (Olentangy and Scioto 

Rivers; Alum, Big Walnut and Deer 

Creeks). Currently, remnant populations 

remain in six streams, making the 

snuffbox precariously close to 

extirpation throughout this once rich 

system. 


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Olentangy River—The Olentangy 

River is a major headwater tributary of 

the Scioto River, draining central Ohio 

and flowing south to its confluence in 

Franklin County. OSUM snuffbox 

records date to the 1870s, although most 

are from the 1950s and 1960s. The 

snuffbox was reported from 15 of 31 

mainstem sites collected during a 

1960–61 survey, when it appeared 

‘‘fairly common’’ in the lower river 

(Stein 1963, p. 138). A single L 

individual in southern Delaware County 

and two FD specimens in eastern 

Marion County were found among 30 

sites in 1989, with R shells at 7 other 

sites (Hoggarth 1990, pp. 20–27). The 

small population has declined 

(Hoggarth 1990, p. 14), and viability is 

unknown. 

Big Darby Creek—Big Darby Creek is 

one of the major tributaries draining the 

northwestern portion of the Scioto River 

system in central Ohio. Dozens of large 

OSUM lots of snuffbox date to the late 

1950s; six Pickaway County collections 

in 1962 alone had 250 L and FD 

specimens. Watters (1990, p. 4; 1994, 

p. 100) surveyed 42 mainstem sites in 

1986 and 49 sites in 1990. Combining 

the data from both years, 80 L and FD 

snuffbox were collected at 22 sites 

(Watters 1994, p. 101). The population 

in 1990 occurred in a reach from 

approximately RM 11.5 to RM 42.5. The 

snuffbox was recruiting (Watters 1994, 

p. 101); four individuals during both 

1986 and 1990 were 2 to 5 years of age. 

The overall population trend over the 

past 40 years has been downward. 

Between 1986 and 1990, the number of 

L and FD specimens was reduced from 

54 to 16 and its distribution declined 

from 17 to 8 sites. Two FD specimens 

were found at sites in Franklin (1996) 

and Pickaway (2000) Counties, and 

three other sites produced only R 

specimens (OSUM records). This 

historically large snuffbox population 

has declined to marginal status and its 

viability is questionable. 

Little Darby Creek—Little Darby Creek 

is the major tributary in the Big Darby 

Creek system, flowing in a southeasterly 

direction to its confluence in 

southwestern Franklin County, Ohio. 

The 25 OSUM lots for this species are 

small (fewer than five specimens per 

lot), date to the early 1960s, and 

represent lower mainstem sites in 

Madison County. Single FD and R 

specimens were collected in 1999 from 

a Union County site (OSUM 66740), 

where L individuals were collected in 

1964 (Stein 1966, p. 23). This site 

yielded only R specimens in 1990 

(Watters 1990, Appendix A.11; 1994, 

p. 102). Overall, the snuffbox was 

historically known from 35 river mi 

(56 river km). The well documented 

OSUM collection history illustrates the 

steady decline of a snuffbox population 

nearing extirpation. 

Salt Creek—Salt Creek is an eastern 

tributary in the Scioto River system, 

south-central Ohio. All records (OSUM) 

were collected in the lower mainstem 

(Ross County) beginning in 1958. A 

single L individual from 1987 represents 

the last known record. The mussels in 

this system ‘‘have been heavily 

impacted, apparently by the towns of 

Adelphi and Laurelville’’ (Watters 1992, 

p. 78). The current status of this 

snuffbox population is unknown. 

Scioto Brush Creek—Scioto Bush 

Creek is a small, western tributary of the 

lower Scioto River in Scioto County, 

south-central Ohio. The snuffbox was 

discovered here in the 1960s (Watters 

1988a, p. 45). Three L and FD 

specimens from 2 sites and R shells 

from 2 other sites were collected during 

a 1987 survey covering 11 sites (Watters 

1988a, pp. 210–220). The snuffbox 

population, collectively known from 

five fragmented sites along the lower 

two-thirds of stream, is small, and its 

viability is unknown. 

South Fork Scioto Brush Creek— 

South Fork Scioto Brush Creek is a 

small tributary of Scioto Brush Creek, in 

the lower Scioto River system. A single 

snuffbox was found during a survey of 

five sites in 1987 (Watters 1988a, pp. 

210–220). The South Fork and Scioto 

Brush Creek populations can be 

considered a single population unit, the 

viability of this unit is uncertain. 

Kinniconick Creek—Kinniconick 

Creek is a small, southern tributary of 

the Ohio River in northeastern 

Kentucky. Snuffbox was reported L from 

4 of 15 sites sampled in 1982 with R 

shells from an additional 2 sites (Warren 

et al. 1984, pp. 48–49). Single FD and 

L snuffbox were collected in 2001 and 

2004, respectively, from sampling 

efforts at several sites (Butler 2007, 

p. 51), and a single FD specimen was 

found while resurveying four sites in 

2005 (Butler 2007, p. 51). The snuffbox 

declined in the past few decades, is 

considered rare, and its viability is 

uncertain. 

Little Miami River—The Little Miami 

River is a northern tributary of the Ohio 

River in southwestern Ohio, flowing 

south into the latter at the eastern fringe 

of the Cincinnati metropolitan area. 

Snuffbox records from the Little Miami 

date to the mid-1800s, but most 

collections are from the past several 

decades. Seven FD specimens were 

found at 4 of 46 mainstem sites 

surveyed during 1990–91, with 10 R 

shells at 6 other sites (Hoggarth 1992, 

p. 265). The FD specimens were found 

in approximately 20 river mi (32.2 river 

km), mostly in Warren County. Current 

viability of this small population is 

unknown. 

Licking River—The Licking River is a 

southern tributary of the Ohio River in 

northeastern Kentucky, flowing in a 

northwesterly direction to its 

confluence across from Cincinnati. The 

snuffbox occurred at 13 of 60 historical 

mainstem sites below Cave Run 

Reservoir (Laudermilk 1993, p. 45) and 

a preimpoundment site in the reservoir 

footprint (Clinger 1974, p. 52). The 

population extended approximately 50 

river mi (80.5 river km). All collections 

of snuffbox are small in number (Butler 

2007, p. 52). A single L individual and 

a FD specimen were found at 2 sites and 

R shells were reported from 7 other sites 

among 49 sites sampled in 1991 

(Laudermilk 1993, p. 45). Single L and 

FD snuffbox were collected in 1999 

(Cicerello 2003, pers. comm.), and a 

single L individual was found in 2006 

(Butler 2007, p. 53). The snuffbox has 

become very rare, sporadic in 

occurrence, and its viability is 

questionable. 

Slate Creek—Slate Creek is a southern 

tributary of the Licking River below 

Cave Run Dam in east-central Kentucky. 

Historically, the snuffbox was 

considered ‘‘extremely abundant 

throughout the stream’’ (Taylor and 

Spurlock 1983) and collectively known 

from six sites (Laudermilk 1993, p. 45). 

Seventeen D specimens were recorded 

from a site in 1987 (Cicerello 2003, pers. 

comm.). A single FD and seven R 

specimens were found at three sites 

sampled in 1991 (Butler 2007, p. 53), 

when it was considered ‘‘occasional’’ in 

distribution (Laudermilk 1993, p. 45). 

Twelve L individuals were found in 

1992 (Cicerello 2003, pers. comm.). 

Subsequent sampling has produced no 

additional snuffbox; two sites and four 

sites yielded only R specimens in 2001 

and 2002, respectively (Cicerello 2005, 

pers. comm.). If extant, the population 

is marginal at best, with unlikely 

viability. 

Stillwater River—The Stillwater River 

is a 67-mi (108-km), western tributary of 

the Great Miami River draining 

southwestern Ohio. The species was 

collectively known from eight sites 

throughout the River (Watters 1988a, 

pp. 59–71; OSUM records). One FD 

specimen below Englewood Dam in 

Montgomery County was found among 

18 sites surveyed in 1987, with R shells 

from 5 other sites (Watters 1988a, pp. 

59–71). No other information on the 

small population is available, and its 

viability is unknown. 

Middle Fork Kentucky River—The 

Middle Fork is one of three headwater 


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tributaries (with the North and South 

Forks) forming the Kentucky River, 

flowing in a northerly then westerly 

direction and draining a portion of 

southeastern Kentucky. The snuffbox 

was first reported in 1966. Three L 

individuals and a R shell were found at 

three sites in 1996, and a single L 

individual was collected from another 

site in 1997 (Cicerello 2003, pers. 

comm.). All sites occur within a 

10-river-mi (16-river-km) reach above 

Buckhorn Reservoir in Leslie County. 

This small population has unknown 

viability. 

Red Bird River—The Red Bird River is 

a north-flowing headwater tributary of 

the South Fork Kentucky River in Clay 

County, southeastern Kentucky, forming 

the latter at its confluence with Goose 

Creek. Ten FD specimens were recorded 

from two sites in 1988, and three L and 

one FD snuffbox were collected from 

four sites in 1995 (Cicerello 2003, pers. 

comm.). This small population occurs 

sporadically in the lower 20 river mi 

(32 river km), and viability is unknown 

(Cicerello 2003, pers. comm.; 2006, pers. 

comm.). 

Red River—The Red (or North Fork 

Red) River is a westerly flowing 

tributary of the upper Kentucky River in 

eastern Kentucky. No L snuffbox were 

found in surveys of the 9-river-mi 

(15-river-km) reach of the Wild River 

section during surveys of 1980, 1986, 

and 1991 (Houp 1980, p. 56; 1993, 

p. 96), but two FD and one L snuffbox 

were found at three sites in 1988, while 

five L individuals were found in 1996 

(Cicerello 2006, pers. comm.). Mostly 

males have been found since 2002, and 

they are being held in captivity for 

future culture efforts (Butler 2007, 

p. 55). A small population persists over 

a 10-river-mi (16-river-km) reach in the 

lower section of the Red River Gorge 

Geological Area of the Daniel Boone 

National Forest in Menifee, Wolfe, and 

Powell Counties (Cicerello 2006, pers. 

comm.). Viability of this population is 

unknown. 

Rolling Fork Salt River—The Rolling 

Fork is a major southern tributary of the 

Salt River in central Kentucky, flowing 

in a northwesterly direction to join the 

Salt near its mouth. The snuffbox was 

first reported in 1958 (Rosewater 1959, 

p. 62). Seven FD specimens and a single 

L subadult were collected in 1988 from 

four sites in Larue, Marion, and Nelson 

Counties (Cicerello 2003, pers. comm.; 

Haag 2006, pers. comm.). A survey of 12 

mainstem and 30 tributary sites in the 

Rolling Fork system in 1998–99 yielded 

no evidence of the snuffbox, prompting 

an investigator to consider it extirpated 

(Akers 2000, p. 13), but occasional 

specimens may still be found (Butler 

2007, p. 55). The species is sporadically 

distributed over 40 river miles of the 

upper river (Cicerello 2006, pers. 

comm.). If it is still extant, the viability 

of this small population is unknown. 

Green River—A major southern 

tributary of the lower Ohio River, the 

Green River flows in a westerly 

direction and drains west-central 

Kentucky. Ortmann (1926, p. 182) 

considered the snuffbox to be well 

distributed over the system, but not 

abundant. Large museum collections of 

snuffbox were taken from Munfordville 

during 1961–66, but only six R shells 

were reported there in 1967. The 

snuffbox has been rare since. Five L and 

FD snuffbox were collected at 4 of 42 

sites during 1987–89 sampling in 

Mammoth Cave National Park (Cicerello 

and Hannan 1990, pp. 16–17). Three L 

and six FD snuffbox were reported in 

the upper Green River from 1984–90 

(Cicerello 2003, pers. comm.). A single 

L individual was collected in Taylor 

County in 1989 (Layzer 2009, pers. 

comm.), but no evidence of the snuffbox 

was reported at numerous other sites in 

1999, 2000, 2001, and 2003 (Cicerello 

2006, pers. comm.). Once abundant and 

occurring over 200 river mi (322 river 

km), the species has become 

exceedingly rare since the 1960s. 

Current snuffbox viability is unknown, 

and it may be nearing extirpation from 

the entire Green River system, where it 

was formerly known from eight 

tributaries. 

Wabash River System—The Wabash 

River is the second largest sub-basin 

within the Ohio River system, the 

watershed of the 350-mi (563-km) river 

encompassing much of Indiana, westcentral Ohio, and southeastern Illinois. 

The mainstem and at least 27 streams 

had one of the largest snuffbox 

population clusters. The species persists 

today as seven small populations in the 

system; the viability of these 

populations is unknown (Butler 2007, p. 

57). 

Salamonie River—The Salamonie 

River is a southern tributary of the 

upper Wabash River, flowing in a 

northwesterly direction and draining 

east-central Indiana. Two historical 

museum records were found. Nine sites 

were surveyed during 1993–94 without 

finding any evidence of the snuffbox 

(ESI 1995, p. 19). The snuffbox was 

rediscovered in 2004 above Salamonie 

Reservoir, where two L individuals at 

one site and FD shells, including a very 

small juvenile, were found at another 

site 2 mi (3 km) away (Fisher 2005, pers. 

comm.). The small population is 

considered to be recruiting and viable at 

some level. 

Tippecanoe River—The largest 

tributary of the upper Wabash River 

system, the Tippecanoe River drains 

north-central Indiana and flows 

westerly then southerly before joining 

the Wabash near Lafayette. Nearly all 

records of the snuffbox were made in 

the past 20 years. Two weathered shells 

were found in the lower mainstem 

among 16 sites sampled in 1987 

(Cummings et al. 1987, p. 25; Cummings 

and Berlocher 1990, p. 93) and 30 sites 

in 1991–92 (ESI 1993, p. 68). One L 

individual and over 32 FD specimens 

were found at a site at the upper end of 

Freeman Reservoir during a 1993 

drawdown that may have contributed to 

their demise (Fisher 2003, pers. comm.). 

A single FD specimen was found below 

Shafer Reservoir among 13 sites 

sampled in 2003 (ESI 2003, p. 9). The 

viability of this declining population is 

unknown, but it appears close to 

extirpation (Fisher 2003, pers. comm.). 

Embarras River—The Embarras River 

is a southerly flowing, western tributary 

of the lower Wabash River in 

southeastern Illinois. Museum lots 

represent collections dating to 1956 and 

contain snuffbox from nine mainstem 

and two tributary sites. A total of 9 L 

and 15 FD specimens were collected at 

four sites in 1986 in Coles and Douglas 

Counties (Cummings et al. 1988, p. 8). 

Although overall mussel abundance at 

the 21 sites sampled in both 1956 and 

1986 dropped 86 percent, the snuffbox 

was one of only five species that 

showed relatively stable population size 

over the 30-year period (Cummings et 

al. 1988, p. 9). Additional L and FD 

snuffbox from museum collections were 

recorded from single sites in 1988. 

Three L and eight FD snuffbox were 

found at two sites in 1992, and one L 

and three FD were found at three of six 

sites surveyed during 2001–2002. Since 

1986, the small snuffbox population has 

occurred sporadically at six sites over 

50 river mi (80 river km) of the upper 

river. The species was reported as 

significant and viable by Butler (2007 

pers. comm.), but has declined to some 

extent. Recent surveys, however, 

documented only one L individual in 

2005 and one L and one FD in 2008, 

indicating that the Embarras River 

population may be closer to a marginal 

population than a significant one 

(Tiemann 2009, pers. comm.). 

Sugar Creek—Sugar Creek is a 

tributary in the upper East Fork White 

River system, draining central Indiana 

east and south of Indianapolis. A single 

L individual from one site, FD 

specimens from seven sites, and R shells 

from an additional eight sites were 

reported in 1990 (Harmon 1992, pp. 40– 

41 1998). The snuffbox population 


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occurred sporadically over 35 river mi 

(56 km) to near the mouth. Only R shells 

were found while resampling some 

historical sites in 1995, 1998, and 2001 

(Butler 2007, p.59). It is questionable 

whether the population remains extant. 

Buck Creek—Buck Creek is a 

southerly flowing, western tributary of 

Sugar Creek in the upper East Fork 

White River system east of Indianapolis. 

A FD snuffbox was found near the 

mouth and R specimens at an upstream 

site in 1990 (Harmon 1992, p. 41. 

Similar to the parent stream population 

in Sugar Creek, the snuffbox may 

already be extirpated in Buck Creek 

(Fisher 2003, pers. comm.). 

Muscatatuck River—The Muscatatuck 

River is a large, westerly flowing 

tributary of the upper East Fork White 

River in southeastern Indiana. The 

snuffbox was first reported from the 

stream by Daniels (1903, p. 646). FD 

specimens (unknown number) were 

recorded at a site downstream from 

Graham Creek that was sampled in 1988 

(Harmon 1989, p. 118). Status and 

viability of snuffbox in the Muscatatuck 

River are unknown. 

Graham Creek—Graham Creek flows 

southwesterly to join Big Creek in 

forming the Muscatatuck River in the 

East Fork White River system in 

southeastern Indiana. The species was 

found FD (numbers unknown) at six 

sites over 10 river mi (16 river km) of 

the lower stream in Jennings County in 

1988 (Harmon 1989, p. 117), and a 

single FD specimen was found in 1990 

(Harmon 1998). Viability of these small 

population is unknown. 

Cumberland River System—Snuffbox 

populations are known from the 

mainstem Cumberland River and 6 of its 

tributaries. With few exceptions, most 

mainstem records were made prior to 

the 1920s when the species was locally 

common (Wilson and Clark 1914, p. 45). 

The snuffbox is considered extirpated 

from the mainstem. Currently, a single 

tributary population may be extant, but 

is considered not viable. The species is 

likely to become extirpated from the 

entire river system in the foreseeable 

future. 

Buck Creek—Buck Creek is a 

southerly flowing, northern tributary of 

the upper Cumberland River below 

Cumberland Falls in southeastern 

Kentucky. One D valve was found at a 

site in 1981 (Clarke 1981b, Appendix), 

and two L and one FD snuffbox were 

reported from three sites during 1983– 

84 (Schuster et al. 1989, p. 82). The 

species was also reported L from a lower 

mainstem site among seven sites 

sampled from 1987–90 (Layzer and 

Anderson 1992, p. 16). A recent survey 

found only R shells at 3 of 23 sites 

(Hagman 2000, p. 21). If extant, the 

declining snuffbox population in Buck 

Creek is likely to become extirpated in 

the foreseeable future. 

Tennessee River System 

The Tennessee River is the largest 

tributary of the Ohio River, draining 

seven southeastern States and joining 

the Ohio near its mouth in western 

Kentucky. The snuffbox originally was 

known from throughout all but the 

lower section of river and 17 of its 

tributaries. Hundreds of miles of large 

river habitat on the mainstem have been 

lost under nine reservoirs, with 

additional dams on several tributaries 

(Clinch, Holston, and Elk Rivers) 

(Tennessee Valley Authority (TVA) 

1971, p. 4). The loss of mussel resources 

has been substantial (Watters 2000, p. 

262). Muscle Shoals, the 53-river-mi 

(85-river-km) reach in northwestern 

Alabama, historically harbored 69 

mussel species, the most diverse mussel 

fauna ever known (Garner and 

McGregor 2001, p. 155). The 

construction of three dams (Wilson in 

1925, Wheeler in 1930, and Pickwick 

Landing in 1940) inundated most of the 

mussel beds. No L snuffbox have been 

reported at Muscle Shoals for around 

100 years (Garner and McGregor 2001, 

p. 162). The snuffbox may persist in the 

mainstem at a very low density and in 

only five tributaries. The Clinch River 

maintains a stronghold population, but 

highly restricted populations persist in 

the other streams. 

Clinch River—The 350-mi (563-km) 

Clinch River is a major tributary of the 

upper Tennessee River originating in 

southwestern Virginia, and flowing in a 

southwesterly direction to its 

confluence near Knoxville in 

northeastern Tennessee. No other river 

in North America has extant 

populations of more federally 

endangered (15) and candidate (4) 

species of mussels than does the upper 

Clinch River above Norris Reservoir. 

The snuffbox was reported from nine 

sites by Ortmann (1918, pp. 601–606). 

Museum records from Hancock County, 

Tennessee, during 1965–71 documented 

a very large population of snuffbox. The 

snuffbox is generally distributed from 

RM 170 to RM 195 in Hancock County, 

but is sporadic in Virginia (RM 213– 

235), where it has recently declined 

(Butler 2007, p. 62). The snuffbox 

population is recruiting, viable, and 

currently stable, although decreased in 

size and range from 40 years ago. The 

Clinch River ranks among the six 

stronghold snuffbox populations 

rangewide. 

Powell River—The Powell River is the 

major tributary of the upper Clinch 

River flowing in a southwesterly 

direction parallel to and northwest of 

the Clinch River in southwestern 

Virginia and northeastern Tennessee. 

The snuffbox was reported at three sites 

by Ortmann (1918, pp. 597–598), five 

sites during 1973–78 by Dennis (1981, 

p. 3), four sites from 1975–78 by 

Ahlstedt and Brown (1979, p. 42), and 

four Virginia sites in 1988–89 by 

Wolcott and Neves (1994, p. 7). Large 

collections attest to its former 

abundance. The species was found L 

and FD in the Powell River, Tennessee, 

during 1989–90 (Hubbs et al. 1991, 

Appendix A). Johnson (2008) collected 

two L individuals at RM 95. The 

population has declined, viability is 

questionable, and its extirpation may be 

imminent (Butler 2007, p. 63). 

Tennessee River—The snuffbox 

originally was known from all but the 

lower section of the river. Butler (2007, 

p. 61) reported the snuffbox as ‘‘believed 

to be extirpated from the entire 

Tennesssee River.’’ However, Yokley 

(2002, p. 1) collected a single FD male 

in 2002 at the U.S. 231 Bridge, Madison 

and Morgan Counties. In 2006, one L 

female was found at the same location, 

though it was the only snuffbox out of 

8,978 mussels collected at the site 

(Yokley 2006, p. 1). Nothing further is 

known about the status of the snuffbox 

in the Tennessee River mainstem. 

Paint Rock River—The Paint Rock 

River is a southerly flowing, northern 

tributary of the southern bend of the 

Tennessee River in northeastern 

Alabama and adjacent Tennessee. The 

snuffbox was first reported from one of 

six mainstem sites by Ortmann (1925, p. 

359). No evidence of snuffbox was 

found in two surveys during 1965–67 

(Isom and Yokley 1973, p. 444) and a 

1980 survey (Butler 2007, p. 64). Twelve 

L and FD snuffbox were found at four 

sites between RMs 13 and 21 (Ahlstedt 

1995–96, p. 70). The species was again 

absent from 10 upper mainstem sites 

surveyed in 2002 (Godwin 2002, p. 9). 

Four FD specimens of varying sizes 

were found at lower river sites in 2002 

(Fraley 2003, pers. comm.; Smith 2005, 

pers. comm.) and 2003–2006 (Freeman 

2006, pers. comm.). One L and 11 FD 

specimens were found at RM 21 in 

2005, and 2 L and 16 FD were collected 

at RM 31 in 2007 (Gangloff 2007, pers. 

comm.). In July 2008, Freeman (2008, 

pers. comm.) observed multiple age 

classes (sizes) of FD snuffbox in 

middens between RM 34.7 and 32.5. 

Fobian et al. (2008, p. 14) collected 21 

L snuffbox at 7 sites and FD specimens 

at 8 sites between RM 46.7 and 13.1. 

The stronghold snuffbox population 

exists between RMs 13 and 44, and is 


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recruiting, viable, and has clearly 

improved since 1980. 

Elk River—The Elk River is a large, 

northern tributary flowing 200 river mi 

(322 river km) in a southwesterly 

direction in the southern bend of the 

Tennessee River in south-central 

Tennessee and north-central Alabama. 

Snuffbox collections have been 

sporadic. The species was found at 2 

sites in the mid-1960s (Isom et al. 1973, 

p. 440), and a single L individual was 

found among 108 sites sampled in 1980 

(Ahlstedt 1983, p. 47). Single specimens 

were also reported from 4 sites sampled 

in the lower river in 1997 (Madison and 

Layzer 1998, Table 6) and 16 sites 

sampled in 1999 (Service 1999, p. 3). A 

very large FD specimen was found at 

RM 51 among 4 sites sampled in 2001 

(Hubbs 2002, p. 5; Butler 2007, p. 65). 

A single L and a FD snuffbox were 

found at a site in Giles County during 

qualitative sampling events at five sites 

in 2005 (Ahlstedt et al. 2006). Ford 

(2008, pers. comm.) reported collecting 

FD specimens at Stairstep Shoals in 

Giles County, Tennessee, in July 2007. 

The small snuffbox population has 

recently recruited, exhibits some level 

of viability, and its numbers appear 

relatively stable in recent history. 

Duck River—The Duck River is the 

downstream-most large tributary of the 

Tennessee River draining south-central 

Tennessee and flowing 285 river miles 

(459 river km) west to its confluence 

near the head of Kentucky Reservoir. 

The snuffbox historically occurred 

throughout the Duck River and, based 

on museum records, was locally 

common 40 to 50 years ago, but was 

absent in surveys from RM 180 

downstream in the mid-1970s (Ahlstedt 

1981, p. 62; Dennis 1984, p. 38). Two L 

individuals were collected from 2 of 99 

sites surveyed in 1979 (Butler 2007, p. 

66). A single L individual was 

discovered in Maury County among 72 

sites sampled during 2000–03 (Ahlstedt 

et al. 2004, p. 119), but none were found 

at 11 lower sites surveyed in 2000 

(Schilling and Williams 2002, p. 409). 

The snuffbox is very rare, and its 

viability is uncertain. 

Lower Mississippi River Sub-Basin 

The Lower Mississippi River Subbasin includes 954 miles of the 

Mississippi River from its confluence 

with the Ohio River at Cairo, Illinois, to 

its mouth in the Gulf of Mexico. The 

snuffbox is known from a single stream 

in this sub-basin, outside of the White 

River system. 

St. Francis River—The St. Francis 

River is a major tributary of the lower 

Mississippi with its headwaters in 

southeastern Missouri, and flowing 

south into northeastern Arkansas. The 

only Arkansas records available for this 

450-mi (724-km) river are from 1964, 

located approximately 1 mi southwest 

of Parkin in Cross County (Bates and 

Dennis 1983, p. 63; Harris et al. 2007, 

p. 10). Snuffbox records exist for Butler, 

Wayne, and Stoddard Counties, 

Missouri, where it was considered 

‘‘locally abundant’’ (Oesch 1984, p. 235). 

The species is known from above 

Wappapello Reservoir, but was absent 

from Missouri surveys conducted below 

Wappapello Dam in 1983 (Bates and 

Dennis 1983, p. 63) and 1986 (Ahlstedt 

and Jenkinson 1991, p. 240). Twelve L 

snuffbox were sampled at sites in 2002 

(Hutson and Barnhart 2004, pp. 84–85). 

Live individuals were found during 

collections at RM 172.1 in 2005 and 

2006 (Butler 2007, p. 67). The snuffbox 

is restricted to a 10-mi (16-km) reach 

(RM 172.1–182.0) on the northeastern 

edge of the Ozark Plateaus in the 

vicinity of Sam A. Baker State Park, 

Wayne County (Hutson and Barnhart 

2004, p. 85). This medium-sized 

snuffbox population appears to be stable 

and viable, but restricted in distribution. 

White River System—The 690-mi 

(1,110-km) White River is a large 

tributary system of the western bank of 

the Mississippi River. A snuffbox 

population once occurred in the 

mainstem and six of its larger 

tributaries. The last record from the 

mainstem in Arkansas is pre-1921 

(Harris et al. 2007, p. 10). Highly 

restricted populations persist in four 

streams. 

Buffalo River—The Buffalo River is a 

large, eastward-flowing tributary of the 

middle White River in north-central 

Arkansas. The snuffbox was not found 

during surveys in 1910 (26 sites; Meek 

and Clark 1912, p. 13) and 1995 (40 

sites; Harris 1996, p. 9), but two L 

individuals were found at a single site 

among 60 sites surveyed in 2006 

(Matthews 2007, pers. comm.). The 

small population occurs in the lower 

river in Marion County, and its viability 

is unknown. 

Black River—The Black River is the 

largest tributary in the White River 

system, draining much of southeastern 

Missouri and northeastern Arkansas 

before flowing in a southerly direction 

into the White River near Newport, 

Arkansas. A long but sporadic collection 

history for the snuffbox appears in the 

300-mi (483-km) Black River. A single, 

approximately 4-year-old L male was 

collected at RM 65.5, Wayne County, 

among 51 Missouri sites sampled in 

2002 (Hutson and Barnhart 2004, p. 

154). The species has become extirpated 

from the lower river on the Mississippi 

Embayment, including Arkansas. The 

snuffbox appears rare but viable at some 

level. 

Spring River—The Spring River is a 

large tributary of the Black River that 

drains the eastern Ozark Plateaus in 

south-central Missouri and northeastern 

Arkansas. Based on pre-1986 records, 

the snuffbox was known in low 

numbers from at least four sites in 

approximately 20 river mi (34 river km) 

of the lowermost mainstem in Arkansas 

(Harris and Gordon 1987, p. 53). A 

single L adult male was found in 

Lawrence County in 2005, and 

represents the first L specimen found in 

Arkansas in more than 20 years (Butler 

2007, p. 69). Further, 53 FD snuffbox 

were collected in four large muskrat 

middens (Harris et al. 2007, p. 15). The 

extent of the population is not known, 

but it is probably limited to relatively 

few miles in the lower mainstem in 

Lawrence and Randolph Counties. This 

population appears small, and its status 

and viability are unknown. 

Strawberry River—The Strawberry 

River is a western tributary of the Black 

River draining a portion of the 

southeastern Ozark Plateaus in 

northeastern Arkansas. The only 

snuffbox records were from around 1983 

and 1997 in the middle mainstem in 

Sharp County (Butler 2007, p. 69). No 

other details on these collections or the 

status of the population are known. 

Considering the dearth of records, the 

snuffbox appears to be very rare in the 

Strawberry River, and of unknown 

viability. 

Summary of Snuffbox Population 

Estimates and Status 

The snuffbox has declined rangewide 

and appears to be extant in 74 of 208 

streams and lakes of historical 

occurrence, a 65 percent decline in 

occupied streams. Realistically, much 

more than 65 percent of the habitat 

historically available for this species no 

longer supports its populations. Habitat 

losses measured in the thousands of 

miles have occurred rangewide. Since 

multiple streams may comprise single 

snuffbox population segments (for 

example, the French Creek system), the 

actual number of extant populations is 

somewhat less. Extant populations, with 

few exceptions, are highly fragmented 

and restricted to short reaches. The 

elimination of this species from scores 

of streams and thousands of miles of 

stream reaches indicates catastrophic 

population losses and a precipitous 

decline in overall abundance. It is 

reasonable to estimate that total range 

reduction and overall population losses 

for the snuffbox each approximate, if 

not exceed, 90 percent. 


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Previous Federal Action 

We identified the rayed bean as a 

Category 2 species in a notice of review 

published in the Federal Register on 

May 22, 1984 (49 FR 21664). The rayed 

bean remained a Category 2 species in 

subsequent notices including January 6, 

1989 (54 FR 554), November 21, 1991 

(56 FR 58804), and November 15, 1994 

(59 FR 58982). Prior to 1996, a Category 

2 species was one that we were 

considering for possible addition to the 

Federal List of Endangered and 

Threatened Wildlife but for which 

conclusive data on biological 

vulnerability and threats were not 

available to support a proposed rule. We 

stopped designating Category 2 species 

in the February 28, 1996, Notice of 

Review (61 FR 7596). We now define a 

candidate species as a species for which 

we have on file sufficient information to 

propose it for protection under the Act. 

We designated the rayed bean as a 

candidate species on May 4, 2004 (69 

FR 24876). 

We identified the snuffbox as a 

Category 2 species in the notice of 

review published in the Federal 

Register on November 21, 1991 (56 FR 

58804). The snuffbox remained a 

Category 2 in the subsequent notice on 

November 15, 1994 (59 FR 58982) but 

was dropped from the list in the 

February 28, 1996, Notice of Review (61 

FR 7596), when we stopped designating 

Category 2 species. The snuffbox is not 

currently listed as a candidate species 

for listing. 

Summary of Information Pertaining to 

the Five Factors 

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 Their Habitat or Range. 

Both species have experienced 

significant curtailment of their occupied 

habitats (see Background, above). The 

rayed bean has been eliminated from 

about 74 percent of the streams it 

historically occurred in. This species 

has also been eliminated from long 

reaches of former habitat in hundreds of 

miles of the Maumee, Ohio, Wabash, 

and Tennessee Rivers and from 

numerous stream reaches in their 

tributaries. The snuffbox has been 

eliminated from about 65 percent of the 

streams in which it historically 

occurred. Furthermore, extant 

populations, with few exceptions, are 

highly fragmented and restricted to 

short reaches. Available records indicate 

that 33 percent of streams considered to 

harbor extant populations of the 

snuffbox are represented by only one or 

two recent L or FD individuals. The 

primary cause of range curtailment for 

both species has been modification and 

destruction of river and stream habitats, 

primarily by the construction of 

impoundments. 

Impoundment—Impoundments result 

in the dramatic modification of riffle 

and shoal habitats and a resulting loss 

of mussel resources, especially in larger 

rivers. Neves et al. (1997, pp. 63–64) 

and Watters (2000, pp. 261–262) 

reviewed the specific effects of 

impoundments on freshwater mollusks. 

Dams interrupt a river’s ecological 

processes by modifying flood pulses; 

controlling impounded water 

elevations; altering water flow, 

sediments, nutrients, and energy inputs 

and outputs; increasing depth; 

decreasing habitat heterogeneity; 

decreasing stability due to subsequent 

sedimentation; blocking host fish 

passage; and isolating mussel 

populations from fish hosts. Even small, 

low-head dams can have some of these 

effects on mussels. 

The reproductive process of riverine 

mussels is generally disrupted by 

impoundments, making the rayed bean 

and snuffbox unable to successfully 

reproduce and recruit under reservoir 

conditions. Population losses due to 

impoundments have likely contributed 

more to the decline and imperilment of 

the rayed bean and snuffbox than has 

any other single factor. Neither species 

occurs in reservoirs lacking riverine 

characteristics, and only the snuffbox 

persists in large rivers with dams (Ohio 

River), and then only in sections 

retaining riverine characteristics 

(generally tailwaters). Both species, 

however, historically occurred in the 

wave-washed shallows of several glacial 

lakes, an environment very different 

from that found in impoundments. 

Stream habitat throughout major 

portions of the range of both species has 

been impounded. The majority of the 

Tennessee and Cumberland River 

mainstems 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 is 

either directly impounded by U.S. Army 

Corps of Engineers (Corps) structures or 

otherwise impacted by cold tailwater 

released from dams. Watters (2000, pp. 

262–263) summarizes the tremendous 

loss of mussel species from various 

portions of the Tennessee and 

Cumberland River systems. The rayed 

bean has been eliminated from the 

Tennessee River system and the 

snuffbox, once widespread throughout 

both systems, now persists in only five 

Tennessee River tributaries and one 

Cumberland River tributary. 

This impoundment scenario is similar 

in many other parts of the range of the 

rayed bean and snuffbox, and includes 

numerous navigational locks and dams 

(Ohio, Allegheny, Muskingum and 

Green Rivers), major dams (Shenango, 

Elk, Walhonding, Scioto, Little Miami, 

Green, Nolin, Barren, Tippecanoe, 

Wabash, Mississinewa, Salamonie, and 

Duck Rivers), and low-head dams (Pine, 

Belle, Clinton, Huron, Maumee, 

Auglaize, Sandusky, Mahoning, 

Tuscarawas, Walhonding, Scioto, 

Olentangy, Wabash, Mississinewa, East 

Fork White, West Fork White, and Duck 

Rivers; and Middle Island, Big Walnut, 

Alum, Big Darby, Little Darby, Sugar, 

and Richland Creeks) that have 

contributed to the loss of the species’ 

habitat. Sediment accumulations behind 

dams of all sizes generally preclude the 

occurrence of the rayed bean and 

snuffbox. 

Dredging and Channelization— 

Dredging and channelization activities 

have profoundly altered riverine 

habitats nationwide. Hartfield (1993, pp. 

131–141), Neves et al. (1997, pp. 71–72), 

and Watters (2000, pp. 268–269) 

reviewed the specific effects of 

channelization on freshwater mollusks. 

Channelization impacts a stream’s 

physical (accelerated erosion, reduced 

depth, decreased habitat diversity, 

geomorphic instability, and riparian 

canopy loss) and biological (decreased 

fish and mussel diversity, changed 

species composition and abundance, 

decreased biomass, and reduced growth 

rates) characteristics (Hartfield 1993, p. 

131; Hubbard et al. 1993, pp. 136–145). 

Channel construction for navigation has 


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been shown to increase flood heights 

(Belt 1975, p. 189). This is partially 

attributed to a decrease in stream length 

and increase in gradient (Hubbard et al. 

1993, p. 137). 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 

benthic (bottom-dwelling) organisms 

such as the rayed bean and snuffbox. 

The only known rayed bean 

populations that remain in navigation 

channels are in the upper two 

navigation pools of the Allegheny River. 

Activities associated with navigation 

channels may have contributed to the 

elimination of the rayed bean from the 

Ohio, lower Allegheny, and Muskingum 

Rivers, and potentially others. Channel 

maintenance operations for barge 

navigation have impacted habitat for the 

snuffbox in several large rivers. Impacts 

associated with barge traffic, which 

include construction of fleeting areas, 

mooring cells, docking facilities, and 

propeller wash, also disrupt habitat. 

Navigation maintenance activities may 

continue to adversely affect this species 

in the upper Ohio River. Hundreds of 

miles of rayed bean (Olentangy, 

Salamonie, Mississinewa, Vermilion, 

North Fork Vermilion, Embarras Rivers) 

and snuffbox (Grand, Kankakee, 

Sangamon, Kaskaskia, Olentangy, 

Salamonie, Mississinewa, Eel, 

Vermilion, and North Fork Vermilion, 

Embarras, Paint Rock, and St. Francis 

Rivers; and Tonawanda, Killbuck, 

Chickamauga, and Bear Creeks) streams 

were dredged and channelized decades 

ago, and some populations have been 

eliminated from these streams. The 

entire length of the Kankakee River in 

Indiana was channelized by 1917. In 

addition, hundreds of drains (formed 

from ditching low-gradient creeks and 

swales) were created around 100 years 

ago in Illinois, Michigan, and other 

midwestern States. Stream 

channelizations were attempts to reduce 

flooding, drain low-lying areas, and 

‘‘improve’’ storm flow runoff. 

Chemical Contaminants—Chemical 

contaminants are ubiquitous throughout 

the environment and are considered a 

major threat in the decline of freshwater 

mussel species (Cope et al. 2008, p. 451; 

Richter et al. 1997, p. 1081; Strayer et 

al. 2004, p. 436; Wang et al. 2007, p. 

2029). Chemicals enter the environment 

through both point and nonpoint 

discharges, including spills, industrial 

sources, municipal effluents, and 

agricultural runoff. These sources 

contribute organic compounds, heavy 

metals, pesticides, and a wide variety of 

newly emerging contaminants to the 

aquatic environment. As a result, water 

and sediment quality can be degraded to 

the extent that mussel populations are 

adversely impacted. 

Chemical spills can be especially 

devastating to mussels because they 

may result in exposure of a relatively 

immobile species to extremely elevated 

concentrations that far exceed toxic 

levels and any water quality standards 

that might be in effect. Some notable 

spills that released large quantities of 

highly concentrated chemicals resulting 

in mortality to mussels include: massive 

mussel kills on the Clinch River at 

Carbo, Virginia, occurred from a power 

plant alkaline fly ash pond spill in 1967 

and a sulfuric acid spill in 1970 

(Crossman et al. 1973, p. 6); 

approximately 18,000 mussels of several 

species including 750 individuals from 

three endangered mussel species were 

eliminated from the upper Clinch River 

near Cedar Bluff, Virginia, in 1998, 

when an overturned tanker truck 

released 1,600 gallons (6,056 liters) of a 

chemical used in rubber manufacturing 

(Jones et al. 2001, p. 20; Schmerfeld 

2006, p. 12); and an ongoing release of 

sodium dimethyl dithiocarbamate, a 

chemical used to reduce and precipitate 

hexachrome, starting in 1999 impacted 

approximately 10 river miles (16 km) of 

the Ohio River and resulted in an 

estimated loss of one million mussels, 

including individuals from two 

federally listed species (DeVault 2009, 

pers. comm.; Clayton 2008, pers. 

comm.). These are not the only 

instances where chemical spills have 

resulted in the loss of high numbers of 

mussels (Brown et al. 2005, p. 1457; 

Neves 1991, p. 252; Jones et al. 2001, p. 

20; Schmerfeld 2006, pp. 12–13), but are 

provided as examples of the serious 

threat chemical spills pose to mussel 

species. The rayed bean and snuffbox 

are especially threatened by chemical 

spills because these spills can occur 

anywhere there are highways with 

tanker trucks, industries, or mines and 

where these overlap with rayed bean 

and snuffbox distribution. 

Exposure of mussels to lower 

concentrations of contaminants more 

likely to be found in aquatic 

environments can also adversely affect 

mussels and result in the decline of 

freshwater mussel species. Such 

concentrations may not be immediately 

lethal, but over time, can result in 

mortality, reduced filtration efficiency, 

reduced growth, decreased 

reproduction, changes in enzyme 

activity, and behavioral changes to all 

mussel life stages. Frequently, 

procedures which evaluate the ‘‘safe’’ 

concentration of an environmental 

contaminant (for example, national 

water quality criteria) do not have data 

for freshwater mussel species or exclude 

data that is available for freshwater 

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; Wang, 2007a, pp. 2041– 

2046; Valenti 2005, pp. 1244–1245; 

Valenti 2006, pp. 2514–2517; March 

2007, pp. 2068–2073) and juveniles 

(Bartsch et al. 2003, p. 2561; Augspurger 

et al. 2003, p. 2569; Mummert et al. 

2003, p. 2549, Wang, 2007b, pp. 2053– 

2055, Wang, 2007a, pp. 2041–2046, 

Valenti 2005, pp. 1244–1245; Valenti 

2006, pp. 2514–2517; March 2007, pp. 

2068–2073) 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, 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. 2007a, p. 2087; 

Bringolf et al. 2007c, p. 2101; Wang et 

al. 2007a, p. 2029; Wang et al. 2007b, p. 

2036; Wang et al. 2007c, p. 2048). 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 sources (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) (Goudreau et al. 1993, 

p. 212; Hickey and Martin 1999, p. 44; 

Augspurger et al. 2003, p. 2569; 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, high level of toxicity, and 

because the highest concentrations 


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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 Environmental Protection 

Agency’s established ammonia water 

quality criteria (EPA 1985, p. 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 and 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 (Naimo 

1995, pp. 351–355; Keller and Zam 

1991, p. 543; Jacobson et al. 1997, p. 

2390; 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 

2007a, pp. 2036–2047; Wang 2007b, pp. 

2048–2056). 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’s 1996 chronic 

water quality criterion of 15 ug/L 

(hardness 170 mg/L) for copper (Wang 

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 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 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 greater than 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 (http://

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 (112-km) 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 and 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 

spaces in the substrate where lower 

dissolved oxygen concentrations are 

more likely than on the sediment 

surface where adults tend to live 

(Sparks and 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 

mussels, 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 rayed bean and snuffbox. 

A potential, but undocumented, threat 

to freshwater mussel species, including 

rayed bean and snuffbox, 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 (NPDES)) 

and non-permitted sites throughout the 

country. Permitted discharge sites are 


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ubiquitous in watersheds with rayed 

bean and snuffbox 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 a 

declining population of snuffbox; 

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 the rayed bean and 

snuffbox. This information indicates the 

potential for contaminants to contribute 

to declining rayed bean and snuffbox 

populations—from spills that are 

immediately lethal to species to chronic 

contaminant exposure, which results in 

death, reduced growth, or reduced 

reproduction of rayed bean and 

snuffbox. 

Mining—The low pH commonly 

associated with coal mine runoff can 

reduce glochidial encystment rates, thus 

impacting mussel recruitment (Huebner 

and Pynno ̈nen 1992, p. 2350). 

Additionally, adverse impacts from 

heavy metal-rich drainage from coal 

mining and associated sedimentation 

has been documented in portions of 

historical rayed bean and snuffbox 

habitat in the upper Ohio River system 

in western Pennsylvania (Ortmann 

1909c, p. 97), West Virginia, and 

southeastern Ohio. Likewise, coal 

mining has impacted rayed bean habitat 

in the upper Tennessee River system, 

Virginia (Kitchel et al. 1981, p. 21), and 

snuffbox habitat in eastern Kentucky 

(lower Ohio and Mississippi River 

systems in southeastern Illinois and 

western Kentucky; upper Cumberland 

River system in southeastern Kentucky 

and northeastern Tennessee; and upper 

Tennessee River system in southwestern 

Virginia) (Ortmann 1909c, p. 103; Neel 

and Allen 1964, pp. 428–430; Kitchel et 

al. 1981, p. 21; Anderson et al. 1991, pp. 

6–7; Gordon 1991, p. 2; Bogan and Davis 

1992, p. 2; Layzer and Anderson 1992, 

pp. 91–94; Ahlstedt and Tuberville 

1997, p. 75; Milam et al. 2000, p. 53; 

Warren and Haag 2005, p. 1394). Acid 

mine drainage was implicated in the 

mussel die-off in the Little South Fork 

Cumberland River, Kentucky (Anderson 

et al. 1991, pp. 6–7; Layzer and 

Anderson, 1992, p. 94; Ahlstedt and 

Saylor 1995–96, pp. 92–93; Warren and 

Haag 2005, p. 1394). Tailings pond 

failures have also impacted aquatic 

resources (Powell River, Virginia; Butler 

2007, p. 83). A decline of the snuffbox 

and other imperiled mussels in the 

Powell River was blamed on coal 

mining impacts (Ahlstedt and 

Tuberville 1997, p. 75). Increased 

mining activities in the upper Clinch 

River system is resulting in ‘‘blackwater’’ 

events (Jones and Neves 2004, p. 2). 

Anecdotal evidence suggests that coal 

fines are increasing in the Clinch River 

reach that harbors a stronghold snuffbox 

population (Butler 2007, p. 84). A coalfired power plant planned for the upper 

Clinch River in Virginia would further 

increase mining in the Clinch and 

Powell watersheds. 

Currently, coal mining activities occur 

only in the Elk River in West Virginia 

(Douglas 2010, pers. comm.). However, 

if coal mining activities are reinitiated 

in western Pennsylvania, they could 

become a threat to populations of both 

species in the lower French Creek and 

the Allegheny River. 

Instream and alluvial (clay, silt, sand, 

or other material deposited by running 

water) gravel mining has been 

implicated in the destruction of several 

mussel populations (Hartfield 1993, pp. 

135–136; Brown and Curole 1997, pp. 

239–240). Negative impacts associated 

with gravel mining include stream 

channel modifications (altered habitat, 

disrupted flow patterns, sediment 

transport), water quality modifications 

(increased turbidity, reduced light 

penetration, increased temperature), 

macroinvertebrate population changes 

(elimination, habitat disruption, 

increased sedimentation), and changes 

in fish populations (impacts to 

spawning and nursery habitat, food web 

disruptions) (Kanehl and Lyons 1992, 

pp. 26–27; Roell 1999, p. 5). Gravel 

mining may continue to be a localized 

threat to rayed bean and snuffbox 

populations (Kankakee, Bourbeuse, 

Walhonding, Elk (Tennessee), and 

Strawberry Rivers; Big Darby and Buck 

(Kentucky) Creeks). 

Other mining activities that impact 

snuffbox populations include mining for 

metals (lead, cadmium, zinc) in 

Missouri. Mining has been implicated in 

the decline of mussels from the upper 

St. Francis River (Hutson and Barnhart 

2004, pp. 86–87). Lead and barite 

mining is common in the Big River, a 

Meramec River tributary. A tailingspond blowout discharged 81,000 cubic 

yards of mine tailings in 1977 that 

impacted approximately 80 river mi 

(129 river km) (Buchanan 1980, p. 9; 

Roberts and Bruenderman 2000, p. 24). 

As of 2000, high levels of heavy metals 

were still detected in the system 

(Roberts and Bruenderman 2000, p. 24) 

and may continue to hinder stream 

recovery. Forty-five tailings ponds and 

numerous tailings piles remain in the 

watershed (Roberts and Bruenderman 

2000, p. 24). 

Oil and gas production may have 

contributed to the decline of the rayed 

bean and snuffbox in certain drainages 

(Sangamon River in the upper 

Mississippi River system; Slippery Rock 

and Connoquenessing Creeks in the 

upper Ohio River system; Green, 

Kentucky, Salamonie, and Mississinewa 

Rivers in the lower Ohio River system) 

(Ortmann 1909c, p.104; Schanzle and 

Cummings 1991, p. 1; ESI 1995, p. 39; 

Cicerello 1999, p. 11). Pollutants 

include brines, high levels of potassium, 

and numerous organic compounds 

(Imlay 1971, p. 39). An increasing 

demand for domestic energy resources 

is expected to accelerate oil and gas 

exploration in certain rayed bean and 

snuffbox streams in the foreseeable 

future. 

Siltation—Excessive sedimentation 

affects an estimated 46 percent of all 

U.S. streams (Judy et al. 1984), 

including the majority of the streams 

with extant rayed bean and snuffbox 

populations. Sedimentation has been 

implicated in the decline of mussel 

populations nationwide, and is a threat 

to rayed bean and snuffbox (Kunz 1898, 

p. 328; Ellis 1936, pp. 39–40; Marking 

and Bills 1979, p. 204; Vannote and 

Minshall 1982, pp. 4105–4106; Dennis 

1984, p. 212; Wolcott and Neves 1990, 

pp. 74–75; Brim Box 1999, p. 79; Fraley 

and Ahlstedt 2000, p. 194; Poole and 

Downing 2004, pp. 119–120). Specific 

biological impacts include reduced 

feeding and respiratory efficiency from 

clogged gills, disrupted metabolic 

processes, reduced growth rates, limited 

burrowing activity, and physical 

smothering (Ellis 1936, pp. 39–40; 

Stansbery 1971, p. 6; Imlay 1972, p. 76; 

Marking and Bills 1979, p. 210; Vannote 

and Minshall 1982, p. 4105; Waters 

1995, p. 7). 

Studies indicate that excessive 

sediment level impacts are sublethal, 

with detrimental effects not 

immediately apparent (Brim Box and 

Mossa 1999, p. 101). Physical habitat 

effects include altered suspended and 

bed material loads, and bed sediment 

composition associated with increased 

sediment production and run-off; 

clogged interstitial habitats and reduced 

interstitial flow rates and dissolved 

oxygen levels; changed channels in 

form, position, and degree of stability; 

altered depth or width-depth ratio that 

affects light penetration and flow 

regime; aggraded (filling) or degraded 

(scouring) channels; and changed 

channel positions that dewater mussel 

beds (Vannote and Minshall 1982, p. 

4105; Gordon et al. 1992, pp. 296–297; 

Kanehl and Lyons 1992, pp. 26–27; 

Brim Box and Mossa 1999, p. 102). 


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Interstitial spaces in the substrate 

provide essential habitat for juvenile 

mussels. When clogged, interstitial flow 

rates and spaces may become reduced 

(Brim Box and Mossa 1999, p. 100), thus 

reducing juvenile habitat availability. 

The rayed bean burrows deep into 

interstitial substrates, making it 

particularly susceptible to degradation 

of this habitat. Sediment may act as a 

vector for delivering contaminants such 

as nutrients and pesticides to streams. 

Juveniles can readily ingest 

contaminants adsorbed to silt particles 

during normal feeding activities. These 

factors may explain, in part, why so 

many mussel populations, including 

those of the rayed bean and snuffbox, 

appear to be experiencing recruitment 

failures. 

Agricultural activities produce the 

most significant amount of sediment 

that enters streams (Waters 1995, pp. 

17–18). Neves et al. (1997, p. 65) stated 

that agriculture (including both 

sediment and chemical run-off) affects 

72 percent of the impaired river miles 

in the country. Unrestricted access by 

livestock is a significant threat to many 

streams and their mussel populations 

(Fraley and Ahlstedt 2000, p. 193). Soil 

compaction for intensive grazing may 

reduce infiltration rates and increase 

run-off, and trampling of riparian 

vegetation increases the probability of 

erosion (Armour et al. 1991, pp. 8–10; 

Trimble and Mendel 1995, pp. 238–239; 

Brim Box and Mossa 1999, p. 103). 

The majority of extant rayed bean and 

snuffbox populations are threatened by 

some form of agricultural runoff (e.g., 

nutrients, pesticides, sediment). The 

Maumee River system, for example, has 

a drainage area that contains 

approximately 89 percent agricultural 

land (Sanders 2002, p. 10.1). The 

decline of rayed bean and snuffbox in 

this system may be largely attributed to 

stream habitat impacts resulting from 

intensive farming and associated runoff. 

The rayed bean and snuffbox once 

occurred in the Maumee River 

mainstem, as well as in up to nine of its 

tributaries. Currently, the snuffbox is 

extirpated from the Maumee River 

system and the rayed bean is only found 

in distinct but small reaches of the St. 

Joseph River, Fish Creek, Swan Creek, 

and Blanchard River. All of these 

remaining populations (which comprise 

about 20 percent of all remaining rayed 

bean populations rangewide) are 

currently threatened by ongoing 

agricultural activities. This scenario is 

echoed across the remaining extant 

range of the rayed bean and snuffbox. 

Other Activities Affecting Rayed Bean 

and Snuffbox Habitat—Activities 

associated with urbanization can be 

detrimental to stream habitats (Couch 

and Hamilton 2002, p. 1) and were 

summarized by Feminella and Walsh 

(2005, pp. 585–587). Developmental 

activities may impact streams and their 

mussel fauna where adequate 

streamside buffers are not maintained 

and erosion of impacted land is allowed 

to enter streams (Brainwood et al. 2006, 

p. 511). Types of development may 

include highway construction, parking 

lots, building construction, general 

infrastructure (utilities, sewer systems), 

and recreation facilities. Factors 

impacting rayed bean and snuffbox 

populations in urban and suburban 

areas include lawn care chemicals 

(Conners and Black 2004, pp. 366–367), 

sedimentation, toxic effluents, domestic 

sewage, road salts, and general runoff. 

Impervious surfaces are detrimental to 

mussel habitat by altering various 

hydrological factors, including: 

Increased volumes of flow, annual flow 

rates, peak flows and duration, and 

temperature; decreased base flow; and 

changes in sediment loadings (Galli 

1991, p. 28; EPA 1997, p. 4; DeWalle et 

al. 2000, p. 2655; Myers-Kinzie et al. 

2002, p. 822). These factors result in 

flooding, erosion, channel widening, 

altered streambeds, channel instability, 

riparian and instream habitat loss, and 

loss of fish populations (EPA 1997, p. 

4). As little as 10 percent of a watershed 

being impervious can cause channel 

instability and a host of other stream 

habitat effects (Booth 1991, p. 98; Booth 

and Reinelt 1993, p. 549). Impervious 

surfaces may reduce sediment input 

into streams but result in channel 

instability by accelerating stormwater 

runoff, which increases bank erosion 

and bed scouring (Brim Box and Mossa 

1999, p. 103). Stream channels become 

highly unstable as they respond to 

increased flows by eroding a groove in 

the bottom of the channel (incising), 

which increases the force of the water 

against the channel (shear stress) and 

bed mobilization (Doyle et al. 2000, p. 

156). Hydrological variability influences 

the distribution of mussels in streams, 

with distinct communities associated 

with hydrologically flashy and 

hydrologically stable streams (Di Maio 

and Corkum 1995, p. 669). High shear 

stress, peak flows, and substrate 

movement limits mussel communities, 

reduces abundance (particularly for 

juveniles), and increasingly dislodges 

mussels and moves them downstream 

(Layzer and Madison 1995, p. 337; 

Myers-Kinzie et al. 2002, p. 822; 

Gangloff and Feminella 2006, p. 70). 

Recruitment is also significantly 

reduced in high discharge years 

(Howard and Cuffey 2006, p. 688). Most 

rayed bean and snuffbox streams have 

been impacted by general 

developmental activities and increased 

impervious surface levels (Butler 2007, 

p. 88; Butler 2002, p. 25). 

All rayed bean or snuffbox streams are 

crossed by bridges and roads. Effects 

from these structures were reviewed by 

Wheeler et al. (2005). Categories of 

impacts include primary effects 

(construction), secondary effects (postconstruction), and indirect effects 

(development associated with highway 

presence) (Angermeier et al. 2004, pp. 

21–24). Culverts act as barriers to fish 

passage (Wheeler et al. 2005, p. 149), 

particularly by increasing flow velocity 

(Warren and Pardew 1998, p. 637). 

Stream channels become destabilized 

when culverted or improperly bridged 

by interrupting the transport of woody 

debris, substrate, and water (Wheeler et 

al. 2005, p. 152). 

Anthropogenic activities can lower 

water tables, making rayed bean, 

snuffbox, and other mussel populations 

susceptible to depressed flow levels. 

Water withdrawals for irrigation, 

municipal, and industrial water 

supplies are an increasing concern. U.S. 

water consumption doubled from 1960 

to 2000 and is likely to increase further 

(Naiman and Turner 2000, p. 960). 

Therefore, we anticipate water 

withdrawals and potential stream 

dewatering to be a threat to rayed bean 

and snuffbox in the foreseeable future. 

We have identified a number of 

threats to the habitat of the rayed bean 

and snuffbox which have operated in 

the past, are impacting the species now, 

and will continue to impact the species 

in the foreseeable future. On the basis of 

this analysis, we find that the present 

and threatened destruction, 

modification, or curtailment of the 

species’ habitats is a threat to the rayed 

bean and snuffbox throughout all of 

their range. Based on our analysis of the 

best available information, we have no 

reason to believe that the present or 

threatened destruction, modification, or 

curtailment of rayed bean or snuffbox 

habitat will change in the foreseeable 

future. The decline of the freshwater 

mussels in the eastern United States is 

primarily the result the long-lasting 

effects of habitat alterations such as 

impoundments, channelization, 

chemical contaminants, mining, and 

sedimentation. Although efforts have 

been made to restore habitat in some 

areas, the long-term effects of large-scale 

and wide-ranging habitat modification, 

destruction, and curtailment will last far 

into the foreseeable future. 


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B. Overutilization for Commercial, 

Recreational, Scientific, or Educational 

Purposes 

The rayed bean and snuffbox are not 

commercially valuable species. Rare 

species like the rayed bean and snuffbox 

may increasingly be sought by lay and 

experienced collectors. Most stream 

reaches inhabited by these species are 

restricted, and their populations are 

generally small. Although scientific 

collecting is not thought to represent a 

significant threat, localized populations 

could become impacted and possibly 

extirpated by over-collecting, 

particularly if this activity is 

unregulated. Native Americans were 

known to harvest the rayed bean for 

food, but because of its size, utilization 

rates were very low (Bogan 1990, p. 

134). Localized declines of snuffbox 

from use as bait by fishermen has been 

noted (Cumberland River; Wilson and 

Clark 1914, p. 45), although it is 

unlikely that exploitation activities have 

eliminated any snuffbox populations. 

On the basis of this analysis, we find 

that overutilization for commercial, 

recreational, scientific, or educational 

purposes is not now a threat to the 

rayed bean or snuffbox in any portion of 

their 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 and 

Brunner 2007). However, mussel dieoffs have been documented in rayed 

bean and snuffbox 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 and 

Brunner 2007). Generally, parasites are 

not suspected of being a major limiting 

factor (Oesch 1984, p. 16), but a recent 

study provides contrary evidence. 

Reproductive output and physiological 

condition were negatively correlated 

with mite and trematode abundance, 

respectively (Gangloff and Feminella 

2004). 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 native mussel 

fauna, including rayed bean and 

snuffbox (Strayer 1999b, p.88). 

The muskrat (Ondatra zibethicus) is 

cited as the most prevalent mussel 

predator (Kunz 1898, p. 328; Hanson et 

al. 1989, p. 15). 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. The snuffbox ranked fourth 

among 12 species in a St. Croix River 

muskrat midden, being nearly four 

times more abundant than in 

quantitative surveys (Tyrrell and 

Hornbach 1998, p. 304). Numbers were 

too low to determine selectivity indices 

or statistics. 

Muskrats were not thought to be a 

threat to the rayed bean by West et al. 

(2000, pp. 255–256), due to their general 

selection of mussels larger than 1.4–1.6 

in (3.6–4.1 cm) long (Convey et al. 1989, 

p. 656; Hanson et al. 1989, p. 24). Neves 

and Odom (1989, pp. 938–939) also 

noted that muskrats did not select for 

small mussels. Nevertheless, some 

muskrat predation on the rayed bean 

has recently been documented in 

Cassadaga Creek, New York, but is 

generally considered insignificant. 

Other mammals (raccoon (Procyon 

lotor), mink (Mustela vison), river otter 

(Lutra Canadensis), striped skunk 

(Mephitis mephitis), hog (Sus scrofa), rat 

(Rattus spp.)), amphibians (hellbender 

(Cryptobranchus alleganiensis)), turtles, 

aquatic birds, and fishes (freshwater 

drum (Aplodinotus grunniens), redear 

sunfish (Lepomis microlophus)) feed on 

mussels (Kunz 1898, p. 328; Meek and 

Clark 1912, p. 6; Neck 1986, p. 64; 

Tyrrell and Hornbach 1998, p. 301). 

Hydra, non-biting midge larvae, 

dragonfly larvae, crayfish, and 

especially flatworms are invertebrate 

predators on newly metamorphosed 

juveniles (Zimmerman and Neves 2003, 

p. 28; Klocker and Strayer 2004, p. 174). 

The overall threat posed by these 

predators on the rayed bean and 

snuffbox is not considered significant. 

Studies indicate that in some 

localized areas, disease and predation 

may have a negative impact on mussel 

populations. However, based on our 

analysis of the best available 

information, we do not believe that 

disease or predation is a significant 

threat to the overall status of rayed bean 

or snuffbox, nor do we believe that it is 

likely to become a significant threat in 

the foreseeable future. 

D. The Inadequacy of Existing 

Regulatory Mechanisms 

Most States with extant rayed bean 

and snuffbox populations prohibit 

collection of mussels without a State 

collecting permit. However, 

enforcement of this permit requirement 

is difficult. 

Sources of nonpoint source pollution 

include timber clearcutting, clearing of 

riparian vegetation, urbanization, road 

construction, and other practices that 

allow bare earth to enter streams (The 

Nature Conservancy 2004, p. 13). 

Current laws do not adequately protect 

rayed bean and snuffbox habitat from 

nonpoint source pollution, as the laws 

to prevent sediment entering waterways 

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 rayed bean or 

snuffbox during Federal activities, or to 

ensure that Federal projects will not 

jeopardize their continued existence. 

Point source discharges within the 

range of the rayed bean and snuffbox 

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 

rayed bean and snuffbox 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 U.S. 

Army Corps of Engineers), from being 

fully used or effective. 

Despite these existing regulatory 

mechanisms, the rayed bean and 

snuffbox 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 

rayed bean and snuffbox and, therefore, 

that the inadequacy of existing 

regulatory mechanisms is a threat to 

these species throughout all of their 

range. 

Based on our analysis of the best 

available information, we have no 

reason to believe that the 

aforementioned regulations, which 

currently do not offer adequate 

protection to the rayed bean and 

snuffbox, will be improved in the 

foreseeable future. 


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E. Other Natural or Manmade Factors 

Affecting Its Continued Existence 

Other factors have played a role in the 

decline of rayed bean and snuffbox 

populations. Reduced numbers of host 

fish have an indirect impact by 

contributing to reduced recruitment 

(Watters 1996, p. 83; Khym and Layzer 

2000, p. 183). Factors associated with 

climate change likely to affect regional 

mussel populations include changes in 

stream temperature regimes and 

precipitation levels that may indirectly 

result in reduced habitat and declines in 

host fish stocks (Hastie et al. 2003, p. 

44). Remedial (such as flood control 

structures) and preventative (for 

example, more renewable energy from 

hydroelectric facilities to reduce 

greenhouse gas emissions) measures to 

address climate change issues (Hastie et 

al. 2003, p. 45) may impact rayed bean 

and snuffbox populations in the future. 

Population Fragmentation and 

Isolation—The majority of the 

remaining populations of the rayed bean 

and snuffbox are generally small and 

geographically isolated. 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 

1993–94, p. 257). Furthermore, this 

level of isolation makes natural 

repopulation of any extirpated 

population unlikely without human 

intervention. Population isolation 

prohibits the natural interchange of 

genetic material between populations, 

and small population size reduces the 

reservoir of genetic diversity within 

populations, which can lead to 

inbreeding depression (Avise and 

Hambrick 1996, p. 461). 

The Scioto River system provides a 

good example of the impacts of 

population fragmentation and isolation. 

Historically, the rayed bean and 

snuffbox were widespread and locally 

abundant in the mainstem and 

numerous tributaries. The Scioto River 

became highly contaminated over a 

century ago (Trautman 1981, p. 33; 

Yoder et al. 2005, p. 410), and these 

species eventually died out in the 

mainstem and most tributaries. The 

population segments that persist have 

become increasingly isolated due to 

impoundments and other factors; all are 

very small, highly fragmented, and 

appear to be on a trend towards 

extirpation. 

Many rayed bean and snuffbox 

populations are potentially below the 

effective population size (EPS) required 

to maintain genetic heterogeneity and 

population viability (Soule ́ 1980, p. 

162). Isolated populations eventually 

die out when population size drops 

below the EPS or threshold level of 

sustainability. Recruitment reduction or 

failure is a potential problem for many 

small rayed bean and snuffbox 

populations rangewide, a condition 

likely exacerbated by their reduced 

range and increasingly isolated 

populations. Evidence of recruitment 

has not been documented in many 

populations, indicating that recruitment 

reduction or outright failure is possible. 

Many populations of both species may 

be experiencing the bottleneck effect of 

not attaining EPS. Small, isolated, below 

EPS-threshold populations of shortlived species (most host fishes) 

theoretically die out within a decade or 

so, while below-threshold populations 

of long-lived species (like the rayed 

bean and snuffbox) might take decades 

to die out even given years of total 

recruitment failure. 

We find that fragmentation and 

isolation of small remaining populations 

of the rayed bean and snuffbox are 

current and ongoing threats to both 

species throughout all of their range that 

will continue into the foreseeable 

future. 

Exotic Species—Various exotic or 

nonnative species of aquatic organisms 

are firmly established in the range of the 

rayed bean and snuffbox. The exotic 

species that poses the most significant 

threat to the rayed bean and snuffbox is 

the zebra mussel (Dreissena 

polymorpha). The invasion of the zebra 

mussel poses a threat to the mussel 

fauna 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. 616). 

Strayer (1999b, pp. 77–80) reviewed in 

detail the mechanisms by 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. 

Another way zebra mussels may 

impact native mussels 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). Additionally, an 

indirect impact is the proliferation of 

aquatic plants from increased water 

clarity in lakes, which in turn has 

prompted managers to increase the use 

of herbicides that may threaten mussels 

via food reduction (Marangelo 2005b, 

pers. comm.). 

Zebra mussels are thoroughly 

established in the Great Lakes drainages 

and much of the Ohio River system, 

overlapping much of the current range 

of the rayed bean and snuffbox. Zebra 

mussels have eliminated populations of 

the rayed bean in Lakes Erie and 

Tippecanoe and the Detroit River. The 

greatest current potential for zebra 

mussels to impact the rayed bean and 

snuffbox are in the Lake St. Clair 

drainages, Allegheny River, Tippecanoe 

River, French Creek, and Lake 

Maxinkuckee. In addition, there is longterm potential for zebra mussel 

invasions into other systems that 

currently harbor rayed bean and 

snuffbox populations. However, zebra 

mussels are not always a serious threat 

to rayed bean and snuffbox (Tippecanoe 

River, Fisher 2005, pers. comm.; Clinton 

River, Butler 2007, p. 94; French Creek, 

Butler 2007, p. 94). Significant but 

highly fluctuating zebra mussel 

populations remain largely restricted to 

navigational waterways, although 

smaller streams have also had their 

mussel fauna virtually eliminated by 

them (Martel et al. 2001, p. 2188). At 

least two of the stronghold snuffbox 

populations (Wolf River and French 

Creek) presently have low numbers of 

zebra mussels. 

The Asian clam (Corbicula fluminea) 

has spread throughout the range of the 

rayed bean and snuffbox since its 

introduction in the mid-1900s. Asian 

clams compete with native mussels, 

especially juveniles, for food, nutrients, 

and space (Neves and Widlak 1987, p. 

6; Leff et al. 1990, p. 415) and may 

ingest sperm, glochidia, and newly 

metamorphosed juveniles of native 

mussels (Strayer 1999b, p. 82; Yeager et 

al. 2001, p. 257). Dense Asian clam 

populations actively disturb sediments 

that may reduce habitat for juvenile 

mussels (Strayer 1999b, p. 82). 

Asian clam densities vary widely in 

the absence of native mussels or in 

patches with sparse mussel 

concentrations, but 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 


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67578 Federal Register / Vol. 75, No. 211 / Tuesday, November 2, 2010 / Proposed Rules 

(Vaughn and Spooner 2006, p. 335). The 

invading clam therefore appears to 

preferentially invade sites where 

mussels are already in decline (Strayer 

1999b, p. 82; Vaughn and Spooner 2006, 

p. 332) 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 can exacerbate unionid 

imperilment through competition and 

impeding mussel population expansion 

(Vaughn and Spooner 2006, p. 335). 

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 1999b, pp. 87–88). This 

species is an aggressive competitor of 

similar sized benthic fish (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 (Strayer 1999b, p. 88; Janssen and 

Jude 2001, p. 325). Round gobies may 

therefore have indirect effects on the 

rayed bean and snuffbox through 

negative impacts to their host fishes. 

Additional exotic species will 

invariably become established in the 

foreseeable future (Strayer 1999b, pp. 

88–89). These include Limnoperna 

fortunei, a biofouling mussel (an animal 

that undesirably accumulates on wetted 

surfaces) from southeast Asia that has 

already spread to Japan and South 

America, and ‘‘probably will have strong 

effects’’ on native mussels (Strayer 

1999b, p. 89). 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 

1999b, p. 88). 

Exotic species, such as those 

described above, are an ongoing threat 

to the rayed bean and snuffbox—a threat 

that is likely to increase as these exotic 

species expand their occupancy within 

the range of the rayed bean and 

snuffbox. 

Summary of Threats 

The decline of the rayed bean and 

snuffbox (described by Butler 2002, 

2007) is primarily the result of habitat 

loss and degradation (Neves 1991, p. 

252). These losses have been well 

documented since the mid-19th century 

(Higgins 1858, p. 551). Chief among the 

causes of decline are impoundments, 

channelization, chemical contaminants, 

mining, and sedimentation (Neves 1991, 

pp. 260–261; 1993, p. 4–5; Williams et 

al. 1993, p. 7; Neves et al. 1997, pp. 60– 

72; Watters 2000, p. 269). These 

stressors have had profound impacts on 

rayed bean and snuffbox populations 

and their habitat. 

The majority of the remaining 

populations of the rayed bean and 

snuffbox are generally small and 

geographically isolated (Butler 2002, 

2007). The patchy distributional pattern 

of populations in short river reaches 

makes those populations much more 

susceptible to extirpation from single 

catastrophic events, such as toxic 

chemical spills (Watters and Dunn 

1993–94, p. 257). Furthermore, this 

level of isolation makes natural 

repopulation of any extirpated 

population virtually impossible without 

human intervention. Various nonnative 

species of aquatic organisms are firmly 

established in the range of the rayed 

bean and snuffbox; however, the exotic 

species that poses the most significant 

threat to the rayed bean and snuffbox is 

the zebra mussel (Dreissena 

polymorpha) (Butler 2002, p. 27; 2007, 

p. 93). 

Proposed Determination 

Section 3 of 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 an endangered 

species within the foreseeable future 

throughout all or a significant portion of 

its range.’’ We find that the rayed bean 

and snuffbox are presently in danger of 

extinction throughout their entire range, 

based on the immediacy, severity, and 

scope of the threats described 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. Therefore, on the basis of the 

best available scientific and commercial 

information, we propose listing the 

rayed bean and snuffbox 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 rayed bean and 

snuffbox occur throughout their range. 

Therefore, we assessed the status of the 

species throughout their entire range. 

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 the 

species throughout their entire range. 

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 results in 

public awareness and conservation by 

Federal, State, Tribal, and local 

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 by Federal agencies and the 

prohibitions against certain activities 

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. 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, selfsustaining, 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 


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final recovery plan will be available on 

our Web site (http://www.fws.gov/

endangered), or from our Ohio 

Ecological Services Field Office (see FOR

FURTHER INFORMATION CONTACT). 

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. 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. 

If this species is listed, funding for 

recovery actions will be available from 

a variety of sources, including Federal 

budgets, State programs, and cost share 

grants for non-Federal landowners, the 

academic community, and 

nongovernmental organizations. 

Additionally, under section 6 of the Act, 

we would be able to grant funds to the 

States of Illinois, Indiana, Kentucky, 

Michigan, New York, Ohio, 

Pennsylvania, Tennessee, Virginia, and 

West Virginia for management actions 

promoting the conservation of the rayed 

bean and to the States of Alabama, 

Arkansas, Illinois, Indiana, Iowa, 

Kansas, Kentucky, Michigan, 

Minnesota, Mississippi, Missouri, New 

York, Ohio, Pennsylvania, Tennessee, 

Virginia, West Virginia, and Wisconsin 

for the conservation of the snuffbox. 

Information on our grant programs that 

are available to aid species recovery can 

be found at: http://www.fws.gov/grants. 

Although the rayed bean and snuffbox 

are only proposed for listing under the 

Act at this time, please let us know if 

you are interested in participating in 

recovery efforts for these species. 

Additionally, we invite you to submit 

any new information on these species 

whenever it becomes available and any 

information you may have for recovery 

planning purposes; if you submit 

information after the date listed in the 

DATES section above, you will need to 

send it to the street address provided in 

the FOR FURTHER INFORMATION CONTACT

section. 

Section 7(a) of the Act 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 being 

designated. Regulations implementing 

this interagency cooperation provision 

of the Act are codified at 50 CFR part 

402. Section 7(a)(4) requires Federal 

agencies to confer informally with us on 

any action that is likely to jeopardize 

the continued existence of a proposed 

species or result in destruction or 

adverse modification of proposed 

critical habitat. If a species is listed 

subsequently, section 7(a)(2) of the Act 

requires Federal agencies to ensure that 

activities they authorize, fund, or carry 

out are not likely to jeopardize the 

continued existence of such a species or 

to destroy or adversely modify its 

critical habitat. If a Federal action may 

affect a listed species or its critical 

habitat, the responsible Federal agency 

must enter into formal consultation with 

us. 

Federal agency actions that may 

require conference or consultation as 

described in the preceding paragraph 

include the issuance of permits for 

reservoir construction, stream 

alterations, wastewater facility 

development, water withdrawal 

projects, pesticide registration, 

agricultural assistance programs, 

mining, road and bridge construction, 

and Federal loan programs. Activities 

will trigger consultation under section 7 

of the Act if they may affect the rayed 

bean or snuffbox, or both species, 

addressed in this proposed rule. 

Jeopardy Standard 

Prior to and following listing and 

designation of critical habitat, if prudent 

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 populations(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. 

Section 9 Take 

The Act and implementing 

regulations set forth a series of general 

prohibitions and exceptions that apply 

to all endangered and threatened 

wildlife. If we finalize listing of the 

rayed bean and snuffbox, these 

prohibitions would be applicable to the 

rayed bean and snuffbox. The 

prohibitions of section 9(a)(2) of the Act, 

codified at 50 CFR 17.21 for endangered 

wildlife, 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, capture, or collect, or to attempt 

any of these), import or export, deliver, 

receive, carry, transport, or ship in 

interstate or foreign 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 possess, sell, deliver, carry, 

transport, or ship any such wildlife that 

has been taken illegally. Further, it is 

illegal for any person to attempt to 

commit, to solicit another person to 

commit, or to cause to be committed, 

any of these acts. Certain exceptions 

apply to our agents and State 

conservation agencies. 

We may issue permits to carry out 

otherwise prohibited activities 

involving endangered wildlife under 

certain circumstances. We codified the 

regulations governing permits for 

endangered species at 50 CFR 17.22. 

Such permits are available for scientific 

purposes, to enhance the propagation or 

survival of the species, or for incidental 

take in the course of otherwise lawful 

activities. 

It is our policy, published in the 

Federal Register on July 1, 1994 (59 FR 

34272), to identify, to the maximum 

extent practicable at the time a species 

is listed, those activities that would or 

would not constitute a violation of 

section 9 of the Act and associated 

regulations at 50 CFR 17.21. The intent 

of this policy is to increase public 

awareness of the effect of this proposed 

listing on proposed and ongoing 

activities within a species’ range. We 

believe, based on the best available 

information, that the following actions 

will not result in a violation of the 

provisions of section 9 of the Act, 

provided these actions are carried out in 

accordance with existing regulations 

and permit requirements: 

(1) Activities authorized, funded, or 

carried out by Federal agencies (e.g., 

bridge and highway construction, 

pipeline construction, hydropower 

licensing, etc.), when such activities are 

conducted in accordance with the 

consultation and planning requirements 

for listed species under section 7 of the 

Act. 

(2) Any action carried out for 

scientific research or to enhance the 

propagation or survival of the rayed 

bean or snuffbox that is conducted in 


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accordance with the conditions of a 50 

CFR 17.22 permit. 

(3) Any incidental take of rayed bean 

or snuffbox resulting from an otherwise 

lawful activity conducted in accordance 

with the conditions of an incidental take 

permit issued under 50 CFR 17.22. NonFederal applicants may design a habitat 

conservation plan (HCP) for the species 

and apply for an incidental take permit. 

HCPs may be developed for listed 

species and are designed to minimize 

and mitigate impacts to the species to 

the greatest extent practicable. 

We believe the following activities 

would be likely to result in a violation 

of section 9 of the Act; however, 

possible violations are not limited to 

these actions alone: 

(1) Unauthorized killing, collecting, 

handling, or harassing of individual 

rayed bean or snuffbox, or both species, 

at any life stage. 

(2) Sale or offer for sale of rayed bean 

or snuffbox in addition to delivering, 

receiving, carrying, transporting, or 

shipping in interstate or foreign 

commerce any rayed bean or snuffbox. 

(3) Unauthorized destruction or 

alteration of the species’ habitat 

(instream dredging, channelization, 

impoundment, streambank clearing, 

discharge of fill material) that actually 

kills or injures individual rayed bean or 

snuffbox by significantly impairing their 

essential behavioral patterns, including 

breeding, feeding, or sheltering. 

(4) Violation of any discharge or water 

withdrawal permit within these species’ 

occupied ranges that results in the death 

or injury of individual rayed bean or 

snuffbox by significantly impairing their 

essential behavioral patterns, including 

breeding, feeding, or sheltering. 

(5) Discharge or dumping of toxic 

chemicals or other pollutants into 

waters supporting the species that 

actually kills or injures individual rayed 

bean or snuffbox by significantly 

impairing their essential behavioral 

patterns, including breeding, feeding, or 

sheltering. 

We will review other activities not 

identified above on a case-by-case basis 

to determine whether they may be 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 

of the Act to the Field Supervisor of the 

Service’s Ohio Ecological Services Field 

Office (see FOR FURTHER INFORMATION

CONTACT section). Requests for copies of 

regulations regarding listed species and 

inquiries about prohibitions and permits 

should be addressed to the U.S. Fish 

and Wildlife Service, Ecological 

Services 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 meaning 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, Federal action agency’s and the 

applicant’s obligation is not to restore or 

recover the species, but to implement 

reasonable and prudent alternatives to 

avoid destruction or adverse 

modification of critical habitat. 

For inclusion in a critical habitat 

designation, the habitat within the 

geographical area occupied by the 

species at the time it was listed must 

contain the physical and biological 

features essential to the conservation of 

the species, and be included only if 

those features may require special 

management considerations or 

protection. Critical habitat designations 

identify, to the extent known using the 

best scientific and commercial data 

available, habitat areas that provide 

essential life cycle needs of the species 

(areas on which are found the physical 

and biological features (PBFs) laid out 

in the appropriate quantity and spatial 

arrangement for the conservation of the 

species). Under the Act and regulations 

at 50 CFR 424.12, we can designate 

critical habitat in areas outside the 

geographical area occupied by the 

species at the time it is listed only when 

we determine that those areas are 

essential for the conservation of the 

species and that designation limited to 

those areas occupied at the time of 

listing would be inadequate to ensure 

the conservation of the species. 

Section 4 of the Act requires that we 

designate critical habitat on the basis of 

the best scientific and commercial data 

available. Further, our Policy on 

Information Standards Under the 

Endangered Species Act (published in 

the Federal Register on July 1, 1994 (59 

FR 34271)), the Information Quality Act 

(section 515 of the Treasury and General 

Government Appropriations Act for 

Fiscal Year 2001 (Pub. L. 106–554; H.R. 

5658)), and our associated Information 

Quality Guidelines, provide criteria, 

establish procedures, and provide 

guidance to ensure that our decisions 

are based on the best scientific data 

available. They require our biologists, to 

the extent consistent with the Act and 

with the use of the best scientific data 

available, to use primary and original 

sources of information as the basis for 

recommendations to designate critical 

habitat. 

When we are determining which areas 

should be designated as critical habitat, 

our primary source of information is 

generally the information developed 

during the listing process for the 

species. Additional information sources 

may include the recovery plan for the 

species, articles in peer-reviewed 

journals, conservation plans developed 

by States and counties, scientific status 

surveys and studies, biological 

assessments, or other unpublished 

materials and expert opinion or 

personal knowledge. 

Habitat is often dynamic, and species 

may move from one area to another over 

time. Furthermore, we recognize that 

critical habitat designated at a particular 

point in time may not include all of the 

habitat areas that we may later 

determine are necessary for the recovery 

of the species. For these reasons, a 


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critical habitat designation does not 

signal that habitat outside the 

designated area is unimportant or may 

not be required for recovery of the 

species. 

Areas that are important to the 

conservation of the species, but are 

outside the critical habitat designation, 

will continue to be subject to 

conservation actions we implement 

under section 7(a)(1) of the Act. Areas 

that support populations are also subject 

to the regulatory protections afforded by 

the section 7(a)(2) jeopardy standard, as 

determined on the basis of the best 

available scientific information at the 

time of the agency action. Federally 

funded or permitted projects affecting 

listed species outside their designated 

critical habitat areas may still result in 

jeopardy findings in some cases. 

Similarly, critical habitat designations 

made on the basis of the best available 

information at the time of designation 

will not control the direction and 

substance of future recovery plans, 

habitat conservation plans (HCPs), or 

other species conservation planning 

efforts if new information available at 

the time of these planning efforts calls 

for a different outcome. 

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 

habitat at the time we determine that a 

species is 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 the rayed bean or 

snuffbox, 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 rayed bean and 

snuffbox 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 the rayed bean and snuffbox, 

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 

designation of critical habitat is prudent 

for the rayed bean and snuffbox. 

Critical Habitat Determinability 

As stated above, section 4(a)(3) of the 

Act requires the designation of critical 

habitat concurrently with the species’ 

listing ‘‘to the maximum extent prudent 

and determinable.’’ Our regulations at 50 

CFR 424.12(a)(2) state that critical 

habitat is not determinable when one or 

both of the following situations exist: 

(i) Information sufficient to perform 

required analyses of the impacts of the 

designation is lacking, or 

(ii) The biological needs of the species 

are not sufficiently well known to 

permit identification of an area as 

critical habitat. 

When critical habitat is not 

determinable, the Act provides for an 

additional year to publish a critical 

habitat designation (16 U.S.C. 

1533(b)(6)(C)(ii)). 

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 essential to the 

conservation of the species. These 

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 physical and biological features 

essential for the conservation of the 

rayed bean and snuffbox because 

information on those features for 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 the rayed bean and 

snuffbox have not been observed for 

decades in many of their historical 

locations, and much of the habitat in 

which they still persist 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 the 

rayed bean and snuffbox. 

Key features of the basic life history, 

ecology, reproductive biology, and 

habitat requirements of most mussels, 

including the rayed bean and snuffbox, 

are unknown. Species-specific 

ecological requirements have not been 

determined (for example, minimum 

water flow and effects of particular 

pollutants). Population dynamics, such 

as species’ interactions and community 

structure, population trends, and 

population size and age class structure 

necessary to maintain long-term 

viability, have not been determined for 

these species. Of particular concern to 

both species is that many of the 

remaining rayed bean and snuffbox 

populations consist of very low 

densities, which limit our ability to 

investigate their population dynamics. 

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 longterm viability. As we are unable to 


srobinson on DSKHWCL6B1PROD with PROPOSALS3
67582 Federal Register / Vol. 75, No. 211 / Tuesday, November 2, 2010 / Proposed Rules 

identify many physical and biological 

features essential to the conservation of 

the rayed bean and snuffbox, we are 

unable to identify areas that contain 

these features. Therefore, although we 

have determined that the designation of 

critical habitat is prudent for the rayed 

bean and snuffbox, because the 

biological and physical requirements of 

these species are not sufficiently known, 

we find that critical habitat for the rayed 

bean and snuffbox 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 

list rayed bean and snuffbox as 

endangered and our proposal regarding 

critical habitat for this species. 

We will consider all comments and 

information we receive during the 

comment period on this proposed rule 

during preparation of a final 

rulemaking. Accordingly, our final 

decision may differ from this proposal. 

Public Hearings 

The Act provides for one or more 

public hearings on this proposal, if 

requested. Requests must be received 

within 45 days after the date of 

publication of this proposal in the 

Federal Register (see DATES). Such 

requests must be sent to the address 

shown 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 

accommodation, in the Federal Register 

and local newspapers at least 15 days 

before the hearing. 

Persons needing reasonable 

accommodation to attend and 

participate in a public hearing should 

contact the Ohio Ecological Services 

Field Office at 614–416–8993, ext. 22, 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. 

Required Determinations 

Clarity of 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. 

Paperwork Reduction Act (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. This rule 

will 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 (42 

U.S.C. 4321 et seq.) 

We have determined that we do not 

need to prepare an environmental 

assessment, as defined under the 

authority of the National Environmental 

Policy Act of 1969, 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 proposed rule is available on the 

Internet at http://www.regulations.gov 

or upon request from the Field 

Supervisor, Ohio Ecological Services 

Field Office (see FOR FURTHER

INFORMATION CONTACT section). 

Author 

The primary author of this proposed 

rule is Angela Boyer of the Ohio 

Ecological Services Field Office (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 Federal 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. Amend § 17.11(h) by adding new 

entries for ‘‘Mussel, rayed bean’’ and 

‘‘Mussel, snuffbox’’ in alphabetical order 

under CLAMS to the List of Endangered 

and Threatened Wildlife as follows: 

§ 17.11 Endangered and threatened 

wildlife. 

* * * * * 

Species 

Historic range 

Vertebrate 

population 

where endangered or 

threatened 

Status When listed Critical 

habitat 

Special 

rules Common name Scientific name 

* * * * * * * 

.


srobinson on DSKHWCL6B1PROD with PROPOSALS3
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Species 

Historic range 

Vertebrate 

population 

where endangered or 

threatened 

Status When listed Critical 

habitat 

Special 

rules Common name Scientific name 

CLAMS

* * * * * * * 

Mussel, rayed bean ..... Villosa fabalis .............. U.S.A. (IL, IN, KY, MI, 

NY, OH, PA, TN, 

VA, WV, WI).

NA E .................... NA NA 

* * * * * * * 

Mussel, snuffbox .......... Epioblasma triquetra ... U.S.A. (AL, AR, IL, IN, 

IA, KS, KY, MI, MN, 

MS, MO, NY, OH, 

PA, TN, VA, WV, 

WI).

NA E .................... NA NA 

* * * * * * * 

* * * * * Dated: October 15, 2010. 

Gary D. Frazer, 

Acting Director, U.S. Fish and Wildlife 

Service. 

[FR Doc. 2010–27413 Filed 11–1–10; 8:45 am] 

BILLING CODE 4310–55–P 


srobinson on DSKHWCL6B1PROD with PROPOSALS3