Hemistena lata - (Rafinesque, 1820)
Cracking Pearlymussel
Other English Common Names: Cracking Pearly Mussel
Synonym(s): Lastena lata (Rafinesque, 1820)
Taxonomic Status: Accepted
Related ITIS Name(s): Hemistena lata (Rafinesque, 1820) (TSN 80350)
Unique Identifier: ELEMENT_GLOBAL.2.109635
Element Code: IMBIV20010
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
Image 11992

Public Domain

 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Hemistena
Genus Size: A - Monotypic genus
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Concept Reference
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Concept Reference: Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.
Concept Reference Code: B98TUR01EHUS
Name Used in Concept Reference: Hemistena lata
Taxonomic Comments: This species is in a monotypic genus closely related to the genus Elliptio.
Conservation Status
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NatureServe Status

Global Status: G1
Global Status Last Reviewed: 29Apr2009
Global Status Last Changed: 25Nov1996
Rounded Global Status: G1 - Critically Imperiled
Reasons: This species has been severely reduced through habitat degradation in range to only three widely disjunct populations on two rivers, one of which may no longer be viable; and all of which are represented by a small number of individuals.
Nation: United States
National Status: N1 (25Nov1996)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S1), Illinois (SX), Indiana (SX), Kentucky (SX), Ohio (SX), Pennsylvania (SX), Tennessee (S1), Virginia (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE, XN: Listed endangered, nonessential experimental population (28Sep1989)
Comments on USESA: Listed Endangered throughout its range, except in the free-flowing reach of the Tennessee River from the base of Wilson Dam downstream to the backwaters of Pickwick Reservoir and the lower 5 RM of all tributaries to this reach in Colbert and Lauderdale Counties, Alabama. Here is it listed as an experimental, non-essential population. (Federal Register, 14 June 2001).

The USFWS, in cooperation with the State of Tennessee and Conservation Fisheries, Inc., announced a final rule to reintroduce this species into its historical habitat in the free-flowing reach of the French Broad River below Douglas Dam to its confluence with the Holston River, Knox County Tennessee, and in the free-flowing reach of the Holston River below Cherokee Dam to its confluence with the French Broad River (Federal Register, 13 September 2007). The proposed rule for this action was published on June 13, 2006.

U.S. Fish & Wildlife Service Lead Region: R4 - Southeast
IUCN Red List Category: CR - Critically endangered
American Fisheries Society Status: Endangered (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: 100-250 square km (about 40-100 square miles)
Range Extent Comments: This species was once widely distributed in the Ohio, Cumberland, and Tennessee river systems. It ranged in the Ohio River from Ohio downstream to Illinois. In Indiana and Illinois, it was historically known from the White, Wabash (Fisher, 2006), and Tippecanoe Rivers (Cummings and Berlocher, 1990). Kentucky records show it once inhabited the Upper Cumberland, Big South Fork, Green, and Kentucky Rivers. It was historically collected in Tennessee from the Tennessee, Cumberland, Powell, Clinch, Holston, Elk, Duck, and Buffalo Rivers. In Alabama it existed in the Tennessee River presumably across the northern part of the state but all museum records are from Muscle Shoals but is reduced to one population on the Elk River (Williams et al., 2008). Portions of the Powell, Clinch, and Holston Rivers in Virginia also once suppported the species (USFWS, 1990). It has been extirpated from most of its former range but some viable populations may persist in the upper Clinch River in Hancock Co., Tennessee (Parmalee and Bogan, 1998), and Scott Co., Virginia; as well as the Elk River in Lincoln Co., Tennessee (USFWS, 1990). A population in the Powell River in Hancock Co., Tennessee, that was barely surviving in 1979 (3 of 78 sites over 98 river miles) likely is no longer extant or if so, viable (USFWS, 1990).

Area of Occupancy: 26-2,500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 1 - 5
Number of Occurrences Comments: This species has been reduced to possibly 3 reproducing populations in the Tennessee River system in Virginia (Scott Co.) and Tennessee (Hancock Co.) in the upper Clinch River and Elk River (USFWS, 1990). In Tennessee, viable populations survive in the upper Clinch River (Hancock Co.) in east Tennessee (Parmalee and Bogan, 1998). It is also extant in Elk River, Alabama and possibly at the state line in Tennessee, where densities are very low and viability is questionable (Mirarchi et al., 2004; Williams et al., 2008; Hubbs, 2002).

Population Size: 50 - 1000 individuals
Population Size Comments: Apparently this species is declining at 3 remaining sites with population numbers very low likely below 1000 individuals (USFWS, 1990; 2006).

Number of Occurrences with Good Viability/Integrity: Very few (1-3)
Viability/Integrity Comments: Barr et al. (1994) determined (based on 1981 survey data) that viable populations exist in The Elk and Clinch Rivers. The Clinch River population is the largest and covers the largest river length (a few sites but over 140 or so miles) (USFWS, 1990). The population in the Elk River is not high enough to maintain long-term genetic viability (USFWS, 1990). A population in the Powell River in Hancock Co., Tennessee, that was barely surviving in 1979 (3 of 78 sites over 98 river miles) likely is no longer extant or if so, viable. The Green River population has been extirpated since 1966 (USFWS, 1990).

Overall Threat Impact: High
Overall Threat Impact Comments: Impoundments, siltation and pollution leading to water quality and habitat deterioration. Inadequate sewage treatment, coal mining, oil and gas drilling and poor land-use practices. The Powell River watershed was mined extensively for coal, and coal mining impacts are still present, especially in the upper reaches. The lower reaches of the Powell River have large deposits of coal fines and silt. The Clinch River has been adversely affected by pollution and land use practices along the river have contributed to the decline of water quality and loss of mussel populations. It has also experienced some impacts from coal mining. Toxic spills have historically produced mussel kills in the river. Although suitable habitat exists in the Elk River, cold water releases from Tims Ford Reservoir and pollution from unknown sources in the lower Elk River have impacted mussel fauna and reduced density (USFWS, 1990).

Short-term Trend: Decline of >70%
Short-term Trend Comments: This species has declined substantially in Illinois to the point of extinction and was formerly found in the Wabash River (Cummings and Mayer, 1997). In Tennessee, it has been extirpated from most of its former range, although local and apparently viable populations survive, although the population in the Elk River persisted only until about 1981 (Parmalee and Bogan, 1998). The Elk River population has declined to near absence, the Green River population has not yielded specimens since 1966 despite surveys in the late 1980s, live specimens were taken in the Tennessee River below Pickwick Dam in the 1970s but only relict specimens after that, the Powell River population is also likely extirpated (USFWS, 1990).

Long-term Trend: Decline of >90%
Long-term Trend Comments: This species was once widely distributed in the Ohio, Cumberland, and Tennessee river systems. It was reported from the lower and upper Elk River, Tennessee (Isom et al., 1973). It is now reduced to about 3 (possibly 2) viable populations and some nonessential experimental populations reintroduced (USFWS, 1990; 2006). This species is extirpated from Pennsylvania (Bogan, 1993; Spoo, 2008) where it formerly occurred in the Ohio drainage. It is extirpated from Indiana where it was reported in the Wabash River in the last century (Fisher, 2006), and the Tippecanoe River early in the century (Cummings and Berlocher, 1990). It is extirpated in Ohio where it was formerly known from the Tuscarawas River at New Philadelphia an dCoshocton, the Scioto River at Circleville and Columbus, Ohio River at Cincinnati and Marietta, and Buffington Island on the Ohio River adjacent to Meigs County; having last been seen in 1911 (Watters et al., 2009).

Intrinsic Vulnerability: Highly vulnerable
Intrinsic Vulnerability Comments: All populations are geographically isolated from one another restricting natural interchange of genetic material and all have extremely small population size with questionable viability at best (the Clinch River population being an exception) (USFWS, 1991).

Other NatureServe Conservation Status Information

Inventory Needs: Define extent, numbers and viability in existing populations.

Protection Needs: Continue Clinch Valley Bioreserve initiatives to improve water quality and land-use practices, through variety of governmental and private partners.

Distribution
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Global Range: (100-250 square km (about 40-100 square miles)) This species was once widely distributed in the Ohio, Cumberland, and Tennessee river systems. It ranged in the Ohio River from Ohio downstream to Illinois. In Indiana and Illinois, it was historically known from the White, Wabash (Fisher, 2006), and Tippecanoe Rivers (Cummings and Berlocher, 1990). Kentucky records show it once inhabited the Upper Cumberland, Big South Fork, Green, and Kentucky Rivers. It was historically collected in Tennessee from the Tennessee, Cumberland, Powell, Clinch, Holston, Elk, Duck, and Buffalo Rivers. In Alabama it existed in the Tennessee River presumably across the northern part of the state but all museum records are from Muscle Shoals but is reduced to one population on the Elk River (Williams et al., 2008). Portions of the Powell, Clinch, and Holston Rivers in Virginia also once suppported the species (USFWS, 1990). It has been extirpated from most of its former range but some viable populations may persist in the upper Clinch River in Hancock Co., Tennessee (Parmalee and Bogan, 1998), and Scott Co., Virginia; as well as the Elk River in Lincoln Co., Tennessee (USFWS, 1990). A population in the Powell River in Hancock Co., Tennessee, that was barely surviving in 1979 (3 of 78 sites over 98 river miles) likely is no longer extant or if so, viable (USFWS, 1990).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: endemic to a single nation

U.S. & Canada State/Province Distribution
United States AL, ILextirpated, INextirpated, KYextirpated, OHextirpated, PAextirpated, TN, VA

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Lauderdale (01077)*, Limestone (01083)
IN Knox (18083)*, Morgan (18109)*
KY Campbell (21037)*, Carroll (21041)*, Cumberland (21057)*, Edmonson (21061)*, Garrard (21079)*, Hart (21099)*, Jefferson (21111)*, Jessamine (21113)*, Kenton (21117)*, McCreary (21147)*, Monroe (21171)*, Pulaski (21199)*, Russell (21207)*, Wayne (21231)*
TN Anderson (47001)*, Claiborne (47025)*, Decatur (47039)*, Giles (47055), Grainger (47057)*, Hancock (47067), Hardin (47071), Lincoln (47103), Maury (47119)*, Wayne (47181)*
VA Lee (51105), Russell (51167)*, Scott (51169)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
05 Upper Ohio-Shade (05030202)*, Tuscarawas (05040001)*, Upper Scioto (05060001)*, Lower Scioto (05060002)*, Middle Ohio-Laughery (05090203)+*, Lower Kentucky (05100205)+*, Upper Green (05110001)+*, Middle Wabash-Deer (05120105)*, Tippecanoe (05120106)*, Lower Wabash (05120113)+*, Upper White (05120201)+*, Upper Cumberland (05130101)*, Upper Cumberland-Lake Cumberland (05130103)+*, South Fork Cumberland (05130104)+*, Silver-Little Kentucky (05140101)+*
06 Holston (06010104)*, Watts Bar Lake (06010201)*, Upper Clinch (06010205)+, Powell (06010206)+, Lower Clinch (06010207)+*, Middle Tennessee-Chickamauga (06020001)*, Upper Elk (06030003)+, Lower Elk (06030004)+, Pickwick Lake (06030005)+*, Lower Tennessee-Beech (06040001)+, Upper Duck (06040002)*, Lower Duck (06040003)+*, Buffalo (06040004)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A freshwater mussel - cracking pearlymussel.
Reproduction Comments: The glochidial host is not known.
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, CREEK, MEDIUM RIVER, Riffle
Special Habitat Factors: Benthic
Habitat Comments: This species is abundant in sand, gravel, and cobble substrates in swift currents or mud and sand in slower currents (Gordon and Layzer, 1989).
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: This species was listed as federally endangered in the U.S. in 1989 and a recovery plan created (USFWS, 1990). Recovery objectives include: (1) preserve present populations and occupied habitat, (2) determine threats to the species, conduct research necessary for the species' management and recovery, and implement management where needed, (3) search for additional populations and/or habitat suitable for reintroduction efforts, (4) determine, through research, the feasibility of augmenting extant populations and reestablishing the cracking pearlymussel into historic habitat and reintroduce where feasible, (5) develop and implement cryogenic techniques to preserve the species' genetic material until such time as conditions are sutable for reintroduction, (6) develop and implement a program to monitor population levels and habitat conditions of presently established populations as well as newly discovered, introduced, or expanding populations, (7) annually assess overall success of the recovery program and recommend action (modify recovery objectives, delist, continue to protect, implement new measures, or other studies, etc.). Nonessessential Experimental Populations (NEPs) have been established in the Tennessee River below Wilson Dam, Colbert and Lauderdale Cos., Alabama (USFWS, 2001). Nonessessential Experimental Populations (NEPs) have also been proposed for reintroduction into the free-flowing reach of the French Broad River below Douglas Dam (Knox and Sevier Cos., Tennessee) to its confluence with the Holston River, Knox Co., Tennessee, and in the free-flowing reach of the Holston River below Cherokee Dam to its confluence with the French Broad River (Knox, Grainger, and Jefferson Cos., Tennessee), where this species currently does not exist (USFWS, 2006).
Biological Research Needs: Biological and ecological studies to determine life history and population traits.
Population/Occurrence Delineation
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Group Name: Freshwater Mussels

Use Class: Not applicable
Minimum Criteria for an Occurrence: Occurrences are based on some evidence of historical or current presence of single or multiple specimens, including live specimens or recently dead shells (i.e., soft tissue still attached and/or nacre still glossy and iridescent without signs of external weathering or staining), at a given location with potentially recurring existence. Weathered shells constitute a historic occurrence. Evidence is derived from reliable published observation or collection data; unpublished, though documented (i.e. government or agency reports, web sites, etc.) observation or collection data; or museum specimen information.
Mapping Guidance: Based on the separation distances outlined herein, for freshwater mussels in STANDING WATER (or backwater areas of flowing water such as oxbows and sloughs), all standing water bodies with either (1) greater than 2 km linear distance of unsuitable habitat between (i.e. lotic connections), or (2) more than 10 km of apparently unoccupied though suitable habitat (including lentic shoreline, linear distance across water bodies, and lentic water bodies with proper lotic connections), are considered separate element occurrences. Only the largest standing water bodies (with 20 km linear shoreline or greater) may have greater than one element occurrence within each. Multiple collection or observation locations in one lake, for example, would only constitute multiple occurrences in the largest lakes, and only then if there was some likelihood that unsurveyed areas between collections did not contain the element.

For freshwater mussels in FLOWING WATER conditions, occurrences are separated by a distance of more than 2 stream km of unsuitable habitat, or a distance of more than 10 stream km of apparently unoccupied though suitable habitat. Standing water between occurrences is considered suitable habitat when calculating separation distance for flowing water mussel species unless dispersal barriers (see Separation Barriers) are in place.

Several mussel species in North America occur in both standing and flowing water (see Specs Notes). Calculation of separation distance and determination of separation barriers for these taxa should take into account the environment in which the element was collected. Juvenile mussels do not follow this pattern and juveniles are typically missed by most standard sampling methods (Hastie and Cosgrove, 2002; Neves and Widlak, 1987), therefore juvenile movement is not considered when calculating separation distance.

Separation Barriers: Separation barriers within standing water bodies are based solely on separation distance (see Separation Distance-suitable, below). Separation barriers between standing water bodies and within flowing water systems include lack of lotic connections, natural barriers such as upland habitat, absence of appropriate species specific fish hosts, water depth greater than 10 meters (Cvancara, 1972; Moyle and Bacon, 1969) or anthropogenic barriers to water flow such as dams or other impoundments and high waterfalls.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 10 km
Alternate Separation Procedure: None
Separation Justification: Adult freshwater mussels are largely sedentary spending their entire lives very near to the place where they first successfully settled (Coker et al., 1921; Watters, 1992). Strayer (1999) 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. Movement occurs with the impetus of some stimulus (nearby water disturbance, physical removal from the water such as during collection, exposure conditions during low water, seasonal temperature change or associated diurnal cycles) and during spawning. Movement is confined to either vertical movement burrowing deeper into sediments though rarely completely beneath the surface, or horizontal movement in a distinct path often away from the area of stimulus. Vertical movement is generally seasonal with rapid descent into the sediment in autumn and gradual reappearance at the surface during spring (Amyot and Downing, 1991; 1997). Horizontal movement is generally on the order of a few meters at most and is associated with day length and during times of spawning (Amyot and Downing, 1997). Such locomotion plays little, if any, part in the distribution of freshwater mussels as these limited movements are not dispersal mechanisms. Dispersal patterns are largely speculative but have been attributed to stream size and surface geology (Strayer, 1983; Strayer and Ralley, 1993; van der Schalie, 1938), utilization of flow refuges during flood stages (Strayer, 1999), and patterns of host fish distribution during spawning periods (Haag and Warren, 1998; Watters, 1992). Lee and DeAngelis (1997) modeled the dispersal of freshwater into unoccupied habitats as a traveling wave front with a velocity ranging from 0.87 to 2.47 km/year (depending on mussel life span) with increase in glochidial attachment rate to fish having no effect on wave velocity.

Nearly all mussels require a host or hosts during the parasitic larval portion of their life cycle. Hosts are usually fish, but a few exceptional species utilize amphibians as hosts (Van Snik Gray et al., 2002; Howard, 1915) or may metamorphose without a host (Allen, 1924; Barfield et al., 1998; Lefevre and Curtis, 1911; 1912). Haag and Warren (1998) found that densities of host generalist mussels (using a variety of hosts from many different families) and displaying host specialists (using a small number of hosts usually in the same family but mussel females have behavioral modifications to attract hosts to the gravid female) were independent of the densities of their hosts. Densities of non-displaying host specialist mussels (using a small number of hosts usually in the same family but without host-attracting behavior) were correlated positively with densities of their hosts. Upstream dispersal of host fish for non-displaying host specialist mussels could, theoretically, transport mussel larvae (glochidia) over long distances through unsuitable habitat, but it is unlikely that this occurs very often. D. Strayer (personal communication) suggested a distance of at least 10 km, but a greater distance between occurrences may be necessary to constitute genetic separation of populations. As such, separation distance is based on a set, though arbitrary, distance between two known points of occurrence.

Date: 18Oct2004
Author: Cordeiro, J.
Notes: Contact Jay Cordeiro (jay_cordeiro@natureserve.org) for a complete list of freshwater mussel taxa sorted by flow regime.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 29Apr2009
NatureServe Conservation Status Factors Author: Cordeiro, J. (2009); Lipford, M. (1991)
Element Ecology & Life History Edition Date: 01May2007
Element Ecology & Life History Author(s): Cordeiro, J.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Bogan, A.E. 1993a. Workshop on freshwater bivalves of Pennsylvania. Workshop hosted by Aquatic Systems Corporation, Pittsburgh, Pennsylvania, held at Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, 6-7 May 1993. 80 pp.

  • Fisher, B.E. 2006. Current status of freshwater mussels (Order Unionoida) in the Wabash River drainage of Indiana. Proceedings of the Indiana Academy of Science, 115(2): 103-109.

  • Gordon, M.E. and J.B. Layzer. 1989. Mussels (Bivalvia: Unionoidea) of the Cumberland River review of life histories and ecological relationships. U.S. Fish and Wildlife Service Biological Report, 89(15): 1-99.

  • Howard, A.D. 1915. Some exceptional cases of breeding among the Unionidae. The Nautilus 29:4-11.

  • Isom, B.G., P. Yokley, Jr., and C.H. Gooch. 1973. Mussels of Elk River Basin in Alabama and Tennessee- 1965-1967. American Midland Naturalist 89(2):437-442.

  • Lefevre, G. and W.T. Curtis. 1912. Studies on the reproduction and artificial propogation of fresh-water mussels. Bulletin of the Bureau of Fisheries 30:102-201.

  • Mirarchi, R.E., J.T. Garner, M.F. Mettee, and P.E. O'Neil. 2004b. Alabama wildlife. Volume 2. Imperiled aquatic mollusks and fishes. University of Alabama Press, Tuscaloosa, Alabama. xii + 255 pp.

  • Moyle, P. and J. Bacon. 1969. Distribution and abundance of molluscs in a fresh water environment. Journal of the Minnesota Academy of Science 35(2/3):82-85.

  • Parmalee, P.W. and A.E. Bogan. 1998. The freshwater mussels of Tennessee. University of Tennessee Press, Knoxville, Tennesee. 328 pp.

  • Spoo, A. 2008. The Pearly Mussels of Pennsylvania. Coachwhip Publications: Landisville, Pennsylvania. 210 pp.

  • Strayer, D. 1983. The effects of surface geology and stream size on freshwater mussel (Bivalvia, Unionidae) distribution in southeastern Michigan, U.S.A. Freshwater Biology 13:253-264.

  • Strayer, D.L. 1999a. Use of flow refuges by unionid mussels in rivers. Journal of the North American Benthological Society 18(4):468-476.

  • Strayer, D.L. and J. Ralley. 1993. Microhabitat use by an assemblage of stream-dwelling unionaceans (Bivalvia) including two rare species of Alasmidonta. Journal of the North American Benthological Society 12(3):247-258.

  • Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.

  • U.S. Fish and Wildlife Service (USFWS). 2001. Endangered and Threatened Wildlife and Plants; establishment of nonessential experimental population status for 16 freshwater mussels and 1 freshwater snail (Anthony's Riversnail) in the free-flowing reach of the Tennessee River below the Wilson Dam, Colbert and Lauderdale Counties, Alabama. Federal Register, 66(115): 32250-32264.

  • U.S. Fish and Wildlife Service (USFWS). 2006. Endangered and threatened wildlife and plants; establishment of nonessential experimental population status for 15 freshwater mussels, 1 freshwater snail, and 5 fishes in the lower French Broad River and in the lower Holston River, Tennessee; Proposed Rule. Federal Register, 71(113): 34195-34230.

  • Van der Schalie, H. 1938a. The naiad fauna of the Huron River in southeastern Michigan. Miscellaneous Publication of the Museum of Zoology, University of Michigan 40:7-78.

  • Watters, G.T. 1992a. Unionids, fishes, and the species-area curve. Journal of Biogeography 19:481-490.

  • Williams, J. D., A. E. Bogan, and J. T Garner. 2008. Freshwater mussels of Alabama & the Mobile Basin in Georgia, Mississippi, & Tennessee. University of Alabama Press, Tuscaloosa, Alabama. 908 pages.

  • Williams, J.D., M.L. Warren, Jr., K.S. Cummings, J.L. Harris, and R.J. Neves. 1993b. Conservation status of freshwater mussels of the United States and Canada. Fisheries 18(9): 6-22.

References for Watershed Distribution Map
  • Barr, W.C., S.A. Ahlstedt, G.D. Hickman, and D.M. Hill. 1993-1994. Cumberlandian mollusk conservation program. Activity 8: Analysis of macrofauna factors. Walkerana 7(17/18):159-224.

  • Biological Resources Division, USGS. 1997. Database of museum records of aquatic species. Compiled by J. Williams (USGS-BRD, Gainesville, FL).

  • Cummings, K.S. and J.M. Berlocher. 1990. The naiades or freshwater mussels (Bivalvia: Unionidae) of the Tippecanoe River, Indiana. Malacological Review 23:83-98.

  • Hubbs, D. 2002. Monitoring and management of endangered mussels. 2001-02 Annual Report Project 7365, Tennessee Wildlife Resources Agency, Nashville, Tennessee. 3 pp.

  • Parmalee, P.W. and A.E. Bogan. 1998. The Freshwater Mussels of Tennessee. University of Tennessee Press: Knoxville, Tennessee. 328 pp.

  • U.S. Fish and Wildlife Service (USFWS). 1990h. Cracking pearlymussel (Hemistena (= Lastena) lata) recovery plan. U.S. Fish and Wildlife Service, Atlanta, Georgia. 25 pp.

  • Watters, G.T., M.A. Hoggarth, and D.H. Stansbery. 2009b. The Freshwater Mussels of Ohio. Ohio State University Press: Columbus, Ohio. 421 pp.

  • Williams, J.D., A.E. Bogan, and J.T. Garner. 2008. Freshwater Mussels of Alabama & the Mobile Basin in Georgia, Mississippi & Tennessee. University of Alabama Press: Tuscaloosa, Alabama. 908 pp.

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