Toxolasma cylindrellus - (I. Lea, 1868)
Pale Lilliput
Other English Common Names: Pale Lilliput Naiad, Pale Lilliput Pearly Mussel
Taxonomic Status: Accepted
Related ITIS Name(s): Toxolasma cylindrellus (I. Lea, 1868) (TSN 80361)
Unique Identifier: ELEMENT_GLOBAL.2.116497
Element Code: IMBIV43020
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Toxolasma
Genus Size: C - Small genus (6-20 species)
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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: Toxolasma cylindrellus
Taxonomic Comments: In an unpublished study of molecular systematics, Campbell and Harris (2006) found this species to be valid and most closely related to Toxolasma lividus.
Conservation Status
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NatureServe Status

Global Status: G1
Global Status Last Reviewed: 19May2009
Global Status Last Changed: 15Sep1997
Rounded Global Status: G1 - Critically Imperiled
Reasons: This is a severly declining regional endemic known from one (possibly two) viable extant populations whose low numbers make it expecially vulnerable to extinction. Any impact to the species is significant.
Nation: United States
National Status: N1 (15Sep1997)

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), Georgia (SX), Tennessee (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (14Jun1976)
U.S. Fish & Wildlife Service Lead Region: R4 - Southeast
IUCN Red List Category: CR - Critically endangered
Convention on International Trade in Endangered Species Protection Status (CITES): Appendix I
American Fisheries Society Status: Endangered (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: <100 square km (less than about 40 square miles)
Range Extent Comments: This is a small stream species restricted to the tributaries of the Tennessee River, but not the Tennessee River proper (USFWS, 1984). One exception is a single record from Swamp Creek, Whitfield County, Georgia (recorded in the late 1800s; likely now extirpated); absent from the upper Tennessee drainage. Ortmann collected specimens he thought were Toxolasma cylindrellus from Little Pigeon River, Sevier County, Tennessee. H. D. Athearn collected specimens from the Sequatchie and Little Sequatchie Rivers in Marion County, Tennessee back in the 1950s (Bogan and Parmalee, 1983; Parmalee and Bogan, 1998). It was found in the Paint Rock River and in Larkin Fork of the Paint Rock River (Bogan and Parmalee, 1983) and in the Estill Fork of the Paint Rock River (McGregor and Shelton, 1995). It has also been found in the Flint River and Indian Creek, Madison County, Alabama and in the Elk River, Franklin County, Tennessee. It was also found in the Duck River from Hickman County upstream to Bedford County, Tennessee (Bogan and Parmalee, 1983; Parmalee and Bogan, 1998). Currently it is believed to be extant only in the Paint Rock River Drainage in Jackson County, Alabama and adjacent Franklin Co., Tennessee (Bogan and Parmalee, 1985; Williams et al., 2008). It is found in the mainstem of the Paint Rock River and each of its major tributaries: Larkin Fork, Estill Fork, and Hurricane Creek (McGregor and Shelton, 1995) (Doug Shelton, pers. obs., 1996). The largest population was found in the Estill Fork from the Tennessee state line downstream for about one mile (Shelton, 1997) but its viability is now questionable. The Duck River population is believed to have been extirpated (Steve Ahlstedt, pers. comm. May, 1995), although Parmalee and Bogan (1998) believe it may still be hanging on.

Area of Occupancy: 1-125 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 1 - 5
Number of Occurrences Comments: Formerly widespread throughout the Tennessee River drainage it is now believed to exist only in the Paint Rock River drainage in Jackson Co. Alabama and adjacent Franklin Co., Tennessee (one viable population at Hurricane Creek and a likely non-viable population in Estill Fork- Ahlstedt, 1996) and in the Duck River of Tennessee at Big Rock Creek (population may be extirpated) (Parmalee and Bogan, 1998; USFWS, 1984; Williams et al., 2008). It was thought to still exist above Guntersville Lake Possibly in Mobile Basin stream in northwest Georgia (it was known from Swamp Creek in Whitfield Co. in the late 1800s) although it is believed extirpated in that state (J. Wisniewski, GA NHP, pers. comm., January 2007). The only likely remaining population in the entire range is the upper Paint Rock River population (Williams et al., 2008) mentioned above.

Population Size: 1 - 1000 individuals
Population Size Comments: One viable population exists in the Estill Fork of the Paint Rock River. This one population appears to be limited to a reach of stream less than one mile in length. Dead shells are occasionally found throughout the upper Paint Rock River and its tributaries (Doug Shelton, pers. obs. 1995). Even in the 1970s and 1980s, few specimens were collected (2 specimens in Paint Rock River below Hurricane Creek; 6 fresh dead specimens at Estill Fork near Freedom Bridge) and numerous surveys have uncovered no specimens at all in the Paint Rock River, Hurricane Creek, and Estill Fork (see USFWS, 1984).

Number of Occurrences with Good Viability/Integrity: None (zero)

Overall Threat Impact: Very high - high
Overall Threat Impact Comments: Globally, threats to this species include alteration and destruction of stream habitat (due to impoundment for flood control, navigation, hydroelectric power, and recreation), siltation (due to strip mining, coal-washing, dredging, farming, logging, and road construction), pollution (caused by municipal, agricultural, and industrial waste discharges) (USFWS, 1984). Regionally, the most imminent threat to the largest population is the practice of the locals to ride four-wheel recreational vehicles in the stream. This has occurred with some regularity at the site in recent years. The reluctance of state and federal authorities to intervene may well jeopardize this last viable population. Silviculture upstream of this site in Franklin County, Tennessee may also threaten his population. (Doug Shelton - personal observation, 1995). Other sites within the drainage may be threatened by poor agricultural practices (Ahlstedt, 1991) and poor silvicultural practices (Doug Shelton - personal observation, 1995).

Short-term Trend: Decline of >70%
Short-term Trend Comments: This rare species is declining. It is extirpated from most of its historic range. Its historic and current rarity hinder the evaluation of decline in populations and their sizes. Even in the 1970s and 1980s, few specimens were collected (2 specimens in Paint Rock River below Hurricane Creek; 6 fresh dead specimens at Estill Fork near Freedom Bridge) and numerous surveys have uncovered no specimens at all in the Paint Rock River, Hurricane Creek, and Estill Fork (see USFWS, 1984).

Long-term Trend: Decline of >90%
Long-term Trend Comments: This species possibly occurred historically in a Mobile Basin stream in northwest Georgia (it was known from Swamp Creek in Whitfield Co. in the late 1800s) although it is believed extirpated in that state (J. Wisniewski, GA NHP, pers. comm., January 2007). In Alabama, it was previously endemic to the middle reaches of the Tennessee River system (a few mainstem records but most from tributaries) across much of northern Alabama, but is now extirpated except for the Paint Rock River population (Mirarchi, 2004; Williams et al., 2008).

Intrinsic Vulnerability: Highly vulnerable
Intrinsic Vulnerability Comments: The low numbers of individuals and the few populations of this species render it especially vulnerable to extinction.

Environmental Specificity: Narrow. Specialist or community with key requirements common.
Environmental Specificity Comments: This species is buried in firm rubble, gravel, and sand substrates in shallow riffles and shoals. It is a small stream species restricted to Tennessee River tributaries most often in clean, fast-flowing water in riffle areas in substrates that contain relatively firm rubble, gravel, and sand substrates swept free from siltation (USFWS, 1984). It is sometimes found in sand among aquatic grasses along the shoreline (Doug Shelton - personal observation, 1995).

Other NatureServe Conservation Status Information

Inventory Needs: Survey historic sites to determine the current presence/absence of the species at these sites. Estimate the number of individuals in the existing populations.

Protection Needs: Proper planning and involvement of representatives from various state and federal agencies could save the species.

Distribution
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Global Range: (<100 square km (less than about 40 square miles)) This is a small stream species restricted to the tributaries of the Tennessee River, but not the Tennessee River proper (USFWS, 1984). One exception is a single record from Swamp Creek, Whitfield County, Georgia (recorded in the late 1800s; likely now extirpated); absent from the upper Tennessee drainage. Ortmann collected specimens he thought were Toxolasma cylindrellus from Little Pigeon River, Sevier County, Tennessee. H. D. Athearn collected specimens from the Sequatchie and Little Sequatchie Rivers in Marion County, Tennessee back in the 1950s (Bogan and Parmalee, 1983; Parmalee and Bogan, 1998). It was found in the Paint Rock River and in Larkin Fork of the Paint Rock River (Bogan and Parmalee, 1983) and in the Estill Fork of the Paint Rock River (McGregor and Shelton, 1995). It has also been found in the Flint River and Indian Creek, Madison County, Alabama and in the Elk River, Franklin County, Tennessee. It was also found in the Duck River from Hickman County upstream to Bedford County, Tennessee (Bogan and Parmalee, 1983; Parmalee and Bogan, 1998). Currently it is believed to be extant only in the Paint Rock River Drainage in Jackson County, Alabama and adjacent Franklin Co., Tennessee (Bogan and Parmalee, 1985; Williams et al., 2008). It is found in the mainstem of the Paint Rock River and each of its major tributaries: Larkin Fork, Estill Fork, and Hurricane Creek (McGregor and Shelton, 1995) (Doug Shelton, pers. obs., 1996). The largest population was found in the Estill Fork from the Tennessee state line downstream for about one mile (Shelton, 1997) but its viability is now questionable. The Duck River population is believed to have been extirpated (Steve Ahlstedt, pers. comm. May, 1995), although Parmalee and Bogan (1998) believe it may still be hanging on.

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, GAextirpated, TN

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Jackson (01071), Lauderdale (01077)*, Madison (01089)*
GA Whitfield (13313)*
TN Coffee (47031)*, Franklin (47051), Grundy (47061)*, Lewis (47101)*, Marion (47115)*, Marshall (47117), Maury (47119)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Conasauga (03150101)*
06 Sequatchie (06020004)+*, Guntersville Lake (06030001)*, Wheeler Lake (06030002)+, Upper Elk (06030003)+*, Lower Elk (06030004)*, Pickwick Lake (06030005)+*, Upper Duck (06040002)+, Lower Duck (06040003)+, Buffalo (06040004)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
U.S. Distribution by Watershed (based on multiple information sources) Help
Ecology & Life History
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Basic Description: A small freshwater mussel or bivalve mollusk with a tawny or yellowish-green shell.
Diagnostic Characteristics: This species can be confused with TOXOLASMA LIVIDUS which is found along with it. TOXOLASMA CYLINDRELLUS is larger and more elongate. The periostracum is usually lighter in color. The nacre of TOXOLASMA LIVIDUS is usually a dark purple throughout. The nacre of TOXOLASMA CYLINDRELLUS is a purplish bronze inside the pallial line and a lighter cream outside the pallial line.
Reproduction Comments: The glochidial host is not known.
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): CREEK, High gradient, Pool, Riffle
Special Habitat Factors: Benthic
Habitat Comments: This species is buried in firm rubble, gravel, and sand substrates in shallow riffles and shoals. It is a small stream species restricted to Tennessee River tributaries most often in clean, fast-flowing water in riffle areas in substrates that contain relatively firm rubble, gravel, and sand substrates swept free from siltation (USFWS, 1984). It is sometimes found in sand among aquatic grasses along the shoreline (Doug Shelton - personal observation, 1995).
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 1976 and a recovery plan (USFWS, 1984) created. Recovery objectives include: (1) preserve populations and presently used habitat with emphasis on the Paint Rock River, Estill Fork, and Hurricane Creek, (2) determine the feasibility of introducing the species back into rivers within its historic range and introduce where feasible, (3) conduct life history studies not covered (i.e. fish hosts, age and growth, reproductive biology, longevity, natural mortality factors, and population dynamics), (4) determine the number of individuals required to maintain a viable population, (5) investigate the necessity for habitat improviement and identify techniques and sites for improvement to include implementation, (6) develop and implement a program to monitor population levels and habitat conditions of presently established populations as well as introduced and expanding populations, (7) assess overall success of recovery program and recommend action.
Biological Research Needs: 1. Determine the extent of current populations within the Paint Rock River drainage. 2. Determine fish host(s). 3. Describe glochidia. 4. Determine if culturing of the species is a viable option for conservation. 5. Assess sites for possible relocation should culturing of the species prove successful.
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: 19Dec2011
NatureServe Conservation Status Factors Author: Cordeiro, J. (2011); Shelton, Douglas N. (1997)
Management Information Edition Date: 13Jun2007
Management Information Edition Author: Cordeiro, J.
Element Ecology & Life History Edition Date: 13Jun2007
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
Help
  • Ahlsted, S.A. 1984. Twentieth Century changes in the freshwater mussel fauna of the Clinch River (Tennessee and Virginia). M.S. Thesis. Wildlife and Fisheries, University of Tennessee, Knoxville, Tennessee. 102 pp.

  • Ahlstedt, S. A. 1991. Status survey for federally listed endangered freshwater mussel species in the Paint Rock River system, northeastern Alabama. Norris Tennessee Valley Authority file report submitted to U.S. Fish and Wildlife Service. 26 pp.

  • Bogan, A.E. and P.W. Parmalee. 1983. Tennessee's rare wildlife. Vol. 2: The mollusks. Tennessee Wildlife Resources Agency and the Tennessee Conservation Department: Nashville, Tennessee. 123 pp.

  • Campbell, D. and P. Harris. 2006. Report on molecular systematics of poorly-known freshwater mollusks of Alabama. Report to the Alabama Department of Conservation and Natural Resources, Montgomery, Alabama. 34 pp.

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

  • 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.

  • McGregor, S. W., and D. N. Shelton. 1995. A qualitative assessment of the unionid fauna of the headwaters of the Paint Rock and Flint Rivers of north Alabama and adjacent areas of Tennessee, 1995. Geological Survey of Alabama, Environmental Geology Division, in cooperation with the Alabama Department of Conservation and Natural Resources. 23 pp. + appendix.

  • 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.

  • Mirarchi, R.E., et al. 2004a. Alabama Wildlife. Volume One: A Checklist of Vertebrates and Selected Invertebrates: Aquatic Mollusks, Fishes, Amphibians, Reptiles, Birds, and Mammals. University of Alabama Press: Tuscaloosa, Alabama. 209 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.

  • Shelton, D. N. 1997. The Paint Rock River Initiative. Pages 68-71 in K. S. Cummings, A. C. Buchanan, C. A. Mayer, and T. J. Naimo (editors). Conservation and Management of Freshwater Mussels II: Initiatives for the Future. Proceedings of a UMRCC Symposium, 16-18 October, 1995, St. Louis, Missouri. Upper Mississippi River Conservation Committee, Rock Island, Illinois.

  • 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) (Ahlstedt, S.) 1984. Recovery plan for the pale lilliput pearly mussel; Toxolasma (Carunculina) cyllindrellus (Lea, 1868) U.S. Fish and Wildlife Service, Region 4, Atlanta, Georgia.

  • 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
  • Ahlstedt, S.A. 1995-1996. Status survey for federally listed endangered freshwater mussel species in the Paint Rock River system, northeastern Alabama, U.S.A. Walkerana 8(19):63-80.

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