Quadrula intermedia - (Conrad, 1836)
Cumberland Monkeyface
Other English Common Names: Cumberland Monkey-face Pearly Mussel
Taxonomic Status: Accepted
Related ITIS Name(s): Quadrula intermedia (Conrad, 1836) (TSN 80070)
Unique Identifier: ELEMENT_GLOBAL.2.116132
Element Code: IMBIV39070
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Quadrula
Genus Size: D - Medium to large genus (21+ 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: Quadrula intermedia
Taxonomic Comments: Quadrula intermedia has been synonymized in the past with Quadrula tuberosa and Quadrula sparsa (Ortmann, 1918). The relationship to Quadrula tuberosa, Quadrula intermedia, and Quadrula sparsa is questionable; however, both Q. tuberosa and Q. sparsa have been found historically within the range of Q. intermedia and no intergrade specimens have been found (USFWS, 1984). Q. tuberosa may be the form of Q. intermedia occurring in big rivers, where a more obese form typically occurs in many unionid species. Quadrula sparsa is currently recognized by Turgeon et al. (1998). Distributional records became confused when Ortmann (1914; 1918) lumped Quadrula sparsa and Quadrula tuberosa under Quadrula intermedia (Bogan and Parmalee, 1983; Parmalee and Bogan, 1998). Historical records for Q. tuberosa from the Cumberland River are therefore also included with historical records for Q. sparsa (USFWS, 1984) but if these are distinct species, then Q. sparsa was not present in the headwaters of the Cumberland River (Parmalee and Bogan, 1998). At Muscle Shoals, Morrison (1942) described a more inflated form as Quadrula biangulata and shells similar to this form have been recovered from archaeological remains in the lower Clinch River, Tennessee, and identified as Quadrula sparsa (Parmalee and Bogan, 1986). Parmalee and Bogan (1998) placed Q. biangulata in the synonymy of Quadrula quadrula. Whether or not the biangulata form represents a valid species is unclear as it is conchologically different from typical Q. intermedia but appears to represent one extreme in range of variation of the species. Williams et al. (2008) tentatively place it in synonymy with Q. sparsa.
Conservation Status
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NatureServe Status

Global Status: G1
Global Status Last Reviewed: 19Dec2011
Global Status Last Changed: 16Sep1997
Rounded Global Status: G1 - Critically Imperiled
Reasons: This species has been extirpated from nearly all of its former range. A few extant occurrences exist on the Duck, Elk, and Powell Rivers, and these do not appear to have healthy populations with possibly the only viable one left on the Powell River (a small 0.8 km linear stretch). The species is very close to global extinction and is continually threatened with habitat degradation and pollution.
Nation: United States
National Status: N1 (16Sep1997)

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 (SX), Tennessee (S1), Virginia (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE, XN: Listed endangered, nonessential experimental population (14Jun1976)
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 it is 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, 12 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: EN - 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: 1000-5000 square km (about 400-2000 square miles)
Range Extent Comments: Historically, this species was widespread in the upper Tennessee River system (Tennessee, Elk, Duck, Holston, north and south fork Holston, Nolichucky, French Broad, Tellico, Clinch, Powell Rivers) (Simpson, 1914; Ortmann, 1918) in Tennessee, Alabama, and Virginia, and possibly in the Cumberland River system (Cumberland, Big South Fork Cumberland, Caney Fork) where its former occurrence remains uncertain because the closely related Quadrula tuberosa was also reported there (USFWS, 1984). It is likely that Ortmann's 1918-1925 records for the Cumberland River system were probably Quadrula tuberosa, here recognized as a synonym of Quadrula intermedia (USFWS, 1984). Since 1960, it has been found in large tributaries of the Tennessee River including the Duck, Clinch, Elk and Powell Rivers. Since 1970, it has been found only in the Clinch, Powell and Tellico Rivers (USFWS, 1984). It was recently found alive in the Duck River in Tennessee (Louis Levine, pers. comm. 10/7/1997). It appears to be extirpated from Alabama, although reintroduction efforts are underway (Mirarchi et al., 2004).

Area of Occupancy: 501-12,500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 1 - 5
Number of Occurrences Comments: There are very few recent records for this species. Surveys of the Duck River in the 1970s uncovered only about a dozen live specimens (rm 173, 171, 163) plus a few more in the early 1980s (rm 179); surveys of the Elk River in during the 1970s-1980s found a few live specimens (rm 106, 110); while Powell River surveys during the same time found live specimens in a few locationis (rm 99, 106, 117, 130, 167) (USFWS, 1984). A 210 km survey of the Elk River from the Alabama border through Tennessee in 1980 found this species at a few sites in Lincoln Co., Tennessee (Ahlstedt, 1983). Evidence since 1984 (USFWS, 1984) suggests populations from the Powell River (approximately 31 miles extending from Tennessee into Lee Co., Virginia) are declining significantly but represent the healthiest remaining population and the Duck River, Tennessee, has too few specimens to be viable. Other historical sites in Tennessee including the Elk River, Lincoln and Franklin Cos., are likely extirpated (Parmalee and Bogan, 1998). An August 1997 survey of the Duck River population in Tennessee only found two very old individuals, previously one was found in 1995, and none in 1993 and 1994 surveys (Watson, 1998). No other surveys in other Tennessee rivers uncovered this species. It appears since 1989, range in the Powell River has expanded by 3.8 linear km and overall the species occurs in portions of a 100 km reach of the river, however age studies indicate little recruitment outside a small 0.8 km linear stretch of the river in recent years (Johnson, 2011).

Population Size: 1 - 1000 individuals
Population Size Comments: Extant populations are scattered and have typically supported low numbers (Bogan and Parmalee, 1983). An August 1997 survey of the Duck River population in Tennessee only found two very old individuals, previously one was found in 1995, and none in 1993 and 1994 surveys (Watson, 1998). A 210 km survey of the Elk River from the Alabama border through Tennessee in 1980 found this species at a few sites in Lincoln Co., Tennessee (only a few specimens total) (Ahlstedt, 1983).

Number of Occurrences with Good Viability/Integrity: None (zero)
Viability/Integrity Comments: Barr et al. (1994) determined (based on 1981 survey data) that viable populations exist in Powell River at Buchanan Ford (pop. est. 115) and at Fletcher Fork (pop. est. 90). The only remaining viable population is likely that in the upper Powell River but that population is declining (Parmalee and Bogan, 1998).

Overall Threat Impact: Very high - high
Overall Threat Impact Comments: Threats include impoundment (for flood control, navigation, hydroelectric power production, and recreation) including Norris Dam and Columbia Dam, siltation (due to strip mining, coal washing, dredging, farming, logging, and road construction), and pollution (municipal, agricultural, and industrial) from sawdust (logging), coal mine acids, toxic wastes, gravel dredging, fertilizers, pesticides, chemical spills and discharges (USFWS, 1984).

Short-term Trend: Decline of >70%
Short-term Trend Comments: Hubbs (2002) reported relict shells only in the Elk River (RM 105.4), Tennessee. It appears since 1989, range in the Powell River has expanded by 3.8 linear km and overall the species occurs in portions of a 100 km reach of the river, however age studies indicate little recruitment outside a small 0.8 km linear stretch of the river in recent years (Johnson, 2011).

Long-term Trend: Decline of >90%
Long-term Trend Comments: The range of this species was formerly restricted to the upper Tennessee River system but was never abundant (Simpson, 1914; USFWS, 1984). It was reported from the upper Elk River, Tennessee (Isom et al., 1973). It is extirpated from the Upper Clinch, and North and South Fork Holston Rivers (USFWS, 1984) and has not been seen in Alabama in the Tennessee River downstream of Muscle Shoals since the river was impounded in the early 1900s (Mirarchi, 2004; Williams et al., 2008). Still, it has declined significantly throughout its range now found in only 2 or 3 rivers (most not viable).

Intrinsic Vulnerability: Highly vulnerable

Environmental Specificity: Narrow. Specialist or community with key requirements common.
Environmental Specificity Comments: This species inhabits shallow riffle and shoal areas of headwater streams and bigger rivers. It prefers clean, fast-flowing water in shoal conditions, and has never been found in the ponded stretches of rivers, nor is it known from small streams (USFWS, 1984). It has been found living in a sand and gravel substrate in 6 inches to 2 feet of water (Bogan and Parmalee, 1983).

Other NatureServe Conservation Status Information

Inventory Needs: Search for more EOs and verify old. Continued monitoring of the population.

Protection Needs: See Recovery Plan

Distribution
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Global Range: (1000-5000 square km (about 400-2000 square miles)) Historically, this species was widespread in the upper Tennessee River system (Tennessee, Elk, Duck, Holston, north and south fork Holston, Nolichucky, French Broad, Tellico, Clinch, Powell Rivers) (Simpson, 1914; Ortmann, 1918) in Tennessee, Alabama, and Virginia, and possibly in the Cumberland River system (Cumberland, Big South Fork Cumberland, Caney Fork) where its former occurrence remains uncertain because the closely related Quadrula tuberosa was also reported there (USFWS, 1984). It is likely that Ortmann's 1918-1925 records for the Cumberland River system were probably Quadrula tuberosa, here recognized as a synonym of Quadrula intermedia (USFWS, 1984). Since 1960, it has been found in large tributaries of the Tennessee River including the Duck, Clinch, Elk and Powell Rivers. Since 1970, it has been found only in the Clinch, Powell and Tellico Rivers (USFWS, 1984). It was recently found alive in the Duck River in Tennessee (Louis Levine, pers. comm. 10/7/1997). It appears to be extirpated from Alabama, although reintroduction efforts are underway (Mirarchi et al., 2004).

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 ALextirpated, TN, VA

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Colbert (01033)*, Jackson (01071)*, Lauderdale (01077)*, Limestone (01083)*
TN Claiborne (47025), Franklin (47051)*, Giles (47055)*, Hamilton (47065)*, Hancock (47067), Hawkins (47073)*, Knox (47093)*, Lincoln (47103), Marshall (47117), Maury (47119), Monroe (47123)*, Sullivan (47163)*
VA Lee (51105), Russell (51167)*, Scott (51169)*, Washington (51191)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
06 North Fork Holston (06010101)+*, South Fork Holston (06010102)+*, Holston (06010104)+*, Nolichucky (06010108)*, Watts Bar Lake (06010201)*, Lower Little Tennessee (06010204)+*, Upper Clinch (06010205)+*, Powell (06010206)+, Lower Clinch (06010207)*, Middle Tennessee-Chickamauga (06020001)+*, Guntersville Lake (06030001)+*, Wheeler Lake (06030002)*, Upper Elk (06030003)+, Lower Elk (06030004)+*, Pickwick Lake (06030005)+*, Lower Tennessee-Beech (06040001)*, Upper Duck (06040002)+, Lower Duck (06040003)+*
+ 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 medium-size freshwater mussel or bivalve mollusk with a greenish-yellow to yellowish-green shell that darkens with age.
Reproduction Comments: Glochidial hosts include streamlined chub (Erymystax dissimilis) and blotched chub (Erymystax insignis) (Hill, 1986; Yeager and Saylor, 1995).
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, High gradient, MEDIUM RIVER, Moderate gradient, Riffle
Special Habitat Factors: Benthic
Habitat Comments: This species inhabits shallow riffle and shoal areas of headwater streams and bigger rivers. It prefers clean, fast-flowing water in shoal conditions, and has never been found in the ponded stretches of rivers, nor is it known from small streams (USFWS, 1984). It has been found living in a sand and gravel substrate in 6 inches to 2 feet of water (Bogan and Parmalee, 1983).
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 created (USFWS, 1984).

The recovery plan (USFWS, 1984) lists the following objectives: (1) preserve populations and presently used habitat with emphasis on the Duck, Elk, and Powell Rivers, (2) determine the feasibility of introducing the species back into rivers within its historic range and introduce where feasible, (3) determine the number of individuals required to maintain a vialbe population, (4) investigate the necessity for habitat improvement and, if feasible and desirable, identify techniques and sites for improvement to include implementation, (5) develop and implement a program to monitor population levels and habitat conditions of presently established populations as well as introduced and expanding populations, (6) assess overall success of recovery program and recommend action.

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 (USFWS, 2001).

The USFWS, in cooperation with the State of Tennessee and Conservation Fisheries, Inc., proposes 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 (USFWS, 2006).

Biological Research Needs: 1. Determine aspects of life history in regard to spawning period and fish host identification. 2. Determine if culturing is a viable means of conservation. 3. Determine if fish host is in need of similar culturing. 4. Assess potential sites for reintroduction should culturing 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: 20Jun2007
Management Information Edition Author: Cordeiro, J.
Element Ecology & Life History Edition Date: 19Dec2011
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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  • Ahlstedt, S.A. 1983. The molluscan fauna of the Elk River in Tennessee and Alabama. American Malacological Bulletin 1:43-50.

  • Ahlstedt, S.A. 1984. Twentieth century changes in the freshwater mussel fauna of the Clinch River (Tennessee and Virginia). M.S. Thesis, The University of Tennessee, Knoxville, Tennessee. 102 pp.

  • Ahlstedt, Steven A. 1986. Activity 1: Mussel Distribution Surveys. Cumberlandian Mollusk Conservation Program. TVA.

  • 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.

  • Hill, D.M. 1986. Cumberlandian mollusk conservation program, activity 3: identification of fish hosts. Office of Natural Resources and Economic Development, Tennessee Valley Authority, Knoxville, Tennessee. 55 pp.

  • 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.

  • Johnson, M.S. 2011. A quantitative survey of the freshwater mussel fauna in the Powell River of Virginia and Tennessee, and life history study of two endangered species, Quadrula sparsa and Quadrula intermedia. M.S. Thesis, Virginia Polytechnic Institution. 171 pp.

  • 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.

  • 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.

  • Morrison, J.P.E. 1942. Preliminary report on mollusks found in the shell mounds of the Pickwidk Landing basin in the Tennessee River valley. Bureau of American Ethnology Bulletin, 129: 339-392.

  • 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.

  • Ortmann, A.E. 1914. Studies in naiades (in partim). The Nautilus, 28: 28-34.

  • Ortmann, A.E. 1918c. The nayades (freshwater mussels) of the Upper Tennessee drainage. With notes on synonymy and distribution. Proceedings of the American Philosophical Society 57: 521-626.

  • Parmalee, P.W. and A.E. Bogan. 1986. Molluscan remains from aboriginal middens at the Clinch River Breeder Reactor Plant Site, Roan County, Tennessee. American Malacological Bulletin 4(1):25-37.

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

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

  • Simpson, C.T. 1914. A Descriptive Catalogue of the Naiades or Pearly Fresh-water Mussels. Bryant Walker: Detroit, Michigan. 1540 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) (Ahlstedt, S.). 1984. Recovery plan for the Cumberland monkeyface pearly mussel; Quadrula intermedia (Conrad, 1836). U.S. Fish and Wildlife Service, Region 4, Atlanta, Georgia. 35 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.

  • Watson, S.N., Jr. 1998. Lillard Mill mussel survey, 1997. Triannual Unionid Report, 14: 7-8.

  • 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.

  • Yeager, B.L. and C.F. Saylor. 1995. Fish hosts for four species of freshwater mussels (Pelecypoda: Unionidae) in the Upper Tennessee River drainage. American Midland Naturalist, 133(1): 1-6.

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.

  • Hubbs, D. 2002. Monitoring and management of endangered mussels. 2001-02 Annual Report Project 7365, Tennessee Wildlife Resources Agency, Nashville, Tennessee. 3 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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Citation for data on website including State Distribution, Watershed, and Reptile Range maps:
NatureServe. 2018. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia. Available http://explorer.natureserve.org. (Accessed:

Citation for Bird Range Maps of North America:
Ridgely, R.S., T.F. Allnutt, T. Brooks, D.K. McNicol, D.W. Mehlman, B.E. Young, and J.R. Zook. 2003. Digital Distribution Maps of the Birds of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Bird Range Maps of North America:
"Data provided by NatureServe in collaboration with Robert Ridgely, James Zook, The Nature Conservancy - Migratory Bird Program, Conservation International - CABS, World Wildlife Fund - US, and Environment Canada - WILDSPACE."

Citation for Mammal Range Maps of North America:
Patterson, B.D., G. Ceballos, W. Sechrest, M.F. Tognelli, T. Brooks, L. Luna, P. Ortega, I. Salazar, and B.E. Young. 2003. Digital Distribution Maps of the Mammals of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Mammal Range Maps of North America:
"Data provided by NatureServe in collaboration with Bruce Patterson, Wes Sechrest, Marcelo Tognelli, Gerardo Ceballos, The Nature Conservancy-Migratory Bird Program, Conservation International-CABS, World Wildlife Fund-US, and Environment Canada-WILDSPACE."

Citation for Amphibian Range Maps of the Western Hemisphere:
IUCN, Conservation International, and NatureServe. 2004. Global Amphibian Assessment. IUCN, Conservation International, and NatureServe, Washington, DC and Arlington, Virginia, USA.

Acknowledgement Statement for Amphibian Range Maps of the Western Hemisphere:
"Data developed as part of the Global Amphibian Assessment and provided by IUCN-World Conservation Union, Conservation International and NatureServe."

NOTE: Full metadata for the Bird Range Maps of North America is available at:
http://www.natureserve.org/library/birdDistributionmapsmetadatav1.pdf.

Full metadata for the Mammal Range Maps of North America is available at:
http://www.natureserve.org/library/mammalsDistributionmetadatav1.pdf.

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