Plethobasus cicatricosus - (Say, 1829)
White Wartyback
Other English Common Names: White Warty-back Pearly Mussel
Taxonomic Status: Accepted
Related ITIS Name(s): Plethobasus cicatricosus (Say, 1829) (TSN 80227)
Unique Identifier: ELEMENT_GLOBAL.2.119203
Element Code: IMBIV34010
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Plethobasus
Check this box to expand all report sections:
Concept Reference
Help
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: Plethobasus cicatricosus
Taxonomic Comments: Thomas Say originally described Unio cicatricosus in 1829. Isaac Lea described Unio varicosus later the same year. Both names were widely used in the literature; however, varicosus Lea was pre-occupied by varicosus Lamarck, 1819. Frierson (1911) synonomized Unio cicatricosus Say with Unio aesopus Green, 1827 (= Plethobasus cyphyus Rafinesque, 1820) and renamed Lea's shell as Unio detectus Frierson, 1911. He also named two additional species Unio cicatricoides and Unio comptertus as being part of this complex. In 1919, Ortmann synonomized all of Frierson's names under cicatricosus Say. Three years later Ortmann and Walker (1922) sunk cicatricosus into cyphyus. Recent authors have not followed Ortmann and Walker (1922) and recognize Plethobasus cicatricosus as distinct (Parmalee, 1967; Bogan and Parmalee, 1983; Cummings and Mayer, 1992; Turgeon et al., 1998).
Conservation Status
Help

NatureServe Status

Global Status: G1
Global Status Last Reviewed: 07May2009
Global Status Last Changed: 17Mar1998
Rounded Global Status: G1 - Critically Imperiled
Reasons: Long-term viability of this mussel (if in fact it still exists) is doubtful. No known living populations have been found in numerous surveys of the Tennessee River since the 1960s although a fresh dead specimens were found in 1979 and 1982. The only known extant population might occur in tailwaters of Wilson Dam on Tennessee River where it is rare; otherwise all occurrences throughout its entire historic range appear extirpated.
Nation: United States
National Status: N1 (17Mar1998)

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

Other Statuses

U.S. Endangered Species Act (USESA): LE, XN: Listed endangered, nonessential experimental population (14Jun1976)
Comments on USESA: Listed Endangered throughout its range. 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: 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: Historically, this species was widely distributed in the Ohio, Cumberland, and Tennessee drainages including the Wabash, Ohio, Kanawha, Cumberland, Holston, and Tennessee rivers (USFWS, 1984). No live specimens have been found in the Wabash, Ohio, Kanawha, Cumberland, or Holston rivers since around the turn of the century. It possibly still exists in a very short reach of the Tennessee River mainstem below Pickwick Dam near Savannah, Tennessee, near river mile 207 because all of the most recent historical records were from the original Tennessee River channel (USFWS, 1984; Parmalee and Bogan, 1998). The only other potential extant population is in the tailwaters of Wilson Dam on the Tennessee River where it is rare (Mirarchi et al., 2004). It historically occurred in the Wabash mainstem in Indiana (Fisher, 2006).

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

Number of Occurrences: 1 - 5
Number of Occurrences Comments: The only recent record was the collection of fresh dead shells from a commercial shell pile in 1979 and 1982 below Pickwick Dam near Savannah, Tennessee (USFWS, 1984). Both collections were older specimens. Currently the only known extant population might occur in the tailwaters of Wilson Dam on the Tennessee River in northwest Alabama where it is rare (Mirarchi et al., 2004).

Population Size: 1 - 1000 individuals
Population Size Comments: No live specimens have been collected in over 30 years and no estimates of population size or abundance have been made but what few live specimens have been found have been single old individuals (USFWS, 1989).

Number of Occurrences with Good Viability/Integrity: None (zero)
Viability/Integrity Comments: In the remaining 18 km of the Tennessee River downstream of Wilson Dam, evidence of recruitement has been observed but the species is very rare (Williams et al., 2008).

Overall Threat Impact: Very high - high
Overall Threat Impact Comments: Smith (1971) ranked the causes of extirpation or declines in fish species as follows: siltation, drainage of bottomland lakes, swamps, and prairie marshes, desiccation during drought, species introductions, pollution, impoundments, and increased water temperatures. All of these factors render habitats unsuitable, cause extirpations, and lead to the isolation of populations thereby increasing their vulnerability to extirpation for many aquatic species (including mussels) throughout North America. Zebra mussels, Dreissena polymorpha, have destroyed mussel populations in the Great Lakes and significantly reduced mussels in many of the large rivers of eastern North America. Zebra mussels have the potential to severely threaten other populations especially if they make their way into smaller streams. Pollution through point (industrial and residential discharge) and non-point (siltation, herbicide and fertilizer run-off) sources is perhaps the greatest on-going threat to this species and most freshwater mussels. Lowered dissolved oxygen content and elevated ammonia levels (frequently associated with agricultural runoff and sewage discharge) have been shown to be lethal to some species of freshwater naiads (Horne and McIntosh, 1979). Residential, mineral and industrial development also pose a significant threat. Rotenone, a toxin used to kill fish in bodies of water for increased sport fishery quality, has been shown to be lethal to mussels as well (Heard, 1970). Destruction of habitat through stream channelization and maintenance and the construction of dams is still a threat in some areas. Impoundments reduce currents that are necessary for basic physiological activities such as feeding, waste removal and reproduction. In addition, reduced water flow typically results in a reduction in water oxygen levels and a settling out of suspended solids (silt, etc.), both of which are detrimental. Dredging of streams has an immediate effect on existing populations by physically removing and destroying individuals. Dredging also affects the long-term recolonization abilities by destroying much of the potential habitat, making the substrates and flow rates uniform throughout the system. Natural predators include raccoons, otter, mink, muskrats, turtles and some birds (Simpson, 1899; Boepple and Coker, 1912; Evermann and Clark, 1918; Coker et al., 1921; Parmalee, 1967; Snyder and Snyder, 1969). Domestic animals such as hogs can root mussel beds to pieces (Meek and Clark, 1912). Fishes, particularly catfish, Ictalurus spp. and Amieurus spp., and freshwater drum, Aplodinotus grunniens, also consume large numbers of unionids.

USFWS (1984) lists the following reasons for decline: impoundment (for flood control, navigation, hydroelectric power production, and recreation) causing altered temperature regime, extreme water level fluctuations, reduced turbidity, seasonal oxygen deficits, and high concentrations of heavy metals; siltation (from strip mining, coal washing, dredging, farming, logging, and road construction) causing increased silt transport, increased turbidity, erosion of bank habitat, elimination of host fish; pollution (municipal, agricultural, and industrial waste discharges).

Short-term Trend: Decline of >70%
Short-term Trend Comments: Distribution has been greatly diminished with local and state extirpations, e.g. the Wabash in Illinois (Cummings and Mayer, 1997) with all recent records bordering on historical and only one or two specimens found in the Tennessee River (USFWS, 1989).

Long-term Trend: Decline of >90%
Long-term Trend Comments: No live specimens have been found in the Holston River, Cumberland River, Ohio River, Wabash River, and Konawha River since before the turn of the 20th Century (USFWS, 1989). It is extirpated from Kentucky but formerly occurred in the Ohio and lower Cumberland rivers (Cicerello and Schuster, 2003). It historically occurred in the Wabash mainstem in Indiana (Fisher, 2006) but only old weathered shells have been found there recently and it is believed extirpated from the state (Ron Hellmich, IN NHP, pers. comm., March 2008). In Ohio, it is extirpated but historically occurred in the Ohio River at Marietta dn Clarington (turn of 20th Century) and Ohio River at Cincinnati and Manchester, and lower Muskingum River (Watters et al., 2009). It is extirpated from the McAlpine dam pool in the Ohio River at Louisville, Kentucky (and Indiana) (Watters and Flaute, 2010).

Intrinsic Vulnerability: Highly vulnerable
Intrinsic Vulnerability Comments: If any populations still remain, they are isolated and extremely small with poor viability in the Tennessee River with poor dispersal capability (USFWS, 1989).

Environmental Specificity: Very narrow. Specialist or community with key requirements scarce.
Environmental Specificity Comments: This species is restricted to riffles in large rivers; most of which are now impounded.

Other NatureServe Conservation Status Information

Inventory Needs: The historical distribution is reasonably well known. A concentrated search for this species should be conducted to assess its current status. If live animals are found an effort to artificially propagate the mussel should be undertaken.

Distribution
Help
Global Range: (<100 square km (less than about 40 square miles)) Historically, this species was widely distributed in the Ohio, Cumberland, and Tennessee drainages including the Wabash, Ohio, Kanawha, Cumberland, Holston, and Tennessee rivers (USFWS, 1984). No live specimens have been found in the Wabash, Ohio, Kanawha, Cumberland, or Holston rivers since around the turn of the century. It possibly still exists in a very short reach of the Tennessee River mainstem below Pickwick Dam near Savannah, Tennessee, near river mile 207 because all of the most recent historical records were from the original Tennessee River channel (USFWS, 1984; Parmalee and Bogan, 1998). The only other potential extant population is in the tailwaters of Wilson Dam on the Tennessee River where it is rare (Mirarchi et al., 2004). It historically occurred in the Wabash mainstem in Indiana (Fisher, 2006).

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, TN, WVextirpated

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Colbert (01033), Lauderdale (01077)
IN Fountain (18045)*, Gibson (18051)*, Knox (18083)*, Marion (18097)*, Parke (18121)*, Posey (18129)*, Tippecanoe (18157)*, Vermillion (18165)*, Vigo (18167)*, Warren (18171)*
KY Oldham (21185)*
TN Anderson (47001)*, Claiborne (47025)*, DeKalb (47041)*, Grainger (47057)*, Hardin (47071), Jefferson (47089)*, Perry (47135), Smith (47159)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
05 Little Muskingum-Middle Island (05030201)*, Upper Ohio-Shade (05030202)*, Muskingum (05040004)*, Upper Kanawha (05050006)*, Lower Kanawha (05050008)*, Ohio Brush-Whiteoak (05090201)*, Middle Ohio-Laughery (05090203)*, Middle Wabash-Deer (05120105)+*, Middle Wabash-Little Vermilion (05120108)+*, Vermilion (05120109)+*, Middle Wabash-Busseron (05120111)+*, Lower Wabash (05120113)+*, Upper White (05120201)+*, Caney (05130108)+*, Lower Cumberland-Old Hickory Lake (05130201)+*, Lower Cumberland (05130205)*, Red (05130206)*, Silver-Little Kentucky (05140101)+*, Highland-Pigeon (05140202)+*
06 Holston (06010104)+*, Powell (06010206)+*, Lower Clinch (06010207)+*, Middle Tennessee-Chickamauga (06020001)*, Guntersville Lake (06030001)*, Wheeler Lake (06030002)*, Pickwick Lake (06030005)+, Lower Tennessee-Beech (06040001)+
+ 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
Help
Basic Description: An oblong or elongate freshwater mussel, light brown, with a row of large tear-shaped tubercles running down the side of the shell.
General Description: SHELL EXTERIOR: Shell oblong, elongate, thick, and moderately inflated. Anterior end rounded, posterior end broadly rounded. Dorsal and ventral margins curved. Umbos low, directed forward and not elevated above the hinge line. Shell smooth, except for a single row of large pustules or knobs on the center of the valve, running from below the umbo to the edge. Periostracum yellow or light brown in juveniles, becoming chestnut to dark brown in adults. Faint green rays present on the umbo. Length to five inches.

SHELL INTERIOR: Pseudocardinal teeth large and well developed; two in the right valve, one in the right, with a smaller tooth on either side. Lateral teeth rather short, straight or slightly curved; two in the left valve, one in the right Beak cavity relatively shallow. Nacre white, iridescent posteriorly (Cummings and Mayer, 1992).

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, Riffle
Special Habitat Factors: Benthic
Habitat Comments: This species was presumed to inhabit shoals and riffles in large rivers like the Tennessee (Bogan and Parmalee, 1983).
Phenology Comments: Life history information is not known.
Economic Attributes Not yet assessed
Help
Management Summary
Help
Stewardship Overview: This species was listed as federally endangered in the U.S. in 1976 and a recovery plan created (USFWS, 1984).

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).

The recovery plan (USFWS, 1984) outlines the following strategy: (1) preserve any known populations and presently used habitat, (2) determine the feasibility of introducing the species back into rivers within its historic range and introduce where feasible, (3) conduct life history studies, (4) determine the number of individuals required to maintain a viable population, (5) investigate the necessity for habitat improvement and, if feasible and desirable, 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: In order to effectively manage mussel species it is necessary to work out certain life history characteristics first. Because of their unusual life-cycle and dependence on fish for completion of that cycle, it is imperative that the host species for the white wartyback be ascertained. Life history studies need to be done to identify age and size at sexual maturity, recruitment success, age class structure, and other important life history parameters.

Research is needed to assess the success of watershed protection on mussel populations. Abundance and distribution of selected species needs to be monitored in order to ascertain how species abundances change over time. From that we can assess what land-use changes, conservation practices, and physical/chemical parameters are correlated with, and possibly responsible for, the biological changes.

Population/Occurrence Delineation
Help
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
Help
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
Help
Authors/Contributors
Help
NatureServe Conservation Status Factors Edition Date: 07May2009
NatureServe Conservation Status Factors Author: Cordeiro, J. (2009); Cummings, K. S. (1998)
Element Ecology & Life History Edition Date: 14Feb2007
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
  • Boepple, J.F. and R.E. Coker. 1912. Mussel resources of the Holston and Clinch rivers of eastern Tennessee. Bureau of Fisheries Document 765. 13 pp.

  • Bogan, A.E. and P.W. Parmalee. 1983. Tennessee's Rare Wildlife: Volume II: The Mollusks. Report to the Tennessee Natural Heritage Program and Tennessee Wildlife Resources Agency: Nashville, Tennessee. 123 pp.

  • Cicerello, R.R. and G.A. Schuster. 2003. A guide to the freshwater mussels of Kentucky. Kentucky State Nature Preserves Commission Scientific and Technical Series 7:1-62.

  • Coker, R.E., A.F. Shira, H.W. Clark, and A.D. Howard. 1921. Natural history and propagation of fresh-water mussels. Bulletin of the Bureau of Fisheries [Issued separately as U.S. Bureau of Fisheries Document 839] 37(1919-20):77-181 + 17 pls.

  • Cummings, K.S. and C.A. Mayer. 1992. Field Guide to Freshwater Mussels of the Midwest. Illinois Natural History Survey Manual 5, Illinois. 194 pp.

  • Cummings, K.S. and C.A. Mayer. 1997. Distributional checklist and status of Illinois freshwater mussels (Mollusca: Unionacea). Pages 129-145 in: K.S. Cummings, A.C. Buchanan, C.A. Mayer, and T.J. Naimo (eds.) Conservation and management of freshwater mussels II: initiatives for the future. Proceedings of a UMRCC Symposium, October 1995, St. Louis, Missouri. Upper Mississippi River Conservation Committee, Rock Island, Illinois.

  • Evermann, B.W. and H.W. Clark. 1918. The Unionidae of Lake Maxinkukee. Proceedings of the Indiana Academy of Science 1917:251-285.

  • 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.

  • Frierson, L.S. 1911. Remarks on Unio varicosus, Cicatricosus and Unio compertus, new species. The Nautilus, 25(5): 51-54 + 2 plates.

  • 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.

  • Heard, W.H. 1970. Eastern freshwater mollusks. 1. The south Atlantic and Gulf drainages. In: A.H. Clarke (ed.) Rare and endangered molluscs of North America. Malacologia 10:1-56.

  • Horne, F.R. and S. McIntosh. 1979. Factors influencing distribution of mussels in the Blanco River of central Texas. The Nautilus 94(4):119-133.

  • 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.

  • Meek, S. E., and H.W. Clark. 1912. The mussels of the Big Buffalo Fork of White River, Arkansas. Report and Special Papers of the U.S. Fish Commission [Issued separately as U.S. Bureau of Fisheries Document 759] 1911:1-20.

  • 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.

  • Natural Resources Commission. 2014. Roster of Indiana Animals, Insects, and Plants That Are Extirpated, Endangered, Threatened or Rare. Information Bulletin #2 (Sixth Amendment. 20pp.

  • Ortmann, A.E. 1919. Monograph of the naiades of Pennsylvania. Part III. Systematic account of the genera and species. Memoirs of the Carnegie Museum 8(1):1-385.

  • Ortmann, A.E. and B. Walker. 1922. On the nomenclature of certain North American naiades. Occasional Papers of the Museum of Zoology, University of Michigan, 112: 1-75.

  • Parmalee, P.W. 1967. The freshwater mussels of Illinois. Illinois State Museum, Popular Science Series 8:1-108.

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

  • Simpson, C.T. 1899. The pearly fresh-water mussels of the United States; their habits, enemies, and diseases, with suggestions for their protection. Bulletin of the U.S. Fish Commission [Issued separately as U.S. Bureau of Fisheries Document 413] 18(1898):279-288.

  • Smith, P.W. 1971. Illinois streams: A classification based on their fishes and an analysis of factors responsible for disappearance of native species. Illinois Natural History Survey Biological Notes 76:1-14.

  • Snyder, N. and H. Snyder. 1969. A comparative study of mollusk predation by Limpkins, Everglade Kites, and Boat-tailed Grackles. Eighth Annual Report of the Cornell Laboratory of Ornithology 8:177-223.

  • Stansbery, D.H. 1964. The mussel (muscle) shoals of the Tennessee River revisited.American Malacological Union Annual Report, 1964: 25-28.

  • 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). 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.

  • Watters, G.T. and C.J.M. Flaute. 2010. Dams, zebras, and settlements: The historical loss of freshwater mussels in the Ohio River mainstem. American Malacological Bulletin 28:1-12.

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

  • 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) (Ahlstedt, S.A.) 1984. Recovery plan for the white warty-back pearly mussel (Plethobasus cicatricosus) (Say 1829). U.S. Fish and Wildlife Service, Region 4, Atlanta, Georgia. 43 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.

Use Guidelines & Citation

Use Guidelines and Citation

The Small Print: Trademark, Copyright, Citation Guidelines, Restrictions on Use, and Information Disclaimer.

Note: All species and ecological community data presented in NatureServe Explorer at http://explorer.natureserve.org were updated to be current with NatureServe's central databases as of March 2018.
Note: This report was printed on

Trademark Notice: "NatureServe", NatureServe Explorer, The NatureServe logo, and all other names of NatureServe programs referenced herein are trademarks of NatureServe. Any other product or company names mentioned herein are the trademarks of their respective owners.

Copyright Notice: Copyright © 2018 NatureServe, 4600 N. Fairfax Dr., 7th Floor, Arlington Virginia 22203, U.S.A. All Rights Reserved. Each document delivered from this server or web site may contain other proprietary notices and copyright information relating to that document. The following citation should be used in any published materials which reference the web site.

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.

Restrictions on Use: Permission to use, copy and distribute documents delivered from this server is hereby granted under the following conditions:
  1. The above copyright notice must appear in all copies;
  2. Any use of the documents available from this server must be for informational purposes only and in no instance for commercial purposes;
  3. Some data may be downloaded to files and altered in format for analytical purposes, however the data should still be referenced using the citation above;
  4. No graphics available from this server can be used, copied or distributed separate from the accompanying text. Any rights not expressly granted herein are reserved by NatureServe. Nothing contained herein shall be construed as conferring by implication, estoppel, or otherwise any license or right under any trademark of NatureServe. No trademark owned by NatureServe may be used in advertising or promotion pertaining to the distribution of documents delivered from this server without specific advance permission from NatureServe. Except as expressly provided above, nothing contained herein shall be construed as conferring any license or right under any NatureServe copyright.
Information Warranty Disclaimer: All documents and related graphics provided by this server and any other documents which are referenced by or linked to this server are provided "as is" without warranty as to the currentness, completeness, or accuracy of any specific data. NatureServe hereby disclaims all warranties and conditions with regard to any documents provided by this server or any other documents which are referenced by or linked to this server, including but not limited to all implied warranties and conditions of merchantibility, fitness for a particular purpose, and non-infringement. NatureServe makes no representations about the suitability of the information delivered from this server or any other documents that are referenced to or linked to this server. In no event shall NatureServe be liable for any special, indirect, incidental, consequential damages, or for damages of any kind arising out of or in connection with the use or performance of information contained in any documents provided by this server or in any other documents which are referenced by or linked to this server, under any theory of liability used. NatureServe may update or make changes to the documents provided by this server at any time without notice; however, NatureServe makes no commitment to update the information contained herein. Since the data in the central databases are continually being updated, it is advisable to refresh data retrieved at least once a year after its receipt. The data provided is for planning, assessment, and informational purposes. Site specific projects or activities should be reviewed for potential environmental impacts with appropriate regulatory agencies. If ground-disturbing activities are proposed on a site, the appropriate state natural heritage program(s) or conservation data center can be contacted for a site-specific review of the project area (see Visit Local Programs).

Feedback Request: NatureServe encourages users to let us know of any errors or significant omissions that you find in the data through (see Contact Us). Your comments will be very valuable in improving the overall quality of our databases for the benefit of all users.