Leptodea fragilis - (Rafinesque, 1820)
Fragile Papershell
Taxonomic Status: Accepted
Related ITIS Name(s): Leptodea fragilis (Rafinesque, 1820) (TSN 80182) ;Medionidus mcglameriae van der Schalie, 1939 (TSN 80264)
French Common Names: leptodée fragile
Unique Identifier: ELEMENT_GLOBAL.2.827431
Element Code: IMBIV24010
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Leptodea
Genus Size: B - Very small genus (2-5 species)
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Concept Reference
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Concept Reference: 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.
Concept Reference Code: B08WIL01EHUS
Name Used in Concept Reference: Leptodea fragilis
Taxonomic Comments: Williams et al. (2008) place Medionidus mcglameriae in the synonymy of Leptodea fragilis, based on its thin shell, slight dorsal wing, blade-liek pseudocardinal teeth and absence of corrugations on the posterior slope.
Conservation Status
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NatureServe Status

Global Status: G5
Global Status Last Reviewed: 05May2009
Global Status Last Changed: 25Nov1996
Rounded Global Status: G5 - Secure
Reasons: This species has a very large range throughout the entire Mississippi River drainage into the Gulf of Mexico and Great Lakes and St. Lawrence River system. It is secure throughout its range.
Nation: United States
National Status: N5 (16Jul1998)
Nation: Canada
National Status: N4 (01Aug2017)

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 (S5), Arkansas (S5), Georgia (SNR), Illinois (S4), Indiana (S4), Iowa (S2), Kansas (S3), Kentucky (S4S5), Louisiana (S5), Michigan (SNR), Minnesota (SNR), Mississippi (S5), Missouri (S4), Nebraska (SNR), New York (S3), North Dakota (SNR), Ohio (S5), Oklahoma (S4), Pennsylvania (S2S3), South Dakota (S5), Tennessee (S5), Texas (S4), Vermont (S2), Virginia (S1), West Virginia (S3), Wisconsin (S3)
Canada Ontario (S4), Quebec (S2)

Other Statuses

IUCN Red List Category: EX - Extinct
American Fisheries Society Status: Currently Stable (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: This species is extremely wide-ranging occurring throughout all of the Mississippi River drainage; Gulf of Mexico drainage from Alabama to Texas (Burch, 1975) and the entire Great Lakes-St. Lawrence system in Canada (Clarke, 1981). Estimated range extent based on centroids of mapped level 8 hydrobasins (2015).

Number of Occurrences: > 300
Number of Occurrences Comments: It is throughout S Minnesota: Minnesota, St. Croix, Mississippi drainages below/above St. Anthony Falls, some southern streams (Sietman, 2003). It is common throughout Illinois (Cummings and Mayer, 1997; Schanzle and Cummings, 1991; Sietman et al., 2001; Schanzle et al., 2004; Tiemann et al., 2005). In Indiana: Wabash (Fisher, 2006), Tippecanoe (Cummings and Berlocher, 1990), East Fork White (Harmon, 1992), St. Joseph, St. Mary's, Maumee (Pryor, 2005). It is widely distributed in Ohio (Watters, 1992; 1995; Watters et al., 2009); Black (Lyons et al., 2007), Lower Maumee (Grabarciewicz, 2008); Raccoon (Hoggarth et al., 2007). In South Dakota, it is in the Minnesota, mid-upper Big Sioux (Skadsen and Perkins, 2000), Lake Sharpe Flats and Oahe (Shearer et al., 2005), James (Perkins and Backlund, 2003), Vermillion, and Missouri River (Backlund, 2000). It is in North Dakota in the James (live) and Apple (shells) Rivers (K. Duttenheefner, ND NHP, pers. comm., 2010). In Vermont, it is in Lake Champlain and tributaries (Fichtel and Smith, 1995); Missisquoi, Lamoille, Winooski, Poultney Rivers, Otter Creek. It was recently (upper Tonawanda Creek) in the Tonawanda basin, New York (Marangelo and Strayer, 2000). In Wisconsin, it is abundant mostly Mississippi and lower Wisconsin Rivers (Mathiak, 1979). In West Virginia, it is in the Upper Ohio/Kanawha (Zeto et al., 1987; Morris and Taylor, 1992). In Mississippi, it is in all drainages except Pascagoula (Jones et al., 2005; Darden et al., 2002). In Louisiana it is in most drainages (Vidrine, 1993). It is in Arkansas in the Ouachita (Posey et al., 1996; Posey, 1997), St. Francis (Ahlstedt and Jenkinson, 1991), Poteau (Vaughn and Spooner, 2004), Cache and White (Christian, 1995; Christian et al., 2005; Gordon, 1982; Gordon et al., 1994); and lower Arkansas (Gordon, 1985). In Texas, it is from the Colorado basin N and E (Howells et al., 1996); incl. Village Creek (Hardin/Tyler/Polk Cos.) (Bordelon and Harrel, 2004). It is throughout Tennessee: Cumberland, Tennessee, Mississippi drainages (Parmalee and Bogan, 1998). In Alabama, it is throughout the Tennessee and Mobile basins (Ahlstedt, 1996; Mirarchi, 2004; McGregor and Garner, 2004; Luxapallila to Mississippi- Johnson and Ahlstedt, 2005; Tombigbee and Alabama- Williams et al., 1992; McGregor et al., 1999), absent from Tallapoosa above the Fall Line and Gulf Coast E of the Mobile Basin (Williams et al., 2008). It is in Kentucky statewide (Cicerello and Schuster, 2003): Middle Green (Gordon, 1991), Barren (Cochran and Layzer, 1993); Black Warrior in Tuscaloosa and Greene/Hale Cos. (Williams et al., 1992). In the Coosa basin, Georgia, it is historical from the Coosa and Conasauga; recently Etowah, Oostanaula, and Coosawattee (Williams and Hughes, 1998). In Kansas, it is widespread in the Kansas, Arkansas, Neosho, Verdigris, and Marais des Cygnes drainages; Spring below Empire Lake (Couch, 1997; Tiemann, 2006). Oklahoma: "Oklahoma City"; Red, Washita, Blue, Clear, Lower Boggy, Kiamichi, Arkansas, Little (Vaughn and Taylor, 1999), Verdigris (Boeckman and Bidwell, 2008), Neosho, Chickaskia, N and Deep Fork Canadian; Neosho and Spring (Branson, 1966); Lake Texoma, Kiamichi rivers; Mountain Fork (Spooner and Vaughn, 2007); Big and Middle Caney; Blue River and Tenkiller Reservoir (Branson, 1984; Vaughn, 2000). In the Little Blue basin it is in Kansas and Nebraska (Hoke, 2004); Big Blue (SE NE and NE KS) in Big and Little Blue Rivers near the border (Hoke, 2005); also Platte River, Nebraska (Sarpy/Hall/Buffalo Cos.) (Freeman and Perkins, 1992). It is in the Lake Michigan, Huron, St. Clair (Badra and Goforth, 2003), and Kalamazoo River, Michigan (Mulcrone and Mehlne, 2001). In Canada, it was historically throughout the lower Great Lakes- St. Lawrence of Ontario (wider than indicated by Clarke, 1981) (Metcalfe-Smith et al., 2003) and Quebec but declined recently (extirpated Lake Erie?) due to zebra mussels (Metcalfe-Smith and Cudmore-Vokey, 2004).

Population Size: >1,000,000 individuals

Number of Occurrences with Good Viability/Integrity: Very many (>125)
Viability/Integrity Comments: The best Ohio populations are in the Little Miami, Scioto, and Muskingum Rivers (Watters et al., 2009).

Short-term Trend: Relatively Stable (<=10% change)
Short-term Trend Comments: Recently, Sietman (2003) reports this species expanding in the Mississippi River above St. Anthony Falls, Minnesota. Som decline has been indicated in the Great Lakes due to zebra mussel invasion (possibly extirpated from Lake Erie) (Metcalfe-Smith and Cudmore-Vokey, 2004). In Tennessee, it occurred in Reelfoot Lake and the Wolf, Nolichucky, and Swquatchie Rivers prior to 1960 (Parmalee and Bogan, 1998).

Long-term Trend: Decline of <30% to increase of 25%
Long-term Trend Comments: What was formerly considered Medionidus mcglameriae was described in 1939 (from specimens collected in 1935) (Van der Schalie, 1939) from the Tombigbee River (Mobile River basin) at Epes in Sumter Co., Alabama (types in MZUM and UMMZ) and is only known from a few specimens (USFWS, 2000; Burch, 1975; Johnson, 1977; Williams et al., 1992). It is now considered a synonym of Leptodea fragilis (Williams et al., 2008).

Intrinsic Vulnerability: Not intrinsically vulnerable

Environmental Specificity: Broad. Generalist or community with all key requirements common.
Environmental Specificity Comments: This species is tolerant of a variety of aquatic habitats.

Other NatureServe Conservation Status Information

Distribution
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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) This species is extremely wide-ranging occurring throughout all of the Mississippi River drainage; Gulf of Mexico drainage from Alabama to Texas (Burch, 1975) and the entire Great Lakes-St. Lawrence system in Canada (Clarke, 1981). Estimated range extent based on centroids of mapped level 8 hydrobasins (2015).

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: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AL, AR, GA, IA, IL, IN, KS, KY, LA, MI, MN, MO, MS, ND, NE, NY, OH, OK, PA, SD, TN, TX, VA, VT, WI, WV
Canada ON, QC

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Baldwin (01003), Bibb (01007), Blount (01009), Clarke (01025), Dallas (01047), Greene (01063), Jackson (01071), Jefferson (01073), Lamar (01075), Limestone (01083), Madison (01089), Marshall (01095), Morgan (01103)*, Perry (01105), Pickens (01107), Sumter (01119), Tuscaloosa (01125), Wilcox (01131)
IA Allamakee (19005), Appanoose (19007), Buena Vista (19021), Carroll (19027), Cedar (19031), Cherokee (19035), Clay (19041), Clayton (19043), Clinton (19045), Des Moines (19057), Dickinson (19059), Dubuque (19061), Greene (19073), Hamilton (19079), Henry (19087), Jackson (19097), Johnson (19103), Lee (19111), Linn (19113), Louisa (19115), Lyon (19119), Muscatine (19139), Sac (19161), Scott (19163), Story (19169), Wapello (19179), Washington (19183), Webster (19187), Woodbury (19193)
NY Clinton (36019), Erie (36029), Franklin (36033), Niagara (36063), Orleans (36073), St. Lawrence (36089), Washington (36115)
OH Coshocton (39031), Franklin (39049), Madison (39097), Montgomery (39113), Paulding (39125), Pickaway (39129), Sandusky (39143), Scioto (39145), Wood (39173)
PA Allegheny (42003), Armstrong (42005), Beaver (42007), Butler (42019), Erie (42049), Warren (42123)*, Washington (42125)*, Westmoreland (42129)
VA Lee (51105)*, Russell (51167), Scott (51169), Wise (51195)*
VT Addison (50001), Chittenden (50007), Franklin (50011), Grand Isle (50013), Rutland (50021)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Cahaba (03150202)+, Middle Alabama (03150203)+, Lower Alabama (03150204)+, Buttahatchee (03160103)+, Sipsey (03160107)+, Locust (03160111)+, Sucarnoochee (03160202)+
04 Upper Maumee (04100005)+, Cedar-Portage (04100010)+, Buffalo-Eighteenmile (04120103)+, Niagara (04120104)+, Lake Erie (04120200)+, Oak Orchard-Twelvemile (04130001)+, Grass (04150304)+, St. Regis (04150306)+, Mettawee River (04150401)+, Otter Creek (04150402)+, Winooski River (04150403)+, Lamoille River (04150405)+, Missiquoi River (04150407)+, Lake Champlain (04150408)+
05 Middle Allegheny-Tionesta (05010003)+*, Middle Allegheny-Redbank (05010006)+, Lower Allegheny (05010009)+, Lower Monongahela (05020005)+*, Upper Ohio (05030101)+, Tuscarawas (05040001)+, Walhonding (05040003)+, Muskingum (05040004)+, Upper Scioto (05060001)+, Lower Scioto (05060002)+, Upper Great Miami (05080001)+, Little Scioto-Tygarts (05090103)+, Ohio Brush-Whiteoak (05090201)+
06 Upper Clinch (06010205)+, Powell (06010206)+*, Wheeler Lake (06030002)+, Lower Elk (06030004)+
07 Coon-Yellow (07060001)+, Grant-Little Maquoketa (07060003)+, Turkey (07060004)+, Apple-Plum (07060005)+, Copperas-Duck (07080101)+, Upper Wapsipinicon (07080102)+, Flint-Henderson (07080104)+, South Skunk (07080105)+, Skunk (07080107)+, Lower Cedar (07080206)+, Lower Iowa (07080209)+, Middle Des Moines (07100004)+, Boone (07100005)+, North Raccoon (07100006)+, Lower Des Moines (07100009)+
10 Rock (10170204)+, Little Sioux (10230003)+, Boyer (10230007)+, Upper Chariton (10280201)+
+ 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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Reproduction Comments: A known glochidial host is the freshwater drum (Aplodinotus grunniens) (Howard, 1913; Wilson, 1916; Howard and Anson, 1922; Cummings et al., 1993). Baker (1928) lists the species as being bradytictic; the reproductive period lasting from the end of August to about mid-July. Sietman et al. (2009) confirmed freshwater drum (Aplodinotus grunniens) as a host species.
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, CREEK, High gradient, Low gradient, MEDIUM RIVER, Moderate gradient, Pool, Riffle
Lacustrine Habitat(s): Deep water, Shallow water
Special Habitat Factors: Benthic
Habitat Comments: This species is tolerant of a variety of aquatic habitats and can be found in small streams in strong current with coarse gravel and sand substrates but also rivers or river-lakes possessing slow current and a firm substrate composed of sand and mud. It can occur at depths of up to 15 or 20 feet but reaches greatest population density at normal water levels of three feet or less in areas such as shallow embayments (Parmalee and Bogan, 1998).
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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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: 05May2009
NatureServe Conservation Status Factors Author: Cordeiro, J.
Element Ecology & Life History Edition Date: 21May2007
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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  • Burch, J.B. 1975. Freshwater unionacean clams (mollusca: pelecypoda) of North America. Malcological Publications. Hamburg, Michigan. 204 pp.

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  • Perkins III, K. and D.C. Backlund. 2003. A survey for winged mapleleaf (Quadrula fragosa) and scaleshell (Leptodea leptodon) in the James River, South Dakota. South Dakota Game, Fish and Parks, Pierre, South Dakota, Report GFP 2003-17. 21 pp.

  • Posey, W.R., III, J.L. Harris, and G.L. Harp. 1996b. An evaluation of the mussel community in the Lower Ouachita River. Report to the Arkansas Game and Fish Commission, Arkansas. 28 pp.

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  • Schanzle, R.W., G.W. Kruse, J.A. Kath, R.A. Klocek, and K.S. Cummings. 2004. The freshwater mussels (Bivalvia: Unionidae) of the Fox River basin, Illinois and Wisconsin. Illinois Natural History Biological Notes, 141: 1-35.

  • Sietman, B.E., K. Bloodsworth, B. Bosman, A. Lager, M. Lyons, M.C. Hove, and S.I. Boyer. 2009. Freshwater drum confirmed as a suitable host for Leptodea, Potamilus, and Truncilla species. Ellipsaria 11(3):18-19.

  • Skadsen, D.R. and K. Perkins III. 2000. Unionid mussels of the Big Sioux River and tributaries: Moody, Minnehaha, Lincoln, and Union Counties, South Dakota. GFP Report 2000-9 to the South Dakota Department of Game, Fish, and Parks, Pierre, South Dakota. 52 pp.

  • Smith, Douglas G. 1985. A study of the distribution of freshwater mussels (mollusca: pelecypoda: Unionda) of the Lake Champlain drainage in northwestern New England. American Midland Naturalist 114: 19-29.

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

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  • Strayer, David L. and K.J. Jirka. 1997. The Pearly Mussels (Bivalva: Unionoidea) of New York State. New York State Museum Memoir 26. The New York State Education Department.

  • Strayer, David L., Kurt J. Jirka, and Kathryn J. Schneider. 1991. Recent collections of freshwater mussels (Bivalvia: Unionidae) from western New York. Walkerana 5(12): 63-72.

  • 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.

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  • Watters, G. Thomas. 1994. An Annotated Bibliography of the Reproduction and Propogation of the Unionoidea (Primarily of North America). Ohio Biological Survey, College of Biological Sciences, The Ohio State University. In cooperation with Ohio Division of Wildlife. 158 pp.

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  • 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.

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  • Posey II, W.R. 1997. Location, species composition and community estimates for mussel beds in the St. Francis and Ouachita Rivers, Arkansas. M.S. Thesis, Arkansas State University. 178 pp.

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