Obliquaria reflexa - Rafinesque, 1820
Threehorn Wartyback
Taxonomic Status: Accepted
Related ITIS Name(s): Obliquaria reflexa Rafinesque, 1820 (TSN 80164)
French Common Names: obliquaire trois cornes
Unique Identifier: ELEMENT_GLOBAL.2.109254
Element Code: IMBIV30010
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Obliquaria
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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: Obliquaria reflexa
Conservation Status
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NatureServe Status

Global Status: G5
Global Status Last Reviewed: 12Apr2007
Global Status Last Changed: 25Nov1996
Rounded Global Status: G5 - Secure
Reasons: This species occurs throughout most of the Mississippi River drainage from western Pennsylvania north into Michigan and Minnesota, southwest to eastern Kansas, Oklahoma, and Texas; and in the Coosa-Alabama River and Tombigbee River systems in the southeast. Although once recorded from Lake Erie adn its tributaries, recently this species has been confirmed to be likely extirpated from the main channel of the Detroit River between Lake St. Clair and Lake Erie, Michigan/Ontario; due to zebra mussel invasion. Otherwise it is considered stable throughout its range.
Nation: United States
National Status: N5 (16Jul1998)
Nation: Canada
National Status: N1 (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 (S4), Illinois (S4), Indiana (S3), Iowa (S1), Kansas (S3), Kentucky (S4S5), Louisiana (S5), Michigan (S1), Minnesota (SNR), Mississippi (S5), Missouri (S4), Ohio (S2), Oklahoma (S3), Pennsylvania (SH), South Dakota (S1), Tennessee (S5), Texas (S4), West Virginia (S3), Wisconsin (S3)
Canada Ontario (S1)

Other Statuses

Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Threatened (03May2013)
Comments on COSEWIC: This rare species historically occurred in the Great Lakes drainages including Lake St. Clair, western Lake Erie, and the Grand, Thames, and Detroit rivers. The species has not been found since 1992 in Lake St. Clair and the Detroit River and may be extirpated there due largely to the impacts of Zebra and Quagga mussels. It was last recorded from the Canadian side of Lake Erie in 1997. Pollution (sediment loading, nutrient loading, contaminants and toxic substances) related to both urban and agricultural activities represents a high and continuing threat at the three remaining riverine locations.
Designated Threatened in May 2013.

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 occurs throughout most of the Mississippi River drainage from western Pennsylvania north into Michigan and Minnesota, southwest to eastern Kansas, Oklahoma, and Texas; and in the Coosa-Alabama River and Tombigbee River systems in the southeast (Parmalee and bogan, 1998). Although once recorded from Lake Erie and its tributaries (Clarke, 1981), recently this species has been confirmed to be likely extirpated from the main channel of the Detroit River between Lake St. Clair and Lake Erie, Michigan/Ontario; due to zebra mussel invasion (Schloesser et al., 2006).

Number of Occurrences: > 300
Number of Occurrences Comments: In Minnesota, it is widespread and abundant in the Mississippi River below St. Anthony Falls (recently expanded above the falls), and common in the lower St. Croix River; uncommon in the Minnesota River (Sietman, 2003). In Illinois, it is distributed and common in the larger rivers (Cummings and Mayer, 1997; Schanzle and Cummings, 1991); and much of Indiana including Wabash (Fisher, 2006). In Ohio, it is sporadic in southern rivers (Muskingum, Ohio Brush Creek) (Watters, 1992; 1995; Hoggarth et al., 2007; Watters et al., 2009), but may be extirpated from Lake Erie drainages. Recently the James (Perkins and Backlund, 2003) and Big Sioux River, South Dakota (Backlund, 2000). Oklahoma: Blue, Muddy Boggy, Kiamichi, Little (Vaughn and Taylor, 1999), Verdigris (Boeckman and Bidwell, 2008), Neosho, North Fork Canadian, Poteau Rivers; large creeks of the Arkansas drainage (not Chickaskia drainage); Neosho, Poteau, Lake Texoma, Big Caney (Washington Co.), Salt (Osage Co.); Washita, Blue, Kiamichi, Little and Glover Rivers and Pennington Creek; Tenkiller Ferry Reservoir (Illinois River) (Branson, 1984; Vaughn, 2000). In Kansas, it is in the Marais des Cygnes, Fall, Elk, Verdigris, Cottonwood, and Neosho Rivers but is likely extirpated from the Walnut and Spring Rivers (Couch, 1997). In Texas, it occurs from the Trinity drainage basin into systems north and east (Howells et al., 1996) incl. Village Creek drainage of Hardin, Tyler, and Polk Cos. in SE Texas in 2001-2002 (Bordelon and Harrel, 2004). It is threatened in Ohio and was recently found in Ohio Brush Creek (Matter et al., 2006). Although formerly thought to be extirpated in Pennsylvania (Bogan, 1993), it may still occur in the Upper Ohio basin (PA NHP, pers. comm., 2007) where it was widespread incl. Lower Monongahela basin (Ortmann, 1919). In West Virginia, it occurs in the Upper Ohio/Kanawha (Zeto et al., 1987). In Mississippi, it occurs in the Mississippi River North, Big Black, Yazoo, Tennessee, Pearl, and Tombigbee drainages (Jones et al., 2005). In Louisiana, it is widespread in the Pearl, Tqngipahoa, and Amite Rivers (Brown and Banks, 2001); upper Mississippi, Tensas, Boeuf, Bayou Bartholomew, Ouachita, Black, Little, Caddo, Bayou Pierre, Bayou Teche, Calcasieu, and Sabine and Neches Rivers (Vidrine, 1993). It occurs in Arkansas in the Ouachita (Posey et al., 1996; Posey, 1997), Poteau (Vaughn and Spooner, 2004), St. Francis (Ahlstedt and Jenkinson, 1991), Cache and White Rivers (Christian, 1995; Christian et al., 2005; Gordon, 1982; Gordon et al., 1994); and lower Arkansas (Gordon, 1985). In Tennessee, it is throughout the Cumberland and Tennessee Rivers and major tributaries and is relict in the Hatchie River in west Tennessee. It has expanded its range upstream in the Cumberland and Tennessee Rivers (in reservoirs) (Parmalee and Bogan, 1998). McGregor and Garner (2004) listed it in the Bear Creek drainage in Alabama/Mississippi. In Alabama, it is common throughout the Tennessee River system and Mobile basin (Ahlstedt, 1996; Mirarchi, 2004; Williams et al., 2008; McGregor et al., 1999). It is in Kentucky in the Middle Green and Barren Rivers (Cochran and Layzer, 1993), but is distributed to occasional nearly statewide (Cicerello and Schuster, 2003). In the Coosa River basin in Georgia, it occurs in the Coosa, Etowah, Oostanaula, and Conasauga River drainages (Williams and Hughes, 2001). This species was recently collected from the Black Warrior River in Tuscaloosa and Greene/Hale Cos. and upper Tombigbee River in Sumter and Greene Cos., Alabama (Williams et al., 1992). In Wisconsin, it is known from the Wisconsin and Mississippi Rivers (Mathiak, 1979). It is in Michigan in St. Clair drainage (Badra and Goforth, 2003). In Canada, it is rare with a few specimens in recent years in the Sydenham River, Ontario (Metcalfe-Smith et al., 2003), and historically in the lower Thames and Grand Rivers and western Lake Erie, all Ontario (Metcalfe-Smith and Cudmore-Vokey, 2004).

Population Size: >1,000,000 individuals
Population Size Comments: It was reported to be the fourth most abundant mussel species in Wheeler Reservoir in Alabama in 1991 with an estimated population of 44,590,000 individuals (Ahlstedt and McDonough, 1993).

Number of Occurrences with Good Viability/Integrity: Many to very many (41 to >125)

Short-term Trend: Relatively Stable to increase of <25%
Short-term Trend Comments: Sietman (2003) indicates this species has recently expanded above St. Anthony Falls on the Mississippi River in Minnesota. It has expanded its range upstream in the Cumberland adn Tennessee Rivers (in reservoirs) in Tennessee (Parmalee and Bogan, 1998). Likely this species no longer occurs in the lower Thames and Grand Rivers and western Lake Erie in Ontario (Metcalfe-Smith and Cudmore-Vokey, 2004).

Long-term Trend: Decline of <30% to increase of 25%
Long-term Trend Comments: Skadsen and Perkins (2000) reported subfossil material in the Big Sioux River in Lincoln and Union Cos., South Dakota.

Environmental Specificity: Broad. Generalist or community with all key requirements common.
Environmental Specificity Comments: This species is typical of the large rivers where there is moderately strong current and a stable substrate composed of gravel, sand, and mud. Although found at depths of up to 20 feet, it seems to do well at a depth of no more than four to six feet often in shallow, sand- and mud-bottom river embayments with little or no current. It also occurs in many reservoirs (Parmalee and Bogan, 1998).

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 occurs throughout most of the Mississippi River drainage from western Pennsylvania north into Michigan and Minnesota, southwest to eastern Kansas, Oklahoma, and Texas; and in the Coosa-Alabama River and Tombigbee River systems in the southeast (Parmalee and bogan, 1998). Although once recorded from Lake Erie and its tributaries (Clarke, 1981), recently this species has been confirmed to be likely extirpated from the main channel of the Detroit River between Lake St. Clair and Lake Erie, Michigan/Ontario; due to zebra mussel invasion (Schloesser et al., 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: 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, OH, OK, PA, SD, TN, TX, WI, WV
Canada ON

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Bibb (01007), Blount (01009), Colbert (01033), Dallas (01047), Greene (01063), Jackson (01071), Jefferson (01073), Limestone (01083), Madison (01089), Marshall (01095), Monroe (01099), Morgan (01103), Perry (01105), Pickens (01107), Shelby (01117)*, Tuscaloosa (01125)
IA Allamakee (19005), Clayton (19043), Clinton (19045), Des Moines (19057), Dubuque (19061), Jackson (19097), Johnson (19103), Lee (19111), Louisa (19115), Muscatine (19139), Scott (19163)
MI Allegan (26005)*, Berrien (26021)*, Monroe (26115), Ottawa (26139)*, Saginaw (26145), St. Clair (26147), Van Buren (26159)*, Wayne (26163)*
OH Adams (39001), Athens (39009), Brown (39015)*, Clermont (39025), Defiance (39039), Erie (39043)*, Franklin (39049), Gallia (39053), Hamilton (39061), Henry (39069), Lawrence (39087), Lorain (39093)*, Lucas (39095), Meigs (39105), Morgan (39115), Muskingum (39119), Ottawa (39123), Pickaway (39129), Pike (39131)*, Ross (39141), Sandusky (39143)*, Scioto (39145), Warren (39165), Washington (39167), Wood (39173)
OK Cherokee (40021), LeFlore (40079), Pushmataha (40127)
PA Allegheny (42003)*, Beaver (42007)*, Washington (42125)*, Westmoreland (42129)*
SD Hutchinson (46067), Yankton (46135)
WV Cabell (54011), Jackson (54035), Marshall (54051), Mason (54053), Pleasants (54073), Tyler (54095), Wetzel (54103), Wood (54107)
* 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)+, Sipsey (03160107)+, Locust (03160111)+
04 St. Joseph (04050001)+*, Black-Macatawa (04050002)+*, Kalamazoo (04050003)+*, Lower Grand (04050006)+*, Tittabawassee (04080201)+, Saginaw (04080206)+*, St. Clair (04090001)+, Detroit (04090004)+*, Ottawa-Stony (04100001)+*, Raisin (04100002)+*, Lower Maumee (04100009)+, Cedar-Portage (04100010)+, Sandusky (04100011)+*, Black-Rocky (04110001)+*, Lake Erie (04120200)+
05 Lower Monongahela (05020005)+*, Upper Ohio (05030101)+*, Upper Ohio-Wheeling (05030106)+, Little Muskingum-Middle Island (05030201)+, Upper Ohio-Shade (05030202)+, Hocking (05030204)+, Muskingum (05040004)+, Upper Scioto (05060001)+, Lower Scioto (05060002)+, Paint (05060003)+, Raccoon-Symmes (05090101)+, Little Scioto-Tygarts (05090103)+, Ohio Brush-Whiteoak (05090201)+, Little Miami (05090202)+, Middle Ohio-Laughery (05090203)+*
06 Wheeler Lake (06030002)+, Lower Elk (06030004)+, Bear (06030006)+
07 Coon-Yellow (07060001)+, Grant-Little Maquoketa (07060003)+, Turkey (07060004)+, Apple-Plum (07060005)+, Copperas-Duck (07080101)+, Flint-Henderson (07080104)+, Lower Iowa (07080209)+
10 Lower James (10160011)+
11 Illinois (11110103)+, Kiamichi (11140105)+
+ 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: Barnart and Baird (2000) observed a natural infestation of the glochidia of this species on goldeye (Hiodonta alosoides) with numerous glochidia that had grown while encysted. Other hosts include Ericymba buccata (silverjaw minnow), Luxilus chrysocephalus (common shiner), and Rhynichthys cataractae (longnose dace) (Watters et al., 1998; 1999).
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, 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 typical of the large rivers where there is moderately strong current and a stable substrate composed of gravel, sand, and mud. Although found at depths of up to 20 feet, it seems to do well at a depth of no more than four to six feet often in shallow, sand- and mud-bottom river embayments with little or no current. It also occurs in many reservoirs (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: 07May2009
NatureServe Conservation Status Factors Author: Cordeiro, J.
Element Ecology & Life History Edition Date: 12Apr2007
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. and T.A. McDonough. 1993. Quantitative evaluation of commercial mussel populations in the Tennessee River portion of Wheeler Reservoir, Alabama. Pages 38-49 in K.S. Cummings, A.C. Buchanan, and L.M. Koch (eds.) Conservation and Management of Freshwater Mussels. Proceedings of a UMRCC Symposium, 12-14 October 1992, St. Louis, Missouri.l Upper Mississippi River Conservation Committee, Rock Island, Illinois.

  • Barnhart, M.C. and M.S. Baird. 2000. Fish hosts and culture of mussel species of special concern. Annual Report to U.S. Fish and Wildlife Service, Columbia, Missouri, and Missouri Department of Conservation, Jefferson City, Missouri. 39 pp.

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

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  • COSSARO. 2013. Ontario Species at Risk Evaluation Report for Threehorn Wartyback (Obliquaria reflexa). June 2013 (final). 14pp.

  • Christian, A.D. 1995. Analysis of the commercial mussel beds in the Cache and White Rivers in Arkansas. M.S. Thesis, Arkansas State University. 210 pp.

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  • Cochran, T.G. II and J.B. Layzer. 1993. Effects of commercial harvest on unionid habitat use in the Green and Barren Rivers, Kentucky. Pages 61-65 in K.S. Cummings, A.C. Buchanan, and L.M. Koch (eds.) Conservation and Management of Freshwater Mussels: Proceedings of a UMRCC Symposium, 12-14 October, 1992, St. Louis, Missouri. Upper Mississippi River Conservation Committee, Rock Island, Illinois. 189 pp.

  • Ecological Specialists, Inc. 1996. Unionid Mussel Survey of the Blue River, Indiana. Prepared for The Nature Conservancy. 23 pp.

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

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  • McGregor, S.W., T.E. Shepard, T.D. Richardson, and J.F. Fitzpatrick, Jr. 1999. A survey of the primary tributaries of the Alabama and Lower Tombigbee rivers for freshwater mussels, snails, and crayfish. Geological Survey of Alabama, Circular 196. 29 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.

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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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  • 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.

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  • 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.

  • 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.

  • Vaughn, C.C. 2000. Changes in the mussel fauna of the middle Red River drainage: 1910 - present. Pages 225-232 in R.A. Tankersley, D.I. Warmolts, G.T. Watters, B.J. Armitage, P.D. Johnson, and R.S. Butler (eds.). Freshwater Mollusk Symposia Proceedings. Ohio Biological Survey, Columbus, Ohio. 274 pp.

  • Vaughn, C.C. and C.M. Taylor. 1999. Impoundments and the decline of freshwater mussels: a case study of an extinction gradient. Conservation Biology, 13(4): 912-920.

  • Vaughn, C.C. and D.E. Spooner. 2004. Status of the mussel fauna of the Poteau River and implications for commercial harvest. American Midland Naturalist, 152: 336-346.

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

  • Watters, G.T. 1992b. Distribution of the Unionidae in south central Ohio. Malacology Data Net 3(1-4):56-90.

  • Watters, G.T. 1995a. A field guide to the freshwater mussels of Ohio. revised 3rd edition. Ohio Department of Natural Resources, Division of Wildlife, Columbus, Ohio. 122 pp.

  • Watters, G.T., O'Dee, S.H., Chordas, S., and J. Reiger. 1998c. Potential hosts for Lampsilis reeviana brevicula, Obliquaria reflexa. Triannual Unionid Report 16: 21-22.

  • Watters, G.T., S.W. Chordas, S.H. O'Dee, and J. Reiger. 1999. Host identification studies for six species of Unionidae. Pages 75-76 in Program Guide & Abstract of the First Symposium of the Freshwater Conservation Society, 17-19 March 1999, Chattanooga, Tennessee. 92 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 pages.

  • Williams, J.D. and M.H. Hughes. 1998. Freshwater mussels of selected reaches of the main channel rivers in the Coosa drainage of Georgia. U.S. Geological report to U.S. Army Corps of Engineers, Mobile District, Alabama. 21 pp.

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  • Williams, J.D., S.L.H. Fuller, and R. Gracea. 1992a. Effects of impoundment on freshwater mussels (Mollusca: Bivalvia: Unionidae) in the main channel of the Black Warrior and Tombigbee Rivers in western Alabama. Bulletin of the Alabama Museum of Natural History 13:1-10.

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  • Backlund, D.C. 2000. Summary of current known distribution and status of freshwater mussels (Unionoida) in South Dakota. Central Plains Archaeology, 8(1): 69-77.

  • Badra, P.J. and R.R. Goforth. 2003. Freshwater mussel surveys of Great Lakes tributary rivers in Michigan. Report Number MNFI 2003-15 to the Michigan Department of Environmental Quality, Coastal Zone Management Unit, Lansing, Michigan. 40 pp.

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  • Branson, B.A. 1984. The mussels (Unionacea: Bivalvia) of Oklahoma- Part 3: Lampsilini. Proceedings of the Oklahoma Academy of Science, 64: 20-36.

  • Brown, K.M. and P.D. Banks. 2001. The conservation of unionid mussels in Louisiana rivers: diversity, assemblage composition and substrate use. Aquatic Conservation: Marine and Freshwater Ecosystems, 11(3): 189-198.

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  • Couch, K.J. 1997. An Illustrated Guide to the Unionid Mussels of Kansas. Karen J. Couch. [Printed in Olathe, Kansas]. 124 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.

  • Galbraith, H.S., D.E. Spooner, and C.C. Vaughn. 2008. Status of rare and endangered freshwater mussels in southeastern Oklahoma. The Southwestern Naturalist, 53(1): 45-50.

  • Gordon, M.E. 1982. Mollusca of the White River, Arkansas and Missouri. The Southwestern Naturalist, 27(3): 347-352.

  • Gordon, M.E. 1985. Mollusca of Frog Bayou, Arkansas. The Nautilus, 99(1): 6-9.

  • Gordon, M.E., S.W. Chordas, G.L. Harp. and A.V. Brown. 1994. Aquatic Mollusca of the White River National Wildlife Refuge, Arkansas, U.S.A. Walkerana, 7(17/18): 1-9

  • Hoggarth, M.A., D.A. Kimberly, and B.G. Van Allen. 2007. A study of the mussels (Mollusca: Bivalvia: Unionidae) of Symmes Creek and tributaries in Jackson, Gallia and Lawrence Counties, Ohio. Ohio Journal of Science 107(4):57-62.

  • Howells, R.G., R.W. Neck, and H.D. Murray. 1996. Freshwater Mussels of Texas. Texas Parks and Wildlife Press: Austin, Texas. 218 pp.

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  • 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.

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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 pp.

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