Reginaia ebenus - (Lea, 1831)
Ebonyshell
Synonym(s): Fusconaia ebena (I. Lea, 1831) ;Fusconaia ebenus (I. Lea, 1831)
Taxonomic Status: Accepted
Related ITIS Name(s): Fusconaia ebena (I. Lea, 1831) (TSN 80046)
Unique Identifier: ELEMENT_GLOBAL.2.121112
Element Code: IMBIV17060
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Reginaia
Genus Size: B - Very small genus (2-5 species)
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Concept Reference
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Concept Reference: Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.
Concept Reference Code: B98TUR01EHUS
Name Used in Concept Reference: Fusconaia ebena
Taxonomic Comments: According to Williams et al. (2017), "Several studies showed that the genus Fusconaia as portrayed by Turgeon et al. (1998) was polyphyletic (Lydeard et al. 2000; Serb et al. 2003; Campbell et al. 2005; Campbell and Lydeard 2012a, 2012b; Pfeiffer et al. 2016)." Based on these results, they reassign three species recognized by Turgeon et al. (1998) to other genera: F. succissa to Cyclonaias, F. barnesiana to Pleuronaia, and F. ebenus to Reginaia. Reginaia was described to accommodate F. ebenus and two other species (Campbell and Lydeard 2012a). Williams et al. (2017) follow Watters et al. (2009) in correcting the spelling of the species name ebena to ebenus.
Conservation Status
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NatureServe Status

Global Status: G4G5
Global Status Last Reviewed: 19Dec2011
Global Status Last Changed: 25Nov1996
Rounded Global Status: G4 - Apparently Secure
Reasons: This species is extremely widespread in North America occurring in large rivers throughout most of the central and eastern part of the country. It is considered stable throughout most of its range with minimal threats from overharvest in some areas where it is taken commercially for the cultured pearl industry. An exception is the upper Mississippi River where it has gone from one of the most dominant species to one of the rarest.
Nation: United States
National Status: N4N5 (16Jul1998)

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 (S4S5), Arkansas (S3S4), Georgia (SNR), Illinois (S2), Indiana (S3), Iowa (SX), Kentucky (S4S5), Louisiana (S3), Minnesota (S1), Mississippi (S4), Missouri (S1), Ohio (S1), Oklahoma (S1), Tennessee (S4S5), West Virginia (S1), Wisconsin (S1)

Other Statuses

IUCN Red List Category: LC - Least concern
American Fisheries Society Status: Currently Stable (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: 200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Range Extent Comments: This species is distributed through Ohio in the Ohio River, north to Wisconsin and Minnesota, west to Oklahoma and east Texas, souththrough Alabama and Tennessee and into Mississippi, and northern Louisiana (Parmalee and Bogan, 1998). New York reports are false (Strayer and Jirka, 1997) as are Texas reports (Howells et al., 1996).

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

Number of Occurrences: 81 to >300
Number of Occurrences Comments: There are approximately 120 occurrences reported from Alabama, Illinois, Wisconsin, Minnesota, Ohio, Missouri, Louisiana, West Virginia, and Oklahoma; but the species is common enough in many states that many more occurrences are not tracked by Natural Heritage Programs. The species is likely extirpated from Iowa, likely due to the construction of dams and unregulated harvest (Coker et al., 1921; Coker, 1930). In the Upper Mississippi River only 94 live specimens have been collected since 1975 from 27 sites in the (23 sites in the river proper, 3 sites in the lower St. Croix River, and 1 site in the Wisconsin River) (Kelner and Sietman, 2000). In Minnesota, this species is known only to be extant in the lower St. Croix River (Sietman, 2003) where Kelner and Sietman (2000) note only 3 remaining sites. In Illinois, it is generally distributed in the Ohio and sporadic in the Mississippi and Wabash Rivers (Cummings and Mayer, 1997). In the Wabash River in Indiana, it no longer occurs in the mainstem but is restricted to tributaries (Fisher, 2006). In Wisconsin, Mathiak (1979) reported it to be very rare from the upper Mississippi River in a few sites only. In Ohio, it is extirpated from all but the Ohio River (Watters, 1995; Watters et al., 2009). In Kentucky, it is generally distributed in the Ohio River and the western half of the state (Cicerello and Schuster, 2003). In Mississippi, it occurs in the Mississippi River South, Big Black, Yazoo, Pearl, Pascagoula, and Tombigbee drainages (Jones et al., 2005). In Louisiana, this species has been reported from the Pearl and Amite Rivers (Brown and Banks, 2001), upper Mississippi River, Atchafalaya River and basin, Tensas River, Boeuf River, Bayou Baretholomew, and the Red River (Vidrine, 1993). This species was recently collected in St. Francis (Ahlstedt and Jenkinson, 1991), Ouachita (Posey et al., 1996), Cache and White Rivers, Arkansas (Christian, 1995; Christian et al., 2005; Gordon et al., 1994). In Missouri, it has been found only in the Meramec River, Mississippi River, and Osage River, and in the Little Black River (all only a few valves and all historical) (Oesch, 1995). In Tennessee, it occurs in the Hatchie River in the west and the main Tennessee and Cumberland Rivers (in reservoirs) in middle Tennessee (Parmalee and Bogan, 1998). In the Coosa River basin in Georgia, it is known historically from the Etowah River drainage but has not been collected live recently (Williams and Hughes, 1998). McGregor and Garner (2004) recently documented this species in the Bear Creek drainage in Alabama/Mississippi. 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; McGregor et al., 1999). In Alabama, it is in the Tennessee River drainage and Mobile basin and is common and widespread but uncommon upstream of Muscle Shoals (Williams et al., 2008).

Population Size: >1,000,000 individuals

Number of Occurrences with Good Viability/Integrity: Unknown

Overall Threat Impact Comments: Decline of this species in Wisconsin is attributed to loss of host fish with the construction of impoundments on the Mississippi River (Mathiak, 1979). Similarly, overharvest during the pearl button era, pollution, and dams, which block the migration of its primary host fish, skipjack herring (Alosa chrysochloris), are the reasons for the species' demise in Minnesota (Kelner and Sietman, 2000; Sietman, 2003). Excessive harvesting by commercial shellers has had some detrimental effects on some local populations (Parmalee and Bogan, 1998). In the upper Mississippi River, it was a historically dominant species, representing as much as 75% of pre-historic shell middens and 80% of the late 19th and early 20th Century mussel beds but populations were decimated by unregulated harvest, pollution, and construction of dams to the point where it is now rare and distributed disproportionately among reaches (Kelner and Sietman, 2003).

Short-term Trend: Relatively Stable (<=10% change)
Short-term Trend Comments: This species is considered currently stable with viable populations occurring throughout major stretches of the Cumberland River and middle and lower sections of the Tennessee River (Parmalee and Bogan, 1998). In Alabama, it is in the Tennessee River drainage and Mobile basin and is common (locally abundant in Coastal Plain reaches of large Mobile Basin rivers)and widespread but uncommon upstream of Muscle Shoals but expanding (Williams et al., 2008). This contrasts to its situation in the Coosa River where it has disappeared since the river was impounded.

Long-term Trend: Decline of <30% to relatively stable
Long-term Trend Comments: In Minnesota, this species was historically dominant in the Upper Mississippi River prior to impoundment but is currently only known from the lower St. Croix River (Sietman, 2003). In the upper Mississippi River, it was a historically dominant species, representing as much as 75% of pre-historic shell middens and 80% of the late 19th and early 20th Century mussel beds but populations were decimated by unregulated harvest, pollution, and construction of dams to the point where it is now rare and distributed disproportionately among reaches (only 94 live specimens collected since 1975 from 27 sites in the Upper Mississippi River = 23 sites, lower St. Croix River = 3 sites, and Wisconsin River = 1 site) (Kelner and Sietman, 2000). Warren (1995) confirmed occurrences first cited by Starrett (1971) in the Illinois River near Havana of premodern shells indicating the species once occurred in the Illinois River but no longer does (not since 1912). In Ohio, it is extirpated from the Scioto River where it formerly occurred up to Columbus, the lower Muskingum River, and much of the Ohio River (Watters et al., 2009).

Intrinsic Vulnerability: Not intrinsically vulnerable

Environmental Specificity: Moderate to broad.
Environmental Specificity Comments: This species inhabits large rivers and prefers swift water and stable sandy or gravely shoals (Cummings and Mayer, 1992). Parmalee and Bogan (1998) list is occurring in current at depths of 10 to 15 feet or more. A course sand and gravel substrate provides the most suitable habitat, although this species thrives in rivers composed of sand, silt, and mud.

Other NatureServe Conservation Status Information

Distribution
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Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) This species is distributed through Ohio in the Ohio River, north to Wisconsin and Minnesota, west to Oklahoma and east Texas, souththrough Alabama and Tennessee and into Mississippi, and northern Louisiana (Parmalee and Bogan, 1998). New York reports are false (Strayer and Jirka, 1997) as are Texas reports (Howells et al., 1996).

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, AR, GA, IAextirpated, IL, IN, KY, LA, MN, MO, MS, OH, OK, TN, WI, WV

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Baldwin (01003), Bibb (01007), Clarke (01025), Colbert (01033), Dallas (01047), Greene (01063), Lauderdale (01077), Limestone (01083)*, Monroe (01099), Morgan (01103)*, Perry (01105), Pickens (01107), Sumter (01119)*
IA Allamakee (19005)*, Clinton (19045)*, Muscatine (19139)*
IL Adams (17001), Crawford (17033)*, Gallatin (17059), Hancock (17067)*, Hardin (17069)*, Jersey (17083)*, Lawrence (17101), Madison (17119)*, Massac (17127), Mercer (17131)*, Morgan (17137), Peoria (17143), Pike (17149), Pope (17151)*, Pulaski (17153), Rock Island (17161)*, Scott (17171), Tazewell (17179), Wabash (17185), Wayne (17191), White (17193)
LA Madison (22065), Morehouse (22067), Richland (22083)*, St. Tammany (22103)
MN Carver (27019), Chisago (27025), Dakota (27037), Goodhue (27049), Hennepin (27053), Houston (27055), Ramsey (27123), Scott (27139), Sibley (27143), Wabasha (27157), Washington (27163), Winona (27169)
MO Butler (29023), Clark (29045), Cole (29051), Franklin (29071), Gasconade (29073), Jefferson (29099), Lewis (29111), Lincoln (29113), Maries (29125), Marion (29127), Mississippi (29133), Osage (29151), Pike (29163), Ralls (29173), Ripley (29181), St. Charles (29183), St. Louis (29189), Wayne (29223)
OH Brown (39015), Clermont (39025), Hamilton (39061), Lawrence (39087), Pike (39131)*, Scioto (39145)
OK McCurtain (40089)
WI Adams (55001)*, Buffalo (55011)*, Columbia (55021)*, Crawford (55023), Grant (55043), Iowa (55049), Juneau (55057)*, La Crosse (55063)*, Pierce (55093), Polk (55095), Richland (55103), Sauk (55111)*, St. Croix (55109), Trempealeau (55121)*
WV Cabell (54011), Mason (54053), 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)+, Sucarnoochee (03160202)+*, Lower Pearl. Mississippi (03180004)+
05 Upper Ohio-Shade (05030202)+, Lower Scioto (05060002)+*, Raccoon-Symmes (05090101)+, Little Scioto-Tygarts (05090103)+, Ohio Brush-Whiteoak (05090201)+, Middle Ohio-Laughery (05090203)+, Middle Wabash-Busseron (05120111)+, Lower Wabash (05120113)+, Little Wabash (05120114)+, Lower Ohio-Bay (05140203)+, Lower Ohio (05140206)+
06 Wheeler Lake (06030002)+*, Pickwick Lake (06030005)+
07 Twin Cities (07010206)+, Lower Minnesota (07020012)+, Lower St. Croix (07030005)+, Rush-Vermillion (07040001)+, Cannon (07040002)+*, Buffalo-Whitewater (07040003)+, La Crosse-Pine (07040006)+, Coon-Yellow (07060001)+, Grant-Little Maquoketa (07060003)+, Castle Rock (07070003)+*, Lower Wisconsin (07070005)+, Copperas-Duck (07080101)+*, Flint-Henderson (07080104)+*, Lower Des Moines (07100009)+, Bear-Wyaconda (07110001)+, The Sny (07110004)+, Peruque-Piasa (07110009)+, Lower Illinois-Lake Chautauqua (07130003)+, Lower Illinois (07130011)+, Meramec (07140102)+, Bourbeuse (07140103)+, Big (07140104)+
08 Lower Mississippi-Memphis (08010100)+, Bayou Bartholomew (08040205)+, Boeuf (08050001)+*, Tensas (08050003)+
10 Lower Osage (10290111)+, Lower Gasconade (10290203)+
11 Upper Black (11010007)+, Current (11010008)+, Mountain Fork (11140108)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: a freshwater mussel
Reproduction Comments: The primary glochidial host is Alosa chrysochloris (skipjack herring) (Surber, 1913; Howard, 1914; 1916; Coker et al., 1921; Howard and Anson, 1922) and possibly Micropterus salmoides (largemouth bass), Pomoxis annularis (white crappie), and Pomoxis nigromaculatus (black crappie) (Howard, 1914). Payne and Miller (2000) found that rapid and large spring rises in the lower Ohio River discharge in 1981 and 1990 were immediately followed by rapid and large declines in mussel density for this species. These rises in spring flow woincided with the expected spawning peak of the host fish Alosa chrysochloris. The rapid return to low flow and depositional conditions was appropriately timed to enhance successful settlement of juvenile Fusconaia ebena after their parasitic stage on A. chrysochloris gills.
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, High gradient, MEDIUM RIVER, Moderate gradient
Special Habitat Factors: Benthic
Habitat Comments: This species inhabits large rivers and prefers swift water and stable sandy or gravely shoals (Cummings and Mayer, 1992). Parmalee and Bogan (1998) list is occurring in current at depths of 10 to 15 feet or more. A course sand and gravel substrate provides the most suitable habitat, although this species thrives in rivers composed of sand, silt, and mud.
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: 19Dec2011
NatureServe Conservation Status Factors Author: Cordeiro, J. (2011); Whittaker, J.C. (1994)
Element Ecology & Life History Edition Date: 19Dec2011
Element Ecology & Life History Author(s): Cordeiro, J.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • 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.

  • Coker, R. E. 1919. Fresh-water mussels and mussel industries of the United States. Bulletin of the U.S. Bureau of Fisheries 36:13-89.

  • 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 United States Bureau of Fisheries 37:78-181.

  • Cummings, K. S., and C. A. Mayer. 1992. Field guide to freshwater mussels of the Midwest. Illinois Natural History Survey Manual No. 5. 194 pp.

  • 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. 1992. Field guide to freshwater mussels of the Midwest. Ill. Nat. Hist. Surv. Manual 5. 194pp.

  • Cummings, Kevin S. et al. 1992. Survey of the Freshwater Mussels (Mollusca: Unionidae) of the Wabash River Drainage. Final Report. INHS Center for Biodiversity Tech. Rep. 1992 (1):210 pp.

  • Dawley, C. 1947. Distribution of aquatic mollusks in Minnesota. American Midland Naturalist 38:671-697.

  • 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.

  • 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

  • Howard, A.D. 1914. Experiments in propagation of fresh-water mussels of the Quadrula group. Report of the U.S. Commission of Fisheries for 1913. Appendix 4: 1-52 + 6 plates. [Issued separately as U.S. Bureau of Fisheries Document No. 801].

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

  • Howard, A.D. 1916. An artificial infection with glochidia on the river herring. Transactions of the American Fisheries Society, 46: 93-100.

  • Howard, A.D. and B.J. Anson. 1922. Phases in the parasitism of the Unionidae. Journal of Parasitology, 9(2): 68-82.

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

  • Kelner, D. E., and B. E. Sietman. 2000. Relic populations of the ebony shell, Fusconaia ebena (Bivalvia: Unionidae), in the Upper Mississippi River drainage. Journal of Freshwater Ecology 15(3):371-377.

  • Kelner, D.E. and B.E. Sietman. 2000. Relic populations of the ebony shell, Fusconaia ebena (Bivalvia: Unionidae), in the upper Mississippi River drainage. Journal of Freshwater Ecology, 15(3): 371-377.

  • 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.

  • Mathiak, H.A. 1979. A river survey of the unionid mussels of Wisconsin, 1973-1977. Sand Shell Press: Horicon, Wisconsin. 75 pp.

  • McGregor, S.W. and J.T. Garner. 2004. Changes in the freshwater mussel (Bivalvia: Unionidae) fauna of the Bear Creek system of northwest Alabama and northeast Mississippi. American Malacological Bulletin, 18(1/2): 61-70.

  • 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.

  • Oesch, R.D. 1995. Missouri Naiades. A Guide to the Mussels of Missouri. Second edition. Missouri Department of Conservation: Jefferson City, Missouri. viii + 271 pp.

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

  • Parmalee, P.W. 1967. The fresh-water mussels of Illinois. Ill. State Mus., Popular Sci. Series Vol. VIII. 108pp.

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

  • Payne, B.S. and A.C. Miller. 2000. Recruitment of Fusconaia ebena (Bivalvia: Unionidae) in relation to discharge of the lower Ohio River. American Midland Naturalist 144: 328-341.

  • 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.

  • Sietman, B. E. 2003. Field guide to the freshwater mussels of Minnesota. Minnesota Department of Natural Resources, St. Paul, Minnesota. 144 pp.

  • Starrett, W. C. 1971. A survey of the mussels (Unionacea) of the Illinois River: a polluted stream. Illinois Natural History Survey Bulletin, 30(5): 267-403.

  • 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.

  • Strayer, D.L. and K.J. Jirka. 1997. The Pearly Mussels of New York State. New York State Museum Memoir 26. The University of the State of New York. 113 pp. + figures.

  • Surber, T. 1913. Notes on the natural hosts of fresh-water mussels. Bulletin of the United States Bureau of Fisheries, 32: 101-116.

  • 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.

  • Warren, R.E. 1995. Premodern Pleurobema rubrum (Rafinesque 1820) from the Illinois River. Transactions of the Illinois State Academy of Science 88(1-2): 5-12.

  • Watters, G. T. 1994. An annotated bibliography of the reproduction and propagation of the Unionoidea (Primarily of North America). Ohio Biological Survey Miscellaneous Contributions No. 1, Columbus, Ohio. 158 pp.

  • 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.

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

  • 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.

  • Williams, J. D., A. E. Bogan, R. S. Butler, K. S. Cummings, J. T. Garner, J. L. Harris, N. A. Johnson, and G. T. Watters. 2017. A revised list of the freshwater mussels (Mollusca: Bivalvia: Unionida) of the United States and Canada. Freshwater Mollusk Biology and Conservation 20:33-58.

  • 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., A. E.Bogan, R. S. Butler, K. S.Cummings, J. T. Garner, J. L. Harris, N. A. Johnson, and G. T. Watters. 2017. A revised list of the freshwater mussels (mollusca: bivalvia: unionida) of the United States and Canada. Freshwater Mollusk Biology and Conservation 20:33-58.

  • Williams, J. D., R. S. Butler, G. L. Warren, and N. A. Johnson.  2014a.  Freshwater Mussels of Florida.  University of Alabama Press, Tuscaloosa, Alabama. 498 pp.

  • 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.

  • 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.

References for Watershed Distribution Map
  • Ahlstedt, S.A. and J.J. Jenkinson. 1991. Distribution and abundance of Potamilus capax and other freshwater mussels in the St. Francis River system, Arkansas and Missouri, U.S.A. Walkerana, 5(14): 225-261.

  • 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.

  • Christian, A.D., J.L. Harris, W.R. Posey, J.F. Hockmuth, and G.L. Harp. 2005. Freshwater mussel (Bivalvia: Unionidae) assemblages of the lower Cache River, Arkansas. Southeastern Naturalist, 4(3): 487-512.

  • 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. 1930. Keokuk Dam and the fisheries of the upper Mississippi River. Bulletin of the Bureau of Fisheries, 45: 87-139.

  • 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.

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

  • Jones, R.L., W.T. Slack, and P.D. Hartfield. 2005. The freshwater mussels (Mollusca: Bivalvia: Unionidae) of Mississippi. Southeastern Naturalist, 4(1): 77-92.

  • Oesch, R.D. 1984a. Missouri Naiades: a Guide to the Mussels of Missouri. Jefferson City, Missouri: Conservation Commision of the State of Missouri. 270 pp.

  • Ouachita Ecoregional Assessment Team. 2003. Ouachita Mountains ecoregional assessment. Report to the Arkansas Nature Conservancy, Little Rock, Arkansas. 41 pp. + app.

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

  • Sietman, B.E. 2003. Field Guide to the Freshwater Mussels of Minnesota. Minnesota Department of Natural Resources: St. Paul, Minnesota. 144 pp.

  • Vidrine, M.F. 1993. The Historical Distributions of Freshwater Mussels in Louisiana. Gail Q. Vidrine Collectibles: Eunice, Louisiana. xii + 225 pp. + 20 plates.

  • 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. 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.

  • 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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