Villosa fabalis - (I. Lea, 1831)
Rayed Bean
Taxonomic Status: Accepted
Related ITIS Name(s): Villosa fabalis (I. Lea, 1831) (TSN 80201)
Unique Identifier: ELEMENT_GLOBAL.2.116088
Element Code: IMBIV47050
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Villosa
Genus Size: C - Small genus (6-20 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: Villosa fabalis
Taxonomic Comments: This species is a member of the widespread genus Villosa but was formerly placed in one or more of the following genera; Unio, Eurynia, Margarita, Margaron, Micromya, and Lemiox (Bogan and Parmalee, 1983). Synonyms include Unio capillus Say, 1831 and Unio lapillus Say, 1832 (Williams et al., 2008). Preliminary analysis indicates that the genus Villosa is not a monophyletic group, based on a molecular phylogenetic analysis and includes as many as six distinct clades (Buhay et al., 2003).
Conservation Status
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NatureServe Status

Global Status: G2
Global Status Last Reviewed: 21May2009
Global Status Last Changed: 31May2006
Rounded Global Status: G2 - Imperiled
Reasons: This species is declining throughout its range to where 78% of streams formerly occupied no longer contain viable populations. Distribution is greatly fragmented and only a small percentage of former populations are known to exist. Remaining rayed bean populations are small and geographically isolated making them susceptible to a single catastrophic event and limiting potential for making natural repopulation or any genetic interchange between disjunct populations. Long-term viability of extant populations is questionable, particularly in the presence of introduced competitors.
Nation: United States
National Status: N2 (09Nov2005)
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 (SX), Illinois (SX), Indiana (S1), Kentucky (SX), Michigan (S1S2), New York (S1), Ohio (S1), Pennsylvania (S1S2), Tennessee (S1), Virginia (SX), West Virginia (S1)
Canada Ontario (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (14Feb2012)
U.S. Fish & Wildlife Service Lead Region: R3 - North Central
Canadian Species at Risk Act (SARA) Schedule 1/Annexe 1 Status: E (05Jun2003)
Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Endangered (25Apr2010)
Comments on COSEWIC: Reason for designation: This freshwater mussel is one of the smallest in Canada. It is found in two rivers in southern Ontario; more than 99% of the estimated total population is found in the Sydenham River. The original COSEWIC assessment (2000) concluded that it had been extirpated from most of its Canadian range and was confined to one river but a new, albeit small, population was discovered in 2004 in the North Thames River. Thirteen live individuals were found between 2004 and 2008 in this river. The main limiting factor is the availability of shallow, silt-free riffle habitat. Both riverine populations are in areas of intense agriculture and urban development, subject to siltation and pollution. Invasive Zebra Mussels have rendered much of the historic habitat unsuitable and pose a continuing threat to one of the last remaining populations.

Status history: Designated Endangered in April 1999. Status re-examined and confirmed in May 2000 and April 2010.

IUCN Red List Category: EN - Endangered
American Fisheries Society Status: Special Concern (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: 20,000-200,000 square km (about 8000-80,000 square miles)
Range Extent Comments: The rayed bean was historically known from 106 streams, lakes, and some man-made canals in 11 states (Alabama, Illinois, Indiana, Kentucky, Michigan, New York, Ohio, Pennsylvania, Tennessee, Virginia, and West Virginia) and 1 Canadian province (Ontario). It is widespread in the Ohio River drainage from headwaters in western New York and Pennsylvania downstream to near the mouth of the Ohio River (Cummings and Mayer, 1992; Strayer and Jirka, 1997; Watters et al., 2009). The southernmost range is Alabama (historical only) (Mirarchi et al., 2004; Williams et al., 2008) and Tennessee (Parmalee and Bogan, 1998). The mussel occurred in parts of the upper (i.e., Lake Michigan drainage) and lower Great Lakes system, and throughout most of the Ohio and Tennessee River systems. It also occurred in western Lake Erie and its tributaries, including the Maumee River and tributaries to Lake St. Clair (Strayer, 1980) and the St. Clair River (Hoeh and Trdan, 1985) with the easternmost records form western New York (Strayer and Fetterman, 1999). In Canada, it is only still extant in the Sydenham and in low numbers the North Thames in Ontario (Cudmore et al., 2004; COSEWIC, 2010).

Area of Occupancy: 2,501 to >12,500 4-km2 grid cells
Area of Occupancy Comments: Canadian EOO is 678 sq. km, AOO 180 sq. km (COSEWIC, 2010).

Number of Occurrences: 21 - 80
Number of Occurrences Comments: Currently, this species is limited to ten streams in the lower Great Lakes system; and twelve streams and one lake in the Ohio River system. Weathered shells as well as a few fresh dead specimens are reported from Tippecanoe River (Cummings and Berlocher, 1990) and Sugar Creek (east fork White River drainage) in central Indiana (Harmon, 1992). A population may also exist in the mainstem of the St. Joseph River in Allen Co., Indiana (Pryor, 2005). In Ohio, living specimens are known from the Stillwater River, Fish Creek, Big and Little Darby Creeks, Scioto Brush Creek, Little Miami River, East Fork Little Miami River, Walhonding River, Blanchard River, and Swan Creek but may be more widely distributed than current range suggests (Watters, 1992; Grabarkiewicz and Crail, 2006; Grabarkiewicz, 2008; Watters et al., 2009). This species is very rare in Muddy Creek (French Creek drainage) in the Erie NWR in Crawford Co., Pennsylvania (Mohler et al., 2006). This species is also known from the Clinton River drainage in Michigan (Strayer, 1980; Trdan and Hoeh, 1993) and recently Pine and Belle Rivers (St. Clair-Detroit drainage) (Badra and Goforth, 2003). In Canada, it is extant in two rivers in Ontario: the middle reach of the Sydenham River (Metcalfe-Smith and Cudmore-Vokey, 2004; West et al., 2000) and in the North Thames River (Cudmore et al., 2004) (COSEWIC, 2010). It was known from the upper Elk River, Tennessee (Isom et al., 1973).

Population Size: >1,000,000 individuals
Population Size Comments: Abundance in mussels is difficult to estimate and no estimates of population size or abundance have been made for this species in the U.S. Hoggarth (1998) reports that at least in the upper Blanchard River, this species is the third most common unionid species found. Smith and Crabtree (2010) found this species at 7 of 32 sites (7 with recruitment) along the entire length of Pennsylvania's French Creek; one of the more abundant species in French Creek with 8.3% relative abundance. Of the 22 records in the Thames and Sydenham Rivers in Canada, 2 are represented by single specimens only at upstream localities. More recent surveys in Canada have revealed higher densities than previously thought. Three Sydenham River populations had >3 individuals per sq. m and were the most abundant species present (total est. population 1,555,000 +- 658,800). The recently discovered North Thames River population (total est. population 4300) is much less dense (COSEWIC, 2010).

Number of Occurrences with Good Viability/Integrity: Few (4-12)
Viability/Integrity Comments: Relatively few streams are thought to harbor long-term viable populations (e.g., Sydenham, Blanchard, Allegheny Rivers; French Creek). The best remaining populations are found in the far northern portions of its range, including the middle reach of the East Sydenham River in Ontario, where it remains fairly abundant (Metcalfe-Smith and Cudmore-Vokey, 2004). Another excellent population is that in Swan Creek (Lower Maumee basin) in Ohio (Grabarciewicz, 2008). The rayed bean appears to be extirpated from the southern unglaciated portion of its range (see USFWS, 2003).

Overall Threat Impact: Very high - high
Overall Threat Impact Comments: The greatest threat to this species is habitat loss or degradation. Smith (1971) ranked the causes of extirpation or declines in fish species and aquatic habitats as follows: siltation, drainage of bottomland lakes, swamps, and prairie marshes, desiccation during drought, species introductions, pollution, impoundments, and increased water temperatures. All of these factors render mussel habitats unsuitable, cause extirpations, and lead to the isolation of populations thereby increasing their vulnerability to extirpation. Dreissenid mussels (Zebra mussel, Dreissena polymorpha, and Quagga mussel, D. rostiformis) have caused dramatic declines in mussel populations in the Great Lakes and significantly reduced mussels in many of the large rivers of eastern North America. Zebra mussels have the potential to severely threaten other populations especially if they make their way into smaller streams. Pollution through point (industrial and residential discharge) and non-point (siltation, herbicide and fertilizer run-off) sources is perhaps the greatest on-going threat to this species and most freshwater mussels. Lowered dissolved oxygen content and elevated ammonia levels (frequently associated with agricultural runoff and sewage discharge) have been shown to be lethal to some species of freshwater naiads (Horne and McIntosh, 1979). Residential, mineral and industrial development also pose a significant threat. Rotenone, a toxin used to kill fish in bodies of water for increased sport fishery quality, has been shown to be lethal to mussels as well (Heard, 1970). Destruction of habitat through stream channelization and maintenance and the construction of dams is still a threat in some areas. Impoundments reduce currents that are necessary for basic physiological activities such as feeding, waste removal and reproduction. In addition, reduced water flow typically results in a reduction in water oxygen levels and a settling out of suspended solids (silt, etc.), both of which are detrimental. Dredging of streams has an immediate effect on existing populations by physically removing and destroying individuals. Dredging also affects the long-term recolonization abilities by destroying much of the potential habitat, making the substrates and flow rates uniform throughout the system. Natural predators include raccoons, otter, mink, muskrats, turtles and some birds (Simpson, 1899; Boepple and Coker, 1912; Evermann and Clark, 1918; Coker et al., 1921; Parmalee, 1967; Snyder and Snyder, 1969). Domestic animals such as hogs can root mussel beds to pieces (Meek and Clark, 1912). Fishes, particularly catfish, Ictalurus spp. and Amierus spp., and freshwater drum, Aplodinotus grunniens, also consume large numbers of unionids. Butler (2003) lists habitat loss and degradation (caused by impoundments, channelization, chemical contaminants, mining, sedimentation) as the chief threat for this species, in particular. The decline of the rayed bean in the Great Lakes drainages and the Ohio and Tennessee River systems and other mussel species in the eastern United States is primarily the result of habitat loss and degradation. These losses have been well documented since the mid-19th century (Higgins, 1858). Chief among the causes of decline are impoundments, channelization, chemical contaminants, mining, and sedimentation (Williams et al., 1993; Neves, 1993; Neves et al., 1997; Watters, 2000). Other threats include: (1) overutilization for commercial, recreational, scientific, or educational purposes, although this species is not a commercially valuable species; (2) disease or predation (little known); (3) inadequacy of existing regulatory mechanisms; (4) geographic isolation and genetic isolation; (5) invasive species (Asiatic clam, zebra mussel, black carp). The threats to the rayed bean are significant and present throughout the species range, and thus are high in magnitude. Threats to the remaining populations are imminent and will likely not lessen in the future (Butler, 2003; U.S. Fish and Wildlife Service, 2003).

Short-term Trend: Decline of 50-70%
Short-term Trend Comments: The species has been eliminated from 78% of the total number of streams and other water bodies from which it was historically known (22 streams and a lake currently compared to 106 water bodies historically). This species has also been eliminated from long reaches of former habitat in hundreds of miles of the Maumee, Ohio, Wabash, and Tennessee Rivers and from numerous stream reaches in their tributaries. In addition, the species is no longer known from the States of Illinois, Kentucky, Tennessee, Virginia, Alabama, and West Virginia, representing over half of the states from which it was formerly known. Canadian populations have always been rare and have been reduced from three rivers in southwestern Ontario to only the middle reach of the East Sydenham River (Metcalfe-Smith and Cudmore-Vokey, 2004) and possibly the North Thames with a single live specimen found in 2004 (Cudmore et al., 2004). More recent surveys in Canada have revealed higher densities than previously thought, however, with three Sydenham River populations having >3 individuals per sq. m (most abundant species present); although the North Thames River population is several orders of magnitude less dense (COSEWIC, 2010). Relatively few streams are thought to harbor longterm viable populations (e.g., Sydenham, Blanchard, Allegheny Rivers; French Creek) (see Butler, 2003; U.S. Fish and Wildlife Service, 2003; COSEWIC, 2010). Specimens were last seen in the Elk River, Tennessee, in the 1870s (Isom et al., 1973; van der Schalie, 1973; Parmalee and Bogan, 1998) and Duck River, Alabama, in 1982, despite reports in many earlier surveys (Ahlstedt et al., 2004) and the species is represented elsewhere in Alabama historically by a single prehistoric record from a Tennessee River shell midden in Jackson Co. (Mirarchi et al., 2004; in appendix 1.2 published separately). Contrary to this, Williams et al. (2008) claims there are no historic records from Alabama but distribution includes upper reaches of the Tennessee River drainage and upper Elk River in Tennessee, and presume it occurred in intervening reaches of northern Alabama. There is one prehistoric record from the Tennessee River near Bridgeport (Warren, 1975).

Long-term Trend: Decline of 50-70%
Long-term Trend Comments: It has been eliminated from 78% of the total number of streams and other water bodies from which it was historically known (22 streams and a lake currently compared to 106 water bodies historically). Historical locations include the following stream systems (with tributaries): upper Great Lakes system (Pigeon River); lower Great Lakes system (Black [Mill Creek], Pine, Belle, Clinton [North Fork Clinton River], Sydenham, South Branch Thames, Detroit, Rouge, Huron, Raisin [Macon Creek], Maumee [St. Joseph River (West Branch St. Joseph; Fish, Cedar Creeks; Feeder Canal to St. Joseph River), Auglaize (Ottawa, Blanchard Rivers)], Sandusky [Tymochtee, Wolf Creeks] Rivers; Lake Erie); Ohio River system (Ohio River [Allegheny River (Chautauqua Lake outlet; Chautauqua Lake; Olean, Cassadaga, Conewango, Oil, French [Cussewago Creek], Crooked Creeks), West Fork, Beaver (Shenango, Mahoning Rivers; Pymatuning Creek) Rivers; Middle Island Creek; Muskingum (Walhonding, Mohican Rivers), Elk, Scioto (Olentangy River; Mill, Alum, Whetstone, Big Walnut [Walnut Creek], Big Darby [Little Darby Creek], Deer, Sugar, Scioto Brush, Cedar Creeks; Buckeye Lake; Ohio and Erie Canal), Little Miami (East Fork Little Miami River), Stillwater, South Fork Licking, North Fork Elkhorn Creek, Eagle Creek, Brashears Creek, Green (Nolin, Barren Rivers), Wabash (Salamonie, Mississinewa, Tippecanoe Rivers [Tippecanoe, Winona Lakes; Lake Maxinkuckee], Vermilion [Salt Fork Vermilion, Middle Fork Vermilion, North Fork Vermilion Rivers], Embarras, Sugar Creek, White [West, East Forks White; Big Blue Rivers; Walnut, Mill, Fall, Sugar Creeks])] Rivers); and Tennessee River system [Tennessee River (Holston [North, South Forks Holston River], Nolichucky [Lick Creek], Clinch [North Fork Clinch, Powell Rivers], Elk [Richland Creek], Duck Rivers)] (USFWS, 2003; Butler, 2003; Ahlstedt et al., 2004). Historically in Canada from the Thames, Sydenham, and Detroit Rivers and western Lake Erie in southwestern Ontario (Metcalfe-Smith and Cudmore-Vokey, 2004; West et al., 2000), but now only known from the Sydenham and Thames Rivers (COSEWIC, 2010). This species has also been eliminated from long reaches of former habitat in hundreds of miles of the Maumee, Ohio, Wabash, and Tennessee Rivers and from numerous stream reaches in their tributaries. In addition, the species is no longer known from the States of Illinois, Kentucky, Tennessee, Virginia, West Virginia, and Alabama, representing over half of the states from which it was formerly known (see U.S. Fish and Wildlife Service, 2003; Butler, 2003; Ahlstedt et al., 2004). In Ohio, it was historically widely distributed in the Scioto and Sandusky Rivers and Bass Islands of Lake Erie (Watters et al., 2009). In West Virginia, it was historically known from the Monongahela River, Twelvepole Creek, Middle Island Creek, and West Fork River (Taylor and Horn, 1983). The only record from Alabama is a prehistoric shell midden along the Tennessee River near Bridgeport (Warren, 1975; Williams et al., 2008).

Intrinsic Vulnerability: Moderately vulnerable to not intrinsically vulnerable.
Intrinsic Vulnerability Comments: Because it is a small species often buried in the sediment, survey efforts may be biased because specimens are overlooked. Dispersal capability is relatively limited as glochidia on fish hosts. Three of the known fish hosts are darters; most darters have low dispersal ability (White in Strayer and Jirka 1997; White et al., 1996; Butler, 2003, COSEWIC, 2010). Extant Canadian populations are isolated from one another and from populations in the U.S. by large distances (40-700 km) and, although genetic isolation is a concern over these distances (Zanatta and Murphy, 2006), there is currently no genetic data on variation between and within populations.

Environmental Specificity: Moderate. Generalist or community with some key requirements scarce.
Environmental Specificity Comments: The decline in the overall range of this species suggests that it is not tolerant of poor water quality. Individuals are sensitive to pollution, siltation, habitat perturbation, inundation, and loss of glochidial hosts (Butler, 2003, however, the species has a broad habitat tolerance in terms of flow and substrate preference (COSEWIC, 2010).

Other NatureServe Conservation Status Information

Inventory Needs: Historical distribution is reasonably well known. Additional survey work in its historical range may be warranted since historical range is so large. Periodic status surveys are needed to monitor changes in the remaining populations of this mussel. Enough survey work completed to date to consider for candidate status. Continue to monitor for zebra mussel invasions into native habitat.

Protection Needs: Few areas are currently protected.

Distribution
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Global Range: (20,000-200,000 square km (about 8000-80,000 square miles)) The rayed bean was historically known from 106 streams, lakes, and some man-made canals in 11 states (Alabama, Illinois, Indiana, Kentucky, Michigan, New York, Ohio, Pennsylvania, Tennessee, Virginia, and West Virginia) and 1 Canadian province (Ontario). It is widespread in the Ohio River drainage from headwaters in western New York and Pennsylvania downstream to near the mouth of the Ohio River (Cummings and Mayer, 1992; Strayer and Jirka, 1997; Watters et al., 2009). The southernmost range is Alabama (historical only) (Mirarchi et al., 2004; Williams et al., 2008) and Tennessee (Parmalee and Bogan, 1998). The mussel occurred in parts of the upper (i.e., Lake Michigan drainage) and lower Great Lakes system, and throughout most of the Ohio and Tennessee River systems. It also occurred in western Lake Erie and its tributaries, including the Maumee River and tributaries to Lake St. Clair (Strayer, 1980) and the St. Clair River (Hoeh and Trdan, 1985) with the easternmost records form western New York (Strayer and Fetterman, 1999). In Canada, it is only still extant in the Sydenham and in low numbers the North Thames in Ontario (Cudmore et al., 2004; COSEWIC, 2010).

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 ALextirpated, ILextirpated, IN, KYextirpated, MI, NY, OH, PA, TN, VAextirpated, WV
Canada ON

Range Map
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U.S. Distribution by County Help
State County Name (FIPS Code)
AL Lauderdale (01077)*
IL Vermilion (17183)*
IN Adams (18001), Allen (18003), Bartholomew (18005), Carroll (18015), Cass (18017), De Kalb (18033), Delaware (18035)*, Fountain (18045)*, Fulton (18049)*, Grant (18053)*, Greene (18055), Hamilton (18057)*, Howard (18067), Huntington (18069), Johnson (18081), Kosciusko (18085), Lagrange (18087)*, Lawrence (18093), Madison (18095)*, Marshall (18099)*, Miami (18103), Owen (18119), Parke (18121)*, Posey (18129)*, Pulaski (18131), Randolph (18135), Shelby (18145)*, Tippecanoe (18157), Vermillion (18165)*, Wabash (18169), Wells (18179)*, White (18181)
KY Boone (21015)*, Campbell (21037)*, Gallatin (21077)*, Grayson (21085)*, Green (21087)*, Hardin (21093)*, Hart (21099)*, Kenton (21117)*, Owen (21187)*, Pendleton (21191)*, Scott (21209)*, Spencer (21215)*, Warren (21227)*
MI Hillsdale (26059), Lenawee (26091)*, Macomb (26099)*, Monroe (26115)*, Oakland (26125), St. Clair (26147), Wayne (26163)
NY Cattaraugus (36009), Chautauqua (36013)
OH Auglaize (39011)*, Brown (39015), Champaign (39021)*, Clermont (39025), Coshocton (39031), Delaware (39041)*, Franklin (39049)*, Hancock (39063), Hardin (39065), Logan (39091), Lucas (39095), Madison (39097)*, Marion (39101)*, Montgomery (39113), Morrow (39117)*, Muskingum (39119)*, Ottawa (39123)*, Pickaway (39129)*, Putnam (39137)*, Ross (39141)*, Scioto (39145), Union (39159), Warren (39165), Williams (39171), Wyandot (39175)
PA Armstrong (42005), Clarion (42031), Crawford (42039), Erie (42049), Forest (42053), Lawrence (42073)*, Mercer (42085), Venango (42121), Warren (42123)
TN Greene (47059)*, Lincoln (47103)*, Marshall (47117)*
VA Scott (51169), Washington (51191)*
WV Clay (54015)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
04 St. Joseph (04050001)+, Pigeon-Wiscoggin (04080103)*, Pine (04080202), St. Clair (04090001)+, Lake St. Clair (04090002)+, Clinton (04090003)+, Detroit (04090004)*, Huron (04090005)+, Ottawa-Stony (04100001)+*, Raisin (04100002)+*, St. Joseph (04100003)+, St. Marys (04100004)+, Upper Maumee (04100005)+, Auglaize (04100007)+, Blanchard (04100008)+, Lower Maumee (04100009)+, Cedar-Portage (04100010)+*, Sandusky (04100011)+, Lake Erie (04120200)
05 Upper Allegheny (05010001)+, Conewango (05010002)+, Middle Allegheny-Tionesta (05010003)+, French (05010004)+, Middle Allegheny-Redbank (05010006)+, West Fork (05020002)*, Shenango (05030102)+, Mahoning (05030103)+*, Beaver (05030104)+*, Little Kanawha (05030203)*, Tuscarawas (05040001)*, Mohican (05040002)+*, Walhonding (05040003)+, Muskingum (05040004)+, Licking (05040006)*, Elk (05050007)+*, Upper Scioto (05060001)+, Lower Scioto (05060002)+, Paint (05060003)*, Upper Great Miami (05080001)+, Twelvepole (05090102)*, Ohio Brush-Whiteoak (05090201)+*, Little Miami (05090202)+, Middle Ohio-Laughery (05090203)+*, Licking (05100101)+*, South Fork Licking (05100102)+*, Lower Kentucky (05100205)+*, Upper Green (05110001)+*, Barren (05110002)+*, Upper Wabash (05120101)+, Salamonie (05120102)+, Mississinewa (05120103)+, Eel (05120104)+, Middle Wabash-Deer (05120105)+, Tippecanoe (05120106)+, Wildcat (05120107)+, Middle Wabash-Little Vermilion (05120108)+*, Vermilion (05120109)+*, Sugar (05120110)+, Middle Wabash-Busseron (05120111)*, Embarras (05120112)*, Lower Wabash (05120113)+*, Upper White (05120201)+*, Lower White (05120202)+, Driftwood (05120204)+, Flatrock-Haw (05120205)*, Upper East Fork White (05120206)+, Muscatatuck (05120207), Lower East Fork White (05120208)+, Salt (05140102)+*, Lower Ohio-Little Pigeon (05140201)*
06 North Fork Holston (06010101)+*, South Fork Holston (06010102)*, Holston (06010104)*, Nolichucky (06010108)+*, Watts Bar Lake (06010201)*, Upper Clinch (06010205)*, Powell (06010206)*, Upper Elk (06030003)+*, Lower Elk (06030004)*, Pickwick Lake (06030005)+*, Upper Duck (06040002)+*, Lower Duck (06040003)*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A very small, solid, freshwater mussel that is dark green in color with numerous wavy lines on the shell.
General Description: The following description of the rayed bean is summarized from Cummings and Mayer (1992), Parmalee and Bogan (1998), and West et al. (2000). The rayed bean is a small mussel usually less than 4.5 cm in length. Shell outline is elongate or ovate in males and elliptical in females, and moderately inflated in both sexes, but more so in females. The valves are thick and solid. The anterior end is rounded in females and bluntly pointed in males. Females apparently are generally smaller than males. Dorsally, the shell margin is straight, while the ventral margin is straight to slightly curved. The beaks are slightly elevated above the hingeline, with sculpture consisting of double loops with some nodules. No posterior ridge is evident. Surface texture is smooth and sub-shiny. Surface color is green, yellowish-green, or brown, with numerous wavy dark green rays of various widths (sometimes obscure in older, blackened specimens). Internally, the left valve has two pseudocardinal teeth that are triangular, relatively heavy, and large, and two short heavy lateral teeth. The right valve has a low triangular pseudocardinal tooth, with possibly smaller secondary teeth anteriorly and posteriorly, and a short, heavy, and somewhat elevated lateral tooth. The color of the nacre (mother-of-pearl) is silvery white or bluish and iridescent posteriorly (see USFWS, 2003).
Diagnostic Characteristics: From Williams et al. (2008):
Length to 40 mm, males often larger than females; moderately thick; males somewhat compressed, females moderately inflated; male outline elliptical, female outline oval; posterior margin narrowly rounded to bluntly pointed; anterior margin rounded; dorsal margin convex; ventral margin straight to slightly convex; posterior ridge low, rounded; posterior slope moderately steep; umbo broad, somewhat compressed, elevated slightly above hinge line, umbo sculpture double-looped ridges, interrupted in middle, nodulous posteriorly; periostracum olive to brown, with numerous thin, wavy, green rays. Pseudocardinal teeth low, triangular, 2 divergent teeth in left valve, 1 tooth in right valve, usually with blade-like accessory denticle anteriorly; lateral teeth short, thick, straight, 2 in left valve, 1 in right valve; inteerdentum short, narrow; umbo cavity shallow, open; nacre white to bluish white.

Villosa fabalis may appear similar to species of Lampsilis, although they can be distinguished by the irregular row of papillae on the mantle edge (Ortmann, 1912) as opposed to the mantle flaps characteristic of Lampsilis. Key characters for distintuishing the rayed bean are its small size, thick valves, unusually heavy teeth for a small mussel, and color pattern (Butler, 2003; U.S. Fish and Wildlife Service, 2003). It resembles Villosa trabalis but is less elongate and has a more rounded posterior margin; also the umbo is broader than V. trabalis and it is less inflated (Williams et al., 2008). It resembles Villosa iris but has a thicker shell and larger more triangular pseudocardinal teeth; also the periostracum of V. iris from Alabama populations is generally more yellow, with less wavy rays than those of V. fabalis (Williams et al., 2008). It may resemble small Villosa vanuxemensis, but V. fabalis is more elongate, with a more pointed posterior margin and thicker shell; also the periostracum of V. fabalis typically has more distinctive rays than V. vanuxemensis, which may lack rays altogether; also shell of V. fabalis is white, whereas that of V. vanuxemensis is purple or salmon (Williams et al., 2008). It may superficially resemble Toxolasma lividum, but periostracum of V. fabalis is typically rayed and T. lividum is not; also male V. fabalis are usually less inflated and shell nacre is white instead of purple; live V. fabalis has papillate mantle folds just ventral to the incurrent aperture and the folds are replaced by caruncles in T. lividum (Williams et al., 2008).

Reproduction Comments: Villosa fabalis is reported to be a long-term breeder in that it holds glochidia overwinter for spring release (Ortmann, 1909). Typically gravid females occur in July and August (Ortmann, 1909) although gravid females have also been collected during mid to late May (Ortmann, 1919). Females display papillate mantle folds at certain times to attract potential fish hosts (Williams et al., 2008). The glochidial fish hosts have not been studied, however they include the Tippecanoe darter (Etheostoma tippecanoe) (White in Strayer and Jirka 1997; White et al., 1996; Butler, 2003). Additional hosts were found for Canadian populations including rainbow darter (Etheostoma caeruleum), greenside darter (Etheostoma blennioides), mottled sculpin (Cottus bairdii), and largemouth bass (Micropterus salmoides) (Woolnough, 2002).
Ecology Comments: Mussel literature consists primarily of taxonomy, systematics, distribution, and life history of the taxa. Villosa fabalis appears to be a species negatively affected by eutrophication, siltation, and invasive species. The species has a broad habitat tolerance in terms of flow and substrate preference, but a limited host range with host species that have low dispersal abilities (COSEWIC, 2010). Refer to the General Freshwater Mussel ESA for general ecology of mussels.
Habitat Type: Freshwater
Non-Migrant: Y
Locally Migrant: N
Long Distance Migrant: N
Mobility and Migration Comments: Adults are essentially sessile. Passive movement downstream may occur. Dispersal is accomplished by glochidia encysted on host fish (Williams et al., 2008). Some freshwater mussel species have developed elaborate methods to increase chances of glochidial contact with potential host fish. Some species package glochidia into bundles resembling food items called conglutinates (Ortmann, 1912; Lefevre and Curtis, 1912; Utterback, 1931; Morrison, 1973; Haag and Straton, 2003; Haag and Warren, 2003). Conglutinates are believed to aid in delivering glochidia to potential host fishes. Long-term brooders have evolved modifications to the posterioventral mantle margin that are utilized to lure potential host fish and often resemble food items (e.g. fishes, crayfish, insect larvae, worms) for the hosts (Kraemer, 1970; Haag et al., 1999). Females of Villosa fabalis display papillate mantle folds at certain times to attract potential fish hosts (Williams et al., 2008).
Riverine Habitat(s): CREEK, MEDIUM RIVER, Riffle
Lacustrine Habitat(s): Shallow water
Special Habitat Factors: Benthic
Habitat Comments: From USFWS (2003):
"The following habitat requirements are generally summarized from Watters (1988), Parmalee and Bogan (1998), and West et al. (2000). The rayed bean is generally known from smaller headwater creeks, but records exist in larger rivers. They are usually found in or near shoal or riffle areas, and in the shallow wave-washed areas of glacial lakes, including Lake Erie. In Lake Erie, it is generally associated with islands in the western portion of the lake. Substrates typically include gravel and sand. It is oftentimes associated with vegetation (e.g., water willow, Justicia americana; water milfoil, Myriophyllum sp.) in and adjacent to riffles and shoals. Specimens are typically buried among the roots of the vegetation (see Butler, 2003; Ortmann, 1919; La Rocque, 1967; U.S. Fish and Wildlife Service, 2003)."

Williams et al. (2008) note that it occurs primarily in flowing water of small to large streams but may also be found in small or medium rivers and occasionally natural lakes (in areas subject to frequent wave action).

Adult Food Habits: Detritivore
Immature Food Habits: Parasitic
Food Comments: Little is known about food and feeding of freshwater mussels with early information summarized by Fuller (1974). Stomach content analysis indicates freshwater mussels generally feed on mud, desmids, diatoms, rotifers, flagellates, and other unicellular organisms (Lefevre and Curtis, 1910; 1912; Wilson and Clark, 1912; Allen, 1914; 1921; Evermann and Clark, 1918, 1920; Coker et al., 1921; Howard, 1922; Churchill and Lewis, 1924). Glochidia (larval form) of freshwater mussels are typically parasitic on fish. Adult mussels are filter filters. General literature has assumed that plankton constitutes the majority of food. Studies indicate that detritus is the primary energy source (James, 1987).
Adult Phenology: Circadian
Phenology Comments: Freshwater mussel life cycles are very complex. Males release sperm into the water column to be taken in by females via the incurrent aperture and fertilization is internal (Yokley, 1972). Fertilized eggs are brooded in marsupial spaces between the gills to complete development into glochidia (Lefevre and Curtis, 1910; Heard, 1975). Glochidia are discharged from the excurrent siphon and most are obligate parasites on fish or amphibians. While encysted on the host, glochidia transform into juvenile mussels with organs adapted for a free-living existence are partially developed. Mussel species are either short-term brooders, which produce gametes over an extended period, from autumn to the following winter; or long-term brooders, which produce gametes over an extended period, usually from late winter or early spring to early summer (Haggerty et al., 1995; Garner et al., 1999). Most species of freshwater mussels are long-lived with many species living 30 to 70 years (Bauer, 1992).

Villosa fabalis is reported to be a long-term breeder in that it holds glochidia overwinter for spring release (Ortmann, 1909). Villosa fabalis individuals may live for 11 years or more, but only reach 40 mm in length. They grow quickly during the first three years, then abruptly slow with growth rates fairly uniform across populations. Female characteristics become apparent in the shells as early as the second year (Watters et al., 2009).

Length: 4.5 centimeters
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: This species was listed as a federal candidate species for endangered species status in the U.S. (USFWS, 2003). Management strategies: 1) Maintain high quality Villosa fabalis habitat, consisting of riffle areas of streams with good water quality. 2) Monitor and regulate land use in the watershed to prevent siltation of streams. 3) Prevent any modifications to streams such as dredging and impoundment, as well as modifications to the natural fish fauna in areas where the species may occur.
Restoration Potential: Upgrading of water quality undoubtedly will help in the recovery of sites, providing the fish host is still present. Dredging for sand and gravel as well as channel modifications in good mussel habitats should be halted, as well as any management schemes which would alter the natural fish population.
Preserve Selection & Design Considerations: Small streams or tributaries with good water quality would probably make the best preserve areas for the species. Because the species is usually found buried in the substrate, it may not be as dependent on specific minimum flow rates in its habitat as some other mussels. Acquisition of land on either side of a stream would help insure that pollutant/siltation runoff to the stream would be minimal, although the entire watershed would need to be monitored to minimize siltation effects.
Management Requirements: Maintenance of flowing water in riffle areas with suitable water quality. Stream modifications such as dredging and impoundment should be avoided, and no alterations to the natural fish communities should be allowed.

Construction, mining, and agricultural activities in stream watersheds should be closely monitored in order to minimize siltation and acid runoff to streams. Point sources should be closely checked as well to insure compliance with discharge permit regulations (permitting standards may need upgrading in some cases).

Monitoring Requirements: Because Villosa fabalis is most often reported from shallow water habitats, wading and hand picking specimens from the substrate is probably the most effective sampling method. Care must be taken to thoroughly search the substrate, since the species is relatively small and could easily be overlooked.
Management Research Needs: Specific ecological requirements of the mussel (microhabitat preferences, identification of the glochidial fish host), and the effects of particular pollutants need to be determined.
Biological Research Needs: In order to effectively manage mussel species it is necessary to work out certain life history characteristics first. However the small population size of this species may hinder its study. Because of their unusual life-cycle and dependence on fish for completion of that cycle the host species for the rayed bean should be ascertained if possible. Life history studies need to be done to identify age and size at sexual maturity, recruitment success, age class structure, and other important life history parameters. Research is needed to assess the success of watershed protection on mussel populations. Abundance and distribution of selected species needs to be monitored in order to ascertain how species abundances change over time. From that we can assess what land-use changes, conservation practices, and physical/chemical parameters are correlated with, and possibly responsible for, the biological changes. Propogation technology should be developed. Little known about life history.
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: 21May2009
NatureServe Conservation Status Factors Author: Cordeiro, J. (2009); Cummings K. (2000); Lauritsen, D. (1998)
Management Information Edition Date: 16Apr1987
Management Information Edition Author: Lauritsen, Diane
Element Ecology & Life History Edition Date: 11Dec2007
Element Ecology & Life History Author(s): Cordeiro, J.

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

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

  • Ahlstedt, S.A., J.R. Powell, R.S. Butler, M.T. Fagg, D.W. Hubbs, S.F. Novak, S.R. Palmer, and P.D. Johnson. 2004. Historical and current examination of freshwater mussels (Bivalvia: Margaritiferidae, Unionidae) in the Duck River basin of Tennessee. Final report submitted to the Tennessee Wildlife Resources Agency, contract FA-02-14725-00, Tennessee. 212 pp.

  • Allen, W.R. 1914. The food and feeding habits of freshwater mussels. Biological Bulletin 27(2):127-147.

  • Allen, W.R. 1921. Studies of the biology of freshwater mussels. Experimental studies of the food relations of certain Unionidae. Biological Bulletin 40(4):21-241.

  • Bauer, G. 1992. Variation in the life span and size of the fresh water pearl mussel. Journal of Animal Ecology 61:425-436.

  • Boepple, J.F. and R.E. Coker. 1912. Mussel resources of the Holston and Clinch rivers of eastern Tennessee. Bureau of Fisheries Document 765. 13 pp.

  • Bogan, A.E. and P.W. Parmalee. 1983. Tennessee's rare wildlife. Vol. 2: The mollusks. Tennessee Wildlife Resources Agency and the Tennessee Conservation Department: Nashville, Tennessee. 123 pp.

  • Buhay, J.E. and W.R. Haag. 2003. Molecular systematics of the freshwater mussel genus Villosa (Bivalvia: Unionidae). Third Biennial Symposium, Freshwater Mollusk Conservation Society, 16-19 March 2003, Meeting Program and Abstracts, pages 15-16.

  • Burch, J.B. 1975. Freshwater unionacean clams (mollusca: pelecypoda) of North America. Malcological Publications. Hamburg, Michigan. 204 pp.

  • Burch, J.B. 1975c. Freshwater Unionacean Clams (Mollusca: Pelecypoda) of North America: Biota of Freshwater Ecosystems, Identification Manual No 11. Environmental Protection Agency, Washington, D.C. 176 pp.

  • Butler, R.S. 2003a. Status assessment for the rayed bean, Villosa fabalis, occurring in the Mississippi River and Great Lakes systems. Unpublished report prepared by the Ohio River Valley Ecosystem Team Mollusk Subgroup, Asheville, North Carolina, March 2003. 65 pp.

  • COSEWIC. 2010. COSEWIC assessment and status report on the rayed bean Villosa fabalis in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, Canada. vii + 40 pp.

  • Churchill, E.P., Jr. and S.I. Lewis. 1924. Food and feeding in fresh-water mussels. Bulletin of the U.S. Bureau of Fisheries 39(1923-1924):439-471.

  • 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.

  • Clarke, A.H. 1981a. The Freshwater Molluscs of Canada. National Museum of Natural Sciences, National Museums of Canada, D.W. Friesen and Sons, Ltd.: Ottawa, Canada. 446 pp.

  • Clarke, A.H. 1981a. The freshwater mollusks of Canada. National Museum of Natural Sciences, National Museums of Canada, D. W. Friesen and Sons, Ltd.: Ottawa, Canada. 446 pp.

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

  • 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 5, Illinois. 194 pp.

  • Cummings, K.S. and J.M. Berlocher. 1990. The naiades or freshwater mussels (Bivalvia: Unionidae) of the Tippecanoe River, Indiana. Malacological Review 23:83-98.

  • 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.

  • Dennis, S.D. 1984. Distributional analysis of the freshwater mussel fauna of the Tennessee River system, with special reference to possible limiting effects of siltation. Ph.D. Thesis. Virginia Polytechnic Institute and State University, Blacksburg, Virginia. 247 pp.

  • Dextrase, A.J. 1999. COSSARO Candidate V, T, E Species Evaluation Form: Rayed Bean (Villosa fabalis). Natural Heritage Section, Ontario Ministry of Natural Resources. 6 January, 3 pp.

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

  • Evermann, B.W. and H.W. Clark. 1920. Lake Maxinkukee, a physical and biological survey. Indiana Department of Conservation, Indianapolis, Indiana. 512 pp.

  • Fuller, S.L.H. 1974. Chapter 8: Clams and mussels (Mollusca: Bivalvia). Pages 215-273 in: C.W. Hart, Jr. and S.L.H. Fuller (eds.) Pollution Ecology of Freshwater Invertebrates. Academic Press: New York. 389 pp.

  • Garner, J.T., T.M. Haggerty, and R.F. Modlin. 1999. Reproductive cycle of Quadrula metanevra (Bivalvia: Unionidae) in the Pickwick Dam tailwater of the Tennessee River. American Midland Naturalist 141:277-283.

  • Haag, W.R. and J.L. Staton. 2003. Variation in fecundity and other reproductive traits in freshwater mussels. Freshwater Biology 48:2118-2130.

  • Haag, W.R. and M.L. Warren, Jr. 2003. Host fishes and infection strategies of freshwater mussels in large Mobile Basin streams, USA. Journal of the North American Benthological Society 22(1):78-91.

  • Haag, W.R., M.L. Warren Jr., and M. Shillingsford. 1999. Host fishes and host-attracting behavior or Lampsilis altilis and Villosa vibex (Bivalvia: Unionidae). American Midland Naturalist 141:149-157.

  • Haggerty, T.M., J.T. Garner, G.H. Patterson, and L.C. Jones, Jr. 1995. A quantitative assessment of the reproductive biology of Cyclonaias tuberculata (Bivalvia: Unionidae). Canadian Journal of Zoology 73:83-88.

  • Harmon, J.L. 1992. Naiades (Bivalvia: Unionidae) of Sugar Creek, east fork White River drainage, in central Indiana. Malacology Data Net 3(1-4):31-42.

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

  • Heard, W.H. 1975. Sexuality and other aspects of reproduction in Anodonta (Pelecypoda: Unionidae). Malacologia 15(1):81-103.

  • Higgins, F. 1858. A catalogue of the shell-bearing species, inhabiting the vicinity of Columbus, Ohio, with some remarks thereon. 12th Annual Report, Ohio State Board of Agriculture for 1857: 548-555.

  • Hoggarth, M.A., D.L. Rice, and D.M. Lee. 1995. Discovery of the federally endangered freshwater mussel, Epioblasma obliquata obliquata (Rafinesque, 1820) (Unionidae), in Ohio. Ohio Journal of Science 95(4):298-299.

  • Hoggarth, M.A., D.L. Rice, and T.L. Grove. 1998. The correlation of mussels with fish in the upper Bloanchard River in Hardin and Hancock Counties, Ohio, with special regard to the rayed bean (Villosa fabalis). [Poster presentation]. Triannual Unionid Report 14:23.

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

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

  • Howard, A.D. 1922. Experiments in the culture of freshwater mussels. Bulletin of the U.S. Bureau of Fisheries 38(1924):63-89.

  • James, M.R. 1987. Ecology of the freshwater mussel Hyridella menziesi in a small oliogotrophic lake. Archives of Hydrobiology 108:337-348.

  • Kraemer, L.R. 1970. The mantle flap in three species of Lampsilis (Pelecypoda: Unionidae). Malacologia 10:255-282.

  • LaRocque, A. 1966-70. Pleistocene Mollusca of Ohio. Bureau of the Geological Survey of Ohio, 62(1-4): 113-356.

  • Lefevre, G. and W.C. Curtis. 1910. Reproduction and parasitism in the Unionidae. Journal of Experimental Zoology 9(1):79-115 + 5 plates.

  • Lefevre, G. and W.T. Curtis. 1912. Studies on the reproduction and artificial propogation of fresh-water mussels. Bulletin of the Bureau of Fisheries 30:102-201.

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

  • Mirarchi, R.E., 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.

  • Mohler, J.W., P. Morrison, and J. Haas. 2006. The mussels of Muddy Creek on Erie National Wildlife Refuge. Northeastern Naturalist 13(4):569-582.

  • Morrison, J.P.E. 1973. The families of pearly fresh- water mussels. Bulletin of the American Malacological Union, Inc., March, 1973:45-46.

  • Moyle, P. and J. Bacon. 1969. Distribution and abundance of molluscs in a fresh water environment. Journal of the Minnesota Academy of Science 35(2/3):82-85.

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

  • Neves, R.J. 1993. A state-of-the unionid address. Pages 1-10 in K.S. Cummings, A.C. Buchanan, and L.M. Koch (eds.) Conservation and management of freshwater mussels. Proceedings of a UMRCC symposium, October 1992, St. Louis, Missouri. Upper Mississippi River Conservation Committee, Rock Island, Illinois.

  • Neves, R.J., A.E. Bogan, J.D. Williams, S.A. Ahlstedt, and P.W. Hartfield. 1997. Status of aquatic mollusks in the southeastern United States: a downward spiral of diversity. Pages 43-85 in G.W. Benz and D.E. Collins (eds.) Aquatic Fauna in Peril: the Southeastern Perspective. Special Publication 1, Southeast Aquatic Research Institute, Chattanooga, Tennessee.

  • Ortmann, A.E. 1909a. The destruction of the fresh-water fauna in western Pennsylvania. Proceedings of the American Philosophical Society 48(191):90-110.

  • Ortmann, A.E. 1909b. The breeding season of Unionidae in Pennsylvania. The Nautilus 22:91-95, 99-103.

  • Ortmann, A.E. 1912. Notes upon the families and genera of the najades. Annals of the Carnegie Museum 8(2):222-365.

  • Ortmann, A.E. 1919. A monograph on the Naiades of Pennsylvania. Memoirs of the Carnegie Museum. 8:1-384.

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

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

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

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

  • Pryor, W.W. 2005. Distribution of the native freshwater mussels in the rivers of Allen County, Indiana. Report to the St. Joseph River Watershed Initiative, Fort Wayne, Indiana. 71 pp.

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

  • Simpson, C.T. 1914. A Descriptive Catalogue of the Naiades or Pearly Fresh-water Mussels. Bryant Walker: Detroit, Michigan. 1540 pp.

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

  • Smith, T.A. and D. Crabtree. 2010. Freshwater mussel (Unionidae: Bivalvia) distributions and densities in French Creek, Pennsylvania. Northeastern Naturalist 17(3):387-414.

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

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

  • Stansbery, D.H. 1976b. Naiad mollusks. Pages 42-52 in H. Boschung (ed.). Endangered and threatened plants and animals of Alabama. Bulletin of the Alabama Museum of Natural History 2:1-92.

  • Strayer, D. 1980. The freshwater mussels (Bivalvia: Unionidae) of the Clinton River, Michigan, with comments on man's impact on the fauna, 1870-1978. The Nautilus 94(4):142-149.

  • 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.

  • Strayer, David L. 1995. Some collections of freshwater mussels from Schoharie Creek, Tonawanda Creek, and the Allegheny basin in New York in 1994. Unpublished report. New York Natural Heritage Program, New York State Department of Environmental Conservation. Latham, NY.

  • 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.

  • Taylor, R.W. and K.J. Horn. 1983. A list of freshwater mussels suggested for designation as rare, endangereed or threatened in West Virginia. Proceedings of the West Virginia Academy of Science (Biology Section) 54:31-34.

  • Trdan, R.J. and W.R. Hoeh. 1993. Relocation of two state-listed freshwater mussel species (Epioblasma torulosa rangiana and Epioblasma triquetra) in Michigan. Pages 100-105 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.

  • Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. 2nd Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp.

  • U.S. Fish and Wildlife Service (USFWS). 2010. Endangered and threatened wildlife and plants; listing the rayed bean and snuffbox as endangered; proposed rule. Federal Register 75(211):67552-67583.

  • Utterback, W.I. 1931. Sex behavior among naiades. West Virginia Academy of Science 5:43-45.

  • 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. 1975. Prehistoric uionacean (freshwater mussel) utilization at the Widows Creek Site (1JA305), northeast Alabama. MA Thesis, University of Nebraska, Department of Anthropology, Lincoln, Nebraska. 245 pp.

  • Watters, G. Thomas. 1996. 1996 Survey of the Mussels of the Fish Creek Drainage. Final Report to the Indiana Chapter of The Nature Conservancy.

  • Watters, G.T. 1988. A survey of the freshwater mussels of the St. Joseph River system, with emphasis on the federally endangered white cat's paw pearly mussel. Unpublished report, Indiana Department of Natural Resources, West Lafayette, Indiana. 127 pp.

  • 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. 2000. Freshwater mussels and water quality: a review of the effects of hydrologic and instream habitat alterations. Pages 261-274 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.

  • West, E., J. L. Metcalfe-Smith, and S. K. Staton. 1998. Status of the Rayed Bean, Villosa fabilis, in Canada. DRAFT COSEWIC Status report prepared by Aquatic Ecosystem Protection Branch, National Water Research Institute Environment Canada, Burlington, ON. 29 pp + maps and appendices.

  • West, E.L., J.L. Metcalf-Smith, and S.K. Staton. 2000. Status of the Rayed Bean, Villosa fabalis (Bivalvia: Unionidae), in Ontario and Canada. Canadian Field-Naturalist 114 (2): 248-258.

  • West, E.L., J.L. Metcalfe-Smith, and S.K. Staton. 2000. Status of the rayed bean, Villosa fabalis (Bivalvia: Unionidae), in Ontario and Canada. The Canadian Field-Naturalist 114:248-258.

  • White, L.R., B.A. McPheron, and J.R. Atauffer, Jr. 1996. Molecular genetic identification tools for the unionids of French Creek, Pennsylvania. Malacologia 38(1-2):181-202.

  • 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, and J.T. Garner. 2008. Freshwater Mussels of Alabama & the Mobile Basin in Georgia, Mississippi & Tennessee. University of Alabama Press: Tuscaloosa, Alabama. 908 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., 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.

  • Woolnough, D.A. 2002. Life history of endangered freshwater mussels of the Sydenham River, southwestern Ontario, Canada. MS Thesis, University of Guelph, Ontario. 128 pp.

  • Yokley, P. 1972b. Life history of Pleurobema cordatum (Rafinesque, 1820) (Bivalvia:Unionacea). Malacologia 11:351-364.

  • Zanatta, D.T. and R.W. Murphy. 2006. Evolution of active host-attraction strategies in the freshwater mussel tribe Lampsilini (Bivalvia: Unionidae). Molecular Phylogenetics and Evolution 41:195-208.

References for Watershed Distribution Map
  • 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.

  • Cudmore, B., C.A. MacKinnon, and S.E. Madzia. 2004. Aquatic species at risk in the Thames River watershed, Ontario. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2707. 123 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.

  • Grabarkiewicz, J. and T. Crail. 2006. Freshwater Mussels of the Maumee Drainage. A Compendium and Guide to the Unionids of the Maumee River and Tributaries. Lucas Soil and Water Conservation District, Maumee, Ohio. 61 pp.

  • Grabarkiewicz, J.D. 2008. Three years of unionid surveys in Swan Creek, Lower Maumee River watershed, Lucas Co., OH. Final Report to the Ohio Division of Wildlife, Toledo Naturalists' Association, and Metroparks of the Toledo Area, Toledo, Ohio. 18 pp. + app.

  • Graf, D.L. 2002. Historical biogeography and late glacial origin of the freshwater pearly mussel (Bivalvia: Unionidae) faunas of Lake Erie, North America. Occasional Papers on Mollusks 6(82):175-211.

  • Isom, B.G., P. Yokley, Jr., and C.H. Gooch. 1973. Mussels of Elk River Basin in Alabama and Tennessee- 1965-1967. American Midland Naturalist 89(2):437-442.

  • Master, L. L. 1996. Synoptic national assessment of comparative risks to biological diversity and landscape types: species distributions. Summary Progress Report submitted to Environmental Protection Agency. The Nature Conservancy, Arlington, Virginia. 60 pp.

  • Metcalfe-Smith, J.L. and B. Cudmore-Vokey. 2004. National general status assessment of freshwater mussels (Unionacea). National Water Research Institute / NWRI Contribution No. 04-027. Environment Canada, March 2004. Paginated separately.

  • Metcalfe-Smith, J.L., J. Di Maio, S.K. Staton, and S.R. De Solla. 2003. Status of the freshwater mussel communities of the Sydenham River, Ontario, Canada. American Midland Naturalist 150:37-50.

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

  • U.S. Fish and Wildlife Service (USFWS). 2003. Candidate assessment and listing priority assignment form: Villosa fabalis. U.S. Fish and Wildlife Service, Ohio Field Office, Reynoldsburg, Ohio. 23 pp.

  • Van der Schalie, H. 1973. The mollusks of the Duck River drainage in central Tennessee. Sterkiana 52:45-56.

  • Watters, G.T., M.A. Hoggarth, and D.H. Stansbery. 2009b. The Freshwater Mussels of Ohio. Ohio State University Press: Columbus, Ohio. 421 pp.

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