Cyprogenia stegaria - (Rafinesque, 1820)
Fanshell
Taxonomic Status: Accepted
Related ITIS Name(s): Cyprogenia irrorata irrorata (TSN 80275) ;Cyprogenia irrorata pusilla Simpson (TSN 80276) ;Cyprogenia irrorata Lea (TSN 80274) ;Cyprogenia stegaria (Rafinesque, 1820) (TSN 80277)
Unique Identifier: ELEMENT_GLOBAL.2.110462
Element Code: IMBIV10020
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Cyprogenia
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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: Cyprogenia stegaria
Taxonomic Comments: No allozymic or other genetic data are available and the phylogenetic relationships of this species or genus to other lampsilines have not been investigated. Formerly placed in the genera Obovaria, and Unio. Often found under the name Cyprogenia irrorata. Other common names include: Eastern fanshell, Ohio fanshell, pimpleback, ringed wartyback. Similar species include: Western fanshell, Cyprogenia aberti, and the pimpleback, Quadrula pustulosa. Genetic analysis indicating C. aberti is not a monophyletic group and may comprise 2 and possibly 5 distinct taxa, one of which includes the federally endangered Cyprogenia stegaria (Serb, 2003; 2006). In Serb's analysis, there are 2 major clades of fanshells, A and B. Cyprogenia from the Black River (White River system) are closely related to C. stegaria from the Clinch River in Tennessee, and form one clade within clade A. Cyprogenia from the upper Arkansas system also form one well-supported clade within clade A. A portion of the Cyprogenia from the Ouachita and St. Francis River systems form the final clade in clade A. Cyprogenia from the St. Francis, White and Ouachita River systems form the two clades within Clade B. Serb (2006) surmises that the best fit scenario to the phylogenetic results obtained is that C. aberti contains cryptic biological diversity including distinct lineages (species) of unionids on separate evolutionary paths including two distinct non-interbreeding entities occurring sympatrically in the Ouachita System with reproductive barriers. She does not go so far as to separate the entities into distinct species yet but suggests other independent data (such as life history traits) are needed to support designation of multiple species. Based on morphology, the above work by Serb, and differences in host specificity and conglutinate morphology, Eckert (2003) and Barnhart and Eckert (2004) believe the type population in the Verdigris River system should retain the name Cyprogenia aberti. Further study of genetics and host requirements of populations in the Black River and populations east of the Mississippi are needed.
Conservation Status
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NatureServe Status

Global Status: G1Q
Global Status Last Reviewed: 28Apr2009
Global Status Last Changed: 30Jan1998
Rounded Global Status: G1 - Critically Imperiled
Reasons: This species is rare throughout its limited range in the Tennessee and Ohio river systems and extremely rare in Virginia. Tremendous declines (>80%) have occurred over the past century and particularly in the past 10 years to the point where very few populations remain that are viable. Most of the remaining fanshell populations are small and all are goegraphically isolated from each other. It is threatened by water quality degradation and habitat loss.
Nation: United States
National Status: N1 (30Jan1998)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S1), Illinois (S1), Indiana (S1), Kentucky (S1), Ohio (S1), Pennsylvania (SH), Tennessee (S1), Virginia (S1), West Virginia (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE, XN: Listed endangered, nonessential experimental population (21Jun1990)
Comments on USESA: The FWS is currently monitoring the following populations of the fanshell:

1) Listing status: Endangered
Population location: Alabama , Illinois , Indiana , Kentucky , Ohio , Tennessee , Virginia , West Virginia

2) Listing status: Experimental Population, Non-Essential
Population location: U.S.A. (TN - specified portions of the French Broad and Holston Rivers; see 17.85(b)(1))
Listing status: Experimental Population, Non-Essential

U.S. Fish & Wildlife Service Lead Region: R4 - Southeast
IUCN Red List Category: CR - Critically endangered
American Fisheries Society Status: Endangered (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: 250-1000 square km (about 100-400 square miles)
Range Extent Comments: This species was historically considered endemic to the eastern highlands east of the Mississippi River. It was historically widely distributed in the Tennessee, Cumberland, and Ohio River systems (Parmalee and Bogan, 1998), although it has become very rare in recent years. In the Ohio drainage it has been recently found in: the deep channel of the Ohio River between Cincinnati and Pittsburgh (Johnson, 1980); the lower Muskingum and Walhonding Rivers, Ohio (Stansbery et al., 1982); the Salt and Licking Rivers, tributaries of the Ohio (Stansbery, pers. comm.); the Green River, Kentucky (USFWS, 1991) the Kanawha River, West Virginia (Stansbery, pers. comm.); the Allegheny River, Pennsylvania (Dennis, 1970); and the lower Clinch River in Scott County (Neves, 1991). It has recently been suggested that Cyprogenia aberti is not a monophyletic group and may comprise 2 and possibly 5 distinct taxa, one of which includes the federally endangered Cyprogenia stegaria (Serb, 2003; 2006) (see Element Management).

Area of Occupancy: 126-2,500 4-km2 grid cells
Area of Occupancy Comments: An estimated 760 km or rivers throughout the U.S. currently contain populations of this species, which represents <10% of its historic range (R. Biggins, USFWS, pers. comm., in Jones and Neves, 2001).

Number of Occurrences: 1 - 5
Number of Occurrences Comments: Only three reproducing populations are known: the upper Green and Licking Rivers in Kentucky (Cicerello and Schuster, 2003) and the Clinch River in Tennessee (USFWS, 1991). The only Ohio River extant population in Ohio is from Muskingum Island in the Ohio River near the confluence of the Ohio and Muskingum Rivers (anonymous, 1996; Watters et al., 2009) at the Belleville dam pool (Watters and Flaute, 2010) and the Markland dam pool in eastern Indiana (Watters and Flaute, 2010), but weathered shells are known from Killbuck Creek (Muskingum tributary) (Watters et al., 2009). A few other apparently non-reproducing populations are known from Indiana (Tippecanoe River- Cummings and Berlocher, 1990; Fisher, 2006; as shell only; a few tributaries of thw Wabash such as lower East Fork White River- Fisher, 2006), West Virginia (upper Kanawha River- Taylor and Horn, 1983), and Illinois (Wabash River in 1984- Cummings and Mayer, 1997). It has been collected in Kentucky in the Middle Green and Barren Rivers (Cochran and Layzer, 1993). Also, a small reproducing population may exist in the Tennessee River below Pickwick Landing Dam (Parmalee and Bogan, 1998). In the Alabama and Mobile basin, it occurred historically across the state of Alabama and the Elk River, it is known to be extant only in Wilson Dam tailwaters with questionable viability (Williams et al., 2008).

Population Size: Unknown
Population Size Comments: Due to problems obtaining a unbiased and complete sample, abundance in mussels is always difficult to estimate, and no estimates of population size or abundance have been made for this species. Jenkinson and Ahlstedt (1988) found only two individuals in 348 quantitative samples taken from the Clinch River.

Number of Occurrences with Good Viability/Integrity: Very few (1-3)
Viability/Integrity Comments: Barr et al. (1994) determined (based on 1981 survey data) that viable populations exist in the Clinch River. The population in the Green River is likely the best of the three remaining reproducing populations. Although the Clinch River population extends over 86 river miles, the fanshell is believed to comprise less than 1% of mussels there (USFWS, 1991).

Overall Threat Impact: Very high - high
Overall Threat Impact Comments: Smith (1971) ranked the causes of extirpation or declines in fish species as follows: siltation, drainage of bottomland lakes, swamps, and prairie marshes, desiccation during drought, species introductions, pollution, impoundments, and increased water temperatures. All of these factors render habitats unsuitable, cause extirpations, and lead to the isolation of populations thereby increasing their vulnerability to extirpation for many aquatic species (including mussels) throughout North America. The loss of many historic populations was likely due to the impacts of impoundments, navigation projects, pollution, and habitat alterations, such as gravel and sand dredging, that directly affected the species and reduced or eliminated its fish host (USFWS, 1991). Zebra mussels, Dreissena polymorpha, have destroyed mussel populations in the Great Lakes and significantly reduced mussels in many of the large rivers of the eastern North America and has the potential to severely threaten and other populations especially if it makes its 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. Destruction of habitat through stream channelization and maintenance and the construction of dams although slowed in recent years is still a threat in some areas. Impoundments reduce currents that are necessary for the most 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. 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). Natural predators include raccoons, otter, mink, muskrats, turtles and some birds, which feed heavily upon freshwater mussels (Simpson, 1899; Boepple and Coker, 1912; Evermann and Clark, 1918; Coker et al., 1921; Parmalee, 1967; Snyder and Snyder, 1969). Domestic animals such as hogs can root mussel beds to pieces (Meek and Clark, 1912). Fishes, particularly catfish, Ictalurus spp. and Amieurus spp., and freshwater drum, Aplodinotus grunniens also consume large numbers of unionids.

The distribution and reproductive capacity of this species has been seriously impacted by the construction of impoundments and navigation facilities, dredging for channel maintenance, sand and gravel mining, and water pollution. The three reproducing populations are threatened by a variety of factors. The Green River has been degraded by runoff from oil and gas exploration and production sites and by alteration of stream flows by an upstream reservoir. Land use practices along the Clinch River have contributed to a decline in water quality and mussel populations. The Clinch River has also experienced some adverse impacts from coal mining, and the river has been subjected to two mussel kills resulting from toxic substance spills from a riverside coal-fired power plant. At least 30 collecting sites on the Clinch River once contained more than 18 different species of freshwater mussels (Ahlstedt, 1984). Now, the mussel abundance in the Clinch River has decreased from an average of 11.64 mussels per square meter in 1979, to 6 mussels per square meter by 1988 (USFWS, 1991). The fanshell has been known from about 86 river miles of the Clinch River; yet, this species comprised less than 1 percent of the mussels collected at 11 sampling sites in 1979 and 1988 (USFWS, 1990).

Short-term Trend: Decline of >70%
Short-term Trend Comments: This species has experienced tremendous declines. Considered to be Endangered by the freshwater mussel subcommittee of the endangered species committee of the American Fisheries Society (Williams et al., 1993). In the Midwest, the fanshell was widely distributed and relatively common in the Wabash and Ohio rivers. Today it is extremely rare with live occurences only in the upper and lower Wabash. An estimated 760 km or rivers throughout the U.S. currently contain populations of this species, which represents <10% of its historic range (R. Biggins, USFWS, pers. comm., in Jones and Neves, 2002). Although once widespread in Alabama in the Tennessee River and Elk River, it is only extant, with questionable viability, in the Wilson Dam tailwaters.

Long-term Trend: Decline of 70-90%
Long-term Trend Comments: Since the turn of the century, this species has undergone a substantial reduction in its range. It was historically widely distributed in the Ohio, Wabash, Cumberland, and Tennessee Rivers and their larger tributaries in Pennsylvania, Ohio, West Virginia, Illinois, Indiana, Kentucky, Tennessee, Alabama, and Virginia (USFWS, 1991). In Ohio, it is now extirpated from the Tuscarawas River, Scioto River, Little Miami River, Great Miami River, and nearly so in the Ohio River (Watters et al., 2009). Although historically reported from the Holston, Powell, Little Tenenssee, and Duck Rivers in Tennessee, it is likely now extirpated there (Parmalee and Bogan, 1998). It is extirpated from Pennsylvania (Bogan, 1993; Spoo, 2008) where it formerly occurred in the Middle Allegheny-Redbank, Upper Ohio, and Lower Allegheny drainages (Ortmann, 1919). It was last seen in Alabama in the tailwaters of Wilson Dam but was historically also known from the Tennessee River across northern Alabama and probably the Elk River (Mirarchi et al., 2004). It is extirpated from the McAlpine dam pool in the Ohio River at Louisville, Kentucky (and Indiana) (Watters and Flaute, 2010).

Intrinsic Vulnerability: Unknown

Environmental Specificity: Moderate. Generalist or community with some key requirements scarce.
Environmental Specificity Comments: Characteristic habitat is medium to large streams (Dennis, 1984). It has been found in river habitats with gravel substrates and a strong current, in both deep and shallow water (Ortmann, 1919; Parmalee, 1967).

Other NatureServe Conservation Status Information

Inventory Needs: Many of the suspected non-reproducing populations need to be investigated. Many of the sites were only vistited once during a routine survey of the river. Detailed site investigations where live fanshells have been found in recent years should be conducted. Periodic surveys of known populations should be done to monitor the status of the remaining populations. An inventory of existing museum records should be compiled to provide information on historical sites and potential new ones.

Distribution
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Global Range: (250-1000 square km (about 100-400 square miles)) This species was historically considered endemic to the eastern highlands east of the Mississippi River. It was historically widely distributed in the Tennessee, Cumberland, and Ohio River systems (Parmalee and Bogan, 1998), although it has become very rare in recent years. In the Ohio drainage it has been recently found in: the deep channel of the Ohio River between Cincinnati and Pittsburgh (Johnson, 1980); the lower Muskingum and Walhonding Rivers, Ohio (Stansbery et al., 1982); the Salt and Licking Rivers, tributaries of the Ohio (Stansbery, pers. comm.); the Green River, Kentucky (USFWS, 1991) the Kanawha River, West Virginia (Stansbery, pers. comm.); the Allegheny River, Pennsylvania (Dennis, 1970); and the lower Clinch River in Scott County (Neves, 1991). It has recently been suggested that Cyprogenia aberti is not a monophyletic group and may comprise 2 and possibly 5 distinct taxa, one of which includes the federally endangered Cyprogenia stegaria (Serb, 2003; 2006) (see Element Management).

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, IL, IN, KY, OH, PA, TN, VA, WV

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Colbert (01033)*, Jackson (01071)*, Lauderdale (01077)*, Madison (01089)*, Marshall (01095)*
IL White (17193)*
IN Bartholomew (18005), Carroll (18015), Cass (18017), Daviess (18027), Dubois (18037), Fountain (18045), Gibson (18051), Greene (18055), Harrison (18061)*, Jackson (18071), Knox (18083), Lawrence (18093), Marion (18097)*, Martin (18101), Monroe (18105), Morgan (18109), Owen (18119), Parke (18121), Pike (18125), Posey (18129)*, Sullivan (18153)*, Tippecanoe (18157), Vermillion (18165), Vigo (18167)*, Wabash (18169), Warren (18171), Washington (18175), White (18181)
KY Allen (21003), Barren (21009), Boone (21015)*, Boyd (21019)*, Bracken (21023), Butler (21031), Campbell (21037), Carter (21043)*, Christian (21047)*, Cumberland (21057)*, Edmonson (21061), Fleming (21069), Green (21087), Greenup (21089), Harrison (21097), Hart (21099), Henderson (21101)*, Henry (21103)*, Jefferson (21111)*, Kenton (21117), Larue (21123), Lawrence (21127)*, Lewis (21135)*, Livingston (21139)*, Lyon (21143)*, Marshall (21157)*, Mason (21161)*, Mercer (21167)*, Monroe (21171)*, Muhlenberg (21177), Nelson (21179), Nicholas (21181), Ohio (21183), Oldham (21185)*, Owen (21187)*, Pendleton (21191), Powell (21197)*, Robertson (21201), Russell (21207)*, Spencer (21215)*, Todd (21219)*, Warren (21227), Wayne (21231)*, Woodford (21239)*
OH Coshocton (39031), Meigs (39105), Morgan (39115), Washington (39167)
PA Allegheny (42003)*, Armstrong (42005)*, Beaver (42007)*, Westmoreland (42129)*
TN Anderson (47001)*, DeKalb (47041)*, Decatur (47039), Hancock (47067), Hardin (47071), Loudon (47105)*, Marion (47115)*, Meigs (47121), Putnam (47141)*, Rhea (47143), Roane (47145)*, Sevier (47155)*, Smith (47159)*, Trousdale (47169)*, Wilson (47189)*
VA Lee (51105)*, Scott (51169)
WV Fayette (54019), Jackson (54035), Wood (54107)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
04 Little Calumet-Galien (04040001)*, St. Joseph (04050001)*, St. Joseph (04100003)*, St. Marys (04100004)*, Upper Maumee (04100005)*, Auglaize (04100007)*
05 Middle Allegheny-Redbank (05010006)+*, Lower Allegheny (05010009)+*, Upper Ohio (05030101)+*, Upper Ohio-Wheeling (05030106)*, Little Muskingum-Middle Island (05030201), Upper Ohio-Shade (05030202)+, Tuscarawas (05040001)*, Walhonding (05040003)+, Muskingum (05040004)+, Upper Kanawha (05050006)+, Upper Scioto (05060001)*, Lower Scioto (05060002)*, Big Sandy (05070204)+*, Upper Great Miami (05080001)*, Whitewater (05080003)*, Little Scioto-Tygarts (05090103)+, Ohio Brush-Whiteoak (05090201)+*, Little Miami (05090202)*, Middle Ohio-Laughery (05090203)+*, Licking (05100101)+, South Fork Licking (05100102)+*, Upper Kentucky (05100204)+*, Lower Kentucky (05100205)+*, Upper Green (05110001)+, Barren (05110002)+, Middle Green (05110003)+, Lower Green (05110005)+*, 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)+, Eel (05120203)+*, Driftwood (05120204)*, Flatrock-Haw (05120205)*, Upper East Fork White (05120206)+, Muscatatuck (05120207)*, Lower East Fork White (05120208)+, Patoka (05120209)*, Upper Cumberland (05130101)*, Upper Cumberland-Lake Cumberland (05130103)+*, Upper Cumberland-Cordell Hull (05130106), Caney (05130108)+*, Lower Cumberland-Old Hickory Lake (05130201)+, Lower Cumberland (05130205)+*, Red (05130206)+*, Silver-Little Kentucky (05140101)+, Salt (05140102)+*, Rolling Fork (05140103)+, Blue-Sinking (05140104)*, Lower Ohio-Little Pigeon (05140201)+*, Highland-Pigeon (05140202)*
06 North Fork Holston (06010101), Holston (06010104)*, Pigeon (06010106)*, Lower French Broad (06010107)+*, Watts Bar Lake (06010201), Lower Little Tennessee (06010204)*, Upper Clinch (06010205)+, Powell (06010206)+*, Lower Clinch (06010207)+*, Middle Tennessee-Chickamauga (06020001)+, Guntersville Lake (06030001)*, Wheeler Lake (06030002)+*, Lower Elk (06030004)*, Pickwick Lake (06030005)+, Lower Tennessee-Beech (06040001)+, Upper Duck (06040002)*, Kentucky Lake (06040005)*, Lower Tennessee (06040006)+*
07 Kankakee (07120001)*, Iroquois (07120002)*, Chicago (07120003)*, Des Plaines (07120004)*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
U.S. Distribution by Watershed (based on multiple information sources) Help
Ecology & Life History
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Basic Description: Rounded in shape with numerous pustules, elevated growth lines, and broken green rays.
General Description: SHELL: Rounded, solid, and moderately inflated. Anterior margin rounded, posterior margin bluntly rounded or truncated. Ventral margin broadly rounded. Umbos not elevated above the hinge line. Beak sculpture, if visible, of a few weak ridges. Growth lines appear as distinct elevated ridges. Numerous pustules usually concentrated in the center, but occasionally covering the entire surface of the shell. Periostracum usually greenish yellow, with a pattern of dark green rays made up of numerous smaller broken lines or dots. Length to 3 inches. Pseudocardinal teeth relatively large and serrated; two in the left valve, one in the right. Lateral teeth roughened, straight to slightly curved, heavy and very short. Interdentum wide. Beak cavity shallow to moderately deep. Nacre white, iridescent posteriorly (Cummings and Mayer, 1992).

ANIMAL: "Color of soft parts whitish. Abdominal sac and mantle suffused with black. Edge of mantle brown with black spots, this mottling extending all around. Marsupium, when charged, red, (according to Sterki) sometimes white. Branchial, anal and supra-anal as usual, the latter two separated by a very short mantle-connection. Branchial with papillae, anal finely crenulated. In front of the branchial, the inner edge of the mantle has a series of fine crenulations which soon disappear, the edge becoming smooth. Palpi normal, posterior margins connected at base only. Gills short and broad, the inner much wider than the outer throughout its whole length. Diaphragm normal, inner lamina of inner gills free from abdominal sac, except at the anterior end. Anterior attachment of gills as usual. Septa and water-tubes in both gills normally developed, the latter moderately wide in the male and the non-marsupial gills of the female. Marsupium formed by a section in the middle of the outer gill: in fact this section is a little more toward the anterior end of the gill. Ovisacs few (three to eight in my specimens; up to eleven reported by other authors; Simpson gives for the genus twenty-three as maximum), hardly different in width from the rest of the water-tubes, that is to say in the longitudinal direction. But, when charged, they swell somewhat in the transverse direction, so as to become subcylindrical. The ovisacs project to an extreme degree beyond the edge of the gill. Although they begin near the base of the gill, and although a considerable part is enclosed between the original laminae of the gill, this part is very small when compared with the prolonged portion. The latter curves backward in a circle, and is rolled up spirally, the spiral forming about one and a half to two turns, but only the posterior ovisacs complete the whole revolution, while the anterior ones stop earlier, the first after completing the circle about once. The distal parts of the spiral wind up in the direction toward the median line of the body, so that in a view from the outside, they are hidden under the outer gill and the first whorl of the marsupium. The ova fill the ovisacs in the shape of closely packed masses, forming distinct and very solid placentae, red n color, rarely white." (Ortmann, 1912 313-314).

Diagnostic Characteristics: A white nacre is characteristic except posteriorly, where it becomes iridescent (Simpson, 1914).
Reproduction Comments: The mussel is reported as a long-term breeder (holds glochidia overwinter for spring release) (Ortmann, 1919), and the glochidial host has been reported to be banded sculpin, Cyprogenia stegaria; greenside darter, Etheostoma blennioides; mottled sculpin, Cottus bairdi; Tennessee snubnose darter, Etheostoma simoterum; banded darter, Etheostoma zonale; Tengerine darter, Percina aurantiaca; blotchside logperch, Percina burtoni; logperch, Percina caprodes; and Roanoke darter, Percina roanoka (Schulz and Marbain, 1998; Jones and Neves, 2001l 2002). Glochidia are contained in red, worm-like conglutinates that resemble oligochaetes. These are about 20 to 80 mm long and are released through the excurrent aperture. Estimated fecundity is 22,357 to 63,459 glochidia/mussel (Jones and Neves, 2001).
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, MEDIUM RIVER
Special Habitat Factors: Benthic
Habitat Comments: Characteristic habitat is medium to large streams (Dennis, 1984). It has been found in river habitats with gravel substrates and a strong current, in both deep and shallow water (Ortmann, 1919; Parmalee, 1967).
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: This species was listed as a U.S. federal endangered species in 1990. A recovery plan has been drafted (USFWS, 1991). Management priorities for CYPROGENIA STEGARIA: 1. Maintain high quality C. STEGARIA habitat. 2. Monitor and regulate land use upstream to prevent siltation of streams. Recovery plan (USFWS, 1991).

A current taxonomic challenge to Cyprogenia aberti was brought about by a phylogenetic analysis indicating C. aberti is not a monophyletic group and may comprise 2 and possibly 5 distinct taxa, one of which includes the federally endangered Cyprogenia stegaria (Serb, 2003; 2006). In Serb's analysis, there are 2 major clades of fanshells, A and B. Cyprogenia from the Black River (White River system) are closely related to C. stegaria from the Clinch River in Tennessee, and form one clade within clade A. Cyprogenia from the upper Arkansas system also form one well-supported clade within clade A. A portion of the Cyprogenia from the Ouachita and St. Francis River systems form the final clade in clade A. Cyprogenia from the St. Francis, White and Ouachita River systems form the two clades within Clade B. Serb (2006) recommends that Cyprogenia aberti from the Arkansas, Black, and White drainages be manaed as separate units for conservation. This may also influence the conservation management plans for Cyprogenia stegaria as its close phylogenetic affinity with the Black River Cyprogenia aberti may affect its federal status as well as the status of the Black River population of C. aberti.

Restoration Potential: Many areas of historical occurrences cannot be restored because of river impoundment. Other former habitats can probably be recovered if water quality is sufficiently upgraded and mussels are reintroduced, provided the appropriate fish host is present.
Preserve Selection & Design Considerations: In terms of species density and definition of appropriate habitat, the occurrences located in the upper Clinch River may be easier to protect. Concerted efforts to maintain water quality (monitor point source dischargers) and prevent siltation (land use in the watershed should be monitored) are required.
Management Requirements: This mussel requires flowing water in either riffle areas or deep water of rivers with suitable water quality. River modifications such as dredging and impoundment should be avoided.

As it is found in larger rivers that have been extensively modified, it would be difficult to manage many occurrences because of conflicting interests. The upper Clinch River, including Pendleton Island, may be the best location to focus management efforts. Water quality should be carefully monitored, upstream impoundments should be prohibited, and wise land use practices in the watershed should be encouraged to minimize siltation resulting from construction and agriculture.

Monitoring Requirements: Historical versus present records indicate that C. STEGARIA has become increasingly more rare. A monitoring program for the species would aid in the development of a management plan, as well as provide basic biological and ecological data. Refer to the General Freshwater Mussel ESA.
Management Research Needs: Specific ecological requirements of the mussel (minimum water flow required, substrate preferences, influences of water temperature and food quality on growth rates, etc.), as well as the effects of particular pollutants should be determined.
Biological Research Needs: In order to effectively manage mussel species it is necessary to work out certain life history characteristics first. Because of their unusual life-cycle and dependence on fish for completion of that cycle, it is imperative that the host species for the fanshell be ascertained. The only reported host was the artificially induced laboratory infection on the goldfish, CASSASIUS AURATUS (Chamberlain, 1934). 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 abundanceOs change over time. From that we can assess what land-use changes, conservation practices, and physical/chemical parameters are correlated with, and possibly responsible for, the biological changes.

Population/Occurrence Delineation
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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: 28Apr2009
NatureServe Conservation Status Factors Author: Cordeiro, J. (2009); Morrison, M.; K.S. Cummings (1998)
Management Information Edition Date: 01Aug1986
Management Information Edition Author: Lauritsen, Diane
Element Ecology & Life History Edition Date: 16Oct2006
Element Ecology & Life History Author(s): Cordeiro, J. (2006); LAURITSEN, D. (1991)

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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References for Watershed Distribution Map
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