Strophitus undulatus - (Say, 1817)
Creeper
Other English Common Names: Squawfoot
Taxonomic Status: Accepted
Related ITIS Name(s): Strophitus undulatus tennesseensis Frierson (TSN 80153) ;Strophitus undulatus undulatus (Say, 1817) (TSN 80152) ;Strophitus undulatus (Say, 1817) (TSN 80151)
French Common Names: strophite nodule
Unique Identifier: ELEMENT_GLOBAL.2.107752
Element Code: IMBIV42030
Informal Taxonomy: Animals, Invertebrates - Mollusks - Freshwater Mussels
 
Kingdom Phylum Class Order Family Genus
Animalia Mollusca Bivalvia Unionoida Unionidae Strophitus
Genus Size: A - Monotypic genus
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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: Strophitus undulatus
Taxonomic Comments: The taxonomic status of Strophitus in western Gulf Coast drainages is unclear as Strecker (1931) reported S. subvexus from eastern Texas but Howells et al. (1996) considered all individuals in the state to be S. undulatus. Meanwhile, Vidrine (1993) suggested that Strophitus from western Louisiana (i.e., those in the Calcasieu and Sabine Rivers) represent an undescribed species similar to S. subvexus. Williams et al. (2008) confined S. subvexus to the Tombigbee and Black Warrior River drainages and S. connasaugensis to eastern drainages of the Mobile Basin, however individuals resembling S. subvexus can sometimes be found in eastern reaches of the Mobile basin and individuals resembling S. connasaugaensis can occasionally be found in the Black Warrior and Tombigbee River drainages.
Conservation Status
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NatureServe Status

Global Status: G5
Global Status Last Reviewed: 01Aug2017
Global Status Last Changed: 25Nov1996
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G5 - Secure
Reasons: The species is extremely wide ranging being one of the few North American mussels to occur on both sides of the Appalachian Mountains. It is distributed throughout the Mississippi River and Great Lakes systems, northern Atlantic Coast drainages, and parts of Canadian Interior Basin. Presumably, it occurred throughout the Tennessee River system historically. In Alabama, it reaches its southern limit in a short reach of Bear Creek, Colbert Co., where it may not be viable, otherwise it is considered stable throughout its wide range.
Nation: United States
National Status: N5 (16Jul1998)
Nation: Canada
National Status: N5 (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 (S1), Arkansas (S4), Colorado (SNR), Connecticut (SU), Delaware (S1), District of Columbia (SNR), Georgia (SNR), Illinois (S4), Indiana (S4), Iowa (S2), Kansas (S2), Kentucky (S4S5), Louisiana (S2), Maine (SNR), Maryland (S2), Massachusetts (S3), Michigan (SNR), Minnesota (SNR), Mississippi (S1), Missouri (S4), Nebraska (SNR), New Hampshire (S3S4), New Jersey (S3), New York (S4), North Carolina (S3), North Dakota (SNR), Ohio (S5), Oklahoma (S3), Pennsylvania (S5), Rhode Island (S1), South Carolina (S2), South Dakota (S3), Tennessee (S5), Texas (S1), Vermont (S3), Virginia (S3S4), West Virginia (S3), Wisconsin (S4)
Canada Manitoba (S5), New Brunswick (SU), Nova Scotia (S1), Ontario (S5), Quebec (S3), Saskatchewan (SU)

Other Statuses

American Fisheries Society Status: Currently Stable (01Jan1993)

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: The creeper is distributed very widely throughout the Ohio and Mississippi Rivers and Great Lakes systems, northern Atlantic Coast drainages, and parts of Canadian Interior Basin as far west as Texas and Saskatchewan (Burch, 1975; Parmalee and Bogan, 1998) and historically into Kansas and Nebraska (Hoke, 2005). Presumably, it occurred throughout the Tennessee River system historically. Colorado reports are in error/unconfirmed (Wu, 1989; Cordeiro, 1999; Clarke et al., 2003).

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

Number of Occurrences: > 300
Number of Occurrences Comments: In Alabama, it is only in Bear Creek (Colbert Co.) (Mirarchi et al., 2004; Williams et al., 2008). Tennessee (statewide): upper Clinch, Powell, Watauga, Little, Elk, Big S Fork Cumberland, Stones, Harpeth to Hatchie and W (Parmalee and Bogan, 1998). In Kentucky, Green River E (Cicerello and Schuster, 2003). In Mississippi: Mississippi River S, Yazoo, and Tennessee basins (Jones et al., 2005). It is in the White (rare) (Gordon, 1982; Christian, 1995), lower Arkansas (Gordon, 1985), Ouachita (Posey et al., 1996; Posey, 1997), and Poteau (Vaughn and Spooner, 2004), Arkansas. It is in all Illinois drainages (Cummings and Mayer, 1997; Schanzle and Cummings, 1991; Sietman et al., 2001; Schanzle et al., 2004; Tiemann et al., 2005). In Indiana: E. Fork White (Harmon, 1992), Muscatatuck (Harmon, 1989); Tippecanoe (Cummings and Berlocher, 1990). It is throughout Ohio (Watters, 1995; Lyons et al., 2007; Grabarciewicz, 2008; Hoggarth et al., 2007; Watters et al., 2009). In Louisiana, it is in the N (Cypress Creek, Bayou D'Arbonne, Tensas, Little Corney Bayou, Dugdemona, Sabine, Bayou Pierre) (Vidrine, 1993). In West Virginia: Upper Ohio (Zeto et al., 1987), Upper Potomac (Clayton et al., 2001; Taylor, 1985). In Maryland, it is in the Upper Potomac, Washington Metro, Chester, Choptank drainages (Bogan and Proch, 1995); lower Susquehanna (Ashton, 2009). In Maine, it is uncommon in most basins (absent from Knox, Lincoln, York Cos.) (Nedeau et al., 2000). In Rhode Island, it is in the middle to S (Raithel and Hartenstein, 2006). In Vermont, it is widely distributed; rarely common (Fichtel and Smith, 1995). Connecticut: Connecticut, Housatonic, Thames drainages (Nedeau and Victoria, 2003; J. Cordeiro, pers. obs., 2006). Massachusetts: most drainages (uncommon exc. Connecticut tribs.) (Smith, 2000). In the Delaware basin, it is in all drainages (Middle Delaware- Mongaup- Broadhead, Upper Delaware, E Branch Delaware); New York to Pennsylvania (Strayer and Ralley, 1991). In Minnesota, it is abundant statewide (Sietman, 2003); Red and Lake of the Woods; not Lake Superior (Graf, 1997; Cvancara, 1970). In South Carolina, it is widespead in the Savannah, Cooper-Santee, and Pee Dee basins (Bogan and Alderman, 2004); Great Pee Dee (Catena Group, 2006); Santee (Alderman, 2006). In South Dakota, it is in the Minnesota, Big Sioux (Skadsen and Perkins, 2000), Lake Kampeska, Vermillion (Backlund, 2000), and James (Perkins and Backlund, 2003). In North Carolina, it is widespread in Atlantic slope (Broad, Pee Dee, Cape Fear, Neuse, Pamlico, Roanoke); French Broad (Tennessee basin) (Bogan, 2002; LeGrand et al., 2006- 36 Cos.). Oklahoma: Illinois, Kiamichi, Neosho, Poteau, Clear Boggy Rivers (Branson, 1983); Mountain Fork (Spooner and Vaughn, 2007), Little and Red Rivers (Vaughn and Taylor, 1999; Vaughn, 2000). In Texas, from Guadalupe drainage N and E (Howells et al., 1996) and Cypress Bayou NE (Mather and Bergmann, 1994). In Wisconsin, it is widespread and abundant (Mathiak, 1997). In Kansas, it is in the Neosho, Verdigris, Spring, and Marais des Cygnes basins; historical in Wakarusa (Tiemann, 2006), Ninnescah (AR), Smoky Hill to lower Republican (KS), and S Fork Big Nemaha (MO) (Couch, 1997). In the Little Blue, weathered in Kansas/Nebraska (Hoke, 2004); common in Kansas part of Big Blue (most weathered); and in Big Hemana (E) (Hoke, 2005); also Platte (Hall, Buffalo Cos.) and Niobrara (Sioux Co.) (Freeman and Perkins, 1992; 1997). It is in the Clinton drainage, Michigan (Strayer, 1980; Goodrich and Van der Schalie, 1939) and Lakes Michigan, Huron, St. Clair drainages (Badra and Goforth, 2003). In Canada, it is widespread/abundant in Ontario; sensitive in Manitoba (Assiniboine drainage- Watson, 2000; Pip, 2006) and Quebec, at risk in Nova Scotia (Cumberland, Cape Breton Cos.) (Athearn and Clarke, 1962; Clarke and Rick, 1964); and information lacking on Saskatchewan and New Brunswick (Shediac River- Athearn, 1961) (Metcalfe-Smith and Cudmore-Vokey, 2004).

Population Size: >1,000,000 individuals
Population Size Comments: This species was recently collected in the Opequon River (Potomac watershed) in West Virginia (Vila et al., 2003). Smith and Crabtree (2010) found this species at 8 of 32 sites (unknown how many with recruitment) along the entire length of Pennsylvania's French Creek.

Number of Occurrences with Good Viability/Integrity: Very many (>125)

Short-term Trend: Relatively Stable (<=10% change)
Short-term Trend Comments: This species is extremely rare or extirpated from the North Fork Holston River, Virginia (Jones and Neves, 2007).

Long-term Trend: Decline of <30% to increase of 25%
Long-term Trend Comments: It is historically known from the Upper Clinch River, Virginia (Jones et al., 2001).

Intrinsic Vulnerability: Not intrinsically vulnerable
Intrinsic Vulnerability Comments: Widespread, though not usually common; generalist.

Environmental Specificity: Broad. Generalist or community with all key requirements common.
Environmental Specificity Comments: Habitat generalist.

Other NatureServe Conservation Status Information

Distribution
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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) The creeper is distributed very widely throughout the Ohio and Mississippi Rivers and Great Lakes systems, northern Atlantic Coast drainages, and parts of Canadian Interior Basin as far west as Texas and Saskatchewan (Burch, 1975; Parmalee and Bogan, 1998) and historically into Kansas and Nebraska (Hoke, 2005). Presumably, it occurred throughout the Tennessee River system historically. Colorado reports are in error/unconfirmed (Wu, 1989; Cordeiro, 1999; Clarke et al., 2003).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AL, AR, CO, CT, DC, DE, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, NC, ND, NE, NH, NJ, NY, OH, OK, PA, RI, SC, SD, TN, TX, VA, VT, WI, WV
Canada MB, NB, NS, ON, QC, SK

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
DE Kent (10001)
IA Allamakee (19005), Black Hawk (19013), Bremer (19017), Buchanan (19019), Carroll (19027), Cerro Gordo (19033), Chickasaw (19037), Clayton (19043), Clinton (19045), Delaware (19055), Des Moines (19057), Dubuque (19061), Floyd (19067), Franklin (19069), Greene (19073), Hamilton (19079), Hardin (19083), Howard (19089), Humboldt (19091), Jackson (19097), Johnson (19103), Jones (19105), Linn (19113), Louisa (19115), Mitchell (19131), Muscatine (19139), Sac (19161), Scott (19163), Story (19169), Washington (19183), Webster (19187), Winneshiek (19191)
KS Allen (20001), Anderson (20003), Bourbon (20011), Chase (20017), Chautauqua (20019), Cherokee (20021), Coffey (20031), Cowley (20035), Dickinson (20041), Elk (20049), Franklin (20059), Greenwood (20073), Jackson (20085), Labette (20099), Lyon (20111), Marion (20115), Marshall (20117), Miami (20121), Montgomery (20125), Morris (20127), Neosho (20133), Shawnee (20177), Wabaunsee (20197), Wilson (20205)
LA Claiborne (22027), Lincoln (22061)*, Madison (22065), Rapides (22079), Union (22111)*
MA Berkshire (25003), Bristol (25005)*, Franklin (25011), Hampden (25013), Hampshire (25015), Middlesex (25017), Norfolk (25021), Plymouth (25023), Worcester (25027)
MD Allegany (24001), Caroline (24011), Carroll (24013), Cecil (24015), Frederick (24021), Garrett (24023), Harford (24025), Howard (24027), Kent (24029), Montgomery (24031), Prince Georges (24033), Queen Annes (24035), Talbot (24041), Washington (24043)
MS Coahoma (28027), Copiah (28029), Issaquena (28055), Marshall (28093), Montgomery (28097), Tishomingo (28141)
NC Alamance (37001), Alexander (37003), Anson (37007), Buncombe (37021), Burke (37023), Cabarrus (37025), Caswell (37033), Catawba (37035), Chatham (37037), Davidson (37057)*, Durham (37063), Edgecombe (37065), Forsyth (37067), Franklin (37069), Gaston (37071), Granville (37077), Halifax (37083), Harnett (37085)*, Henderson (37089), Johnston (37101), Jones (37103), Lee (37105), Mecklenburg (37119), Montgomery (37123), Moore (37125), Nash (37127), Orange (37135), Person (37145), Pitt (37147), Polk (37149)*, Randolph (37151), Richmond (37153), Rockingham (37157), Stanly (37167), Surry (37171), Transylvania (37175), Union (37179), Vance (37181), Wake (37183), Warren (37185), Wilson (37195), Yadkin (37197)
NJ Bergen (34003), Gloucester (34015), Hunterdon (34019), Mercer (34021), Morris (34027), Passaic (34031), Salem (34033), Somerset (34035), Sussex (34037), Warren (34041)
OK LeFlore (40079), McCurtain (40089), Pushmataha (40127)
SC Chesterfield (45025), Edgefield (45037), Lancaster (45057), McCormick (45065), Richland (45079), Saluda (45081)
SD Brookings (46011), Davison (46035), Deuel (46039), Grant (46051), Hamlin (46057), Hutchinson (46067), Lincoln (46083), McCook (46087)*, Minnehaha (46099), Moody (46101), Roberts (46109), Turner (46125)*, Union (46127), Yankton (46135)
TN Hancock (47067)
TX Anderson (48001), Angelina (48005), Bexar (48029), Cass (48067), Cherokee (48073), Harrison (48203), Marion (48315), Nacogdoches (48347), Panola (48365), San Augustine (48405)
VA Scott (51169)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Nashua (01070004)+, Concord (01070005)+, Middle Connecticut (01080201)+, Miller (01080202)+, Deerfield (01080203)+*, Chicopee (01080204)+, Westfield (01080206)+, Farmington (01080207)+, Charles (01090001)+, Cape Cod (01090002)+, Blackstone (01090003)+, Narragansett (01090004)+*, Quinebaug (01100001)+, Housatonic (01100005)+
02 Rondout (02020007)+, Hackensack-Passaic (02030103)+, Raritan (02030105)+, Middle Delaware-Mongaup-Brodhead (02040104)+, Middle Delaware-Musconetcong (02040105)+, Lower Delaware (02040202)+, Cohansey-Maurice (02040206)+, Lower Susquehanna (02050306)+, Chester-Sassafras (02060002)+, Choptank (02060005)+, Patuxent (02060006)+, North Branch Potomac (02070002)+, Cacapon-Town (02070003)+, Conococheague-Opequon (02070004)+, Middle Potomac-Catoctin (02070008)+, Monocacy (02070009)+, Middle Potomac-Anacostia-Occoquan (02070010)+
03 Middle Roanoke (03010102)+, Lower Dan (03010104)+, Upper Tar (03020101)+, Fishing (03020102)+, Lower Tar (03020103)+, Upper Neuse (03020201)+, Contentnea (03020203)+, Lower Neuse (03020204)+, Haw (03030002)+, Deep (03030003)+, Upper Cape Fear (03030004)+*, Upper Yadkin (03040101)+, Lower Yadkin (03040103)+, Upper Pee Dee (03040104)+, Rocky, North Carolina, (03040105)+, Lower Pee Dee (03040201)+, Lynches (03040202)+, Upper Catawba (03050101)+, Upper Broad (03050105)+*, Lower Broad (03050106)+, Saluda (03050109)+, Stevens (03060107)+
05 Youghiogheny (05020006)+
06 Upper French Broad (06010105)+, Upper Clinch (06010205)+, Bear (06030006)+
07 Upper Minnesota (07020001)+, Lac Qui Parle (07020003)+, Coon-Yellow (07060001)+, Upper Iowa (07060002)+, Grant-Little Maquoketa (07060003)+, Turkey (07060004)+, Apple-Plum (07060005)+, Maquoketa (07060006)+, Lower Wisconsin (07070005)+, Copperas-Duck (07080101)+, Upper Wapsipinicon (07080102)+, Flint-Henderson (07080104)+, South Skunk (07080105)+, Skunk (07080107)+, Upper Cedar (07080201)+, Shell Rock (07080202)+, Winnebago (07080203)+, West Fork Cedar (07080204)+, Middle Cedar (07080205)+, Lower Cedar (07080206)+, Upper Iowa (07080207)+, Lower Iowa (07080209)+, Middle Des Moines (07100004)+, Boone (07100005)+, North Raccoon (07100006)+
08 Lower Mississippi-Greenville (08030100)+, Coldwater (08030204)+, Big Sunflower (08030207)+, Bayou D'arbonne (08040206)+, Tensas (08050003)+, Upper Big Black (08060201)+, Bayou Pierre (08060203)+, Bayou Teche (08080102)+
09 Bois De Sioux (09020101)+
10 Lower James (10160011)+, Vermillion (10170102)+*, Upper Big Sioux (10170202)+, Lower Big Sioux (10170203)+, Boyer (10230007)+, Lower Smoky Hill (10260008)+, Upper Kansas (10270101)+, Middle Kansas (10270102)+, Delaware (10270103)+, Lower Big Blue (10270205)+, Upper Marais Des Cygnes (10290101)+, Little Osage (10290103)+, Marmaton (10290104)+
11 Kaw Lake (11060001)+, Upper Verdigris (11070101)+, Fall (11070102)+, Middle Verdigris (11070103)+, Elk (11070104)+, Caney (11070106)+, Neosho headwaters (11070201)+, Upper Cottonwood (11070202)+, Lower Cottonwood (11070203)+, Upper Neosho (11070204)+, Middle Neosho (11070205)+, Spring (11070207)+, Kiamichi (11140105)+, Upper Little (11140107)+, Caddo Lake (11140306)+
12 Middle Sabine (12010002)+, Upper Neches (12020001)+, Upper Angelina (12020004)+, Lower Angelina (12020005)+, Upper San Antonio (12100301)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
U.S. Distribution by Watershed (based on multiple information sources) Help
Ecology & Life History
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Reproduction Comments: The creeper is a long-term brooder, with eggs fertilized in the summer and glochidia released the following spring. Van Snik Gray et al. (2002) found glochidia released 26 April to 2 June in laboratory conditions mimicking Pennsylvania stream habitats. Laboratory studies by Ellis and Keim (1918) revealed plains killifish (Fundulus zebrinus) and green sunfish (Lepomis cyanellis) as glochidial hosts as well as creek chub (Semotilus atromaculatus) according to Baker (1928). Later studies by Hillegass and Hove (1997) found the following fish to be suitable hosts: black bullhead (Ameiurus melas), largemouth bass (Micropterus salmoides), green sunfish (Lepomis cyanellus), black bullhead (Ameirus natalis), and yellow perch (Perca flavescens). Previous studies by Hove (1995) and Hove et al. (1997) also revealed fathead minnow (Pimephales promelas), spotfin shiner (Cyprinella spiloptera), walleye (Stizostedion vitreum), and bluegill (Lepomis macrochirus) as glochidial hosts. Wicklow and Beisheim (1998) found the following fish to be suitable hosts: the longnose dace (Rhinichthys cataractae), fallfish (Semotilus corporalis), golden shiner (Notemigonus crysoleucas), and slimy sculpin (Cottus cognatus). They also found that the larvae of the northern two-lined salamander (Eurycea bislineata) was a suitable glochidial host. Watters et al. (1998a; 1998b; 1999) documented other hosts including fantail darter (Etheostoma flabellare), banded darter (Etheostoma zonale), sand shiner (Notropis lundibundus), bluntnose minnow (Pimephales notatus), and white crappie (Pomoxis annularis). Van Snik Gray et al. (1999; 2002) found 15 hosts (10 of which were previously unknown) bringing the total number to 29, second only to Pyganodon grandis at 32. New host species reported by Van Snik Gray et al. (1999; 2002) include: Acipenser oxyrhynchus- Atlantic sturgeon, Campostoma anomalum- central stoneroller, Luxilus cornutus- common shiner, Nocomis micropogon- river chub, Rhinichthys atratulus- blacknose dace, Oncorhynchus mykiss- rainbow trout, Salvelinus fontinalis- brook charr, Ambloplites rupestris- rock bass, Etheostoma olmstedi- tessellated darter, and Notophthalmus viridescens- red-spotted newt. Recent studies (include those mentioned above) are contrary to early reports (Lefevre and Curtis, 1911) that this species may metamorphose without a host. This species utilizes amphibians as well as fish for hosts. Cliff et al. (2001) futher documented central stoneroller (Campestoma anomalum), brook stickleback (Culaea inconstans), rainbow darter (Etheostoma caeruleum), Iowa darter (Etheostoma exile), Johnny darter (Etheostoma nigrum), channel catfish (Ictalurus punctatus), pumpkinseed (Lepomis gibbosus), burbot (Lota lota), smallmouth bass (Micropteris dolomieu), spottail shiner (Notropis hudsonius), logperch (Percina caprodes), blackside darter (Percina maculata), slenderhead darter (Percina phoxocephala), northern redbelly dace (Phoxinus eos), black crappie (Pomoxis nigromaculatus), blacknose dace (Rhinichthys atratulus), creek chub (Semotilus atromaculatus), and central mudminnow (Umbra limi). New host fish confirmation from Watters et al. (2005): rainbow darter (Etheostoma caeruleum), red-eared sunfish (Lepomis microlophus). Watters et al. (2006) include Ameirus nebulosus brown bullhead, Etheostoma zonale banded darter, and Percina caprodes logperch as hosts. Hove (1995) and Hove et al. (1997) list Cyprinella spiloptera spotfin shiner
Habitat Type: Freshwater
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Riverine Habitat(s): BIG RIVER, CREEK, Low gradient, MEDIUM RIVER, Moderate gradient, Pool, Riffle
Lacustrine Habitat(s): Shallow water
Special Habitat Factors: Benthic
Habitat Comments: This species is a habitat generalist, with a wide distribution. It is usually found in streams and rivers in a range of flow conditions (rarely in high-gradient streams of mountainous regions) but can tolerate lakes and ponds, particularly in outlets.
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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Population/Occurrence Delineation
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Group Name: Freshwater Mussels

Use Class: Not applicable
Minimum Criteria for an Occurrence: Occurrences are based on some evidence of historical or current presence of single or multiple specimens, including live specimens or recently dead shells (i.e., soft tissue still attached and/or nacre still glossy and iridescent without signs of external weathering or staining), at a given location with potentially recurring existence. Weathered shells constitute a historic occurrence. Evidence is derived from reliable published observation or collection data; unpublished, though documented (i.e. government or agency reports, web sites, etc.) observation or collection data; or museum specimen information.
Mapping Guidance: Based on the separation distances outlined herein, for freshwater mussels in STANDING WATER (or backwater areas of flowing water such as oxbows and sloughs), all standing water bodies with either (1) greater than 2 km linear distance of unsuitable habitat between (i.e. lotic connections), or (2) more than 10 km of apparently unoccupied though suitable habitat (including lentic shoreline, linear distance across water bodies, and lentic water bodies with proper lotic connections), are considered separate element occurrences. Only the largest standing water bodies (with 20 km linear shoreline or greater) may have greater than one element occurrence within each. Multiple collection or observation locations in one lake, for example, would only constitute multiple occurrences in the largest lakes, and only then if there was some likelihood that unsurveyed areas between collections did not contain the element.

For freshwater mussels in FLOWING WATER conditions, occurrences are separated by a distance of more than 2 stream km of unsuitable habitat, or a distance of more than 10 stream km of apparently unoccupied though suitable habitat. Standing water between occurrences is considered suitable habitat when calculating separation distance for flowing water mussel species unless dispersal barriers (see Separation Barriers) are in place.

Several mussel species in North America occur in both standing and flowing water (see Specs Notes). Calculation of separation distance and determination of separation barriers for these taxa should take into account the environment in which the element was collected. Juvenile mussels do not follow this pattern and juveniles are typically missed by most standard sampling methods (Hastie and Cosgrove, 2002; Neves and Widlak, 1987), therefore juvenile movement is not considered when calculating separation distance.

Separation Barriers: Separation barriers within standing water bodies are based solely on separation distance (see Separation Distance-suitable, below). Separation barriers between standing water bodies and within flowing water systems include lack of lotic connections, natural barriers such as upland habitat, absence of appropriate species specific fish hosts, water depth greater than 10 meters (Cvancara, 1972; Moyle and Bacon, 1969) or anthropogenic barriers to water flow such as dams or other impoundments and high waterfalls.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 10 km
Alternate Separation Procedure: None
Separation Justification: Adult freshwater mussels are largely sedentary spending their entire lives very near to the place where they first successfully settled (Coker et al., 1921; Watters, 1992). Strayer (1999) demonstrated in field trials that mussels in streams occur chiefly in flow refuges, or relatively stable areas that displayed little movement of particles during flood events. Flow refuges conceivably allow relatively immobile mussels to remain in the same general location throughout their entire lives. Movement occurs with the impetus of some stimulus (nearby water disturbance, physical removal from the water such as during collection, exposure conditions during low water, seasonal temperature change or associated diurnal cycles) and during spawning. Movement is confined to either vertical movement burrowing deeper into sediments though rarely completely beneath the surface, or horizontal movement in a distinct path often away from the area of stimulus. Vertical movement is generally seasonal with rapid descent into the sediment in autumn and gradual reappearance at the surface during spring (Amyot and Downing, 1991; 1997). Horizontal movement is generally on the order of a few meters at most and is associated with day length and during times of spawning (Amyot and Downing, 1997). Such locomotion plays little, if any, part in the distribution of freshwater mussels as these limited movements are not dispersal mechanisms. Dispersal patterns are largely speculative but have been attributed to stream size and surface geology (Strayer, 1983; Strayer and Ralley, 1993; van der Schalie, 1938), utilization of flow refuges during flood stages (Strayer, 1999), and patterns of host fish distribution during spawning periods (Haag and Warren, 1998; Watters, 1992). Lee and DeAngelis (1997) modeled the dispersal of freshwater into unoccupied habitats as a traveling wave front with a velocity ranging from 0.87 to 2.47 km/year (depending on mussel life span) with increase in glochidial attachment rate to fish having no effect on wave velocity.

Nearly all mussels require a host or hosts during the parasitic larval portion of their life cycle. Hosts are usually fish, but a few exceptional species utilize amphibians as hosts (Van Snik Gray et al., 2002; Howard, 1915) or may metamorphose without a host (Allen, 1924; Barfield et al., 1998; Lefevre and Curtis, 1911; 1912). Haag and Warren (1998) found that densities of host generalist mussels (using a variety of hosts from many different families) and displaying host specialists (using a small number of hosts usually in the same family but mussel females have behavioral modifications to attract hosts to the gravid female) were independent of the densities of their hosts. Densities of non-displaying host specialist mussels (using a small number of hosts usually in the same family but without host-attracting behavior) were correlated positively with densities of their hosts. Upstream dispersal of host fish for non-displaying host specialist mussels could, theoretically, transport mussel larvae (glochidia) over long distances through unsuitable habitat, but it is unlikely that this occurs very often. D. Strayer (personal communication) suggested a distance of at least 10 km, but a greater distance between occurrences may be necessary to constitute genetic separation of populations. As such, separation distance is based on a set, though arbitrary, distance between two known points of occurrence.

Date: 18Oct2004
Author: Cordeiro, J.
Notes: Contact Jay Cordeiro (jay_cordeiro@natureserve.org) for a complete list of freshwater mussel taxa sorted by flow regime.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 31Dec2011
NatureServe Conservation Status Factors Author: Cordeiro, J.
Element Ecology & Life History Edition Date: 31Dec2011
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).

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