Epioblasma rangiana

(I. Lea, 1838)

Northern Riffleshell

G1Critically Imperiled Found in 1 roadless area NatureServe Explorer →
G1Critically ImperiledGlobal Rank
Critically endangeredIUCN
Very highThreat Impact
Northern riffleshell (Epioblasma rangiana). Photo by U.S. Fish & Wildlife Service, Public Domain (U.S. Government Work), via ECOS.
U.S. Fish & Wildlife Service, https://www.usa.gov/government-works
Identity
Unique IDELEMENT_GLOBAL.2.118738
Element CodeIMBIV16184
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryInvertebrate Animal
IUCNCritically endangered
CITESAppendix II
Endemicoccurs (regularly, as a native taxon) in multiple nations
KingdomAnimalia
PhylumMollusca
ClassBivalvia
OrderUnionoida
FamilyUnionidae
GenusEpioblasma
Synonyms
Dysnomia sulcata delicata(Conrad, 1836)Dysnomia torulosa rangiana(I. Lea, 1838)Epioblasma bilobaRafinesque, 1831Epioblasma sulcata delicata(Conrad, 1836)Epioblasma torulosa rangiana(I. Lea, 1838)
Other Common Names
Épioblasme ventrue (FR)
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.
Taxonomic Comments
This taxon was previously recognized as a subspecies of Epioblasma torulosa but was elevated to species status by Williams et al. (2017). They noted that "Turgeon et al. (1998) recognized eight subspecies of Epioblasma in three nominal species: florentina (three), obliquata (two), and torulosa (three). A conclusive assessment of the taxonomic status of these taxa may be impossible at this time because half are considered extinct (E. florentina florentina, E. f. curtisii, E. torulosa torulosa, and E. t. gubernaculum). Cummings and Berlocher (1990) found no evidence of intergradation between E. t. torulosa and E. t. rangiana and both taxa co-occurred at many sites; based on this evidence, we elevate these subspecies to species status."
Conservation Status
Rank MethodLegacy Rank calculation - Biotics v1
Review Date2019-10-09
Change Date2019-10-09
Edition Date2019-10-09
Edition AuthorsJackson, D. R. (2019); Cordeiro, J. (2007); Bier, C., T. G. Watters,and K.S. Cummings (1998)
Threat ImpactVery high
Range Extent250-5000 square km (about 100-2000 square miles)
Number of Occurrences6 - 20
Rank Reasons
This once widespread species historically occurred throughout much of the Ohio River watershed, but its range has been dramatically reduced to eight to ten populations dispersed across four states and one province; some of these populations may have low viability. Chief factors in decline are the damming of rivers, which floods habitat and fragments populations; erosion and siltation from strip mining, logging, and farming; and pollution from agricultural and industrial runoff. Invasive mussels pose an additional serious threat.
Range Extent Comments
The species' range centers around the U.S. Midwest and the Ohio River drainage. Range states include Illinois, Indiana, Kentucky, Michigan, New York, Ohio, Pennsylvania, and West Virginia. Additionally, it occurs in the Canadian province of Ontario within the Lake Huron drainage. As with most naiads, the species' present range is a remnant of its former distribution. E. rangiana was once thought to be a subspecies of E. torulosa that replaced Epioblasma t. torulosa in the headwaters of the Ohio River drainage. However, Cummings and Berlocher (1990) found that both taxa co-occurred at many sites (without intergradation).

NOTE: the following pre-2008 summary contains useful information but was written before E. rangiana was elevated to species status; it therefore may include data for E. torulosa and hence should be reevaluated.
Historically, this entity was recorded from the mainstem of the Ohio River (Stansbery and Cooney, 1985) but has apparently been extirpated from there. The Ohio State University Museum of Zoology has records for the White and Wabash Rivers in Indiana but it has not been collected there in recent times (Clark, 1976; Cummings et al., 1991). Taylor and Hughart (1981) presumed that it was no longer present in the Elk River of West Virginia. Currently extant in only seven streams: the Green River in Kentucky, French and LeBoeuf Creeks and the Allegheny River in Pennsylvania, the Detroit River in Michigan (possibly extirpated - see below), Big Darby Creek in Ohio (USFWS, 1993), and at least one additional river in Ontario (Metcalfe-Smith et al., 1998). Staton et al. (2000) lists historical and present distribution. Recently found in Conewanto Creek near Warren, Warren Co., Pennsylvania, where it was previously thought to be extirpated (Evans and Smith, 2005). In Illinois, it formerly occupied the Vermilion, Wabash, and Ohio rivers but is now extirpated in the state (Cummings and Mayer, 1997). Recently this species has been confirmed to be likely extirpated from the main channel of the Detroit River between Lake St. Clair and Lake Erie, Michigan/Ontario, due to zebra mussel invasion (Schloesser et al., 2006).
Occurrences Comments
Once widespread in the Ohio River basin and into southern Canada in the Lake Huron drainage, this species now exists in eight to ten isolated populations, most of which are small and peripheral and with few signs of reproduction.

NOTE: the following paragraph was written prior to 2010 when the taxon was still considered a subspecies of E. torulosa. It is retained here for its detailed notes on occurrence in multiple river systems.

Epioblasma torulosa rangiana, the only remaining extant subspecies, was formerly widespread in the Ohio River basin (including Ohio River system in Ohio River at Cincinnati, Little Miami, Scioto and tribs., Muskingham, Auglaize, Sandusy, and Olentangy Rivers, and Big Darby Creek- a tributary of the Scioto; Beaver River in Ohio and Pennsylvania; Lake Erie drainage in Fish Creek, and Kanawha River, West Virginia; Wabash River and tributaries-Tippecanoe River and the Eel, Blue, and White Rivers) (Parmalee and Bogan, 1998) as well as having been collected historically from the Allegheny River drainage in Pennsylvania above Pittsburgh and from the West Fork River (Ortmann, 1913), a tributary of the Monongahela River, Harrison Co., West Virginia (Parmalee and Bogan, 1998; USFWS, 1994). This subspecies now exists in eight to ten isolated populations, most of which are small and peripheral and with little signs of reproduction. It is known from the Kentucky, Licking, and Green River drainages in Kentucky (Johnson, 1978), but is likely only still extant in the Upper Green. In the Maumee River drainage, it was last seen in Fish Creek (St. Joseph drainage) in Ohio and Indiana and Big Darby Creek in Ohio (Watters et al., 2009), where live individuals were very rare (Watters, 1995; Grabarkiewicz and Crail, 2006). Recently found in Conewango Creek near Warren, Warren Co., Pennsylvania, where it was previously thought to be extirpated (Evans and Smith, 2005). It also occurs in the Allegheny River, Pennsylvania (Jones, 2004). Epioblasma torulosa rangiana occurs in Muddy Creek (French Creek drainage) in the Erie NWR in Crawford Co., Pennsylvania (Mohler et al., 2006) and elsewhere in the French Creek drainage. Weathered shells are reported from the Tippecanoe River (shell only- Cummings and Berlocher, 1990), Sugar Creek (east fork White River drainage) in central Indiana (Harmon, 1992) as well as throughout most central Indiana drainages but no living specimens are known recently (USFWS, 1994) and Fisher (2006) lists it as extirpated from the Wabash drainage. As such, it is likely extirpated from the rest of Indiana (IN NHP, pers. comm., 2009) with the last remaining population in the Tippecanoe drainage (USFWS, 1994). Specimens from the Black River (St. Clair drainage), Michigan, were relocated to the Detroit River in 1992 (Trdan and Hoeh, 1993). It is historically known from the Clinton River drainage in Michigan (Strayer, 1980). In Canada, it historically occurred in the Cedar River and currently occurs in the Sydenham River in the Northern Lake Erie drainage (Metcalfe-Smith et al., 2003) and also the Ausable River in the Eastern Lake Huron drainage (Staton et al., 2000).
Threat Impact Comments
Members of the genus Epioblasma seem to be particularly sensitive to impoundment; most species are riffle/run inhabitants and cannot tolerate other habitats. For example, impoundment of the Clinch River in Tennessee by the Norris Reservoir resulted in the extirpation of the majority of species below the dam (Ahlstedt 1984), and construction of the Wilson Dam on the Tennessee River eliminated 20 of the original 22 Cumberlandian naiad species (Stansbery 1971). 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 Zebra mussel, Dreissena polymorpha, has 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 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 development, mineral extraction, and industrial development also pose significant threats. 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 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 long-term recolonization abilities by destroying much of the potential habitat by making substrates and flow rates uniform throughout a 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 (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. USFWS (1994) lists the following reasons for mussel decline: siltation (from agriculture, construction, and forestry runoff), impoundment (including dam construction and maintenance), instream sand and gravel mining (for channelization), pollutants (pesticides and fertilizers, heavy metals, ammonia from wastewater, acid-mine runoff, and invasive species (zebra mussel, quagga mussel).
Ecology & Habitat

Description

SHELL: Shell elongate, thick anteriorly, much thinner posteriorly, and compressed. Anterior margin rounded, posterior-ventral margin broadly rounded in females, indented in males. Umbos low, somewhat turned forward. Tubercles or knobs on the lateral surface absent. A wide, shallow sulcus or depression present between the posterior and medial ridges. Periostracum yellowish brown with numerous fine green rays, obscure in older shells. Adult size to 2 inches. Pseudocardinal teeth triangular, divergent, and roughened; two in the left valve, one in the right, with two smaller teeth on either side. Lateral teeth moderately long, straight, and roughened; two in the left valve, one in the right with a much smaller tooth below. Beak cavity moderate to shallow. Nacre white, iridescent posteriorly (Cummings and Mayer, 1992).

ANIMAL: Unknown.

Habitat

Ortmann (1919: 334) reported that this species was "always found...on riffles, on a bottom of firmly packed and rather fine gravel, in swiftly flowing, shallow water or coarse gravel" and Clarke (1981: 362) gave its habitat as "highly oxygenated riffle". Preferred habitat appears to require swiftly moving water. The high oxygen concentrations in swift streams may be necessary for survival. It is a species of riffle areas of smaller streams, and as such has fared better than larger river species, which have been heavily impacted by dredging and impoundment. Of the eleven or so species of naiads thought to be extinct in 1971 by Stansbery, most were from this latter type of habitat and all were species of Epioblasma.

Reproduction

Ortmann (1912: 358) reported Epioblasma torulosa rangiana to be gravid in September and Clarke (1981: 352) said it was a "longterm breeder, gravid from late summer to following spring". The female uses the posterior portion of the outer gill as marsupia. It is widely believed that the "brood pouch" swellings of the female shell are caused by the expanded gravid gills, but Stansbery (pers. comm.) and Hoggarth (pers. comm.) do not believe this to be the case.

Watters (1996) reports the following fish as suitable glochidial hosts: the mottled sculpin (Cottus bairdi), banded darter (Etheostoma zonale), bluebreast darter (Etheostoma camurum), and brown trout (Salmo trutta).

Based upon counts of annular growth lines, this species may reach 15+ years of age. It is not known at what ages reproductive maturity begins and ends. Because of the rarity of live material (and their enforced protection), it is not known if existing populations are reproductively active. Because of their small size, it is not known if juveniles are present in any of the populations. It must be emphasized that existing populations may be large, healthy, and reproductively active and still be in imminent danger of extinction if the host fish is not present in the range.
Other Nations (2)
CanadaN1
ProvinceRankNative
OntarioS1Yes
United StatesN2
ProvinceRankNative
IllinoisS1Yes
West VirginiaS1Yes
MichiganS1Yes
KentuckyS1Yes
OhioS1Yes
New YorkS1Yes
IndianaS1Yes
PennsylvaniaS2Yes
Threat Assessments
ThreatScopeSeverityTiming
1 - Residential & commercial developmentLarge - restrictedModerate - slightHigh (continuing)
1.1 - Housing & urban areasLarge - restrictedModerate - slightHigh (continuing)
1.2 - Commercial & industrial areasLarge - restrictedModerate - slightHigh (continuing)
2 - Agriculture & aquacultureRestricted (11-30%)Serious - slightHigh (continuing)
2.1 - Annual & perennial non-timber cropsRestricted (11-30%)Serious - slightHigh (continuing)
3 - Energy production & miningRestricted (11-30%)Extreme - seriousHigh (continuing)
3.2 - Mining & quarryingRestricted (11-30%)Extreme - seriousHigh (continuing)
7 - Natural system modificationsLarge - restrictedSerious or 31-70% pop. declineHigh (continuing)
7.2 - Dams & water management/useLarge - restrictedSerious or 31-70% pop. declineHigh (continuing)
8 - Invasive & other problematic species, genes & diseasesLarge (31-70%)UnknownHigh (continuing)
8.1 - Invasive non-native/alien species/diseasesLarge (31-70%)UnknownHigh (continuing)
8.2 - Problematic native species/diseasesRestricted - smallSlight or 1-10% pop. declineHigh (continuing)
9 - PollutionLarge (31-70%)Serious or 31-70% pop. declineHigh (continuing)
9.1 - Domestic & urban waste waterLarge - restrictedSerious - moderateHigh (continuing)
9.2 - Industrial & military effluentsLarge (31-70%)Serious or 31-70% pop. declineHigh (continuing)
9.3 - Agricultural & forestry effluentsLarge - restrictedSerious - moderateHigh (continuing)

Roadless Areas (1)
Pennsylvania (1)
AreaForestAcres
Allegheny FrontAllegheny National Forest7,430
References (69)
  1. 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.
  2. 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.
  3. Bogan, A.E. 1997. A resolution of the nomenclatural confusion surrounding <i>Plagiola </i>Rafinesque, <i>Epioblasma </i>Rafinesque, and <i>Dysnomia </i>Agassiz (Mollusca: Bivalvia: Unionidae). Malacological Review 30:77-86.
  4. Clark, C.F. 1976. The freshwater naiads of the lower end of the Wabash River, Mt. Carmel, Illinois to the South. Sterkiana, (61): 1-14.
  5. 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.
  6. 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.
  7. Crabtree, D.L. and T.A. Smith. 2009. Population attributes of an endangered mussel, <i>Epioblasma torulosa rangiana</i> (northern riffleshell), in French Creek and implications for its recovery. Northeastern Naturalist 16(3):339-354.
  8. 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.
  9. 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.
  10. 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.
  11. Cummings, K.S., C.A. Mayer, and L.M. Page. 1991. Survey of the freshwater mussels (Mollusca: Unionidae) of the Wabash River drainage. Phase III: White River and selected tributaries. Illinois Natural History Survey, Center for Biodiversity, Technical Report, 1991(3): 47 pp + appendices.
  12. D. B. Abrell. Indiana Department of Natural Resources. Personal communication.
  13. Evans, R. and T. Smith. 2005. Northern riffleshells located in Conewango Creek, Allegheny River basin, Pennsylvania, with a note on new records for the zebra mussel. Ellipsaria, 7(3): 6-7.
  14. Evermann, B.W. and H.W. Clark. 1918. The Unionidae of Lake Maxinkukee. Proceedings of the Indiana Academy of Science 1917:251-285.
  15. Fisher, B.E. 2006. Current status of freshwater mussels (Order Unionoida) in the Wabash River drainage of Indiana. Proceedings of the Indiana Academy of Science, 115(2): 103-109.
  16. Freshwater Mollusk Conservation Society (FMCS). 2023. The 2023 checklist of freshwater bivalves (Mollusca: Bivalvia: Unionida) of the United States and Canada. Considered and approved by the Bivalve Names Subcommittee October 2023. Online: https://molluskconservation.org/MServices_Names-Bivalves.html
  17. 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.
  18. Graf, D.L. and K.S. Cummings. 2021. A 'big data' approach to global freshwater mussel diversity (Bivalvia: Unionoida), with an updated checklist of genera and species. Journal of Molluscan Studies 87(1):1-36.
  19. 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.
  20. Hill, D.M. 1986. Cumberlandian mollusk conservation program, activity 3: identification of fish hosts. Office of Natural Resources and Economic Development, Tennessee Valley Authority, Knoxville, Tennessee. 55 pp.
  21. Hoggarth, M. A. 1985. Museum of Zoology, Ohio State University. Personal communication.
  22. 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.
  23. Johnson, R.I. 1978. Systematics and zoogeography of <i>Plagiola </i>(= <i>Dysnomia </i>= <i>Epioblasma</i>), an almost extinct genus of freshwater mussels (Bivalvia: Unionidae) from middle North America. Bulletin of the Museum of Comparative Zoology, 148(6): 239-320.
  24. Jones, J.W. 2004. A holistic approach to taxonomic evaluation of two closely related endangered freshwater mussel species, the oyster mussel (<i>Epioblasma capsaeformis</i>) and tan riffleshell (<i>Epioblasma florentina walkeri</i>) (Bivalvia: Unionidae). MS Thesis, Virginia Polytechnic Institute and State University, Blacksburg, Virginia. 178 pp.
  25. 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.
  26. Metcalfe-Smith, J.L., G.L. Mackie, S.K. Staton, and E.L. West. 1998. Current status of rare species of freshwater mussels in southern Ontario. Triannual Unionid Report, 14: unpaginated.
  27. 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.
  28. 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.
  29. MolluscaBase eds. 2024. MolluscaBase. Accessed at https://www.molluscabase.org
  30. 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.
  31. Neves, R.J. and J.C. Widlak. 1987. Habitat ecology of juvenile freshwater mussels (Bivalvia: Unionidae) in a headwater stream in Virginia. American Malacological Bulletin 5:1-7.
  32. Ortmann, A.E. 1912. Notes upon the families and genera of the najades. Annals of the Carnegie Museum 8(2):222-365.
  33. Ortmann, A.E. 1913. The Alleghenian divide, and its influence on the freshwater fauna. Proceedings of the American Philosophical Society, 52(210): 287-390.
  34. 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.
  35. Parmalee, P.W. 1967. The freshwater mussels of Illinois. Illinois State Museum, Popular Science Series 8:1-108.
  36. Parmalee, P.W. and A.E. Bogan. 1998. The Freshwater Mussels of Tennessee. University of Tennessee Press: Knoxville, Tennessee. 328 pp.
  37. Schloesser, D.W., J.L. Metcalfe-Smith, W.P. Kovalak, G.D. Longton, and R.D. Smithee. 2006. Extirpation of freshwater mussels (Bivalvia: Unionidae) following the invasion of dreissenid mussels in an interconnecting river of the Laurentian Great Lakes. American Midland Naturalist, 155: 307-320.
  38. 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.
  39. 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.
  40. Smith, T.A. and D. Crabtree. 2010. Freshwater mussel (Unionidae: Bivalvia) distributions and densities in French Creek, Pennsylvania. Northeastern Naturalist 17(3):387-414.
  41. 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.
  42. Spoo, A. 2008. The Pearly Mussels of Pennsylvania. Coachwhip Publications: Landisville, Pennsylvania. 210 pp.
  43. Stansbery, D.H. 1970. Eastern freshwater mollusks (I): The Mississippi and St. Lawrence River systems. Malacologia, 10(1): 9-22.
  44. Stansbery, D. H. 1971. Rare and endangered freshwater mollusks in eastern United States. Pages 5-18 in S.E. Jorgensen, and R.W. Sharp. Proceedings of a symposium of rare and endangered mollusks (naiads) of the United States. U.S. Department of the Interior: Twin Cities, Minnesota. 79 pp.
  45. Stansbery, D. H., and J. D. Cooney. 1985. Survey of the unionid mollusks of the Ohio River in the vicinity of the William H. Zimmer Station (Ohio River miles 442.6 to 445.6). Final Report submitted to Cincinnati Gas and Electric Company, Columbus and Southern Ohio Electric Company, Dayton Power and Light Company. 33 pp.
  46. Stansbery, D. H. Museum of Zoology, Ohio State University. Personal communication.
  47. Staton, S.K., J.L. Metcalfe-Smith, and E.L. West. 2000. Status of the northern riffleshell, <i>Epioblasma torulosa rangiana</i> (Bivalvia: Unionidae), in Ontario and Canada. The Canadian Field-Naturalist, 114: 224-235.
  48. 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.
  49. 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.
  50. Taylor, R.W. and R.C. Hugart. 1981. The freshwater naiads of Elk River, West Virginia, with a comparison of earlier collections. The Nautilus, 95(1): 21-25.
  51. Trdan, R.J. and W.R. Hoeh. 1993. Relocation of two state-listed freshwater mussel species (<i>Epioblasma torulosa rangiana</i> and <i>Epioblasma triquetra</i>) 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.
  52. Trdan, R. J., W.R. Hoeh, and S.E. Cordoba. 1987. Viable populations of <i>Dysnomia torulosa</i>, an endangered freshwater mussel, in the Black River of Sanilac County, Michigan. Unpublished report to MDNR.
  53. 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.
  54. U.S. Fish and Wildlife Service (USFWS). 1985d. Recovery plan for tubercled-blossom pearly mussel <i>Epioblasma </i>(= <i>Dysnomia</i>) <i>torulosa torulosa</i> (Rafinesque, 1820), turgid-blossom pearly mussel <i>Epioblasma </i>(= <i>Dysnomia</i>) <i>turdigula</i> (Lea, 1858), and yellow-blossom pearly mussel <i>Epioblasma </i>(= <i>Dysnomia</i>) <i>florentina florentina</i> (Lea, 1857). U.S. Fish and Wildlife Service, Atlanta, Georgia. 39 pp.
  55. U.S. Fish and Wildlife Service (USFWS). 1993. Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for the Northern Riffleshell Mussel (<i>Epioblasma torulosa rangiana</i>) and the Clubshell Mussel (<i>Pleurobema clava</i>) (Final Rule). Federal Register, 58(13): 5638-5642.
  56. U.S. Fish and Wildlife Service (USFWS). 1994. Clubshell (Pleurobema clava) and northern riffleshell (Epioblasma torulosa rangiana) recovery plan. U.S. Fish and Wildlife Service, Hadley, Massachusetts. 58 pp.
  57. U.S. Fish and Wildlife Service (USFWS). 2022. Endangered and Threatened Wildlife and Plants; Technical Amendments for Southeastern Mussels, Snails, and a Reptile. Federal Register 87(33):8960-8967.
  58. U.S. Fish and Wildlife Service (USFWS). 2022. Endangered and Threatened Wildlife and Plants; Technical Corrections for Four Midwest Mussel Species. Federal Register 87(52):15143-15146.
  59. Watters, G.T. 1994. An annotated bibliography of the reproduction and propagation of the Unionoidea (primarily of North America). Ohio Biological Survey Miscellaneous Contributions, 1: 1-158.
  60. Watters, G.T. 1995a. A field guide to the freshwater mussels of Ohio. revised 3rd edition. Ohio Department of Natural Resources, Division of Wildlife, Columbus, Ohio. 122 pp.
  61. Watters, G.T. 1996. Hosts for the northern riffle shell (<i>Epioblasma torulosa rangiana</i>). Triannual Unionid Report 10: 14.
  62. Watters, G.T. 2008. Northern riffleshells transplanted to Big Darby Creek, Ohio. Ellipsaria, 10(2): 7.
  63. Watters, G.T. and C.J.M. Flaute. 2010. Dams, zebras, and settlements: The historical loss of freshwater mussels in the Ohio River mainstem. American Malacological Bulletin 28:1-12.
  64. Watters, G.T., M.A. Hoggarth, and D.H. Stansbery. 2009b. The Freshwater Mussels of Ohio. Ohio State University Press: Columbus, Ohio. 421 pp.
  65. Wells, S.M., R.M. Pyle, and N.M. Collins. 1983. The IUCN Invertebrate Red Data Book. IUCN, Gland, Switzerland. 632 pp.
  66. 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.
  67. Williams, J. D., A. E. Bogan, R. S. Butler, K. S. Cummings, J. T. Garner, J. L. Harris, N. A. Johnson, and G. T. Watters. 2017. A revised list of the freshwater mussels (Mollusca: Bivalvia: Unionida) of the United States and Canada. Freshwater Mollusk Biology and Conservation 20:33-58.
  68. 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.
  69. Zanatta, D.T. and R.W. Murphy. 2007. Range-wide population genetic analysis of the endangered northern riffleshell mussel, <i>Epioblasma torulosa rangiana</i> (Bivalvia, Unionoida). Conservation Genetics 8:1393-1404.