Simpsonaias ambigua

(Say, 1825)

Salamander Mussel

G1Critically Imperiled (G1G2) Found in 8 roadless areas NatureServe Explorer →
G1Critically ImperiledGlobal Rank
VulnerableIUCN
Very high - highThreat Impact
Identity
Unique IDELEMENT_GLOBAL.2.114652
Element CodeIMBIV41010
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryInvertebrate Animal
IUCNVulnerable
Endemicoccurs (regularly, as a native taxon) in multiple nations
KingdomAnimalia
PhylumMollusca
ClassBivalvia
OrderUnionoida
FamilyUnionidae
GenusSimpsonaias
Synonyms
Simpsoniconcha ambigua(Say, 1825)
Other Common Names
Mudpuppy Mussel (EN) Mulette du Necture (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.
Conservation Status
Rank Method Rank calculation - Biotics v2
Review Date2023-09-06
Change Date2023-09-06
Edition Date2023-09-06
Edition AuthorsCordeiro, J. (2006); Whittaker, J.C. (1998); rev. R. L. Gundy (2023)
Threat ImpactVery high - high
Range Extent200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Number of Occurrences21 - 80
Rank Reasons
This species has faced steep historical declines and continues to decline. Range extent has declined by 30%, the number of occurrences has declined by 40%, and area of occupancy has been greatly reduced by numerous local extirpations. Approximately 66 occurrences remain extant, of which 57 are expected to become extirpated or functionally extirpated by 2043. It is highly threatened by pollution, sedimentation, and erosion. The host for its larvae (glochidia) and sole means of long-distance dispersal, the mudpuppy (Necturus maculosus), is also declining.
Range Extent Comments
This species lives in the Great Lakes region of the United States and Ontario, Canada. Historically, this species was known from the St. Croix River on the Wisconsin/Minnesota border west to western New York and southeast to Arkansas and Missouri (GBIF 2023, McMurray 2015, Morris and Burridge 2006, USFWS 2023). The historical range extent was approximately 1,000,000 km² (GBIF 2023, GeoCat 2023). Drastic declines and numerous local extirpations have reduced its range. It appears to be extirpated from Arkansas, Iowa, Kansas, Missouri, New York, North Carolina, Virginia, and West Virginia (GBIF 2023, McMurray 2015, Morris and Burridge 2006, USFWS 2023). These extirpations have reduced range extent to approximately 700,000 km² (GBIF 2023, GeoCat 2023).
Occurrences Comments
There are approximately 66 occurrences (USFWS 2023). See area of occupancy comments for a breakdown of extant locations.
Threat Impact Comments
Declining water quality due to agriculture and development is one of the primary threats to this species. Pollution is a primary threat. Pollutants such as ammonia, chloride, copper, potassium, nitrogen, and phosphorous likely have serious impacts on this species (Hinck et al. 2011, Kost et al. 2012, Salerno et al. 2018, USFWS 2023). Commonly encountered pesticides may pose only a minimal risk (Salerno et al. 2020). Sedimentation, often due to shorelines where vegetation has been removed, is also a primary threat throughout its range (Hinck et al. 2011, Kost et al. 2012, Morris and Burridge 2006, USFWS 2023). Invasive plants can also decrease the stability of shorelines, contributing to erosion (Kost et al. 2011). The invasive zebra mussel (Dreissena polymorpha) threatens the salamander mussel by competition for space, including zebra mussels colonizing the outer shell surface at densities high enough to incapacitate the host mussel, and is present in multiple waterways occupied by the salamander mussel (Hinck et al. 2011, Metcalfe-Smith and Cudmore-Vokey 2004, Smith and Meyer 2010). The invasive round goby (Neogobius melanostomus) may also represent a threat, possibly through disruption of local food webs, and is present in multiple waterways including the Sydenham River, Ontario and French Creek, Pennsylvania (Clark 2021, Ho 2011, McDaniel et al. 2009, Poos et al. 2010). This species is susceptible to warming temperatures and drought conditions predicted to occur with global climate change (Lee et al. 2011). Its host, the mudpuppy (Necturus maculosus), is also declining due to collection for biological supply companies, siltation, lampricide application, habitat degradation, and toxic algal blooms (Hoffman et al. 2014, McDaniel et al. 2009, Stapleton et al. 2018).
Ecology & Habitat

Description

Clarke (1985) gives the following description. "Shell lenticular-ovate, moderately inflated, unsculptured, with only rudimentary hinge teeth; up to about 48 mm long, 22 mm high, and 16 mm wide. Much thicker (2.9 mm) anteriorly than posteriorly (0.5 mm). Anterior margin more or less evenly rounded; ventral margin flatly rounded, flat, or slightly concave just posterior of center; posterior margin also more or less evenly rounded and similar to anterior margin or a little broader; and dorsal parallel with ventral margin except indented in front of the umbones. Maximum inflation (and maximum height) both behind the middle of the shell. Beaks somewhat pointed, directed inwardly and toward the anterior, not inflated, located about 24% to 26% the distance from anterior to posterior, and projecting only a little above the hinge line. Area of posterior ridge convex and inflated but no distinct posterior ridge is discernable in most species. Area of posterior slope slightly concave near margin. Growth increments indicated by dark concentric periostracal bands covering barely discernable concentric grooves. Additional post-juvenile sculpturing consisting only of concentric threads (especially anteriorly), a few obscure radial lines posteriorly, and low lines and grooves of growth. Periostracum predominately brown or yellowish brown but blackish posteriorly in some specimens. Rarely one sees faint traces of narrow rays over the center of the shell, but most specimens, including juveniles, are entirely unrayed. Ligament rather long or of medium length, of moderate thickness, brown, and fragile when dry. Hinge teeth unusual, small, and incomplete. The right valve has a single, small, low, rounded, slightly elongated pseudocardinal tooth that arises from the shell wall (not from a thickened hinge plate as in other specimens) just in front of the umbone. The left valve of some specimens has an even smaller tooth that arises below and posterior to the umbone, i.e., in the same position as the interdental projection in Lasmigona species. This tooth, where present, is low, short, somewhat flange-like, rounded, and irregular. There are no articulating lateral hinge teeth, although the edge of the shell is a little thickened below the ligament and, in some specimens, a poorly defined lateral ridge may be present. Beak cavities somewhat excavated but not deep, and with a variable number of small, irregular muscle scars within. Major anterior muscle scars small and shallow but well marked; pallial band well marked, located quite far from the margin (and more clearly defined) anteriorly but closer to the margin posteriorly, and in some specimens with several tiny, parallel, collabral ridges within the anterior portion; and posterior muscle scars very lightly etched and located distinctly forward of the most posterior portion of the pallial band loop. Nacre bluish white, iridescent posteriorly, with yellowish, salmon, or purplish suffusions in the center and near the beak cavities, and thin in a narrow band around the edge of the shell with the periostracal color showing through. Beak sculpture composed of about six parallel, predominately inverted, V-shaped ridges, apparently corresponding to the middle portions of the double-looped ridges in Lasmigona . The earliest two ridges ore obscure in available material: they appear to be single-looped and especially expanded anteriorly. Later ridges are inverted V- shaped, with anterior arms short and directed ventrally in their proximal portions but curving anteriorly distally, and with posterior arms also short and more or less straight and parallel with the ligament or slightly curved upward. In some specimens the bars are in the shape of shallow V's and the later bars are simple undulating ridges but in most specimens the bars are deeply indented centrally."

Habitat

Although occasionally found elsewhere, there is little doubt the preferred habitat is under large, flat stones in areas of swift current in medium to large rivers and lakes (Call 1900, Howard 1915, Buchanan 1980, Clarke 1985, Oesch 1984, Oesch 1995, Parmalee and Bogan 1998). Its presence is linked to the mudpuppy (Necturus maculosus) and it is usually rare though can be abundant in patches. This species generally occupies rivers, but can also be found in creeks, streams, and lakes on a variety of substrates (mud, silt, sand, gravel, cobble or boulder) in areas of swift current (Cudmore et al. 2004).

Ecology

Refer to the General Freshwater Mussel ESA for general ecology of mussels.

Reproduction

Howard (1915, 1951) demonstrated that the host is the mudpuppy (Necturus maculosus). He suspected "that Necturus eats the adult mussel and in seeking food visits one rock after another. In satisfying its appetite it becomes infected with the mussel glochidia, nourishing them, and when they have matured serves as a transporting and distributing agent for the young mussels". Glochidia were found deeply imbedded in the external gills of the mudpuppy, which was confirmed as a host by Bequaert et al. (1998). There is some evidence that the glochidia are released in the fall (Clarke 1985).

Simpson (1914) described gravid specimens; "Brood pouch filling the entire outer gills and forming enormously thickened pads, the upper part finely vertically striate, the lower part of different texture, lighter colored, wrinkled and granular on the surface: embryos very large; outer and inner gills nearly alike in size, the latter free from the abdominal sac, all united to the mantle to their posterior ends..."
Other Nations (2)
United StatesN3
ProvinceRankNative
New YorkS1Yes
IndianaS2Yes
MissouriS1Yes
MichiganS1Yes
IllinoisS1Yes
ArkansasS1Yes
KentuckyS2Yes
West VirginiaS2Yes
OhioS2Yes
MinnesotaS1Yes
IowaSXYes
WisconsinS2Yes
PennsylvaniaS1Yes
TennesseeS1Yes
CanadaN1
ProvinceRankNative
OntarioS1Yes
Threat Assessments
ThreatScopeSeverityTiming
8 - Invasive & other problematic species, genes & diseasesLarge - restrictedModerate - slightHigh (continuing)
8.1 - Invasive non-native/alien species/diseasesLarge - restrictedModerate - slightHigh (continuing)
9 - PollutionPervasive - largeExtreme - seriousHigh (continuing)
9.1 - Domestic & urban waste waterPervasive - largeSerious - moderateHigh (continuing)
9.3 - Agricultural & forestry effluentsPervasive - largeExtreme - seriousHigh (continuing)
9.3.1 - Nutrient loadsPervasive - largeExtreme - seriousHigh (continuing)
9.3.2 - Soil erosion, sedimentationPervasive - largeExtreme - seriousHigh (continuing)
11 - Climate change & severe weatherPervasive (71-100%)Moderate - slightModerate - low
11.2 - DroughtsPervasive (71-100%)Moderate - slightModerate - low

Roadless Areas (8)
Idaho (1)
AreaForestAcres
Bear CreekCaribou-Targhee National Forest118,582
Illinois (2)
AreaForestAcres
Bay CreekShawnee National Forest120
Burke BranchShawnee National Forest6,231
Kentucky (1)
AreaForestAcres
WolfpenDaniel Boone National Forest2,835
Wisconsin (4)
AreaForestAcres
09011 - Flynn Lake Study AreaChequamegon-Nicolet National Forest5,951
09012 - Round Lake Study AreaChequamegon-Nicolet National Forest3,707
09157 - Chase CreekChequamegon-Nicolet National Forest6,140
09183 - Shoe Lake IslandsChequamegon-Nicolet National Forest7
References (110)
  1. Athearn, H.D. 1992. New records for some species of Alasmidontini. Malacology Data Net, 3(1-4): 90-91.
  2. Baker, F.C. 1902. The Mollusca of the Chicago Area, the Pelecypoda and Gastropoda. Chicago Academy of Sciences Nautral History Survey Bulletin 3(1/2): 417 pp + 36 pls.
  3. Baker, Richard J. (Natural Heritage and Nongame Research Program, MN Department of Natural Resources). 1997. Review and annotation of fish and mussel watershed distribution maps. Review requested by Ruth Mathews, TNC. November 1997.
  4. Barnhart, C., F. Riusech, and M. Baird. 1998. Hosts of salamander mussel (<i>Simpsonaias ambigua</i>) and snuffbox (<i>Epioblasma triquetra</i>) from the Meramec River system, Missouri. Triannual Unionid Report, 16: 34.
  5. Bogan, A.E. and D.D. Locy. 2009. Current distribution of the salamander mussel, <i>Simpsonaias ambigua</i> (Say, 1825), in Pennsylvania. Ellipsaria 11(3):11-12.
  6. Bolton, M. J. 2008. Discovery of a population of <i>Cylonaias tuberculata</i> (Rafinesque), the purple wartyback mussel (Bivalvia: Unionidae), in the Olentangy River, Delaware County, Ohio. Ohio Journal of Science 108(3):44-46.
  7. Buchanan, A.C. 1980. Mussels (naiades) of the Meramec River Basin. Missouri Department of Conservation, Aquatic Series, 17: 1-68.
  8. Burch, J.B. 1975a. Freshwater unionacean clams (Mollusca: Pelecypoda) of North America. Malacological Publications: Hamburg, Michigan. 204 pp.
  9. Call, R.E. 1900. A descriptive illustrated catalogue of the mollusca of Indiana. Annual Report of the Indiana Department of Geology and Natural Resources, 24: 335-535.
  10. Cicerello, Ronald R. (Kentucky State Nature Preserves). 1997b. Review and annotation of mussel watershed distribution maps. Review requested by Ruth Mathews, TNC. September 1997.
  11. Cicerello, Ronald R. (Kentucky State Nature Preserves). 1997c. Review and annotation of mussel watershed distribution maps. Review requested by Larry Master, TNC. June 1997.
  12. Cicerello, R.R. and G.A. Schuster. 2003. A guide to the freshwater mussels of Kentucky. Kentucky State Nature Preserves Commission Scientific and Technical Series 7:1-62.
  13. Clark, C.F. 1988. Some fresh-water mussels from the Red River drainage, Kentucky. Malacology Data Net, 2(3/4): 100-104.
  14. 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.
  15. Clarke, A.H. 1985. The tribe Alasmidontini (Unionidae: Anodontinae). Part II: <i>Lasmigona </i>and <i>Simpsonaias</i>. Smithsonian Contributions to Zoology, 399: 1-75.
  16. Clark, K. H., J. M. Wisor, S. J. Mueller, C. Bradshaw-Wilson, E. W. Boyer, and J. R. Stauffer, Jr. 2021. Status of freshwater mussels (Unionidae) in the French Creek Watershed, USA at the onset of invasion by round goby, <i>Neogobius melanostomus</i>. Water 13, 3064. https://doi.org/10.3390/w13213064.
  17. Cudmore, B., C.A. MacKinnon, and S.E. Madzia. 2004. Aquatic species at risk in the Thames River watershed, Ontario. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2707. 123 pp.
  18. Cummings, Kevin S. (Illinois Natural History Survey). 1997. Review and annotation of mussel watershed distribution maps. Reviews requested by Christine O'Brien, USGS-BRD. May and July 1997.
  19. 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.
  20. 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.
  21. 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.
  22. 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.
  23. 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
  24. Frierson, L.S. 1927. A classified and annotated checklist of the North American naiades. Baylor University Press. Waco, Texas. 111 pp.
  25. Gangloff, M.M. and G.W. Folkerts. 2006. New distributional record for <i>Simpsonaias ambigua</i> (Say) (salamander mussel; Bivalvia: Unionidae) in the Duck River, central Tennessee. Southeastern Naturalist, 5(1): 53-56.
  26. Gordon, M.E. and J.B. Layzer. 1989. Mussels (Bivalvia: Unionoidea) of the Cumberland River review of life histories and ecological relationships. U.S. Fish and Wildlife Service Biological Report, 89(15): 1-99.
  27. Graf, D.L. 2002. Historical biogeography and late glacial origin of the freshwater pearly mussel (Bivalvia: Unionidae) faunas of Lake Erie, North America. Occasional Papers on Mollusks 6(82):175-211.
  28. 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.
  29. Harmon, J.L. 1989. Freshwater bivalve mollusks (Bivalvia: Unionidae) of Graham Creek, a small southeastern Indiana stream. Malacology Data Net, 2(5/6): 113-121.
  30. 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.
  31. Harris, J.L. and M.E. Gordon. 1987. Distribution and status of rare and endangered mussels (Mollusca: Margaritiferidae, Unionidae) in Arkansas. Proceedings of the Arkansas Academy of Science, 41: 49-56.
  32. Harris, J.L., P.J. Rust, A.C. Christian, W.R. Posey II, C.L. Davidson, and G.L. Harp. 1997. Revised status of rare and endangered Unionacea (Mollusca: Margaritiferidae, Unionidae) in Arkansas. Journal of the Arkansas Academy of Science, 51: 66-89.
  33. Harris, J.L., W.R. Posey II, C.L. Davidson, J.L. Farris, S.R. Oetker, J.N. Stoeckel, B.G. Crump, M.S. Barnett, H.C. Martin, M.W. Matthews, J.H. Seagraves, N.J. Wentz, R. Winterringer, C. Osborne, and A.D. Christian. 2009. Unionoida (Mollusca: Margaritiferidae, Unionidae) in Arkansas, third status review. Journal of the Arkansas Academy of Science 63:50-86.
  34. Hart, J. 2012. Freshwater mussel populations of the Monongahela River, PA and evaluation of the ORSANCO Copper Pole Substrate Sampling Technique using GIS interpolation with geometric means. M.S. thesis. Marshall University, Huntington, West Virginia. 138 pp.
  35. Havlik, M.E. and D.H. Stansbery. 1978. The naiad mollusks of the Mississippi River in the vicinity of Prairie du Chien, Wisconsin. Bulletin of the American Malacological Union, 1977: 9-12.
  36. Heath, D. and L. Kitchel. 1997. Review and annotation of mussel watershed distribution maps by the Wisconsin Department of Natural Resources. Review requested by Ruth Mathews, TNC. September 1997.
  37. Hinck, J. E., C. G. Ingersoll, N. Wang, T. Augspurger, M. C. Barnhart, S. E. McMurray, A. D. Roberts, and L. Schrader. 2011. Threats of habitat and water-quality degradation to mussel diversity in the Meramec River Basin, Missouri, USA. US Geological Survey Open File Report 2011–1125. 18 pp.
  38. Ho, E. 2011. Conserving a species by accessing its host: Potential to expand the range of Simpsonaias ambigua into known Necturus maculosus habitats and range. Thesis Part II. University of Waterloo, Ontario, Canada. 10 pp.
  39. Hoffman, A. S., J. R. Robb, and B. E. Fisher. 2014. Recent records for mudpuppies (<i>Necturus maculosus</i>) in Indiana with notes on presumed declines throughout the Midwest. Proceedings of the Indiana Academy of Science 123(1): 1-6.
  40. Hoftyzer, E., J. D. Ackerman, T. J. Morris, and G. L. Mackie. 2008. Genetic and environmental implications of reintroducing laboratory-raised unionid mussels to the wild. Canadian Journal of Fisheries and Aquatic Sciences 65:1217-1229.
  41. Hoggarth, M.A., D.A. Kimberly, and B.G. Van Allen. 2007. A study of the mussels (Mollusca: Bivalvia: Unionidae) of Symmes Creek and tributaries in Jackson, Gallia and Lawrence Counties, Ohio. Ohio Journal of Science 107(4):57-62.
  42. Hove, M. B. Sietman, M. Berg, D. Hornbach, and S. Boyer. 2013. Chippewa River mussel bed holds federally endangered <i>Plethobasus cyphyus</i>. Ellipsaria 15(2):21-23.
  43. Howard, A. D. 1915. Some exceptional cases of breeding among the Unionidae. The Nautilus 29:4-11.
  44. Howard, A.D. 1951. A river mussel parasitic on a salamander. Natural History Miscellanea, 77: 1-6.
  45. Indiana Natural Heritage Data Center. 2006-2007. Personal communication with Jay Cordeiro (NatureServe) about freshwater mussel distribution in Indiana in 2006 and 2007.
  46. Johnson, Paul (Alabama Aquatic Biodiversity Center). 2006. Personal communication with Jay Cordeiro (NatureServe) about the distribution of <i>Simpsonaias ambigua </i>in Tennessee.
  47. Johnson, R.I. 1980. Zoogeography of North American Unionacea (Mollusca: Bivalvia) north of the maximum Pleistocene glaciation. Bulletin of the Museum of Comparative Zoology, Harvard University 149(2): 77-189.
  48. Kost, M. A., Y. M. Lee, P. J. Badra, M. J. Monfils, M. R. Penskar, and H. D. Enander. 2012. Natural features inventory and management recommendations for Port Huron State Game Area. Michigan Natural Features Inventory Report Number 2012-05, Lansing, MI. 82 pp.
  49. Lambert, T. D., L. C. Hinz Jr., and Y. Cao. 2016. Identifying regional priority areas for focusing conservation actions in streams and grasslands: conservation planning. Illinois Natural History Survey Technical Report 2016 (31). 60 pp.
  50. Lee, Y., M. R. Penskar, P. J. Badra, B. J. Klatt, and E. H. Schools. 2011. Climate change vulnerability assessment of natural features in Michigan’s coastal zone – Phase I: assessing rare plants and animals. Michigan Natural Features Inventory Report No. 2011-18, Lansing, MI. 80 pp.
  51. Lefevre, G. and W. T. Curtis. 1912. Studies on the reproduction and artificial propagation of fresh-water mussels. Bulletin of the Bureau of Fisheries 30:102-201.
  52. Master, L. L. 1996. Synoptic national assessment of comparative risks to biological diversity and landscape types: species distributions. Summary Progress Report submitted to Environmental Protection Agency. The Nature Conservancy, Arlington, Virginia. 60 pp.
  53. Mathiak, H.A. 1979. A river survey of the unionid mussels of Wisconsin, 1973-1977. Sand Shell Press: Horicon, Wisconsin. 75 pp.
  54. McDaniel, T. V., P. A. Martin, G. C. Barrett, K. Hughes, A. D. Gendron, L. Shirose, and C. A. Bishop. 2009. Relative abundance, age structure, and body size in mudpuppy populations in southwestern Ontario. Journal of Great Lakes 35(2):182-189.
  55. McMurray, S. E. 2015. Status assessments of <i>Cumberlandia monodonta</i>, <i>Alasmidonta viridis</i>, <i>Simpsonaias ambigua</i>, <i>Lampsilis rafinesqueana</i>, and <i>Quadrula cylindrica cylindrica</i> in Missouri. Missouri Department of Conservation, Columbia, MO. 27 pp.
  56. Metcalfe-Smith, J.L. and B. Cudmore-Vokey. 2004. National general status assessment of freshwater mussels (Unionacea). National Water Research Institute / NWRI Contribution No. 04-027. Environment Canada, March 2004. Paginated separately.
  57. 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.
  58. Michigan Natural Features Inventory. 2007. Personal communication with Jay Cordeiro (NatureServe) about the distribution of <i>Simpsonaias ambigua</i> in Michigan.
  59. MolluscaBase eds. 2024. MolluscaBase. Accessed at https://www.molluscabase.org
  60. Morris, T. J. and M. Burridge. 2006. Recovery Strategy For Northern Riffleshell, Snuffbox, Round Pigtoe, Mudpuppy Mussel and Rayed Bean in Canada.[Proposed]. Species at Risk Act Recovery Strategy Series. Fisheries and Oceans Canada, Ottawa, x + 76 pp.
  61. Moyle, P., and J. Bacon. 1969. Distribution and abundance of molluscs in a fresh water environment. Journal of the Minnesota Academy of Science 35(2/3):82-85.
  62. Oesch, R.D. 1984a. Missouri Naiades: a Guide to the Mussels of Missouri. Jefferson City, Missouri: Conservation Commission of the State of Missouri. 270 pp.
  63. Oesch, R.D. 1995. Missouri Naiades. A Guide to the Mussels of Missouri. Second edition. Missouri Department of Conservation: Jefferson City, Missouri. viii + 271 pp.
  64. 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.
  65. Panozzo, B. 2020. Champaign County Ecological Assessment. Department of Urban and Regional Planning, University of Illinois Champain-Urbana. 18 pp.
  66. Parmalee, P.W. 1967. The freshwater mussels of Illinois. Illinois State Museum, Popular Science Series 8:1-108.
  67. Parmalee, P.W. and A.E. Bogan. 1998. The Freshwater Mussels of Tennessee. University of Tennessee Press: Knoxville, Tennessee. 328 pp.
  68. Pennsylvania Natural Heritage Program. 2007. Personal communication with Jay Cordeiro (NatureServe) about the distribution of <i>Simpsonaias ambigua</i> in Pennsylvania.
  69. Poos, M., A. J. Dextrase, A. N. Schwalb, and J. D. Ackerman. 2010. Secondary invasion of the round goby into high diversity Great Lakes tributaries and species at risk hotspots: potential new concerns for endangered freshwater species. Biological Invasions 12:1269-1284.
  70. Porto-Hannes, I., K. McNichols-O’Rourke, M. Goguen, M. Fang, and T. J. Morris. 2021. Sampling protocol for the freshwater mussel <i>Simpsonaias ambigua</i> (salamander Mussel) in Canada. Canadian Technical Report of Fisheries and Aquatic Sciences 3411: vii + 60 p.
  71. Price, A. L., D. K. Shasteen, and S. A. Bales. 2012a. Freshwater mussels of the Des Plaines River and Lake Michigan tributaries in Illinois. Illinois Natural History Survey Technical Report 2012 (10). Champaign, Illinois. 16 pp.
  72. Price, A. L., D. K. Shasteen, and S. A. Bales. 2012b. Freshwater mussels of the Kankakee River in Illinois. Illinois Natural History Survey Technical Report 2012 (12). Champaign, Illinois. 16 pp. + appendix.
  73. Price, A. L., S. A. Bales, and D. K. Shasteen. 2012c. Freshwater mussels of the Sangamon River. Illinois Natural History Survey Technical Report 2012 (39). Champaign, Illinois. 21 pp. + appendix.
  74. Reid, S. M., and T. J. Morris. 2017. Tracking the recovery of freshwater mussel diversity in Ontario rivers: evaluation of a quadrat-based monitoring protocol. Diversity 9, 5; doi:10.3390/d9010005
  75. Salerno, J., C. J. Bennett, E. Holman, P. L. Gillis, P. K. Sibley, and R. S. Prosser. 2018. Sensitivity of multiple life stages of 2 freshwater mussel species (Unionidae) to various pesticides detected in Ontario (Canada) surface waters. Environmental Toxicology and Chemistry 37(11): 2871-2880.
  76. Salerno, J., P. L. Gillis, H. Khan, E. Burton, L. E. Deeth, C. J. Bennett, P. K. Sibley, and R. S. Prosser. 2020.Sensitivity of larval and juvenile freshwater mussels (Unionidae) to ammonia, chloride, copper, potassium, and selected binary chemical mixtures. Environmental Pollution 256, 113398.
  77. Sargent, Barbara (West Virginia Natural Heritage Program). 1997. Review and annotation of mussel watershed distribution maps. Review requested by Ruth Mathews, TNC. September 1997.
  78. Schneider, Kathryn (New York Natural Heritage Program). 1997. Review and annotation of fish and mussel watershed distribution maps. Review requested by Larry Master and Ruth Mathews, TNC. September 1997.
  79. Shasteen, D. K., S. A. Bales, and A. L. Price. 2012. Freshwater mussels of the Embarras River basin and minor Wabash tributaries. Illinois Natural History Survey Technical Report 2012 (30). Champaign, Illinois. 20 pp. + appendix.
  80. Sherman, Renée A. (University of Michigan, Museum of Zoology). 1997. Review and annotation of mussel watershed distribution maps. Review requested by Christine O'Brien, USGS-BRD. June 1997.
  81. Sietman, B.E. 2003. Field Guide to the Freshwater Mussels of Minnesota. Minnesota Department of Natural Resources: St. Paul, Minnesota. 144 pp.
  82. Simpson, C.T. 1914. A Descriptive Catalogue of the Naiades or Pearly Fresh-water Mussels. Bryant Walker: Detroit, Michigan. 1540 pp.
  83. Smith, T. A., and E. S. Meyer. 2010. Freshwater mussel (Bivalvia: Unionidae) distributions and habitat relationships in the navigational pools of the Allegheny River, Pennsylvania. Northeastern Naturalist 17(4):541-564.
  84. Spoo, A. 2008. The Pearly Mussels of Pennsylvania. Coachwhip Publications: Landisville, Pennsylvania. 210 pp.
  85. Stansbery, D.H. 1970. Eastern freshwater mollusks (I): The Mississippi and St. Lawrence River systems. Malacologia, 10(1): 9-22.
  86. Stapleton, M. M., D. A. Mifsud, K. Greenwald, J. Boase, M. Bohling, A. Briggs, J. Chiotti, J. Craig, G. Kennedy, R. Kik IV, J. M. Hessenauer, D. Leigh, E. Roseman, A. Sedman, J. Sutherland, and M. Thomas. 2018. Mudpuppy assessment along the St. Clair-Detroit River System. Herpetological Resource and Management Technical Report. 110 pp.
  87. Stegmann, E. 2020. Habitat selection and host detection in the salamander mussel, <i>Simpsonaias ambigua</i>. M.S. thesis. Missouri State University, Springfiled, MO. 47 pp.
  88. Stodola, A. P., S. A. Douglass, and D. K. Shasteen. 2014. Historical and current distributions of freshwater mussels in Illinois. Illinois Natural History Survey Technical Report 2014 (37). Champaign, Illinois. 82 pp.
  89. Strayer, D. 1983. The effects of surface geology and stream size on freshwater mussel (Bivalvia, Unionidae) distribution in southeastern Michigan, U.S.A. Freshwater Biology 13:253-264.
  90. Strayer, D. L. 1999. Use of flow refuges by unionid mussels in rivers. Journal of the North American Benthological Society 18(4):468-476.
  91. Strayer, D. L., and J. Ralley. 1993. Microhabitat use by an assemblage of stream-dwelling unionaceans (Bivalvia) including two rare species of <i>Alasmidonta</i>. Journal of the North American Benthological Society 12(3):247-258.
  92. Strayer, D.L. and K.J. Jirka. 1997. The Pearly Mussels of New York State. New York State Museum Memoir 26. The University of the State of New York. 113 pp. + figures.
  93. Trauth, S. E., B. A. Wheeler, W. R. Hiler, R. L. Lawson, H. C. Martin, and A. D. Christian. 2007. Current distribution and relative abundance of the crayfish, mussels, and aquatic salamanders of the Spring River, Arkansas. Final Report to the Arkansas Game and Fish Commission. 85 pp.
  94. 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.
  95. U.S. Fish and Wildlife Service (USFWS). 2011. Endangered and threatened wildlife and plants; partial 90-day finding on a petition to list 404 species in the southeastern United States as threatened or endangered with critical habitat. Federal Register 76(187):59836-59862.
  96. U.S. Fish and Wildlife Service (USFWS). 2023. Endangered and Threatened Wildlife and Plants; Endangered Species Status for Salamander Mussel and Designation of Critical Habitat. Proposed rule. Federal Register 88(161):57224-57290.
  97. U.S. Fish and Wildlife Service (USFWS). 2023. National Listing Workplan. Online. Available: https://www.fws.gov/project/national-listing-workplan
  98. U.S. Fish and Wildlife Service (USFWS). 2025. Endangered and Threatened Wildlife and Plants; Review of Species That Are Candidates for Listing as Endangered or Threatened; Annual Notification of Findings on Resubmitted Petitions; Annual Description of Progress on Listing Actions. Candidate notice of review (CNOR). Federal Register 90(209):48912-48937.
  99. Van der Schalie, H. 1938. The naiad fauna of the Huron River in southeastern Michigan. Miscellaneous Publication of the Museum of Zoology, University of Michigan 40:7-78.
  100. Watson, E.T., J.L. Metcalfe-Smith, and J. Di Maio. 2000. Status of the Mudpuppy Mussel (<i>Simpsonaias ambigua</i>) in Canada. Draft Report for the Committee on the Status of Endangered Wildlife in Canada. 46 pp.
  101. Watters, G.T. 1992b. Distribution of the Unionidae in south central Ohio. Malacology Data Net 3(1-4):56-90.
  102. Watters, G. T. 1992. Unionids, fishes, and the species-area curve. Journal of Biogeography 19:481-490.
  103. Watters, G. T. 1993. A Guide to the Freshwater Mussels of Ohio. Revised Edition prepared for The Division of Wildlife, Ohio Department of Natural Resources. 106 pp.
  104. 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.
  105. Watters, G.T., M.A. Hoggarth, and D.H. Stansbery. 2009b. The Freshwater Mussels of Ohio. Ohio State University Press: Columbus, Ohio. 421 pp.
  106. West Virginia Natural Heritage Program (WV NHP). 2007. Personal communication with Jay Cordeiro (NatureServe) about the distribution of <i>Alasmidonta varicosa</i> and <i>Simpsonaias ambigua</i> in West Virginia.
  107. 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.
  108. Williams, J. D., M. L. Warren, Jr., K. S. Cummings, J. L. Harris, and R. J. Neves. 1993. Conservation status of freshwater mussels of the United States and Canada. Fisheries 18(9):6-22.
  109. Wisconsin Natural Heritage Program. 2007. Personal communication with Jay Cordeiro (NatureServe) about the distribution of <i>Simpsonaias ambigua</i> in Wisconsin.
  110. Womble, K. I., G. R. Dinkins, J. Brian Alford, and M. H. Harris. 2020. New species distribution record for <i>Simpsonaias ambigua</i> (Say) (salamander mussel, Bivalvia: Unionidae) in the Harpeth River, Tennessee. Notes of the Southeastern Naturalist 19(1):24-28.