(DRAFT) - Taxonomy
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
TAXONOMY
NAME - MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
OTHER COMMON NAMES - MUSSEL, PEARLY, BLOSSOM, TUBERCLED-;BLOSSOM and TUBERCLED-
ELEMENT CODE -
CATEGORY - Aquatic Molluscs
PHYLUM AND SUBPHYLUM - MOLLUSCA,
CLASS AND SUBCLASS - BIVALVIA,
ORDER AND SUBORDER - UNIONOIDA,
FAMILY AND SUBFAMILY - UNIONIDAE,
GENUS AND SUBGENUS - EPIOBLASMA,
SPECIES AND SSP - TORULOSA, TORULOSA
SCIENTIFIC NAME - EPIOBLASMA TORULOSA TORULOSA
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Tubercled-blossom Pearly Mussel
Epioblasma torulosa torulosa (Rafinesque, 1820)
KINGDOM: Animal GROUP: Clam
PHYLUM: Mollusca CLASS: Bivalvia
ORDER: Unionoida FAMILY: Unionidae
E. t. torulosa is the inflated big river form of E. torulosa, and
it exhibits considerable ecophenotypic variation (01). It is a
medium-sized species, reaching 90 mm in length, with a shell outline
described as irregularly ovate, elliptical, or obovate (02). Valves
are inequilateral and solid with a short hinge ligament; unbos are
full and turned slightly forward in the anterior third of the shell
(02). The periostracum has many distinct growth lines and is smooth
and shiny, straw to yellow-green in color with numerous fine green
rays. Posterior and medial ridges on the valves may vary from smooth
to possessing elevated knobs.
Sexual dimorphism occurs in this species. The posterior ridge on
male shells is low and narrowly rounded, separated from the medial
ridge by a broad furrow that ends ventrally in an emargination between
the ridges (02). Females are typically larger than males and possess
rounded marsupial swellings extending from the mid-ventral margin to
Taxonomy - 1 (DRAFT) - Taxonomy
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
the posterior end. These swellings are flattened and usually dark
green with occasional, small radial furrows (03). The beak cavity of
the species is shallow, and the pallial line and muscle scars are
prominent. Nacre color ranges from white to salmon-red (02).
The form torulosa apparently graded into the headwater form
gubernaculum in the Tennessee River near Knoxville. Clinal variation
of these two forms has been discussed in early studies, particularly
the relationship between shell inflation, tuberculation, and stream
size (04). Typically, the form torulosa has a row of prominent knobs
along the medial ridge whereas the form gubernaculum had reduced or no
knobs present.
Some controversy still exists among malacologists as to the valid
generic name for this group of species; Epioblasma, Dysnomia, or
Plagiola. All three have been used in the recent literature. A
discussion of these generic names was presented in 1978 (02).
Detailed descriptions and photographs of this species are
included in previous publications (01,02,03).
Taxonomy - 2 (DRAFT) - Status
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
STATUS
Coded Status
Alabama; Federal Endangered
Alabama; Unofficially Listed
Kentucky; Federal Endangered
Kentucky; State Recognized
Tennessee; Federal Endangered
Tennessee; State Listed
West Virginia; Federal Endangered
West Virginia; Unofficially Listed
E: Federal Endangered
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The tubercled-blossom pearly mussel (Epioblasma torulosa
torulosa) has been designated an Endangered species pursuant to the
Endangered Species Act of 1973 (50 CFR 17.11; P.L. 93-205, 87 Stat.
884; 16 U.S.C. 1531-1540), as amended. The species has this status
wherever found including the States of Alabama, Illinois, Kentucky,
Ohio, Tennessee, and West Virginia. The Federal Register listing
included Kentucky, Illinois, Indiana, West Virginia, and Tennessee.
This subspecies is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild animal
(alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
DOD (COE) -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public land under their control. Also responsible for
Status - 1 (DRAFT) - Status
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
management/recovery on Department of Defense lands.
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Alabama and West Virginia
UNOFFICIAL LIST: (Alabama) Extirpated, Bulletin No.2,
Endangered and Threatened Plants and Animals of
Alabama, Alabama Museum of Natural History.
(West Virginia) Federally Endangered, Special
Animal List, Wildlife Resources Division, WV
Department of Natural Resources.
STATE: Illinois, Kentucky, Ohio, Tennessee, and West Virginia
DESIGNATED STATUS: Endangered (IL, TN); Recognized Endangered
(OH, KY, WV)
ADMINISTRATIVE AGENCY: (Illinois) Department of Conservation;
(Kentucky) Department of Fish and
Wildlife Resources;
(Ohio) Department of Natural Resources;
(Tennessee) Wildlife Resources Agency;
(West Virginia) Department of Natural
Resources.
STATE STATUTES: (Illinois) IL Revised Statutes, Chapter 8,
Sec. 331-340, IL Endangered Species Protection
Act of 1972;
(Kentucky) KY Fish and Wildlife Codes, 150.183,
Importing, transporting, or possessing
endangered species of wildlife;
(Ohio) Annotated Code 1501:31-23-01, Eff. July
11, 1980;
(Tennessee) TN Code Annotated, Chapter 9,
Sec. 51-901 through 51-912, Proclamations 75-15,
76-4, and amendments;
(West Virginia) Reorganization Plan Numbered 4
of 1970 as cited in Appendix A of the Limited
Authorities Cooperative Agreement with the U.S.
Fish and Wildl. Service.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
The tubercled-blossom pearly mussel is listed in appendix I of
CITES and is listed as Endangered in the IUCN Red Data Book, 1983.
ECONOMIC STATUSES:
There are no known commercial values for this species, however,
its sedentary nature and filter feeding habits may make it useful as a
water quality indicator. It has been shown that mussels accumulate
Status - 2 (DRAFT) - Status
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
such pollutants as heavy metals and pesticides.
74/10/17:39 FR 37078/37079 - Notice of review
75/09/26:40 FR 44329/44333 - Proposed rule
76/06/14:41 FR 24062/24067 - Final rule
76/06/16:41 FR 24367/42378 - Proposal - CITES I
77/02/22:42 FR 10484/10485 - Final - CITES I
81/02/27:46 FR 14652/14658 - Five year review
Status - 3 HABITAT ASSOCIATIONS
HABITAT - AQUATIC
INLAND AQUATIC
LAND USE -
Streams and Canals
NATIONAL WETLAND INVENTORY CODES
NWI NWICLS NWIMOD NWISPEC
Riverine, upper perennial UB1
Riverine, upper perennial SB1
COMMENTS ON HABITAT ASSOCIATIONS -
E. t. torulosa is a riverine species typically found in rivers
with shallow sand and gravel shoals and rapid current (11). This
species does not occur in the impounded sections of rivers. It is
undoubtedly sedentary, intolerable of silt and water pollution, and
required ample water flow and stable substrate for survival. Eight
species of Epioblasma, now extinct, were recorded from riffles of
large rivers (11).
Habitat of the glochidia is initially within the gills of the
female, then in the water column, and finally attached to a suitable
fish host. Any alteration of these life stage-specific habitats
during its life cycle would affect the survival of remnant
populations. Habitat requirements or associations for the juvenile
stage are unknown. Specific environmental requirements for the adults
are also unknown.
Habitat Associations - 1 (DRAFT) - Food Habits
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
FOOD HABITS
TROPHIC LEVEL -
OMNIVORE
LIFESTAGE FOOD FOOD PART
General Detritus
General Bacteria
General Phytoplankton
General Diatoms
General Zooplankton
General Fish
Food Habits - 1 (DRAFT) - Environment Associations
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Aquatic Features: Pool areas
Environment Associations - 1 (DRAFT) - Life History
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
LIFE HISTORY
FOOD HABITS:
There are no known interspecific differences in feeding among
freshwater mussels (05). The glochidia are probably obligate
parasites on the gills or fins of fish. Adult mussels are
filter-feeders and consume particulate matter suspended in the water
column. Identifiable stomach contents almost invariably include mud,
desmids, diatoms, other unicellular algae, protozoa, and zooplankters
(12,13). No study of the food habits of the tubercled-blossom has
ever been conducted; required/preferred foods are therefore unknown.
HOME RANGE/TERRITORY:
Non-territorial. The species is sedentary during its adult life
and generally occurs in rivers with others of its kind.
PERIODICITY:
No information on this subject exists for E. t. torulosa, but
other Epioblasma exhibit a seasonal periodicity (14). During the
glochidial release period in spring, specimens are readily visible in
the stream bottom. After this period, these species burrow in the
substrate with only siphons exposed.
MIGRATION PATTERNS:
Non-migratory.
COVER/SHELTER REQUIREMENTS:
Based on available reports of Epioblasma spp. (01), the species
probably occurs in river shoals with moderate to swift current and
sand-gravel substrate. No species-specific information on
cover/shelter is available in the literature, except that it was
collected in water varying from inches to about 6 feet deep (15,16,
17).
REPRODUCTIVE SITE REQUIREMENTS:
The tubercled-blossom is a riffle and run species that occurs in
swift flowing, medium-sized streams with stable substrate (01).
Since this species is apparently sedentary and movement is
inconsequential, this is the type of environment where all life
processes would be carried out, including reproduction.
REPRODUCTIVE CHARACTERISTICS:
The reproductive cycle of E. t. torulosa is presumably similar to
other Epioblasma spp. (01). During spawning males discharge sperm
into the water column, and the sperm are taken in by the females
during siphoning. Eggs are fertilized in the female's suprabranchial
cavity or gills, which serve as marsupia for embryos developing to the
parasitic stage (glochidia). Glochidia are released by the female to
attach to a suitable host for metamorphosis to the free-living
juvenile stage.
The tubercled-blossom is probably a long-term breeder (01,16),
spawning in late summer, retaining glochidia through fall and winter,
and releasing glochidia the following spring/summer. The fish host(s)
Life History - 1 (DRAFT) - Life History
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
are unknown.
PARENTAL CARE:
Except for retention of the glochidia within the gills of the
female prior to release, no parental care occurs.
POPULATION BIOLOGY:
No information exists on the population biology of this species.
SPECIES INTERRELATIONSHIPS:
The required fish hosts for the parasitic glochidia of this
species are unknown.
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 2 (DRAFT) - Management Practices
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Developing/maintaining stream structures
Beneficial Maintaining/Controlling Water Flow
Beneficial Controlling/Restricting Mining
Beneficial Controlling/Restricting Road Maintainance Actions
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Controlling pollution [thermal, chemical, physical]
Beneficial Controlling/Restricting Pesticide Use
Beneficial Controlling/Restricting Herbicide Use
Beneficial Controlling/Restricting Agricultural Practice
Beneficial Stocking captive-reared wild-strain animals
Beneficial Transplanting wild animals
Beneficial Transplanting Wild Eggs/Wild Seeds
Beneficial Restricting Poaching
Beneficial Controlling/Restricting Noncommercial Harvest
Adverse Inherent Reproductive Characteristics
Existing Inherent Reproductive Characteristics
Adverse Low Gene Pool
Existing Low Gene Pool
Adverse Underground Mines
Existing Underground Mines
Adverse Surface Mines
Existing Surface Mines
Adverse Siltation
Existing Siltation
Adverse Water Temperature Alteration
Existing Water Temperature Alteration
Adverse Dissolved Oxygen Reduction
Existing Dissolved Oxygen Reduction
Adverse Water Level Fluctuation
Existing Water Level Fluctuation
Adverse Reservoirs
Existing Reservoirs
Adverse Migration barriers
Existing Migration barriers
Adverse Dredging
Existing Dredging
Adverse Applying herbicides
Existing Applying herbicides
Adverse Applying pesticides
Existing Applying pesticides
Adverse Environmental Contamination/Pollution
Existing Environmental Contamination/Pollution
Adverse Erosion
Existing Erosion
Adverse
Existing
Management Practices - 1 (DRAFT) - Management Practices
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
COMMENTS ON MANAGEMENT PRACTICES -
The mussel fauna in most streams of the Tennessee River drainage
has been impacted by dam construction, channelization, siltation, and
water polluation , which directly affect all mussel species (01,05).
The change from lotic to lentic conditions because of dams has also
altered the species composition of the fish community, thus
jeopardizing the reproductive success of some mussel species by
eliminating required fish hosts. Although siltation has been
suggested as the most significant adverse affect of impoundments on
mussels (05), other detrimental factors include lowered water
temperatures, pH changes, oxygen depletion, and dewatering of mussel
beds below dams (05). Mussel deaths can occur in substrate on which
0.6 - 2.5 cm of silt has accumulated, resulting in interference with
feeding and suffocation (06). The siphoning period and metabolic rate
of mussels can also be affected by such contaminants as heavy metals
and agricultural chemicals (07).
Water pollution has probably contributed to the decline of this
species and other Epioblasma spp. Acid mine drainage in tributaries
of the Cumberland River has nearly eliminated the most diverse group
of Epioblasma (08). Numerous tributaries in the upper Tennessee River
drainage were polluted already by the twentieth century, and the
mussel fauna was in a decline at that time (09).
The genus Epioblasma has been considered the most highly
developed and recently evolved group of mussel species, with at least
8 of them that have recently become extinct (10). The decline of E.
t. torulosa and extinction of other Epioblasma can not be explained,
but appears to be symptomatic of a general synecological problem that
exists between Epioblasma and chronic environmental changes that have
occurred and apparently are continuing in eastern rivers. Because
this species is so rare (possibly extinct), the causes for its decline
will remain conjectural.
Substantial mussel die-offs that have occurred over the past 5
years (since 1982) also pose a direct threat to this species. If
such a die-off occurred in the Kanawha River it may well lead to
extinction if the species still exists there.
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1984. Recovery Plan for
Tubercled-blossom, Turgid-blossom and Yellow-blossom Pearly Mussels.
U.S. Fish and Wildlife Service, Atlanta, GA. 39 pp.
Recovery actions for the tubercled-blossom pearly mussel include:
(1) Conduct intensive surveys to determine if the species is still
extant.
(2) Once found, identify essential habitat in need of protection.
(3) Utilize existing regulations to protect the species and its
habitat (e.g., controlling take and other factors adversely affecting
the habitat). Habitat restoration or improvement may be necessary
to improve conditions for survival of the species (i.e., restoration
Management Practices - 2 (DRAFT) - Management Practices
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
of stream banks and channels and assuring optimal clean water flow).
(4) Determine present and forseeable threats to the species and
strive to minimize and/or eliminate them. These negative impacts
typically include dredging, strip mining, industrial development, road
and bridge construction, installation and operation of sewage
treatment plants, and the use of herbicides, pesticides and farm
fertilizers.
(5) Utilize artificial culture techniques to produce juvenile mussels
for reintroduction in augmenting populations.
Except for enforcement of regulations, there are no recovery
efforts presently underway for this species.
Management Practices - 3 (DRAFT) - References
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 U.S. Fish and Wildlife Service. 1983. Agency Draft Recovery Plan
for the Tubercled-blossom pearly mussel [Epioblasma (=Dysnomia)
torulosa torulosa (Rafinesque, 1820)], Turgid-blossom pearly mussel
[Epioblasma (=Dysnomia) turgidula (Lea, 1858)], yellow-blossom
pearly mussel [Epioblasma (=Dysnomia) florentina florentina (Lea,
1857). U.S. Fish and Wildlife Service, Asheville, NC. 38 pp.
02 Johnson, R.I. 1978. Systematics and zoogeography of Plagiola
(=Dysnomia=Epioblasma) an almost extinct genus of freshwater
mussels (Bivalvia:Unionidae) from middle North America. Bull. Mus.
Comp. Zool. 148:239-321.
03 Bogan, A.E. and P.W. Parmalee. 1983. Tennessee's rare wildlife
volume II: the mollusks. Tenn. Wildl. Resour. Agency, Tenn. Dep.
Conserv., and Tenn. Heritage Prog., Univ. Tenn., Knoxville, TN.
123 pp.
04 Ball, G.H. 1922. Variation in freshwater mussels. Ecology
3:93-121.
05 Fuller, S.L.H. 1974. Clams and mussels (Mollusca:Bivalvia).
Pages 215-273 In: C.W. Hart and S.L.H. Fuller (eds.), Pollution
Ecology of Freshwater Invertebrates. Academic Press, NY.
06 Ellis, M.M. 1931. Some factors affecting the replacement of the
commercia, fresh-water mussels. U.S. Bur. Fish. Circ. N0.7.
10 pp.
07 Salanki, J. 1979. Behavioural studies in mussels under changing
environmental conditions. Symp. Biol. Hung. 19:169-176.
08 Neel, J.K. and W. Allen. 1964. The mussel fauna of the upper
Cumberland Basin before its impoundment. Malacologia 1:427-459.
09 Ortmann, A.E. 1918. The nayades (freshwater mussels) of the upper
Tennessee drainage with notes on synonymy and distribution. Proc.
Am. Phil. Soc., Phila. 57:521-626.
10 U.S. Fish and Wildlife Service. 1984. Recovery Plan for the Tan
Riffle Shell Mussel (Epioblasma walkeri). U.S. Fish and Wildlife
Service. Atlanta, GA. 59 pp.
11 Stansbery, D.H. 1971. Rare and endangered mollusks in eastern
United States. Pages 5-18. IN: S.E. Jorgenson and R.E. Sharp
(eds.), Proceedings of a symposium on rare and endangered mollusks
(naiades) of the U.S. Bur. Sport Fish. Wildl., U.S. Fish and
Wildlife Service, Wash., D.C.
12 Coker, R.E., A.F. Shira, H.W. Clark, and A.D. Howard. 1921.
Natural history and propagation of fresh-water mussels. Bull. U.S.
Bur. Fish. 37:77-181.
13 Churchill, E.P., Jr. and S.I. Lewis. 1924. Food and feeding in
freshwater mussels. Bull. U.S. Bur. Fish. 39:439-471.
14 Neves, R.J. 1984. Personal observations. Va. Coop. Fish. Res.
Unit, VPI & SU, Blacksburg, VA.
15 Hickman, M.E. 1937. A contribution to mollusca of east Tennessee.
M.S. Thesis, Univ. Tenn. 165 pp.
16 Ortmann, A.E. 1919. A monograph of the naiades of Pennsylvania.
Part 3. systamatic account of the genera and species. Mem.
Carnegie Mus. 8:1-389.
References - 1 (DRAFT) - References
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
17 Parmalee, P.W. 1967. The freshwater mussels of Illinois. Ill.
State Mus. Pop. Ser. 8:1-108.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 U.S. Fish and Wildlife Service. 1983. Agency Draft Recovery Plan
for the Tubercled-blossom pearly mussel [Epioblasma (=Dysnomia)
torulosa torulosa (Rafinesque, 1820)], Turgid-blossom pearly mussel
[Epioblasma (=Dysnomia) turgidula (Lea, 1858)], yellow-blossom
pearly mussel [Epioblasma (=Dysnomia) florentina florentina (Lea,
1857). U.S. Fish and Wildlife Service, Asheville, NC. 38 pp.
02 Stansbery, D.H. 1976. Naiad mollusks. Pages 42-52. IN:
Endangered and threatened plants and animals of Alabama. Bull.
Ala. Mus. Nat. Hist. No.2.
03 Jenkinson, J.J. 1981. Endangered or threatened aquatic mollusks
of the Tennessee River system. Bull. Am. Malacol. Union
1980:43-45.
04 Stansbery, D.H. 1980. The naiad mollusks of the Kanawha River
below Kanawha Falls with special attention to endangered species
(Bivalvia:Unionoida:Unionidae). Tech. Rep., Ohio State Univ. Mus.
Zool., Columbus, OH. 16 pp.
05 Clarke, A.H. 1982. Survey of the freshwater mussels of the upper
Kanawha River (RM 91-95), Fayette County, West Virginia, with
special reference to Epioblasma torulosa (Rafinesque) and Lampsilis
abrupta (Say) [=Lampsilis orbiculata (Hildreth), of authors].
Final Rep., U.S. Fish and Wildlife Service, Newton Corner, MA.
45 pp.
06 Taylor, R.W. 1983. A survey of the freshwater mussels of the
Kanawha River. Final Rep., Corps of Engineers, Huntington, WV.
62 pp.
07 Stansbery, D.H. 1976. Status of endangered fluviatile mollusks in
central North America: Epioblasma turgidula (Lea, 1958). Ohio
State Univ. Res. Found. Rep., U.S. Fish and Wildlife Service, Bur.
Sport Fish. Wildl. 12 pp.
08 Bogan, A.E. and P.W. Parmalee. 1983. Tennessee's rare wildlife
volume II: the mollusks. Tenn. Wildl. Resour. Agency, Tenn. Dep.
Conserv., and Tenn. Heritage Prog., Univ. Tenn., Knoxville, TN.
123 pp.
09 Ortmann, A.E. 1925. The naiad fauna of the Tennessee River system
below Walden Gorge. Am. Midl. Nat. 9:321-372.
10 Stansbery, D.H. 1972. A preliminary list of the naiad shells
recovered from the Buffalo site. Rep. archeol. Invest. No.5, WV.
Geol. Econ. Surv., Morgantown, WV.
11 Johnson, R.I. 1978. Systematics and zoogeography of Plagiola
(=Dysnomia=Epioblasma) an almost extinct genus of freshwater
mussels (Bivalvia:Unionidae) from middle North America. Bull. Mus.
Comp. Zool. 148:239-321.
12 Isom, B.G., P.Yokley, Jr., and C.H. Gooch. 1973. Mussels of the
Elk River Basin in Alabama and Tennessee 1965-1967. Am. Midl. Nat.
89:442-446.
13 Morrison, J.P.E. 1942. Preliminary report on molluscan fauna in
the Tennessee River Valley. Bur. Am. Ethol. Bull. 129:337-392.
References - 2 (DRAFT) - References
Species MUSSEL, PEARLY, BLOSSOM, TUBERCLED-
Species Id ESIS404015
Date 13 MAR 96
14 van der Schalie, H. 1939. Additional notes on the naiades
(freshwater mussels) of the lower Tennessee River. Am. Midl. Nat.
22:452-457.
References - 3