r,

of the
FLORIDA STATE MUSEUM

Biological Sciences
Volume 16 1972 Nu mber 4

THE UNIONIDAE (MOLLUSCA: BIVALVIA} OF

PENINSULAR FLORIDA

Richard I. Johnson

1 1. -'I , , . ,1 -t, 4
5' 1

I. r .
-

_ 1 1,-4 · + I - .-

4 4, 1 4,1-1 - 1 . 4

UNIVERSITY OF FLORIDA GAINESVILLE



Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, Bio-
logical Sciences, are published at irregular intervals. Volumes contain about
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Publication date: 5 June, 1972 Price: $1.40



THE UNIONIDAE (MOLLUSCA: BIVALVIA) OF
PENINSULAR FLORIDA

RICHARD I. JOHNSON

SYNOPSIS: This is a study of the Unionidae, or fresh-water mussels, of penin-
sular Florida, considered here as that region below the Suwannee River sys-
tem in the west and the St. Marys River system in the east. As thus defined
the area has a unionid fauna of 12 species belonging to 6 genera; 9 of these
species also occur in the Apalachicolan region to the west and north where
there are 49 species in 17 genera; 2 of these also occur in the Southern At-
lantic Slope region, which has 37 species and 11 genera. One species is clear-
ly of Southern Atlantic Slope origin, while two others are endemic. The pau-
city of the fauna and distribution of the species give credence to the geological
evidence that most, of peninsular Florida was inundated sometime during the
Pliocene or early Pleistocene, and that it has since been repopulated by
Unionidae mostly from the west and north.

TABLE OF CONTENTS
Introduction .................................. ..182
Acknowledgements ........ .. .. ..... ..... 182
Geology and Topography of Peninsular Florida ... 183

Geology and Topography ................ 183
Marine Shore Lines . ........... . 183
Drainage Systems ...... .... .. ... 186

Distribution of the Florida Unionidae . 191
Regions ......... ... ..... . .. .191
Generic Affinities ................ .... .. 194
Summary and Analysis of Distribution . .. 194

A Revision of the Florida Unionidae ... .. .. 196
Systematic Section .. . ................ . 198

Elliptio .. ······199 Carunculina .. 230
Uniomerus ..........220. Villosa .. 234
Anodonta ....... 225 Lampsilis , 242

Bibliography ......... ..,. ... 244
Index to Relevant Unionid Taxa ............... .247

Richard I. Johnson is an Associate in Malacology at the Museum of Compara-
tive Zoology, Harvard University, Cambridge, Massachusetts. Manuscript ac-
cepted 10 May 1971.- Ed.

Johnson, Richard I. 1972. The Unionidae (Mollusca: Bivalvia) of Peninsular
Florida. Bull. Florida State Mus., Biol. Sci., Vol. 16, No. 4, pp. 181-

181



182 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No.. 4

INTRODUCTION

In peninsular Florida the Unionidae offer two advantages as a group
of animals for zoogeographie study. There are a reasonable number of
species, most of which are clearly distinguishable; they have a limited mode
of distribution, being unable to pass over land from one drainage system
to another. Their ability to move between drainage systems is dependent
on the mobility of fishes to which the glochidia attach themselves. For this
reason the distribution of the species of Unionidae may afford evidence of
former stream confluences and of lowland flooding in the coastal regions
that were, or are. reduced to base level.

The Unionacea of most of the Apalachicolan region were studied by
Clench and Turner (1956). They supported the idea that the fauna was
distributed by mechanical means, though th.Qy were vague as to what
these mechanical means might have been.

Clench and Turner described 32 of the 49 species of Apalachicolan
Unionacea. The distribution of this fauna was reinterpreted (Johnson,
1970), and the Southern Atlantic Slope species were revised. The distri-
bution of these faunas gave evidence of a former confluence of the head-
waters of the Alabama-Coosa, Apalachicola, and Savannah river systems.
Exception was taken to Clench and Turner's theory of distribution so
far as the Unionacea are concerned, as the zoogeographic evidence failed
to show that unionid distribution is fortuitous. In peninsular Florida the
unionids offer no examples of stream capture that can be demonstrated
specifically, but their distribution does not indicate that it occurred by
any passive agents other than natural fish hosts.

ACKNOWLEDGEMENTS

I am grateful to the following people for allowing me to examine the col-
lections under their care, and for permitting me to borrow all relevant types:
Harald A, Rehder, Joseph Rosewater and Joseph P. E. Morrison, United
States National Museum; R. Tucker Abbott, Delaware Natural History Mu-
seum, Greenville (formerly of the Academy of Natural Sciences, Philadelphia);
Henry van der Schalie and John B. Burch, Museum of Zoology, University of
Michigan; Joshua C. Dicki»son, Jr,, Florida State Museum, Gainesville,
Florida.

Herbert D. Athearn, Cleveland, Tennessee; William H. Heard, Florida
State University, Tallahassee; and Fred G. Thompson, F16rida State Mu-
seum, Gainesville, are gratefully acknowledged for material, presented to the
Museum of Comparative Zoology, which proved useful in this study.

Finally, thanks are offered to Kenneth J. Boss, M.K. Jacobson, and
Robert C. Bullock who read the manuscript.



1972 JOHNSON: FLORIDA UNIONIDAE 183

GEOLOGY AND TOPOGRAPHY OF PENINSULAR FLORIDA AS
RELATED TO THE UNIONID FAUNA.

GEOLOGY AND TOPOGRAPHY
Present day peninsular Florida occupies only part of a much larger

unit, the Florida Plateau. This platform is nearly 500 miles long and
from 250 to 400 miles wide. The Plateau is part of the continent and
is probably an extension of the metamorphic rocks of the Georgia Pied-
mont that are buried under some 4,000 feet of sedimentary rocks that
are mostly limestone.

During the millions of years it has been in existence, the Plateau
has been alternately dry land or covered by shallow seas. It appears to
be one of the world's more stable areas. There is no faulting, with just a
slight doming in the north central portion. The Plateau is nearly level,
the highest part (near Haines City, Polk County) being little more than
325 feet above sea level. Nearly two-thirds of the state is below the 50-
foot contour.

MARINE SHORE LINES
Cooke (1945: 248) recognized seven Pleistocene shore lines in penin-

sular Florida, but MacNeil (1950:99), basing his identification of ma-
rine shore lines on the coexistence of shore-line searps, regarded only
four of these as peaks of Pleistocene flooding (Table 1)  Russell (1957:
427-428), On the basis of the complete melting of the polar ice, cast
doubt on the extent of Pleistocene flooding, as did Oaks and Coch
(1963) on the basis of cores made in Virginia. They postulated six
cycles of Pleistocene seas with maximum heights 45 feet above present
levels.

The highest recognized marine shore line in peninsular Florida is
at a level of from 215 to 270 feet (depending on the authority) above
the present one. All of ·Florida was inundated except for several small
islands in the vicinity of Polk County. Cooke (1945: 273, fig. 43)
thought this flooding, which 16rmed the Brandywine terrace (Citronelle
formation in the southeast), took place in the early Pleistocene during
the Aftonian interglacial stage, but Alt and Brooks (1965: 408), on the
basis of new geological evidence, concluded that this flooding took place
during the Upper Miocene. Laessle (1968) later confirmed this dating
with botanical evidence. It is not possible to tell if any of the present
fresh-water mollusks have persisted since the Upper Miocene. 1

The highest Pleistocene shore line recognized by MacNeil (1950,
pl. 1), the Okefenokee or Sunderland of Cooke (1945, 278, fig. 43), not
specifically recognized by Alt and Brooks (1965) or Alt (1968), was formed.
sometime during the Pliocene when the sea level was 150 feet higher

' Orange (Ocall) Island referred to by Clench and Turner (1956. 1()4) was a land mass
separated fron, the continent by the Suwannie Strait during the late Oligocene (Vaughn,
1910: 156) and its existence appears to have no bearing on the present mollusean fauna.



TABLE 1.- CORRELATION OF MARINE SHORE LINES
184 

B
U

LLE
T

IN
- F

LO
R

ID
A
 S

TAT
E
 

M
U

S
E

U
M
 

V
 

X
V

I 
N

o. 
4

Alt and Brooks (1965) and Alt (1968) Cooke)1946) MacNeil.(1950)

Age Altitude Shore Line Altitude Shore Line Altitude Stage
(feet) (feet) (feet)

Sangamon 5-10 (Silver Bluff) 5 Silver Bluff 8-10 Post-Wisconsin
Yarmouth interglacial 25-30 Pamlico 25 Pamlico 25-35 Mid-Wisconsin glacial

recessionl
Aftonian interglacial 45-50 Talbot 42
Late Pliocene or

Early Pleistocene 70-80 Penholoway 70 Wicomico 100 Sangamon interglacial
Pliocene 90-100 Wicomico 100

Sunderland 170 Okefenokee 150 Yarmouth intel·glacial
Upper Miocene 215-250 ~-Coharie 21L

~Brandywine 270 (not recognized) Aftonian interglacial



1972 JOHNSON: FLORIDA UNIONIDAE 185

than the present level. All that remained of the peninsula was part of
Trail Ridge, which formed a large, pear-shaped promontory in Bradford
and Clay Counties; there were three irregular, roughly parallel ridges in
Polk and Highlands Counties in Central Florida; high hills between
Dade City and Brooksville in Pasco and Hernando Counties stood as
islands; to the north numerous Small hills stood above the 150 foot level;
and a large expanse of rocks of the Hawthorne Formation formed an is-
land farther north in Alachua County.

The Wicomibo Shore line (Cooke, 1945: 281, fig. 44) is the least sharply
defined of the shores recognized by MacNeil, which might indicate that the
sea stood at this level» for a comparatively short time. It was formed during
the Pliocene (Alt, 1968: 92) when the sea levelwas 90 to 100 feet higher
than the present level. Florida was again reduced to a number of islands
in Pasco, Hemando, Citrus, Sumter, and Marion Counties. Hubbell
(1954. 48, 49 [inl Olson et al; 1956: 86), in sophisticated papers on the
flightless dung beetle, genus. Mycot~i:pes, concluded on zoogeographical
evidence that the five species that now live on "islands" of sandy plains
6r hills separated by marshes or other nonsandy habitats, evolved on ac-
tual islands in the interglacial seas and that some land areas persisted in
Florida throughout the Pleistocene. Swift (1970: 325) said of a total pri-
mary freshwater fauna of 47 species that now inhabits either the St. Johns
or Suwannee rivers or both, "Only three species of primary fresh water
fishes apparently arose in south or central Florida and supported the
view of moderate Pleistocene flooding. Thompson (1968: 15), on the basis
of the distribution of 35 species of Floridian Hydrobiidae, a group of fresh
and brackish water snails, suggested that a peninsula persisted through-
out the Pleistocene and refuted the marine origin of any of the teri·aces
other than the Pamlico. My interpretation of his data, (with the exception
of the two species Hylalopyrgus brevissimus (Pilsbry ) and H. aequico-
status (Pilsbry) that may have had refugia, or else speciated on islands in
the Wicomico sea) is that many of the species migrated into peninsular
Florida after Wicomico flooding. A number of the species have not fully
occupied the older part of the peninsula, nor penetrated beyond the Pam-
lico Terrace, which suggests a rather recent repopulation probably from
the west and north. Among the 12 species of Unionidae, aside from Ellip-
tio buckle!~i (Lea), which may have persisted on the peninsula prior to
the Pliocene 1 , and Villosa amygdala ( Lea ), which may have speciated
from V. lienosa (Conrad)  on one of the larger land masses, the remaining
species have repopulated the peninsula since Wicomico flooding.

The Pamlico Shore Line (Cooke, 1945: 297, fig. 47), the best pre-
served of the Pleistocene shores, was formed during the Yarmouth inter-
glacial stage when the sea was 20 to 30 feet above present levels. At this
time the shape of Florida was much as it is today, except that the

1 Unio caloo.wzen.%-iN Dall (1895. Trans. Wagner Free Inst. Sci„ 3 (3): 688, pl. 2.3, figs.
5, 12b found iii the Pliocene mari. of the Caloosahatchie River) is either E. buekle!/1
C Lea) or very close to it.



186 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

peninsula was narrower and shorter, terminating near Lake Okeechobee.
Off the southwestern end of the peninsula was a large oval island. A long,
wide lagoon, including the present St. Johns River, extended southward
from Orange Bluff on St. Marys River to Sanford, and was separated
from the open ocean by a chain of large islands.

The shore extended much farther out on the continental shelf as
little as 11,000 years ago (Emery, 1967, fig. 9). At that time it may have
been easier for Unionidae to disperse along a largely baseleveled coast,
which might explain the presence of one unionid, Elliptio dariensis (Lea),
found Only in the  Altamaha and St. Johns river systems. The distribution of
some species of Hydrobiidae (Thompson, 1968), presently restricted to
the ocean side of the Pamlico shore, offers striking evidence of repopulation
and rapid speciation in this area.

DRAINAGE SYSTEMS
Peninsular Florida  (Figure 1) averages over 50 inches of rain a year.

Much of this sinks into the ground, as the soil is loose and sandy, and is
stored up as a great reservoir of ground water, some of which seeps to the
surface in artesian springs. These springs usually rise through deep vertical
holes in the underlying limestone and result from rain that fell on a higher
level. Most of the isolated springs have no Unionidae in them, but those
that form the sources of rivers often have at least Elliptio icterina (Conrad)
or E. buckle!/i (Lea). Many of the springs contain endemic species of Hy-
drobiidae (Thompson, 1968).

Wherever the surface of the ground dips below the water table, lakes
are formed and, when there is an outlet at a lower level, water flows away
as a surface stream. Many lake basins are the result of the dissolution of
underlying limestone, though some occ"py former sea floor depressions.

Particularly in the highlands the landscape is dotted with solution
impressions. Some of these basins lie between the limits of fluctuation of
the water table, and while they contain water in the wet seasons, during

FIGURE 1, Drainages of peninsular Florida and relevant ones in the Apalachi-
colan and Southern Atlantic Slope regions. The major drainage areas of peninsular
Florida are indicated by the dashed lines (after U.S. Dept. Interior, Geol. Survey.
1960. Water Supply paper 1304, pl. 1).

APALACHICOLAN REGION: 1. Apalachieola River, 2. Ochlockonee R., 3, St.
Marks R., 4. Aucilla R., 5. Econfna R.,6. Suwannee R., 7. St. Marys R.,
8. Satilla R.

SOUTHERN ATLANTIC $LOPE REGION: 9. Altamaha River,
PENINSULAR FLORIDA REGION: 10. Waccasassa River, 11. Withlacoochee R.,

12. Pithlachascotee R., 13. Hillsborough R.,.14. Atafia R., 15. Myakka R.,
16. Peace R.

KISSIMMEE RIVER SYSTEM AND EVERCLADES: 17. Caloosahatchee River, 18.
Fisheating Creek, 19. Lake Okeechobee, 20. Istokpoga R., 21, Kissimmee R.

ST. JoHNs RIvER SYSTEM: 22. Econlockhatchee River, 23. Wekiva R., 24.
Oklawaha R., 25. Haw Creek, 26. Rice Cr., 27. Black Cr., 28. Julington Cr., 29.
St. Johtis R



1972 JOHNSON: FLORIDA UNIONIDAE 187

times of drought they may be completely dry. Many of the lake basins
are simple sinks that have always been tributary to the groundwater sup-
ply, while others at one time or another have formed part of the surface
drainage and are therefore connected with the river systems. The history
of the lakes is complicated by the fluctuations of the sea level during the
Pleistocene . A number of presently isolated lakes contain Elliptio buckleyi

4--1

9

7
--.-4-

29
28

3
4 5  '~ 26

1 2 : :,0 0 25

\ 24 1\10
23

iT ",0 2213
1 ,:.¥,0'20 1A

15



188 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

(Lea). Specimens of this species from some of the larger sandy-bottomed
lakes, such as Griffin, Eustis, Harris, and Dora that are subject to rather
heavy wave action, show considerable ecophenotypic variation and were
named Unio cunninghami by Wright.

SUWANNEE RIVER SYSTEM. -This systein originiztes in soutliwester'n
Georgia and flows over a large portion of the Florida pehinsula, but the
unionid fauna is Apalachicol:in (Johnson, 1970: 267,269). The headwaters
are above maximum Pliocene flooding, and some species may have had
refugia there during that time. At present the S.uwiumee River has its head-
w~ters on the Sunderland or Okefenokee ternice in the very acid Okefeno-
kee Swamp. No unionids are known from the Suwannee above its con-
Huence on the, west with the Withlacoochee River (not of the Withla-
coochee system described below), a sandy-bottomed stream. On the east
the Suwannee is joined by the Santa Fe, which with its principal
tributary, the New River, drams much of the north central part of the
peninsula.

WACCASASSA RIVER SYSTEM. -- The Waccasass:1 River, with its largest
tributary; Otter Creek, is a small system in Levy Cotility. It is poorly con-
nected with the swamps and ponds of Gilchrist County, and is the first
entirely peninsular system flowing into the Gulf of Mexico. These spring-
fed streams flow over limestone , and Elliptic) icterinct (Conrad) and Vil /osa
uibex (Conrad) kire the only Unionidae found iii them.

WITHLACOOCHEE RIVER SYSTEM. -The Withlacoochee River origi-
nates in Polk County and follows a generally northerly course past Lake
Tsala Apopka and enters the Gulf at latitude 29'N. The lake connects in
several places with the river, which flows northwestward along its north-
ern edge; it has a very intricate shoreline and contains many islands. There
is no 1:urge expanse of water. At the beginning of the Pleistocene, during
the Aftonian interglacial stage when the sea- was some 42 feet higher, it
was probably a broad, partly silted bay or estuary, separated from the Gulf
by narrow straits at Dunnellon. On the eastern side of the river is Lake
Paiiasoffkee, a large body of water connected to the river. Vaughan (1910:
149) suggested that the Withlacoochee River may have captured the head-
waters of the Hillsborough and Oklawaha rivers. If this is true, any ex-
change of the fauna must have occurred before Lampsilis teres (Rafin -
esque) reached this system, as this is now its southern terininus.

SMALLER WEST COAST DRAINAGE SYSTEMS. -In Citrus, Hernando,
Pasco, and Pinellas Counties are a number of small streams and ponds of
recent origin that do not belong to any general, well defined system. In
Hernando County the very short Weekiwachee River is formed by Weeki-
wachee Springs. It contains only Uniomerus tetralasmus (Say). Further
south, in Pasco County, is the Pithlachascotee River, ' which contains
Elliptio jagensis (Lea ) and U. tetralasmus. In Lake Jovita (Clear Lake ),
Pasco County, and Lake Tarpon at Tarpon Springs, Pinellas County,
only Elliptio buckleyi (Lea) occurs.



1972 JOHNSON: FLORIDA UNIONIDAE 189

Hillsborough County contains several small systems, including the
Hillsborough, Alafia, and Little Manatee river systems, all of which dis-
charge into the Gulf in the vicinity of Tampa. As can be seen from Table
2, the fauna of the Hillsborough River is sufficiently extensive to suggest
stream capture with the Withlacoochee River.

The Alafia River system appears to contain only three species,
Elliptio icterina (Conrad), E. ja!/ensis (Lea), and Uniomems tetralasmu.5
(Say), while no unionids have yet been found in the Little Manatee or
Manatee river systems.

Below the Manatee River, in Manatee County, is another small
system, the Myakka, which contains six species.

PEACE RIVER SYSTEM. - The l,eike River system rises in Pc)lk County
among a number of lakes that it drains on the wests ide of the 150-foot con-
tour. It flows from north to south :incl enters the (:illf at Charlotte Harbor.
As Table 2 shows, its fauna is ver.y simil:ir to that of the Myakka River.

KISSIMMEE RIVER SYSTEM AND EVERGLADES. -Tile Kissiminee River,
whose headwaters are a number of large lakes in Oranke·County, and its
principal tributary, the Istokpoga River, drain into Lake Okeechobee.
Cooke (1939: 107) suggested that Lake Istokpoga, Lake Kissimmee and
perhaps other lakes of the Kissimmee Basin such as Lake Okeechobee,

TABLE 2.-DISTRIBUTION OF UNIONIDAE IN PENINSULAR FLORIDA

AND ADJACENT REGIONS.

Drainages Species'

1 2 3 4 5 6 7 8 9 10 11 12

Apalachicolan X XXXXXXXX
Waccasassa X X
Withlacoochee XXX XXXXXX
Hillsborough X XXXXXXX
Myakka X XX XXX
Peace XXXXX XXXX
Kissimmee and

Everglades XXXXX X X
St. Johns XXXXXX XXXX
St. Marys X X X
Altamaha X X XX X

11. Elliptio dariensis; 2. E. icterina; 3. E. buckleyi; 4. E. jagensis, 5. Unio-
merus tetralasmus; 6. Anodonta couperiana; 7. A. peggyae; 8. Carunculina
parva; 9. Villosa uillosa; 10. V. vibex; 11. V. am!/~dala; 12. Lampsilis teres.



190 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

may have been hollows ih the [Pliocene] sea bottom; in any event, the
formet :fre on higher ten:~ces than the latter, and older.

Lake Okeechobee, one of the larger lakes in North America, is from
25 to 31 miles across. The lake is very shallow, and was not more than
15 to 20 feet deep before canals and dikes were built. Its principal out-
let is now the Ca106sahatchee River, which dfains westward to the Gulf
at Fort Myers. In former times during the rainy season it often over-
flowed its southern bank, which was only a few inches higher than the
surrounding country, and flowed south over the Everglades.

ST. JOHNS RIVER SYSTEM. --The St. J~ihIst River is unique among the
rivers of the United States, as it flows from south to north for nearly 2()0
miles, :ind its headwaters are less than 20 feet above sea level. It is a rela-
tively new river - its upper valley al)ove Lake Harne>, did not come into
existence until the late Pleistocene when a barrier island, now the east
bank of the river, accutnulated in the Panilico Sea. From Lake Hellen
B l aze s t h e r i v e r wan de rs t h ro u gh gras s y 111 a t s h e s b r o ke n by th e e x p a n s i o n s
of Lakes Sawgrass, Washington, Winder, and Poinsett. Between Lakes
Harney ariel George the river ch:innel is little more than 100 yards wide and
is 8 to 20 feet cleep, broken b>, the expansi(ms of Lake Mom·oe about 8 feet
deep, Lake Beresford 5 to 10 feet deep, and Lake Dexter 2 to 10 feet deep.
At Lake George the river expands to :1 width of 6 to 7 miles and 111£11,itains
a remarkably uniform depth of 9 to l l feet for all ofits 11-mile length. Tidal
effects are still felt over 103 miles from the mouth. At Palatka the tidal
range is still 2.5 feet. Beck (1965: 118) pc,inted out thatthe degree of salin-
ity varies -in the river unexpectedly from the discharge of mesohaline
springs in Marion County and furthey noted that, "this stream cannot be
included in any of the designated types, as it has reaches of swamp-:ind-
bog characteristics, others that have sand-bottomed characteristics, mid
stretches not comparable to either. The chemical characteristics of this
river defy summarizing, for anything reported for one i-each would be un-
true of reaches a few miles upstream or downstream." In any event, the
salinity is sufficient for a brackish water mactrid bivalve, Rangia cuileata
(J. E. Gray), to live us far upstream as L:ike Hai Iiev, Seminole County.
Eltilitio jiileizsis ( Lea) and Uniomerns tetra /asnitis (Say ) attain great size
in the big shallow lakes. Many of the specie.4 of Elliptic, show considerable
ecophenotypic variation. Aniong the springs th;it flow directly into the St.
Johns River are Benson's niiner:11 spring and Blue Springs, both in Volusia
Cozinty, mid Alexander Springs in Lake County. All contain endemic
species of Hydrobiidae.

WEKIVA RIVER DRAINAGE. -Among the smaller rivers that drain into

the St. Johns River is the Wekiva River of Orange and Seminole Counties.
This sinall system has its sources in Seminole and Rock Springs in Orange
County and in Wekiwa [fic] and Sanlando Springs in Seminole County.
These are all sulphur springs, :md each supports endemic species of Hy-
drobiidae. Specimens of Elliptio icterina (Conrad) from these slirings



1972 JOHNSON: FLORIDA UNIONIDAE 191

tend to be heavy, inflated, and produced basally with a golden perio-
stracum, but specimens in the spring runs, become compressed and sub-
1-liomboidal, ending in a broad biangulation below the·medial line with the
periostracizifii usuall> s·ellowish or brownish, sometimes with green rays.

OKLAWAHA RIVER DRAINAGE-The largest tributary oftlie St: Johns
River is the Oklawaha, which enters from the west between Lake George
and Palatka where the water is still tidal. Its principal source is the group
of large lakes that includes Griffin, Eustis, Harris, and Dora. all mostly
in Lake Cozint>·, and which are separated from one another by peaty muck.
The· Oklawalia is joined by Orange Creek whose source is Orange Lake in
Alachua County. Cooke (1939: 110) suggested th.it some of the headwaters
have been in existence ever since the Sunderland teri·ace [Upper Mio-
cene] emerged .from the sea, though most of the lakes and all of the lower
reaches are of more.recent origin.

BLACK CREEK DRAINAGE.-This small system enters the St, Johns
River from the west between Palatka and Jacksonville. It appears to be of
Pleistocene origin :is it flows over the Talbot and Pamlic·o terraces. The
unionid fauna is now effectively separated froni the St. Johns by saltwater.
Specimens of Elliptio icterina (Coniad ) closely resemble those of the St .
Marys River, the next system to the north, whose waters are rather iwid.
Black Creek contains Elliptic) dariensis (Lea) which is found in the Alta-
Illalla River, Georgia, to the north, but not in the intervening.St. Marys (,1
Satilla river systenis.

JULINGTON CREEK DRAINAGE.-This small s>stem enters the St.
Johns River from the east a few nfiles soutli of Jacksonville, It is of recent
origin like the St. Johns itself. Its uni(mid fatin:, is remarkable because of
the large size that individual specimens attain.

DISTRIBUTION OF THE FLORIDA UNIONIDAE

REGIONS

In order to understand the distribution of the Unionidae of penin-
sular Florida and to emphasize the paucity of species found there, a few
remarks on the unionid fauna of the regions to the west and north are
needed.

The Apalachicolan region has been generally regarded by previous
authors (H. and A. van der Schalie, 1950: 450; Clench and Turner, 1956)
as consisting of the river systems from the Escambia to the Suwannee
that· flow into the Gulf of Mexico. Although they flow into the Atlantic
Ocean, the St. Marys and Satilla river systems ate now also included in
this region, as their modest unionid faunas consist entirely of species
found in the Apalachicolan region, the dominant species in them being
Elliptio crassidens crassidens (Lamarck) (Johnson, 1970: 305) and E. c.
downiei (Lea) (Johnson, 1970:307) respectively. The former is abundant
in the Interior Basin and is found in the Alabama-Coosa and Aoalachi-



192 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

cola river systems. It is missing in the Ochlockonee and Suwannee river
systems and from peninsular Florida and the Southern Atlantic slope
regions, both as defined below. E.c. crassidens' appears in the Pliocene of
peninsular Florida, indicating its presence in this general area for a long
time.

The Apalachicolan region as defined above has a unionid fauna of
49 species. Of these, 19 have affinities with species to the west; 9 are en-
demic to the region; 3 others extend into peninsular Florida; and 11 more
are restricted to individual river systems. Three species that were probably
once endemic to the Apalachicolan region have spread into the Atlantic
Slope region, and four Atlantic Slope species have spread in the opposite
direction through a onetime confluence of the headwaters of the Apalachi-
cola and Savannah river systems (Johnson, 1970: 268, text fig. 1)

To the east and north of the Apalachicolan region is the Southern
Atlantic Slope region, which extends from the Altamaha River system in
Georgia to the James River system in Virginia. This fauna contains 37
species, of which 4 are found iii the Apalachicolan region as well.

The peninsular Florida region, defined here ds a separate region, is
that area below the Suwannee River system in the west and the St. Marys

' Described as : Elliptio pochur){!cm Pilsbr; 1953 [ in ] Ols .son . A . A . and A . Harbison .
Pliocene Molt. Southern Florida. Ac·ad. X,it. Sci, Phila. Monog. 8 p. 447. PI. 65, fig. 8
(St. Petersburg [Pinellas Co.]. Floricla. h„int>·lie ANSP 18.588).

FIGURE 2, Species of Unionidae that appear to have migrated into peninsular Flor-
ida subsequent to maximum Pliocene.flooding, mostly from the west.

A. Uniomerus tetralasmus (Say). Widely distributed in the Interior Basin, West
Gulf Coastal region, Alabama-Coosa River system, and Apalachieolan region: Rio
Grande River system, Texas, east to the Suwannee River system; Florida; Peninsu-
lar Florida; Southern Atlantic Slope: Altamaha River system, Georgia,·north to the
Nottaway River of the Chowan River system, North Carolina. As this species is ab-
sent in the two intervening river systems between the St. Johns and the Altamaha,
it probably spread into Florida from the west.

B. Villosa Cibex (Cbnrad). West Gulf Coastal region, Alabama-Coosa River
system and Apalachicolan region: Pearl River system, Mississippi, east to the Su-
wannee River system, Florida; Peninsular· Florida; Southern Atlantic Slope: Alta-
maha River system, Georgia, north to the coastal ponds of the Cape Fear River
system, North Carolina. As this species is absent in the two intervening river sys-
tems between the St. Johns and Altamaha, it probably spread into Florida from
the west.

C. Elliptic (Elliptio) icterina (Conrad). Apalachicolan region: Escambia River
system, Florida, east to the St. Marys River system, Georgia, Peninsular Florida;
Southern Atlantic Slope: Altamaha River system, Georgia, nofth to the White Oak
River, North Carolina. The ecophenotypic variation in this species suggests that it
entered Florida from both the west and north.

D. Carunculina prirva (Barnes). Widely distributed in the Interior Basin,
Apalachicolan region, and Peninsular Florida. It is replaced in the Southern AtIan-
tic Slope region by C. pulla (Conrad) and must have reached peninsular Florida
from the west.



1972 JOHNSON: FLORIDA UNIONIDAE 193

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194 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

River system in the northeast, The v.alidity of this area as a region was
recently substantiated by Gilbert and Bailey (1972) who found that fol-
lowing Pliocene reconnection of insular Florida and the rest of the
southeastern United States, the freshwater fish, Notropis emiliae penin-
sularia, invaded the neighboring river systems to the north , the Suwan-
nee, Ochlockonee, St. Marys, and Satilla, where it came into contact
and integrated with the nominate subspecies.

GENERIC AFFINITIES
Only six genera of Unionidae occur in peninsular Florida: Elliptio,

Uniomerus, Anodonta, Camnculina, Villosa, and Lampsilis. These six
genera are also represented by species in the Southern Atlantic Slope and
Apalachicolan regions. The Southern Atlantic Slope region has 11
genera, 9 of which are represented in the Apalachicolan region where the
total is 17 geriera.

SUMMARY AND ANALYSIS OF DISTRIBUTION
Twelve species comprise the uni6nid fauna of the peninsular Florida

FIGURE 3. Species of Unionidae that appear to have migrated into peninsu-
lar Florida probably since the formation of the Pamlieo Terr;ice, during the Yar
mouth interglacial stage:

A . Open circles : Lampsilis (Lampsilis) ter€s ( Raftnesque), Widely distri -
buted in the Interior Basin; West Gulf Coastal region, Alabama-Coosa River
system, Apalachicolan region, and Peninsular Florida; Northern Mexico, Rio
Grande River system, Texas, east to the Withlacoochee River system, Florida.
This species' shallow penetration into the peninsula Shows its recent arrival,
Solid circles: Eltiptio (Elliptio) dariensis (Lea). Known only from the Altamaha
River system, Georgia, and the St. Johns River system, Florida. It appears to
have spread southward from the Altamaha rather recently when the shord line
extended farther to the east. In any event it entered the St. Johns subse-
quent to the formation of the Pamlico Terrace, as the St. Johns River lies east
of it.

Species of Unionidae that appear to have migrated into peninsula'r Florida
from the Apalachicolan or Atlantic Slope regionsj

B. Open circles: Anodonta couperiana Say. Apalachicolan region: Apalachi-
cola, Ochlockonee, and St, Marys River systems; Peninsular Florida; Atlantic Slope
region: Altamaha River system Georgia, north to the Cape F.ear River system,
North Carolina. The apparent absence of this species from the Suwannee and
Withlacoochee river systems suggests that it entered the Apalachicola River
system from a former confluence with  the Savannah River System and that it
sfread jnto the Florida peninsula from the north. Closed circles: Anodonta
peggyae Johnson. Apalachicolan region : Choctawhatchee River system, east
to the Suwannee River system; Peninsular Florida: Withlacoochee and Hills-
borough River systems.

C. Elliptio (Elliptic) jagensis (Lea). Apalachicolan region: St. Marks and Su-
wannee River systems; Peninsular Florida.

D. Villoaa uillosa (Wright). Apalachicolan region: Apalachicola River system,
east to the St. Marys River system, Georgia; Peninsular Florida.



1972 JOHNSON FLORIDA UNIONIDAE 195

region (Table 2). Nine are also found in the Apalachicolan region, and
among these four are clearly of more western origin, while three may have
had their origin in the Apalachicolan region. Of two species found in
both the Atlantic Slope and Apalachicolan regions, one appears to have
spread into the peninsula from both the west and north, the other from
the north. One other species is from the Atlantic Slope region. Two
species are endemic.

0
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196 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No, 4

Unidinerus tetralas'mus, Villosa vibex, Caruneulina paTua, and Lainp-
silis teres extend west beyond the Apalachicolan region . Two of these ,
U, tetralasmits and V. uibex also occur in the Southern Atlantic Slope
region, but as they are absent  in two intervening river systems (St. Marys
and Satilla) between the Altamaha and St. johns, they probably found
their way into Florida from the west. Carunculina parua and L. teres
clearly came from the west.

Villosci ui/losa, Eliiptio ja!/ensis, and Anodonta peggyae may have
had their origin in the Apalachicolan region or in peninsular Florida.
There is evidence that E/liptio icterina and Andonta couperiana spread
into the Apalachicolan region from a former conHuence with the Apa-
lachicola and Savannah rivers, E. icterina reached peninsular Florida
from both the west and north, and A. couperiana reached it from the
north . Elliptio dariensis, though its distribution is now discontinuous ,
evidently spread into the St. Johns River system from the Altamaha
River system of the Atlantic Slope . Villow amygcialci and Elliptio buck-
te!/i are endemic to peninsular Florida. The former is closely related to
V. lienosa of the Apalachicolan region, and E. buck/egi is close to E.
icterina.

A REVISION OF THE FLORIDA UNIONIDAE

In spite of the provincial restrictions of this paper each of the
species studied has been completely monographed, including those that
occur elsewhere. The synonymy of each species is believed to be com-
plete, and while the modern species concept has been assiduously ap-
plied, infallibility of judgement is not claimed. If there are composite
species , they will probably be found among the ubiquitous ELI.iptio .

Most of the Unionidae of Florida were described by Isaac Lea and
Timothy A. Conrad before the middle of the last centuij  Between 1883

. and 1934 the Wrights, father and son, described 52 species of mollusks,
mostly Unionidae from Florida (Johnson, 1967). The Wrights were un-
sophisticated naturalists who redescribed mahy of the species, some of
which.had already been described several times over.

Simpson (1892: 405-406) discussed the collectors of Florida Union-
idae up to that time. Many collections of Florida unioni(is have been
made since, including extensive ones by. G.W. Van Hyning and E.P.
St. John. These specimens are iii the Florida State Museum in Gaines-
ville. In 1962 my family and i, accompanied by Samuel L.H. Fuller, spent
6 weeks collecting in peninsular Florida at some 140 stations. Our col-
letting was facilitated by drought conditions.

The primary systematic studies of the Floridian Unionidae are those
of Charles T. Simpson. His Notes on the Unionidae of Florida and the
Southeastern States (1892) was the first attempt at a revision of these
species. A non-critical revision was later made by H. von Ihering, 03



1972 JOHNSON: FLORIDAUNIONIDAE 197

Unioniclos dit Florida (1895 ). Simpson subsequently published a SYnop,sis
of the Naiades ( 1900 ), which was expanded into the Descriptive Catalogue
of the Naiades (1914 ). These works were revisions ofthe naiades on a world -
wide basis. Simpson's conclusions concerning most ofthe Floridian species re-
mained unchanged until the publication of Frierson's, A Classijied and An-
notated Check List Of the North American Naiades (1927) in which some of
Simpson's synonymies were modified. Haas in Supe,jamilia Unionacea (1969a)
essentially followed Frierson and largely ignored the more recent work of
Clench and Turner, The Freshwater Mollusks OfA/abama, Georgia and Florida
from the Escambia to the Suwannee River (1956). In Supelfamil!/ Union-
(leed, Haas ( 1969 b ) only discussed the genera.

The anatomically-based classification of the genera o f Unionacea proposed
by Ortmann (1911, 1912) is used here. Since Ortmann's time, a few necessary
taxonomic changes have been made, but no one has substantially modified his
concepts of the North American genera until recently. Heard and Guckert,
(1970) in a work on the higher classification of Unionacea, interpreted the
phylogenetic relationships of these animals on reproductive features and not
on shell characters. They placed Elliptio and Uniomerus, as well as C!/clo-
naias, Hemistena, Lexingtonia, Plethobasus and Pleurobema, in the subfamily
Pleurobeminae. and put Elliptio buckle!/ i (Lea ) in the genus Popenaias

f f

BA

\r

FIGURE 4. Two species that may have survived on island refugia du ring Wico-
mico flooding in the Pliocene:

A. Villosa amggdata (Lea). Peninsular Florida. Replaced in the Apalachicolan
region by lienosa (Conrad) and in the Atlantic Slope region by delumbus (Conrad).

B. Emptio (Elliptio) bucklegi (Lea). Peninsular Florida.



198 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Frierson in a new subfamily Popenaindinae . These families differ in a single
way, the Pleurobeminae are tachytictic (i.e. the larvae have a short term in-
cubation) whereas the Popenaiadinae are bradytictic (i.e. the larvae have a
long term incubation).

Fuller (1971: 141) in a phylogenetic list of the Savannah River system
Unionidae rejected the work of Heard and Guckert (1970) reverting to the
scheme ofOrtmann. I,am not yet able to evaluate the validity of Plet,robeminae
and Popenaiadinae, and , as they do not appear to be germane to the under-
standing of the zoogeography of the fauna. in this study, they are ignored here.
This is not to imply that I regard shell characters or anatomical structures as
more important than reproductive features.

Valentine and Stansbery (1971: 13) also discussed the subfamilies of
Unionidae, bilt their paper was written before the new system of Heard and
Guckert appeared.

SYSTEMATIC SECTION

The following abbreviations have been used in the text, and figure
captions.

ANSP - Academy of Natural Sciences of Philadelphia, Pennsylvania
UF- Florida State Museum, University of Florida, Gainesville,

Florida
MCZ - Museum of Comparative Zoology, Cambridge, Massachu-

setts
UMMZ-Museum of Zoology, University of Michigan, Ann Arbor,

Michigan
USNM -United States National Museum, Washington, D.C.
SYNONYMY. -For ease of reference full citations are included for each

taxon. Elsewhere in the text references are abbreviated and may be found
in the bibliography.

Isaac Lea often rushed brief Latin descriptions of his new species
into print, a practice common in his time. These were subsequently fol-
lowed by adequate descriptions and figures, which were then reprinted as
"Observations on the Genus Unio." Only page references are included
here for this work, as the plates and figures were not renumbered, but are
the same as in the preceeding reference. Lea generally gave several locali-
ties where each of his .species had been found and did not select types, but
he always figured a single specimen for which he gave measurements. In
lieu of the use of the word 'type,' under Article 73 (b) Int. Code Zool.
Nomen. (1964), this is an "equivalent expression" and these specimens are
regarded as holotypes. During the early part of the century W. B. Marshall
located most of these figured specimens in the USNM.

With a few mentioned exceptions, all the types and type localities
were relocated. The latter are often rendered more specific from data on
original labels, by references to standard atlases, modern county maps, or
U.S. Geological Survey maps. These additional data are given in brackets.



1972 JOHNSON: FLORIDA UNIONIDAE 199

Unless specifically mentioned to the contrary, all extant types have
been examined. Almost none of Say's primary types have survived, and
many of Conrad's are missing, whereas most of Lea's have been located.

Only pertinent references are included. Simpson (1914) and Clench
and Turner (1956) are referred to only when the present synonymy is simi-
lar, or when the differences ate easily reconcilable.

DESCRIPTIONS.-Within the formal descriptiml, novelty of language
from one to another is introduced only when it serves to elucidate the dif-
ferences between the species. The discussion of the various characters
follows the same sequence throughout. The measurements are only in-
tended to convey the general size of specimens from a given station or to
illustrate sexual dimorphism when relevant.

HABITAT.-(Given only as observed in the Florida peninsula,
REMARKS. -These are designed primarily to aid iii differentiating

one species from another within the Apalachicolan, Southern Atlantic
Slope, and peninsular Florida regions.

RANGE.-The ralige covers the total area even for widely distributed
species.

SPECIMENS EXAMINED. -The records, limited to the area of the stildy,
are based mostly on specimens in the major collections mentioned above
under abbreviations. All specimens listed have been examined. Some-
times the same records are found in several museums, and those in the
MCZ are given preference. It is to be assumed that all records are in
this museum, unless specifically mentioned to the contrary.

Insofar as possible, the records are arranged from headwaters to the
mouth of the rivers, and from west to east- or south to north.

FIGURES.-When available, types are generally used to illustrate the
various species. Often more than one illustration is included to show intra-
specific variation. Some of the data on the plate captions are not repeated
elsewhere.

SUPERFAMILY UNIONACEA Thiele 1935
Family UNIONIDAE (Fleming 1828) Ortinann 1911

Subfamily UNIONINAE (Swainson 1840) Ortmann 1910
Genus Elliptio Rafinesque

Subgenus Elliptio RaRnesque

Elliptic Rafinesque, 1819, Jour. de Physique, de Chimie, d'Hist. Nat. (Paris),
88: 426 [nome,i nudum]. Rafinesque, 1820, Ann. 66n, des Sci, Phy-
siques (Bruxelles), 5: 291. Species listed: E. nigra Rafinesque, E. cms:sa
( Say), E . viridis Rafinesque, E. fasciata Rafinesque.
TYPE SPECIEF-Unio nigra Rafinesque. Subsequent designation, Ortmann,
1912 , Ann . Carnegie Mus ., 8 : 266 . The previous use of Unio crassidens
Lamarck as type species' by Simpson, (1900, Proc. U.S. Natl. Mus., 22: 700)
is invalid because Rafnesque did not include Lamarck's name in his list
of species, H. B. Baker (1964, Nautilus, 78: 33) pointed out that Rafinesque
consistently used Elliptio as feminine (e.g. E. nigra) and that therefore
the name should be thus treated.



200 BULLETIN FLORIDA STATE MUSEUM Vol, XVI No. 4

Speciation within Elliptio has Occurred primarily in the Apalachi-
colan, peninsular Florida, and Southern Atlantic Slope regions. In these
regions the species of Elliptio are the most abundant Unionidae. Elliptios
are often found in environments where no other Unionidae live, as some
of them have an unusually wide environmental tolerance, even to silting
and pollution, As a consequence some of the species have developed many
ecophenotypes, and at a given station there often appears to be less inter-
specific variation than intraspecific variation between localities. This has
led to a plethora of names applied to the several species.

Elliptio (Elliptio) darien.gis (Lea)
Figures 3A, 5A-C

Unio dariensia Lea 1842, Trans. Amer. Philos. Soc., 8: 246 pl. 26, fig. 61 ([Alta-
maha River] near Darien [Mcintosh Co,], Georgia; figured holotype
USNM 85691). Lea, 1842, Obs. Unio, 3: 84.

Unio monroen.sis Lea 1843, Dese. Twelve Uniones. (Lake Monroe, Florida).
Lea, 1846, Trans. Amer. Philos, Soc., 9: 279 pl. 41, fig. 8, figured holo-
type USNM 85169. Lea, 1848, Obs. Unio, 4: 37.

Unio websterii B.H. Wright 1888, Proc. Acad. Nat. Sci. Phila., p. 113, pl. 2,
fig. 2 (Lake Woodruff, Volusia Co., Florida; lectotype USNM 125697,
selected by Johnson, 1967, Oce. Papers on Moll., 3: 10, pl. 7, fig. 2)

Onto hartwrightii B.H. Wright 1896, Nautilus, 9: 121, pl 2, figs. 4-6 (Lake Beres-
ford [Volusia Co.], Florida; holotype USNM 151031, refgured by John-
son, 1967, 0ce, Papers on Moll., 3: 6, pl, 7, fig. 1 and the type locality
further restricted on the basis of the original label to: [St. Johns River],
Blue Springs [3 mi. S Lake Beresford, Volusia Co.], Florid:i).

Elliptio (Elliptio) dariensia (Lea). Johnson, 1970, Bull. Mus. Comp. Zool., 140
(6): 310, pl. 6.
DESCRIPTION. -Shell often large, exceeding 100 mm in length. Out-

line subrhomboidal or subtrapezoidal. Valves rather flat to subinflated,
thin but strong, inequilateral. Ailterior end regularly rounded; posterior
end occasionally a little produced iind slightly biangulate, but more often
obliquely truncated. Vetitral margin straight m- slightly curved. Dorsal
margin straight, forming a sharp angle with the obliquely descending
posterior margin. Posterior ridge usually very sharp with a faint secondary
ridge above. Posterior slope rather broad, but well defined, with numerous
wrinkles on it. Umbos full to inflated, but rather low, located in the anter-
ior third of the shell, their sculpture consisting of five or six double-looped
bars, slightly more elevated and angular behind the sinus. Periostracum
smooth and yellowish with fine green rays when young, becoming darker
greenish-yellow, dark chestnut, or blackish; generally smooth on the
disk but sometimes roughened, especially on the posterior slope.

Left valve with two heavy, rough pseudocardinal teeth, the more
anterior one slightly smaller and. a little lower. Hinge line rather short
and broad with two short slightly curved. granular, lateral teeth. Right



1972 JOHNSON: FLORIDA UNIONIDAE 201

B

C 
-I.

*h

Figure 51 . Elliptio ( Elliptio) dariemis ( Lea ). A . Holotype of Unio monroen.sis Lea.
Lake Monroe, Florida , USNM 851691 , L 69, H 42, W 26 , B . Holotype of Uiii()
hartwrightii B. H. Wright. [St, Johns River], Blue Springs [3 mi. S of Lake Beres-
ford , Volusia Co.], Flor icla . USNM 151031 . L 78, H 47 , W 31 . C . Lectotype of Unio
websteri B. H. Wright. Lake Woodruff, Volusia Co., Florida. USNM 125697. L 96,
H 57, \V 33.
' All figures slightly reduced. Mcasurement: in mni; L = length, H = height. W = width.



202 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

valve with one chunky, serrated pseudocardinal; one lateral tooth. Beak
cavities shallow, with a few dorsal muscle scars under the hinge plate.
Anterior adductor muscle scars deep, posterior ones and palial line dis-
tinct. Nacre generally purple, sometimes white, occasionally yellow.

MEASUREMENTS,-L 106 nim, H 60 min, W 37 mm (St, Johns River
[town of] Lake Monroe, Seminole Co.); L 96 mm, H 57 mm, W 33 mm
(Lake Woodruff, Volusia Co., lectotype of U. websteri Wright); L 78 mm,
H 47 mm, W 31 mm (St. Johns River, Blue Springs, 3 mi. S of Lake Beres-
ford, Volusia Co., holotype of U. hartwrightii Wright).

HABITAT. -Generally found in s:md, in flowing water.
REMARKS. -Elliptio dariensis (Lea) is restricted to the St. Johns River

system, Florida, and the Altamaha River system, Georgia. In the latter
system it reaches its greatest size, some individuals exceeding 130 mm in
length. It is close to Elliptio c. crassidens (Lamarck) of the Apalachicolan
region, but the shell of dariensis is never as ponderous; the posterior ridge
is consistently much sharper, especially in immature individuals; and the
posterior slope, though generally wrinkled, does not have the strong
radial pattern of E. c. crassidens.

The especially sharp posterior ridge and relatively thin shell with
its yellowish, finely green-rayed periostracum that becomes dark chestnut,
separate this species from any other in peninsular Florida.

RANGE.- Peninsular Florida: St. Johns River system. Southern At-
lantic Slope: Altamaha River system, Georgia.

SPECIMENS EXAMINED.-ST. JOHNS RIvER DRAINAGE. Econlockhatchee
River, near confluence with St. Johns River; St. Johns River, 1 mi. ESE of
Osceola; St. Johns River, 4 mi. E of Sanford; St. johns River [town of] Lake
Monroe; Lake Monroe; all Seminole Co, Lake Beresford, Lake Woodruff, both
Volusia Co. 'St. Johns River, Georgetown; Putnam Co. (FSM). LAKE REGIoN
DR~NAGE. Lake Virginia, near entrance of Lake Sue Canal, Winter Park;
Orange Co. BLACK CREEK DRAINAGE. North Fork, Biack Creek, 14 mi. SW
of Orange Park, Clay Co.

Elliptio (Elliptio) icterina (Conrad)
Figures 2C, 6A-H

Unio icterinus Conrad, [May] 1834, New Fresh Water Shells United States,
p. 41, pl. 6, fig. 5 (muddy shore, Savannah River, opposite Augusta [Rich-
mond Co.], Georgia; figured holotype ANSP 41381). Published m May,
teste Conrad, 1853, Proc. Acad. Nat. Sci.  Phila., 6, 244, and not disputed
by Lea, 1854, Proc. Acad. Nat. Sci. Phila., 7: 336-349. Conrad, 1836, Mono-
graphy Unionidae, no. 4, p. 39, pl. 18, fig, 2,

Unto raveneli Conrad, [May] 1834, New Fresh Water Shells United States, p.
39, pl. 6, fig. 4 (Wateree Canal; since found in the small creeks near
Cooper River; vicinity of Santee Canal; all South Carolina; 2 syntypes
ANSP 41370, the smaller one agrees with Conrad's description, but is not
the figured specimen, which appears to be Elliptio complanata (Lightfoot).
The second specimen is Elliptio lanceolata (Lea); non Unio ravenelianus
Lea, 1834).



1972 JOHNSON: FLORIDA UNIONIDAE 203

Unto watereensis Lea 1836, Synopsis Unionidae, p. 31, New name for Unio
raueneli Conrad, 1834, non Unio ravenelianus Lea, 1834. As pointed out
by Simpson, 1900, Proc. U.S. Nat. Mus., 22: 748; this change was un-
hecessary.

Unio confertus Lea, [August or September] 1834, Trans. Amer. Philos. Soc.,
5: 103, pl. 16, fig. 47 (Santee Canal, South Carolina, type not in USNM
[presumed lost]), Lea, 1834, Obs. Unio, 1: 215. Published in Augu st or
September 2834, teste Lea, 1854, Proc, Acad, Nat. Soi, Phila., 7: 244.

Unio lugubris Lea 1834, Trans. Amen Philos. Soc., 6: 30, pl. 9, fig. 25 ([Alta-
maha River], Hopeton, near Darien [Mcintosh C6.], Georgia; figured holo-
type USNM 85638). Lea, 1838, Obs. Unio, 2: 30, non Say, 1832.

Unio geddingsianus Lea 1840, Proc. Amer. Philos. Soc., 1: 285 (Congaree
River, South Carolina). Lea, 1842, Trans. Amer. Philos. Soc., 8: 202, pl,
11, fig. 15; figured holotype USNM 85650. Lea, 1842, Obs. Unio, 3: 40.

Unio fuscatus Lea 1843, Dese. Twelve Uniones (Black Creek, Florida). Lea,
1846, Trans. Amer. Philos. Soc,, 9: 277, pl. 40, fig. 4, figured holotype
USNM 85243. Lea, 1848, Obs. Unio, 4: 35.

Unio occultus Lea 1843, Desc Twelve Uniones (Black Creek; Lake Monroe;
both Florida). Lea, 1846, Trans. Amer. Philos, Soc., 9: 279, pl. 41, fig. 7,
figured holotype USNM 85247, from Black Creek. Lea, 1848, Obs. Unio,4: 37.

Unio limatulus Conrad 1849, Proc. Acad. Nat. Sci. Phila., 4: 154 (Savannah
River, [Georgia]; type not in ANSP [presumed lost] ). Conrad, 1850, Jour.
Acad. Nat. Sci. Phila., (2) 1: 276, pl. 37, fig. 9. Conrad, 1853, Proc. Acad.
Nat. Sci. P.hila., 6: 251.

Unio tuomeyi Lea 1852, Trans. Amer. Philos, Soc., 10: 256, pl. 13, fig. 4
(Abbeville District [Savannah River drainage], South Carolina, figured
holotype USNM 85669). Lea, 1852, Obs. Unio, 5: 12.

Unio tohiteianus Lea 1852, Trans. Amer. Philos. Soc., 10: 258, pl. 14, fig. 8
(near Savannah [Chatham Co.], Georgia; figured holotype USNM 85658).
Lea, 1852, Obs. Unio, 5: 14.

Unio barrattii Lea 1852, Trans. Amen Philos. Soc., 10: 256, pl. 13, fig, 5 (Abbe-
ville District [Savannah River drainage], South Carolina; figured holo-
type USNM 86010). Lea, 1852, Obs. Unio, 5: 12.

Unio pullutis Lea 1856, Proc. Acad. Nat. Sci. Phila.,8: 262 (Creeks near Col-
umbus [Muscogee Co.], Georgia). Changed to:

Unio pullatus Lea 1858, Jour, Acad. Nat. Sci. Phila., (2) 4: 57, pl. 8, fig. 39;
figured holotype USNM 86020. Lea, 1858, Obs. Unio, 6: 57.

Unio coruscus Gould 1856, Proc. Boston Soc. Nat. Hist., 6: 15 (River Saiht
John's, near Lake Beresford, Florida; measured holotype MCZ 169097,
figured by Frierson, 1911, Nautilus, 25, pl. 1, figs. 1-3 and by Johnson,
1964, U.S. Natl. Mus., Bull. no. 239, p. 60, pl. 32, fig. 3)

Unio micans Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 85 (Catawba River,
Gaston Co.; Deep River, Gulf [Chatham Co.] ; both North Carolina).
Lea, 1862, Jour. Acad. Nat. Sci. Phila,, (2) 5: 59, pl. 3, fg. 207, figured
holotype USNM 85077 from the Catawba River. Lea, 1862, Obs. Unio,8: 63.'

Unio obnubiMs Lea 1857, Proc, Acad. Nat: Sci. Phila., 9: 169 (Buckhead Creek,



204 BULLETIN FLORIDA STATE MUSEUM Vol XVI No. 4

Burke Co., Georgia). Lea, 1858, Jour: Acad. Nat Sci. Phila., (2) 4: 84, pl

17, fig. 64; figured holotype USNM 85646. Lea, 1858, Obs, Umo 6: 84.

Unio opacus Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 169 (Buckhead Creek,

Burke Co., Georgia), Lea, 1858, Jour, Ac.id, Nat, Sci. Phila., (2) 4: 86, pl

18, fig. 66; figured holotype USNM 85546. Lea, 1858, Obs. Unio, 6: 86.

Unio sinzilis Lea 1857, Proc. Acad, Nat. Sci. Phila., 9: 169 (Buckhead Creek,

Burke Co., Georgia). Lea, 1858, Jour. Aead. Nat. Sci. Phila., (2) 4: 91, pl

19, fig. 71, Agured holotype USNM 85653. Lea, 1858, Obs. Unio, 6: 91.

Unto sub/atus Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 169 (IChattahoochee

River], Uehee Bar, below Columbus [Muscogee Co.], Georgia). Lea, 1858

Jour. Acad. Nat. Sci. Phila., (2) 4: 82, pl. 16, fig. 62; figured holotype USNM

85897. Lea, 1858, Obs. Unio, 6: 82.
Unio viridicatus Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 170 (Buckhead Creek,

Burke Co., Georgia). Lea, 1858, Jour. Acad. Nat. Sci. Phila., (2) 4: 87, pl

18, fig. 67; fgured holotype USNM 85551. Lea, 1858, Obs. Unio, 6: 87.

Unio letricus Lea 1857, Proc. Acad, Nat. Sci. Phila., 9: 170 (Flint River, near Al-

bany [Dougherty Co.], Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila., (2)

4: 195, pl. 22, fig. 78; figured holotype USNM 85655. Lea, 1859, Obs. Unio,

7: 13.
Unio aequatits Lea 1857, Proc. Acad. Nat, Sci. Phila,, 9: 170 (Buckhead Creek,

Burke Co., Georgia). Lea, 1858, jour. Acad. Nat. Sci. P.hila., (2) 4: 89, pl. 19,

fig. 69; fgured holotype USNM 85561. Lea, 1858, Obs, Unio, 6: 89.

Unio aquilus Lea 1857, Proc. Acad. Nat. Sci. Phila., 9: 172 (Flint River, Macon

[County], Georgia). Lea, 1858, Jour. Acad. Nat. Sci. Phila., (2) 4: 92, pl. 20,

fig. 72; figured holotype USNM 85993. Lea, 1858, Obs. Unio, 6: 92.

Unio viridiradiatus Lea 1859, Proc . Acad . Nat. Sci . Phila ,, 11 : 154 ( Big Uchee

River [ = Creek, Russell Co., Alabama] near Columbus, Georgia). Lea, 1860,

Jour. Acad. Nat. Sci. Phila., (2) 4: 336, pl, 53, fig. 161; figured holotype USNM

86018), Lea, 1880, Obs. Unio, 8. 18.

FicuRE 6 1. Elliptio (Elliptio) icterina (Conrad). A. Holotype of Unio ft,Scottis

Lea. Black Creek, Florida. USNM 85243. L 43, H 23, W 13. B. Holotype of Unio

occultus Lea. Black Creek, Florida. USNM 85247, L 52, H 38, W 19, C. Holotype

of Umo simpsom B. H. Wr{ght. Like Woodruff, Volusia Co., Florida. USNM

151038. L 59, H 31, W 16. D. Holotype of Unto frganus B.H. Wright, Lake Ashby,

Volusia Co., Florida . USNM 151082 . L 44 , H 25 , W 16 . E . Lectotype of Unio

burtchiant,s S.H. Wright. St. Marys River, Nassau Co., Florida. USNM 149653.

L 52, H 29, W 18. F. Lectotype of Unio diazensis S. H. Wright. Lake Diaz, Vo-

lusia Co ., Florida , USNM 149652 , L 34 , H 20 , W 13 . G . Holotype of Elliptio may -

webbae B, H. Wright. Near Seminole Springs [3.4 mi. NE Sorrento], 15 mi.

SE of Eust-is [Lake Co.], Florida. USNM 425354. L 52, H 30, W 23. H. Holo-

type of Unio notani B. H. Wright. Creek flowing into St. Johns River, near Pa-

latka [Putnam Co.], Florida, USNM 151030, L 71, H 36, W 22.

Elliptio (Emptio) bucklegi (Lea). 1. Holotype of Unto pinei B. H. Wright. Un-

named lake in the Witthacoochee [Withlacoochee] River region of Hernando Co.,

Florida. USNM 150127. L 74, H 34, W 22.
1 Slightly reduced; measurements in ninb L ·= length, H = height, W = width.



1972 JOHNSON: FLORIDA UNIONIDAE 205

B
4- w-----ig

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I.

%--  V.'.L-'fe 41~ --1----Il

.*54/loil/'ll..../
ri *•8-'- 5 .1--/Uma~

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i.·

J

f- V I.

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14 '4-.&6«
S'*



206 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Unio hepaticus Lea 1859, Proc. Acad. Nat. Sci. Phila., 11: 154 (Salkahatchie River,
South Carolina), Lea, 1860, Jour. Acad. Nat. Sci. Phila., (2).4: 348, pl. 57,
fig. 173; figured holotype USNM 85559. Lea, 1860, Obs. Unio, 8:30.

Unio uiridans Lea 1859, Proc. Acad. Nat, Sci. Phila., 11: 170 (near Columbus
[Muscogee Co.], Georgia). Lea, 1860, Jour. Acad. Nat. Sci. Phila., (2) 4: 337,
pl. 54, fig. 162; fgured holotype USNM 85579. Lea, 1860, Obs. Unio, 8: 19.

Unio uerutus Lea 1859, Proc. Aead. Nat. Sci. Phila., 11: 171 (Flat Rock Creek, near
Columbus [Muscogee Co.], Georgia). Lea, 1860, Jour. Acad. Nat. Sci.
Phila., (2) 4: 335, pl. 53, fig. 160; figured holotype USNM 85899. Lea, 1860,
Obs. Unio, 8: 17.

Unio ocmulgeensis Lea 1861, Proc. Acad. Nat. Sci. Phila., 13: 38 (Little Ocmulgee
River, Lumber City [Telfair Co.], Georgia). Lea, 1862, Jour. Acad. Nat. Sci.
Phila., 5: 95, pl. 14, fig. 243; fgured holotype USNM 85901. Lea, 1862, Obs.
Unio, 8: 99.

Unio merceri Lea 1862, Proc, Acad. Nat. Sci. Phila., 14: 169 (Lee Co., [Flint
River drainage], Georgia). Lea, 1862, Jour. Acad. Nat. Sci. Phila., (2) 5: 209,
pl. 31, fig. 278; figured holotype USNM 86057. Lea, 1863, Obs. Unio, 9.31.

Unio lucidu,s Lea 1863, Proc. Acad. Nat. Sci. Phila., 15: 192 (Livingston's Creek,
Brunswick Co., [NE corner of Columbus Co., Cape Fear River drainage],
North Carolina). Lea, 1866, Jour. Acad. Nat. Sci. Phila., (2) 6: 9. pl. 2, fig. 6,
figured holotype USNM 85242. Lea, 1867, Obs. Unio, 11: 13.

Unio lioingstonensis Lea 1863, Proc. Acad. Nat. Sci. Phila., 15: 192 (Livings-
ton's Creek, Brunswick Co., [NE corner of Columbus Co., Cape Fear
River drainage] North Carolina). Lea, 1866, Jour. ·Acad. Nat. Sci. Phila.,
6: 14, pl. 4, fig. 11; figured holotype USNM 85536. Lea, 1867, Obs. Unio,
11: 18.

Unio ablatus [ sic ] Lea 1863, Proc. Acad . Nat. Sci . Phila., 15 : 193 (Long
Creek, Gaston Co., North Carolina). Changed to:

Unio oblatus Lea 1866, Jour. Acad. Nat. Sci. Phila., (2) 6. 13, pl. 4, fig. 10;
figured.holotype USNM 86001. Lea, 1867, Obs. Unio, 11: 17.

Unio radiolus Lea 1871, Proc. Acad. Nat, Sci. Phila., 23: 192 (Ogeechee River,
Liberty Co,, Georgia). Lea, 1874, Jour. Acad. Nat. Sci. Phila., (2) 8: 21,
pl. 6, fig. 18; figured holotype USNM 85621. Lea, 1874, Obs. Unio, 13:
25.

Unio cuspitatus Lea 1872, Proc. Acad. Nat. Sci. Phila., 24: 159 (Buckhead
Creek, Burke Co., Georgia; Abbeville District [Savannah River drainage],
South Carolina). Lea, 1874, Jour. Acad: Nat. Sci. Phila., (2) 8: 43, pl. 14,
fig. 50; figured holotype USNM 86014, from Buckhead Creek. Lea, 1874,
Obs. Unio, 13: 47.

Unio hastatus Lea 1873, Proc. Aead. Nat. Sci. Phila,, 25: 423 (New Market,
Abbeville District [Savannah River drainage], South Carolina). Lea,
1874, Jour. Acad. Nat. Sci. Phila., (2) 8: 56, pl. 19, fig. 54; figured holotype
USNM 86013. Lea, 1874, Obs. Unio, 13: 60.

Unio frganus B.H. Wright 1888, Proc. Acad. Nat. Sci. Phila., p. 113, pl. 2, fig.
1 (Lake Ashby, Volusia Co., Florida; figured holotype USNM 151032, re-
figured by Johnson, 1967, Occ. Papers on Moll, 3: 6, pl. 8, fig. 5)

Unio nolani B. H. Wright 1888, Proc. Aead. Nat. Sci. Phila., p. 116, pl. 4, fig.



1972 JOHNSON: FLORIDA UNIONIDAE 207

11 (a creek flowing into St. Johns River, near Palatka [Putnam Co.],
Florida; holotype USNM 151030, refgured by Johnson, 1967, Oce. Papers
on Moll., 3: 7, pl. 10, fig. 4)

Unio simpsoni B. H. Wright 1888, Proc. Acad. Nat. Sci. Phila., p. 117, pl. 5, fig.
1 (Lake Woodruff, Volusia Co., Florida; holotype USNM 151038, refigitr-
ed by Johnson. 1967, Occ. Papers on Moll., 3..8, pl. 8, fig. 2).

Unio burtchianus S.H. Wright 1897, Nautilus, 10: 137 (St. Marys River, Nassau
Co., Florida; lectotype USNM 149653, selected by Johnson, 1967, Oce.
Papers on Moll., 3: 5, pl. 8, fig. 4, possibly the specimen figured by Simp-
son, 1900, Proc. Acad. Nat. Sci. Phila., p. 80, pl. 4, fig. 8).

Unio diazensis S.H. Wright 1897, Nautilus, 11: 5 (Lake Diaz, Volusia Co.,
Florida; lectotype USNM 149652, selected by Johnson, 1967, Occ. Papers
on Moll.,3: 6, pl. 8, fig. 6).

Unio dispalans B.H. Wright 1899, Nautilus, 13: 50 (Suwannee River, Florida;
holotype USNM 159986, figured by Simpson, 1900, Proc. Acad. Nat. Sci.
Phila., p. 80, pl. 1, fig. 9. refigured by Johnson, 1967, Oce, Papers on Moll.,
3: 6, pl 8, fig. 3)

Unto singularis B.H. Wright 1899, Nantilus, 13: 75 (Spring Creek, [a branch
of the Flint River], Decatur Co., Georgia; measured holotype USNM
159988, figured by Johnson, 1967, Occ. Papers on Moll., 3.8, pl. 5, fig, 7)

Elliptio niagwebbae B.H. Wright 1934, Nautilus 48: 28; 47, pl. 13, figs. 5-8
(near Seminole Springs [3.4 mi. NE Sorrento], 15 miles SE Eustis [Lake
Co.1, Florida, refigured by Johnson 1967, Occ, Papers on Moll., 3: 7,pl. 10, fig. 3).

Elliptio cylindraceus Frierson 1927, Cheek List North American Naiades, p
29, new name for Unio lugubris Lea, 1838, non Say, 1832.

Elliptio strigosus (Lea). partim, Clench and Turner, 1956, Bull. Florida State
Mus., 1: 165.

Elliptio (Elliptio) icterina (Conrad). Johnson, 1970, Bull. Mus. Comp, Zool.,
140 (6): 325, pl. 9, figs. 2-10, pl. 10„figs. 1-3.
DESCRIPTION. -Shell generally small to medium, seldom reaching

over 100 mm in length. Outline variable; subquadrate, to subelliptical,
sometimes rather pointed. Valves subinflated, subsolid to very solid, in-
equilateral. Anteri(ir end regularly rounded; posterior end generally bi-
angulate near the base, though sometimes rather produced and pointed.
Ventral margin straight or slightly arcuate, sometimes obliquely descend-
ing. Dorsal margin slightly curved or almost straight, meeting the oblique-
ly descending posterior margin in a more or less distinct angle. Hinge
ligament long and low. Posterior ridge bfoadly rounded, generally faintly
double. Posterior slope flat to slightly concave, sometimes with very
faint radial sculpture present. Umbos broad and full but very low, located
in the anterior quarter of the shell, their sculpture consisting of several
double-looped ridges. Disk surface generally flat or slightly concave
when an umbonal-ventral sulcus is present. Periostracum generally fine
and shiny, though sometimes heavy and rough, black, brownish-black,
yellowish-brown, or bright yellow, chestnut, often with numerous very
fine green rays.



208 BULLETIN FLORIDA STATE MUSEUM VoI. XVI No. 4

Left valve with two stumpy pseudocardinal teeth, one in front of the
other, often of about equal height. Hinge line short and narrow; two long,
straight lateral teeth. Right valve with two roughly parallel pseudocard-
inals, the posterior one apt to be serrated and chunky, the more anterior
one low and vestigial; one lateral tooth. Beak cavities very shallow, with
a few dorsal muscle scars. Anterior and posterior adductor muscle scars
and pallial line all distinct. · Nacre generally purplish, though sometimes
salmon, bluish-white, or pinkish; posteriorly iridescent.

MEASUREMENTS. - L 85 inm, H 41 inm, W 29 nim (Black Creek, 2 mi.
E of [town of] Kingsley Lake, Clay Co.); L 76 mm, H.39 mm, W 27 mm
(Lake Beresford, Volusia Co.); L 56 mm, H 30 mm, W 17 mm (Magnesia
Springs, 3.5 mi. W of Hawthorne, Alachiia Co.)

HABITAT.-Found iii hikes, ponds, sinall streanis and large rivers in
nearly every type of substrate. Elliptio ilterina (Conrad) is sometimes
found with E. buckle!/i (Lea) and other Unionidae, but like E. bucklegi
it is often found alone. One or the other of these species is generally
more abundant than other unionids at a given station.

REMARKS. -Elliptio icterina (Comrad) is a highly variable species and
a number of popillaticins have been named, soine·sever:11 times over. While
some populations are often more or less identifiable, there is usually a
gradual transition between one river system and the next, such that while
specimens from extremes of the range bear little resemblance to one
another, there appears to be no point at which subspecies can be sepa-
rated. There is often a great deal of ecophenotypieal variation, even at
what appears to be a single station, the extremes usually connected by
intergrades.

In the Apalachicolan region E. icterina can be confused with E.
comlita,lara (Lightfoot) (Johnson, 1970: 314) and E. arctata (Conrad)
(Johnson, 1970:331). The latter is a rare species outside the Alabama
River system. It is distinctly and consistently arcuate with compressed
valves, whereas icterina has a generally straight or curved ventral margin,
is bluntly or acutely pointed posteriorly, and when occasionally produced
postbasally, the valves are somewhat inflated. Elliptio icterina occurs
with E. complanata in the Chattahoochee Riyer system, and it can be
distinguished from E. complanata by its less rhomboidal. more elongate,
often pointed shape.

The most common form E. icterina takes in the Apalachicolan re-
gion is subrhomboidal to subelliptical, sometimes appearing quite pointed
posteriorly if the biangulated posterior ridge ends near the medial line. The
tendency to be pointed is more prevalent in specimens from the Apala-
chieolan region and western rivers of peninsular Florida than in those
from the Southern Atlantic Slope. The similarity between the popula-
tions of this species from the Chattahoochee River and the upper Savan-
nah River, first noted in the .localities of some of the taxa Isaac Lea de-
scribed, affords evidence of the commingling of the headwaters of. these
two systems. The shell form just described includes most of the taxa Simp-



1972 JOHNSON: FLORIDA UNIONIDAE 209

son (1914, 2: 661) grouped under U. tuome!/i Lea and Clench and
Turner (1956: 165-169) under El/.iptio strigosus (Lea).

Noteworthy are the populations from Moccasin Creek 6f Econfina
Creek, Bay Co., Florida, on the Gulf Coast; Black Creek, Florida; St.
Marys and Canoochee Rivers, Georgia; all on the Atlantic Slope, which
resemble one another more than they do those from the several inter-
vening river systems. The shells from these rivers tend to be solid, to
be more uniformly biangulate posteriorly, and to have a rather charac-
teristic yellowish-brown to shiny chestnut pei·iostracum often with fine
dark green rays.

in northern Florida is a smaller ecophenotype that lives in lakes. It
has a heavy shell and tends to be generally oval.

In the Wekiva and Oklawaha Rivers of the St. Johns River system,
Florida, occurs a very thin, compressed subrhomboidal ecoI,henotype,
the shells ·of which end in a broad biangulation below the medial line,
but which tend to become heavier, more inflated, and produced post-ba-
sally toward the headwaters of the streams and in springs. This shell form
occurs again in abundance in Buckhead Creek of the Ogeechee River
system, Georgia which, like these Floridian rivers, is rich in carbonates;
it also occurs. in Brier Creek of the Savannah River system, Georgia, and
the Salkahatchie River, South Carolina. That the shape of the shell is
environmentally controlled is illustrated by the close resemblance of
shells from Magnesia Springs, 3 mi. W Hawthorne, Alachua Co., Florida,
the headwaters of the Ogeechee River, and Cedar Spring, 2 mi. SE Bamberg,
Bamberg Co., South Carolina. Though from widely separated drainage
systems, shells from these stations bear a closer resemblance to one an-
other than to specimens from other stations in their respective drainage
systems. The shell form just described includes most of the taxa Simpson
(1914) grouped under Unio obnubilis Lea (p. 641) and some of those
under Unio confertus Lea Cp. 639).

Some specimens of E, icterina, especially from the tidal areas of At-
lantic Slope rivers, are elongate and inflated with a tendency -for the
ventral margin to be slightly arcuate. The periostracum is often rough and
black. This shell form includes most of the taxa Simpson (1914: 639)
grouped under Unio confertus Lea.

On the Atlantic Slope of Georgia, E. icterina (Conrad) is most easily
confused with E. complanata (Lightfoot) with which it is associated at
many stations, but complanata is quite consistently rhomboidal, and the
valves are less apt to be inflated. The periostracum of icterina is sometimes
bright yellow or chestnut and is generally more shiny and smooth than
that of complanata.

In peninsular Florida, E. icterina can be confused with E. buckle!/i
(Lea), under which see Remarks. Morrison (1972: 38) regarded Elliptio ic-
terina (Conrad) 1834 as a synonym of E. cong,imea (Lea) 1831, but as
Johnson ( 1970 : 309 ) pointed out , congamea, with its fine ridges radiating
from the upper posterior ridge to the dorsal margin , is closer to E , cras-



210 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

sidens cm.ysidens (Lamarck) than to icterina, which is more similar to E.
complanata ( Lightfoot ) (Johnson , 1970 : 328 ). As both icterina and con-
gamea are good species, Morrison's att€impt to resurrect moene/i as the
taxon for the former is unnecessary.

RANGE. -Apalachicolan region: Escanibia River system, east to the
St. Marys River system, Georgia. Peninsular Florida. Southern Atlantic
Slope: Altamaha River system, Ceorgia, north to the White Oak River,
North Carolina.

~PECIMENS EXAMINED.-WACCASASSA RIVER ~YSTEM: Blue Springs, 3 mi.

W Bronson; Waccasassa River, 4. mi. NE [town of] Otter Creek, both Levy Co.
WITHLACOOCHEE RivER SYSTEM: Withlacoothee River, 9 mi. N D.,de City; With-

lacoochee River, 1 mi. NW Lacoochee, both Pasco Co, Little Withlacoochee
River, Rerdell; Withlacoochee River, Istachatta; both Hernando Co. Lake
Ts.ila, Apopka, Citrus Co.; Withlacoochee River, Dunn ellon; Marion Co.;
Sulphur Spring, near Withlacoochee River, N Red Level; Citrus Co. HiLLS-
BOROUGH RIVER SYSTEM: Itchepackesassa Creek; Hillsborough River, Morris
Bridge, 14 mi. NE Tampa; Lake Thonotosassa, 2.5 mi. WNW Antioch;
Hillsborough River, 4 mi. NE Temple Terrace; all.Hillsborough Co. ALAFIA
RIVER SYSTEM: Fishhawk Creek, 2 mi. S Lithia; Hillsborough Co. PEACE RIVER
SYSTEM: Lake Gibson, Cibsonia; Polk Co. Payne Creek, 1.1 mi. S Bowling
Green, Charley Apopka Creek, 1.5 mi. N Gardner; both Hardee Co. Horse
Creek, 8 mi. W Arcadia; Charlie Creek, 3 mi. SW Nocatee; Prairie Creek;
Peace River, 1.5 mi. below Arcadia;, all De Soto Co, KISSIMMEE RIVER SYS-
TEM AND EVERCLADES: KISSIMMEE RIVER DRAINAGE: Lake Verona,
Avon Park; Highlands Co. Alligator Lake, 4 mi. SE Ashton, Osceola Co. Weo-
hyakapka Creek, 5 mi. ESE Hesperides; Lake Rosalie, 4 mi. ENE Hesperides;
both Polk Co. LAKE OKEECHOBEE DRAINAGE: Seven Mile Tower Road,
3.5 mi. S Tamiami Trail, Everglades National Park, canal, W of Bridge no.
22, Tamiami Trail; both Dade Co. Snake Creek, 9 mi. W. Hallandale; Broward
Co. Loxahatchee Creek, 6 mi. W Jupiter; Palm Beach Co. ST. JOHNS
RIVER SYSTEM: ST. JoHNs RivER DRAINAGE: Ditch near Lake Washing-
ton, 6 mi. W Eau Callie; Lake Poinsett, 10 mi. WNW Coca; both Brevard Co.
Puzzle Lake, 7 mi. SE Geneva; Little Econlockhatchee River, 3.5 mi. S Oviedo;
Econlockhatchee River, near confluence with St. Johns River; Lake Horseshoe,
Chuluota; Lake Catherine, Chuluota; Lake Jessup; Gee Creek, near North
Orlando; St, Johns River, Sanford; St. johns River, [town of] Lake Monroe;
all Seminole Co. Lake Ashby, 8 mi. NE Osteen; St. Johns River, near Lake
Beresford; Spring Garden Lake, 1 mi. NW De Leon Springs; all Volusia Co.
Alexander Spring Creek, 5 mi. S Astor Park; Lake Co. Lake Kerr, 3 mi. SW
Kerr City; Marion Co. Crescent Lake; Lake Stella, Crescent City; both Putnam
Co. Lake Howell; Seminole Co, WEKIvA RIvER DRAINACE. Seminole Springs, 15
mi. SE Etistis, Lake Co. Rock Springs Creek; Wekiva River [Seminole Co,],
12 mi. NW Winter Park; Wekiva River, 10 mi. below Rock Springs, all Orange
Co. Wekiva River, 7 mi. WSW [town of] Lake Monroe, Seminole Co. Wekiva
River, Rutland Ferry, 7 mi. S Sorrento, Lake Co. OKLAWAHA RIVER DRAINAGE.

Lake Minneola, Clermont; Lake Lucy, 2 mi. N Groveland; both Lake Co. Mill
Creek, Apopka, Orange Co. Lake Harris; Sumter Co. Lake Eustis, Tavares;
Lake Saunders; Lake Griffin, all Lake Co. Lake Weir, Oklawaha; Juniper
Creek, 12 mi, E Lynn; Lake Lou, 5 mi. NE Lynn; Orange Creek, 1 mi. N Orange
Springs; Oklawaha River, Eureka Springs; all Marion Co. Magnesia Spring,



1972 JOHNSON: FLORIDA UNIONIDAE 211

3.5 mi. W Hawthorne; Cross Creek, 5 mi. NW Island Grove; Hatchet Creek,
Hawthorne Road, NE Gainesville; Hogtown Creek, 3 mi. SW Gainesville; all
Alachua Co. Morris Lake 12 mi. N Johnson; Lake Winnott, 4 mi. S Melrose;
Redwater Lake, 4 mi. W Johnson; Little. Orange Creek; all Putnam Co.
REE CREEK DRAINAGE: Rice Creek, 7.3 mi. WNW Palatka, Putnam Co.
HAw CREEK DRAINAGE: Lake Dias, 5 mi, ENE De Leon Springs, Volusia Co.
Lake Disston, 5 mi. SW Deanville, Little Haw Creek, 2 mi. SW Deanville; both
Flagler Co. BLACK CREEK DRAINAGE, Black Creek, 2 mi. E [town ofl Kingsley
Lake; North Fork, Black Creek, 14 mi. SW Orange Park; South Fork, Black
Creek, 1 Ini. SW Middleburg; South Fork, Black Creek, 8 mi. E Kingsley;
Black Creek, 10 mi. NW Green Cove Springs; all Clay Co.

Elliptio (Elliptic) buckleyi (Lea)
Figures 48, BI, 7A-G

Unio buckleyi Lea 1843, Dese. Twelve Uniones (Lake George and Lake Mon-
roe, Florida). Lea, 1846, Trans. Amer. Philos. Soc. 9: 276, pl. 39, fig. 2, figured
holotype USNM 85236, from Lake George. Lea, 1848, Obs. Unio, 4.34.

Unio cunninghanti B. H. Wright 1883, Proc. Acad. Nat. Sci. Phila., p. 58, pl. 1,
figs. 1-4 (Lakes of Sumter Co., Florida; lectotype ANSP 41348a, selected by
Johnson, 1967, Occ. Papers on Moll,, 3. 5, pl. 13, fig, 2 and the type locality
restricted to Lake Harris, Yalaha, Lake Co., Florida).

Unio orcuttii S. H. Wright 1888, West American Scientist, 4: 60, pl. 3 (Manatee
River [Manatee Co.] ; West Coast of Florida, Lake Myakka; figured holo-
type USNM 309971, refigured by Johnson, 1967, Oce. Papers on Moll., 3: 7,
pl. 9, fig. 1 and the type locality restricted to [Horse Creek, near] Mana-
tee River [Manatee Co. ], Florida. As this species has not been found
subsequently in either the Manatee or Horse Creek drainages, the type
locality is probably erroneous, and is here restricted to Upper Myakka
Lake, Sarasota Co., Florida.)

Unio dalli B. H. Wright 1888, Proc. Acad. Nat. Sci. Phila., p. 119, pl. 6, fig. 1 (Lake
Beresford, Volusia Co., Florida; figured holotype USNM 151037, refgured
by Johnson, 1967, Oce. Papers on Moll,, 3: 5, pl, 10, fig, 2).

Unto dorei B. H. Wright 1888, Proc. Acad. Nat. Sci., Phila., 115, pl. 3, fig. 1 (Lake
Monroe, Florida; holotype USNM 151034, refigured by Johnson, 1967.
Occ. Papers on Moll.,.3: 6, pl. 9, fig. 2).

Unio hinklegi B. H. Wright 1888, Proc. Acad. Nat, Sci. Phila., p. 117, pl 4, fig. 3
(Lake Monroe, Florida; holotype USNM 151033, refigured by Johnson,
1967, Occ. Papers on Moll., 3: 7, pl, 11, fig. 1)

Unio ferrissii Marsh 1891 , Nautilus , 5 : 30.( a small creek near Palatka [ Putnam
Co.], Florida; location of type not known, but figured by Simpson, 1892,
Proc. U. S. Natl. Mus., 15: 423, pl. 66, figs. 1-2).

Unio pi,lei B. H. Wright 1897, Nautilus, 11: 40 (unnamed lake in Witthacoo-
chee [Withlacoochee] River region of Hernando Co., Florida, holotype
USNM 150127, figured by Simpson, 1900, Proc. Acad. Nat. Sci. Phila., p. 80,
pl. 3, fig. 1, and refigured by Johnson, 1967, Oce. Papers 08 Moll.,3 8, p. 10,
fig. 5).

Unio ( Unio) caloosaensis Dall 1898, Trans. Wagner Free Inst. Sci., 3 (4): 688, pl



212 BULLET.IN FLORIDA STATE MUSEUM Vol. XVI No. 4

25, figs. 5, 12 b (Pliocene maris of the Caloosahatchie River, Florida, type
USNM 107745 [not seen]).

Unio subluridus Simpson 1900, Proc. U. S. Natl. Mus., 15: 432, pl. 73, figs. 3-4
(Orange Springs, Volusia [Marion] Co., Florida; figured holotype USNM
104002).

Unio tenuisculus Frierson 1911, Nautilus, 25: 29, pl. 1, figs. 4-6 (Reedy Lake,
Polk Co., Florida, lectotype, here selected, UMMZ 96321, portrayed in
figs. 4,6)

Unio (Elliptio) sanctrumjohanium B. H. Wright 1933, Nautilus, 47: 17, pl. 1
(Lake Druid, near Floral City [Citrus Co.] Florida; holotype USNM 424738,
refigufed by Johnson, 1967, Occ. Papers on Moll., 3: 8, pl. 9, fig. 3).

Unio (Elliptio) webbianus B. H. Wright 1934, Nautilus, 47: 94, pl. 10, figs. 1-2
(Lake Consuelo [or Little. Lake, SE Floral City, just outside village limits]
near Floral City, Citrus Co., Florida; holotype USNM 424923, refgured
by Johnson, 1967, Occ. Papers on Moll., 3: 10, pl. 9, fig.  4).

Unio (Elliptio) webbianus hartii B. H. Wright 1934, Nautilus, 47: 95, pl. 10, figs.
3-4 (Lake Consuelo [or Little Lake, SE Floral City, just outside village
limits] near Floral City, Citrus Co., Florida; holotype USNM 424925, re-
figured by Johnson, 1967, Otc. Papers on Moll.. 3: 6, pl. 9, fig. 5)

Popenaias buckleyi (Lea). Heard and Guckert, 1970, Malacologia, 10: 340, 348.

DESCRIPTION. -Shell generally small to medium, seldom reaching
100 mm in length. Outline somewhat variable though generally subtrape-
zoidal, sometimes obovate. Valves subinflated to inflated, subsolid to very
solid, inequilateral. Anterior end often subtruncate, sometimes regularly
rounded; posterior end slightly biangulate near the base, but not very pro-
duced. Ventral margin straight or slightly curved. Dorsal  margin straight,
sometimes forming a wing and a sharp angle with the obliquely descending
posterior margin hinge ligament short and low. Posterior ridge often rather
sharp, though sometimes rounded. Posterior slope rather flat, occasionally
with faint radial sculpture. Umbos rather full and high, located in the
anterior third of the shell, their sculpture consisting of several double-
looped ridges. Disk surface fiat. Periostracum varying from fine and shiny

FIGuRE 71. Ellipcio (Elliptic) buckle!/i (Lea). A. Holotype of Unio buckleyi Lea. Lake
George, Florida. USNM 85236. L 93, H 48, W 38 (slightly reduced). B. Lecto-
type of Unio cunninghami B. H. Wright. [Lake Harris, Yalaha, Lake Co.], Flor-
ida. ANSP 41348a. L 48, H 29, W 24 (nat. size, reversed image). C. Holotype of
Unio dallii B. H. Wright. Lake Beresford, Volusia Co., Florida. USNM 1511037. L
63, H 35, W 21 (slightly reduced). D. Holotype of Unio orcuttii S. H. Wright. [Up-
per Lake Myakka, Sarasota Co., Florida]. USNM 308971. L 58, H 34, W 25
(slightly reduced). E. Lectotype of Unio tenuisculus Frierson. Reedy Lake, Polk
Co., Florida, UMMZ 96312, L 36.5 H 20, W 14.5 (nat: size, reversed image);. F.
Holotype of Unio dorei B. H. Wright. Lake Monroe, [Volusia Co.], Florida.
USNM 151034,.L 65, H 40, W 31 (slightly reduced). G. Holotype of Unio sublur-
idus Simpson. Orange Springs, Volusia [Marion] Co., Florida. USNM 104002.
L 41, H 25, W 15 (slightly  reduced),
1 Measurements in mm, L = length, H = height, W = width.



1972 JOHNSON: FLORIDA UNIONIDAE 213

4

.i

A .5 1~1

!92~

F  G ~--awfllyle,



214 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

yellowish or greenish with numerous darker green rays to rough and black
or brownish.

Left valve with two rather variable, stumpy pseudocardinal teeth,
often of equal height, but with the hinder one sometimes higher. Hinge line
very short and narrow; two long, straight lateral teeth. Right valve with two
roughly parallel pseudocardinals, the posterior one apt to be serrated and
chunky, the more anterior one low and vestigial; one lateral tooth. Beak
cavities very shallow with a few dorsal muscle scars. Anterior and pos-
terior muscle scars and pallial line all distinct. Nacre purplish, salmon,
bluish white, or pinkish, posteriorly iridescent.

MEASUREMENTS, - L 94 171111, H 50 min, W 36 min (Lake George, holo-
type of U. buckle!/i Lea); L 73 mm, H 38 mm, W 25 mm (Lake Monroe,
Volusia Co., holot>,pe of Unio hinkle!/i Wright); L 48 nini, H 29 min, W 24
mm (Lake Harris, Yalaha, Lake Co., lectotype of Unio cunninghami
Wright).

HABITAT. -Found .iii lakes, ponds, small streams, and lizrge rivers, in
nearly every type of substrate, but seems to prefer sandy bkittoins.

REMARKS. - E//iptio buckleui (Lea) is endemic to peninsular Florida
and exhibits considerable variation, but it can be confused only with the
considerably more variable forms of the more widely spread E. icterina
(Conrad). E. buckle!/i tends to be rather consistently subtrapezoidal, less
apt to be produced posteriorly and generally more inflated with a distinct
posterior ridge. Many specimens are characterized by the anterior dorsal
margin forming a sharp angle with a somewhat truticated anterior margin.

While many examples from favorable environments are shiny yellow-
ish-brown or greenish, sometimes with fine green rays, others, especially
from some of the larger lakes, tend to have coarse brownish or blackish
periostracum, are dwarfed with heavier shells, and have a tendency for the
posterior end to extend below the base line. These forms from the larger
lakes were named U. cunninghami Wright. A diminutive population in
Reedy Lake, Polk Co., was named U. tenuisculus Frierson.

RANC:E.- Peninsular Florida.

SPECIMENS EXAMINED.-WITHLACOOCHEE RivER SYSTEM: Lake, 6 mi. NNW

Panasoffkee; Lees Lake, Panasoffkee; both Sumter Co. Lake Druid, near
Floral City; Little Lake, SE Floral City; Lake Tsala Apopka, Hernando; all
Citrus Co. Withlacoochee River, near Inglis; Levy Co. MYAKKA RIVER SYS-

TEM: Myakka River, near Manatee [River]; Manatee Co. Upper Lake Myakka;
Myakka River between upper athd lower lakes, both Sarasota Co. PEACE

RlvER SYSTEM: Lake Agnes, Polk· City, Lake Gibson, Gibsonia; Lake Alfred.
[town of] Lake Alfred; Lake Rochelle, 1 mi. S [town of] .Lake Alfred; Lake
Fannie, 3 mi. SE [town of] Lake Alfred; Lake Hamilton, 4 mi. SSW Haines,
Citv. Lake Howard, Winter Haven; Lake Silver, Winter Haven; Lake Roy,
Winter Haven; Lake M¢Leod [town· of] Eagle Lake; Dinner Lake, 1,5 mi, ESE
Waverly; all Polk Co. KISSIMMEE RIVER SYSTEM AND EvERGLADES: ISTOKPOCA

RIVER DRAINAGE: Lake Lee, Waverly; Lake Starr, [town of] Lake of the

Hills; Lake Amoret, Highland Park; Reedy Lake, 2.5 mi. E Frostproof, Lake
Arbuckle; Silver Lake, Frostproof; atl Polk Co. Lake Brent,vood, 2 mi. N



1972 JOHNSON: FLORIDA UNIONIDAE 215

Avon Park, Lake  Viola, N Avon Park; Lake Verona, Avon Park; Lake Lotela,
2 mi. SE Avon Park; Bonnet Lake, 3 mi. N Sebring; Dinner Lake, 2 mi. N
Sebring; Lake Jackson (Rex Beach Lake), 2 mi, NW Sebring; Lake  June in
Winter, 3 mi. W [town of] Lake Placid, Lake Istokpoga, 6 mi. E [town of] Lake
Placid; Istokpoga Canal, 4.6 mi. NW Cornwall; all Highlands Co. KISSIMMEE
RIvER DRAINAGE: East Tohovekaliga Lake, Narcoossee,  Emerald Lake, 1 mi.
SE St. Cloud; Alligat*r Lake, 4 mi. SE Asht., T6hopekaliga Lake, Kissimmee;
Lake Kissimmde; all Osceola Co, Lake Hatchineka; Lake Aurora, Hesperida;
Lake Rosalie, 4 mi. NE Hesperides; Tiger Lake, 7 mi. ESE Hesperides; Weo,
hyakapa Lake,.4 mi. SE Hesperides; all Polk Co. Lake Marion, 3 mi. WNW
Kenansville, Osceola Co. Kissimmee River, Basinger; Gum Tree Slough, near
Basinger; Kissimmee River, 10 mi. S Basinger; Taylor Creek; all. Okeechobee
Co. LAKE ~KEECHOBEE DRAIMAGE: Fisheating Creek, Palmdale; Fisheating
Creek, 1 mi. S Lakeport; Caloosahatchie River, Moore Haven; all Glades Co.
Lake Okeechobee, 2 mi. N Canal Point; Lake Osborne, 2.5 mi. NW Lantana;
Clear Lake, West Palm Beach; all Palm Beach Co. Lake Okeechobee, 2.4 mi:
N Port Mayaca, Martin .Co. ST: JOHNS RivER SYSTEM.-ST, JOHNS -RIVER
DRAINAGE: St. Johhs Marsh; ditch near Lake Washington, 6 mi. W Eau Callie;
Lake Poinsett, Rockledge; all Brevard Co. Puzzle Lake, 7 mi. SE Geneva, Lake
Harney, 3 mi. NE Geneva; tributary of Econlockhatehee River, 2.5 mi ENE
Oviedo; Lake Jessup, 5 mi. N Oviedo, St. johns River [town of] Lake Monroe;
all Seminole Co. Lake Monroe, near Enterprise; Lake Woodruff, both Volusia
Co. Econlackhatchee ,River, E Orlando, Orange Co. :St, Johns River, Crows
Bluff, St. Johns River, Aston Lake George; all Lake Co, OKLAwAHA RIVER
DRAINACE: Lake Builer, Windermere; Lake Bessie, Windermere; Lake Down,
Windermere; all Orange Co. Lake' Lucy, 2 mi. N Groveland; Pine Island Lake,
6 mi, S Groveland; Lake Minnehaha, 2 mi. S Clermont, Lake Minneola, Cler-
mont; Lake Apopka, 2.5 mi.  S Monteverde; all Lake Co. Lake Carleton, 2 mi.
W Tangerine, Ofange Co. Lake Griffin, Leesburg; Lake Beauclaire, 2 mi. S
Mt. Dora; Lake Dora, Tavares; Lake Gertrude, Mt. Dora; Lake Saunders, 1 mi.
E Tavares; Lake Harris, 4. mi. NW Tavares; Silver Lake, 5 mi. NW Tavares;
Lake Griffin, Leesburg,  Lake Eustis, Tavares; Lake Yale, Grand Island;
[ Haines Creek] 5 mi. W Eustis; Oklawaha River, all Lake Co: Lake. Weir,
Oklawaha; Oklawaha River; Lake Kerr, 3 mi,.SW Kerr City; all Marion Co.
Cross Creek, 5 mi. NW Island Grove, Alachua Co. Little Orange Creek, 3 mi.
E Hawthorne, ditch, W of bridge, Orange Creek, Orange Spring; both Putnam
Co. HAw CREEK DRAINAGE: Lake Stella, Crescent City, Putnam Co. JULINGTON
CREEK DRAINAGE. Durbin Creek, 1 mi. WSW Bayard; lake on Julington Creek,
2 mi: W Bayard, both Duval Co.

E/.liptio (Elliptio)jagensis (Lea)
Figures 3C, 8 A-F, 9A

Unio jagensis Lea 1838 Trans. Amer. Philos. Soc., 6: 28, pl. 9, fig: 23 (Flor-
ida; Tgured hol6type USNM 86031). Lea 1838 Obs. Unio, 2: 28.

Unio aheneus Lea 1843, Desc. Twelve Unione5 (Black Creek, Florida), Lea,
1846, Trans. Amer. Philos. Soc., 9: 280, pl. 41, fig. 9, figured holotype
USNM 86030. Lea, 1848, Obs. Unio, 4; 38.

Unio waltoni B.H, Wright 1888, Proc. Acad. Nat. Sci. Phila., p. 114, pl 2, fig. 3



216 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

B - ~f~

/5 .r««~~ r-

D --

02-49.90-:-*'.lir.
- .: 7.'-

-

- ~irs*lul,

FIGURE 81. Elliptio ( Elliptio) jagensis (Lea). A. Holotype of Unio aheneus Lea. Black
Creek, Florida. USNM 86030. L 53, H 24, W 14. B. Lectotype of Unio buxtoni
B. H. Wright. Lakelets of Marion Co., Florida. USNM 150131. L  46, H 20, W 15.
C. Lectotype of Unio oscari B. H. Wright. A creek from Lake Osceola, Winter
Park, [Orange Co. ], Florida. USNM. 123526. L 59, H 26, W 23. D. Lectotype of
Unio waltoni B. H. Wright. Lake Woodruff, Volusia Co., Florida. USNM 91145.
L 78, H 28, W 16. E. Holotype of Unio jagensis Lea. Florida. USNM 86031. L 652
H 32, W 22, based on single valve. F. Holotype of Unto tryoni B. H. Wright. Lake

Woodruff [Spring Garden Lake], near De Leon Springs, Volusia Co., Florida.
USNM 151036. L 98, H 45, W 32.
1 Slightly reduced, measurements in mm, L = length, H = height, W = width.



1972 JOHNSON: FLORIDA UNIONIDAE 217

(Lake Woodruff, Volusia Co., Florida; lectotype USNM 91145 selected
by Johnson, 1967, Oce, Papers on Moll., 3: 9, pl: 11, fig. 5).

Unto marshii B. H. Wright 1888, Proc. Acad. Nat. Sci, Phila., p. 118, pl. 5, fig, 2
(Lake Woodruff, Volusia Co., Florida; holotype USNM 151028 from [St.
Johns River] Blue Springs [3 mi. S Lake Beresford], refigured, and the
type locality thus restricted, by Johnson, 1967, Oce. Papers on Moll., 3
7, pl. 12, fig. 2).

Unio trgoni B: H. Wright 1888, Proc. Acad. Nat. Sci, Phila., p. 120 pl. 6, fig. 2
(Lake Woodruff [Spring Garden Lake] near DeLeon Springs, Volusia
Co., Florida; holotype USNM 151036, refigured by Johnson, 1967, Occ
Papers on Moll., 3. 9, pl. 12, fig. 1).

Unio oscari B. H. Wright 1892, Nautilus, 5: 124 (a creek from Lake Osceola,
at Winter Park [Orange Co.], Florida). Wright, 1896, Nautilus, 9: 122,
pI. 2, figs. 1-3, lectotype USNM 123526, selected by Johnson, 1967, Occ.
Papers on Moll, 3: 8, pl, 11, Fig. 3.

Unio yuttoni B. H. Wright 1897, Nautilus, 11: 56 ( [Smith] Lake, near Candler,
Marion Co., Florida; holotype USNM 150129 figured by Johnson, 1967,
Oce. Papers on Moll., 3: 9, pl. 11, fig. 2).

Union buxtoni B. H, Wright 1897, Nautilus, 11: 55 (Lakelets of Marion Co.,
Florida; lectotype USNM 150131, selected by Johnson, 1967, Occ, Papers
on Moll. 3: 5, pl: 11, fig. 4).

DESCRIPTION. -- Shell generally medium in size, not exceeding 70 min
in length over most of its range, except in the St, Johns River system
where specimens often reach 90 mm. Outline elongate-trapezoidal, or
elongate-oval, often a little more than twice as long as high. Valves
usually compressed or sub-compressed, occasionally subinflated, thin to
subsolid, inequilateral. Anterior end regularly rounded; posterior end
more or less sharply pointed, often below the medial line. Ventral margin
straight or slightly curved. Dorsal margin straight, generally forming a
sharp angle with the obliquely descending posterior slope. Hinge liga-
ment rather long and low. Posterior ridge subangular with a secondary
ridge above it; the ridges sometimes cause the point to be slightly biangu-
late. Umbos very low, located in the anterior fourth of the shell, their
sculpture consisting of corrugated, longitudinal ridges. Disks usually flat,
rarely slightly concave, because of a slight umbonal-ventral sulcus. Pei·io-
stracum fine, subshiny, greenish yellow, with very fine green rays of vary-
ing width, especially when young or from favorable habitats, often be-
coming rough and black with age.

Left valve with two stumpy pseudocardinal teeth, one in front of the
other, both somewhat triangular; the hinder one may be vestigial. Hinge
line rather short and narrow; two long, straight lateral teeth. Right valve
with two roughly parallel pseudocardinals, the posterior one inclined to
be trial-Igular, serrated, and chunky, the more anterior one low and ves-
tigial; one lateral tooth. Beak cavities very shallow, with a few dorsal
muscle scars. Anterior and posterior adductor muscle scars and pallial



218 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

line all distinct. Nacre white, bluish-white, pinkish, purplish, posteriorly
iridescent.

MEASUREMENTS. - L 93 111111, H 43 nim, W 28117111 (Lake Beresfoicl, Vo-
lusia Co.); L 93 mm, H 33 mm, W 22 inm (Lake Woodruff, Volusia Co.,
paralectotype of U. waltoni Wright); L 57 mm, H 28 inm, W 17 mm
(Little Haw Creek, 2 mi S of Deanville, Flagier Co.).

HABITAT. - G.enerally lives iii sand, but sometimes in mud, usually
where the current is not very swift.

REMARKS, -Elliptio jauensis (Lea) can only be confused witli E. ic-
terina (Conrad) and E. buckle!/i (Lea) in peninsular Florida, but it is
usually easily distinguished from either of them because it is more elon-
gate, being generally over twice as long as high.

The shell varies considerably in the ratio of height to length, the
sharpness of the posterior point, whether the point is above or below the
medial line, the degree of inflation, and the thickness of the valves. When
they are visible the very fine green rays of varying width are characteristic
and do not resemble those of any other Floridian unionid.

E. jal/ensis is close to E. lanceolata (Lea)  (Johnson, 1970: 333),
which is found in the Apalachicolan and Atlantic Slope regions, but not
in peninsular Florida. While both species exhibit many similar variations,
jayensis is generally smaller, with a tendency to be higher posteriorly.
The dorsal margins are not generally parallel, and jagensis has more
numerous, finer, darker green rays.

E. jagensis (Lea) was described from a single, thin, not very char-
acteristic valve, probably from the St. Johns River system where the species
is most abundant and where individuals reach the greatest size and
diversity of form, especially in the several great lakes of the system.

RANGE. -Apalachicol:,11 region: St. Marks River system. Suwannee

River system. Peninsular Florida.
~PECIMENS EXAMINED.-PITHLACHASCOTEE RIVER SYSTEM': Pithlachascotee

River, 2 mi. W Gowers Corner; Pasco Co. HILLSBOROUGH RIVER SYSTEM: Hills-

borough River, Morris Bridge, 14 mi. NE Tampa; Hillsborough Co. ALAFIA

RIVER SYSTEM: Fishhawk Creek, 2 mi. S Lithia, Hillsborough Co. PEACE RrVER

SYSTEM: Peace River Drainage. Peace River, Arcadia, De Soto Co. KISSIMMEE
RIVER SYSTEM: AND EVERCLADES: ISTOKPOCA DRAINACE. Tohopekaliga

Lake, Kissimmee; Osceola Co. Lake Istokpoga, 6 mi. E [towh of] Lake Placid;

FIGURE 91. Elliptio (Elliptio) jagensis (Lea): A. Holotype of Unio marshil B. H.
Wright. [St. Johns River] Blue Springs [3 mi. S of Lake Beresford, Volusia Co.,
Florida]. USNM 151028. L 90, H  47, W 33 (slightly reduced).

Uniomerus tetralcismus (Say). B. Holotype of Unio blandingianus Lea. St.
Johns River, Florida, USNM 85715. L 57, H 35, W 22 (slightly reduced). C.
Holotype of Unio jewettii Lea. Sink of Noonanis [Newnan'sl Lake, [Alachua
Co.],Florida. USNM. 85374. L 49, H 28, W 17 (slightly reduced). D. Lecto-
type of Unio paludicolus Gould. Everglades of Florida. MCZ 169278. L 46, H
39, W 14 (1.1 X): E. Holotype of Unio buddiant,s Lea. Lake George, Florida.
USNM 85606. L 97, H 53, W 31 (slightly reduced).
1 Measurements in mm, L = length, H = height, W = width.



1972 JOHNSON: FLORIDA UNIONIDAE 219

Istokpoga Canal, 4.5 mi. NW Corwell; both Highlands Co. KISSIMMEE RIVER
DRAINAGE, Lake Arbuckle; Lake Rosalie, 4 mi. NE Hesperides; Tiger Lake,
7 mi. ESE Hesperides; all Polk Co. LAKE OKEE(HoBEE DRAINAGE. Fisheating
Creek, 1 mi. S Palmdale; Caloosahatchie Biver, above lock, Moore Haven; all
Glades Co. Cal-~als, West Palm Beach, Palm Beach Co. Lake Okeechobee.
ST. JOHNS RIVER SYSTEM: ST. JoHNs RIvER DRAINAGE. Little Econlockhatchee
River, 3.5 mi. S Oviedo; Econlockhatchee River, near confluence with St. Johns
River; Gee Creek, near North Orlando; St. Johns River, 4 mi. E Sanford,

B



220 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

all Seminole Co. Lake Ashby, 8 mi. NE Osteen; Lake Woodruff, both
Volusia Co. LAKE REGION DRAINAGE. Lake Maitland, Winter Park; Creek
from Lake Osceola, Winter Park; both Orange Co. OKLAWAHA RIVER
DRAINACE: Smith Lake, 1 mi. W Candler; Oklawaha River, 1 mi. E White
Ferry; Oklawaha River, Moss Bluff 5 mi. NE Lynn, Orange Creek, 1 mi. N
Orange Sprihgs; all Marion Co. Redwater Lake, 4 mi. W Johnson; Putnam Co.
HAw CREEK DRAINAGE: Lake Disston, 5 mi. SW Deanville; Little Haw Creek,
2 mi. SW Deanville; both Flagler Co. BLACK (REEK DRAINAGE: Black Creek,
2 mi. E [town of] Kingsley Lake; North Fork, Black Creek, 14 mi. SW Orange
Park; Black Creek, 10 mi. NW Green Cove Springs; all Clay Co. JULINCTON
CREEK DRAINAGE. Durbin Creek, 1 mi. WSW Bayard; lake on Julington Creek,
2 mi. W Bayard; both Duval Co.

Genus Uniomerus Conrad
Uniomerus Conrad 1853, Proc, Acad. Nat. Sci. Phila., 6: 268. Species listed:

U. decliuis Say, camptodon Say, subcroceus Conrad, sayii Ward, ri-
oularis Conrad, perrectus [sic] Conrad, symmetricus Lea, excultus
Conrad.

Type species, Unio tetralasmus Say. Subsequent designation, Simpson,
1900, Proc. U.S. Natl. Mus., 22: 739. Since U. excultus Conrad is includ-
ed in Simpson's synonymy of U. tetralasmus, the subsequent selection of
the former by Cleneh and Turner (1956, Bull. Florida State Mus., 1: 176)
is invalid under Article 69 (a) (iv) of the Int. Code Zool. Nomen. (1964).
Ortmann, 1912, Ann. Carnegie Mus.,8: 272.
Frierson (1927: 34-35) listed a number of species and subspecies under

Uniomerus. Like Elliptio, Uniomerus, has a wide range of environmental
tolerance, and while there are a number of ecophenotypes, the genus appears
to be monotypic. This view is supported by Fuller (1971: 142)

Uniomerus tetmlasmus (Say)
Figure 2A, 9 B-E.

Unio tetralasmus Say, [September] 1831, American Conchology, no. 3 [no
pagination], pl. 23 (Bayou St. John [not located], near New Orleans,
Louisiana; type not in ANSP [lost]).

Unio obesus Lea 1831, Trans. Amer. Philos. Soc., 4: 96, 108, pl, 13, fig. 26
(York River, Virginia [corrected to Georgia, Maj. Leconte on p. 108] ;
figured holotype USNM 85366, labeled, "Little Ogeechee River [ Hancock·
Co], Georgia; Maj. Leconte"). Lea. 1834, Obs. Unio, 1: 106, 118. Clench
and Turner (1956, Bull. Florida State Mus., 1: 178) did not see the type,
and their restriction of the type locality is invalid.
Lea (1854, Proc. Acad. Nat. Sci. Phila., 7:243) claimed that this description
appeared toward the end of 1831. It was reported during 1832 (Jan.- March
number: Amen Jour. Sci., 22: 169 [probably appeared in April] ). There
is no way to be sure which name has priority, but Say's name is certainly
better known, and should be used for this species.

Unio dectivis Say, 1831 [1832], Transylvania Jour. Medicine, 4: 527 (Bayou



1972 JOHNSON: FLORIDA UNIONIDAE 221

Teche, Louisiana). Say, 1832, American Conchology, no. 4 [no pagination],
pl. 35; 3 syntypes ANSP 41698 from Mr. Barabino, all smaller than fig-
ured type).

Unto camptodon Say; 1832, American Conchology, no, 5 [no pagination], pl
42 (opposite to New Orleans [ Jefferson Parish, Louisiana], in ponds;
type not. in ANSP [lost])

Unio· geometr.icus Lea 1832,  Trans. Amer. Philos. Soc., 5: 38, pI. . 4, fig. 10
(Bayou Teche, L.6uisiana; figured holotype USNM 85712 [not seen])
Lea, 1834, Obs. Unio, 1: 150.

Unio blandingianus Lea 1834, Trans. Amer. Philos. Soc., 5: 101, pl. 15, fig.
44 (St. Johns River, Florida; figured holotype USNM 85715). Lea, 1834,
Obs. Unio, 1: 213.

Unio dectivis Conrad, 1836, non Say. See under: Unio rivularis Conrad, 1853.
Unio excultus Conrad 1838, Monography Unionidae, no. 11, p. 99, pl. 55, fig.

1 (New Orleans [Orleans Parish], Louisiana; ty~e.ANSP 20427 [lost]).
Unio sayii Ward 1839, [in  Tappan], Amer. Jour. Sci., 35: 268, pl. 3, fig. 1

(Walnut Creek, and Ohio Canal, near Cirdeville [Pickaway Co.], Ohio;
[location of type' unknown]).

Unio paratellus Conrad 1841, Proc. Acad. Nat. Sci. Phila., 1: 20 non Sowerby
1840. Changed to:

Unio porrectus Conrad 1854, Jour. Acad. Nat. Sci. Phila. (2).2. 296, pl. 26,
fig, 7 (,[Pearl River] Jackson [Hinds Co.], Mississippi; figured holotype
ANSP 42847).

Unio buddianus Lea 1843, Desc. Twelve Uniones (Lake George and Lake
Monroe, Florida). Lea, 1845, Trans. Amer. Philos. Soc., 9: 277, pl. 40,
fig, 5, figured holotype USNM 85606, from Lake George. Lea, 1848, Obs,
Unio, 4: 35.

Unio symmetricus Lea 1845, Trans. Amer. Philos. Soc., 10: 73, pl. 4, fig. 11
(Red River, Alexandria [Rapides Parrish], Louisiana; figured holotype
USNM. 85604), Lea, 1848, Obs: Unio, 4: 47,

Unio paludicolus Gould 1845, Proc. Boston Soc. Nat. Hist., 2: 53 (Everglades
of Florida; lectotype MCZ 169278, selected by Johnson, 1964, U.S. Natl.
Mus., Bull. 239, p. 121, pl, 31, fig. 3).

Unio ineptus Lea 1852, Trans. Amer. Philos. Soc., 10: 261, pl. 15, fig. 12 (Abbe-
ville District [Savannah River drainage], South Carolina; figured holo-
type USNM 85326). Lea, 1852, Obs. Unio, 5: 17.

Unto hebes Lea 1852, Trans. Amer. Philos. Soc., 10: 267, pl. 18, fig. 21 (Oconee
River, near Athens [Clarke Co. 1, Georgia; figured holotype USNM 85383).
Lea, 1852, Obs. Unio, 5: 23.

Unio rivularis Conrad 1853, Proc. Acad. Nat. Sci. Phila., 6: 257. New name
for Unio decliuis Conrad 1836, non Say 1831, [ in] Monography Union-
idae, no. 5, p.. 45, pl 23, fig. 1 (small creek in Greene Co., Alabama; figured
holotype ANSP 42852).

Unio paludicolor Conrad 1853, Proc: Acad. Nat. Sci. Phila., 6: 254. Error for
U paludicolits Gould.

Unio subcroceus Conrad 1854, Jour. Acad. Nat. Sci. Phila., (2) 2: 297, pl. 27,
fig. 1 (one of the tributaries to Canadian River, Arkansas; figured holotype
ANSP 41406a



222 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Unio manubius Gould.1855, Proc. Boston Soc. Nat. Hist., 5: 229 (Chihuahua,
60 mi. from Camp Ringgold = Rio Agualeguas, 3 mi. NE General Trevino,
Nuevo Leon [State, Mexico], teste Taylor, 1967, Veliger, 10: 154; holotype
MCZ 169447, fgured by Johnson, 1964, U.S, Natl. Mus,, Bull, 239, p. 108,pl. 32, fig. 5).

Unio columbensis Lea 1857, Proc. Acad. Nat. Sci. Phila:, 9: 31 (Creeks near
Columbus [Muscogee Co.], Georgia). Lea, 1858, Jour. Acad. Nat. Sci.
Phila., (2) 4: 75, pl. 14, fig. 55; figured holotype USNM 85360. Lea, 1858,
Obs. Unio, 6: 75.

Unio jamesianus Lea 1857, Proc, Acad, Nat, Sci, Phila., 9: 84 C  [Pearl ~ River]
Jackson [Hinds Co.], Mississippi). Lea, 1858, Jour, Acad, Nat. Sci. Phila.,
(2) 4: 53, pl. 6, fig. 35; figured holotype USNM 85365). Lea, 1858, Obs.
Unio, 6: 52.

Unio plantii Lea 1857, Proc. Aead. Nat. Sci. Phila., 9: 171 (Flint River, near
Macon [Co.], Georgia), Lea, 1859, Jour, Acad, Nat, Sci. Phila., (2) 4:
192, pl. 21, fig, 76; figured holotype USNM 85005. Lea, 1859, Obs. Unio,
7: 10. [Known only from the holotype which is a pathological specimen.].

Unio cicur Lea 1861, Proc. Acad. Nat. Sci. Phila., 13: 39 (Little Ocmulgee
River, Georgia). Lea„ 1862, Jour. Acad. Nat. Sci. Phila,, (2) 5: 93, pl, 13,
fig. 241; figured holotype USNM 85532. Lea, 1862, Obs. Unio, 8:97.

Unio squalidus Lea 1863, Proc, Acad. Nat. Sci. Phila., 15: 192 (Neuse Rivers
near Raleigh [Wake Co. ] ; Roanoke River, near Wheldon [Halifax Co.],
Deep River,; all North Carolina), Lea, 1866, Jour. Acad, Nat, Sci, Phila,,
(2) 6: 22, pl, 7, fig. 20; figured holotype USNM 85376, from Roanoke
River. Lea, 1867, Obs. Unio, 11: 26.

Unio, electrinus Reeve 1865, Conch, Iconica, 16, Unio, pl. 25, fig: 121 ([locality
unknown] Cuming coin, type, British Mui. (Nat, Hist,) [lost]).

Unio bisselianus Lea 1867, Proc. Acad, Nat. Sci. Phila., 15: 81, (Bissels
Pond, Charlotte [Mecklenberg Co.], North Carolina). Lea, 1868, Jour,
Acad. Nat. Sci. Phila., (2) 6: 277, pl. 37, fig, 90; figured holotype USNM
85373. Lea, 1869, Obs. Unio, 12: 37.

Unio jewetta Lea 1867, Proc. Acad. Nat, Sci. · Phila,, 11: 81 (sink of Noo-
nan's [Newnansl Lake [Alachua Co.], Florida). Lea, 1868, Jour. Acad,
Nat. Sci. Phila., (2) 6: 276, pl. 37, fig. 89; figured holotype USNM 85374.
Lea, 1869, Obs. Unio, 12: 36.

Unio rioicolus Conrad 1868, Amer. Jour. Conch., 4: 280, pl, 18, Rg. 4 (brook
near Tampa [Hillsborough Co.], Florida; figured holotype ANSP 41411).

Unio pawensis Lea 1868, Proc. Acad, Nat. Sci. Phila., 20: 161 (Paw Creek
[Mecklenberg Co.], Beaver Co. [ = Creek, Gaston Cd.], Catawba Run
[Gaston and Mecklenberg Cos,] ). Lea, 1868, Jour. Acad. Nat. Sci. Phila.,
(2) 6: 302, pl. 45, fig, 114, figured holotype USNM 85380, labeled, "Beaver
Creek, [into?] Catawba Run, North Carolina". Lea, 1869, Obs. Unid,
12: 62.

Uniomerus obesus (Lea). Clench and Turner, 1956, Buil. Florida State Mus.,
1: 177, pl. 5, fig. 2.

Unionierus tetralasmus (Say). La Rocque, 1967, Ceol. Survey, Ohio, Bull., 62(2): 178 fig. 67.



1972 JOHNSON: FLORIDA UNIONIDAE 223

Uniomerus tetralas,nus camptodon (Say). La Rocque, 1967, Geol. Survey,
Ohio, Bull., 62 (2): 180.

Uniomerus tetralasmus sagi (Ward), La Rocque, 1967, Geol. Survey, Ohio,
Bull., 62 (2): 181.

Uniomerus tetralasmus (Say)  Johnson, 1970, Bull. Mus. Comp. Zool., 140
(6): 339, pl. 12, figs 1-6.

Uniomerus tetralasmus (Say). Valentine and Stansbery, 1971, Sterkiana, no.
42, p. 22.
DESCRIPTION. - Shell generally medium, though sometinies large

reaching over 114 mm in length. Outline rhomboidal or elongate rhom-
boidal. Valves subinflated or inflated, subsolid. Anterior end regularly
rounded or slightly truncated; posterior end usually somewhat produced.
Ventral margin incurved. Dorsal margin curved, generally forming a
sharp angle with the almost straight posterior margin. Hinge ligament,
long and narrow, located posterior to the umbos. Posterior ridge rounded,
ending in a point or feeble biangulation at the base of the shell, some-
times rendering older specimens a bit arcuate. P.osterior slope often with
two radial sulci. Umbos low to slightly elevated, located in the anterior
quarter of the shell, their sculpture consisting of fiv.e or six heavy ridges
that form a rounded angle on the posterior ridge, in front of which
they tend to be corrugated. Periostracum generally black and slightly
rough, but with a satiny sheen over most of the surface. Sometimes the
surface is smooth and shiny, especially in the umbonal area, and may
then be brownish-yellow, or yellowish mixed with green, but not rayed.

Left valve with two ragged, subequal pseudocardinal teeth, and two
straight lateral teeth. Right valve with one triangular pseudocardinal
often with a vestigial tooth above it; one lateral tooth. Beak cavities com-
pressed, but with several muscle scars; ariterior adductor muscle scars
deep, posterior ones faint. Pallial line distinct. Nacre white, bluish-
white, or pinkish to lurid purple.

MEASUREMENTS.-L 119 min, H 70 min, W 45 min (St. Johns River
[town ofl, Lake Monroe, Seminole Co.); L 98 mm, H 58 mm, W 41 mm
(Lake on Julington Creek, 2 mi W of Bayard, Duval Co.); L 58 min, H 36
mm, W 23 mm (Lake Louisa, 6 mi. SSE of Clermont, Lake Co.).

HABITAT. - Generally lives in smaller streams and ponds iii sand,

usually where the current is not swift.
REMARKS. - Uniomerits tetralasmus (Say) can be confused in the

Apalachicolan and Southern Atlantic Slope regions with Elliptio com-
planata (Lightfoot) and E. icterina (Conrad). In peninsular Florida, U.
tetralasmus is quite distinct from all species except E. icterina. In gen-
eral, tetralasmus is more inflated, proportionately higher, more acutely
angular where the dorsal margin meets the posterior one, and very often
has a characteristic satiny periostracum. The yellowish brown, unrayed
periostracum and bluish white or pinkish nacre differentiate tetralasmus
from icterina, which has a sometimes bright yellow or chestnut, often
rayed periostracum and more variety in nacre color.



224 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Uniomerus tetralasmus (Say) is generally common throughout the
Apalachicolan region, peninsular Florida, and Southern Atlantic Slope
rivers of Georgia. It becomes scarce in the Carolinas, where the periostra-
cum is more apt to be smooth, similar to specimens from the Interior
Basin. In peninsular Florida this species achieves its greatest size in the
St. Johns River system.

Under the name Uniomerus obesus, Valentine and Stansbery (1971:
22) call attention to the ecophenotypic variation of tetralasmus, which
they regard as a coastal species ranging from North Carolina to southern
Florida to Texas, by mentioning the occurrence of a population Of obesus
in Arkansas.

RANGE. - I,iterior B:~siii: Mississippi drainage, generally north to
about latitude 40', Ohio River. West Gulf Coastal region, Alabama-
Coosa River system, and Apalachicolan region: Rio Grande River system,
Texas, east to the Suwannee River system, Florida, Pehinsular Florida.
Southern Atlantic Slope: Altamaha River system, north to the Nottaway
River of the Chowan River system, North Carolina.

~PECIMENS EXAMINED.-WITHLACOOCHEE RIVER ~YSTEM:-Withlacoochee
River, 1 mi. W Lacoochee; Pasco Co. Little Withlacoochee River, Rerdell,
Hernando Co. Lake, 6 mi NNW Panasoffkee; Sumter Co. Lake Tsala Apopka,
Floral City, Citrus Co. Withlacoochee River, Dunnellon; Marion Co. WEEKI-
WACHEE RIVER DRAINAGE: Mud Springs, 3.5 mi. NW Berkeley; Hernando
CO. PITHLACHASCOTEE RIVER SYSTEM:-Pithlachascotee River, 2 mi. W Cow-
ers Corner; Paseo Co. HILLSBOROUGH RIVER SYSTEM:-Hillsborough River, 4
mi. NE Temple Terrace; Hillsborough Co, ALAFIA RIVER SYSTEM:-Branch of
Howells Creek, 3 mi. S Plant City; Hillsborough Co. MYAKKA RIVER SYSTEM:
-Upper Myakka Lake; Myakka River; both Sarasota Co. PEACE RIVER SYS·
TEM:-Haines Lake [town of] Lake Alfred; Lake Hamilton, 4 mi. SSW
Haines City; Silver Lake, Winter Haven, all Polk CO, PEACE RIVER SYSTEM:-
Payne Creek, 4.25 mi. N. Wauchula, Hardee Co. Wares Creek; Manatee Co.
Horse Creek, 8 mi. W Arcadia; Peace River, 1.25 mi. below Arcadia; both De
Soto Co. KISSIMMEE RIVER SYSTEM AND EVERGLADES:-ISTOKPOGA DRAINAGE:
Tohopekaliga Lake, Kissimmee; East Tohopekaliga Lake, Narcoossee; Emerald
Lake, 1 mi. SE St. Cloud; all Osceola Co. KissiMMEE RIVER DRAINAGE: Lake
Rosalie, 4 mi. NE Hesperides; Tiger Lake, 7 mi. ESE Hesperides; both Polk
Co. Gum Tree Slough, near Basinger; Taylor Creek; both Okeechobee Co.
LAKE OKEECHOBEE DRAINAGE  Fisheating Creek, 1 mi, S Lakeport; _ 4 mi. S Fort
Myers, Lee Co. (USNM). Cypress Swamp,. 4 mi. S Monroe Station, Collier
Co. Paradise Key, Royal Palm Park, Everglades National Park, Dade Co. Ca-
nals, West Palm Beach; Palm Beach Co. 10 mi. SW Fellsmere; St. Lucie Co,
(ANSP). ST. JOHNS RIVER SYSTEM:-ST. JOHNS RIVER DRAINAGE: Ditches be-
tween Deer Park and Melbourne; ditch near Lake Washington, 6 mi. W Eau
Callie; Lake Poinsett, 10 mi. WNW Cocoa; all Brevard Co. Lake Harney, 3 mi.
NE Geneva; St. Johns River, 1 mi. ESE Osceola; Econlockhatchee River, near
confluence with St. Johns River, Lake Jessup, 3 mi. N Oviedo; St. Johns
River [town of] Lake Monroe, all Seminole Co, Lake Ashby, 8 mi. NE Osteen;
St. Johns River, Lemon Bluff, 3 mi. SE Osteen; Lake Woodruff, all Volusia
Co. Lake Killarney, Winter Park; canal from Lake Virginia to Lake Sue,
Winter Park; both Orange Co. ~KLAWAHA RIVER DRAINAGE: Lake Louisa, 6



1972 JOHNSON: FLORIDA UNIONIDAE 225

mi. SSE Clermont; Lake Minnehaha, 2 mi. S Clermont; both Lake Co. Indian
Lake, 6 mi. N Silver Springs; Halfmoon Lake, 6 mi. SE Lynn; Lake Lou, 5 mi.
NE Lynn; all Marion Co. Shands Canal, 3 mi. NE Micanopy; Alachua Co.
Orange Creek, 1 mi. N Orange Springs, Marion Co. Little Orange Creek, 3
mi. E Hawthorne, Putnam Co. Hatchet Creek, Hawthorne Road, NE Gaines-
ville; P. rairie Creek, at outlet of Newnans Lake, 4 mi. SE Gainesville; Bivens
Arm, 2 mi. SW Gainesville; all Alachua Co. HAW CREEK DRAINAGE: Little Haw
Creek, 2 mi. SW Deanville; Flagler Co. JULINGTON CREEK DRAINAGE. Durbin
Creek, 1 mi. WSW Bayard; lake on Julington Creek, 2 mi. W Bayard; both
Duval Co.

Subfamily ANODONTINAE (Rafinesque, 1820),.Ortman, 1910.
Genus Anodonta Lamarck

Subgenus Anodonta Lamarck

Anodonta Lamarck 1799, Memoires·de la Soc. d'Hist. Nat. de Paris, p. 87.
Type species: M!/titus c!/gneus Linnaeus. M onotypic.

Placed on the Official List of Generic Names in Zoology (1926) Opinion
94. Reconfirmed (1959) Opinions and Declarations rendered by Int.
Comm. Zool. Nomen., 20 (28) 303-310, Opinion 561.

Subgenus Utterbackia F.C. Baker
Utterbackia F.C. Baker 1927, American Midland Nat., 10: 221, 222. (mis-

spelled as Utterbachia on p. 221)
Type species: Anodonta imbecillis [sic] Say. Original designation. Utterback-

iana Frierson 1927, Check List North American Naiades, p. 17. Type
species, Anodonta suborbiculata Say. Monotypic.
Under the subgenus Lastena Rafinesque (not available for use here

as the type is Anodonta lam Rafinesque , teste Ortmann and Walker ,
(1922: 32), Frierson included all of the tan mentioned in the present
paper except A. suborbiculata Say of the Interior Basin. For this he intro-
duced the subgeneric name Utterbackiana, on the basis that that species
is dioecious, but as mentioned below, this is an unreliable basis for classi-
fication and the shell morphology is clearly, that of Utterbackia.

Morrison (in Walter, 1956: 265) stated that Anodonta imbecilis,
like A. cygnea of Europe and Asia, is monoecious and has flat umbos and
therefore belongs to Anodonta.

Heard (1966: 31) showed clearly that sexuality is an unreliable means
of classifying Anodonta. Neither A. Ct/gnea nor imbecilis are uniformily
hermaphroditic, though each species contains some monoecious individ-
uals. The flat umbos of Utterbackia and Anodonta s.6. appear to be a
convergent character. Utterbackia is quite isolated from Anodonta. In
North America the latter is restricted to the Pacific region. All four
species of Utterbackia are more delicate than cygnea, and individuals of
each of the species may eihibit fine rays toward the umbos, which are
lacking in c!/gnea or any other Anodonta.



226 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

In the Interior Basin are two species of Utterbackia, suborbiculata
and imbecilis, the latter is also found in the Apalachicolan and South-
ern Atlantic Slope regions, but not in peninsular Florida. Speciation has
taken place in the Southeastern states, where two additional species
occur, couperiana and pegg!/ae.

Anodonta (Utterbackia) couperiana Lea
Figures 38, 10 A-B

Anodonta cowperiana [sic] Lea 1840, Proc. Arner. Philos. Soc., 1: 289 (Hope-
ton, near Darien [Mcintosh Co.], Georgia). Changed to:

Anodon ta couperiana Lea 1842, Trans. Amen Philos. Soc., 8. 227, pl. 20, fig
46, figured type, not in USNM [lost]. Lectotype, USNM 86673, selected
by Johnson, 1965, Breviora, Mus, Comp, Zool, no, 213, p. 3, pl. 2, fig. 4.
Lea, 1842, Obs. Unio, 3:65.

Anodonta dunlapiana Lea 1842, Proc. Amen Philos. Soc., 2: 225 (South Caro-
ling). Lea, 1842, Trans. Amen Philos. Soc., 8: 248, pl. 27. fig. 65; figured
type, not in USNM [lost] Lectotype, here selected, USNM 86564, fig lOA,
Charleston, Charleston Co,, South Carolina. Lea, 1842, Obs. Unio,
3:86.

Anodonta cowperiana [sic] Lea. Clench and Turner, 1956, Bull. Florida State
Mus., 1: p. 183, pl. 6, fig 3.
DESCRIPTION.-Shell medimii to large, reaching 93 min in length.

Outline subelliptical to subcircular. Valves somewhat inflated, thin, fra-
gile, and . smooth. Anterior end regularly rounded; posterior end some-
what pointed. Ventral margin broadly curved. Dorsal margih straight
and long, usually forming a distinct wing-like angle where it meets the
obliquely descending posterior margin. Hinge ligament short but promi-
nent. The posterior margin joins the curved. ventral margin at a point
near the medial line. Posterior ridge broadly rounded. Posterior slope
slightly concave. Umbos low and broad, seldom extending above the
dorsal margin, 16cated in the anterior third of the shell, their sculpture
consisting of a number of delicate subconcentric undulations. Periostra-
cum smooth and shiny, except the posterior slope which may be rough-
ened. Surface of the shell straw yellow to yellowish-green usually with
numerous, generally fine green rays, sometimes with distinctly darker
rays on the posterior slope.

No hinge plate or teeth; muscle scars inconspicuous and poorly de-
fined. Nacre bluish-white and it·idescent.

MEASUREMENTS.-L 93 min, H 52 mm„ W 38 min (St. Johns River
[town of] Lake Monroe, Seminole Co.h L 75 mm, H 44 mm, W 31 mm
(same as above); L 64 mm, H 40 mm. W 28 mm (Lake Ashby, 8 mi. NE
of Osteen, Volusia Co.).

HABITAT.  Prefers sandy or muddy bottoms of ponds and slow-mov-
ing streams.

REMARKS. -In peninsular Florida Anodonta couperiana Lea can only
only be confused with A. peggyae Johnson . Both species have umbos that



1972 JOHNSON: FLORIDA UNIONIDAE 227

do not extend above the dorsal margin, but couperiana is elliptical in out-
line and pointed posteriorly, the point ending neal· the medial line, the
dorsal and ventral margins roughly parallel. A. pegg!~ae is subrhomboidal
in 6utline, with a biangulation that ehds near the base. When the dorsal
margin is held straight; the ventral margin often descends obliquely to-
ward the broad posterior basal biangulation.

C
 4&

FIGURE 10'. Anodonta (Utterbackia) couperiana Lea. A. Lectotype of Anodonta dunla-
piana Lea. Charleston, Charleston Co., South Carolina, USNM 86564, L 83, H 46,
W 35. B. Lectotype of Anodo,itd couperiana Lea. Hopeton, near Darien, [ Meln-
tosh Co.], Georgia. USNM 86673. L 67, H 38, W 29.

Anodonta ( Utterbackia) peggyae Johnson. C . Holotype of Anodonta peg -
gyae Johnson . Lake Talquin , Leon Co ., Florida. MCZ 251040 . L 71 , H 43 , W 24 .
D. Paratype of Anodonta pegg!/de Johnson. Same as holotype. MCZ 251041. L 54,
H 33, W 19.
1 Slightly reduced, measurements in mm, L = length, H = height. W = width.



228 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4-

In the Apalachicolan and Southern Atlantic Slope regions, A. coup-
eriana can be confused with A. imbecilis Say, but couperiana differs by
having broad green rays, which are especially fine on the disk, and by
its broadly curved ventral margin, which renders the shell proportion-
ately much higher. In imbecilis the ventral margin is almost straight and
parallel to the dorsal one . The height/length ratio of couperiana is about
2 to that of 1.5 in imbecilia.

A. couperiana is common in central Florida where individuals
tend to be small and often locally abundant. In peninsular Florida
large specimens are found in the St. Johns River system. The largest
specimens occur in the Southern Atlantic Slope region, where the
species is scarce.

The apparent absence of this species from the Suwannee and
Withlacoochee river systems suggests that it entered the Apalachi-
cola River system from a former conRuence with the Savannah River
system, and that it spread into the Florida peninsula from the north.

RANGE.-Apalachicolan region: Apalachicola, Ochlockonee and St.
Marys river systems. Peninsular Florida. Southern Atlantic Slope: Al-
tamaha River system, Georgia, north to the Cape Fear River system,
North Carolina.

SPECIMENS EXAMINED.- MYAKKA RIVER SYSTEM: Upper Lake Myakka; Sar-
.150ta (0. PEACE RIVER SYSTEM: Peace River, Areadia; De Soto Co. KISSIMMEIS
RIVER ~YSTEM AND EVERGLADES:-KISSIMMEE RIVER DRAINAGE: East Toho-
pekaliga Lake, Narcoossee; Tohopekaliga Lake, Kissimmee; Lake Hatchieha,
Lake Marion, 3 mi. WNW Kenansville; all Osceola Co. Lake Rosalie, 4 mi. NE
Hesperides; Lake Howard, Winter Haven; Weohyakapka Lake, Indian Lake
Estates; Tiger Lake, 7 mi. ESE Hesperides; all Polk Co. Lake Bonnet, 3 mi.
N Sebring; Lake Istokpoga; both Highlands Co. LAKE OKEECHOBEE DRAINAGE:
Fisheating Creek, 1 mi. S Palmdale; Glades Co. Canal, 2 mi. NW Lantana;
Lake Osborne, 2 mi. SW Lantana; both Palm Beach Co. ST. JOHNS RIVER
SYSTEM:-ST. JOHNS RIVER DRAINACE: Lake Winder; Brevard Co, Lake Harney
(ANSP), Econlochatchee River, near confluence with St. Johns River; Lake
Jessup, 3 mi. N Oviedo; St. Johns River [town of] Lake Monroe; all Seminole
Co. Lake Ashby, 8 mi. NE Osteen; Lake Woodruff; Spring Garden Lake, 1 mi.
NW De Leon Springs; ali Volusia Co. St. Johns River, Astor; Blue Creek
(ANSP); both Lake Co. Lake George, S end Drayton Island; Putnam Co.
OKLAWAHA RIVER DRAINAGE: John Lake, 1 mi. S Oakland; Orange Co. Eake
Beauclaire, 2 mi. S Mt. Dora (ANSP). Lake Griffin,.Leesburg, both Lake Co,
Lake Eaton, 6 mi. NE Lynn; Marion Co. Cross Creek, 5 mi. NW Island Grove;
Lochloosa Lake, 1 mi. N. Lochloosa, Shands Canal, 3 mi. NE Micanopy;
Prairie Creek, 6 mi. SE Gainesville; Newnans Lake, 6 mi. E Gainesville; River
Styx (USNM), all Alachua Co.

Anodonta (Utterbackia) pegg!jae johnson
Figure 38, 10 C-D

Anodonta imbecitis Say. partim  Clench and Turner, 1956, Bull. Florida State
Mus., 1 : 187, pl . 6, fig . 2 , paratype of Anodonta peggyae, MCZ 251041 .



1972 JOHNSON: FLORIDA UNIONIDAE 229

Anodon ta pegg!/ae Johnson 1965, Breviora, Mus. Comp. Zool., no. 213, pl. 2,
figs. 1-3 (Lake Talquin [formed by a dam on the Ochlockonee River],
Leon County public fishing ground, Leon Co,, Florida, holotype MCZ
251040)
DESCRIPTION, -Shell small to medium, reaching a little over 80 inm in

in length. Outline subrhomboidal, valves slightly inRated, thin,  fragile,
and smooth. Anterior end regularly rounded; posterior end more broadly
rounded and slightly biangulate just above the base. Ventral margin
broadly curved and obliquely descending. Dorsal margin straight or
slightly curved, usually forming a distinct wing-like angle where it meets
the obliquely descending posterior margin. Posterior ridge broadly
rounded, posterior slope sometimes slightly concave. Umbos low and
broad, not extending above the dorsal margin, located in the anterior
third of the shell, their sculpture consisting of seven or eight low, deli-
cate, slightly double-looped undulations. Periostracum smooth and
shiny, except the posterior slope which may be slightly roughened. Sur-
face of the shell straw yellow to yellowish-green, sometimes very dark
green, with numerous, generally fine, green rays over the entire surface.
The rays are distinctly darker on the posterior slope.

No hinge plate or teeth; muscle scars inconspicuous and poorly de-
fined. Nacre bluish-white and iridescent.

MEASUREMENTS. -L 71 111111, H 43111117, W 24 111111.(Lake Tal(litin, Leon
Co:, holotype); L 66 nim, H 35 mm, W 20 mm (Lees Lake, Panasoffkee,
Sumter Co.), L 63 mm, H 32 mm, W 20 mm (same as above).

HABITAT. - Prefers sandy or muddy bottoms of ponds mid slow-mov-
ing strea'ms.

REMARKS.-- In peninsular Florida Anodc)nta peggilae Johnson can be
be confused only with A. couperiana Lea. A. peggyae is subrhomboidal
in outljne with a posterior 6iangulation that ends near the base. When
the dorsal margin is held straight, the ventral margin is often obliquely
descending toward the broad posterior basal biangulation. A. couperiana
is elliptical in outline and pointed posteriorly, the point ending near
the medial line. The dorsal and ventral margins are approximately par-
allel . In the Apalachicolan region , peggyae can also be confused with
A. imbecilis Say, but imbecilis has an elongate elliptical shell, a poster-
ior point ending near the medial line, an almost straight ventral margin
parallel to the dorsal one, and a rather uniformly green periostracum. A.
Pegg!/ae differs from imbecilis in that, with the hinge line held horizontal,
it has a subrhomboidal shell with a less acute point located near the base,
a ventral margin that slopes obliquely from the dorsal one, and a perio-
stracum with numerous green rays that are especially fine on the disk.

RANCE. - Apalachicolan region: Choctawliatelieu River system, east
to the Suwannee River system. Peninsular Florida: Withlacoochee and
Hillsborough riv.er systems.

SPECIMENS EXAMINED.-WITHLACOOCHEE RIVER SYSTEM: Withlacoochee

River, 1 mi. NW Lacoochee, Pasco Co. Little Withlacoochee River, Rerdell,
Hernando Co. Lees Lake, Panasoffkee; lake, 6 mi. NNW Panasoflkee, both



230 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Sumter Co. Lake Tsala Apopka, Floral City, Citrus Co. HILLSBOROUGH RIVER
SYSTEM: Blackwater Creek, 8 mi. N Plant City; Hillsborough River,. 4 mi. NE
Temple Terrace; both Hillsborough Co.

Subfamily LAMPSILINAE (Ihering 1901) Ortman, 191(k
Genus Caruncu/ina Baker

Toxolasma Rafinesque 1831 , Continuation of Monog, Bivalve Shells River
Ohio (Phila.), p. 2 Species listed: Udio cyclips, U. cinerescens, U lividus,
U..#exus, all Rafinesque.

Type species, Unto liuidus Rafinesque. By elimination, Frierson, 1914, Nauti-
lus, 28: 7. Ortmann and Walker, 1922, Occ. Papers, Mus. Zool., Univ.
Mich. no. 112, pp. 54, 55, showed that U. liuidus is a nomia dubia and that
therefore Toxola,ma must be disregarded. Nevertheless, on the mere
statement of Morrison (1969:24) that: " Toxolasmc liuida Ra£ 1831 ( = g/ans
Lea Dec. 1831)", Valentine and Stansbery (1971.: 29) have resurrected this
maine, claiming that it has priority over Cartinculina.

Coruncutina 'Simpson', 1898, [in] F, C. Baker, 1 Bull: Chicago Add. Sci,, 3 (1),
P.109.

Type species, Unio parous Barnes. Monotypic
Carunculina, corrected in the index and on errata sheet, and reconfirmed by

Simpson, 1900, Proc. U. S. Natl. Mus., 22: 563. Ortmann, 1912, Ann. Car-
negie Mus., 8: 377. partim. (under Eurynia)
Call (1896) monographed Caruncutina, and indicated that it in-

cluded only a few very variable species. He probably correctly reduced
to synonymy many of the taxa subsequently recognized as valid species
by Simpson (1914, 1: 148-161) and Frierson (1927: 87-89).

Carunculina parua (Barnes)
Figures 2D, 11 A-B

Unio paruus Barnes 1823, American Jour. Sci. (1) 6: 174, pl. 13, fig. 8 (Fox
River [Wisconsin]; [type presumed lost]).

Unio paulus Lea, 1840, Proc. Amen Philos. Soc., 1: 287 (Chattahoochee River,
Columbus [Muscogee Co.] Georgia). Lea, 1842, Trans. Amer. Philos.
Soc., 8: 213, pl. 15, fig. 29; figured holotype USNM 85274. Lea, 1842, Obs.
Unio, 3: 51.

Unio minor.Lea, 1843, Desc. Twelve Uniones (Lake Monroe and Lake George,
Florida). Lea, 1846, Trans. Amer. Philos. Soc., 9: 276, pl. 39, fig. 3; figured
holotype USNM 85310 from Lake George, Lea, 1848, Obs. Unio, 4: 34,

Unio marginis Lea, 1865, Proc. Acad. Nat. Sci. Phila., 16: 89 (Blue Springs
[Albany] Dougherty Co., Georgia). Lea, 1869 Jour. Acad. Nat. Sci. Phila.,
(2) 6: 225, pl. 31, fig. 69; figured holotype USNM 85295. 1869, Lea, Obs.
Unio, 12: 15.

1 H, B, Baker (1964, Nautilus, 78:33) pointed out that as Simpson chntributed nothing
in the original publication ef this genus (under article 51 (c), 1964 edition of the Inter-
national Code·Zool. Nomen.) tNe'authority for Carunculina must be F. C. Baker.



1972 JOHNSON: FLORIDA UNIONIDAE 231

Unio cromwellii Lea, 1865, Proc. Acad. Nat. Sci. Phila., 16: 89 (Kiokee Creek
near [ W] Albany, Dougherty Co., Georgia) Lea, 1869, Jour, Acad. Nat.
Sci. Phila., (2) 6: 258, pl. 31, fig. 73; figured holotype USNM 85280. Lea,
1869, Obs. Unio, 12: 18.

Unio coroinus Lea, 1868, Proc. Acad. Nat. Sci. Phila., 20: 144 (Flint River,
Georgia; Darien? [ Mcintosh Co., Georgia]; Neuse River, Raleigh [Wake
Co.] North Carolina). Lea 1869, Journ. Acad. Nat. Sci. Phila., (2) 6: 310,
pl. 48, fig. 123; figured holotype USNM 85277 from Flint River, Georgia.
Lea, 1869, Obs. Unio, 12: 70.

Unio stearnsi B. H. Wright, 1888, Check List North American Unionidae
(Portland, Oregon) p. 5, nomen nudum;_ Listed as a synonym of Lamp-
silis minor Lea, by Simpson, 1900. Proc. U:S. Natl. Mus. 22: 562.

Unio si,Lgle!~anus Marsh: 1891, Nautilus, 5: 29 (Palatka, Florida, type figured
by Simpson, 1892, Proc. U.S. Natl. Mus., 15: 426, pl. 68, figs, 4,5, USNM
[not located].

Carunculina parua cdhni · F. C. Baker, 1927, American Midland Nat. 10: 222
(Neosha Mill Pond, Dodge Co., Wisconsin; holotype U. of Illinois Z17341,
figured by Baker, Bull. Univ. Wisconsin, no. 1527, p. 253, pl. 105, Ag, 14),

Corunculina [sic] paula (Lea). Cleneh and Turner, 1956 Bull. Florida State
Mus., 1: 193, pI. 8, fig. 5.

Carunculina parua (Barnes). La Rocque, 1967, Geol. Survey, Ohio, Bull., 62,
(2):263, fig. 153.

Toxolasma parua (Barnes). Valentine and Stansbery, 1971, Sterkiana, no.. 42,
P 29.
DESCRIPTION. - Shell small, seldom reaching over 35 mm in length.

Outline of female obovate; male elliptical. Valves subinflated, generally
thin. Anterior end regularly rounded. Posterior end of females more
broadly rounded and subtruncated below the medial line; males some-
what pointed. Ventral margin straight or slightly curved in males. In fe-
males marsupial swelling causes the margin to be somewhat convex a
little posterior of the center. Dorsal margin slightly curved, usually form-
ing a distinct angle with the obliquely descending posterior margin. Pos-
terior ridge faintly double or, more often, quite in distinct. Umbos prom-
inent, hardly elevated above the hinge line, located in the anterior third
of the shell, their sculpture consisting of several ridges parallel to the
growth lines. Periastracum generally with distinct growth lines, often
satiny, generally blackish, occasionally yellowish or olive, and with very
fine obscure green rays.

Left valve with two raised, triangular, occasionally crenulate
pseudocardinal teeth, one in front of the other. Hinge line short, gener-
ally very narrow, in front of two Short straight lateral teeth.  Right valve
with one rather chunky, triangular pseudocardinal; one lateral tooth.
Beak cavities shallow, with a few dorsal muscle scars under the hinge
plate. Anterior adductor muscle scars well impressed, posterior ones
faint. Pallial line distinct anteriorly. Nacre bluish-white or pink to pur-
plish iridescent.

MEASUREMENTS. - L 39 mm, H .25 mm, W 16 mm (Lake Min-



232 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

nehaha, 2 mi S of Clermont, Lake Co., niale); L 34 mm, H 22 mm, W 15
mm (same as above, female); L 37 mm, H 21 mm, W 17.5 mm (Lake Kil-
larney, Winter Park, Orange Co., male); L 27 mm, H ,17.5 mm, W 14 mm
(same as above, female).

HABITAT, - Lives ill Shallow water near the edges of streams and
ponds, gener:illy in mud, sometimes in sand.

REMARKS. -In peninsulai- Florida some specimens of Cartinculina
parua (Barnes ) can occasionally be confused with Villosa . Both genera
exhibit similar sexual dimorphism, but the species of Villosa are quite dif-
ferent from parua. All three species of Villosa (uillosa, Dibex, and am!/g-
dala) attain much larger size, have thinner, generally yellowish, distinctly
rayed shells, whereas parva is heavier for its size, blackish, and only
rarely obscurely rayed. The hinge teeth in parua tend to be chunky while
those of Villosa tend to be more delicate.

Carunculina parua differs from C. pulla (Conrad) of the Atlantic
Slope region by having an indistinct posterior ridge and a satiny perio-
stracum; C. pulla has a sharp posterior ridge with a second less promi-
nent ridge above it. The periostracum of pulla is generally much
rougher with heavy growth lines.

Call ( 1896 ) first pointed out that except for C . glans (Lea), which
is restricted to the Interior Basin, there appears to be but one other
species of Carunculina iii the Interior Basin, Apalachicolan and
per,insular Florida regions. He was not followed by Simpson (1914:
148-161) or by Clench and Turner (1956: 193). The latter used the name
paula for, this species as this nominal form has its type locality within
the region covered by their study.

RANGE.-Interior Basin generally, from western New York west
Minnesota, south to Arkansas. Apalachicolan region. Peninsular Florida.

FIGURE 111. Cartinculina parua (Barnes). A. Holotype of Unio minor Lea. Lake
George, Florida. USNM 85310. L. 22, H 15, W 10, female (2.5 X). B. Carunculina
parua ( Barnes ). Durbin Creek , 1 mi . WSW Bayard, Dtival Co ., Florida . MCZ
269273. L 35, H 20, W 15, male (nat. size).

Villosa oillosa (B. H. Wright). C. Lectotype of Unio villows B. H. Wright.
Suwannee River [ Luraville], Suwannee Co., Florida. USNM 150503. L 57, H 28,
W 18, female (nat. size). D. Holotype of Lampsilis wrightiana Frierson. Volusia
Co., Florida. UMMZ 91179. L 50, H 27, W 18, male (slightly reduced, reversed
image).

Villosa vibex (Conrad). E. Lectotype of Unio averellii B. H. Wright. Lake
Ashby, Volusia Co., Florida. USNM 91142. L 48, H 30, W 18, female (nat. size).
F. Paralectotype of Unio auerellii B. H. Wright. Same as lectotype. MCZ 252169.
L 42, H 25, W 15, male (nat. size).

Villosa am!/gdalum (Lea). G. Holotype of Unio trossulus Lea: Lake Monroe,
Florida. USNM 84705, L 35, H 21, W 16, male (nat. size). H. Holotype of Unio
am!/gdalum Lea. Lake George, Florida. USNM 86127. L 32, H 22, W 15, female
( nat . size). I . Lectotype of Unio papyraceus Gould . Everglades of Florida. USNM
86125. L. 41, H 25, W 15.2 (nat. size) J  Holotype of Unto vesicularis Lea. Lake
Okeechobee, Florida. USNM 85292, L 32, H 19, W 13 (nat. s ize).
' Measurements in mm, L  = length, H = height, W = width.



1972 JOHNSON: FLORIDA UNIONIDAE 233

A & 3(~»5Sm, 9///1/////A 8 -

C D

- - ~:4
-

-

. 5/~

G

1 -----
Fillay.~



234 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

SPECIMENS EXAMINED.-WITHLACOOCHEE RIVER SYSTEM:-Little Withla-
coochee River, Rerdell, Hernando Co. Lees Lake, Panasoffkee; Sumter Co.
HILLSBOROUGH RivER, SYSTEM: - Hillsborough River, 4 mi. NE Temple Ter-
race, Hillsborough CO. PEACE Ri VER SYSTEM: - Peace River, Areadia; De
Soto Co. KISSIMMEE RIVER ~YSTEM AND EVERCLADES:-KISSIMMEE RIVER,
DRA[NAGE: Tohopekaliga Lake, Kissimmee; Alligator Lake, near Kissimmee;
both Osceola Co, Lake Rosalie, 4 mi, NE Hesperides; Tiger Lake, 7 mi, ESE
Hesperides; both Polk Co, LAKE OKEECHOBEE DRAINAGE: Fisheating Creek, 1
mi. S Palmdale; Glades Co, ST, JOHNS RIVER SYSTEM:-ST. JOHNS RIVER
DRAiNAGE: Econlokhatchee River, near confluence with St. Johns River;
Puzzle Lake, 7 mi. SE Geneva; Lake Jessup, 5 mi. N Oviedo; Wekiva River,
7 mi. WSW [town of] Lake Monroe; all Seminole Co. Lake Ashby, 8 mi. NE
Osteen; Lake Monroe, near Enterprise; both Volusia Co, Lake Killarney,
Winter Park; Orange Co. OKLAWAHA RIVER DRAINAGE: Lake Minneola, Cler-
mont; Lake Minnehaha, 2 mi. S Clermont; b.oth Lake Co. Lake Lou, 5 mi. NE
Lynn; Lake Eaton, 6 mi. NE Lynn; Orange Creek, 1 mi. N Orange Springs;
all Mafion Co. BLACK CREEK DRAINACE: North Fork, Black Creek, 14 mi. SW
Orange Park; South Fork, Black Creek, 1 mi, SW Middleburg; both Clay Co.
JULING-roN CREEK DRAINAGE: Durbin Creek, 1 mi. WSW Bayard; Duval Co.

-Genus Villosa Frierson
Micromya Agassiz 1852, Archiv fur Naturgeschichte, 18 ( 1 ), p . 47 . Species listed :

Unio lapillus Say, Margart'tana fabula Lea, M, curreb,ana Lea, non Mi-
croyin<L Rondani 1840 ( Insecta ),

Type species: Unio lapillus Say. Subsequent designation, Herrmannsen, 1852,
Indicis Generum Malacozoorum, Supp, et Corn, p. 83, Ortmann, 1912,
Ann. Carnegie Mus., 8: 337. partim.

Villosa Frierson, 1927, Check List North American Naiades, pp. 11, 80.
Type species, Unio uillosits Wright. Original designation.

At this writing, it is impossible to tell how many species Villo.sci con-
tains. Frierson (1927: 70-79) included under Lanzpsilis, subgenus Li-
gumia, many taxa that other authors have included under Micromya
C = Villosa). Except for the type species, the taxa Frierson (1927: 80, 81)
listed under Villosa are species of Carunculina, Despite the systematic con-
fusion within this genus, it is clear that the majority of its species occur in
the Interior Basin.

Villosy, villosa (B. H. Wright)
Figures 3D, 11 C-D

Unio villosus B. H. Wright 1898, Nautilus, 12: 32 :Suwannee River [Luraville],
Suwannee Co., Florida; syntype USNM 150503, figured by Simpson, 1900,
Proc. Aead. Nat. Sci. Phila., p. 77, pl, 1 fig..1, selected as lectotype by John-
son, 1967, Oce. Papers on Moll.,3. 9, pl. 8 fig. 1).

Lampsilis villosus (Wright). Simpson, 1914, Cat. Naiades, 1: 143.
Lampsilis wrightiana Frierson 1927, Check List North American Naiades, p . 81

(Volusia Co., Florida; holotype UMMZ 91179, figured by Frierson, 1928,
Nautilus  41: 139, pl. 2, fg 3).



1972 JOHNSON: FLORIDA UNIONIDAE 235

Villosa villosa ( Wright), Clench and Turner , 1956, Bull . Florida State Mus ., 1 :213, pl.  4, fig 2.
DESCRIPTION.-Shell usuall>, small, seldom exceeding 60 111111 in

length. Outline elongate elliptical. Valves subinflated, generally thin, and
translucent. Anterior end regzilarly rounded. Posterior end of females
slightly more broadly rounded; males quite pointed. Ventral margin almost
always broadly curved except in females where a slight marsupial swelling,
somewhat posterior of the center, renders it straight or slightly convex. Dor-
sal margin straight with a very slight, if noticeable, angle where it meets the
obliquely descending posterior margin. Hinge ligament small. Posterior
ridge broadly rounded, double in the male; obscuied by a slight marsupial
swelling in the female. Posterior slope slightly concave. Umbos moderately
swollen, slightly elevated above the hinge line, located in the anterior quar-
ter of the shell, their sculpture consisting of several fine, low, slightly double-
looped ridges. Surface of the shell with irregular growth lines, occasionally
smooth and shiny, but usually covered with either rough or distinctly satiny
periostracum, especially on the posterior slope. Periostracum sometimes ~
subshiny, greenish-yellow, dark greenish, or more often brownish-black,
the entire surface of the shell with broad green rays interspersed with narrow
ones, sometimes only visible in transmitted light.

Left valve with two delicate pseudocardinal teeth, one in front of the
other, the anterior one somewhat triangular, the hind one inclined to be ,
vestigial. Hinge line short and narrow in front of two, short, straight lateral
teeth. Right valve with two triangular, narrow, parallel pseudocardinals
separated by a narrow pit, the more anterior tooth quite vestigial, some-
times absent; one low lateral tooth. Beak cavities shallow, a few dot·sal
muscle scars under the hinge plate. Anterior adductor muscle scars well
impressed, posterior ones faint, if visible, Pallial line distinct anteriotly.
Nacie bluish-white, occasionally yellowish-white, and iridescent, es-
pecially posterior·ly

MEASUREMENTS. -L 57 nim, H 28 mm, W 18 min (Suwannee River,
Luraville, Suwannee Co., lectotype, female): L 54 mm, H 29 mm, W 21
mm (Peace River, near Arcadia, DeSoto Co., male); L 45 mm, H 25 mm,
W 19 mm (same as above, female).

HABITAT. - "Limited to spring-fed streams und clear rivers" (Clench
and Turner 1956: 214), but this report did not include the rather acidic
and muddy St. Marys River, where the species also occurs.

REMARKS . -- Villosa billosa Wright be:irs a resemblance to Villosa
uibex Conrad, which is more broadly distributed and extends over the
whole range of V. uillosa. Both show sexual dimorphism. In the female of
uillosa the post basal swelling generally shows less tendency to extend be-
low the ventral margin, and if somewhat posteriorly pointed, the point is
higher. The male is pointed posteriorly, as in uibex, but the shell is prop01·-
tionately longer. V. villow often has a distinctive roughened pei·iostracum
that produces a satiny Iris ten



236 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

RANGE. -Apalachicolan region: Apalachicola River system, east to
the St. Marys River system, Georgia. Peninsular Florida.

~PECIMENS EXAULNED.-WITHLACOOCHEE RIXER $YSTEM:-Lake Tsala

Apopka, Hernando Co. Withlacoochee River, Dunnellon, Marion Co. HILLS-
BOROUGH RIVER SYSTEM: Hillsborough River, 10 mi. N Tampa, Hillsborough

Co. MYAKKA RIVER SYSTEM: Myakka River, just E Myakka City, Manatee Co.
Upper Myakka LAke, Sarasota Co. PEACE RIVER SYSTEM: Horse Creek, 8 mi.

W Arcadia; ponds along Peace River, 1.25 mi. below Arcadia; both De Soto Co.
ST. JOHNS RIVER SYSTEM: OKLAWAHA RIVER DRA[NAGE: Lake Elistis, Ta-
vares, Lake Co. Cross Creek between Lake Lochloosa and Orange Lake, 19 mi.
S Gainesville, Alachlia Co. JULINGTON CREEK DRAINAGE: Durbin Creek, 1 mi.
WSW Bayard, lake on Julington Creek, 2 mi. W Bayard, both Dtival Co.

Villosa vibex (Conrad)
Figures 2B, 11 E-F

Unio Dibex Conrad, [May] 1834, New Fresh Water Shells United States,
p. 31, p1 4, fig. 3 (Black Warrior River, South of Blount's Spring [Blount
Co.], Alabama; figured holotype ANSP 56488a). Published in May,
teste Conrad, 1853, Proc. Acad. Nat. Sci. Phila., 6: 243.

Unio modioliformis Lea, [August or September] 1834, Trans. Amer. Philos.
Soc., 5: 97, pl. 13, fig. 40 (Santee Canal, South Carolina; probable figured
holotype USNM 85029 [differs slightly from figure] ) Lea, 1834, Obs.
Unio, 1: 209. Published in August or September, teste Lea, 1854, Proc.
Acad. Nat. Sci. Phila,,7: 244,

Unio exiguus Lea 1840, Proc. Amen Philos. Soc., 1: 287 (Chattahoochee River,
near Columbus [ Muscogee Co.], Georgia). Lea, 1842, Trans. Amer.
Philos. Soc., 8: 191, pl. 7, fig. 1; figured holotype USNM 84974. Lea, 1842,
Obs. Unio, 3: 29.

Unio stagnalis Conrad 1849, Proc. Acad. Nat. Sci. Phila,, 4: 152 (inhabits mill
ponds, Ogeechee River, Georgia, J H. Couper [loaned] ). Conrad,
1850, Jour. Acad. Nat. Sci. Phila., (2) 1: 275, pl. 37, fig. 2; figured holotype
MCZ 178778, purchased from J· H. Couper.

Unio preuostianus Lea 1852, Trans. Amer. Philos. Soc., 10: 269,· pl. 19, fig. 24
(Eutowah [Etowah] River [North West], Georgia; figured holo-
type, C. M. Wheatley Colin. in ANSP [lost] ) Lea, 1852, Obs. Unio,
5: 25.

Unio nigrimis Lea 1852, Trans. Amer. Philos. Soc., 10: 284, pl, 24, fig. 44 (West
Florida; figured holotype USNM 86132). Le.1, 1852, Obs. Unio, 5:  40.

Unio gracilior Lea 1856, Proc. Acad. Nat. Sci. Phila., 8: 262 (Buckhead Creek
[Burke Co.], Tobesaulke [Tobasofkee] Creek, near Macon [Bibb Co.] ;
both Georgia), Lea, 1858, Jour.  Acad. Nat. Sci. Phila., (2) 4: 56, pl. 8, fig.
38; figured holotype USNM 85088 [localities not separated] . Lea, 1858,
Obs. Unio, 6: 56.

Unio mtilans Lea 1856, Proc. Acad. Nat, Sci. Phila., 8: 262 (Othealooga [Ooth-
kalooga] Creek, Gordon Co.; Columbus [Muscogee Co.]; both Geor-
gia). Lea, 1858, Jour. Acad. Nat. Sci. Phila., (2) 4: 59, pl. 9, fig. 41; figured
holotype USN.M 85093 from [Oothkalooga] Creek. Lea, 1858, Obs.
Unio, 6: 59.



1972 JOHNSON: FLORIDA UNIONIDAE 237

Unio subellipsis Lea 1856, Proc, Acad. Nat. Sci. Phila., 8: 262 (creeks near
Columbus [Muscogee Co.], Georgia). Lea, 1858, Jour. Acad. Nat. Sci.
Phila., (2) 4: 62, pl: 10, fig. 44; figured holotype USNM 85095. Lea, 1858,
Obs. Unio, 6: 62.

Unio sudus Lea 1857, Proc. Acad, Nat. Sci. Phila., .9: 17.0 (Dry Creek, near
Columbus [Muscogee Co.]; Macon [ Bibb Co.]; both Georgia). Lea,
1859, Jour. Acad. Nat. Sci, Phila., (2).4: 194, pl. 21, fig. 77, figured holo-
type USNM 85155 from Dry Creek. Lea, 1859, 6bs. Unio, 7: 12.

Unio obfuscus Lea 1857, Proc. Acad, Nat. Sci. Phila,, 9: 172 (Flint River, near
Macon [Co.], Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila., (2)
4: 197, pl. 22, fig. 80; figured holotype USNM 85089. Lea, 1859, Obs. Unio,
7: 15.

Unio dispar Lea 1860, Proc. Acad. Nat. Sci. Phila., 12: 305 (Columbus [Mus-
cogee Co.], Georgia). Lea, 1860, Jour. Acad. Nat. Sci. Phila., (2) 4: 327,
pl. 51, fig, 153; figured holotype USNM 85101. Lea, 1860, Obs. Unio, 8: 9.

Unio auerillii B. H. Wright, Unio: 1888, Proc. Acad, Nat, Sci. Phila,, 40: 115,
pl. 3, fig. 4 (Lake Ashby, Volusia Co., Florida, syntype figured by Simp-
son, 1892, Proc. U.S. Natl. Mus., 15: 414, pl. 56, fig. 6 [not located] ).
Lectotype USNM 91142, selected by Johnson, 1967, Oce. Papers on Molt,
3: 5, pl/7, fig. 4.

Villosa uibex (Conrad). Clench and Turner, 1956, Bull. Florida State Mus.,
1: 209, p]  4, fig.  4
DESCRIPTION.-Shell usually sm:ill, not exceeding 60 mm in length,

though occasionally reaching 100 mm. Outline subelliptical. Valves subin-
Rated, generally thin and translucent. Anterior end regularly rounded;
posterior end of females more broadly rounded; males somewhat pointed.
Ventral margin straight or slightly curved in males, often slightly arcuate
in females. Dorsal margin straight with a very slight, if noticeable, angle
where it meets the obliquely descending posterior margin. Hinge ligament
small. Posterior ridge broadly rounded. Posterior slope slightly concave,
occasionally with faint wrinkles and ridges. Umbos moderately swollen,
slightly elevated above the hinge line, located in the anterior quarter of
the shell, their sculpture consisting of several fine, low, slightly double-
looped ridges. Surface of the shell generally rather smooth, but r6ughened
by periostracum posteriorly. Periostracum usually subshiny, greenish-
yellow or yellowish-brown to almost black, the entire surface with numer-
ous, broad, greenish rays, which in darker specimens can be seen in trans-
mitted light.

Left valve with two delicate pseudocardinal teeth, one in front of
the other, the anterior one somewhat triangular, the hind one inclined
to be vestigial. Hinge line rathet long and very narrow in front of two
short, straight lateral teeth. Right valve with two triangular, narrow, par-
allel pseudocardinals sepafated by a narrow pit, the more anterior tooth
vestigial, sometimes absent; one lateral tooth. Beak cavities shallow, a few
dorsal muscle scars under the hinge plate. Anterior adductor muscle scars
well impressed, posterior ones faint, if visible. Pallial line distinct anter-



238 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

iorly  Nacre bluish-white, sometimes pinkish or purple, iride, cent pos-
teriorly.

MEASUREMENTS.-L 76 mm, H 41 mm, W 23 mm (L:ike Minnec,14
Clermont, Lake Co., female); L 73 mm, H 38 mm, W 24 mm (Oklawaha
River, Eureka Springs, Marion Co., male); L 60 mm, H 35 mm, W 22 mm
(same as above, female).

HABITAT.- Lives in small rivers, creeks, and lakes, in mud or soft
Sand, particularly where rich in vegetable detritus.

REMARKS . - In peninsul: Li- Florida Villosa uibex (Conrad ) can be ci, n -
fused with V. am!/gdala (Lea) with which it is often found living. V,  vibex
has broader, less distinct, green rays. Sexual dimorphism is not so appar-
ent iii vibex as it is in amygdala. The males of both species tend to be some-
what pointed posteriorly. Females of uibex tend to be broadly .rounded,
rendering the shell slightly arcuate, while the females of ami/gdala are
greatly inflated, with the posterior margin subangulate dorsally and trun-
cate below. In the Apalachicolan region vibex can be confused with V.
lienosa (Conrad). See remarks ·under V. am!/gdala (Lea )

RANGE.-West Gulf egmstal region, Alabama-Coosa River system and
Apalachicolan region: Pearl River system, Mississippi, east to the Suwan-
nee River system, Florida. Peninsular Florida. Southern Atlantic Slope
region: Altamaha River system, Georgia, north to the coastal ponds of the
Cape Fear River system, North Carolina.

SPECIMENS EXAMINED.-WACCASASSA RIVER SYSTEM: Waccasassa River,
Levy CO, WITHLACOOCHEE RIVER ~YSTEAI: Withlacoothee River, 9 mi. N Dade
City, Withlacoochee River, 1 mi. NW Lacoochee; both Pasco Co. Little With-
lacdochee River, Rerdell, Hernando Co. Shady Brook, 2 mi. S Coleman; Lake,
6 mi. NNW Panasoilkee; both Sumter Co. Withlacoochee River, Dunnellon,
Marion Co. HIt«LSBOROUGH RIVER SYSTEM: Hillsborough River, Morris bridge,
14 mi. NE Tampa; Hillsborough River, 4 mi. NE Temple Terrace; Fishhawk
Creek, 2 mi. S Lithia; all Hillsborough Co. PEACE RIVER SYSTEM: Peace River,
1.4 mi. below Arcadia; De Soto Co. KISSIMMEE RIVER ~YSTEM AND EVER-
GLADES:-KissIMMEE RIVER DRAINAGIE: Reedy Lake; Polk Co. Lake Verona,
Avon Park; Highlands Co. LAKE OKEECHoBEE DRAINAGE: Seven Mile Tower
Road, 3.5 mi. S Tamiami Trail, Everglades Nati. Park; W of Bridge no. '22,
Tamiami Trail; both Dade Co. Snake Creek, 9 mi. W Hallandale; Broward Co.

ST. JOHNS RIVER SYSTEM:-ST. JOHNS RIVER DRAINAGE Ditch near Lake
Washington, 6 mi. W Eau Callie; Lake Poinsett (USNM); both Brevard Co.
Puzzle Lake, 7 mi. SE Geneva; Lake Jessup, St, Johns River at S end of Lake
Monroe; both Seminole Co. Spring Garden Lake, 1 mi. NW De Leon Springs,
Volusia Co. Rice Creek, 7.3 mi. WNW Palatka; Putnam Co. Near St. Augus-
tine; St. Johns Co. (USNM). OKLAWAHA RIVER DRAINAGE. Lake Minneola,
Clermont; Lake Lucy, 2 mi. N Groveland; Lake Eustis, Tavares; Lake Saun-
ders, Tavares; Lake Griffin; all Lake Co. Lake Weir, Oklawaha; Juniper Creek,
12 mi. E Lynn; both Marion Co. Cross Creek, 5 mi. NW Island Grove, Alachua
Co. Orange Creek, 1 mi. N Orange Springs; Oklawaha River, Eureka Springs;
both Marion Co. HAw CREEK DRAINAGE: Lake Dias, 5 mi. ENE.De Leon
Springs, Volusia Co. BLACK CREEK DRAINAGE: North Fork, Black Creek, 14
mi. SW Orange Park, Clay Co.



1972 JOHNSON: FLORIDA UNIONIDAE 239

Villosa amggdola (Lea)
Figures 4A, 11 C-J, 12A

Unio amygdalum Lea, 1843, Desc. Twelve Uniones (Lake George, Florida).
Lea, 1846, Trains. Amer. Philos. Soc., 9: 275, pl. 39, fig. 1, figured holo-
type USNM 86127. Lea, 1848, Obs: Unio, 4: 33.

A

B

C

FIGURE 121. Villosa ami/gdata (Lea). A. Lectotype of Unio lepidus Gould. Lake
Monroe, Florida. MCZ 169223. L 63, H 35, W 25 (1.1 X).

Lampsilis (Lampsilis) teres (Rafinesque). B. Lampsilis (Lampsms) teres (Rafi-
nesque). Sampson Lake, 4 mi. W of Starke, Bradford Co., Florida. MCZ 237516.
L 81, H 41, W 29, female (nat, size). C. Limpsilis (Lampsilis) ·teres (Rafinesque).
Lake Tsala Apopka, Hernando Co., Florida. MCZ 237509, L 78, H 39, W 29, male
(nat. size).
1 Measurements in mm, L = length, H = height, W = width.



240 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

Unio trosculus Lea, 1843, Dese. Twelve Uniones. (Lake Monroe, Florida).
Changed to:

Unio trossulus Lea, 1846, Trans. Amer. Philos. Soc., 9: 278, pl. 40, fig. 6,
figured holotype USNM 84705. Lea, 1848, Obs. Unio, 4: 36.

Unio papraceus Gould, 1845, Proc. Boston Soe. Nat, Hist., 2: 53 (Everglades
of Florida: lectotype USNM 86125 selected by Juhnson, 1964, U.S. Natl.
Mus:,Bull. 239, p. 122, pl. 31, fig. 2).

Unio lepidus Gould 1856, Proc. Boston Soc. Nat. Hist., 6: 15 (Lake Mon-
roe, Florida, lectotype MCZ 169223 selected by Johnson, 1964, U.S Natl.
Mus., Bull. 239, p. 100, pl. 31, fig. 1)

Unio uesicularis Lea 1872, Proc. Acad- Nat. Sci. Phila., 24: 156 (Lake Ochee
chobee [Okeechobee], Florida). Lea, 1874, Jour. Acad. Nat. Sci. Phila.,
(2) 8: 37, pl. 12, fig. 34; figured holotype USNM 85292. Lea, 1874, Obs.
Unio, 13:41.

Unio singleyanus Marsh 1891, Nautilus, 5: 29 (small creek near Pilatka [Pa-
latka, Putnam Co.], Florida: [location of type unknown] figured by Simp-
son, 1892, Proc. U.S. Nati. Mus., 15: 426, pl, 68, fgs. 4,5).

Villosa uibex am!/gdala (Lea). Clench and Turner, 1956, Bull. Florida State
Mus., 1: 211, pl. 4, fig. 1.
DESCRIPTION-Shell usnally sma|1, not exceeding 50 nim iii length,

though occasionally reaching 65 mm. Outline subelliptical. Valves sub-
inflated, generally thin and translucent. Anterior eild regularly rounded,
posterior end of females more broadly rounded, males somewhat pointed.
Ventral margin straight or slightly curved, often slightly arcuate in fe-
males. Dorsal margin straight with a slight angle where it meets the
obliquely descending posterior margin. Posterior ridge broadly rounded,
posterior slope slightly concave. Umbos moderately swollen, somewhat
elevated above the hinge line, located in the anterior quarter of the shell,
their sculpture consisting of several fine, low, slightly double-looped
ridges. Surface of the shell generally rather smooth, but roughened by
periostracum posteriorly. Periostracum generally quite dull, sometimes
sub-shiny, greenish-yellow, sometimes blackish, the surface usually cov-
ered with narrow, light green rays, which in darker specimens can be
seen in transmitted light.

Left valve with two delicate pseudocardinal teeth, one in front of
the other, the anterior one somewhat triangular, the hind one apt to be.
vestigial. Hinge line very narrow, in front of two short, straight lateral
teeth. Right valve with two triangular, narrow, parallel pseudocardinals
separated by a narrow pit, the more anterior tooth vestigial, sometimes
absent; one lateral tooth. Beak cavities shallow, a few dorsal muscle
sears under the hinge plate. Anterior adductor muscle scars well im-
pressed, posterior ones faint, if visible. Pallial line distinct posteriorly.
Nacre almost always uniformly bluish-white and iridescent posteriorly.

MEASUREMENTS. - L 65 111111, H 34 min, \V 23 111,11 (lake on Julington
Creek, 2 mi. W (,f Bayard, Dilval Co., miile); L 55 iitin, H 32 min, W 21 inm
(Lake Eaton 5 mi. NE of Lynn Marion Co., male); I.44 inm H 27 mm, W
17.5111111 (sanie as above, female)



1972 JOHNSON: FLORIDA UNIONIDAE 241

HABITAT-Lives in mud cir soft sand of rivers and lakes, particularly
when rich in vegetable detritus.

REMARKS,- Iii peninsular Florida Villosci amugdma Lea can only be
confused with V. uibex (Conrad) with which it is often found living. The
rays of amvgdala are narrower , generally sharper, :ind of a lighter green
than those of uibex. The surface of am!/gdala is gerierally less shiny, and
the shell is more inflated and generally heavier. Sexual dimorphism is
more apparant iii amilid,ila . The males of both species tend to be some -
what pointed posteriorly, but females of amggdala are more inflated, with
the pc,sterior nmigrn subangulate dorsally and broadly truncate below..
while females of uibex tend to be less inflated and more broadly rounded
posteriorly, rendering the ventral margin slightly arcuate.

Villosa am!/gdala is most closely related to V. lieno.ya (Conrad) which
is found in the interior Basin and extends along the Gulf Coast from Texas
to the Suwannee River, Florida. Both species show similar sexual dimor-
phism, but lienosa has a consistently heavier shell that is less elongate and
less inRated than that of am!/gdala. Unlike the latter, lienosa is infrequently
rayed, and usually has a purplish nacre, while that of am!/gda/a is white.

V. am!/gda/a is replaced in the Atlantic Slope region from the Altama-
ha. River system, Georgia to the Neuse River system, North Carolina, by V.
delumbis (Conrad), which is similarly sexually dimorphic. V. delumbis
has a rather consistently thin shell with a light yellow or light greenish-
yellow periostracum covered by narrow green rays that are characterist-
ically broken by growth rests.

RANGE.-Peninsular Florida
SPECIMENS EXAMINED.-WITHLACOOCHEE RIVER SYSTEM: Lake Jovita (Clear

Lake), St. Leo, Pasco Co. Lees Lake, Panasofikee,-ldke, 6 mi. NNW Panasoffkee;
both Sumter Co. Lake Consuelo, near Floral City; Lake Tsala Apopka, Hernando;
Withlaeoochee River, near Crystal River; all Citrus CO. HILLSBOROUCH

RIVER SYSTEM:-Lake Tarpon, Tarpon Springs; Pinellas Co. Hillsborough
River, 4 mi. NE Temple Terrace, Hillsborough Co. PEACE RIVER SYSTEM: Lake

Hamilton, 4 mi. SSW Haines City; Polk Co, Silver Lake, Tavares, Lake Co.
Horse Creek [De Soto and Hardee Cos.]. Charlie Creek, 3 mi. SW Nocatee; Peace
River 1.25 mi. below Arcadia; both De Soto Co. KISSIMMEE RIVER SYSTEM

AND EVERGLADES:-KISSIMMEE RIVER DRAINAGE. East Tohopekaliga Lake, Nar-
coossee; Lake Tohopekaliga, Kissimmee; Lake Marion, 3 mi. WNW Keansville;
all Osceola Co, Lake Rosalie, 4 mi. NE .Hesperides, Lake Weohyakapka, Indian
Lake Estates, Tiger Lake, 7 mi. ESE Hesperides; all Polk Co. Lake Viola, Avon
Park; Bonnet Lake, 3 mi. N Sebring; both Highlands Co. LAKE OKEECHOBEE
DRAINAGE. Fisheating Creek, 1 mi. S Lakeport; Caloosahatchee River, above

lock, Moore Haven; both Glades Co. Lake Okeechobee, St. Lucie Co. Lake
Okeechobee, 2 mi. N Canal Point; canal from Lake Okeeehobee [town of] South
Bay; both Palm Beach Co. Canal, 10 mi. E Monroe Station; Collier Co, Seven
Mile fire tower, 7 mi. S Tamiami Trail, Everglades National Patk; Dade Co.
Lake Osborne, 2.5 mi. NW Lantana; Palm Beach Co. S·r. JOHNS RIVER SYSTEM:

ST. JoHNs RIvER DRAINAGE.-Ditch, near Deer Park; Osceola Co. Ditch, near
Lake Washington, 6 mi. W Eau Callie; Brevard Co. Puzzle Lake, 7 mi. SE Ge-
neva; Lake Harney, 3 mi. SE Genev:i; Econlockhatchee River, near confluence with



242 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

St. Johns River; Lake Jessup, 3 mi. N Oviedo; St. Johns River, 4 mi. E Sanford,
Lake Monroe, Sanford; ail Seminole Co. Lake Ashby, 8 mi. NE Osteen; St. Johns
River, near Enterprise; Lake Beresford; Spring Garden Lake, 1 mi. NE DeLeon
Springs; Lake Woodruff; all Volusia Co. St. Johns River, Astor; Blue Creek,
above Lake George; both Lake Co. Lake Kerr, 3 mi. SW Kerr City; Marion Co.
OKLAwAHA RIvER DRAINAGE. Black Lake, 3 mi, SW Oakland; John Lake, 1 mi, S
Oakland; both Orange Co. Lake Apopka, 2.5 mi. S Monteverde; Lake Harris,
Tavares; Lake Yale, [town of] Grand Island; Lake Grimn, Leesburg; all Lake
Co. Lake Weir, Oklawaha; Halfmoon Lake, 6 mi. N Lynn; Lake Eaton, 5 mi. NE
Lynn; Oklawaha River, Eureka Springs; all Marion Co, Redwater Lake, 4 mi.
W. Johnson, Putnam Co. JULINGTON CREEK DRAINAGE. Lake on Julington Creek,
2 mi. W Bayard; Duval Co.

Genus Lkimpsilis Rafinesque
Lampsilis Rafnesque 1820, Ann, Gdn, des Sci. Physiques (Bruxelles) 5: 298.

Species listed: Lampsilis cardium Rafinesque, Lampsilis ovam (Say), Lamp-
silisfasciola Rafinesque.

Type specieS, Unio ouatus Say. Subsequent designation, Herrmannsen, 1847,
Indicis Generum Malacozoorum, 1: 575. Ortmann, 1912, Ann. Carnegie, Mus.,
8: 345.

Subgenus Lampsilis
The species of Lampsilis described in this paper belongs to subgenus

Lampsilis. Frierson (1927: 67-86) listed 10 other subgenera, 3 of which are
of his own creation. One of them, Villosa, is now used in place of Microm!/a
Agassiz (see under Villosa). To comment on the other subgenera is not in
the scope of this paper, but on cursory examination, I disagree substant-
ially with Frierson's classification both on a generic and specific level.

Subgenus Lampsilis, while clearly of Interior Basin origin, appears to
have speciated about equally there and in the Apalachicolan and At-
lantic Slope regions.

Lkimpsilis (Lkimpsilis) teres (Rafinesque)
Figure 3A, 12 B, C

Unto teres Rafinesque 1920, Ann. Gen. Sci. Physiques (Bruxelles), 5: 321 (La
riviare Wabash [ Indianal; syntype in Poulson colin., not in ANSP [lost],
figured by Conrad, 1836, Monography Unionidae, no. 6, p. 52, pl. 38, here
selected as type figure, Call, 1900, 24th Ann. Rept. Dept, Geol. and Nat,
Res. Indiana (1899), p. 452, pl. 18.

Unio anodontoides Lea 1831, Trans. Amer. Philos. Soc.,4: 81, pl, 8, fig. 11 (Mis-
sissippi, Alabama, and Ohio rivers; type not in USNM or ANSP [lost]).
Lea, 1831, Obs. Unio, 1: 91.

Unio jioridensis Lea 1852, Trans. Amer. Philos. Soc., 10: 274, pl, 21, fig. 31 (Cha-
chkhi River, West. Florida, figured holotype ANSP. 42081. Clench and
Turner (1956: 202) restricted the type locality to the Choctawatchee River,
1 mi. W Caryville, Holmes Co., Florida). Lea, 1852, Obs, Unio, 5: 30.



1972 JOHNSON: FLORIDA UNIONIDAE 243

Lainpsilis fallaciosus Smith 1899, Bull. U.S. Fish Comm. for 1898, 18: 291, pl, 79
( mouths of narrow arms of [ Mississippi River ],. near Muscatine, [ Musca-
tine Co., Iowa]; [location of type unknown].

Lampsilis anodontoides (Lea). Ortmatin and Walker, 1922, Oce. Papers, Mus.
Zool., Univ. Michfgan, no. 112, p. 60.

Lampsilis anodontoides jioridensis (Lea). Clench and Turner, 1956, Bull. Florida
State Mus,, 1: 201, pl. 3, fig. 1.

Lampsilis anodontoides (Lea). La Rocque, 1967, Geol, Survey, Ohio, Bull.,
62(2): 213, fig. 98.

Lampsilis anodontoides form anodontoides (Lea)  Valentine and Stansbery, 1971,
Sterkiana, no.  42, p. 30.

Ltimpailis anodontoides fullactosa Smith . La Rocque , 1967 , Ceol Survey , Ohio ,
Bull.„62 (2): 215

Lampsills anodontoides form fallaciosa Smith. Valentine and Stansbery, 1971,
Sterkiana, no. 42, p. 30.
DESCRIPTION-Shell medium iii size,.seldom exceeding 1()() min in

length. Outline elliptical or subelliptical. Valves slightly inflated, thin,
but strong, inequilateral. Anterior end regularly rounded; posterior end of
both male and female shell terminating in a pc)int two-thirds of the way up
from the base. Ventral margin straight in males; somewhat arcuate in fe-
males because of marsupial swelling. Dorsal margin straight, forming a
sharp angle with the obliquely descending posterior margin. Posterior ridge
low and rounded, posterior slope slightly concave. Umbos rather full, but
not high, located in the anterior quarter of the shell, their sculpture con-
sisting of numerous ridges looped in the middle but open posteriorly  Perio-
stracum generally smooth except on the posterior slope where it may be
roughened, usually shiny yellow or straw, occasionally brownish, some-
times with fine green rays over the entire surface, but more often, if present,
limited to the posterior end.

Left valve with two subcompressed pseudocardinal teeth, the hind one
somewhat elongated. Hinge line short and narrow, in front of two delicate
straight lateral teeth. Right valve with two triangular, iiarrow pseudocard-
inals separated by a narrow pit, the more atiterior tooth vestigial; one lat-
eral tooth. Beak cavities shallow, a few dorsal muscle scars under the
hinge plate. Anterior adductor muscle scars deep, posterior ones and pallial
line distinct. Nacre white to pale pink, somewhat iridescent.

MEASUREMENTS,-L 90 min, H 41 min, W 30 111111 (Lees Ltike P:iiia-
soffkee, Sumter Co.. male); L 75 min, H 37 imn, W 24 min (Withlacc,c,chee
River,.D unnellon, Marion Co., male); L 73 inm, H 37 11 11 1, W 23 1 11 in (same
ass above, female).

HABITAT, - Li ves i n sand i n either swift or slowl y movi ng water.
REMARKS. - In peninsular Florida, Lanip.silis teres Raftnesque cannot

be confused with any other species. It is distinguished by its long elliptical
shape and shiny yellow, sometimes green rayed periostracum.

Clench and Turner (1956: 158), following Simpson (1914:91), reeog-



244 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

nized floridensis  (Lea ) as a subspecies of L. anodontoides , but admitted that
"young specimens of the typical form would be difficult to separate from

"this subspecies. It is true that specimens of this species tend to be smaller
and thinner toward the southern end of its range, but ~oridensis is not a sub-
species as this concept is currently understood.

It is unfortunate that this species is best known under the name ano-
dontoides, but it has been repeatedly pointed out (Say, 1834, pt. 6 [no
pagination]; Conrad, 1834: 72; F6russac, 1835: 27; Kuester, 1854: 68;
Call, 1899: 452; Utterback, 1916: 442 (179); and Frierson, 1927: 70) that
Rafinesque' s name, teres, has priority. Ortmann and Walker ( 1922 : 60 ),
quoting Pilsbry, offered specious reasons for rejecting this name on the
basis that it might be Lampsilis fallaciosa Smith . The latter is an eco -
phenotypic variant that exhibits green rays on the surface of the shell.
Valentine and Stansbery (1971:30), recognized this variant, but they per-
sist in rising the taxon fallatiosa. The name teres can hardly be considered
nomen oblitum under Article 23 (b) of the International Code (1964).
There is no choice but to recognize it, or ask the Commission to suppress
it.

RANGE.-Interior B:isin generally. West Gulf Coastal region. Ala-
bama-Coosa River system, Apalachicolan region and peninsular Florida;
Northern Mexico, Rio Grande River system, Texas, east to the Withla-
coochee River system, Florida.

~PECIMENS ExAMINED.-Withlacoochee River System:-Withlacoochee
River 9 mi. N Dade City, Withlacoochee River, 1 mi. NW Lacooohee; both
Pasco Co. Little Withlacoochee River, Rerdell; Withlacoochee River, Istachatta
(USNM), both Hernando Co. Lees Lake, Panasofikee, lake, 6 mi. NNW P.ina-
soffkee; both Sumter Co. Lake Tsala Apopka, Floral City; Citrus Co. Withla-
coochee River, Dunnellon; Marion Co.

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1972 JOHNSON: FLORIDA UNIONIDAE 245

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246 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

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1972 JOHNSON: FLORIDA UNIONIDAE 247

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Valentine, Bariy D. and David H. Stansbery. 1971. An introduction to the
Naiades of the Lake Texoma Region, Oklahoma, with notes on the Red
River fauna (Mollused: Unionidae). Sterkiana, no. 42, pp. 1-40.

van der Schalie, Henry and Annette. 1950. The mussels of the Mississippi
River. American Midland Nat.. 44:448-466, maps. 1-2.

Vaughan, Thomas W. 1910. A contribution to the geologic history of the Flo-
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99-185, 15 pls., 6 text figs.

Walter, Waldemar M. 1956. Mollusks of the upper Neuse River Basin, North
Carolina. Jour. Elisha Mitchell Sci. Soc., 72: 262-274, 1 fig.

INDEX TO RELEVANT UNIOMD TAXA
ablatus Lea, Unio cicur Lea, Unio
aequatiLY Lea, Unio columbensis Lea. Unio
aheneus Lea , Unio complanata ( Lightfoot ). Elliptio
am!,SIdala (Le:), Vi/losa confertus Lea, Unio
am!/gdaln C Leal, Villos·n vibe.r conn·cw· Gould, U,Zio
ami/dalum Lea, Unio Cortincu/ina F. C. Baker
Anodonta Lamarek coroinus Lea, Unio
Anodontiii fie Ort man couperiana Lea, Anodo,ita
anodontoides (Lea), Lampsi/8· modontoidey couperlana Lea. Anodonta (Utterbackia)
anodontoideR Len. Unio cou;periana Lea. Anodontri
aqui/ um Lea, Unio craxsidens ( Lamarck ), Elliptio crassiden,
arctata ( Conrad ), Eliiptio cromwellii Lea, Undo
averillii B. H. Wright, Unio cuneata (J. E. Gray). Rangia
barrattii Lea, Unio cunninghami B. H. Wright, Unio
bisselianus Lea, Unio cuspidatu, Lea. Unio
blandingianu.9 Lea, Unio cugnea (Litinaeus),  Anodo,ita
buckle!/1 (Leah Elliptio (:!/gneus Linnaeus, Mutilus
buck/4 ( Lea), Popenaias culindraccus Frierson , Emptio
buckle !/ i Lea , Uitio da// ii B . H , Wright . Unic,
buddianus Lea, Unio dariends (Lea). Elliptio (Elliptio)
burtchinnus S. H. Wright, Unio datiensis Lea, Unio
buxtoni B. H. Wright, Unio decliuis Conrad. Unio
cahni F. C. Baker. Caruncuhina parua de/umbis (Conrad). Vii#m·a
caloosae,1 .· is Dall . Unio (Unio) dia ; ensis. S . H . Wright . Unio
camptodon Say, Unio dispa/ans B . H . Wright. U"iD
camptodon (Say). Uniomerus tetralasmuN dispar Lea, Unio
Carunculina Baker dorei B . H . Wright , Unio



248 BULLETIN FLORIDA STATE MUSEUM Vol. XVI No. 4

downiei ( Lea ). Elliptio crassidens ocmulgeensis Lea. Unio
dun/apiana Lea, Anodonta opacus Lea, Unio
etectrinus Reeve, Unio orcurtii S. H. Wright, Unio
Elliptio Ralinesque oscari B . H . Wright , Unio
excultus Conrad. Unio paludicolor Conrad. Unio
exiguus Lea , Unio paludicolus Gould , Unio
faticic:osc Smith . Lampsili., miodonfoides papraceus Gould . Unio
fallaciosus Smith . Lampsilis parale/ lus Conrad , Unio
ferriwi Marsh, Unio parua (Barnes), Caruncu/ina
.floridensis ( Lea). Lampsi/is anodontoide, parwi ( Barnes ), Toxolasmo
floridensi&· Lea, Unio parous Barnes, Unio
frganus B. H. Wright, Unio paula.(Lea), Coruncu/ma
fuscatus· Lea, Unio paulti.v Lea, Unio
geddingsianus Lea, Unio pawensis Lea, Un,o
geomefricus Lea, Unio peggyae Johnson , Anodonta
glans ( Lea ), Caruncu / ina peggyae Johnson , Anodonta ( Utterbackia)
gracilior Lea, Unio pinet B. H. Wright, Unio
hartii B. H. Wright, Unio (Elhptio) webbianus plantii Lea, Unio
hartwrightii B. H. Wright, Unio Pleurdbeminae
hastatus Lea, Unio Popenaiadinae
hebes Lea . Unio porrectus Conrad . Unio
hepaticus Lea. Unio precostianus Lea, Unio
hinklegi B. H. Wright, Unio pu//a (Conrad), Caruncutina
icterina (Conrad), Eniptio puNatis Lea, Unio
icterina (Conrad), Elliptic (£/liptio) pullatus Lea. Unio
icterinus Conrad, Unio radiolus Lea, Unio
imbecilis Say, Anodonta roueneh Conrad. Unio
imbecilis Say. partim Clench and Turner, Anodonta rivicolus Lea, Unio
imbecillis Say. Anodonta .iuu/aris Conrad, Unio
ineptus Lea, Unio rutilans Lea, Unio
jamesianus Lea, Ullio sanctrumjohanium B. H. Wright,
jagensis ( Lea ), Elliptio ( Elliptio) Unio ( Elliptio)
jagensis Lea. Unio savi (Ward), Uniomerus tetralasmus
jewettii Lea . Unio sayii Ward. Unio
Lampsi/inae Ihering ,·imi/i, Lea, l/nio
lanceolata C Lea ), Elliptio simpsoni B . H . Wright , Unio
lapillus Say, Unio singleganus Marsh, Unio
/epidus Gould. Unio singularia B. H. Wright, Unio
/ienosa (Conrad), villosa squatidus Lea, Unio
fimatulus Conrad, Unio stagnalis Conrad. Unio
liuidus Rafinesque, Unio steamsi B. H. Wright, Unio
livingstonensis Lea. Unio drigos·us (Lea), Elliptio
lucidus Lea, Unio strigosum (Lea). Clench ancl Turner, E//iptio
lugubris Lea, Unio subcroceus Conrad, Unio
manubius Gould, Unio subellip.vis Lea. Unio
marginis Lea,.Unio Nublatus Lea, Unio
marshii B. H. Wright, Unio subturidus Simpson. Unio
magwebbae B . H . Wright , Elliptio . suborbiculata . Say, Anodonte
merceri Lea, Unio sudus Lea, Unio
micans Lea , Unio auttoni B . H . Wright, Unio
Miciom!/a Agassiz symmetricus Lea, Unio
minor Lea, Unio tenuisculus Frierson. Unfo
modiol (formis Lea , Unio teres ( Rafnesque),-Lampsilis ( Lampsilis)
monroensis Lea , Unio t eres Rafinesque , Unio:
nigra Rafinesque, Elliptio refralasmus Say, Unio
nigrinus Lea , Unio fetralasmus (Say). Uniomerus
not,mi B. H. Wright, Unio tetricus Lea, Unio
obesu.i Lea, Unio Toxolasma Rafinesque
obea·us (Lea), Uniomerus frosculus Lea~ Unio
obfuscus Lea, Unio fro.,·sulus Lea, Unio
oblatus Lea, Unio truon, B. H. Wright, Unio
obnubilus Lea, Unio tuomegi Lea, Unio
occuitu.v Lea, Unio Uniomerus Conrad



1972 JOHNSON: FLORIDA UNIONIDAE 249

Unionacea Thiele
Unionidae Fleming
Unioninae Swainson
Utterbac.kia F. C. Baker
Utterbackiana Frierson
uerutu.s Lea, Unio
ve.yicularis Lea, Unio
uibex Conrad. Unio
uibex (Conrad). Villo.ya
Vi/losa Frierson
ui/losus B. H. Wright, Uliio
uilloga (B. H. Wright), Villo.wa
viridans Lea. Unio
uiridicatu.9 Lea, Unio
uiridiradiatus Lea, Unio
wa/toni B. H. Wright. Unio
watereens,3 Lea, Unio
webbiahus B. H. Wright, Unio (Elliptio)
websteri Lea, Unio
whiteianus Lea, Unio
wrightiana Frierson. Lampsilis



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