BIOSTRATIGRAPHY AND PALEOECOLOGY OF THE
PLEISTOCENE INVERTEBRATES FROM THE LEISEY

SHELL PITS, HILLSBOROUGH COUNTY, FLORIDA

Roger W. PorteW, Kevin S. Schindler2, and David Nico13

ABSTRACT

A Pleistocene macro-invertebrate assemblage from two Leisey Shell Corporation pits  located 1 km
inland from Tampa Bay along the Gulf Coast of Florida. is examined. The assemblage is dominated by
marine mollusks and consists of 98 species ofbivalves, 113 species ofgastropods, 3 polyplacophorans, and 3
scaphopods. Other invertebrates identified include 1 poriferan, 3 cnidarians, 16 bryozoans, 1 annelid, 9
a,thropods, and 1 echinoderm. The fossil material was collected at Leisey Shell Pit 1 and I.~isey Shell Pit 3
from three superposed marine shell beds (and spoil) in each pit The strata containing the molluscan faunas
ofthe two lowermost shell horizons (lower shell bed and bone bed) are placed in the Bermont Formation and
contain approximately 3% extinct marine species each. The uppermost unit (upper shell bed) is most similar
to the late Pleistocene Fort Thompson Formation and contains approximately 1% extinct species of marine
mollusks.

At both Leisey pits the base of the lower shell bed contains dark. organic lenses or layers with
freshwater and estuarine mollusks and freshwater and terrestrial vertebrates. The remainder of the lower
shell bed appears to have been deposited under normal marine conditions in relatively quiet waters from 3 to
6 m in depth. The bone bed at each pit (Inisey 1 A and 3A) consists of a thin lens containing an extremely
rich concentration of terrestrial, freshwater, and nearshore marine vertebrate fossils, in addition to the rich,
shallow-marine invertebrate fauna with abundant freshwater gastropods and some terrestrial gastropods. The
molluscan assemblage indicates deposition in a shallow (intertidal to -2 m), protected marine bay or lagoon,
containing abundant marine grass beds and having a large river flowing into iL The upper shell bed at the
Leisey Pits has the least diverse molluscan assemblage, having also been deposited in shallow water
(intertidal to -2 m) under relatively high energy conditions  possibly in a tidal channel.

....  * - ' ' -, Departimn of Naa:ral Scicmes, Florida Museum of
*tural History, University of Florida.  P. 0. Box 117800, Gainesville FL 32611-7800.
- Kevin SA,Wler, 7207 Barcliff Drive, Charlotte NC 28212, U.S.A. (fonierly a Biologist ~ Imerlebnk Palooniology, Depirt=nl of
~atural Scicmes, Florida Muse4m of Natural Histog, University of Florida, P. O. Box 117800, Gaimevilk FL 32611-7800.

IkvW Nicol is Professor Earritus, Dcpartmcal of Geology, Unimrsity of Florida; airient address: 801 14374 Unhirsity Station,
Gainesvil~ FL 32604.

PORTELL, R. W., K. S. SCHINDLER, and D. NICOL. 1995. Biostratigraphy and Paleoecology of
the Pleistocene Invertebrates from the Leisey Shell Pits, Hillsborough Countyy, Florida. Bull.
Florida Mus. Nat. Hist. 37 Pt. I(5):127-164.



128 BULLEr]N FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

RESUMEN

Se examina un ensamblaje Pleistoc6nico de macroinve,lebrados provenientes de dos dep6sitos de
Conchuelas de 1.eisey localizados 1 km tierra adentro de la Bahia de Tampa en la Costa del Golfo en
Florida. El ensamblaje es dominado por moluscos marines  consistiendo de 98 especies de bivatvos, 113
especies de gastripodos, 3 potipla©6foros y 3 scaf6podos. Otros invertebrados identificados incluyen 1
porifero, 3 cnidarios, 16 bryozooa, 1 antlido, 9 artr6podos y 1 equinodermo. El material hil fue colectado
desde tms camas marinas superimpuestas (y estropeadas) del depuito de Conchuelas de kisey 1 y del
dep6sito de Conchuelas de Lcisey 3. En la Forinaci6n Bermont se ubican los estratos de faunas de moluscos
de los doshorizontes de conchuelas mb inferiores (la cama de conchuelas mhs bajay la cama de huesos),
conteniendo cada una aproximadamente 3% de esl)ecies marinas extinguidas. La unidad mds superior (cama
de conchuelas superior) contiene aproximadamente 1% de especies de moluscos marinos extinguides, siendo
la mAs similar a la formaciin Fort Thompson del Pleistoceno Tardio.

En Ambos dep6sitos de Leisey, la base de lacama deconchuelas mdsbaja, inmediatamente por sobre
el contacto con los sedimentos del Mioceno de la Formaci6n Arcadia, contiene capas o lentes organieos
oscuros con moluscos de agua dulce y de estuario, ademas de vertebrados terrestres y de agua dulce. El resto
de las capas de conchuelas mas bajas parecen haber sido depositadas bajo condicioncs marinas normales con
aguas relativamente 1ranquilas de 3 a 6 m de profundidad. La cama de hucsos en Inisey lA y 3A consiste
de lentes delgados que contiene una con©entraci6n extremadamente rica de vertebrados Bsiles terrestres, de
agua dulce, costeros y marinos; ademds de la rica fauna de invertebrados marinos de poca profundidad con
abundantes gastr6podos de agua dulce y algunos gas*6podos terrestres El ensamblaje de moluscos indica
que 6ste fue depositado en una bahia marina o laguna de poca profundidad (intermareal a -2 m), que
contenia abundantes praderas marinas y un gran rio que desembocaba en ella. La cama superior de
conchuelas en los dep6sitos de Leisey contiene el emsamblaje de moluscos menos diverso, la cual tambi6n
fue depositada en aguas de poca profundidad (intermareal a -2 m), bajo condiciones de relativa alta energia.
posiblemente en un r:,n.1 de mareas.

INTRODUCTION

This study summarizes the species composition, biostratigraphy and
paleoecology of the Pleistocene invertebrate fauna from three superposed marine
shell beds at two Leisey Shell Corporation pits (Leisey Shell Pit 1 and Leisey Shell
Pit 3) located about 7 km southwest of Ruskin in southwestern Hillsborough
County, Florida (Sec.15, T325, R18E, Ruskin Quadrangle, USGS 7.5 minute
series; Fig. 1). Both pits are well known for their extremely rich concentrations of
early Pleistocene (Irvingtonian) vertebrate fossils which were collected by the
Vertebrate Paleontology Division staff of the Florida Museum of Natural History
(FLMNH), Frank A. Garcia, and many volunteers from the Tampa Bay Mineral
and Science Club (now Tampa Bay Fossil Club). Their excavations of the main
Leisey Shell Pit 1 and Leisey Shell Pit 3 fossil vertebrate sites were conducted
between July 1983 and September 1984, and between November 1986 and
February 1987, respectively. The vertebrate fossils were collected from thin lenses
within the thick sequences of marine shells at both pits. Because of the potential
for biostratigraphic correlation, based on marine mollusks and terrestrial
mammals, the leaders of the FLMNH excavation crew (G.S. Morgan and R.C.
Hulbert Jr.) collected large samples of invertebrate fossils and bulk matrix from



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 129

0

A
j HILLSBOROUGH

COUNTY

LEISEY SHELL PITS

*.0

B RUSKIN

TAMPA BAY

MYOU

OACH SUN CITY
BAY ; ;„u

0 1"M

Figure 1. (A) General location map ofthe Leisey Shell Pits. Hillsborough County, Florida. (B) Specific
location map ofLeisey Shell Pit 1 and Leisey Shell Pit 3.

not only the thin bone-producing horizons (bone bed) but also the massive shell
beds above (upper shell bed) and below (lower shell bed). This paper is based
primarily on their invertebrate fossil collections and stratigraphic data. Additional



130 BULLEnN FLORIDA MUSEUM NATURAL HISrORY VOL. 37, Fr. I, No. 5

Leisey Shell Pit 3 material was later collected by a field crew from the Invertebrate
Paleontology Division of the FLMNH between February 1989 and May 1990. Both
Leisey Shell Pit 1 and Leisey Shell Pit 3 excavations are now water-filled and no
longer accessible for additional collecting.

Previous studies of Leisey Shell Pit 1 by Hulbert and Morgan (1989) and
Webb et al. (1989), based on preliminary invertebrate identifications, referred the
lower shell bed and bone bed to the Bermont Formation, while the upper shell bed
was referred to the Fort Thompson Formation. Portell et al. (1992), after
compiling a more comprehensive list of fossil mollusks from the three shell units at
Leisey Shell Pit 1, concurred with Hulbert and Morgan (1989) and Webb et at.
(1989) in their placement of these beds within those formations. However, Portell
et al. pointed out that preliminary estimates of extinct species (10% and 12%),
given by Hulbert and Morgan (1989) and Webb et al. (1989), respectively, were too
high and that two Bermont "index" taxa, Miltha carmenae Vokes 1969 and
Fasciolaria okeechobensis Tucker and Wilson 1932, originally reported were not
substantiated. A single specimen of Strombus m®acensis Tucker and Wilson
1933, the only other Bermont "index" taxon reported by Hulbert and Morgan
(1989) and Webb et al. (1989) is now known to have been collected from spoil
(materials out of stratigraphic context) at Leisey Shell Pit 3. Portell et al. (1992)
concurred with the placement of the lower shell bed and bone bed within the
Bermont Formation because their detailed study of Leisey Shell Pit 1 mollusks
revealed the presence of other species, e.g. Semele per/ameUosa Heilprin 1886 and
Conradostrea sculpturata (Conrad 1840), which make their last appearance in the
Bermont Formation elsewhere in Florida, but have never been recorded from the
younger Fort Thompson Formation.

This work represents one of only a few to examine a Bermont invertebrate
assemblage stratigraphically; therefore, it may be useful to help interpret the
complex Plio-Pleistocene stratigraphy of Florida and correlate this unit with
equivalent units outside of Florida. Information contained in this study and
derived from the recently completed Leisey Shell Pit 3 invertebrate faunal list and
the newly updated Leisey Shell Pit 1 invertebrate faunal list (Appendix 1) confirm
most of the previous findings of Hulbert and Morgan (1989), Webb et al. (1989),
and Portell et al. (1992). In addition, the new taxonomic data provided in these
lists will be useful for ongoing fossil mollusk biodiversity compilations (e.g.
Allmon et al. 1993; in press).

ACKNOWLEDGEMENTS

Gary S. Morgan, New Mexico Museum of Natural History (formerly of FLMNH), and Richard C.
Hulbert Jr., Georgia Southern University, were instrumental to this study by providing stratigraphically
collected bulk matrix and by measuring the stratigraphic sections from LEisey Shell Pit 1 and Leisey Shell
Pit 3. Steven D. Emslie, FLMNH, and the late R. Jerry Britt, Jr. assisted in the collection of additional
macro-invertebrate fossils. Fred G. Thompson, Kurt Auffenber& and Kim T. Massey helped with the



PORTELL SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 131

identification of certain molluscan tan and allowed access to the Malacology collections at the FLMNH.
William G. Lyons  Florida Department of Natural Resources, St Petersburg, Florida, and the late Victor A
Zullo, University of North Carolina at Wilmington, kindly identified the polyplacophorans and barnacles.
respectively. Frank J. Maturo, University of Florida. Alan Cheetham, National Museum of Natural History.
and John and Dorothy Soule, University of Southern California, provided identifications for several
bryozoans Photographs of all invertebrate species were taken by Craig W. Oyen. Douglas S. Jones
FLMNH, and William G. Lyons kindly reviewed an earlier draft ofthis manuscript Curation of the 1£isey
invertebrate fossils into the Division of Invertebrate Paleontology of the FLMNH was supported by National
Science Foundation Grants BSR 8711802 and BSR 9002689. This is University of Florida Contribution to
Paleobiology 386.

METHODS AND MATERIALS

The fossiliferous portions of the exposed stratigraphic sections at Leisey Shell
Pit 1 and Leisey Shell Pit 3 were informally divided into three superposed units:
lower shell bed, bone bed, and upper shell bed. The combined thickness of the
three beds at Leisey Shell Pit 1 and Leisey Shell Pit 3 totaled 8.0 m and 6.0 m
respectively (Fig. 2). In order to obtain adequate samples for microvertebrate
analysis, the PLMNH Vertebrate Paleontology crew collected over a metric ton of
matrix, directly from the bone bed of each pit . From this, over a 100 kg sample of
matrix from Leisey Shell Pit 1 and Leisey Shell Pit 3 was washed and all
invertebrate fossils larger than 5.0 mm were removed and identified. A much
smaller sample (about 5 kg) of fine matrix from the bone bed at each pit was sorted
to obtain a representative sample of the micro-mollusks. In addition, bulk samples
of matrix from 5 to 7 kg were collected from the lower and upper shell beds at 1.0
m intervals from three separate stratigraphic sections within each pit. Three
samples from both the lower and upper shell beds were washed through a 1.0 mm
screen, sorted, and identified. Specimens of macro-mollusks were collected in
place from the three main beds in each pit to provide a more accurate picture of the
overall molluscan fauna. We acknowledge that because of the much larger sample
size taken from each bone bed our estimates of the total number of species of the
other units (lower shell bed and upper shell bed) may be comparatively low.

Based on the close proximity of Leisey Shell Pit 1 and Leisey Shell Pit 3 (Fig.
1) and striking similarities in their lithology, faunal preservation, and invertebrate
fauna (especially extinct taxa), the authors consider each specific bed at the two
Leisey pits to be stratigraphic equivalents (i.e. lower shell bed of Leisey Shell Pit 1
= lower shell bed of Leisey Shell Pit 3, etc.). However, we realize that differences
in shell bed thicknesses between the two pits and the occurrence of an
unconformity (above dolomitic "hard layer", Fig. 2) at Leisey Shell Pit 1, and not
Leisey Shell Pit 3, indicate some differences in the accumulation history between
the two pits and, therefore, they may not represent exact time equivalents. In
addition, although the main bone bed from Leisey Shell Pit 1 (Leisey lA) and
Leisey Shell Pit 3 (Leisey 3A) are herein considered stratigraphic equivalents, both



132 BULLEnN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, No. 5

COMPOSITE SECTION COMPOSITE SECTION
LEISEY SHEU PIT I LEISEY SHELL PIT 3

HILLSBOROUGH CO., FLORIDA HILLSBOROUGH CO., FLORIDA

depth below
ground surface

(m)
-0

0.4 4 modern soll zoM 0.4 m, modern soll zone (removed)

E 11 m. unconsolidated
2.4 m, f quartz Band

1O
H

--Id

unconiolldated
guam sand

1.5 m, buff to cream,
UPPER SHELL BED

candy, massive shell
bed_

UPPER SHELLIED 
- ~<Q=<80~BED I

„„pun 
17i&

98 | 
uoliuuod wogues

2 -

'i E
3.0 m, buff, sandy, 0
maasive ahell bed 5 5 LOWER SHELL BED O

i 0
3.3 4 cream to bluish,

- ~ sandy, masslve shell bed
0.4 4 shelly,dolom, -hard layif E
al -4.-* 44, Bod SE€ 8

-I- - - -

1.0 4 darK sandy mud;
g fnahwater shells; bones

LOWER SHEU BED ~

4.3 m, blulah, 1
sandy,

massive shell bed
10Clailieandy

I X mid
5 , modern mean sea level

Ull
tan, hard, massive,

phosphatic dolomlte

El
Figure 2. Composite geologic sections of Leisey Shell Pits 1 and 3, modified slightly from Hulbert and
Morgan (1989).

were lenses with well defined lateral boundaries and clearly differ with respect to
their taphonomic histories.

Identifications were made by comparisons with identified specimens in the
Invertebrate Paleontology and Malacology Collections at the Florida Museum of
Natural History, and by using the descriptions and figures in Abbott (1974) and



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 133

A

tff' 1
3615

,=23„*

B

/ 7/, / /: j 1 jlli: ' /. }fa
i + 'flifm

. 9 #9
--''I:

-

Figure 3. (4 B) Semele perlametiosa Heilprin 1886 from Lower Shell Bed of Leisey Shell Pit 1; UF
13578. Interior and exterior view ofright valve (x 1.0)

01sson and Harbison (1953), among others. Specific names separated by a slash
(/) represent taxonomically ambiguous species that are either indistinguishable to
the authors or represent transitional forms. Typically these taxa, e.g. Anadara
lienosa (Say 1832)6#oridana (Conrad 1867), consist of extinct and extant
equivalents. Basic ecological data of extant species were obtained primarily from
Abbott (1974) and Stanley (1970).



134 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

*Af,13 / 4. - r.

.t. A 4, 0<

D"
r,

= A - ir p

. .4*111, 1 - ~44 a

lil'- i
-

D, 6-4*
f. .

. V A

.

Figure 4. (4 B) Carolinapecten eboreus solanoides (Heilprin 1886) from the lower shell bed of L£isey
Shell Pit 1; UF 13642. Interior and exterior view of left valve (x 0.5). (C, D) Strombus mayacensis
Tucker and Wilson 1933 from spoil materials of Lcisey Shell Pit 3, UF 37952. Apertural and lateral view
of incomplete specimen (x 0.5)

The percentages of extinct taxa, used to help establish placement of the
molluscan assemblages of the lower shell bed, bone bed, and upper shell bed into
the Bermont and Fort Thompson formations, were calculated by excluding all non-
marine species. Also, the aforementioned taxonomically ambiguous species were
judged on the basis of their companion species and therefore considered extant.



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 135

Percentages of extinct mollusks were based solely on materials stratigraphically
collected; spoil materials were not included. All invertebrate fossils discussed in
this paper are housed in the Invertebrate Paleontology Division of the FLMNH.

We follow Harland et al. (1990) in subdividing the Pleistocene into the
early Pleistocene (1.64 to 0.79 Ma), middle Pleistocene (0.79 Ma to 0.13 Ma) and
late Pleistocene (0.13 Ma to 0.01 Ma).

STRATIGRAPHY AND CORRELATION

Miocene strata of the Arcadia Formation underlie the entire Pleistocene
marine shell sequence in the Leisey Shell Pits. The Arcadia sediments consist of a
well-indurated tan to light gray, clayey, phosphatic dolomite containing no
identifiable invertebrate fossils. A major erosional unconformity separates the
Arcadia Formation from the massive, unconsolidated, sandy shell beds that
compose most of the stratigraphic section at Leisey.

Below is a brief stratigraphic description and faunal accounting of the shell
beds (lower shell bed, bone bed, and upper shell bed) along with a short summary
of the more unusual taxa found from spoil. A discussion of the placement of these
beds into their appropriate formations and age of these deposits follows.

Lower Shell Bed.- The lower shell bed at Leisey Shell Pit 1 and Leisey Shell
Pit 3 is composed primarily of marine mollusk shells in a matrix of fine, quartz
sand. Near the base of the lower shell bed at Leisey Shell Pit 1 are irregular lenses
of dark, fine-grained organic sediments containing estuarine, freshwater, and
terrestrial invertebrates, vertebrates, and plants. Similar sediments are found at the
base of the lower shell bed at Leisey Shell Pit 3, not as lenses, but rather as a
distinct layer (up to 1 m thick) containing mainly freshwater and terrestrial
invertebrates and vertebrates.

The combined fauna of the lower shell bed of Leisey Shell Pit 1 and Leisey
Shell Pit 3 consists of the following species distribution: 1 poriferan, 2 cnidarians,
14 bryozoans, 160 mollusks, 1 annelid, 3 arthropods, and 1 echinoderm. This
predominately molluscan assemblage contains 84 species of bivalves (79 marine
and 5 freshwater), 74 species of gastropods (64 marine and 10 nonmarine), and 2
scaphopods. Of the 145 marine mollusk species recorded, only 5 (3%) are extinct.
These are thebivalves: Semele perlamellosa (rig. 3A-B), Carolinapecten eboreus
solarioides (Heilprin 1886) (Fig. 4A-B), Conradostrea sculpturata (Fig. 5A-B),
and Noetia pla(yura (Dall 1898) (Fig. 6A-B), and the gastropod Fermicularia
weberi Olsson and Harbison 1953. K weberi and S per/amellosa were represented
at Leisey Shell Pit 1 as single specimens while the remaining three extinct species
were common at both pits. The only non-molluscan species recognized from this



136 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

»r.,..0~ t# E.;
B

./ i Ff] .», 1< IIT
4.7 4

.

' .4'..C V
U 'i

:;

D
2'OR W p.. -: -s,

8WYG ./. 66 r WS 94I...irY;
- Z ./ 1,Vt ..,· A =. V %456'A ©4. j#*, 1/3. *

' 7'' P ililillillixilili3l://elliFT;'~~~~IL+

Figure 5. (4 B) Conradostrea sculpturata (Conrad 1840) from the lower shell bed of Leisey Shell Pit 1;
UF 13708. Interior and exterior view of left valve (x 1.0). (C, D) Ostreolaequestris (Say 1834) from the
bone bed of Lkisey Shell Pit 3; UF 26935. Interior and exterior view of left valve (x 1.0)

unit as being extinct are the small, scleractinian corals Archohelia limonensis
Vaughan 1919 (Fig. 6E) and Astrangialloridana (Gane 1895).

Bone Bed.- Lying above the lower shell bed is the bone bed, an
unconsolidated, poorly-sorted mixture of marine and freshwater mollusk shells,
bones, fine-grained sand, silt, and dark brown mud. This bed is a well-defined 5-



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 137

30 cm thick layer containing most of the rich vertebrate fauna discussed elsewhere
in this volume. At Leisey Shell Pit 1, there was a 30-50 cm thick layer of
indurated calcareous marl above the bone bed. This "hard layer" contained
abundant freshwater gastropods, especially Planorbella scalaris (Jay 1839), some
fragmented and apparently reworked marine mollusks, fine-grained sand, and
occasional bones of freshwater vertebrates.

The combined fauna of the bone bed at Leisey Shell Pit 1 and Leisey Shell Pit
3 consists of the following species distribution: 1 poriferan, 2 cnidarians, 8
bryozoans, 166 mollusks, 1 annelid, 3 arthropods, and 1 echinoderm. Unlike the
lower shell bed, this unit is dominated by gastropod rather than bivalve species.
All 68 bivalves are marine and of the 93 gastropods, 83 are marine and 10 non-
marine. Two scaphopod and 3 polyplacophoran species are also recorded. Of the
156 marine mollusks recognized, only 4 (3%) are extinct. These include the

bivalves: Noetia platyura, Conradostrea sculpturata, and Carolinapecten eboreus
solarioides and the gastropod Turritella perattenuata Heilprin 1886 (Fig. 6D).
Other extinct taxa recorded from the bone bed include the freshwater gastropod
Stenophysa meigsii (Dall 1890) (Fig. 6C) from Leisey Shell Pit 3 and the coral
Archohelia limonensis from both Leisey pits. Only one of the extinct marine
mollusk taxa (C scu/pturato) occurs in the bone bed at both Leisey pits, while the
remainder are found in either Leisey Shell Pit 1 or Leisey Shell Pit 3, but not both
(see Appendix 1). Another interesting note involves the occurrence of the oysters.
The extinct C sculpturata (Fig. 5A-B) is found in the lower shell bed and bone bed
at both Leisey Shell Pits. The extant Ostreola equestris (Say 1834) (Fig. 5C-D) is
found at Leisey Shell Pit 3 in both the bone bed and upper shell bed. The co-
occurrence of these two taxa, as far as the authors know, has never been
documented and may represent a replacement of the older C. sculpturata by the
younger 0. equestris. The reported occurrence of the extinct bivalve Juhacorbula
scutata (Gardner 1943) from the bone bed at Leisey Shell Pit 1 by Portell et al.
(1992) was in error.

Upper Shell Bed.- Like the lower shell bed and bone bed, the upper shell
bed lacks distinct bedding and is composed predominantly of marine mollusks.
However, in contrast to the bone bed, the upper shell bed has much fewer
freshwater gastropods, is nearly devoid of bones, and has a higher percentage of
fine-grained sand and a lower percentage of mud.

The combined fauna of the upper shell bed consists of the following species
distribution: 1 poriferan, 1 cnidarian, 6 bryozoans, 106 mollusks, 5 arthropods, and
1 echinoderm. The mollusks are represented by 51 marine bivalves, 53 gastropods
(48 marine and 5 nonmarine), and 2 scaphopods. Of the 101 marine mollusks,
only 1 (1%) may be extinct. This is the bivalve Noetia sp. cf. N. platyura, which is
represented by a single fragment and as indicated its identification is tentative.
Like the lower shell bed and bone bed, the extinct coral Archoheha hmonensis is
found in the upper shell bed at both pits.



138 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. 1, No. 5

A B -'ll"wi'Hie
.-6, ~111%*./IU#*liliti'.,3V sgilillii\\\~~,,

ILLI#\\=t
,

C AA Dff;4
'.-

E-4

44 t. ,4 .*

%.
* - 4 %

Figure 6. (A B) Noetia plargura (Da!1 1898) from the lower shell bed of Lkisey Shell Pit 3, UF 37573.
Interior and exterior view of left valve (x 1.0). (C) Stenophysa meigm (Dall 1890) from the bone bed of
Ikisey Shell Pit 3, UF 45869. Apertural view (x 1.0). (D) Turntella peranenuata Heilprin 1886 from the
bone bed ofLeisey Shell Pit 1; UF 10128. Apertural view of incomplete specimen (x 2.0). (E) Archohelia
limonensis Vaughan 1919 from the lower shell bed of Izisey Shell Pit 1, UF 10574. Corallum (x 1.0).



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 139

Spoil.- At Leisey, several tan worth noting were not collected in
stratigraphic context and are known strictly from spoil. These include two bivalve
specimens from Leisey Shell Pit 1 donated to the FLMNH by John Waldrop. The
first  Chione procancellata Mansfield 1932, is known only from the Pliocene
Pinecrest Beds and its equivalents. The second, Anadara scalarina (Heilprin
1886), is known to occur in both the Pliocene Pinecrest Beds and the Plio-
Pleistocene Caloosahatchee Formation and their equivalents. Neither species has
been reported from units younger than the Caloosahatchee. These specimens were
probably derived from a relict pocket of Pinecrest and are not included in the
species counts because they were not part of the stu* materials collected by
FLMNH field crew. As noted earlier, a partial specimen of the gastropod
Strombus mayacensis (Fig. 4C-D) was collected at Leisey Shell Pit 3.

AGE

All strata in the Leisey Shell Pits bounded by the Arcadia Formation below
and the top of the bone bed are referred to the Bermont Formation on the basis of
extinct molluscan tan. DuBar (1974) proposed the name Bermont Formation as
an informal stratigraphic unit for the uppermost fossiliferous marine sands exposed
along Shell Creek in the Bermont Quadrangle, Charlotte County, Florida. DuBar's
(1974) description of the Bermont included scattered excavations and natural
exposures along the Gulf Coast from the type area in Charlotte County northward
to Levy County, along the Caloosahatchee River in Glades and Hendry counties, in
the Lake Okeechobee region, and on the Atlantic coast from Palm Beach County
north to Seminole, Volusia, and Putnam counties.

Earlier names applied to strata DuBar (1974) referred to the Bermont
Formation include: Unit F (DuBar 1962), Unit A (Olsson 1964), and Giales Unit
(Vokes 1963). Hunter (1978) later proposed the informal name Belle Glade
Member (of the Fort Thompson Formation) to include strata from the Belle Glade
Rock Pit in Palm Beach County and surrounding areas south of Lake Okeechobee
that had previously been referred to the Bermont Formation by DuBar (1974).
Hunter considered her Belle Glade Member to be a correlative of the
Okaloakoochee Member of the Fort Thompson Formation ofDuBar (1958).

The Bermont Formation occupies a stratigraphic position between the older
Caloosahatchee Formation and the younger Fort Thompson Formation. At most
localities where the Bermont is exposed, DuBar (1974) found that all or part of the
underlying Caloosahatchee had been removed indicating a period of erosion
between these two formations, presumably corresponding to a marine regression(s)
during a glacial interval(s). The Caloosahatchee Formation was not observed at
the Leisey Shell Pits where the Bermont Formation rests directly upon the
unevenly eroded surface of the Miocene Arcadia Formation. The initiation of



140 BULLErIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. 1, No. 5

Bermont deposition at Leisey is marked by a transgression as the base of the lower
shell bed consists of considerable amounts of dark, organic sediments containing
freshwater and estuarine mollusks as well as freshwater and terrestrial vertebrates.

Most southern Florida formations currently recognized by geologists/
paleontologists are defined by faunal differences based on either Lyellian
percentages (percentage of extinct tan within a particular unit) or the recognition
of certain "index" tan. Lithologic distinctions have played a relatively minor role
and this has led to considerable debate and confusion in the literature (see Scott
1992, p. 21-25). Furthermore faunal comparisons between units have often
ignored taphonomy, facies changes, and paleogeographic differences, factors which
complicate the emerging biozonation schemes for the region.

DuBar (1974) recognized that the Caloosahatchee and Bermont formations
were distinct faunally, that many of the mollusks typical of the Caloosahatchee
were lacking in the Bermont, and that the Bermont contained many unique taxa.
Species listed by DuBar (1962), Vokes (1963), and McGinty (1970) that are
restricted to the Bermont Formation include the bivalves Anadara aequalitas
(Tucker and Wilson 1932) and Miltha carmenae; and the gastropods Fasciolaria
okeechobensis, Fusinus watermani 04. Smith 1936), Haustellum anniae 04. Smith
1940), Lan'rus jucundus (McGinty 1940), L. marwelli Pilsbfy 1939, Melongena
bispinosa 9101\ipi 1844), Murexiella graceae (McGinty 1940), Strombus
mayacensis Tucker & Wilson 1933, TurbineUa hoerlei Vokes 1966, and Fasum
floridanum McGinty 1940. Recently Petuch (1988, 1989) added Lindohva gritlini
Petuch 1988, L. speng/eri Petuch 1988, and Mateo petiti Petuch 1989, and Lyons
(1991) added Latirus caringer (Lamarck 1816) to the list of useful biostratigraphic
indicators for the Bermont Formation. Additional studies by Petuch (1991)
indicate that a considerable number of new "index" taxa will be added to the list of
restricted Bermont species.

At the Leisey Shell Pits, none of these unique taxa were collected from in-
place Bermont sediments (lower shell bed and bone bed). However, a single
broken specimen of Strombus mayacensis, one of the "index" taxa listed above, was
collected from spoil at Leisey Shell Pit 3. DuBar (1974) noted that individuals of
Bermont "index" species are rare and some have never been collected from in-place
deposits. In addition  DuBar (1974; written comm. 1993) stated none of these
"index" tan were found in the type Bermont deposits at Shell Creek. The lack of
"index" taxa and the absence of more tropical species from both Leisey and the
type locality indicate that these two deposits are more closely allied than either is to
the better known, "classic" Bermont deposits surrounding Lake Okeechobee which
contain many more tropical species and all of the "index" taxa mentioned above.

Previous estimates of extinct molluscan taxa for the Bermont Formation
typically ranged from 10% to 30%. Olsson (1969) estimated Unit A (= Bermont
Formation) to contain about 600 species, of which 20% to 30% are extinct. Hoerle
(1970) recognized that approximately 15% of the 434 species collected in the Belle
Glade Rock Pit, Palm Beach County were extinct. However, Lyons (1991) noted



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PrrS PLEISTOCENE INVERTEBRATES 141

that all of the materials collected from Belle Glade Rock Pit were excavated spoil
and therefore it is not known how many of Hoerle's (1970) species may have been
derived from other stratigraphic units. DuBar (written comm. 1993) stated that
when the Bermont molluscan fauna is considered as a whole about 10% of the
species may be extinct. The molluscan assemblage in the lower shell bed and the
bone bed of the Leisey Shell Pits each contains approximately 3% extinct taxa. If
formational assignment was based strictly upon percentages, the entire shell
sequence at Leisey should probably be placed in the Fort Thompson Formation,
which typically contains fewer than 5% extinct taxa (tide Webb et al. 1989).
However, even though the lower shell bed and bone bed at Leisey lack Bermont
"index" taxa and the percentages of extinct species appear low, placement of these
units in the Bermont Formation is justified based on the occurrence of numerous
extinct mollusks that elsewhere in Florida make their last appearance in this unit.
These taxa are the bivalves : Noetia platyura, Conradostrea sculpturata,
Carolinapecten eboreus solarioides, and Semele perlamellosa, and the gastropods:
Turritella perattenuata, Vermicularia weberi, and Stenophysa meigsii.

As mentioned earlier, the Leisey lower shell bed and bone bed lack many of
the more tropical species of mollusks present in the "classic" Bermont faunas (e. g.
Belle Glade Rock Pit). Hulbcrt and Morgan (1989) believed this may reflect the
deposition of the Leisey fauna some 100 km farther north in waters that were
presumably slightly cooler. It is also possible that the Leisey mollusks (and the
type deposits at Shell Creek) may be somewhat younger than those of the more
diverse Bermont faunas in southern Florida and that the assemblages of the lower
shell bed and bone bed at Leisey may represent a temporal intermediate between
the Bermont and Fort Thompson formations.

The upper shell bed is referred to the Fort Thompson Formation because the
fauna contains at least 99% extant tan and all molluscan species that make their
last appearance in the Bermont Formation are absent. Originally named by
Sellards (1919), the "Fort Thompson beds" were described as a series of alternating
marine shell beds and indurated freshwater maris lying unconformably above the
Caloosahatchee Formation between LaBelle and Fort Thompson along the
Caloosahatchee River, Hendry County, Florida. DuBar (1958) described the
marine units of the Fort Thompson in the type area as poorly consolidated tan to
cream-colored sandy shell maris, with the most abundant fossil being the bivalve,
Chione cancellata (Linnaeus 1767). The lithology and faunal composition of the
upper shell bed at the Leisey Shell Pits is very similar to the description of the type
Fort Thompson Formation (except for the lack of indurated freshwater maris).
Outside the type area, the Fort Thompson Formation has been recognized in
numerous localities in southern Florida (DuBar 1958, 1962; Brooks 1968).

The common occurrence of the extinct branching-coral, Archohelia
hmonensis, in the upper shell bed from both Leisey Shell Pits may represent its
first reported occurrence outside the previously known stratigraphic range.



142 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

Weisbord (1974) reported this species from only Caloosahatchee and Unit A
(= Bermont Formation) strata.

As pointed out by Lyons (1991) and Allmon et al. (in press), age
determination of the Plio-Pleistocene units in southern Florida is poorly understood
as are their correlations with deposits outside of the area. Most of these Florida
units are inadequately documented, both lithologically and fa,innily, and are in
need of much more study.

A dichotomy of opinion exists about the age of the Bermont Formation.
Many early workers, including Voices (1963), Taylor (1966), Waller (1969), and
Hoerle (1970), considered the Bermont to be early Pleistocene or late Pliocene or
both. Others workers such as DuBar (1974), DuBar et al. (1991), Petuch (1991),
and Muhs et al. (1992) have placed the Bermont in the middle to late Pleistocene.

DuBar (1974) proposed a minimum age of 0.4 Ma for the Caloosahatchee
Formation and 0.14 to 0.12 Ma for the Fort Thompson Formation. Although
DuBar did not specify an age for the Bermont Formation, it must fall between 0.4
and 0.14 Ma according to his proposed stratigraphic sequence. DuBar et al. (1974)
correlated the Bennont with the Canepatch Formation of South Carolina and the
Flanner Beach Formation of North Carolina based on their similar stratigraphic
positions and molluscan faunas. They stated that all three units are correlative
because each lies directly on apparently correlative early Pleistocene units and is
overlain by seemingly correlative late Pleistocene units.

Blackwelder (1981) placed the age of the Canepatch and Bermont Formations
between 0.5 and 0.4 Ma. Blackwelder assigned the Canepatch Formation to his
Mollusk Zone M2 (Myrtlean Substage) corresponding to the beginning of the
Longian Stage, a time interval proposed to denote the first appearance in the
Atlantic Coastal Plain sequence of essentially modern molluscan assemblages
containing 94-96% living species. As discussed above, typical Bermont faunas
may contain from 70% to 90% extant species, and thus would appear to be older
than Zone M2. Blackwelder's next oldest biozone, Mollusk Zone M3 (Windyan
Substage of the Colerainian Stage), includes molluscan assemblages with only 35%
living species. He placed the James City and Waccamaw Formations of the
Atlantic Coastal Plain and the Caloosahatchee Formation of Florida in Zone M3,
and considered these faunas to be early Pleistocene in age (1.8 to 1.1 Ma), based in
part on helium-uranium dates of 1.9 to 1.7 Ma of corals from the Caloosahatchee
Formation (Bender 1973). Muhs et al. (1992) suggested that these dates are
tentative because of evidence of uranium loss in most of Bender's samples. Lyons
(1991) argued that the Bermont is probably corretative with the Waccamaw
Formation of the Carolinas based on dates provided by Hulbert and Morgan (1989)
and Webb et al. (1989). More recently, Zullo and Harris (1992) correlated the
Bermont with the early Pleistocene James City Formation of the middle Atlantic
Coastal Plain based on sequence stratigraphy.

The Canepatch and equivalent formations of Zone M2 were deposited
following a long depositional hiatus in the middle Atlantic Coastal Plain sequence



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 143

between 1.1 and 0.5 Ma  termed the Windyan hiatus (Blackwelder 1981). The
Bermont molluscan faunas could occur in the time period between 1.1 and 0.5 Ma,
corresponding to Blackweldets Windyan hiatus. Climatic and latitudinal
differences may also have had a significant effect on the timing of extinctions. It is
possible that some extinct species of mollusks survived longer in the more tropical
waters of southern Florida.

Szabo (1985) obtained a uranium-series date of 460 * 100 ka on unaltered
aragonite from corals of the Canepatch Formation but DuBar et al. (1991, p. 602)
stated these corals were probably from the early Pleistocene Waccamaw Formation.
If the Szabo date is correct, then the early Pleistocene age is incorrect; 460 * 100
ka falls within the middle Pleistocene of Harland et al. (1990). Mitterer (1975)
reported an age of 236 ka for the Bermont Formation based on amino acid
racemization of shells of the bivalve Mercenario. Recalculation of Mitterer's data
by Wehmiller and Belknap (1978) placed an age of 0.6 to 0.4 Ma on the Bermont
Formation. Recently, Muhs et al. (1992) obtained uranium-series dates from the
coral, Solenastrea from several Bermont deposits collected near the town of
Okeechobee and from deposits near the southern end of Lake Okeechobee. These
dates range in age from 230-360 ka. They also attempted dating a sample of
Solenastrea sp. cf. S hyades (Dana 1846), found in the bone bed at Leisey Shell
Pit 1; but the much too young date was ascribed to added  secondary uranium.

In summary, the few published dates for the Bermont Formation, along with
the invertebrate biochronology, fail to resolve the age of this unit beyond
suggesting a broad range of ages of between 1.8 and 0.13 Ma (late Pliocene to late
Pleistocene). However, new lines of evidence, such as the mammalian
biochronology (Hulbert and Morgan 1989, Webb et al. 1989, and other papers, this
volume), strontium isotope stratigraphy (Webb et al. 1989, Jones et al., this
volume), and paleomagnetic stratigraphy (Webb et al. 1989, MacFadden, this
volume) suggest an early Pleistocene age (1.6 to 1.0 Ma) for the strata referred to
the Bermont Formation exposed at the Leisey Shell Pits. For a more thorough
review for the geochronology of the Bermont Formation see Lyons (1991) and
Morgan and Hulbert (this volume).

PALEOECOLOGY

The dominant phylum represented by the fossil assemblages of the Leisey
Shell Pits, both in terms of abundance and species diversity, is the Mollusca
(Appendix 1). Of the 248 species of invertebrates identified, 217 (88%) are
mollusks. Consequently, much of the data presented here deal with this group.
General comparisons of species richness, relative abundance, and environment of
deposition between the lower shell bed, bone bed, and upper shell bed are discussed
below.



144 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. 1, No. 5

The majority of mollusks from the Leisey Shell Pits (97%) are extant species.
Therefore, knowledge of the preRrred habitats of modern forms is very useful in
reconstructing the depositional environments. The salinity indicated by the
majority of the mollusks is that of normal sea water, however, freshwater
gastropods (9 species) are extremely abundant in the bone bed and in the indurated
calcareous marl overlying the bone bed at Leisey Shell Pit 1. The freshwater
species Planorbella scalaris is especially common in this unit. Most of these nine
gastropod species are also found in the lower and upper shell beds but are
considerably less abundant, although several freshwater lenses at the base of the
lower shell bed contain tremendous numbers of Fiviparus georgianus (Lea 1834)
and PlanorbeHa duryi (Wetherby 1879). The freshwater gastropod shells from the
indurated calcareous marl and base of the lower shell bed probably accumulated in
a freshwater spring  river, or pond habitat. The remainder of the freshwater
gastropods were probably transported by a river into the shallow marine
environments.

No pennanent hard bottom was present in any of the Leisey units.
Consequently, infaunal species were generally better preserved and represented by
more specimens than epifaunal species. The percentage of infaunal bivalves, for
instance, is 77% of all marine bivalves found at Leisey, a higher-than-average
infaunal element (Nicol 1968). Some epifaunal generg such as Arca, were absent
whne others, such as Chama, Arcinella, Plicatula, and Argopecten were often
poorly preserved. These epifaunal representatives, lacking a hard bottom, probably
attached themselves to dead shells and bones.

The counter-sunk borings of naticid gastropo(Is were commonly seen in
bivalve shells from both Leisey pits. These carnivorous gastropods preyed
preferentially on the small-sized and/or thin-shelled species or the young of larger
species. The mortality in some vulnerable species was great. In a sample of 88
specimens (less than 30.0 mm in length) of Macrocallism nimbosa (Lightfoot
1786) from the bone bed, 29 (33%) were bored by naticids. Specimens of
Pleuromeris tridentata (Say 1%26) and Parvilucina multilineata guomey and
Holmes 1856) were also commonly bored by naticids.

Among other groups of invertebrates, small barnacles (Balanus spp.) and a
small species of branching scleractinian coral (Archoheha limonensis) were most
common. The barnacles (Balanus improvisus Darwin 1854 and B. venustus
Darwin 1854) were more abundant in the lower shell bed and bone bed, while the
corals were more common in the bone bed and upper shell bed. The barnacles
were often found attached to mollusk shells in the lower shell bed and bone bed,
and were occasionally found on bones in the bone bed. Today, Bahmus improvisus
(an estuarine barnacle typically found in lower salinities) and B. venustus (a
subtidal barnacle preferring normal marine salinities) are wide-ranging Atlantic
Basin species (Zullo 1992). Several specimens of massive colonial coral
(So/enastrea sp. cf. S hyades) were found in the bone bed, one of which was
extensively bored by the bivalves Gastrochaena hians (Gmelin 1791) and



PORTELL, SCI{INDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 145

Lithophaga sp. Bryozoans were also commonly found in the Leisey deposits,
usually attached to mollusk shells. Sixteen species, all cheilostomes, were
recovered  including 14 encrusting and 2 free-living forms. A complete analysis of
the'bryozoans is currently underway (Mature and Schindler, in preparation).

Lower Shell Bed.- The lower shell bed has more species of bivalves (84)
than gastropods (74), in contrast to the bone bed, which has more species of
gastropods, and the upper shell bed, in which both groups are almost equally
represented. A diagnostic feature of the lower shell bed, which relates to the
energy of the depositional environment, is the number of different species and
individuals of bivalves preserved as articulated valves. , Twenty-eight species are
represented in this bed by one or more specimens with paired valves. In the bone
bed this is reduced to 12 species; and in the upper shell bed there are only 6
species. Large bivalves such as Mercenaria campechiensis (Gmelin 1791),
Dinocardium robustum Wightfoot 1186), Macrocallista nimbosa, and the thin-
shelled Raeta plicatella (Lamarck 1818), were less apt to be broken in the lower
shell bed sample and were often preserved as articulated valves.

A bed about 20 to 30 cm thick consisting of large numbers of Rangia cuneata
(Sowerby 1831) was found near the base of the lower shell bed in a limited area at
the north end of Leisey Shell Pit 1. A lens of freshwater unionid bivalves occurred
below the bed of Rangia, immediately above the contact with the underlying
Arcadia Formation and also occurred at the base of the lower shell bed at Leisey
Shell Pit 3 (see Bogan and Portell, this volume). Freshwater gastropo(is,
particularly Viviparus georgianus, and Planorbella duryi, were extremely
abundant in several lenses of dark, organic, freshwater sediments at the base of the
lower shell bed. Well preserved articulated specimens of the brackish water oyster,
Crassostrea virgmica (Gmelin 1791), were also present in this basal unit. Neither
these gastropods nor the Crassostrea, however, were found in direct stratigraphic
superposition with each other or with the unionids or Rangia.

The majority of the lower shell bed appears to have been deposited in
relatively quiet water with an estimated depth ranging from 3 to 6 m. The most
likely depositional environment would have been in the deeper regions of a
protected marine embayment similar to present day Tampa Bay (Webb et at. 1989).
The absence of freshwater and estuarine mollusks and freshwater and terrestrial
vertebrates in all but the basal portion of the lower shell bed suggests deposition
farther offshore and in deeper water than the two upper units. It is worth noting
that species more typical of an open continental shelf environment, e.g. Glycymeris
americana (DeFrance 1826) and £ucrassateUa speciosa (A. Adams 1852), were
represented in the lower shell bed and bone bed by a few disarticulated valves
which were possibly cast up by storms.

Bone Bed.- Although gastropod diversity (93 species) exceeds that of
bivalves (68 species) in the bone bed, most of the gastropods are uncommon and



146 BULLEI'IN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

small in size. The bivalves overwhelmingly dominate in both numbers and
average size of individuals. The medium-sized bivalves (length from 10 to 40
mm), Chione cancellata and Carditamera floridana (Conrad 1838), are the two
most abundant marine mollusks in the bone bed, the former species comprising
more than one-half the Volume of mollusk shells in this unit.

Hulbert and Morgan (1989) reported that at Leisey lA (main bone-producing
site), fossils were derived from one of two taphonomically distinct events. The
terrestrial and freshwater elements, including invertebrates, vertebrates, and plants
represent a transported taphocoenose, while a nearshore marine biocoenose is
represented by marine invertebrates and vertebrates. These two taphonomic groups
can be identified not only at Leisey 14 but in the whole, generalized bone bed, as
used in this paper.

One of the most characteristic aspects of the bone bed molluscan fauna, and
the invertebrate representative of the transported taI)hocoenose, is the great
abundance of freshwater gastropods. The freshwater snail Planorbella scalaris
(Jay 1839) is the most abundant gastropod in the bone bed. This species and other
common freshwater snails in the bone bed, Tryonia aequicostata (Pilsbry 1889),
Planorbe#a duryi, and Physella sp., inhabit freshwater streams, springs, marshes
and lakes. These freshwater gastropods were probably transported from inland
areas into a nearshore, shallow marine environment by a large river, probably in
the same way as the terrestrial and freshwater vertebrates reached the site.

Most of the marine invertebrates in the bone bed represent the nearshore
marine biocoenose. An analysis of the 25 most common species of marine
mollusks in the bone bed suggests they were deposited in a nearshore marine
environment with water depths ranging from intertidal to no more than 2 m. Some
of the shells were undoubtedly transported by storms, tides, and currents, as
evidenced by some shell breakage and erosion, the presence of a few specimens of
species that inhabit somewhat deeper water, and the abundance of freshwater
gastropods. The overall well preserved condition of the shells, the common
occurrence of articulated bivalves (though as pointed out earlier, not as common as
in the lower shell bed), and the comparatively high percentage of silt and clay in
the sediment, indicate relatively quiet water conditions and minimal transport.
The majority of the species live today in water approximately 1 m in depth. Over
20% of the gastropo(is in the bone bed, including more than half of the most
common species, occur in association with marine grass beds. The most abundant
marine grass-dwelling gastropods in the bone bed are Rissoina catesbyana Orbigny
1842, Caecum pulchellum Stimpson 1851, Conus jaspideus Gmelin 1791,
Longchaeus crenulatus (Holmes 1859), Bulla striata Bruguiere 1792, and Mitrella
lunata (Say 1826). Two of the most abundant bivalves, Chione cancellata and
Carditamera floridana also occur commonly in marine grass beds. Most of the
remaining gastropods favor a sandy mud bottom or mud flats, as does another
common species of bivalve Anadara transversa (Say 1822). Sand-dwelling species
are present among both the gastropods and bivalves, but are considerably less



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 147

common, which is in accordance with the relatively small fraction of fine-grained
quartz sand in the bone bed sediment.

The absence of rocks or a permanent hard bottom in the bone bed suggests
that ceflain species were probably attached to dead shells and bones. Bryozoans
and oysters (Conradostrea sculpturata) were found attached to some of the bones.

The depositional environment suggested by the marine mollusks of the bone
bed is a shallow (1 to 2 m depth) marine embayment with near normal salinity,
having extensive grass beds, with sediments composed primarily of sand and mud.
A large, slow-moving river probably emptied into the bay, accounting for the
periodic influx, perhaps during floods, of the abundant freshwater gastropods.

Upper Shell Bed.- The upper shell bed has a reduced diversity of both
gastropod (53) and bivalve (51) species when compared to the lower shell and bone
beds. Marine gastropo(Is are comparatively rare in the upper shell bed and most
are either small species or immature specimens of larger species. This unit also
contains the same species of freshwater gastropods found in the bone bed, but they
are much less abundant. At Leisey Shell Pit 1, some of these freshwater gastropo(Is
have been reworked from the underlying freshwater unit, as they have pieces of
indurated marl still attached to them.

Besides shells, the sediment in the upper shell bed consists primarily of fine-
grained quartz sand, with only a very small percentage of silt or clay-sized
particles. The near absence of silt and clay and the high percentage of abraded
shells and shell fragments suggest that the upper shell bed was deposited under
higher energy conditions than the other two beds at Leisey. The number of
bivalves in the upper unit preserved as articulated valves is greatly reduced. The
upper shell bed also seems to have been deposited in the shallowest water depth of
the three units. Shallow-water mollusks that prefer depths ranging from intertidal
to no more than 2 m dominate the upper shell bed. Like the bone bed, but unlike
the lower shell bed, the upper unit contains a number of gastropods that inhabit
marine grass beds in shallow protected bays. Attached epifaunal mollusks,
bryozoans, and barnacles are less common in the upper shell bed, probably
reflecting both the higher energy conditions and the rarity or absence of larger
shells and bones as sites for attachment. The most likely depositional environment
for the upper shell bed was a shallow, nearshore bay or lagoon or perhaps a tidal
channel or inlet leading into a bay.

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___, J. R. Solliday. and J. F. Howard. 1974. Stratigraphy and morphology of Neogene deposits  Neuse
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Hulbert R. C., Jr., and G. S. Morgan. 1989. Stratigraphy, paleoccology, and vertebrate fauna ofthe Leisey
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Hunter, M. E. 1978. What is the Caloosahatchee Mari? Pp. 61-88 in Hydrogeology of south-central
Florida. Southeastern Geol. Soc. Publ. 20.

Lyons, W. G. 1991. Post-Miocene species ofLatirus Montfort, 1810 (Mollusca:Fasciolariidae) of southern
Florida. with a review of regional marine biostratigraphy. Bull. Florida Mus. NaL Hist, Biol. Sci.
35(3):131-308.

McGinty, T. L 1970. Mollusca of the "Glades" Unit of southern Florida: Part I. Introduction and
observations. Tulane Stud. Geol. Paleon. 8:53-56.

Mitterer, R. M. 1975. Ages and diagenetic temperatures of Pleistocene deposits of Florida based on
isoleucine epimerization in Mercenana. Earth Planet Sci. Lett  28:275-282.

Muhs, D. 1, B. J. Szabo, L M©Cartan, P. B. Maal. C. A Bush, and R. B. Halley. 1992. Uranium-series
age estimates of corals from Quaternary marine sediments of southern Florida. Pp. 41-49 in T. M.
Scott and W. D. Allmon. eds. The Plio-Pleistocene stratigraphy and paleontology of southern Florida
Florida Geol. Surv., Spec. Publ. 36. 194 p.

Nicol, D. 1968. Are pelecypods primarily infaunal animals? Nautilus 82:37-43.
Olsson, A A 1964. The geology and stratigraphy ofsouth Florida. Pp. 511-526 in A A  Oisson and R E

Petit, eds. Some Neogene Mollusca from Florida and the Carolinas. Bull. Amer. Paleon
47(217):505-574.
. 1969. A review ofthe late Cenozoic stratigraphy of southern Florida. Pp. 122-34 in J. R  DuBar
and S. S. DuBar, eds. Late Cenozoic stratigraphy of southwestern Florida- Gulf Coast Assoc. Geol.
Societies, Soc. Econ. Paleon. Mineral., Miami Beach. Guidebook Field Trip 4.
, and A Harbison- 1953. Pliocene Mollusca of southern Florida, with special reference to those from
north SL Petersburg; Acad. Nat- Sci. Philadelphia, Monogr. 8:1457.

Petuch, E. J. 1988. Neogenehistory of tropical American mollusks. The Coastal Education and Research
Foundation, Charlottesville, Virginia. 217 pp.

1989. New species of Malea (Gastropoda:Tonnidae) from the Pleistocene of southern Florida
Nautilus 103(3):92-95.



PORTELL, SCHINDLER, NICOL: LEISEY SHELL PITS PLEISTOCENE INVERTEBRATES 149

1991. New gastropods from the Plio-Pleistocene ofsoulhwcstern Florida and the Everglades Basin.
W. H. Dall Paleontological Research Center, Florida Atlantic Univ., Boca Raton. Spec. Publ. 1, 59
PP.

Portell, R. W., K S. Schindler, and G. S. Morgan 1992. ne Pleistoccne molluscan fauna from L£isey
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Plio-Pleistocene stratigraphy and paleontology of southern Florida. Florida Geol. Surv., Spec. Publ.
36.

Sellards, E. H. 1919. Geological sections across the Everglades of Florida. Florida Geol. Surv., 12th Ann.
Rept:67-76.

Scott, T. M. 1992. CAndal plaing stratigraphy: the dichotomy of biostratigraphy and lithostratigraphy - a
philosophical approach to an old problem Pp. 21-25 in T. M. Scott and W. D. Allmon, eds. The
Plio-Pleistocene stratigraphy and paleontology ofsouthem Florida. Florida Geol. Surv., Spec. Publ.
36:1-194.

Stanley, S. M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geol. Soc. Amer.Mem 125. 296 pp.
Szabo, B. J. 1985. Uranium-series dating of fossil corals from marine sediments of southeastern United

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Tulane Stud. Gcol. Paleon. 1(3):93-123.
Waller, T. R. 1969. The evolution oftheA,gopectengibbus stock (Mollusca: Bivalvia), with emphasison

the Tertiary and Quaternary species ofeastern North America. Paleon. Soc. Mem. 3: 1-124.
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Geochronology of a rich early Pleistocene vertebrate fauna, Leisey Shell Pit  Tampa Bay, Florida.
QuaL Res. 32:96-110.

Wehmiller, J. F., and D. F. Belknap. 1978. Alternative kinetic models for the interpretation of amino acid
enantiomeric ratios in Pleistocene mollusks: examples from California, Washington, and Florida
QuaL Res. 9:330-348.

Weisbord, N. E. 1974. Late Cenozoic corals ofsouth Florida. Bull. Amen Paleon. 66(285):259-544.
Zullo, V. A 1992. Review of the Plio-Pleistocene bamacle fauna (Cirrepedia) of Florida. Pp. 117-131 in

T. M. Scott and W. D. Allmon, eds. The Plio-Pleistocene stratigraphy and paleontology of southern
Florida  Florida Geol. Surv., Spec. Publ. 36: 1-194.

- and W. B. Harris. 1992. Sequence stratigraphy of the marine Pliocene and lower Pleistocene
deposits in southwestern Florida: Preliminary assessment Pp. 27-40 in T. M. Scott and W. D.
Allmon, eds. The Plio-Pleistocene stratigraphy and paleontology of southern Florida  Florida Geol.
Sun'., Spec. Publ. 361-194.



APPENDIX 1
150 

B
U

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rIN

 FLO
R

ID
A

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ISTO

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L. 37, F
r. 

N
o. 5

List of fossil invertebrates collected from Lcisey Shell Pits 1 and 3. Occurrence of each species within the three stratigmphic units Oower shell bed,

bone bed, and upper shell bed) and from spoil piles ia noted by an "x- in the appropriate column. Extinct species are preceded by an asterisk (*).

Specific names separated by a slash (0 represent taxonomically ambiguous species that are either indistinguishable to the authors or represent

transitional fbrms.

Leisey 1 Lcisey 3

Lower Upper Lower Upper
ahell Bone shell shell Bone shell

bed bed bed Spoil bed bed bed Spoil

PORIFERA
CLIONIDAE

cf. (Wono F. x x x x x

CNIDARIA
OCULINIDAE

*Archohelia limonensis Vaughan 1919 x x x x x x

RHIZANGHDAE
*Astrangia floddana (Gane 1895) x x

FAVIIDAE
Solenastrea sp. cf. S. hyades (Dana 1846) x

BRYOZOA
MEMBRANIPORIDAE

Membran,bora knuis Desor 1848 x x x x x



CALLOPORIDAE
Antropora leucocypha (Marcus 1937) x x x x

CUPULADRI[DAE
Cupuladda bipomsa Canu & Bassler 1923 x
Discoporella umbellaw depressa (Conrad

1841) x x x x
HIPPOPORINIDAE

Hippoponlna vern'Ui Maturo & Schopf 1968
SMMTINIDAE

Hemisminoidea mateposita (Canu &
Bassler 1923)

X
X
 

X
X

X
X
 

X
X
 

X

X
Parasminind sp. x x
Parasminina nifida morphotype B x

Mature & Schopf 1968
SCHIZOPORELLIDAE

Schizoporetta comum (Gabb & Horn 1862)
Schizoporella erram (Waters 1878) X

X
X
 

*
X

PO
R

TELL. SC
H

IN
D

LER
, N

IC
O

L: LEISEY SH
ELL PITS PLEISTO

C
EN

E IN
VER

TEBR
ATES 

151

Stylopoma minum Canu & Bassler 1923
Slylopoma spongites (Pallas 1766)

CLEIDOCHASMATIDAE
Aimulosia uvulfera (Osburn 1914) x
Cleidochasma comracmm (Waters 1899) x x x x

HIPPOTHOIDAE
Celleporella sp. x

CELLEPORARIIDAE
Cetteporana sp.

MOLLUSCA (POLYPLACOPHORA)
ACANTHOCHHONIDAE

Acanthochimna pygmaea (Pilsbry 1893)
ISCHNOCHITONIDAE

Cl,aempleura apiculata (Say 1834)
/schnochiton sp. X

X
 

X



Appendix 1 Continued
152 

B
U

LLE
TIN

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R

ID
A

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U

S
E

U
M

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A
 

H
IS

TO
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Y
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O
L. 37, F

r. I, N
o. 5

1.cisey 1 Leisey 3

Lower Upper Lower Upper

shell Bone shell shell Bone shell

bed bed bed Spoil bed bed bed Spoil

MOLLUSCA MARINE GASTROPODA)
FISSURELLIDAE

Diodora cayenensis (Lamarck 1822) x x x x x

Diodora meta (von Ihering 1927) x

Diodom sayi (Dall 1889) x

Lucapinetta limatuta (Reeve 1851) x
TROCHIDAE

Cam'ostoma sp.
TURBINIDAE

Turbo castanea Gmelin 1791 x x
RISSOIDAE

Rissoina caksbyana Orbigny 1842 x x

CA~CIDAE
Caecum foridanum Stimpson 1851 x

Caecum imbdcatum Carpenter 1858 x

Caecion putchellum Stimpson 1851 x

Caecum vestitum Folin 1870 x

VrrRINELLIDAE
Cycles#emiscus penmgonus (Gabb 1873) x x

Episcynia inomam (Orbigny 1842) x x x

Teinostoma biscaynense Pitabry & M¢Ginty 1945 x
Vinine#a sp. x x



MODULIDAE
Modulus modulus (Linnaeus 1758)

CERITHUDAE
Bittiolum varium (Pfeiffer 1840) x
Orithium sp. x x x

X
M
 

x 
x
x
 

X
X
 

X

CerW:ium muscanun Say 1832 x
CERrrHIOPSIDAE

Cerithiopsis sp. afT. C. emersoni
(C.B. Adams 1838)

Seita adamsii (H.C. Lea 1845)
TURRWELLIDAE

*Tur,*enaperattenuam Heilprin 1886
*TunigeUa subannulam Heilprin 1886/

acropora Dall 1889 x x x
Vennicula,la fargoi Oisson 1951
* Vermicula,ia weberi Olsson &

Harbison 1953 x

P
O

R
TE

LL SC
H

IN
D

LER
, N

IC
O

L: LE
IS

E
Y

 S
H

E
LL PITS PLEISTO

C
EN

E IN
V

E
R

TE
B

R
ATE

S 
153

TRIPHORIDAE
Triphora metanura (C.B. Adams 1850) x x

EPrrONIIDAE
Epitonium ap. x x x x

STROMBIDAE
Strombus Warns Gmelin 1791 x x x x x x x x
*Strombus mayacensis Tucker &

Wilson 1933 xMEI-ANELLIDAE
Melanda sp. A x x x
Metane# ap. B X

CALYPIRAEIDAE
Catyptmea centralis (Conrad 1841) x
Crepiduln acuteam (Gmelin 1791) x x x x x x





Phy#onomspomum (Gmelin 1791) x x x x x x
Uros*inr perrugam (Conrad 1846)

BUCCINIDAE
Baitya sp. cf. B. parva

(C.B. Adams 1850) x
Canfhands cancellarius (Conrad 1846) x x x x
Cantharus multangulus (Philippi 1848) x

X
 

X
X
 

X
X
 

X
M
 

X
X

X
 

X
 

X
X

Pisania uncta (Conrad 1846) x
COLUMBELLIDAE

Anachis sp. x x x
Anac/its obesa (C.B. Adams 1845) x x
MiweUa lunam (Say 1826) x x

NASSARIIDAE
Nassarius acubis (Say 1822) x
Nassarius albus (Say 1826) x
Nassarius Wba (Say 1822)

X
X
 

X
X

X
 

X
 

X
X

P
O

R
TE

LI. SC
H

IN
D

LER
, N

IC
O

L: LEISEY SH
ELL PITS PLEISTO

C
EN

E IN
VER

TEBR
ATES 

155

MELONGENIDAE
Busycon contrarium (Conrad 1840) x x
Busycon spiratum pyndoldes (Say 1822) x

X
X

X
 

X

X
X

X
 

X
X

X

X
 

X
X

XMetongma corona (Gmelin 1791) x x
FASCIOLARI[DAE

X
X
X
 

X
X

Fasciolar/a litium hunk,ia (G. Perry 1811) x
Fasciolaria Wium Wium G. Fischer 1807 x x
Fasciota,Ya mlipa (Linnaeus-1758) x
Pleumploca gigantea (Kiener 1840) x x x

OLIVIDAE
Oliva sayana Ravenel 1834 x x x
Otivella dealbata (Reeve 1850)
Otive#aforalia (Duclos 1853)
Olivelia mutica (Say 1822)
O#vella pusitta (Marrat 1871)

M 
X

X
X
 

X
 

X
X

X

M
 

X
X
 

X X

MARC]INELLIDAE
Den:imargo ebumeolus (Conrad 1834)



Appendix 1 Continued
156  

B
U

LLE
TIN

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R

ID
A

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U

S
E

U
M

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ATU

R
A

L H
ISTO

R
Y V

O
L. 37, F

r. I, N
o. 5

Leisey l Leisey 3

Lower Upper Lower Upper
shell Bone shell shell Bone shell
bed bed bed Spoil bed bed bed Spoil

/6,alina avenacea (Deshayes 1844) x
Ma,Ninella sp. x x x x x
Marginelia hanteyanum Schwengel 1941 x
Volvwina albolineam (Orbigny 1842)

CANCELLARIIDAE
Trigonostoma tenenon (Philippi 1848) x

CONIDAE
Conus jaspideus Gmelin 1791 x x x x x

TURRIDAE
Crassispira ap. x x

Crassispin, mmpatiasis Bartsch & Rehder 1939 x x
*matosyrinr sp. X

Drillia acumgaia Dall 1890 x
Pyrgocytharapticosa (C.B. Adams 1850) x x

TEREBRIDAE
Terebra sp. X X

Terebm concava (Say 1826)
Terebra distocam (Say 1822)
Terebra protata (Conrad 1846) x

X
X
 

X
X

X

PYRAMIDELL[DAE

X
X
 

XBoonea seminuda (C.B. Adams 1839) x

X
X
 

X

X
 

X
X

Longchaeus crenulam (Holmes 1859)



Odostomia sp.
Odostomia taevigam (Orbigny 1842) x
Odomwomia seminuda (C.B. Adams 1837) x x
Pyramidella crmulam (Holmes 1859) x
Turbonilla ap. x x

X
 

X
X

X
X

X
X
 

X
 

X
X
 

X
X

X
 

X

X
X

X
 

X

ACTEOCINIDAE
Acteocina candei (Orbigny 1842) x x

CYLICHNIDAE
C>Uchnetta bidentam (Orbigny 1841) x

BULLIDAE
Build sulam Bruguiere 1792 x x x x x x

MOLLUSCA ~ON-MARINE GASTROPODA)
VIVIPARIDAE

Viviparus georgianus (Lea 1834) x x x x x xAMPULLARIIDAE
Pomaced sp.

P
O

R
TE

LL, SC
H

IN
D

LER
, N

IC
O

L: LE
IS

E
Y

 S
H

E
LL P

rrS
 PLEISTO

C
EN

E IN
V

E
R

TE
B

R
ATE

S 
157

HYDROBUDAE
Toonia aequicastam (Pilsbry 1889) x x x xhydrobiid gen. and sp. undet. x

PHYSIDAE
Physe#a ap. x x
*Stenophysa meigs/i (Dall 1890)

PLANORBIDAE
Micromenems dilamna avus (Pilsbry 1905) x
Planorbella du,yi (Wetherby 1879) x x x x

X 
X

X
X
 

X

Pianorbetta scalaris (Jay 1839) x x x xSPIRAXIDAE
Eughuidina rosea (Ferussac 1821)

POLYGYRIDAE X
Potygyrs se. X XPolygyra cereolus (Muhlfeld 1816) x
Polygyra avara Say 1818 X



Appendix 1 Continued
158  

B
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LLIO
N

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RIDA M

USEUM
 NA
 

H
ISTO

R
Y V

O
L. 37, F

r. I, N
o. 5

Leisey 1 Leisey 3

Lower Upper Lower Upper
shell Bone shell shell Bone shell
bed bed bed Spoil bed bed bed Spoil

MOLLUSCA (MARINE BIVALVIA)
NUCULIDAE

M,cula proxima Say 1822 x x x

NUCULANIDAE
N,culana acum (Conrad 1832) x x x x x

acm AE
•AB =d.am lienosa (Say 1832)/

foridana (Conrad 1869) x x x x

•801,4 scalarina (Heilprin 1886) x

Anadara transversa (Say 1822) x x x
X 

X
X
 

X
X

X
 

X

X
 

X
X

X
 

X

X
 

X
X

X
 

X

Arropsis adainsi gail 1886) x
Barbatia knera (C.B. Adams 1845)
*Noetia sp. cf. N. pla:yura (Dall 1898) x

*Noeda plaoura (Dall 1898) x x

Noelia ponderosa (Say 1822) x x x

GLYCYMERIDIDAE
Glycymeris ame,icana (DeFrance 1826)

X
X
 

X
X
 

X
X
 

X

64cymeris pecdnam (Gmelin 1791) x x x x x

MYTILIDAE
Brachidonies crusms (Linnaeus 1758) x x x x x x

Li:hophaga 89. x
X

Modiolus sp. x



FrERIIDAE
Pteria cotymbus (Roding 1798) x

PINNIDAE
A#ina sp. x x x x

LIMIDAE
Unta sp.
Limea bronniana Dall 1886 x

OSTREIDAE
*Conrados,rea scidpiumia (Conrad 1840) x x x x x
Crassostrea Wrginica (Gmelin 1791) x x

X
X

X
 

X
X
 

X

Ostreola equesnis (Say 1834) x x
PLICATULIDAE

Plicanda sp. cf. P. gibbosa Lamarck 1801 x x x x x x x
PECI'INIDAE

Aequipecten sp.
A rgopecten inudians (Lamarck 1819) x x x x x x

PO
R

TELL, SC
H

IN
D

LER
, N

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L: LEISEY S
H

E
LL PITS PLEISTO

C
EN

E IN
VER

TEBR
ATES 

159

*Carolinapecten eboreus solarioides
(Henpr~ 1886) x x x x

ANOMIIDAE
Anomia simpler Orbigny 1845

CHAMIDAE
Arcinella comum Conrad 1866 x x x x x x
Chama congregam Conrad 1833 x
Clumu: macerophylla (Gmelin 1791)

X
M

X
 

X
X

X
 

X X
 

X
X
 

X
X

X
X
 

X
X

LUCINmAE
Anodonda sp. cf. A. philippiana (Reeve 1850)
Anodentia alba Link 1807 x x x
Codakia orbicularis (Linnaeus 1758)
Divaricella quadrisukaia (Orbigny 1842)

*
X
 

XLingapensylvanica (Linmeus 1758) x x
*Linga waccamawensis Dall 1903/

amiantus Dall 1901 x x x x
L.ucina nassula (Conrad 1846) x x x x xX

X
 

X



Appendix 1 Continued
160  

B
U

LLE
R

N
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R
ID

A
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U
S

E
U

M
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AT
U

R
A

L H
ISTO

R
Y

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O

L. 37, F
r. I, N

o. 5

Itisey 1 Leisey 3

Lower Upper Lower Upper
shell Bone shell shell Bone shell
bed bed bed Spoil bed bed bed Spoil

Lucina keenae (Chavan 1971)
Panitucina Wanda (Dall & Simpson 1901) x x
Pardlucina multitineaza guomey &

Holmes 1856)
X
 

X
X

X

X
 

X
X

X

Pseudommha~#ondana (Conrad 1833) x
UNGULINIDAE

Diptodonm sp.
LEMONIDAE

Mysellaptanut,*m (Stimpson 1857) x x x

Neaeromya jlo/idana (Dall 1899) x x x x x

LASAEIDAE
Aligena sp.
Parabornia squiU#na Boss 1965 x

CARDn'IDAE
Carditaineraforidana Conrad 1838 x x x x x x

X
 

X
X

X
 

X
X
 

X

Pleuromeris #iden:am (Say 1826) x x x x

Purvmeris pe,plana (Conrad 1841)
CRASSATELLIDAE

Crassinella lum,4,m (Conrad 1834) x x x x x

Eucrussatella speclosa (A. Adams 1852) x x
CARDUDAE

Dinocardium robusium (Lightfoot 1786) x x x x x



odcardium taevigamm (Linnaeus 1758) x x x x x
Trachycanlium egmon#anwn (Shuttleworth

1856) X X X X X X X
Trachycanliwn mu,Ycatum (Linnaeus 1758)

MACTRIDAE
Mactra~wgills Gmelin 1791

X
X
 

*
X

X
X

Mu#nia lageralls (Say 1822) x x x x
Raempticatetta (Lamarck 1818)
Rangia cuneam (Sowerby 1831) x x x x

SOLENIDAE
Ensis minor megisms Pilsbry & MeGinty

1943 X X
TELLINIDAE

Psammowem inms#/am (Say 1826)
Snigma mimbilis (Philippi 1841)
Tentdon, cristam (Recluz 1842) x x

X
 

X
X
 

X
X

X
 

X
X
 

X
 

X
 

X
X

X
X

X
X

X
X

X

Temna sp. X X X X

PO
R

TELL. SC
H

IN
D

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, N

IC
O

L: LEISEY S
H

E
LL PITS PLEISTO

C
EN

E IN
VER

TEBR
ATES  

161

Tellina aequisniam Say 1824
Tellina alkmam Say 1822 x x x

X
 

X
X

Temna magna Spengler 1798
Temna nuens C.B. Adams 1845

SOLECURTnDAE
Solecumis cumingianus (Dunker 1861)
Tagelus divisus (Spengler 1794) x x x x x x

SEMELIDAE
Abra aequalis (Say 1822) x x
um,ingia sp. x
Semele bellastriam (Conrad 1837) x x x x
*Semele periante#oss Heilprin 1886 x
Semele preicua (Pulteney 1799) x x
Semele putpurascens (Gmelin 1791) x



Appendix 1 Continued
162  

BULLEI'IN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, FT. 1, No. 5

Leisey 1 Leisey 3

Lower Upper Lower Upper
shell Bone shell shell Bone shell
bed bed bed Spoil bed bed bed Spoil

VENER-IDAE
Anomakadia aubefiana (Orbigny 1842) x x x x x x
Ozione cancettata (Linnaeus 1767) x x x x x x x
Oione grus (Holmes 1858) x x x x x x
0:ione iniaputpurea (Conrad 1849)
Chione lan'liram (Conrad 1841)
•Chione procancellam Mansfield 1932
Dosinia elegans Conrad 1846

X
M
 

X
X

X
X

X
X
 

X
 

X
Goutdia ce,ina (C.B. Adams 1845)
Macmcallism maculam (Unnaeus 1758)

*
X
 

X
X
X
 

X

Macrocallism nimbosa (Lightfoot 1786) x x x x
Mercenaria campechiensis (Gmelin 1791) x x x

X
X
 

X
X
X
 

X
X

X
 

X
X

X
 

*
K
 

X

Parasmne triquetra (Conrad 1845) x
Pimr ap.
Pimr simpsoni (Dall 1889) x

*
X
 

X
X

Transennella conradina Dall 1883
MYIDAE

Sphenia sp. x

CORBULIDAE
Corbula contracm Say 1822 x x x x x

GASTROCHAENIDAE
Gastrochaena hians (Gmelin 1791) x x x



HIATELLIDAE
Panopea bimincam (Connd 1872) x x x

MOLLUSCA (NON-MARINE BIVALVIA)
UNIONIDAE

Emptoideus stoatianus (Lea 1840) x
Megatondias boR#niana (Lca l 840) x x x
cf. Unerbackia sp.
Unerbackia sp. cf. U. imbecillis (Say 1829)

X
X

X
X
 

X
X

X

Vinosa sp.

MOLLUSCA (SCAPHOPODA)
DENTALUDAE

Denmlium ap. x x x x

PO
R

TELL, S
C

H
IN

D
LE

R
, N

IC
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163

Dentalium antittanon Orbigny 1842
Dentalium Laqueation Verrill 1885 x x x

ANNELIDA
polychaete worm tubes x x x

ARTHROPODA
BALANIDAE

Balanus sp.
Balanus improvisus Darwin 1854 x
Balanus venusms Darwin 1854 x x x x x
Paraconcavus sp.

PYRGOMAT1DAE
Oratoconcha sp. x

CALLIANASSIDAE
Camanassa sp. X X X

LEUCOSHDAE
Pmephona sp. x

PORTUNIDAE



164 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. I, No. 5

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