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*BUiinLLui
of the
FLORIDA
MUSEUM OF
NATURAL HISTORY
PALEONTOLOGY AND GEOLOGY OF THE
LEISEY SHELL PITS,
EARLY PLEISTOCENE OF FLORIDA
PART I
Richard C. Hulbert, Jr., Gary S. Morgan,
and S. David Webb, Volume Editors
Volume 37, Pt. I, Nos. 1-10, pp. 1-344
UNIVERSITY OF FLORIDA
GAINESVILLE
Numbers of the BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY are
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F. WAYNEKING, EDITOR
RHODA J. BRYANT, MANAGING EDITOR
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be addressed to: Managing Editor, Bulletin; Florida Museum of Natural History, University of
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ISSN: 0071-6154
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Publication date: March 14th, 1995
Price $30.00 per set
PALEONTOLOGY AND GEOLOGY OF THE
LEISEY SHELL PITS,
EARLY PLEISTOCENE OF FLORIDA
PART I
Richard C. Hulbert, Jr., Gary S. Morgan,
and S. David Webb, Volume Editors
PREFA CE
For the last two decades, the Leisey Shell Corporation has operated
several shell pits along Tampa Bay southwest of Ruskin in Hillsborough
County, Florida. Like many other mining operations in central and
southern Florida, they excavate shells and sand for use in construction,
primarily as road bedding. As is commonly the case in these shell pits,
the Leisey draglines occasionally unearth vertebrate fossils. Beginning
in the late 1970s and early 1980s several avocational paleontologists
from the Tampa Bay Region, including Frank A. Garcia, Eric
Fernandez, James L. Pendergraft, and James Ranson, periodically
explored the spoil piles and quarry walls of the Leisey Shell Pit and built
up substantial collections of vertebrate fossils. Eric Fernandez and
James Ranson subsequently donated numerous fossils from these early
years of collecting at the Leisey Shell Pit to the Florida Museum of
Natural History (FLMNH, formerly the Florida State Museum).
The situation at the Leisey Shell Pit changed dramatically in July
1983 when a dragline exposed a tremendous concentration of fossil
bones. Fortunately, Frank A. Garcia was collecting fossils in the Leisey
Shell Pit shortly after the dragline exposed this bone bed, now known as
the Leisey Shell Pit 1A Site or Leisey 1A for short (Hulbert and Morgan
1989; Webb et al. 1989). He initially found a great number of bones of
camels, horses, and other large mammals literally covering the spoil
piles. Frank then tracked the fossils to their source, a richly fossiliferous
layer of bones within the exposed stratigraphic section (Garcia 1993).
Shortly afterward he was able to enlist the cooperation of the manager
and owner of the Leisey Shell Pit, Eric Hunter and C. E. "Bud" Leisey,
Jr., respectively, who temporarily ceased quarrying in that area. In all,
the Leisey Shell Corporation held off mining Leisey 1A for 15 months,
thus allowing the nearly complete excavation of the site and recovery of
many thousands of important fossils. For this alone the scientific
community is deeply indebted to the owners of the Leisey Shell Pit.
For the remainder of 1983, Frank A. Garcia, Ronald J. Shrader,
Donald O. Ward, James L. Pendergraft, and their friends excavated
about a quarter of Leisey IA. Of the specimens they recovered, many
were later donated to the FLMNH, including the skull and partial
skeleton of Gymnogyps kofordi Emslie 1988, a new species of large bird
closely related to the California condor. In December of 1983 Frank A.
Garcia donated a large collection of Leisey IA fossils to the FLMNH.
The Museum of Science and Industry in Tampa (MOSI) also houses a
large sample of the specimens collected by Garcia in 1983. Although
paleontologists at the FLMNH became aware of the importance of the
Leisey find in 1983, their field work that year was concentrated on
another site, Haile 21A near Gainesville, which ironically proved to be
very similar to Leisey 1A in age.
In January 1984, S. David Webb and Frank A. Garcia met with
Eric Hunter, William W. Casey, and C. E. "Bud" Leisey, Jr., to plan a
major operation at Leisey in the spring and summer of that year. The
owners of the Leisey Shell Pit transferred their ownership of the fossils
to the FLMNH, and also generously offered to provide heavy equipment
to remove overburden thus exposing a much larger area of the bone bed.
Excavations at Leisey IA resumed April 1, 1984 as a cooperative effort
between the FLMNH, the Tampa Bay Mineral and Science Club, and
the Leisey Shell Corporation. From April until mid-September of 1984
(except for a short break during the hottest weeks of the summer),
several hundred volunteers donated many thousands of person-hours
collecting vertebrate fossils and recording taphonomic data. This
volunteer effort was coordinated by Frank A. Garcia, Rudi Johnson,
Mickey Fowler, Ralph "Tony" Estevez, Ronald J. Shrader, Sue
Bodishbaugh, Red Tincher, and Donald O. Ward. Without the
tremendous contribution of the Leisey volunteers, only a small fraction
of the Leisey 1A Site could have been excavated. It is not possible to
individually list all of the volunteers who freely offered their services
during the spring and summer of 1984, but "you know who you are" and
we are extremely grateful for your many contributions.
Teams of FLMNH paleontologists regularly made the nearly three
hour commute from Gainesville to Leisey during 1984. These field
crews were led by Steven D. Emslie, Richard C. Hulbert, Jr., David
Kendrick, and Gary S. Morgan. Other FLMNH participants included
Mary Ellen Ahearn, Mary B. Ferl, the late Howard H. Converse, Jr.,,
Russell W. McCarty, Ann E. Pratt, S. David Webb, and David B.
Wright. With its combination of high temperatures and humidity,
frequent heavy thunderstorms, and swarms of hungry mosquitoes, the
Florida summer is not the most ideal choice for paleontologic field
work. Despite these hardships, the summer of 1984 is fondly
remembered as one of the most enjoyable and productive in the history
of paleontology in Florida. It remains to date the best example of
cooperation between avocational and professional paleontologists in the
state.
The major scientific importance of Leisey 1A was immediately
apparent, both for the quantity and quality of its specimens. The 1984
dig attracted much attention from the local, state, and national news
media, including a spot on NBC's Today Show and an article in
Newsweek Magazine. Of special note was a lengthy Sunday feature in
the Tampa Tribune, including a full-color poster entitled "Tampa Bay -
One Million B.C." The scientific community was also quickly notified
of the discovery, with presentations in the fall of 1984 at a symposium of
the Institute for Tertiary-Quaternary Studies held in Dallas, Texas, and
at the annual meeting of the Society of Vertebrate Paleontology in
Berkeley, California. However, published descriptions of the site and its
fauna were somewhat slower to appear. This in large part was due to
the enormous number of vertebrate fossils collected over a short time
span, perhaps as many as 50,000 identifiable specimens. Several years
of preparation and curation were necessary before we had a firm grasp
of the fauna as a whole. Indeed, many specimens from the 1984 dig
remain uncatalogued, representing mostly camel and horse postcranials
and isolated turtle shell elements.
Emslie's (1988) description of the extinct Leisey condor was the
first published paper dealing directly with specimens from Leisey 1A,
although Berta (1987) had earlier mentioned the Leisey Smilodon
sample. These were followed by the general overview of Hulbert and
Morgan (1989) that named the Leisey Shell Pit Local Fauna and a
discussion of the site's chronology by Webb et al. (1989). The presence
of Nothrotheriops texanus from Leisey was mentioned by Akersten and
McDonald (1991). The sample of Holmesina from Leisey IA formed an
important component of Hulbert and Morgan's (1993) study of
evolutionary rates in giant armadillos.
Field work continued at Leisey after 1984, although at a less
frantic pace. In the summer of 1986, members of the Tampa Bay Fossil
Club (formed by fossil enthusiasts as a spinoff of the Tampa Bay
Mineral and Science Club) led by Frank A. Garcia excavated what is
now called the Leisey 1B site. This locality was actually the first bone
concentration Garcia discovered at Leisey, but it could not be completely
excavated when it was discovered in January 1981 without damaging a
major road used in the operations of the quarry. In late November 1986
Garcia discovered another major site at Leisey, this time in a newly-
opened quarry on the north side of Gulf City Road. This site is officially
called Leisey Shell Pit 3A, but it is also known as the "Baby Llama
Site." Unlike the other Leisey sites, which have a diverse array of large
mammals, Leisey 3A consists primarily of associated skeletons of
immature individuals of the long-legged llama Hemiauchenia
macrocephala. Leisey 3A was primarily dug by volunteers from the
Tampa Bay Fossil Club, with limited assistance from the FLMNH. In
addition to the abundance of Hemiauchenia, Leisey 3A was rich in small
terrestrial and freshwater vertebrates such as rodents, snakes, and fish.
Excavations at Leisey 3A ceased in February 1987.
Many important specimens of vertebrate fossils were collected at
Leisey between 1987 and 1992, including a partial skeleton of the giant
ground sloth Eremotherium discovered by Frank A. Garcia in March
1992. Steven Beck, D. J. Bethea, Ralph "Tony" Estevez, Wayne Filyaw,
Frank A. Garcia, John Miller, and James L. Pendergraft have all
donated important Leisey vertebrate fossils to the FLMNH over the past
several years. Quarrying in Leisey Shell Pit 3 ceased in 1992, but the
Leisey Corporation acquired another tract of land east of Leisey IA and
mining began there in 1993. Therefore, the potential for significant new
discoveries is certainly great.
Leisey has added an important perspective to vertebrate
paleontology in Florida. After the publication of Webb's Pleistocene
Mammals of Florida in (1974), the focus of vertebrate paleontological
research at the Florida Museum shifted to older Miocene deposits in an
effort to fill gaps in the record. In large measure this was due to the
discovery of the Love Bone Bed in the mid 1970s and the renewed
interest in Thomas Farm and other early Miocene sites in the early
1980s. The near simultaneous discoveries of Leisey IA and Haile 21A
reawakened interest in the early Pleistocene among FLMNH vertebrate
paleontologists. Equally important, before Leisey, interest in Plio-
Pleistocene terrestrial vertebrates from marine shell beds in south
Florida was limited. Instead, the principal places to look for fossils of
this age were always considered to be cave deposits and fissure-fills
exposed by limestone.quarrying in northern Florida (e.g. the Haile and
Reddick sites) or in rivers using SCUBA gear (e.g. the Ichetucknee and
Santa Fe River sites). Leisey changed this attitude, and led to
subsequent important excavations at the Macasphalt (APAC), De Soto,
and Richardson Road shell pits, among others.
The idea for the present volume was conceived shortly after the
completion of the excavations at Leisey IA in the fall of 1984. It was
obvious that serious scientific study of the huge volume of vertebrate
fossils accumulated from Leisey was beyond the capabilities of any
single individual. Therefore, the three compilers of this volume entered
into discussions about the feasibility of organizing a team of specialists
on the various vertebrate groups to study the rich fauna from Leisey IA.
At first those asked to participate in the project were paleontologists
associated with the FLMNH. Several obvious gaps in the coverage of
the vertebrate groups were filled by former University of Florida
students and associates of the FLMNH. By the middle of 1985, most of
the participants had already begun the study of their respective groups.
Discovery of the Leisey 3A Site in late 1986 added a new
dimension to the studies of the Leisey Shell Pit vertebrate fauna, as
small vertebrates were abundant there. The papers on fish and small
mammals were solicited for the Leisey Volume shortly after the
discovery of these rich microvertebrate remains, and the studies of
amphibians and reptiles and birds were both substantially augmented.
Since the inception of the Leisey project, Gary S. Morgan and Richard
C. Hulbert, Jr., had been accumulating geological and stratigraphic data,
as well as collecting samples of invertebrate fossils. These data
eventually spawned two additional papers, an overview of the geology
and vertebrate biochronology of the Leisey Shell Pit and a summary of
the invertebrate paleontology. Paleontologists at FLMNH have begun
multidisciplinary geochronological studies of several paleontological
sites in southern Florida, including Leisey, that have interbedded strata
containing marine invertebrates and terrestrial vertebrates during the
past five years. Two further additions to the Leisey Volume, the ones on
strontium isotope and paleomagnetic stratigraphy, are the results of
these studies.
Even more recently we have continued to invite contributions that
would broaden the coverage of this important site, including studies on
palynology and plant macrofossils and vertebrate taphonomy. Certainly
the most obvious gap in our coverage of the Leisey fauna and flora is the
absence of studies on the various groups of marine microfossils, such as
foraminifera, ostracodes, calcareous nannoplankton, and dinoflagellates.
It should be noted that bulk sediment samples were collected throughout
the stratigraphic section in both the Leisey 1 and 3 pits. These samples
are still available for micropaleontological analysis.
Many present and former students and employees of the FLMNH
have assisted in the curation and preparation of the huge collection of
vertebrate fossils from the Leisey Shell Pit. Along with the editors of
this volume, many others have made significant contributions to the
curation of Leisey vertebrate fossils in the FLMNH collection, including
Steven D. Emslie, David Kendrick, Laura Kozuch, Arthur R. Poyer,
Ann E. Pratt, and Erika H. Simons. Russell W. McCarty, Howard H.
Converse, Jr., Cedric Wynn, Patrick Hylton, and John Church prepared
many of the larger Leisey fossils from plaster jackets. Mary Ellen
Ahearn took the photographs appearing in many of the vertebrate
articles. Dianna C. Carver patiently assisted with manuscript typing.
We would like to extend our special appreciation to the managing editor
of the FLMNH Bulletin, Rhoda J. Bryant, without whose assistance,
expertise, and patience this volume would not have been completed.
We would like to thank the National Science Foundation (NSF)
which provided monetary support for all aspects of the Leisey Project.
Field work at Leisey was supported by NSF Grant EAR 8708045 to S.
David Webb. Curation of the Leisey vertebrates at the FLMNH was
supported by NSF collection improvement grants BSR 8314649 and
BSR 8902822 to S. David Webb and Bruce J. MacFadden, while
curation of invertebrate fossils was supported by BSR 8711802 and
9002689 to Douglas S. Jones. We are especially grateful to the Knight
Foundation, Reed and Barbara Toomey, and James and Lori Toomey,
whose generous financial contributions funded the printing costs for
publication of the Leisey Volume. Additional funding of publication
costs was provided by the Division of Sponsored Research, University of
Florida, and Georgia Southern University.
Last, but certainly not least, we and all of the many hundreds of
professional and avocational paleontologists who have participated in
excavations at the Leisey Shell Pit during the past decade, would like to
extend our deepest thanks and appreciation to the owners, managers,
and employees of the Leisey Shell Corporation, in particular, C. E.
"Bud" Leisey, Jr., Kim Leisey, LeAnn Casey, William W. Casey, Eric
Hunter, and Page Youngblood. The Leisey Shell Corporation did much
more than just cease their operations in the vicinity of the two major
Leisey sites in 1983-1984 and again in 1986-1987. They
wholeheartedly supported the excavations by providing heavy earth-
moving equipment and operators for removing layers of overburden and
a storage trailer to protect equipment and specimens from the elements.
The Leisey Corporation also provided funds for supplies, photographic
and art work for many of the papers published here, and travel expenses
to bring several of the Leisey specialists to Gainesville to study fossils.
The tremendous success of the two major Leisey excavations would
certainly not have been possible without the help of the Leisey Shell
Corporation. The study of paleontology in Florida has benefitted
immeasurably from their generosity.
LITERATURE CITED
Akersten, W. A., and H. G. McDonald. 1991. Nothrotheriops from the Pleistocene of
Oklahoma and paleogeography of the genus. Southwest. Nat. 36(2):178-185.
Berta, A. 1987. The sabrecat Smilodon gracilis from Florida and a discussion of its
relationships (Smilodontini, Felidae, Mammalia). Bull. Florida State Mus., Biol. Sci.
31(1):1-63.
Emslie, S. D. 1988. The fossil history and phylogenetic relationships of condors
(Ciconiiformes: Vulturidae) in the New World. J. Vert Paleon. 8:212-228.
Garcia, F. A. 1993. Miracle at Cockroach Bay ... The Leisey Shell Fossils. Garcia
Paleontology, Inc., St Petersburg, Florida, 32 pp.
Hulbert, R. C., Jr., and G. S. Morgan. 1989. Stratigraphy, paleoecology, and vertebrate fauna
of the Leisey Shell Pit Local Fauna, early Pleistocene (Irvingtonian) of southwestern
Florida. Pap. Florida Paleon. 2:1-20.
___ and 1993. Quantitative and qualitative evolution in the giant armadillo
Holmesina. Pp. 134-177 in R. A. Martin and A. D. Bamosky, eds. Morphological
change in Quaternary mammals of North America. Cambridge Univ. Press,
Cambridge.
Webb, S. D. (editor). 1974. Pleistocene mammals of Florida. Univ. Presses Florida,
Gainesville, 270 pp.
Webb, S. D., G. S. Morgan, R. C. Hulbert, Jr., D. S. Jones, B. J. MacFadden, and P. A.
Mueller. 1989. Geochronology of a rich early Pleistocene vertebrate fauna, Leisey
Shell Pit, Tampa Bay, Florida. Quat Res. 32(1):1-15.
Richard C. Hulbert, Jr.
Gary S. Morgan
S. David Webb
PALEONTOLOGY AND GEOLOGY OF THE LEISEY SHELL
PITS, EARLY PLEISTOCENE OF FLORIDA
VOLUME 37, PART I
TABLE OF CONTENTS
1. Overview of the geology and vertebrate biochronology of the Leisey Shell Pit
Local Fauna, Hillsborough County, Florida
Gary S. Morgan and Richard C. Hulbert, Jr. .................................................... 1
2. Strontium isotope stratigraphy and age estimates for the Leisey Shell Pit faunas,
Hillsborough County, Florida
Douglas S. Jones, Paul A. Mueller, Teresa Acosta,
and Robert D. Shuster .......................... ..... ............................. 93
3. Magnetic polarity stratigraphy and correlation of the Leisey Shell Pits,
Hillsborough County, Florida
Bruce J. MacFadden ............................................ .......... 107
4. Preliminary palynology and macroplant report for the Leisey Shell Pits,
Hillsborough County, Florida
Fredrick J. Rich and Lee Ann Newsom ................................................17
5. Biostratigraphy and paleoecology of the Pleistocene invertebrates from the
Leisey Shell Pits, Hillsborough County, Florida
Roger W. Portell, Kevin S. Schindler, and David Nicol.................................... 127
6. Early Pleistocene freshwater bivalves (Mollusca: Unionidae) from the
Leisey Shell Pits, Hillsborough County, Florida
Arthur Bogan and Roger W Portell........................ ......... .............. 165
7. Taphonomy of the terrestrial mammals of Leisey Shell Pit lA,
Hillsborough County, Florida
Ann E. Pratt and Richard C. Hulbert, Jr ........................ ... ................... 177
8. Chondrichthyes and Osteichthyes from the early Pleistocene Leisey Shell Pit
local fauna, Hillsborough County, Florida
Sylvia Scudder, Erika Simons, and Gary S. Morgan .......................................251
9. Pleistocene amphibians and reptiles from the Leisey Shell Pits,
Hillsborough County, Florida
Peter A M eylan......................... ................................... 273
10. An early Irvingtonian avifauna from Leisey Shell Pits, Hillsborough County, Florida
Steven D. Emslie.................................................... ................299
OVERVIEW OF THE GEOLOGY AND VERTEBRATE
BIOCHRONOLOGY OF THE LEISEY SHELL PIT LOCAL
FAUNA, HILLSBOROUGH COUNTY, FLORIDA
Gary S. Morgan' and Richard C. Hulbert, Jr.2
ABSTRACT
The Leisey Shell Pit Local Fauna was collected from two adjacent commercial shell mines located 7
km southwest of Ruskin and less than 1 km inland from Tampa Bay in Hillsborough County, Florida.
Leisey Shell Pit is one of the most diverse Irvingtonian vertebrate faunas in North America, composed of at
least 203 species: 14 sharks, 9 rays, 50 bony fish, 3 amphibians, 26 reptiles, 52 birds, and 49 mammals.
Leisey IA and Leisey 3A are the largest of the four principal sites that constitute the Leisey Shell Pit Local
Fauna. Leisey 1A has extensive samples of large mammals, in particular: the camelids Palaeolama
mirifica and Hemiauchenia macrocephala, two species of Equus, the primitive mammoth Mammuthus
hayi, the ground sloths Paramylodon harlani and Nothrotheriops texanus, the giant tapir Tapirus haysii,
the peccary Platygonus vetus, the gracile sabercat Smilodon gracilis, the short-faced bear Arctodus
pristinus, and the canid Canis edwardii. The large mammal fauna from Leisey 3A is dominated by
associated juvenile skeletons ofHemiauchenia macrocephala, while its diverse microvertebrate assemblage
contains important samples of freshwater fish, sirens, aquatic snakes, birds, and small mammals.
Sediments in the Leisey Shell Pit are referred to four formations, each of which has produced
vertebrate fossils. At the base of the stratigraphic section is an indurated, tan to light gray dolostone referred
to the Arcadia Formation. A horse tooth identified as "Merychippus" tertius derived from this unit is
indicative of an early Barstovian age (early middle Miocene). Reworked sediments on top of the Arcadia
Formation contain several land mammals, including the horses Nannippus aztecus, Neohipparion eurystyle,
and Cormohipparion ingenuum, whose association is characteristic of Florida late early Hemphillian (late
Miocene) faunas. Phosphatic gravel and spheroidal metaquartzite pebbles in this unit are typical of the upper
Bone Valley Formation. The major concentrations of vertebrate fossils in the Leisey Shell Pit occur in thin,
irregular layers of organic-rich sediment distributed throughout about 7 m of sandy marine shell beds
referred to the Bermont Formation, which unconformably overlies the Arcadia Formation. The large
assemblage of land mammals from these organic units in the Bermont Formation is early Irvingtonian (early
Pleistocene) in age. Shell beds of the Fort Thompson Formation occur in the Leisey section above an
T he senior author is a paleontologist at the New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque NM 87104-
1375 (formerly a Senior Biological Scientist at the Florida Museum of Natural History, P.O. Box 117800, Gainesville FL 32611-7800, USA).
The junior author is an Assistant Professor of Geology at Georgia Southern University, Statesboro GA 31460-8149, USA.
MORGAN, G. S., and R. C. HULBERT, JR. 1995. Overview of the Geology and Vertebrate
Biochronology of the Leisey Shell Pit Local Fauna, Hillsborough County, Florida. Bull. Florida Mus. Nat.
Hist. 37 Pt I(1):1-92.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. 1, NO. 1
erosional unconformity at the top of the Bermont Formation. The Fort Thompson shell beds contain a
distinctly younger mammalian fauna, including the bison Bison sp., the cotton rat Sigmodon hispidus, and
the tapir Tapirus veroensis, typical of the late Pleistocene (Rancholabrean).
Geochronological data derived from paleomagnetic analysis, strontium isotope geochronology, sea
levels, and molluscan and vertebrate biochronology have been incorporated in a multidisciplinary effort to
determine the age of the Bermont Formation at the Leisey Shell Pit. Molluscan biochronology implies an
early or middle Pleistocene age, whereas mammalian biochronology is considerably more precise strongly
favoring an early Pleistocene age between 1.6 and 1.0 Ma. All paleomagnetic samples from the Bermont
Formation at Leisey have reversed polarity and are referred to the Matuyama Chron, indicating an age
greater than 0.78 Ma. The Leisey Bermont section does not correlate to the Jaramillo Subchron of normal
polarity between 1.07 and 0.99 Ma. It is also unlikely the Leisey beds were deposited during the youngest
portion of the Matuyama between 0.99 and 0.78 Ma. The combination of data from magnetic polarity,
mammalian biochronology, and sea level strongly indicates that the Bermont Formation at Leisey is older
than 1.07 Ma and younger than 1.55 Ma. Strontium isotope ratios from Chione cancellata (Mollusca)
shells from Leisey indicate an age between 1 and 2 Ma.
Leisey is younger than Florida's best known earliest Irvingtonian (latest Pliocene) faunas, Inglis IA
and De Soto Shell Pit, based on the presence of five genera unknown in North America before the
Pleistocene: Nothrotheriops, Lutra, Castoroides, Palaeolama, and Mammuthus. Four rodents from
Leisey, Geomys pinetis, Erethizon dorsatum, Sigmodon libitinus, and Ondatra annectens, differ at the
species level from their congeners at Inglis and De Soto. Leisey also lacks various Blancan holdover species
found in the two older faunas, including the dwarf Florida form of Megalonyx leptostomus,
Chasmaporthetes ossifragus, Trigonictis macrodon, and Capromeryx arizonensis. The occurrence of
numerous species at Leisey that are unknown after the early Irvingtonian, including Glyptotherium
arizonae, Pachyarmatherium leiseyi, Holmesina floridanus, Nothrotheriops texanus, Sigmodon libitinus,
and Canis edwardii, further constrains the age of this fauna, ruling out a middle Irvingtonian or younger age
assignment. The Leisey mammalian fauna correlates most closely with the late early Irvingtonian, between
about 1.6 and 1.0 Ma. Other Florida faunas similar in age to Leisey are Haile 16A, Haile 21A, Crystal
River Power Plant, Pool Branch, Payne Creek Mine, Rigby Shell Pit, and Punta Gorda. Of these sites Haile
16A is probably somewhat older (between 1.6 and 1.3 Ma) based on the occurrence of several holdovers
from Florida late Blancan and earliest Irvingtonian faunas, including Sylvilagus webbi, Geomys propinetis,
and Trigonictis.
Western early Irvingtonian faunas that are correlatives of Leisey include: Gilliland, Texas;
Holloman, Oklahoma; Kentuck, Nash, and Wathena in Kansas; Sappa, Nebraska; and Java, South Dakota.
Among these sites, Leisey has the largest number of diagnostic taxa in common with the Gilliland Local
Fauna, including Glyptotherium arizonae, a medium-sized Holmesina floridanus, Nothrotheriops texanus,
Canis edwardii, Tapirus haysii, and primitive Mammuthus. Leisey and other late early Irvingtonian faunas
are younger than earliest Irvingtonian faunas (2.0 to 1.6 Ma) such as Curtis Ranch, Arizona and Inglis 1A
and De Soto Shell Pit of Florida, and are older than middle Irvingtonian faunas (1.0-0.6 Ma), including: the
type Irvington fauna from California; Cudahy, Kansas; Conard Fissure, Arkansas; Cumberland Cave,
Maryland; Port Kennedy Cave, Pennsylvania; Hamilton Cave, West Virginia; and the Florida equivalent
McLeod Limerock Mine.
The occurrence of land mammals in estuarine, freshwater, and terrestrial units within the
predominantly nearshore marine late Pliocene and Pleistocene shell bed sequence of southern peninsular
Florida (Pinecrest Beds, Caloosahatchee Formation, Bermont Formation, and Fort Thompson Formation in
ascending stratigraphic order) has allowed a precision in dating these deposits not previously possible. The
Pinecrest Beds, the uppermost unit of the Tamiami Formation, contain land mammal faunas of late Blancan
age (2.5-2.0 Ma) based on the association of Nannippus and a large suite of Neotropical immigrants,
including Dasypus, Holmesina, Glyptotherium, Glossotherium, Eremotherium, and Neochoerus. Florida
late Blancan faunas derived from the Pinecrest Beds, or found in association with Pinecrest molluscan
faunas, include Macasphalt Shell Pit, Acline Shell Pit, St. Petersburg Times, Kissimmee River, Brighton
Canal, and Lehigh Acres. Vertebrate faunas from the overlying Caloosahatchee Formation lack typical
Blancan forms, including Borophagus, Nannippus, Equus (Dolichohippus), and Rhynchotherium, and
contain taxa typical of earliest Irvingtonian (2.0-1.6 Ma) faunas, such as Inglis IA. The two richest earliest
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 3
Irvingtonian faunas from the Caloosahatchee Formation in southern Florida are the De Soto Shell Pit and
Forsberg Shell Pit. Vertebrate faunas from the Bermont Formation are late early Irvingtonian (1.6-1.0 Ma)
in age, and are typified by the Leisey Shell Pit, as well as Rigby Shell Pit, Crystal River Power Plant, and
Punta Gorda. Vertebrate faunas associated with the Fort Thompson Formation typically contain Bison and
other taxa characteristic of the Rancholabrean Land Mammal Age.
RESUME
La Fauna Local del Dep6sito de Conchuelas de Leisey fue colectada de dos minas de conchuela
comerciales adyacentes y localizadas 7 km al suroeste de Ruskin y a menos de 1 km tierra adentro de la
Bahia de Tampa, en el Condado Hillsborough, Florida. La fauna de vertebrados del Dep6sito de Conchuelas
de Leisey es una de las mis diversas del Irvingtoniano en Norte Am6rica, estando compuesta de por lo
menos 203 species: 14 tiburones, 9 rayas, 50 peces tele6steos, 3 anfibios, 26 reptiles, 52 aves y 49
mamiferos. Leisey IA y Leisey 3A son los mayores de cuatro sitios principles que constituyen la Fauna
Local de los Dep6sitos de Conchuelas de Leisey. Leisey IA contiene vastas muestras de grandes mamiferos,
en particular: los cam6lidos Palaeolama mirifica y Hemiauchenia macrocephala; dos species de Equus;
el mamut primitive Mammuthus hayi; los perezosos terrestres Paramylodon harlani y Nothrotheriops
texanus; el tapir gigante Tapirus haysii; el pecari Platygonus vetus; el tigre dientes de sable grlcil Smilodon
gracilis; el oso de rostro corto Arctodus pristinus; y el cinido Canis edwardii. La fauna de grandes
mamiferos de Leisey 3A es dominada por esqueletos asociados de juveniles de Hemiauchenia
macrocephala, mientras que su diverse ensamblaje de microvertebrados contiene muestras importantes de
peces de agua dulce, salamandras, culebras acuiticas, aves y pequefios mamiferos.
Los sedimentos en el Dep6sito de Conchuelas de Leisey son referidos a cuatro formaciones, cada una
de las cuales ha producido f6siles de vertebrados. En la base de la secci6n estratigrifica se encuentra una
dolostona endurecida marr6n a gris claro, referida como Formaci6n Arcadia. Un diente del caballo
identificado como "Merychippus" tertius derivado de esta unidad es indicative de una edad del Barstoviano
temprano (principios del Mioceno medio). Los sedimentos retrabajados encontrados encima de la Formaci6n
Arcad ia contienen varies mamiferos terrestres, incluyendo los caballos Nannippus aztecus, Neohipparion
eurystyle y Cormohipparion ingenuum cuya asociaci6n es caracteristica de las faunas de fines del
Hemphilliano temprano de Florida (Mioceno tardio). Las gravas fosfiticas y piedrecillas esferoidales de
metaquarzita en esta unidad son tipicas de la Formaci6n superior del Valle Hueso. Las mayores
concentraciones de vertebrados f6siles en el Dep6sito de Conchuelas de Leisey ocurren en delgadas e
irregulares capas de sedimentos ricos en material organica distribuidos a lo largo de aproximadamente 7 m de
camas arenosas de conchuelas marinas referidas como a la Formaci6n Bermont, la cual se ubica sobre la
Formaci6n Arcadia. El vasto ensamblaje de mamiferos terrestres encontrados en estas unidades orgAnicas
en la Formaci6n Bermont pertenece al Irvingtoniano temprano (Pleistoceno temprano). Camas de
conchuelas pertenecientes a la Formaci6n Fort Thompson ocurren en la secci6n Leisey por sobre una
inconformidad erosiva encima de la Formaci6n Bermont Las camas de conchuelas de Fort Thompson
contienen una fauna de mamiferos distintivamente mis j6ven, e incluyen a: el bis6n Bison sp., la rata de
algod6n Sigmodon hispidus y el tapir Tapirus veroensis, todos tipicos del Pleistoceno tardio
(Rancholabreano).
En un esfuerzo multidisciplinario para determinar la edad de la Formaci6n de Bermont en el Dep6sito
de Conchuelas de Leisey, se han incorporado datos geocronol6gicos derivados de anilisis paleomagn6ticos;
geocronologia de is6topos de estroncio; niveles del mar y biocronologia de moluscos y vertebrados. La
biocronologia de moluscos indica una edad del Pleistoceno temprano o medio, mientras que la biocronologia
de mamiferos es considerablemente mas precisa, favoreciendo fuertemente una edad del Pleistoceno
temprano, de entire 1.6 y 1.0 Ma. Todas las muestras paleomagneticas de la Formaci6n Bermont en Leisey
tienen una polaridad revertida y son referidas al Cron de Matuyama, indicando una edad mayor que 0.78
Ma. La secci6n de Bermont en Leisey no se correlaciona con el Subcron de Jaramillo de polaridad normal,
de entire 1.07 y 0.99 Ma. Es tambi6n poco probable que las camas de Leisey hayan sido depositadas durante
la porci6n mis j6ven de Matuyama, entire 0.99 y 0.78 Ma. La combination de datos de polaridad
magn6tica, biocronologia de mamiferos y niveles del mar indican fuertemente que la Formaci6n Bermont en
Leisey es mls antigua que 1.07 Ma y mis reciente que 1.55 Ma. La proporci6n de is6topos de estroncio
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
obtenidos de conchas de Chione cancellata (Mollusca) de Leisey indica una edad de entire 1 y 2 Ma.
En base a la presencia de cinco generous desconocidos en Norte Am6rica antes del Pleistoceno
(Nothrotheriops, Lutra, Castoroides, Palaeolama y Mammuthus), la fauna de Leisey es mas j6ven que
otras faunas mejor conocidas del Irvingtoniano mis temprano de Florida (Plioceno mis tardio): Inglis IA y
el Dep6sito de Conchuelas de De Soto. Cuatro roedores de Leisey difieren a nivel de especie de sus
cong6neres de Inglis y De Soto: Geomys pinetis, Erethizon dorsatum, Sigmodon libitinus y Ondatra
annectens. Leisey tambi6n carece de varies sobrevivientes del Blancano, los cuales se encuentran en las
otras dos faunas mis antiguas: la forma enana de Florida de Megalonyx leptostomus, Chasmaporthetes
ossifragus, Trigonictis macrodon y Capromeryx arizonensis. La presencia de numerosas species en
Leisey desconocidas para despu6s del Irvingtoniano temprano, tales como Glyptotherium arizonae,
Pachyarmatherium leiseyi, Holmesina floridanus, Nothrotheriops texanus, Sigmodon libitinus y Canis
edwardii, constrifien afn mAs la edad de esta fauna, descartando una edad del Irvingtoniano medio o mis
reciente. La fauna de mamiferos de Leisey se correlaciona mis cercanamente con fines del Irvingtoniano
temprano, entire aproximadamente 1.6 y 1.0 Ma. Otras faunas de Florida de similar edad a Leisey son: Haile
16A, Haile 21A, Planta El6ctrica de Crystal River, Pool Branch, Mina Payne Creek, Dep6sito de
Conchuelas de Rigby y Punta Gorda. De estos sitios, Haile 16A es probablemente algo mis antiguo entiree
1.6 y 1.3 Ma) en base a la ocurrencia de varies sobrevivientes de las faunas del Blancano tardio y el mis
temprano Irvingtoniano de Florida, tales como Sylvilagus webbi, Geomys propinetis y Trigonictis.
Entre las faunas Irvingtonianas del oeste que son correlativas a Leisey se incluyen a: Gilliland,
Texas; Holloman, Oklahoma; Kentuck, Nash y Wathena en Kansas; Sappa, Nebraska; y Java, Dakota del
Sur. De entire estos sitios, Leisey compare un ndmero mayor taxas de diagn6sticas con la Fauna Local de
Gilliland, incluyendo Glyptotherium arizonae, una Holmesina floridanus de tamanio medio,
Nothrotheriops texanus, Canis edwardii, Tapirus haysii y un Mammuthus primitive. Leisey y otras faunas
del Irvingtoniano temprano son: mis j6venes que faunas del Irvingtoniano mis temprano (2.0 a 1.6 Ma),
tales como Rancho Curtis, Arizona e Inglis 1A y el Dep6sito de Conchuelas de De Soto en Florida; y: mis
antiguas que faunas del Irvingtoniano medio (1.0-0.6 Ma), incluyendo la Formaci6n tipo de Irvington en
California; Cudahy, Kansas; Fisura Conard, Arkansas; Cueva Cumberland, Maryland; Cueva Puerto
Kennedy, Pennsylvania; Cueva Hamilton, Virginia del Oeste; y el equivalent de Florida, la mina McLeod
Limerock.
La ocurrencia de mamiferos terrestres en unidades estuarinas, de agua dulce y terrestres dentro de la
sequencia de camas de conchuelas predominantemente marinas y costeras de el sur de Florida peninsular
pertenecientes al Plioceno tardio y al Pleistoceno (Camas de Pinecrest, Formaci6n Caloosahatchee,
Formaci6n Bermont y Formaci6n Fort Thompson, en orden estratigrifico ascendente) ha permitido una
precision en la determinaci6n de su edad previamente impossible. Las Camas de Pinecrest, que constituyen la
unidad mis superior de la Formaci6n Tamiami, contienen faunas de mamiferos terrestres del Blancano
tardio (2.5-2.0 Ma), en base a la asociaci6n de Nannippus y una gran variedad de inmigrantes
Neotropicales, que incluyen Dasypus, Holmesina, Glyptotherium, Glossotherium, Eremotherium y
Neochoerus. Las faunas del Blancano tardio de Florida derivadas de las Camas de Pinecrest, o encontradas
en asociaci6n con faunas de moluscos de Pinecrest incluyen el Dep6sito de Conchuelas de Macasphalt, el
Dep6sito de Conchuelas de Acline, St Petesburg Times, Rio Kissimmee, Canal Brighton y Lehigh Acres.
Las faunas de vertebrados de la Formaci6n sobrepuesta de Caloosahatchee carecen de formas Blancanas
tipicas, incluyendo Borophagus, Nannippus, Equus (Dolichohippus) y Rhynchotherium, e incluyen taxas
tipicas del Irvingtoniano mis temprano (2.0-1.6 Ma), tales como Inglis IA. Las faunas mis ricas del
Irvingtoniano mis temprano de la Formaci6n Caloosahatchee en el sur de Florida son el Dep6sito de
Conchuelas de De Soto y el Dep6sito de Conchuelas de Forsberg. Las faunas de vertebrados de la
Formaci6n Bermont son de fines del Irvingtoniano temprano (1.6-1.0 Ma) y son tipificadas por el Dep6sito
de Conchuelas de Leisey, asi como tambien el Dep6sito de Conchuelas de Rigby, la Planta El6ctrica de
Crystal River y Punta Gorda. Las faunas de vertebrados asociadas con la Formaci6n Fort Thompson
contienen tipicamente Bison y otras taxas caracteristicas de la Edad de Mamiferos Terrestres del
Rancholabreano.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 5
INTRODUCTION
A remarkable concentration of early Pleistocene (Irvingtonian) vertebrate
fossils was discovered by Frank Garcia in July 1983 after it was uncovered by a
dragline during routine mining operations at the Leisey Shell Pit. This rich bone
bed, designated Leisey Shell Pit 1A, is located about 7 km southwest of Ruskin in
Hillsborough County, Florida (27042'N latitude, 82030'W longitude). Field teams
from the Florida State Museum (now the Florida Museum of Natural History) and
a large volunteer crew organized by the Tampa Bay Mineral and Science Club
excavated the Leisey Shell Pit 1A Site almost continuously from April to
September 1984. The total areal extent of the bone bed was approximately 2000
m2, about 1300 m2 of which was excavated during 1984, whereas 700 m2 had been
dug by avocational paleontologists in 1983. All fossils recovered in 1984 are
housed in the Vertebrate Paleontology Collection of the Florida Museum of Natural
History, University of Florida (UF).
Frank Garcia discovered a second rich concentration of vertebrate fossils at
Leisey in November 1986, this time in a newly opened pit on the north side of Gulf
City Road, only about 0.5 km north of Leisey 1A (Fig. 1). This site, Leisey Shell
Pit 3A, was dug by Garcia and a volunteer field crew, as well as UF personnel,
between November 1986 and February 1987. Although not as rich as Leisey 1A in
numbers of species or individuals of large mammals, Leisey 3A in many ways
complements the fauna from the original site, particularly in the comparatively
large samples of freshwater and terrestrial microvertebrates. The most
characteristic aspect of the Leisey 3A site is the abundance of the long-limbed
llama, Hemiauchenia macrocephala, especially juvenile individuals.
The Leisey Shell Pit Local Fauna (LF), named by Hulbert and Morgan
(1989), is composed of vertebrate fossils collected from Leisey Shell Pit 1A and
3A, as well as two other smaller sites, Leisey Shell Pit 1B and 3B. The four named
Leisey sites are located in two nearly contiguous shell pits mined by the Leisey
Shell Corporation. These concentrations of vertebrate fossils occur in thin layers
or lenses throughout a stratigraphic interval of about 7 m within the predominantly
marine shell beds of the Bermont Formation. The two shell pits, along with a third
pit owned by the same corporation, have been assigned numbers (Leisey Shell Pit
1-3) and each major concentration of bones within a numbered pit has been given a
letter designation. This is the standard system now used by the UF vertebrate
paleontology program for naming and numbering specific localities within
commercial quarrying operations. For example, Leisey Shell Pit 1A (shortened to
Leisey IA for convenience) specifies only those fossils recovered from the large site
collected in 1983-1984. Fossils not from designated sites, often those collected
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
HILLSBOROUGH
COUNTY
LEISEY SHELL PITS
Figure 1. (A) Map of the Leisey Shell Pit region showing the location of the three Leisey pits and the four
vertebrate-bearing sites mentioned in the text. (B) Map showing general location of the Leisey Shell Pit in
Florida and Hillsborough County.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 7
from excavation spoil piles, are assigned a quarry number, but not a letter (e.g.
Leisey Shell Pit 2). This supercedes a previous system that used roman numerals
to designate quarry numbers (e.g. Coleman IIA is now Coleman 2A).
The maps in Figure 1 show the location of the three Leisey Shell Pits in
southwestern Hillsborough County. Leisey 1 is located in the SWA of section 15
and Leisey 3 is in the S/2 of the NW'A and the S' /of the NE% of section 15, T32S,
RI8E, Ruskin Quadrangle (USGS 7.5 minute series, 1981). Leisey 2 is in the SE'
of section 16, T32S, R18E, Cockroach Bay Quadrangle (1969). The exact
coordinates of the two Leisey sites that have produced the vast majority of the
vertebrate fossils upon which the papers in this volume are based are as follows:
Leisey Shell Pit 1A (NW4, SE'4, SW Sec. 15, T32S, R18E) and Leisey Shell Pit
3A (SW/, SE4, NW% Sec. 15, T32S, R18E). Leisey 1A and 3A were completely
excavated in 1983-1984 and 1986-1987, respectively, and have since been
destroyed by mining operations. Although Leisey 1A and 3A are the largest and
richest sites in the Leisey Shell Pit, there are numerous other Leisey sites, three of
which have been given separate designations (Leisey 1B, 1C, and 3B; see Fig. 1).
Because all three Leisey pits are less than 1 km inland from Tampa Bay to the
west and are below 3 m in elevation, the local water table in this region is very
near the ground surface. Most of the important bone concentrations were actually
below mean sea level (i.e. more than 3 m below the present ground surface). Like
most other commercial shell mining operations along Florida's Gulf Coast, the
water levels in the Leisey pits are maintained at artificially low levels by pumping
so that mining can be conducted above water. The Leisey 1 and 2 pits are no
longer being actively mined and have been allowed to fill with water. Mining
ceased at Leisey 3 in 1992 and it will subsequently flood as well.
There are no in-place samples of vertebrate fossils from Leisey 2 in the UF
collection. Leisey 2 is also the only one of the three pits that contains a substantial
number of Rancholabrean fossils, although Irvingtonian vertebrates are common in
this pit as well. We have been unable to confidently establish the age of many of
the specimens from Leisey 2 because they were collected from spoil piles, and thus
have excluded them from this analysis and from the Leisey Shell Pit LF.
The Leisey Shell Pit LF is one of the richest early Pleistocene (Irvingtonian)
vertebrate faunas in North America, both in terms of numbers of species and
individuals. The vertebrate assemblage from Leisey is composed of 203 species,
including 23 species of cartilaginous fish (14 sharks and 9 rays), 50 species of bony
fish, 3 amphibians, 26 reptiles, 52 birds, and 49 mammals. Table 1 provides a
complete vertebrate faunal list for the Leisey Shell Pit LF. To date, over 15,000
specimens of vertebrate fossils have been catalogued from the Leisey Shell Pit LF,
the majority of which are from Leisey 1A. Most of the mammal papers in this
volume are based primarily on material from Leisey 1A. However, Leisey 3A has a
much richer vertebrate microfauna than does Leisey IA, and consequently the
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
papers on fish, amphibians and reptiles, birds, and small mammals contain
considerable data on fossils from Leisey 3A.
ACKNOWLEDGMENTS
This review represents a compilation of data from a variety of sources, including field studies at
Leisey and other Florida Pliocene and Pleistocene sites by ourselves and many others, study of the extensive
collections of Plio-Pleistocene vertebrate fossils housed in the Florida Museum of Natural History, and a
comprehensive review of the literature on this topic. We thank the many contributors to this volume who
permitted us to briefly summarize their work so that we could provide a complete vertebrate faunal list and a
reasonably comprehensive review of the geochronology and biochronology of the Leisey Shell Pit We are
especially grateful to the authors of the papers on the various vertebrate groups who allowed us to combine
the individual faunal lists from their papers in order to produce a complete vertebrate faunal list for the
Leisey Shell Pit Local Fauna (Table 1).
As with all of the contributors to this volume, we are very grateful to the personnel of the Leisey Shell
Corporation for all of their support and to the hundreds of volunteers who worked with the Florida Museum
of Natural History in the excavation of the Leisey Shell Pit Sites lA and 3A. The Preface to this volume
should be consulted for more extended acknowledgments regarding the assistance provided by the Leisey
Shell Corporation and the dedicated volunteer crews co-ordinated by the Tampa Bay Fossil Club. We would
also like to thank several of our colleagues, including Steven D. Emslie, David Kendrick, Arthur R. Poyer,
and Ann E. Pratt, who over the past ten years have participated "above and beyond the call of duty" in the
excavation of several of the critically important sites discussed here, in particular Leisey, Macasphalt Shell
Pit, Haile 7C, and Haile 21A. The following individuals donated important specimens of vertebrate fossils
to the UF collection from various Florida Blancan and Irvingtonian sites: Stephen Beck, D. J. Bethea, Lelia
and William Brayfield, the late Howard H. Converse, Jr., Daniel Delgado, Ralph "Tony" Estevez, Wayne
Filyaw, Frank A. Garcia, George Heslep, Mitchell Hope, Muriel Hunter, Eric Kendrew, the late Philip
Kinsey, Lany Martin, Thomas Missimer, James Pendergraft, Brian Ridgway, Barbara Toomey, James
Toomey, Reed Toomey, the late Benjamin I. Waller, Suzan Watts, and Steven Wilson. Field work at Leisey
and Macasphalt and curation of vertebrate fossils from these and several other sites discussed here was
supported by National Science Foundation Grants EAR 8708045, BSR 8314649 and BSR 8902822. This
is University of Florida Contribution to Paleobiology Number 401.
ABBREVIATIONS
UF Vertebrate Paleontology Collection, Florida Museum of Natural History, University of Florida,
Gainesville.
LF Local Fauna.
NALMA North American Land Mammal Age.
Ma Mega-anna, millions of years before present on the radioisotopic time scale.
ka kilo-anna, thousands of years before present on the radioisotopic time scale.
cm centimeter.
m meter.
km kilometer.
T Township.
R Range.
P/p upper/lower premolar (e.g. P4 is an upper fourth premolar).
M/m upper/lower molar (e.g. ml is a lower first molar).
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 9
Table 1. Vertebrate faunal list of the early Irvingtonian Leisey Shell Pit Local Fauna, Hillsborough County,
Florida. This list has been compiled from the individual papers on the vertebrate groups published elsewhere
in this volume. Within each family the genera and species are listed in alphabetical, not phylogenetic, order.
Several species of amphibians and reptiles (Meylan, this volume) and birds (Emslie, this volume) are
excluded from this list because they were identified only from Leisey 2 which contains a mixed assemblage
of Irvingtonian and Rancholabrean taxa.
Chondrichthyes
Batoidea
Pristidae
Pristis sp.
Dasyatidae
Dasyatis--2 species
Myliobatidae
Aetobatus narinari
Myliobatis sp.
Rhinoptera bonasus
Rhynchobatidae
Rhynchobatus sp.
Galeomorpha
Orectolobidae
Ginglymostoma cirratum
Ginglymostoma serra
Odontaspidae
Odontaspis taurus
Lamnidae
Carcharodon carcharias
Isurus hastalis
Isurus oxyrinchus
Hemigaleidae
Hemipristis serra
Carcharhinidae
Carcharhinus acronotus
Carcharhinus leucas
Carcharhinus limbatus
Carcharhinus obscurus
Carcharhinus plumbeus
Galeocerdo cuvieri
Negaprion brevirostris
Rhizoprionodon terraenovae
Sphyrna mokarran
Osteichthyes
Semionotiformes
Lepisosteidae
Atractosteus spatula
Lepisosteus cf. L. oculatus
Lepisosteus osseus
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. f, NO. 1
Table 1 Continued
Amiiformes
Amiidae
Amia calva
Elopiformes
Elopidae
Elops saurus
Megalops atlanticus
Anguilliformes
Anguillidae
Anguilla rostrata
Clupeiformes
Clupeidae
genus and species indet.
Salmoniformes
Esocidae
Esox sp.
Cypriniformes
Cyprinidae
Notemigonus chrysoleucas
Catostomidae
Erimyzon cf E. sucetta
Siluriformes
Ariidae
Ariusfelis
Bagre marinus
Ictaluridae
Ameiurus natalis
Ameiurus nebulosus
Batrachoidiformes
Batrachoididae
Opsanus sp.
Atheriniformes
Alherinidae
cf. Menidia sp.
Cyprinodontidae
Cyprinodon variegatus
c Floridichthyes sp.
Fundulus cf. F. grandis
Fundulus seminolis
Fundulus majalis
Exocoetidae
cf. Hyporhamphus sp.
Perciformes
Carangidae
Caranx hippos
cf. Trachinotus sp.
Centrarchidae
Lepomis cf. L. auritus
Lepomis gulosus
Lepomis microlophus
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 11
Table 1 Continued
Micropterus salmoides
Pomoxis nigromaculatus
Centropomidae
Centropomus sp.
Ephippidae
Chaetodipterusfaber
Labridae
Lachnolaimus maximus
Mugilidae
Mugil sp.
Percichthyidae
cf. Morone sp.
Sciaenidae
Bairdiella cf. B. chrysoura
Cynoscion cf. C. nebulosus
Micropogonias undulatus
Pogonias cromis
Sciaenops ocellatus
Sparidae
Archosargus probatocephalus
Calamus sp.
Lagodon rhomboides
Sphyraenidae
Sphyraena barracuda
Pleuronectiformes
Bothidae
genus and species indet.
Tetraodontiformes
Balistidae
Balistes sp.
Diodontidae
Chilomycterus schoepfi
Diodon sp.
Ostraciidae
Lactophrys sp.
Amphibia
Urodela
Sirenidae
Siren lacertina
Anura
Bufonidae
Bufo cf. B. terrestris
Ranidae
cf. Rana sp.
Reptilia
Crocodylia
Alligatoridae
Alligator mississippiensis
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 1 Continued
Squamata
Lacertilia
Anguidae
Ophisaurus compressus
Serpentes
Boidae
cf. Tropidophis sp.
Colubridae
Coluber sp./Masticophis sp.
Elaphe cf. E. obsoleta
Farancia sp.
Lampropeltis getulus
Nerodia cf. N. fasciata
Regina cf. R. alleni
cf. Thamnophis sp.
Viperidae
Agkistrodon piscivorous
Crotalus cf. C. adamanteus
Sistrurus miliarius
Testudines
Cheloniidae
Caretta carettal
Chelonia mydas
Chelydridae
Chelydra serpentina
Macroclemys temmincki
Testudinidae
Gopherus polyphemus
Hesperotestudo crassiscutata
Hesperotestudo mlynarskii
Emydidae
Pseudemys sp.
Terrapene carolina
Trachemys script
Kinostenidae
Kinosternon sp.
Trionychidae
Apaloneferox
Aves
Gaviiformes
Gaviidae
Gavia concinna
Gavia immer
Podicipediformes
Podicipedidae
Podiceps sp.
Podilymbus podiceps
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
Table 1 Continued
Pelecanifonnes
Pelecanidae
Pelecanus cf. P. erythrorhynchos
Phalacrocoracidae
Phalacrocorax-2 species
Anhingidae
Anhinga anhinga
Anhinga sp.
Ciconiifonnes
Ardeidae
Ardea sp.
Casmerodias albus
Egretta cf. E. tricolor
Threskiomnthidae
Ajaia chione new species
Eudocimus leiseyi new species
genus and sp. indet.
Ciconiidae
Ciconia maltha
Ciconia sp.
Teratomithidae
Teratornis cf. T. incredibilis
Teratornis merriami
Vulturidae
Gymnogyps kofordi
Anseriformes
Anatidae
Anabernicula gracilenta
Anas americana
Anas crecca
Anas platyrhynchos
Aythya affinis
Aythya americana
Aythya collaris
Aythya marila
Branta canadensis
Branta dickeyi
Bucephala albeola
Cygnus buccinator
Mergus serrator
Olor sp.
Somateria cf. S. spectabilis
Accipitriformes
Accipitridae
Amplibuteo sp.
Aquila sp.
Buteo cf. B. lineatus
Buteo sp.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 1 Continued
Gallifonnes
Phasianidae
Colinus sp.
Meleagris leopoldi/M. anza
Gruifonnes
Rallidae
Fulica americana
Rallus sp.
Gruidae
Grus canadensis
Grus sp.
Charadriifonnes
Recurvirostridae
Recurvirostra sp.
Scolopacidae
Limosa cf. L.fedora
Alcidae
gen. et sp. indet
Phoenicopteridae
Phoenicopterus copei
Phoenicopterus ruber
Strigiformes
Strigidae
Bubo virginianus
Passerifonnes
Corvidae
Corvus sp.
Mammalia
Xenarthra
Dasypodidae
Dasypus bellus
Pachyarmatherium leiseyi new genus and species
Pampatheriidae
Holmesinafloridanus
Glyptodontidae
Glyptotherium arizonae
Megalonychidae
Megalonyx wheatleyi
Mylodontidae
Paramylodon harlani
Megatheriidae
Eremotherium n. sp.
Nothrotheriops texanus
Insectivora
Soricidae
Blarina cf. B. carolinensis
Carnivora
Canidae
Canis armbrusteri
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 15
Table 1 Continued
Canis edwardii
Urocyon sp.
Phocidae
Monachus tropicalis
Procyonidae
Procyon n. sp.
Ursidae
Arctodus pristinus
Mustelidae
Mustelafrenata
Lutra canadensis
Spilogaleputorius
Felidae
Homotherium n. sp.
Lynx rufus
Miracinonyx inexpectatus
Smilodon gracilis
Rodentia
Castoridae
Castoroides leiseyorum new species
Geomyidae
Geomys pinetis
Erethizontidae
Erethizon dorsatum
Hydrochaeridae
Neochoerus sp.
Muridae
Ondatra annectens
Pedomys n. sp.
Podomys n. sp.
Sigmodon libitinus
Synaptomys sp.
Lagomorpha
Leporidae
Lepus cfL. townsendii
Sylvilagusfloridanus
Perissodactyla
Tapiridae
Tapirus haysii
Equidae
Equus "fraternus"
Equus "leidyi"
Equus (Hemionus) n. sp.
Artiodactyla
Tayassudiae
Mylohyusfossilis
Platygonus vetus
Camelidae
Hemiauchenia macrocephala2
Palaeolama mirifica
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 1 Continued
Cervidae
Odocoileus virginianus
Cetacea
Delphinidae
cf. Stenella sp.
cf. Tursiops sp.
genus and species indet.
Sirenia
Trichechidae
Trichechus sp.
Proboscidea
Mammutidae
Mammut americanum
Gomphotheriidae
Cuvieronius tropics
Elephantidae
Mammuthus hayi
SCollected from Fort Thompson Formation above Leisey IA bone bed.
2 Webb and Stehi (this volume) use H. seymourensis for the Leisey sample ofHemiauchenia.
DEFINITION OF CHRONOLOGIC AND
BIOCHRONOLOGIC BOUNDARIES.
In this section, we discuss the age of the Pliocene/Pleistocene boundary and
the subdivisions of the Pleistocene, as well as the definitions and subdivisions of
the Blancan and Irvingtonian Land Mammal Ages used in this study. This is
necessitated by recently published changes to the geologic and geomagnetic
polarity time scales (Fig. 2). The Pliocene/Pleistocene boundary is now generally
placed at 1.64 Ma, above the top of the Olduvai Normal Subchron of the
Matuyama Chron (Berggren et al. 1985; Harland et al. 1990). This is younger
than the 2.0 Ma date for this boundary recognized until recently by most North
American vertebrate paleontologists (e.g. Kurtdn and Anderson 1980). The older
2.0 Ma date was convenient because it also approximated the boundary between the
Blancan and Irvingtonian Land Mammal Ages. With the recognition of a younger
Pliocene/Pleistocene boundary, the earliest portion of the Irvingtonian (the interval
between 2.0 and 1.6 Ma) is not Pleistocene as has been widely used, but rather
latest Pliocene.
The Pleistocene Epoch is subdivided into the early, middle, and late
Pleistocene (Fig. 2). The early Pleistocene begins at 1.64 Ma and ends at the
boundary between the Matuyama and Brunhes chrons at 0.78 Ma. The middle
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
NALMA GEOM,
OA K^Un
NETIC POLARITY SOUTH FLORIDA
IE SCALE LITHOSTRATIGRAPHY
Figure 2. Chart depicting chronologic correlation between units of the geologic time scale (after Harland et
al. 1990), North American Land Mammal Ages, the geomagnetic polarity time scale (after Valet and
Meynadier 1993), and southwestern Florida stratigraphic formations.
Pleistocene covers the time period from the beginning of the Brunhes Chron at
0.78 Ma until the onset of the last (Sangamonian) interglacial at 132 ka. The late
Pleistocene extends from about 132 ka to the Pleistocene/Holocene boundary at 10
ka. These subdivisions of the Pleistocene follow Harland et al. (1990), as adjusted
for the up-dated geomagnetic polarity time scale of Valet and Meynadier (1993).
Vertebrate paleontologists do not all agree on the definitions and boundaries
of the late Pliocene and Pleistocene North American Land Mammal Ages
(NALMA), including part of the Blancan and all of the Irvingtonian and
Rancholabrean. The subdivisions of these three NALMA are even less well
understood. The two most recent attempts to define and subdivide the Irvingtonian
and Rancholabrean (Lundelius et al. 1987-based on all mammal groups;
Repenning 1987-based only on arvicoline rodents) are not in complete agreement.
AGE
-7
013 nlr nv- FORT THOMPSON FM.
LABREAN
A? BRUNHES
w
z
I--
07, Z
w 0
ca z
C BERMONT FM.
MATUYAMA
SOLDUVAI CALOOSAHATCHEE FM.
u z
0 <
I" Z PINECREST BEDS
Im
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Therefore, it is important to establish our criteria for defining and subdividing
these NALMA, particularly the Irvingtonian since this is the age in which the
Leisey Shell Pit LF occurs.
Repenning (1980; 1987) proposed a biochronology for the Blancan through
the Rancholabrean NALMA based on evolutionary stages and immigration events
of arvicoline rodents, more often termed "microtine" rodents in the extensive
literature on the fossil history of this group. Our use of the term "arvicoline" in
this paper is synonymous with the use of microtine by Repenning (1987) and other
authors. Among the 74 Blancan and Irvingtonian arvicoline faunas discussed by
Repenning (1987), only Cumberland Cave in Maryland is located in eastern North
America (i.e., east of the Mississippi River). Repenning and Grady (1988) have
since described an extensive middle Irvingtonian arvicoline fauna from Hamilton
Cave, West Virginia. The published arvicoline fauna from the late Irvingtonian
Coleman 2A LF, Sumter County, Florida (Martin 1974; Frazier 1977) was not
mentioned by Repenning. Arvicolines are principally a temperate group of
rodents, and consequently they have a limited distribution in subtropical regions
such as Florida. With only three species presently occurring in the state, Microtus
pennsylvanicus, Pitymys pinetorum, and Neofiber alleni, Florida has one of the
poorest arvicoline faunas known from anywhere in the continental United States.
Arvicoline rodents are unknown from Florida Blancan vertebrate faunas and are
uncommon in most Pleistocene sites. Nonetheless, arvicolines are now known
from eight Irvingtonian sites in Florida. A mammalian biochronology that uses
many different types of taxa, in addition to arvicoline rodents (e.g., Lundelius et al.
1987), is much more applicable in Florida.
The Blancan NALMA covers the time interval between 4.5 and 2.0 Ma.
Lundelius et al. (1987) divided the Blancan into the early, middle, and late
Blancan, whereas Repenning (1987) recognized five Blancan subdivisions
(Blancan I-V) based on arvicoline rodents. Only the late Blancan of Lundelius et
al. (1987) or Blancan V of Repenning (1987) is discussed here, as early and middle
Blancan (Blancan I-IV) land mammal faunas are unknown from Florida (Morgan
and Ridgway 1987). The beginning of the late Blancan (about 2.5 Ma) was
marked by the formation of the Panamanian Land Bridge and the first abundant
appearance in North America of Neotropical immigrants, as well as the arrival of
several genera of Eurasian immigrants. The late Blancan immigration events from
both South America and the Old World probably were responses to major
continental glaciation in the Northern Hemisphere and correspondingly low sea
levels between about 3.0 and 2.5 Ma (Shackleton and Opdyke 1977).
Typical Blancan genera found in Florida faunas referred to this NALMA
include Borophagus, Trigonictis, Nannippus, Equus (Dolichohippus), and
Rhynchotherium. Neotropical mammalian genera that first appear in Florida
during the late Blancan include the xenarthrans Dasypus, Glyptotherium,
Holmesina, Eremotherium, and Glossotherium and the caviomorph rodents
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 19
Erethizon and Neochoerus, as well as the giant phororhacid bird Titanis. The co-
occurrence or overlapping range zone of Nannippus and these South American
immigrants defines a narrow time interval between 2.5 and 2.0 Ma, after the
formation of the Panamanian isthmus and the beginning of the Great American
Interchange, but before the extinction of Nannippus (Galusha et al. 1984; Lindsay
et al. 1984; Morgan and Ridgway 1987).
The Blancan/Irvingtonian boundary is usually placed near the base of the
Olduvai Subchron (Lundelius et al. 1987; Repenning 1987), that is now dated at
1.95 Ma (Valet and Meynadier 1993). Lundelius et al. (1987) did not precisely
define this boundary on faunal grounds, but described a transitional period between
about 2.0 and 1.6 Ma, during which late Blancan faunas gradually changed into
early Irvingtonian faunas. However, Repenning (1987) noted a major change in
arvicoline rodent faunas at the Blancan/Irvingtonian boundary (which he placed at
1.9 Ma), with the arrival of a number of Old World immigrants at this time, as
well as the first abundant occurrence of taxa with unrooted teeth. The earliest
Irvingtonian marks the first appearance of arvicoline rodents in the Florida fossil
record (Ondatra idahoensis, Atopomys salvelinus). The ages of the late Blancan
Borchers LF (Kansas) and earliest Irvingtonian Curtis Ranch LF (Arizona) are the
primary basis for determining the date of the Blancan/Irvingtonian boundary
(Lundelius et al. 1987:217). Borchers is located in the top of and just above the
Pearlette B ash bed, which has a K-Ar date of 2.01 Ma (Izett 1981). Curtis Ranch
stratigraphically lies just below the base of the Olduvai Subchron in
geomagnetically reversed sediments. Valet and Meynadier (1993) show a date of
about 1.95 Ma for the base of the Olduvai. These dates tightly constrain the
boundary between the Blancan and Irvingtonian to the interval 2.01 to 1.95 Ma, or
approximately 2.0 Ma (Fig. 2). A somewhat younger age for this boundary, 1.9
Ma, was used by Lundelius et al. (1987) and Repenning (1987), because the then
accepted age for the base of the Olduvai Subchron was 1.88 Ma.
Lundelius et al. (1987) characterized the Irvingtonian by the first appearance
of Equus sensu strict (s.s.), Euceratherium, Mammuthus, Smilodon, and Microtus,
among others. Smilodon is now known to be present in at least four late Blancan
faunas in Florida (Berta 1987). Mammuthus does not reach North America from
the Old World until sometime after 1.6 Ma, and thus its first appearance in the
New World is well above the Blancan/Irvingtonian boundary (Lindsay et al. 1984;
Lundelius et al. 1987). Since the Irvingtonian covers the latest Pliocene and much
of the Pleistocene (between 2.0 and 0.3 Ma), it is useful to subdivide this NALMA
into smaller time units. Lundelius et al. (1987) recognized three subages of the
Irvingtonian: early Irvingtonian (Sappan), middle Irvingtonian (Cudahyan), and
late Irvingtonian (Sheridanian). We use the terms early, middle, and late for these
three subdivisions following the terminology generally applied to other NALMA
(Woodburne 1987). We furthermore find it convenient to recognize four
subdivisions of the Irvingtonian (earliest, late early, middle, and late), at least for
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
our biochronologic analyses of Florida fossil faunas. The boundaries between these
four subdivisions of the Irvingtonian are not well constrained by geochronologic
data in Florida, and thus should be considered approximate and provisional.
The boundary between the earliest and late early Irvingtonian is set at 1.6 Ma
(Fig. 2), to approximate the Plio-Pleistocene boundary. At least some late early
Irvingtonian faunas in Florida come from reversely magnetized sediments that are
correlated to the portion of the Matayuma Chron between the Olduvai and
Jaramillo Subchrons (between 1.79 and 1.07 Ma; Valet and Meynadier 1993).
These faunas also most closely resemble those from the Great Plains (Gilliland,
Holloman, Sappa) that are dated at 1.5 to 1.3 Ma (see discussion below).
The middle Irvingtonian of our usage is the equivalent of the Cudahyan
Subage of Lundelius et al. (1987), and approximates the Irvingtonian II of
Repenning (1987). In the Great Plains, Cudahyan faunas underlie the Pearlette O
ash (0.61 Ma; Izett 1981) and overlie the Pearlette S ash (1.27 Ma; Izett 1981).
The oldest faunas included in the middle Irvingtonian were deposited during the
portion of the Matayuma Chron that postdates the Jaramillo, an interval now dated
at 0.99 to 0.78 Ma (Valet and Meynadier 1993). These include portions of the
Alamosa LF of Colorado (Rogers et al. 1985), and the Irvington LF and Centerville
Beach LF of California (Lindsay et al. 1975; Repenning 1987). We therefore
suggest that the age of the top of the Jaramillo Subchron, ca. 1.0 Ma, be used as the
boundary between the early and middle Irvingtonian. The Cudahy LF of Kansas is
associated with the 0.61 Ma Pearlette O ash, and thus forms a convenient upper
boundary for the middle Irvingtonian (Lundelius et al. 1987).
The late Irvingtonian (or Sheridanian) includes faunas younger than 0.6 Ma,
but which predate the dispersal of Bison into North America, the event that defines
the base of the Rancholabrean. Unfortunately, this dispersal event is poorly
constrained, and estimated to lie between 0.5 and 0.2 Ma (Lundelius et al. 1987).
We therefore use the approximate midpoint of this range, 0.3 Ma, for the
Irvingtonian/Rancholabrean boundary, with the expectation that future work will
more precisely document the arrival of Bison in North America. The 0.3 Ma date
also corresponds to the beginning of the late middle Pleistocene, and the onset of a
major period of continental glaciation (Richmond and Fullerton 1986).
GEOLOGY AND GEOCHRONOLOGY OF THE LEISEY SHELL PITS
Geologic Descriptions of Leisey Strata.-Sediments exposed by mining
operations at the Leisey Shell Pit are tentatively referred to four formations, each of
which has produced vertebrate fossils, as well as a surficial layer of unconsolidated
sand (Fig. 3). In this section we first describe the local geology and stratigraphy of
the three Leisey Shell pits, particularly Leisey 1 and 3, emphasizing the bone-rich
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
COMPOSITE SECTION
LEISEY SHELL PIT I
HILLSBOROUGH CO., FLORIDA
0.4 m, modem soil zone W
2.4 m, 0 E
unconsolidated
quartz sand
UPPER SHELL BED
,.
E
3.0 m, buff, sandy, E
massive shell bed o
W
0.4 m, shelly,dolom., "hard layer" W
0.1im, shall, muddy, BONE BED 0
a.
0 g
LOWER SHELL BED
o
4.3 m, bluish,
sandy,
masalve shell bed
dark, sandy
-mud -
tan, hard, massive, I
phosphatic dolomite 8 -
<
COMPOSITE SECTION
LEISEY SHELL PIT 3
HILLSBOROUGH CO., FLORIDA
depth below
ground surface
(m)
-r-0
0.4 m, modem soil zone (removed)
o E
1.3 m, unconsolidated Z ,
quartz sand .J
I.
UPPER SHELL BED E
1.5 m, buff to cream,
sandy, masalve shell o
bed -
02 In ll sa nd BON BED
W
Z
LU
LOWER SHELL BED
W ,
3.3 m, cream to bluish, j
sandy, massive shell bed o
1.0 m, dark, sandy mud;
freshwater shells; bones
modern mean sea level
Figure. 3. Stratigraphic sections of the Leisey Shell Pit IA and 3A sites, Hillsborough County, Florida,
measured by the authors in 1984 and 1986. See text for detailed description of the sections.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
lenses. We then summarize the geochronology of the Leisey strata based on
vertebrate biochronology and data from other papers in this volume, including
invertebrate biostratigraphy, paleomagnetics, and strontium isotope stratigraphy.
At the base of the stratigraphic section exposed in the Leisey Shell pits is a
well-indurated, tan to light gray, clayey, phosphatic dolostone. Identifiable
invertebrate fossils have not been recovered from this unit, but waterworn sirenian
and cetacean fossils are fairly common. The top of this unit is very uneven and
represents a major erosional unconformity. The most biostratigraphically-useful
fossil found in this bed is a well preserved, slightly worn lower molar (ml or m2)
of a merychippine-grade horse (UF 53819). The crown height and enamel pattern
of UF 53819 closely matches those of lower molars recovered from the Arcadia
Formation at Nichols Mine in Polk County, located about 35 km northeast of
Leisey. Hulbert and MacFadden (1991) tentatively referred this sample to
"Merychippus" tertius, a species otherwise known from the late Hemingfordian
(late early Miocene) of Nebraska. However, other mammals in the Nichols Mine
fauna, principally rodents, indicate a slightly younger, very early Barstovian
(earliest middle Miocene) age (Pratt, Morgan, and Poyer in prep.). Scott (1988)
stated that the Arcadia Formation is widespread throughout the subsurface of
southern Florida, including southwestern Hillsborough County, and his description
of its lithology agrees well with that of the basal Leisey stratum. Accordingly, the
lowermost bed at Leisey is referred to the Arcadia Formation of the Hawthorn
Group.
The Bone Valley Formation unconformably overlies the Arcadia Formation
over a wide area in central Florida (Scott 1988). We use the name Bone Valley
Formation in its traditional, broad sense, and as the equivalent of the Bone Valley
Member of the Peace River Formation of Scott (1988). As mapped in the
subsurface by Scott (1988, figs. 36, 53), the Bone Valley Formation rapidly thins in
western Hillsborough and Manatee counties, and is absent in some sections. The
Bone Valley Formation was not observed in outcrop at Leisey, but several land
mammals characteristic of this unit were recovered from a reworked zone on top of
the Arcadia Formation at the north end of Leisey Pit 1 (Fig. 1). This site, called
Leisey IC, contained phosphate pebbles and gravel and spheroidal metaquartzite
pebbles, all typical of the uppermost unit of the Bone Valley Formation (Pirkle et
al. 1967). Vertebrate fossils from Leisey 1C are heavily waterworn and a dull
black color. Their preservation differs considerably from bones recovered from the
overlying shell beds. Both marine and terrestrial taxa are represented (Hulbert and
Morgan 1989). As is typical in low-elevation deposits of the Bone Valley
Formation in southwestern Florida, isolated horse teeth are the most common
terrestrial vertebrate fossils, and the most useful biochronologically.
Three equid species are recognized from Leisey IC on the basis of diagnostic
dental character states: Nannippus aztecus (UF 43564, 107531, 107532; see
Hulbert 1990 for use of N. aztecus instead of N. minor); Neohipparion eurystyle
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 23
(UF 107528); and Cormohipparion ingenuum (UF 107529, 107530). The
chronologic range of.the first two species is late early Hemphillian through very
late Hemphillian (7.0 to 4.5 Ma; Hulbert 1987, 1990). The range of the latter is
early Clarendonian through late early Hemphillian (11.5 to 6 Ma; Hulbert 1988b).
Their concurrent range is thus constrained to the late early Hemphillian (very late
Miocene). Similarly preserved, low-elevation occurrences of fossiliferous Bone
Valley-type deposits are also known from Manatee (Port Manatee, Manatee County
Dam, and Braden River sites) and Sarasota (Lockwood Meadows) counties. Based
on the biochronology of their equid faunas these five Gulf Coastal sites, termed the
Manatee Fauna (Morgan 1994), are correlated with the late early Hemphillian
(approximately 7 Ma) Withlacoochee 4A and Moss Acres local faunas from
Marion County in northern Florida (Hulbert 1988b), and were probably deposited
just prior to the latest Miocene (Messinian) eustatic drop in sea level that occurred
between 6.7 and 5.2 Ma (Webb and Tessman 1968; Haq et al. 1987; Morgan
1994). Leisey IC represents the northernmost occurrence of these low elevation,
Bone Valley-type deposits along the Florida Gulf Coast.
A rostral or mandibular fragment of the long-beaked dolphin Pomatodelphis
(UF 142238) and four auditory bullae of small cetotheriid mysticetes (UF 88614,
107524, 142240, 142246) from Leisey 1 closely resemble specimens from middle
and late Miocene sites elsewhere in peninsular Florida, in particular the lower unit
of the Bone Valley Formation of Polk and adjacent counties (Morgan 1994).
Abundant large, dense sirenian ribs (UF 142241-142243) from Leisey 1 probably
pertain to the dugongid Metaxytherium, although the ribs are not specifically
diagnostic. The Leisey marine mammal fossils were found near the base of the
section in the Leisey 1 Pit, but were collected out of stratigraphic context and thus
could conceivably belong to either the Barstovian or Hemphillian faunas. The
Metaxytherium-Pomatodelphis-cetothere assemblage (Morgan 1994) characterized
Florida marine mammal faunas from the late early Miocene (late Hemingfordian)
through the late Miocene (late early Hemphillian).
An approximately 8 m thick sequence of massive marine shell beds rests upon
the eroded surface of the Arcadia Formation at Leisey (Fig. 3). Reworked pebbles
and cobbles of dolostone and phosphate from the underlying Arcadia Formation
are common in the base of the shell bed sequence. The shell beds consist primarily
of well preserved mollusc shells in an unconsolidated matrix of fine quartz sand.
In Leisey Shell Pit 1 there are two major superposed shell beds separated by an
erosional unconformity, and locally by a thin layer of indurated freshwater
limestone. These two units are informally designated the lower and upper shell
beds, respectively (Fig. 3). The upper and lower shell beds are similar
lithologically, but differ in the composition of their molluscan faunas (Portell et al.
1992) and by the presence of dark organic-rich silt and clay lenses in the lower
shell bed. The size, stratigraphic position, and faunal composition of these lenses
varies throughout the Leisey pits. All of the major vertebrate concentrations
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
(Leisey 1A, 1B, 3A, and 3B) originate from these organic lenses. One common
type of organic unit encountered in the Leisey pits is composed of a mixture of dark
organic clay and silt, fine to medium-grained sand, well preserved freshwater
mollusc shells (e.g. the gastropods Viviparus and Planorbella and unionid
bivalves), and locally abundant fossil wood. Other organic lenses consist primarily
of shells of the estuarine bivalves Rangia or Crassostrea. The most extensive of
these organic units, Leisey 3B, is not actually a lens but consists of a layer of very
dark-colored clay and silty clay from 20 cm to over 1 m in thickness occurring
widely throughout Leisey Pit 3. Vertebrate fossils are sometimes locally abundant
in Leisey 3B, but were never concentrated to the extent seen in the two main "bone
beds" (Leisey 1A and 3A).
Leisey 1A, the largest and most thoroughly studied organic unit in the lower
shell bed, occurs higher in the section than most of these lenses (approximately 4.5
m above the top of the Arcadia Formation, Fig. 3), and differs from them in
containing predominantly marine molluscs. Leisey lA varies in thickness from 5
to 30 cm over an area of about 2000 m2. It consists of an unconsolidated, poorly
sorted mixture of well preserved mollusc shells, vertebrate bones and teeth,
fossilized mangrove root casts, fine-grained quartz sand, silt, and dark brown mud.
The deposit thins gradually towards its margins, with vertebrate remains becoming
increasingly scarce. Leisey IA is often referred to as a "bone bed" because of the
tens of thousands of vertebrate fossils it contained, although volumetrically, marine
mollusc shells comprised most of the deposit. The taxonomic composition of the
Leisey 1A marine molluscs is generally similar to that of the main lower shell bed
(Portell et al. 1992). The freshwater molluscs of Leisey 1A, in contrast to those
found in the lower organic lenses, often show signs of transport damage. The
mangrove roots probably were not contemporaneous with the deposition of the
Leisey 1A shells and vertebrate fossils, but more likely grew down through the
sediments at a later date. This explains the absence of mangrove pollen in the
Leisey 1A sediments (Rich and Newsom this volume).
Immediately above the Leisey 1A bone bed, and separated from it by a poorly
defined unconformity, is a 30 to 50 cm thick layer of fossiliferous, indurated,
calcareous freshwater limestone. This layer extends beyond the boundaries of
Leisey lA, locally separating the lower and upper shell beds. The limestone
contains great numbers of freshwater gastropod shells (especially Planorbella
scalaris), some poorly preserved and apparently reworked marine mollusc shells,
fine-grained quartz sand, and rare bones. The vertebrate fossils pertain almost
exclusively to freshwater taxa such as emydid and trionychid turtles, alligators, and
water birds. The fauna suggests a freshwater origin for the limestone. Such
freshwater "marls" are commonly interbedded with marine strata in the Plio-
Pleistocene stratigraphic sequence in southern Florida (e.g. DuBar 1958, 1962).
An erosional unconformity above the indurated calcareous marl separates it
from the upper marine shell bed. Reworked fragments of the marl are present in
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 25
the base of the upper shell bed. Like the lower shell bed and bone bed, the upper
shell bed lacks distinct bedding and is composed predominantly of marine mollusc
shells. However, in contrast to the bone bed, the upper shell bed has few
freshwater gastropods, is nearly devoid of bones, lacks fossilized mangrove roots,
and has a higher percentage of fine-grained sand and a lower percentage of silt and
clay. The only notable vertebrate fossils from the upper shell bed in the Leisey 1
pit are a partial plastron of the loggerhead sea turtle, Caretta, and a tooth and
vertebra of Bison. Overlying the upper shell bed is a 2.4 m thick layer of massive
unconsolidated Quaternary quartz sand containing no fossils.
Leisey 1B is a small vertebrate site located several hundred meters south of
Leisey 1A and about 3 m lower in the stratigraphic section. The bones occurred in
a thin lens at the base of the lower shell bed not far above the contact with the
underlying Arcadia Formation. Some of the vertebrate fossils were collected from
a rubble zone lying directly on top of the eroded surface of the Arcadia Formation.
Based on the limited vertebrate assemblage present, the fauna from Leisey 1B is
very similar to that of Leisey 1A. The most diagnostic mammals present in Leisey
1B are the glyptodont Glyptotherium arizonae, the giant armadillo Holmesina
floridanus, the giant tapir Tapirus haysii, and the primitive mammoth Mammuthus
hayi, all indicative of a late early Irvingtonian (early Pleistocene) age.
Leisey 3A occupied about one fourth the area of Leisey lA. It too is located
stratigraphically high within the lower shell bed (Fig. 3). It differs from Leisey IA
in containing a very limited large terrestrial vertebrate fauna overwhelmingly
dominated by juvenile individuals of the llama, Hemiauchenia macrocephala,
many of which consist of associated skeletons. Leisey 3A has a considerably richer
and more diverse terrestrial and freshwater microvertebrate fauna than Leisey 1A.
The most common taxa in the molluscan fauna are the brackish water bivalve
Rangia and the freshwater gastropod Viviparus. No indurated freshwater marl
layer separates Leisey 3A from the overlying shell bed, nor are mangrove roots
present as in Leisey 1A.
Regional Stratigraphic Correlation.--The Pliocene and Pleistocene
stratigraphy of southern Florida is currently undergoing intensive study by a large
number of geologists and paleontologists (e.g. Lyons 1991; DuBar et al. 1991;
Scott and Allmon [eds.] 1992). Clearly, there will be many changes in the
nomenclature of the geologic units recognized in this region (e.g. Waldrop and
Wilson 1990; DuBar at al. 1991; Scott 1992). It is not our intention here to revise
the stratigraphic nomenclature of South Florida, since we are not stratigraphers,
nor do we study molluscs, the primary fossil group upon which the southern
Florida biostratigraphic sequence is based. Our goal is to place the stratigraphic
section exposed in the Leisey Shell Pit into the currently recognized stratigraphic
framework for the region to insure that our work will be comprehensible to present
and future workers. Lyons (1991) has recently summarized the biostratigraphic
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
basis for recognizing the various formational units within the Plio-Pleistocene
shell beds in southern Florida. We follow his subdivision and characterization of
these geologic units.
A century of geologic and paleontologic investigation has resulted in a
generally agreed upon stratigraphic sequence for the extensive Pliocene and
Pleistocene shell beds of southern Florida (e.g. DuBar 1974; Lyons 1991), although
see Scott (1992) for an alternative view. On the basis of stratigraphic superposition
there is little disagreement that the relative ages of the units in question are (from
oldest to youngest): Pinecrest Beds (also known as the Pinecrest Sand Member of
the Tamiami Formation of Hunter 1968; the Fruitville Formation of Waldrop and
Wilson 1990; and the Myakka Member of the Sarasota Formation of DuBar et al.
1991), Caloosahatchee Formation, Bermont Formation, Fort Thompson Formation,
and Coffee Mill Hammock Formation (Fig. 2). These strata are often similar in
their lithological composition, generally consisting of abundant shells of marine
molluscs in a matrix of unconsolidated fine quartz sand, but also including beds of
calcareous marl, indurated freshwater limestone, and dark organic-rich silts and
clays. These lithologies reflect a diversity of nearshore marine, estuarine, and
freshwater depositional environments.
The southern Florida formations composed predominantly of marine shell
beds are primarily differentiated on the basis of the composition of their molluscan
faunas. As suggested by many previous workers, these formations are perhaps
more properly regarded as biostratigraphic units (biozones) because the North
American Stratigraphic Code (1983) requires that formal lithostratigraphic units
be differentiated on the basis of lithology, not fossil content. In the absence of a
formally proposed, well documented biostratigraphic nomenclature for Plio-
Pleistocene units in southern Florida, such as that proposed for the middle Atlantic
Coastal Plain by Blackwelder (1981a), we continue to follow current geological
usage in discussing these units (e.g. Lyons 1991).
The presence of stratigraphically restricted species of molluscs has until
recently been the primary method used to correlate geologic units in southern
Florida. The percentage of extinct species of molluscs in a fauna, termed the
"Lyellian percentage," is another method that has been used to determine relative
age among Florida shell beds. Portell et al. (1992, this volume) identified more
than 200 species of molluscs from the lower shell bed and bone bed from Leisey
Shell Pit 1, approximately 3% of which are extinct. Previously published estimates
of 10-12% extinct species of molluscs from Leisey (Hulbert and Morgan 1989;
Webb et al. 1989) were based upon a preliminary analysis of the molluscan fauna
and are now known to be too high. The Lyellian percentage of extinct species of
molluscs from Leisey is intermediate between faunas of the Bermont and Ft.
Thompson formations (DuBar 1974; Lyons 1991). Bermont molluscan faunas
typically contain from 10-20% extinct species (Hoerle 1970; DuBar 1974; Stanley
1986), compared to those from the younger Fort Thompson Formation which are
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 27
composed of fewer than 5% extinct taxa. The Caloosahatchee Formation, which
directly underlies the Bermont Formation, contains from 50 to 65% extinct species
of molluscs (Olsson and Harbisson 1953; Stanley 1986; Lyons 1991).
DuBar (1958) suggested that the Caloosahatchee Formation was late
Pleistocene in age based primarily on evidence from vertebrate fossils. He later
regarded the Caloosahatchee to be early and middle Pleistocene (1974), whereas
DuBar et al. (1991) placed the Caloosahatchee in the early Pleistocene. We have
re-examined the small sample of mammalian fossils (Holmesina, Eremotherium,
advanced Equus, and Palaeolama) reported from the Caloosahatchee Formation by
DuBar (1958; 1974). These taxa indicate a latest Pliocene or Pleistocene age (i.e.
Irvingtonian or Rancholabrean) but, with the possible exception of the Holmesina,
are not sufficiently diagnostic to provide a more specific indication of the age. The
size of several Holmesina osteoderms collected by DuBar from the Caloosahatchee
Formation is suggestive of a latest Pliocene or early Pleistocene (early
Irvingtonian) age. Within the past five years, an extensive vertebrate fauna has
been collected from the Caloosahatchee Formation in the De Soto Shell Pit in De
Soto County. Preliminary analysis of the De Soto Shell Pit vertebrate fauna (see
discussion below and Table 2) suggests correlation with the well known latest
Pliocene (earliest Irvingtonian) Inglis 1A LF of northern Florida. Lyons (1991)
also considered the Caloosahatchee Formation to be very late Pliocene in age on
the basis of its molluscan fauna.
DuBar (1974) originally proposed the Bermont Formation as an informal
unit, but distinguished the Bermont Formation from the underlying Caloosahatchee
Formation primarily by the absence of most typical Caloosahatchee molluscs, and
by the presence of a few species that are restricted to the Bermont. DuBar (1974)
specifically noted that these two units were similar lithologically. Among the
species of molluscs unique to the Bermont formation (McGinty 1970; DuBar 1974;
Lyons 1991), only the gastropod Strombus mayacensis has been identified from
Leisey (Portell et al. 1992). The lower shell bed at Leisey (including vertebrate
sites Leisey IA and 3A) is tentatively referred to the Bermont Formation on the
basis of compatible lithology, faunal similarity, and the presence of seven extinct
invertebrate taxa (see Portell et al. this volume). In his original description of the
Bermont Formation, DuBar (1974) noted that it was found in widely scattered
exposures from Charlotte County north to Levy County along the West Coast, and
in the vicinity of Lake Okeechobee on the Atlantic Coast.
The best known Bermont molluscan faunas occur 100 km or more south of
Leisey in Charlotte, Glades, Hendry, and Palm Beach counties. The Belle Glade
Rock Pit in Palm Beach County (Hoerle 1970; McGinty 1970), perhaps the best
known Bermont locality, contains over twice the number of molluscan species as
Leisey of which approximately 15% are extinct (Hoerle 1970; Lyons 1991).
However, the fossils from the Belle Glade Rock Pit were collected entirely from
spoil piles, and thus may contain a mixture of faunas (Lyons 1991). The vast
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
majority of the Leisey molluscs were collected from in-place stratigraphic sections.
The reduced number of extinct species at Leisey suggests that its molluscan fauna
may be somewhat younger than other Bermont faunas, and as such is intermediate
in age between this unit and faunas of the Fort Thompson Formation.
Geochronology of the Leisey Shell Pit Beds.-In this section we review
previous attempts to date the Bermont and Fort Thompson formations, and briefly
summarize paleomagnetic (see MacFadden this volume) and stable strontium
isotope (see Jones et al this volume) data from Leisey. The Leisey Shell Pit
provides an important cross-reference point for biochronologies based on marine
molluscan assemblages (e.g., Blackwelder 1981a) and terrestrial mammals (North
American Land Mammal Ages, e.g., Savage and Russell 1983; Woodburne 1987).
Biostratigraphic data from various microfossil groups present in the Leisey strata
(e.g., foraminifera, ostacodes, calcareous nannoplankton) have not yet been
analyzed.
Prior to the 1960s, beds now referred to the Bermont Formation were usually
considered the uppermost strata of the Caloosahatchee Formation (DuBar 1958;
1962). The Bermont Formation was proposed as an informal name in its original
description (DuBar 1974), but has received widespread use in subsequent
publications (e.g. Lyons 1991; DuBar et al. 1991) and was listed as a valid unit in
Swanson et al. (1981). Strata placed in the Bermont Formation by DuBar (1974)
have variously been referred to as Unit F (DuBar 1962), Unit A (Olsson 1964),
Glades Unit (Hoerle 1970; McGinty 1970), the Belle Glade Member of the Fort
Thompson Formation (Hunter 1968), and the Okeechobee Formation (Scott 1992).
There are two prevailing opinions among invertebrate paleontologists
regarding the age of the Bermont Formation. DuBar (1974), Petuch (1988), and
DuBar et al. (1991) considered the Bermont to be middle Pleistocene, falling
between 0.6 and 0.3 Ma. Although Blackwelder (1981b) placed the Bermont in
the late Pleistocene, he did not recognize a three-part subdivision of the
Pleistocene. His proposed age of approximately 0.5-0.4 Ma for the Bermont
Formation would place this unit in the middle Pleistocene as recognized here.
Evidence for a middle Pleistocene age comes primarily from the results of amino
acid racemization studies (Mitterer 1975), from the stratigraphic position of the
Bermont between the supposedly early Pleistocene Caloosahatchee and late
Pleistocene Fort Thompson (DuBar et al. 1991), and indirectly through correlation
with the Canepatch Formation (Oaks and Dubar 1974; Blackwelder 1981a,b;
DuBar et al. 1991). Other invertebrate paleontologists (e.g. Hoerle 1970) have
regarded the Bermont as either late Pliocene or early Pleistocene, citing little
evidence beyond the Lyellian percentage of 10 to 25% extinct species of molluscs.
Bender (1973) obtained dates ranging from 1.9-1.75 Ma for the Caloosahatchee
Formation using the He/U method on corals, which, if accurate, would constrain
the overlying Bermont Formation as younger than 1.7 Ma.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 29
Oaks and DuBar (1974) correlated the Bermont Formation with the
Canepatch Formation of North and South Carolina. The Canepatch is a relatively
securely dated deposit, assigned a middle Pleistocene age on the basis of a
uranium-series date of 460100 ka (Szabo 1985), normal geomagnetic polarity,
and ostracode biochronology (Cronin 1980). Blackwelder (1981a,b) placed the age
of the Canepatch Formation between 0.5 and 0.4 Ma, assigning it to the Myrtlean
Substage of the Longian Stage. Longian Stage molluscan assemblages contain an
essentially modern fauna with less than 5% extinct species. Blackwelder (1981a),
citing Oaks and DuBar's (1974) correlation of the Canepatch and Bermont, also
placed the Bermont in his Myrtlean Substage. Despite these previous age
estimates, several lines of evidence indicate that the Bermont is substantially older
than the Canepatch.
Geochronological data derived from magnetic polarity, strontium isotope
stratigraphy, sea level, and vertebrate chronology are very useful in helping to
determine the age of the Bermont Formation at the Leisey Shell Pit. Samples for
paleomagnetic study were taken through a 5 m section at Leisey, including samples
bracketing the vertebrate-bearing lenses Leisey 1A, 3A, and 3B. All sediments
from both the Leisey 1A and Leisey 3A sites are of reversed magnetic polarity and
lie within the Matuyama Chron (Webb et al. 1989; MacFadden this volume). The
boundary between the Matuyama Chron of generally reversed polarity and the
Brunhes Chron of normal polarity is 0.78 Ma (Cande and Kent 1992; Valet and
Meynadier 1993). The Canepatch Formation of the Carolinas is geomagnetically
normal (Cronin 1980), and thus is not synchronous with the reversed sediments
from Leisey.
Leisey could correlate with the latest Matuyama, between 0.99 and 0.78 Ma,
but other evidence argues against this hypothesis. Most of the uppermost zone of
the Matuyama Chron corresponds with two periods of continental glaciation
(glaciations F and G of Richmond and Fullerton 1986). However, the two Leisey
sites were deposited very close to current sea level and thus would not correlate
with a glacial interval characterized by significantly lowered sea levels.
Furthermore, the marine molluscan fauna from Leisey does not contain evidence of
cooler water forms (Portell et al. 1992). The interval between 1.55 and 1.0 Ma was
a warm period lacking major glacial episodes (Richmond and Fullerton 1986).
Ratios of stable strontium isotopes from biogenic carbonate (shells of the marine
bivalve Chione cancellata) from the Bermont Formation at Leisey correspond to
ages of 2.08 Ma for Leisey 1A and 1.33 Ma for Leisey 3A (Jones et al this volume).
When the 95% confidence intervals for the Leisey strontium isotope values (0.56
Ma) are taken into consideration, Leisey 1A could be as young as 1.52 Ma and
Leisey 3A could be as old as 1.89 Ma, yielding a possible overlap of almost 0.4 Ma
between the two sites. Vertebrate biochronology, as discussed in detail below,
favors a late early Irvingtonian age (1.6-1.0 Ma) for the Bermont at Leisey.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
In summary, both mammalian biochronology and geochronological data
indicate that the Leisey Shell Pit LF is early Pleistocene in age, about 1.6 to 1.0
Ma. Faunal criteria strongly suggest that Leisey is younger than latest Pliocene
faunas such as Inglis lA and De Soto Shell Pit (i.e., younger than 1.6 Ma) and
older than the late middle Irvingtonian McLeod Limerock Mine LF (older than 0.7
Ma). Biochronological data restrict the age of Leisey to the latter half of the early
Irvingtonian or the first half of the middle Irvingtonian. Of these two possibilities
the mammalian fauna is more suggestive of an early Irvingtonian age.
Geochronological data also rule out an early middle Irvingtonian age (between
1.07 and 0.78 Ma). The similarity of Leisey to certain middle Irvingtonian
vertebrate faunas suggests that Leisey falls late in the early Irvingtonian, and
accordingly may be no more than several hundred thousand years older than
Cumberland Cave, Port Kennedy Cave, Hamilton Cave, and other early middle
Irvingtonian faunas. The best estimate for the age of the Bermont Formation at the
Leisey Shell Pit, taking into account all chronologic indicators, is between 1.55 and
1.1 Ma. Moreover, it is most likely that the Leisey 1A and 3A vertebrate sites are
less than 1.3 Ma.
In contrast to the Bermont Formation, there has been relatively little
disagreement concerning the age of the Fort Thompson Formation. Most evidence
points to a middle or late Pleistocene age. In particular, the Coffee Mill Hammock
Member of the Fort Thompson Formation or Coffee Mill Hammock Formation of
some authors is rather securely correlated with the last major interglacial
(Sangamonian) between 130-120 ka (DuBar 1974; Mitterer 1975; Lyons 1991).
This was the last time sea level rose substantially higher than present levels
(Bloom 1983). DuBar (1958:135) listed 13 vertebrate taxa collected in situ in the
Fort Thompson Formation. Contrary to his claims, none of these taxa are strictly
limited to the late Pleistocene (Rancholabrean) as currently understood (Kurt6n
and Anderson 1980; Lundelius et al 1987; Repenning 1987). A late Irvingtonian
age is also possible on faunal grounds alone.
Several specimens of Bison were collected in situ from the upper shell bed at
Leisey Pit 1, which has been referred to the Fort Thompson Formation based on the
lack of extinct species in the molluscan fauna. The occurrence of Bison essentially
defines the Rancholabrean Land Mammal Age (Lundelius et al. 1987). Holmesina
septentrionalis, Sigmodon hispidus, and Tapirus veroensis all were collected from
spoil in Leisey Pit 2, but are presumed to have been derived from the upper shell
bed, as they were not recovered from the lower shell bed at Leisey despite intensive
sampling. Among these three species, Sigmodon hispidus is characteristic of late
Rancholabrean faunas in Florida, whereas the latter two species occur in both late
Irvingtonian and Rancholabrean faunas (<0.6 Ma). However, the overlapping
range zone for these three species plus Bison, occurs only during the
Rancholabrean (0.3-0.12 Ma). Vertebrate fossils from Leisey support previous
determinations that at least the upper units of the Fort Thompson Formation are
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 31
late middle or late Pleistocene in age.
There is a significant gap, corresponding to most of the middle Pleistocene,
between our proposed upper limit for the Bermont beds at the Leisey Shell Pit of
about 1.0 Ma and strata of the Fort Thompson Formation containing Bison that are
0.3 Ma or younger. There are presently no middle or late Irvingtonian vertebrate
faunas known from Florida that occur in stratigraphic superposition with marine
geologic units.
VERTEBRATE BIOCHRONOLOGY OF THE
LEISEY SHELL PIT LOCAL FAUNA
Plio-Pleistocene vertebrate biochronology in Florida.-Table 2 shows the
chronological distribution of 128 species of mammals in Florida during the late
Pliocene and Pleistocene. Five Blancan and 11 Irvingtonian vertebrate faunas are
listed separately in Table 2, whereas the early and late Rancholabrean are
presented as composite faunas derived from a number of different sites. This table
does not include all of the Blancan and Irvingtonian sites known from Florida,
only those with the largest and most chronologically significant faunas, arbitrarily
defined as faunas containing 10 or more species of mammals. Several other
important Florida Blancan and Irvingtonian sites that contain age-diagnostic
mammals are discussed in the text. We caution that Table 2 does not provide
complete mammalian faunal lists for the individual sites, but includes only those
species we consider to be biochronologically diagnostic in Florida. For example,
Insectivora, Chiroptera (with the exception of Desmodus), and many species of
Rodentia are not included in Table 2, although these groups are often common
members of the faunas listed.
We emphasize that the data presented here are based primarily on Florida
faunas and therefore, in the case of certain taxa, may not have widespread
applicability to other North American vertebrate faunas of similar age. Examples
include the late Blancan arrival in Florida of the armadillos Dasypus and
Holmesina and the giant ground sloth Eremotherium, all of South American
origin. These three genera have not been reported elsewhere in North America
prior to the Irvingtonian. Another Neotropical immigrant, the huge flightless bird
Titanis walleri, is unknown outside of Florida, yet its chronological range is
restricted to the late Blancan and earliest Irvingtonian, and it is thus an excellent
biostratigraphic indicator within the state. Whenever possible we attempt to
correlate Florida vertebrate faunas with faunas elsewhere in North America,
especially in instances where the non-Florida faunas have been radiometrically
dated or can be correlated to the magnetic polarity time scale.
32 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 2. Biochronological distribution of Blancan, Irvingtonian, and Rancholabrean mammals in Florida.
An "X" indicates a definite identification, "?" a possible identification, and "-" an absence.
Ear-
Late liest
Blan- Irving-
can tonian
(2.5- (2.0-
2.0 Ma) 1.6 Ma)
Santa Macas- Kis- De
Fe phalt sim- Soto
River Haile Shell mee Haile Inglis Shell
1 15A Pit River 7C 1A Pit
Didelphis virginiana
Dasypus bellus X X X X X X X
Holmesinafloridanus X X X X X X X
Holmesina septentrionalis -
Glyptotherium arizonae X X X X X
Glyptotheriumfloridanum -
Pachyarmatherium leiseyi X -
Glossotherium
chapadmalense X X X X -
Paramylodon harlani X -
Megalonyxjeffersonii -
Megalonyx leptostomus X X X X X X
Megalonyx wheatleyi -
Eremotherium mirabile -
Eremotherium n. sp. 7 X X X
Nothrotheriops texanus -
Desmodus archaeodaptes X
Desmodus stock -
Borophagus diversidens X -
Canis armbrusteri -
Canis dirus -
Canis edwardii X X
Canis latrans -
Canis lepophagus X -
Urocyon cinereoargenteus -
Urocyon minicephalus -
Urocyon n. sp. X -
Procyon lotor -
Procyon n. sp. 7 7 ? X X
Arctodus pristinus X X X -
Tremarctosfloridanus -
Conepatus leuconotus -
Conepatus robustus -
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
Table 2 Extended.
Late Early Late
Early Middle Late Ranch- Ranch-
Irving- Irving- Irving- ola- ola-
tonian tonian tonian brean1 brean2 Recent
(1.6- (1.0- (0.6- (0.3Ma- (130-
1.0 Ma) 0.6 Ma) 0.3 Ma) 130 ka) 10 ka)
Lei- Rig-
sey Payne Crys- by Cole-
Haile Shell Pool Creek tal Shell Haile Mc- man
16A Pit Branch Mine River Pit 21A Leod 2A
34 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 2 Continued.
Ear-
Late liest
Blan- Irving-
can tonian
(2.5- (2.0-
2.0 Ma) 1.6 Ma)
Santa Macas- Kis- De
Fe phalt sim- Soto
River Haile Shell mee Haile Inglis Shell
1 15A Pit River 7C IA Pit
Spilogale putorius -
Spilogale sp. X -
Lutra canadensis -
Satherium piscinarium X X
Trigonictis cookii -
Trigonictis macrodon X X X
Callophoca abscura X X -
Monachus tropicalis X
Dinobastis serus -
Felis amnicola -
Homotherium n. sp. ? ? X -
Lynx rufus -
Lynx sp. X X -
Miracinonyx inexpectatus X X -
Miracinonyx cf. M. trumani -
Panthera atrox -
Panthera onca -
Puma concolor -
Smilodonfatalis -
Smilodon gracilis X X X -
Chasmaporthetes ossifragus X X X
Castor canadensis X -
Castoroides leiseyorum -
Castoroides ohioensis -
Geomyspinetis -
Geomyspropinetis X X X X
Thomomys orientalis -
Zapus sp. -
Erethizon dorsatum -
Erethizon kleini -
Erethizon sp. X -
Hydrochaeris holmesi X X -
Neochoerus dichroplax X X -
Neochoerus pinckneyi -
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
Table 2 Extended/Continued.
Late Early Late
Early Middle Late Ranch- Ranch-
Irving- Irving- Irving- ola- ola-
tonian tonian tonian brean brean Recent
(1.6- (1.0- (0.6- (0.3Ma- (130-
1.0 Ma) 0.6 Ma) 0.3 Ma) 130 ka) 10 ka)
Lei- Rig-
sey Payne Crys- by Cole-
Haile Shell Pool Creek tal Shell Haile Mc- man
16A Pit Branch Mine River Pit 21A Leod 2A
36 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 2 Continued.
Ear-
Late list
Blan- Irving-
can tonian
(2.5- (2.0-
2.0 Ma) 1.6 Ma)
Santa Macas- Kis- De
Fe phalt sim- Soto
River Haile Shell mee Haile Inglis Shell
1 15A Pit River 7C 1A Pit
Neochoerus sp.
Neotomafloridana
Neotoma sp.
Oryzomys palustris
Peromyscus large sp.
Podomysfloridanus
Podomys n. sp.
Sigmodon bakeri
Sigmodon curtisi
Sigmodon hispidus
Sigmodon libitinus
Sigmodon medius
Sigmodon minor
Atopomys salvelinus
Microtus pennsylvanicus
Neofiber alleni
Neofiber leonardi
Ondatra annectens
Ondatra idahoensis
Ondatra zibethicus
Pitymys aratai
Pitymys hibbardi
Pitymys pinetorum
Pedomys n. sp.
Synaptomys australis
Synaptomys n. sp.
Lepus cf L. townsendii
Lepus sp.
Sylvilagus floridanus
Sylvilagus palustris
Sylvilagus webbi
Mylohyusfloridanus
Mylohyusfossilis
Platygonus bicalcaratus
- x
- x x -
- ? X ? ?
x x x -
x x x -
-
- -
- -
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
Table 2 Extended/Continued.
Late Early Late
Early Middle Late Ranch- Ranch-
Irving- Irving- Irving- ola- ola-
tonian tonian tonian brean brean Recent
(1.6- (1.0- (0.6- (0.3Ma- (130-
1.0 Ma) 0.6 Ma) 0.3 Ma) 130 ka) 10 ka)
Lei- Rig-
sey Payne Crys- by Cole-
Haile Shell Pool Creek tal Shell Haile Mc- man
16A Pit Branch Mine River Pit 21A Leod 2A
38 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 2 Continued.
Ear-
Late list
Blan- Irving-
can tonian
(2.5- (2.0-
2.0 Ma) 1.6 Ma)
Santa Macas- Kis- De
Fe phalt sim- Soto
River Haile Shell mee Haile Inglis Shell
1 15A Pit River 7C 1A Pit
Platygonus compressus -
Platygonus cumberlandensis -
Platygonus vetus X
Hemiauchenia blancoensis X X -
Hemiauchenia macrocephala X X X
Hemiauchenia n. sp. X X
Palaeolama mirifica -
Capromeryx arizonensis X X X X
Odocoileus virginianus X X X X X X X
Bison antiquus -
Bison latifrons -
Bovidae, gen. indet.5 X X -
Tapirus haysii -
Tapirus veroensis -
Tapirus n. sp. ? ? ? X X X
Cormohipparion emsliei X X -
Nannippuspeninsulatus X X X X -
Equus (Dolichohippus) sp. X X X X -
Equus "fraternus" -
Equus "leidyi" X X
Equus (Hemionus) n. sp. -
Equus alaskae group -
Equus laurentius group -
Rhynchotherium praecursor X X -
Cuvieronius tropics X X X
Mammut americanum X X
Mammuthus hayi -
Mammuthus columbi -
1 The early Rancholabrean fauna is a composite list compiled from the following sites: Bradenton, Daytona Beach, Haile 7A, Haie BA,
ldsaw, Williston 3A, and Williston 3B.
The late Rancholabrean fauna is a composite list compiled from the following sites: Arredondo 2A, Cutler Hammock, Devils Den,
Ichetucknce River, Melboumne, Monkey Jungle Hammock, Reddick I, Seminole Field, and Vero.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA
Table 2 Extended/Continued.
Late Early Late
Early Middle Late Ranch- Ranch-
Irving- Irving- Irving- ola- ola-
tonian tonian tonian brean brean Recent
(1.6- (1.0- (0.6- (0.3Ma- (130-
1.0 Ma) 0.6 Ma) 0.3 Ma) 130 ka) 10 ka)
Lei- Rig-
sey Payne Crys- by Cole-
Haile Shell Pool Creek tal Shell Haile Mc- man
16A Pit Branch Mine River Pit 21A Leod 2A
3 No longer found in Florida, but still survives elsewhere in North America
4 The Caribbean monk seal has gone extinct within the past 40 years.
5 J. McDonald (pes. comm.) thinks this taxon may be related to Old World Pliocene Bison.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
A quote from Lundelius et al. (1987:218) is particularly appropriate to the
difficulties encountered when trying to correlate Florida Pliocene and Pleistocene
faunas: "Latitudinal, regional, and ecological factors lead to faunal provincialism
that together with lack of radiometric and paleomagnetic control makes correlation
of faunas difficult." Because most Florida Blancan and Irvingtonian vertebrate
faunas lack radiometric and paleomagnetic control, we must rely almost entirely on
biochronologic comparisons with well-dated faunas from the western United
States. However, these correlations may be limited by the faunal provincialism
noted by Lundelius et al. (1987). For instance in eastern North America the late
Blancan and early Irvingtonian (2.5 to 1.0 Ma) are represented almost exclusively
by vertebrate faunas from Florida, many of which appear to sample a more tropical
fauna than typical western faunas of this time period. Despite certain peculiarities
in Florida Blancan and Irvingtonian faunas, there are still more similarities than
differences between Florida and western faunas of this age, and these faunal
similarities will be stressed throughout this paper.
Data on sea level are helpful in placing certain Florida vertebrate faunas in
the context of glacial and interglacial stages. There are two general circumstances
in which information derived from vertebrate fossil sites can provide some
indication of sea level. Low sea level faunas, presumably deposited during glacial
intervals, are indicated by sites collected below present sea level that contain no
evidence of marine vertebrates. By these criteria, the late Blancan St. Petersburg
Times LF and the early Irvingtonian Inglis 1A and Crystal River Power Plant local
faunas are examples of low sea level faunas. High sea level faunas, presumably
reflecting interglacial conditions, are indicated by sites collected well above
modern sea level that contain marine vertebrates. Examples of high sea level
faunas are the late Blancan Haile 15A and Kissimmee River sites, the early
Irvingtonian De Soto Shell Pit, and the early Rancholabrean Daytona Beach and
Oldsmar local faunas. The number of glacial and interglacial intervals, and hence
the number of eustatic sea level changes throughout the late Pliocene and
Pleistocene, appears to be much greater (Shackleton and Opdyke 1977) than the
four glaciations (Nebraskan, Kansan, Illinoian, and Wisconsinan) traditionally
recognized during the Pleistocene. We use the relative sea level position of several
Florida vertebrate fossil sites, in conjunction with other geochronological data, to
help determine the placement of the site relative to the worldwide sea level curve
(e.g. Haq et al. 1987).
No Florida pre-Rancholabrean vertebrate faunas have been radioisotopically
dated, although attempts are underway to obtain uranium-series dates on corals
collected from nearshore marine shell beds in association with vertebrate faunas.
However, uranium-series dates are not reliable for sites older than about one
million years (Szabo 1985). Bender (1973) determined helium/uranium ages on
samples of corals from several Florida Pliocene and Pleistocene marine geologic
units, including the Pinecrest Beds and Caloosahatchee Formation. This dating
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 41
technique has not been applied recently, nor has it been used to directly date any
vertebrate faunas. Other methods of radioisotopic dating (K/Ar, Ar/Ar, etc.) are
not applicable in Florida sites because the surficial geology of this region is devoid
of basalt flows, volcanic ash beds, or other igneous rocks containing minerals
normally used for dating vertebrate-bearing beds in the western United States and
elsewhere.
Long stratigraphic sequences with a paleomagnetic polarity reversal
chronology are unknown from the Florida Plio-Pleistocene. However,
paleomagnetic signatures (i.e. normal or reversed) have now been obtained from
two Florida vertebrate faunas, including Leisey (Webb et al. 1989; MacFadden this
volume) and the late Blancan Macasphalt (=APAC) Shell Pit (Jones et al. 1991).
Most Blancan and Irvingtonian karst deposits in northern Florida (e.g. Haile 15A,
Haile 16A, McLeod, Coleman 2A) were destroyed by mining operations many
years ago, and thus are no longer available for paleomagnetic sampling. In the
future, we intend to obtain a paleomagnetic signature for all Florida Blancan and
Irvingtonian sites containing sediments suitable to this type of analysis.
An integrated approach has been used to determine the age of the vertebrate-
bearing deposits at the Leisey Shell Pit, applying data from paleomagnetic polarity
stratigraphy, strontium isotope chronology, relative sea level position, and
mammalian and molluscan biochronology (see Jones 1992). Many Florida Plio-
Pleistocene vertebrate sites, particularly deposits in the northern portion of the
peninsula, do not lend themselves to this type of integrated analysis. The majority
of northern Florida Blancan and Irvingtonian sites are composed of terrestrial or
freshwater sediments that occur as isolated cave, sinkhole, or fissure fillings in
Eocene marine limestones, and cannot be correlated to a regional stratigraphic
sequence. Most also lack associated marine vertebrate and invertebrate faunas,
thus eliminating the possibility of strontium isotope analysis, determination of
relative sea level position, or comparison with marine molluscan biochronology.
Therefore, sites such as the Leisey Shell Pit are extremely important cross-
references that allow us to compare terrestrial and marine biochronologies, and to
contrast these results with data obtained from other geochronological techniques.
Biochronology of the Leisey Mammalian Fauna.-The Leisey Shell Pit
Local Fauna has one of the richest samples of large terrestrial mammals known
from any Irvingtonian site in North America. Among the 49 mammalian taxa
listed in Table 1, 33 are large mammals. Large herbivores numerically dominate
the Leisey 1A mammalian fauna, in particular the llamas Palaeolama mirifica and
Hemiauchenia macrocephala, two species of the horse Equus, the ground sloths
Paramylodon harlani and Nothrotheriops texanus, the peccary Platygonus vetus,
and the mammoth Mammuthus hayi. The most common large carnivore is the
sabercat Smilodon gracilis. The small mammal fauna consists of one species of
shrew, two lagomorphs, and nine rodents. Five marine mammals have been
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
identified from Leisey, including three odontocete cetaceans, the manatee
Trichechus, and the seal Monachus.
The age of the Leisey Shell Pit LF has been determined principally from its
land mammal fauna. However, the biochronologic significance of selected non-
mammalian taxa from Leisey will also be discussed. The large and well preserved
samples of carnivores, ungulates, and xenarthrans, and to a lesser extent rodents,
from Leisey permit detailed taxonomic and biochronological comparisons of this
fauna with well known Irvingtonian faunas from Florida and elsewhere in North
America (e.g. Kurten and Anderson 1980; Lundelius et al. 1987; Repenning 1987).
Preliminary biostratigraphic analyses of the Leisey Shell Pit vertebrate fauna have
already been published (Hulbert and Morgan 1989; Webb et al. 1989). These
studies and the present analysis indicate that the Leisey Shell Pit LF correlates to
the later half of the early Irvingtonian between 1.6 and 1.0 Ma, and most likely to
the later half of this interval. This age assignment is based on a number of
mammalian taxa, including first appearances of immigrants from both South
America and the Old World, overlapping range zones of certain indicator species,
the evolutionary stage of taxa within certain well known lineages, and the absence
of most taxa characteristic of either Blancan or middle Irvingtonian and younger
faunas.
In addition to age differences, there are many other reasons why taxa may be
absent from a fauna, including collecting bias, taphonomic or paleocological
factors, and differences in biogeography. The faunal differences between the four
individual sites comprising the Leisey Shell Pit LF are probably related to
paleoecological factors. For instance, at Leisey 1A Mammuthus is the dominant
proboscidean and Nothrotheriops and Paramylodon are the only two ground sloths
present, whereas the gomphothere Cuvieronius and the giant ground sloth
Eremotherium are the most common members of these two groups in the Leisey
Shell Pit 3 site. Other faunal anomalies between the Leisey sites, such as the
presence of Glyptotherium arizonae only at the base of the lower shell bed in
Leisey Pit 1, may reflect slight differences in age between the collecting localities.
Xenarthra.--Like many other Florida Pliocene and Pleistocene sites, the
Leisey Shell Pit LF has a diverse sample of xenarthrans, including three genera of
armadillos, one genus of glyptodont, and four genera of ground sloths. Among
Leisey xenarthrans, only the armadillo Dasypus bellus and the ground sloth
Paramylodon harlani are considered conspecific with Rancholabrean species. D.
bellus first appears in Florida during the late Blancan and persists throughout the
remainder of the Pliocene and Pleistocene, going extinct at the end of the
Rancholabrean. Although D. bellus is the only species of Dasypus recognized in
North America during this time period, this armadillo shows a substantial size
increase during its 2.5 million year history (Robertson 1976). The Leisey sample
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 43
ofD. bellus is intermediate in size between small specimens from the late Blancan
and the more typical large Rancholabrean form (Downing and White this volume).
The taxonomic and evolutionary history of the giant armadillo or pampathere
Holmesina has been reviewed by Edmund (1985, 1987) and Hulbert and Morgan
(1993). These studies documented significant evolutionary change in Holmesina
from its first appearance in North America in the late Blancan until its extinction
in the late Rancholabrean. Hulbert and Morgan (1993) referred the Leisey sample
to the smaller of the two recognized species, H. floridanus, rather than to the much
larger H. septentrionalis, which occurs in the middle Irvingtonian through
Rancholabrean. They divided Florida Holmesina samples into four recognizable
groups that are useful in a biochronologic context: 1) very small individuals of H.
floridanus are known only in late Blancan sites, such as the type locality Haile
15A; 2) a slightly larger form referred to H. floridanus occurs in the latest Blancan
Haile 7C LF; 3) an intermediate-sized form also referred to H. floridanus is known
from the early Irvingtonian; and 4) the very large species H. septentrionalis ranges
from the middle Irvingtonian through the Rancholabrean. Intermediate-sized
specimens of H. floridanus, such as those found at Leisey, help restrict the age of
this fauna to early Irvingtonian (2.0 to 1.0 Ma). The oldest record of Holmesina
outside of Florida is from the early Irvingtonian Gilliland LF in Texas (Hibbard
and Dalquest 1966), an approximate temporal equivalent of Leisey.
One of the most intriguing members of the Leisey mammalian fauna is a new
genus and species of large armadillo, Pachyarmatherium leiseyi, named and
described by Downing and White (this volume). The presence of an undescribed
shelled edentate from several Irvingtonian sites in southern Florida has been
known for some time based on a small sample of osteoderms. This taxon had been
confused with Dasypus bellus until the discovery of an extensive sample of small,
but very thick, hexagonal osteoderms from the Leisey Shell Pit.
Pachyarmatherium leiseyi also appears to be a useful biostratigraphic indicator
within the state, as all records are either late Blancan or early Irvingtonian. The
oldest known specimens of P. leiseyi are from the late Blancan Kissimmee River 6
Site and the earliest Irvingtonian Forsberg Shell Pit LF (late Pliocene). Late early
Irvingtonian faunas such as Leisey and Payne Creek represent the youngest known
occurrences of this species.
Glyptotherium has a rather spotty distribution in Florida, both geographically
and chronologically. It is one of the earliest Neotropical immigrants to participate
in the Great American Interchange, where it is first recorded from late Blancan
faunas in Arizona, Texas, and peninsular Florida (Gillette and Ray 1981). Except
for the Florida samples, Gillette and Ray (1981) referred all other North American
Blancan glyptodonts to the small species G. texanum. They tentatively referred
glyptodont osteoderms from the late Blancan Santa Fe River sites and the earliest
Irvingtonian Inglis 1A LF to G. arizonae. G. arizonae is a large, thick-shelled
species reported from three early Irvingtonian faunas in the western United States:
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Curtis Ranch, Arizona (type locality); Holloman, Oklahoma; and Gilliland, Texas
(Gillette and Ray 1981). Glyptodont fossils collected from late Blancan and early
Irvingtonian sites in Florida (including Leisey) in the ten years since of the
publication of Gillette and Ray's (1981) monograph all appear to be referable to G.
arizonae as well. Glyptotherium is rare in the Leisey Shell Pit LF, and is absent
from the two richest sites, Leisey IA and 3A. G. arizonae is represented at Leisey
by a series of associated osteoderms collected from the base of the lower shell bed
in Pit 1 in association with Mammuthus. Glyptotherium apparently was absent in
Florida during the middle and late Irvingtonian. The smaller, thinner-shelled
species, Glyptotheriumfloridanum, is restricted to Rancholabrean faunas in Florida
(Gillette and Ray 1981).
Four species of ground sloths have been identified from Leisey, an
undescribed species of Eremotherium (De Iuliis and Cartelle in prep.), Megalonyx
wheatleyi, Nothrotheriops texanus, and Paramylodon harlani (McDonald this
volume). The new species of Eremotherium is distinguished from the
Rancholabrean species, E. mirabile, by the presence of four well developed claws
on the manus, as opposed to only two claws in the later species (Hulbert et al.
1989). With the exception of the bones of digits 1 and 2 of the manus
(metacarpals, phalanges, and claws), the remaining elements of the skeleton are
very similar in the two species. The new species of Eremotherium has so far been
confidently identified from the latest Blancan Haile 7C LF and the early
Irvingtonian De Soto Shell Pit, Inglis IA, Haile 16A, Leisey Shell Pit, Crystal
River Power Plant, and Payne Creek Mine local faunas. Other records of
Eremotherium of similar age, but lacking the diagnostic skeletal elements of the
manus, include the late Blancan Kissimmee River, Lehigh Acres, and Brighton
Canal local faunas and the early Irvingtonian Pool Branch and Haile 21A local
faunas. The Eremotherium remains from these five sites are tentatively referred to
the new four-clawed form based on their similarity in age to confirmed records of
this species. Eremotherium has been recorded from the base of the lower shell bed
in Leisey Pit 1 and is fairly common in Leisey Pit 3, but is absent from Leisey IA
and 3A. The largest Florida sample ofE. mirabile is from the early Rancholabrean
Daytona Beach LF (Edmund et al. in prep.). E. mirabile was very rare in Florida
during the late Rancholabrean.
McDonald (1977) reviewed the fossil history of Megalonyx in Florida. He
identified a small, endemic Florida form of the typical Blancan species, Megalonyx
leptostomus, from the late Blancan Santa Fe River 1 LF and the earliest
Irvingtonian Inglis IA LF. Additional specimens of this dwarf M. leptostomus
have now been identified from the late Blancan Macasphalt Shell Pit, Kissimmee
River, St. Petersburg Times, and Haile 7C faunas and the earliest Irvingtonian De
Soto Shell Pit LF. Megalonyx has only recently been recorded from the Leisey
Shell Pit LF based on a nearly complete skull from Leisey 3B. This genus is absent
from Leisey 1A and 3A. The Leisey Megalonyx is distinctly larger than the
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 45
Florida material ofM. leptostomus and is here tentatively referred to M. wheatleyi.
According to McDonald (1977), M. wheatleyi is intermediate in size between the
smaller Blancan and earliest Irvingtonian M. leptostomus and the larger late
Pleistocene species M. jeffersonii. Other Florida localities for M. wheatleyi are the
late early Irvingtonian Haile 16A, Payne Creek Mine, and Crystal River Power
Plant local faunas and the middle Irvingtonian McLeod LF (McDonald 1977). M.
wheatleyi appears to be restricted to the late early and middle Irvingtonian in
Florida, whereas the larger M. jeffersonii does not appear until the Rancholabrean.
Additional records of M. wheatleyi include early Irvingtonian faunas from
Holloman, Oklahoma and Vallecito Creek, California and middle Irvingtonian
faunas from Port Kennedy Cave, Pennsylvania (type locality), Cumberland Cave,
Maryland, and Cudahy, Kansas (McDonald 1977).
Nothrotheriops is unknown in North America prior to the early Irvingtonian,
and thus is apparently one of the last genera of South American immigrants to
participate in the Great American Interchange. Irvingtonian specimens of
Nothrotheriops are now regarded as a separate species, N. texanus, distinguished
by its smaller size and more slender cranium from the better known Rancholabrean
species, N. shastensis (see Akersten and McDonald 1991; McDonald this volume).
The oldest well-dated records of N. texanus are from the early Irvingtonian
Vallecito Creek LF, California, Gilliland LF, Texas, El Golfo LF, Sonora, Mexico,
and Leisey Shell Pit and Pool Branch of Florida (Hibbard and Dalquest 1966;
McDonald 1985, this volume). The Nothrotheriops sample from Leisey IA is the
largest Irvingtonian sample of this genus in North America (McDonald this
volume).
"Glossotherium" chapadmalense is one of the first South American
immigrants to reach North America following the formation of the Panamanian
isthmus. Although the systematic relationships remain unresolved, it appears that
the younger species Paramylodon harlani was derived from "G." chapadmalense
sometime during the late Blancan. These two species almost certainly belong in
the same genus (McDonald this volume). "G." chapadmalense is a small species
represented by a partial skeleton from the late Blancan Haile 15A LF (Robertson
1976). Other late Blancan records of "G." chapadmalense include the Santa Fe 1,
Macasphalt Shell Pit, and Kissimmee River local faunas from Florida (Table 2), as
well as Mt. Blanco, Texas (Dalquest 1975) and 111 Ranch, Arizona (Galusha et al.
1984). The oldest Florida record of the larger species, P. harlani, is from the
earliest Irvingtonian Inglis 1A LF. P. harlani is found in Florida throughout the
Irvingtonian and Rancholabrean. McDonald (this volume) demonstrates a gradual
size increase in this species from the early Irvingtonian through the
Rancholabrean.
Carnivora.--Berta (this volume) identified two species of large canids from
the Leisey Shell Pit LF, Canis edwardii and C. armbrusteri. Both of these canids
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
are restricted to North American Irvingtonian faunas. However, prior to the
discovery of Leisey, C. edwardii was known only from early Irvingtonian faunas,
whereas C. armbrusteri was restricted to middle and late Irvingtonian faunas.
These two species are now known to co-occur in two late early Irvingtonian sites in
Florida, Leisey and Haile 21A. C. edwardii also is known from five other Florida
early Irvingtonian faunas, including the earliest Irvingtonian Inglis 1A, De Soto
Shell Pit and Forsberg Shell Pit local faunas and the late early Irvingtonian Crystal
River Power Plant and Rigby Shell Pit local faunas. C. armbrusteri has also been
identified from the late middle Irvingtonian McLeod Limerock Mine and the late
Irvingtonian Coleman 2A LF (Martin 1974; Berta this volume). Florida records of
C. edwardii and C. armbrusteri suggest that the overlapping range zone of these
two species occurs only during a restricted interval of time in the late early
Irvingtonian.
The sample of the sabercat Smilodon gracilis from Leisey is the most
complete known from any Irvingtonian site (Berta this volume). Although the
Leisey specimens are referable to S. gracilis, Berta (1987; this volume) noted that
certain evolutionary changes occurred within this species during the early and
middle Irvingtonian. S. gracilis is also well represented by material from the
earliest Irvingtonian Inglis 1A LF and the middle Irvingtonian Port Kennedy Cave
(type locality) and McLeod Limerock Mine local faunas (Berta 1987; this volume).
The Leisey S. gracilis is intermediate in size and several morphological features
between the samples from the older Inglis IA LF and the younger Port Kennedy
and McLeod sites (Berta 1987; this volume). There are no late Irvingtonian
records of Smilodon from Florida. The larger and more advanced species, S.
fatalis (=S. populator of Berta 1985), first occurs at this time elsewhere in North
America (Berta 1987; Lundelius et al. 1987). S. fatalis appears in Florida during
the early Rancholabrean and is found there throughout the remainder of the
Pleistocene.
A large and possibly undescribed species of the machairodont cat
Homotherium is represented at Leisey by a few isolated elements (Berta this
volume). A partial skeleton of this species, including the skull and mandibles, is
known from the correlative Haile 21A LF. Other fossils of this large Homotherium
have been identified from Inglis IA, Haile 16A, and possibly the late Blancan
Santa Fe River 1 and Kissimmee River faunas. The large Homotherium present in
Florida late Blancan and early Irvingtonian faunas is distinct from Dinobastis
serus (see Berta this volume), a smaller sabercat known principally from the
Rancholabrean, including the Reddick 1A LF in Marion County (Waldrop 1974).
The fossil record of the North American cheetah-like cat, Miracinonyx
inexpectatus, has recently been reviewed (Van Valkenburgh et al. 1990). They
reported M. inexpectatus from faunas of late Blancan through middle Irvingtonian
age, including several specimens from Inglis 1A. Additional Florida records ofM.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 47
inexpectatus not listed by Van Valkenburgh et al. (1990) are from Leisey (Berta
this volume) and the late Blancan Santa Fe River 2.
The large tremarctine bear, Arctodus pristinus, occurs in two late Blancan
faunas and eight Irvingtonian faunas in Florida, including Leisey 1A (Emslie this
volume). The oldest records ofA. pristinus in Florida are isolated teeth from the
late Blancan Kissimmee River and Santa Fe River 1 faunas. Because this species
is found throughout the Irvingtonian in Florida (Table 2), its presence does not
place a fauna more precisely within this NALMA. There appears to be a
chronological separation of tremarctine bears in Florida; Arctodus pristinus is
found in the late Blancan and Irvingtonian and the smaller Tremarctosfloridanus
is restricted to the Rancholabrean. This may apply only in Florida, as the large
bear Arctodus simus is widely distributed in North America during the
Rancholabrean and T. floridanus has been reported from several western Blancan
and Irvingtonian sites (Kurt6n and Anderson 1980).
The river otter Lutra is an old World immigrant that first arrived in North
America during the Irvingtonian (Kurt6n and Anderson 1980; Lundelius et al.
1987). The earliest previously reported North American records of Lutra are from
the middle Irvingtonian Cumberland Cave and Port Kennedy Cave faunas (Kurt6n
and Anderson 1980). The oldest well documented record of Lutra in Florida, and
possibly North America as well, is from the Leisey Shell Pit LF. A well preserved
mandible from Leisey 3B is similar in size to extant L. canadensis, whereas two
associated upper teeth of Lutra from Leisey 1A are distinctly larger than the
modern species. A large extinct otter, L. parvicuspis, described from Cumberland
Cave (Gidley and Gazin 1933), has been synonymized with L. canadensis by most
recent authors (e.g. Kurt6n and Anderson 1980; Berta this volume). L. canadensis
has also been identified from the Crystal River Power Plant, a close faunal
correlate of Leisey.
A large raccoon of the genus Procyon is present in many Florida late Blancan
and Irvingtonian faunas, including Leisey (Procyon n. sp. in Table 2). Klein
(1971) described the Procyon from Inglis 1A LF (listed as "Procyon n. sp." by
Webb and Wilkins 1984) as intermediate in size and morphological features
between the large species P. rexroadensis from the early Blancan Rexroad LF
(Kurten and Anderson 1980) and the living P. lotor. Morgan and Ridgway (1987)
reported a large Procyon from the late Blancan St. Petersburg Times LF. A
Procyon mandible from Leisey 1A tentatively referred to P. lotor (Berta this
volume) is also larger than the modern raccoon and is probably the same as the
Inglis species. The only Irvingtonian record of Procyon listed by Kurt6n and
Anderson (1980) was the small raccoon from Coleman 2A referred to P. lotor by
Martin (1974).
Two genera of seals found in Florida late Pliocene and Pleistocene faunas, the
monachine phocids Monachus and Callophoca, may provide some useful
biochronological information. With the exception of a single bone of the recently
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
extinct Caribbean monk seal, Monachus tropicalis, from the late Rancholabrean
Melbourne LF (Ray 1958), all other Florida records of Monachus are from the
early Irvingtonian (Table 2). Monachus is known from two localities in the
Caloosahatchee Formation of earliest Irvingtonian age, the De Soto Shell Pit LF
and the Ortona Locks along the Caloosahatchee River, and two sites in the
Bermont Formation of late early Irvingtonian age, Leisey Shell Pit and Rigby Shell
Pit. The apparent absence of Monachus in Florida during the remainder of the
Irvingtonian and most of the Rancholabrean may be an artifact of inadequate
sampling of marginal marine faunas during this time interval. Monachine phocids
also are present in four Florida late Blancan faunas, Brighton Canal, Kissimmee
River, Macasphalt Shell Pit, and Richardson Road Shell Pit. These Blancan
specimens appear to represent the large extinct genus, Callophoca, which also
occurs in Florida early Pliocene (late Hemphillian) faunas from the Bone Valley
Formation in central Florida (Morgan 1994).
Rodentia.--Martin (1969) reported several teeth of the giant beaver
Castoroides from the presumed late Blancan Santa Fe River 1B LF. However, the
mammalian fauna from Santa Fe 1B, like many of the Santa Fe faunas, consists of
a mixture of late Blancan and Rancholabrean taxa (see more detailed discussion of
Santa Fe sites below). Based on the absence of Castoroides from all other late
Blancan and earliest Irvingtonian faunas in Florida, it is highly probable that the
Santa Fe 1B giant beaver teeth are Rancholabrean in age. Removing this single
Blancan record, Castoroides first appears in North America during the
Irvingtonian. The oldest well-documented occurrences of Castoroides in Florida
are from the late early Irvingtonian Leisey Shell Pit and Crystal River Power Plant
local faunas. Specimens of Castoroides from Apollo Beach in Hillsborough
County are probably early Irvingtonian in age as well, although this site contains a
mixed assemblage of Irvingtonian and Rancholabrean taxa. Morgan and White
(this volume) describe the Leisey Castoroides as a new species, C. leiseyorum.
The more advanced species Castoroides ohioensis is common in Florida during the
late Rancholabrean, particularly in faunas that sample freshwater depositional
environments (Martin 1969).
Pocket gophers of the genus Geomys are first recorded in Florida from the
late Blancan Haile 15A and Macasphalt Shell Pit local faunas. These late Blancan
specimens were referred to G. propinetis by Morgan and Ridgway (1987), an
extinct species originally described from the early Irvingtonian Inglis IA and Haile
16A local faunas (Wilkins 1984). A small sample of pocket gopher teeth recently
collected from the De Soto Shell Pit, a correlative of Inglis 1A, is here referred to
G. propinetis as well (Table 2). The Geomys sample from the Leisey Shell Pit LF
is intermediate in size between G. propinetis and the extant southeastern pocket
gopher G. pinetis, but otherwise is very similar to the living species (Morgan and
White this volume). Apparently, G. pinetis evolved in Florida during the early
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 49
Irvingtonian, presumably derived from G. propinetis. Specimens of Geomys from
the late Irvingtonian Coleman 2A LF are indistinguishable from G. pinetis (Martin
1974; Wilkins 1984).
There are two caviomorph rodents in the Leisey Shell Pit LF, the capybara
Neochoerus sp. and the porcupine Erethizon dorsatum. Both are Neotropical
immigrants that reached North America in the late Blancan following the
beginning of the Great American Interchange. Two described species of
Neochoerus are known from Florida late Pliocene and Pleistocene faunas, N.
dichroplax from the late Blancan (Ahearn and Lance 1980) and N. pinckneyi from
the Rancholabrean (Ahearn 1981). Although the Neochoerus sample from Leisey
lacks the diagnostic M3, the size and shape of the p4 differ from that of N.
pinckneyi. More complete fossils are necessary to determine whether the Leisey
Neochoerus is referable to N. dichroplax or represents a new species. The only
Florida Blancan record of Erethizon is an associated palate and mandible from
Haile 7C. Frazier (1981) described a small species, E. kleini, from Inglis 1A.
Three well preserved porcupine mandibles from Leisey are much larger than E.
kleini, and appear to be indistinguishable from the living species, E. dorsatum. E.
dorsatum is also present in the early Irvingtonian Haile 16A and Apollo Beach
local faunas and the late Irvingtonian Coleman 2A LF (Frazier 1981).
Martin (1979) reviewed the evolutionary history of the cotton rat Sigmodon.
Six species of cotton rats are recorded from Florida late Pliocene and Pleistocene
faunas. The small, primitive species, S. medius, occurs in two Florida late Blancan
faunas, Haile 15A (Martin 1979) and Macasphalt Shell Pit (Morgan and Ridgway
1987). The similar species, S. minor (conspecific with S. medius according to
Kurten and Anderson 1980), is reported from eastern North America for the first
time based on two teeth from the earliest Irvingtonian De Soto Shell Pit (Morgan
and White this volume). The larger and more progressive species, S. curtisi, was
recorded from the earliest Irvingtonian Inglis 1A LF by Martin (1979) and also
occurs in the correlative De Soto Shell Pit LF (Morgan and White this volume).
Martin (1979) described S. libitinus from the late early Irvingtonian Haile 16A LF.
Morgan and White (this volume) report S. libitinus from the Leisey Shell Pit LF
and two other correlative late early Irvingtonian faunas, Payne Creek Mine and
Haile 21A. S. libitinus is intermediate in certain morphological features between
S. curtisi from the earliest Irvingtonian and S. bakeri from the late Irvingtonian
Coleman 2A LF (type locality) and several early Rancholabrean faunas (Martin
1979). The extant cotton rat S. hispidus is the typical species found in Florida late
Rancholabrean faunas, although it first appears in the early Rancholabrean
Daytona Beach and Haile 8A local faunas.
The oldest arvicoline rodent known from Florida is the primitive muskrat,
Ondatra idahoensis, from the earliest Irvingtonian Inglis IA and De Soto Shell Pit
local faunas. 0. idahoensis occurs in western faunas of similar age, such as Curtis
Ranch in Arizona (Repenning 1987), as well as slightly older late Blancan faunas
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
(Blancan V of Repenning 1987), including Borchers, Kansas and Grand View,
Idaho (the type locality). The larger and more advanced muskrat, 0. annectens, is
present at Leisey Shell Pit, as well as in the correlative Pool Branch and Payne
Creek Mine local faunas from the Bone Valley Region in Polk County. O.
annectens is a rather long-ranging species characteristic of late early and middle
Irvingtonian faunas (Nelson and Semken 1970; Martin and Tedesco 1976;
Repenning 1987). Western early Irvingtonian faunas containing 0. annectens
include Java, South Dakota (Martin and Tedesco 1976; Martin 1989), Kentuck and
Wathena, Kansas (Repenning 1987) and Sappa, Nebraska (Martin and Schultz
1985).
Two other genera of arvicolines are recorded from Leisey, the bog lemming
Synaptomys and the vole Pedomys. About ten complete arvicoline teeth from
Leisey are referred to Pedomys by Morgan and White (this volume). The Leisey
Pedomys are very similar to a large series of jaws and teeth from the slightly older
Haile 16A LF currently being described as a new species by Robert Martin (in
prep.). A third sample of this same species occurs in the correlative Payne Creek
Mine LF (see discussion below). Two teeth from Leisey are referable to
Synaptomys sp. A similar sample of Synaptomys from Haile 16A also is currently
under study by Robert Martin. The Leisey and Haile 16A Synaptomys appear to be
related to S. australis, a large, extinct species present in Florida Rancholabrean
faunas (Martin, pers. comm.).
Perissodactyla.--Hulbert (this volume) describes the Leisey sample of the
giant tapir, Tapirus haysii, and discusses the biochronology of the genus in Florida.
A smaller, undescribed species of Tapirus is present in the earliest Irvingtonian
Inglis 1A and De Soto Shell Pit local faunas, the latest Blancan Haile 7C LF, and
possibly several other late Blancan sites as well (Table 2). T. haysii is known from
the Blancan and Irvingtonian in the western United States, but this species has a
more restricted range in Florida where it has been identified from ten late early or
middle Irvingtonian sites (Hulbert this volume). The tapir from the late
Irvingtonian Coleman 2A LF, although represented only by postcranial elements,
is within the size range of T. veroensis, the common Florida Rancholabrean
species.
The three species of Equus recognized in the Leisey Shell Pit LF (Hulbert,
this volume) are of limited biochronologic use because of the chaotic state of the
taxonomy of North American Equus. None of the Leisey Equus are definitely
conspecific with named species in well-dated western faunas. The absence of the
hipparionine horses Nannippus and Cormohipparion, found in Florida Blancan
faunas, is suggestive of a post-Blancan age for Leisey. The most common horse at
Leisey, Equus "leidyi," occurs throughout the Irvingtonian in Florida. E. "leidyi"
is very similar to, and possibly represents a small eastern subspecies of, E. scotti, a
common late Blancan to early Rancholabrean species with a wide distribution in
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 51
western North American (Winans 1989). The second most common horse at
Leisey is an apparently undescribed form of the subgenus Hemionus. This
subgenus is first known from the late Blancan of western North America (Skinner
1972), where it persisted through the Rancholabrean (E. francisci, Lundelius and
Stevens 1970; Winans 1989). Other than Leisey, this new species is provisionally
identified based on limited samples of isolated teeth from Pool Branch, Apollo
Beach, and Flamingo Waterway in Charlotte County. The rarest of the Leisey
horses is E. "fraternus" which is much better represented at Haile 16A. Both the
relationships and biochronological significance of this species are poorly known.
Artiodactyla.-Two species of peccaries occur in the Leisey Shell Pit LF,
Mylohyus fossilis and Platygonus vetus. In his review of the Leisey tayassuids,
Wright (this volume) refers the sample of Mylohyus to M. fossilis, a species typical
of middle Irvingtonian through late Rancholabrean faunas in the eastern and
central United States. Kinsey (1974) described the small Blancan species M.
floridanus from the late Blancan Haile 15A LF. M. floridanus has since been
reported from the late Blancan Macasphalt Shell Pit as well (Morgan and Ridgway
1987). The Leisey Mylohyus is larger than other specimens of late Blancan or
Rancholabrean Mylohyus examined from Florida.
Wright (this volume) notes that it is difficult to separate the various Blancan
and Irvingtonian species of Platygonus, most of which are larger than the common
Rancholabrean species, P. compressus. Large specimens of Platygonus from
Florida late Blancan faunas (Table 2) generally have been referred to P.
bicalcaratus (Webb 1974a; Morgan and Ridgway 1987). Wright (this volume)
provisionally refers the Leisey Platygonus to P. vetus, a large species known from
many early and middle Irvingtonian faunas, including large samples from Inglis
1A and Haile 21A. An even larger Platygonus from the late Irvingtonian Coleman
2A LF and the early Rancholabrean Haile 7A LF is tentatively referred to P.
cumberlandensis (Martin 1974; Wright this volume). The smaller species, P.
compressus, appears in Florida during the early Rancholabrean.
Camels are the most abundant large mammals in the Leisey Shell Pit LF,
approached in numbers of individuals only by horses. Two camels have been
identified from Leisey, Palaeolama mirifica and Hemiauchenia macrocephala
(Webb and Stehli this volume; they use the species name H. seymourensis for the
Leisey sample of this genus). H. macrocephala is a long-ranging species that first
appeared in the late Blancan and survived until the end of the Pleistocene (Webb
1974b). A larger species, H. blancoensis, has been identified from several Florida
late Blancan sites (Table 2; Webb 1974b; Morgan and Ridgway 1987). The
occurrence of Palaeolama at Leisey represents one of the earliest North American
records of the genus; only the sample from Haile 16A is older. The two major
Leisey sites, Leisey IA and 3A, differ significantly in their camel faunas.
Palaeolama is the most abundant large mammal at Leisey 1A, outnumbering
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Hemiauchenia by more than two to one. In contrast, Hemiauchenia
overwhelmingly dominates the large vertebrate fauna at Leisey 3A and Palaeolama
is absent.
Proboscidea.--The Leisey IA proboscidean sample is predominantly
composed of the mammoth Mammuthus, with only a few, mostly juvenile,
individuals of the mastodon Mammut americanum and several fragmentary
cheekteeth and tusks of the gomphothere Cuvieronius tropicus (see Webb and
Dudley this volume). Leisey 3A is almost devoid of proboscideans, whereas
Cuvieronius is the most common proboscidean in Leisey Shell Pit 3. The presence
of Cuvieronius in Florida faunas was once thought to indicate an early Irvingtonian
or older age (Brooks 1968). However, subsequent discoveries confirm that
Cuvieronius occurs in Florida from the late Blancan throughout the Irvingtonian
and into the early Rancholabrean. Although Cuvieronius is probably the most
ubiquitous proboscidean in Florida Irvingtonian faunas, most of the fossils consist
of isolated and/or fragmentary remains. The only comparatively rich samples of
this genus from Florida are in the early Irvingtonian Punta Gorda LF, Charlotte
County (Brooks 1968; Webb 1974a; see discussion below) and the early
Rancholabrean Daytona Beach LF (Edmund et al. in prep.). The Daytona Beach
site is the youngest recorded Florida occurrence of Cuvieronius.
Leisey Shell Pit 1A has one of the richest known Irvingtonian samples of
Mammuthus (see Webb and Dudley this volume). The Leisey mammoth teeth are
intermediate in morphological features between teeth of M. meridionalis and M.
imperator as defined by Maglio (1973). Webb and Dudley restrict M. meridionalis
to Old World mammoths and refer primitive North American mammoths from the
early Irvingtonian to M. hayi. Webb (1974a) and Webb and Dudley (this volume)
also discuss a sample ofM. hayi from the Punta Gorda LF which is very similar to
the Leisey Mammuthus. The late early Irvingtonian Mammuthus from Leisey and
Punta Gorda are comparable to samples from the Gilliland LF in Texas and the
Holloman LF in Oklahoma, and may be among the earliest mammoths in the New
World. The next youngest Florida fauna containing a measurable sample of
mammoths is the early Rancholabrean Bradenton LF. All Florida Rancholabrean
specimens of Mammuthus are referable to the widespread species, M. columbi.
Sirenia and Cetacea.- The Sirenia and Cetacea are the only groups of
Leisey mammals not covered in the individual taxonomic papers. The Leisey Shell
Pit LF represents the earliest well documented North American record of the
manatee Trichechus. Specimens of Trichechus from Leisey 1A consist of a partial
skull cap (UF 87226), an isolated tooth (UF 87227), and the proximal humerus of a
juvenile (UF 81514 ). Manatee fossils are more common at Leisey 3 and include a
nearly complete skull in the National Museum of Natural History, a partial skull
(UF 124557), a complete humerus (UF 135693), and a metacarpal (UF 129092).
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 53
Domning (1982) reported several supposed late Blancan manatee specimens from
Santa Fe 1B, and noted that they were very similar to the living species T.
manatus. Subsequent study of the Santa Fe 1B fauna (Morgan and Ridgway 1987
and discussion below) has shown that this site contains a mixture of late Blancan
and Rancholabrean taxa. The abundance of manatee fossils in other
Rancholabrean sites along the Santa Fe River suggests that the Santa Fe
Trichechus is late Pleistocene as well. The Leisey Trichechus sample is currently
under study by Daryl Domning.
Cetaceans are rare at Leisey. Two isolated teeth of small delphinids from
Leisey 1A are tentatively identified as Stenella (UF 84919) and Tursiops (UF
84629). Three associated vertebrae (UF 142239) from Leisey 3B represent a much
larger delphinid similar in size to the pilot whale Globicephala.
Biochronology of the Leisey Lower Vertebrate Fauna.- There are several
taxa of lower vertebrates from the Leisey Shell Pit LF, including sharks, turtles,
and birds, that provide some information relating to the age of the site. There is no
established biochronology for any of these groups, and thus the age data are not so
precise as those provided by the better known mammals. Furthermore, several of
these taxa are restricted to Florida sites.
Shark teeth are common in certain units within the late Pliocene and
Pleistocene shell bed sequence of central and southern Florida, but very little has
been published previously on Florida sharks of this age (Scudder et al. this
volume). In his review of Florida fossil sharks, Tessman (1969) did not mention
any specimens younger than samples from the early Pliocene portion of the Bone
Valley Formation.
Three extinct species of sharks, Ginglymostoma serra, Isurus hastalis, and
Hemipristis serra, and the extinct ray Rhynchobatus sp. have been identified from
the Leisey Shell Pit (Scudder et al. this volume). There are no published records of
these species from other Florida Pleistocene faunas. All three of the sharks are
known from the early Pliocene Bone Valley Formation (Tessman 1969), and I.
hastalis and H. serra have been reported from the Pliocene Tamiami Formation
(Morgan and Pratt 1983). It is possible that the teeth of the extinct species of
sharks and rays were reworked into the early Pleistocene Bermont Formation in the
Leisey Shell Pit from underlying Miocene sediments. Most of the Miocene shark
teeth from Leisey are waterworn and have a different color of preservation,
whereas two H. serra teeth from Leisey 1A and one from Leisey 3B were collected
in situ, are in excellent condition, and have the same state of preservation as the
remainder of the shark teeth from this unit.
The two teeth of Isurus hastalis from Leisey are broken and waterworn, and
thus reworking from the underlying Miocene beds is a distinct possibility. A tooth
of the extant shortfin mako shark, 1. oxyrinchus, from the Leisey Shell Pit fauna
apparently represents the first published fossil record of this species in Florida
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
(Tessman 1969). The age of the Ginglymostoma serra teeth from Leisey is
uncertain owing to the rarity of this species in Florida fossil sites, although the
specimens are not obviously reworked. Other Florida records of G. serra are from
the Miocene and Pliocene (Tessman 1969). As is the case with the mako sharks, a
living species of nurse shark, G. cirratum, occurs in the Leisey fauna as well.
Teeth of the living great white shark, Carcharodon carcharias, are relatively
common at Leisey. This shark also has been identified from two late Pliocene sites
in the Pinecrest Beds, the Macasphalt Shell Pit LF in Sarasota County (Waldrop
and Wilson 1990) and the Kissimmee River LF, and from the earliest Irvingtonian
Caloosahatchee Formation at the Cochran Shell Pit in Hendry County (Scudder et
al. this volume). The oldest published record of C. carcharias in Florida is from
the Bone Valley Formation of early Pliocene age (Tessman 1969). The extinct
giant white shark, C. megalodon, was common and widespread in Florida during
the late Miocene and early Pliocene.
The most abundant bony fish in the Leisey Shell Pit LF is the alligator gar,
Atractosteus spatula, a species no longer found in the Florida peninsula (see
Scudder et al. this volume). The closest living population of A. spatula is in the
Florida panhandle (Wiley 1976). Despite its current absence from southern
Florida, A. spatula is common in many Pliocene and early Pleistocene vertebrate
sites in the southern half of the state. The youngest record of A. spatula from
peninsular Florida is the early Rancholabrean Oldsmar LF in Pinellas County
(Scudder et al. this volume).
Three turtles from Leisey provide some evidence for the age of the site.
Auffenberg (1988) described a new species of small land tortoise, Geochelone
mlynarskii, from the late Irvingtonian Coleman 2A LF. Meylan (this volume)
refers the dwarf land tortoise from Leisey to this species, although he transfers it to
the genus Hesperotestudo. Auffenberg (1988) noted that among members of the
lineage of small land tortoises in Florida, termed the incisa group from the
Rancholabrean species H. incisa, the major evolutionary changes occurred between
the Blancan and early Irvingtonian, and between the late Irvingtonian and
Rancholabrean. The Leisey sample is most similar morphologically to the
Irvingtonian members of the incisa group, tentatively referred to H. mlynarskii
(Auffenberg 1988; Meylan this volume).
The most common freshwater turtle in the Leisey Shell Pit LF is Trachemys
scripta, an extant species now absent from central and southern peninsular Florida
(Meylan this volume). Leisey represents one of the earliest records of T. scripta.
Specimens from the earliest Irvingtonian Inglis 1A and De Soto Shell Pit faunas
appear to be referable to the larger extinct species, T. platymarginata (synonymized
with T. idahoensis by Jackson 1988). T. platymarginata originally was described
from the late Blancan Haile 15A and Sante Fe River 1 faunas (Weaver and
Robertson 1967), and is now known from several late Blancan faunas in the
southern half of the state.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 55
Meylan (this volume) documents the widespread occurrence of the alligator
snapping turtle, Macroclemys temmincki, outside of its modern range during the
late Pliocene and Pleistocene. M. temmincki is not currently found south of the
Suwannee River in northern peninsular Florida. Fossil alligator snapping turtles
are known from the late Hemphillian Palmetto Fauna, the late Blancan Macasphalt
Shell Pit and St. Petersburg Times faunas, and the earliest Irvingtonian De Soto
Shell Pit LF, as well as Leisey (Meylan this volume). The youngest fauna from
southern Florida containing M. temmincki is the early Rancholabrean Oldsmar LF.
Several species of birds from Leisey are biochronologically useful (Emslie this
volume). The small extinct loon, Gavia concinna, was previously known in
Florida only from the late Hemphillian Bone Valley Formation in central Florida.
Emslie (this volume) identifies G. concinna from Leisey thereby extending the
biostratigraphic range of this species in Florida by about 3 million years. Leisey is
the type locality for the extinct condor, Gymnogyps kofordi (Emslie 1988). This
species is also known from the early Irvingtonian Haile 16A LF. Earlier records
for the genus Gymnogyps include the late Blancan Macasphalt Shell Pit LF and the
earliest Irvingtonian Inglis 1A LF (Emslie 1988; 1992a). All other fossils of
Gymnogyps from Florida and elsewhere in North America are Rancholabrean in
age and are referable to the living species G. californianus (Emslie 1988).
Emslie (this volume) refers a small sample of fossil turkeys from Leisey to
Meleagris leopoldi/M. anza, following the usage of Steadman (1980). The Leisey
Meleagris is similar to the large sample from Inglis IA referred to M. leopoldi/M.
anza by Steadman (1980). The Inglis and Leisey turkeys are smaller than
specimens of Meleagris from Coleman 2A, a sample recognized by Steadman
(1980) as intermediate between the Inglis turkeys and Rancholabrean specimens of
the living species M. gallopavo.
Summary of Leisey Vertebrate Biochronology.- Knowledge of Florida
Irvingtonian faunas has vastly increased over the past two decades to the point
where Florida may be the most densely sampled geographic region in North
America for the Irvingtonian Land Mammal Age. Owing to the lack of absolute
dates, there is still a degree of uncertainty in correlating Florida Irvingtonian
faunas within the state. Paleoecological and biogeographical factors magnify this
uncertainty when Florida sites are compared to Irvingtonian faunas elsewhere in
the United States. Despite these problems, the correlation of Florida Irvingtonian
faunas both within and outside the state should be accurate within approximately
0.2 Ma.
Four of the genera used by Lundelius et al. (1987) to define the Irvingtonian
NALMA occur in the Leisey Shell Pit LF, including Smilodon, Lepus, Equus s.s.,
and Mammuthus. Other genera identified from Leisey that have their first
appearance in the Irvingtonian are Nothrotheriops, Castoroides, Lutra, and
Palaeolama. Among the Irvingtonian immigrant genera recorded from Leisey,
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Mammuthus and Lutra are Old World in origin and Nothrotheriops is Neotropical.
Smilodon, Castoroides, and Palaeolama evolved in North America, but
nonetheless are important biostratigraphic indicators. Irvingtonian faunas also are
recognized by the absence of characteristic Blancan genera such as Borophagus,
Nannippus, Equus (Dolichohippus), and Rhynchotherium. All of these genera are
known from Florida late Blancan faunas (Morgan and Ridgway 1987), but are
absent from Leisey.
The Leisey Shell Pit LF is younger than the earliest Irvingtonian Inglis 1A
and De Soto Shell Pit local faunas (Table 2; Klein 1971; Webb 1974a; Webb and
Wilkins 1984) based on the presence of Mammuthus and the absence of various
Blancan holdover species found in the two older faunas, including the dwarf
Florida form of Megalonyx leptostomus, Chasmaporthetes ossifragus, Trigonictis
macrodon, and Capromeryx arizonensis. Furthermore, Leisey records the first
occurrence in Florida of at least five genera not presently known from earliest
Irvingtonian faunas: Nothrotheriops, Lutra, Castoroides, Palaeolama, and
Mammuthus. There are also four genera of rodents present at Leisey that differ at
the species level from their congeners at Inglis and/or De Soto (in parentheses),
including Geomys pinetis (G. propinetis), Erethizon dorsatum (E. kleini),
Sigmodon libitinus (S. curtisi), and Ondatra annectens (0. idahoensis).
Leisey is considerably older than the only other thoroughly studied
Irvingtonian fauna from Florida, the late Irvingtonian Coleman 2A LF (Martin
1974). Leisey and Coleman share only a few diagnostic mammalian taxa, as the
latter site actually more closely resembles early Rancholabrean faunas. Examples
of typically Rancholabrean species at Coleman are Didelphis virginiana,
Holmesina septentrionalis, Neofiber alleni, and Tapirus veroensis. Leisey lacks
Didelphis and Neofiber and has the older species, H. floridanus and T. haysii.
There are also two genera of rodents from Leisey that differ at the species level
from their congeners or closely related genera at Coleman (in parentheses),
Sigmodon libitinus (S. baker) and Pedomys n. sp. (Pitymys aratai). Leisey and
Coleman do share several long-ranging Irvingtonian species, including Canis
armbrusteri, Arctodus pristinus, and Equus "leidyi."
A more precise age refinement for Leisey is now possible based on
comparisons with other Irvingtonian faunas from Florida, most of which were
discovered during the past 20 years. Like Leisey, most of these faunas are younger
than Inglis 1A and older than Coleman 2A. These intermediate Irvingtonian
faunas include Haile 16A, Pool Branch, Payne Creek Mine, Crystal River Power
Plant, Rigby Shell Pit, Haile 21A, and McLeod Limerock Mine (see Table 2 and
discussions of the individual faunas below). All of these faunas except McLeod
appear to be late early Irvingtonian in age, and therefore are roughly correlative
with Leisey and between 1.6 and 1.0 Ma. McLeod is somewhat younger that the
other faunas, probably late middle Irvingtonian in age (between 0.8 and 0.6 Ma).
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 57
The presence of three species at Leisey that are restricted to the late Blancan
and/or early Irvingtonian serves to further constrain the age of this fauna,
effectively ruling out a middle Irvingtonian age. The large glyptodont
Glyptotherium arizonae occurs only in early Irvingtonian faunas in the western
United States (Gillette and Ray 1981). In Florida, G. arizonae has been recorded
from the earliest Irvingtonian Inglis and De Soto sites, as well as five late Blancan
localities (Table 2). Leisey is the youngest Florida record of G. arizonae, and the
only fauna in which this glyptodont has been found in association with
Mammuthus. All occurrences of Pachyarmatherium leiseyi (Downing and White
this volume) are from late Blancan and early Irvingtonian sites. Canis edwardii
has been reported only from early Irvingtonian faunas (Kurten and Anderson
1980). Florida specimens of this canid are known from both the earliest
Irvingtonian (Inglis and De Soto) and the late early Irvingtonian (Leisey, Crystal
River, Rigby, and Haile 21A).
At least three species of mammals found at Leisey appear to be restricted to
late early Irvingtonian faunas in Florida. The cotton rat Sigmodon libitinus is
known only from Leisey, Payne Creek, and the type locality Haile 16A LF (Martin
1979), all of which are late early Irvingtonian. These three faunas also share the
same undescribed species of Pedomys (Martin in prep.). Although identified from
only two Florida vertebrate faunas, Leisey and Pool Branch, Nothrotheriops
texanus also appears to be restricted to the late early Irvingtonian. Elsewhere in
North America, the biochronologic ranges of the canids Canis edwardii and C.
armbusteri do not overlap, with the former restricted to the early Irvingtonian and
the latter found in the middle and late Irvingtonian. Their co-occurrence at Leisey
and Haile 21A confirms that C. armbrusteri first appears in the early Irvingtonian,
at least in Florida.
Megalonyx wheatleyi is typically a middle Irvingtonian species, including the
Florida sample from McLeod (McDonald 1977). However, a skull and other
specimens from Leisey appear to be referable to M. wheatleyi, rather than to the
smaller Florida form of M. leptostomus, typical of late Blancan and earliest
Irvingtonian faunas. Leisey, Payne Creek, and Pool Branch all have the extinct
muskrat, Ondatra annectens, a species found in late early and middle Irvingtonian
faunas in the western United States. The large tapir, Tapirus haysii, also occurs
only in the late early and middle Irvingtonian in Florida (Hulbert this volume).
Analysis of several evolutionary lineages, in particular Smilodon and Holmesina,
helps to clarify the age of Leisey. The Leisey representatives of these lineages
further strengthen evidence from other taxa that this site best fits in the latter half
of the early Irvingtonian, between 1.6 and 1.0 Ma.
Comparisons with Irvingtonian faunas in western North America indicate
that Leisey is most similar in age to the Gilliland (Hibbard and Dalquest 1966) and
Rock Creek (Troxell 1915) local faunas from Texas, the Holloman LF from
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 3. List of mammals shared by three correlative late early Irvingtonian local faunas: Leisey Shell Pit,
Florida; Gilliland, Texas; and Holloman, Oklahoma. Only species that occur at Leisey and one of the two
other faunas are listed here. For complete mammalian faunal lists from these sites see Table 1 for Leisey,
Hibbard and Dalquest (1966) for Gilliland, and Dalquest (1977) for Holloman.
Species Leisey Gilliland Holloman
Xenarthra
Holmesinafloridanus X X -
Glyptotherium arizonae X X X
Paramylodon harlani X X X
Nothrotheriops texanus X X -
Carnivora
Canis edwardii X X -
Procyon sp. X X -
Homotherium sp. X X -
Lagomorpha
Sylvilagusfloridanus X X -
Perissodactyla
Tapirus haysii X X X
Equus scow orE. "leidyi"' X X X
Equus (Hemionus) sp. X X X
Artiodactyla
Platygonus vetus X X X
Hemiauchenia macrocephala X X
Odocoileus sp. X X X
Proboscidea
Cuvieronius tropicus X X -
Mammuthus cf. M. hayi X X X
Equus "leidyi" is morphologicay similar to E. scotti and possibly represents a smaller eastem subspecies ofthat species (Hulbert this
volume).
Oklahoma (Dalquest 1977), the Kentuck, Nash, and Wathena local faunas from
Kansas, the Sappa LF of Nebraska (Martin and Schultz 1985; Lundelius et al.
1987), and the Java LF from South Dakota (Martin 1973 1989). Sappa is the type
fauna for the early Irvingtonian Sappan Subage (Schultz et al. 1978; Martin and
Schultz 1985; Lundelius et al. 1987). Sappa, Java, and the three Kansas faunas
mostly consist of small mammals, and thus correlation with Leisey is based either
on absolute ages or comparisons with other faunas of similar age that have both
large and small mammals.
Among the sites listed above, Leisey has the most taxa in common with
Gilliland and Holloman (Table 3). All three faunas record the association of the
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 59
glyptodont Glyptotherium arizonae (Gillette and Ray 1981) and a primitive
Mammuthus (Hibbard and Dalquest 1966; Dalquest 1977; Webb and Dudley this
volume). These probably represent some of the earliest records of mammoths in
North America, whereas G. arizonae is unknown in younger middle Irvingtonian
faunas. The concurrent range zone of Glyptotherium arizonae and Mammuthus cf.
M. hayi seems to occur only during a rather narrow interval of time in the late
early Irvingtonian. Another age-diagnostic large mammal shared by these three
faunas is the giant tapir, Tapirus haysii, a species restricted to late early and
middle Irvingtonian faunas in Florida (Hulbert this volume). Leisey and Gilliland
share several additional mammals that are found primarily in early Irvingtonian
faunas, including the rare ground sloth, Nothrotheriops texanus, a medium-sized
representative of the pampathere Holmesina floridanus, and the canid Canis
edwardii. Hibbard and Dalquest (1966) stated that the Gilliland LF was post-
Blancan and pre-Cudahy in age, or in other words, early Irvingtonian as defined
here and by Lundelius et al. (1987). Although Dalquest (1977) regarded Gilliland
and Holloman as earliest Irvingtonian in age, the presence ofMammuthus indicates
that these two faunas are younger than faunas such as Curtis Ranch now generally
regarded as earliest Irvingtonian (Lundelius et al. 1987).
Leisey and the correlative faunas mentioned above all are considered to be
late early Irvingtonian in age (between 1.6 and 1.0 Ma). These faunas are
somewhat younger than earliest Irvingtonian faunas (2.0 to 1.6 Ma) including:
Curtis Ranch, Arizona; Inglis 1A; and De Soto Shell Pit. Leisey is older than
typical North American middle Irvingtonian (=Cudahyan) faunas (1.0 to 0.6 Ma)
including: the type Irvington fauna from California; Cudahy, Kansas; Conard
Fissure, Arkansas; Vera, Texas; Cumberland Cave, Maryland; Port Kennedy Cave,
Pennsylvania; and Hamilton Cave, West Virginia.
BIOCHRONOLOGY OF FLORIDA LATE PLIOCENE AND
PLEISTOCENE VERTEBRATE FAUNAS
In the following section we present synopses of the most important Blancan
and Irvingtonian vertebrate fossil sites from Florida, with the exception of Leisey
which is discussed in detail above. Each synopsis includes a brief description of
the location and physical setting of the site, discussion of the key mammalian taxa,
reference to other factors such as relative sea level position that pertain to its age,
and citations to the most important published references. Figure 4 is a map of
Florida showing the location of the Blancan and Irvingtonian vertebrate faunas
discussed in the text. More complete information on these sites is available in the
locality files of the UF Vertebrate Paleontology Collections. Following these
synopses we present a brief review of the Rancholabrean NALMA in Florida.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Table 2 is a list of 128 species of biochronologically diagnostic mammals
from the most important Florida late Blancan and Irvingtonian faunas, as well as
composite faunal lists for the early and late Rancholabrean. We have limited the
mammalian faunas listed in Table 2 to those represented by ten or more diagnostic
Figure 4. Map of Florida showing location of Blancan and Irvingtonian vertebrate faunas discussed in text
Late Blancan: 1. Santa Fe River 1, 2, 4A, 8A, and 15A, Columbia County, 2. Haile 15A, Alachua County;
3. St Peterburg Times, Pinellas County, 4. Kissimmee River, Okeechobee County 5. Brighton Canal,
Highlands County, 6. Macasphalt Shell Pit, Sarasota County, 7. Bass Point Waterway, Sarasota County
8. El Jobean, Charlotte County, 9. Acline Shell Pit, Charlotte County 10. Lehigh Acres, Lee County
latest Blancan: 11. Haile 7C, Alachua County earliest Irvingtonian: 12. Inglis IA, Citrus County, 13. De
Soto Shell Pit, De Soto County, 14. Forsberg Shell Pit, Charlotte County late early Irvingtonian: 15. Haile
16A, Alachua County, 16. Haile 21A, Alachua County, 17. Crystal River Power Plant, Citrus County, 18.
Leisey Shell Pit, Hillsborough County, 19. Payne Creek Mine, Polk County, 20. Pool Branch, Polk
County, 21. Rigby Shell Pit, Sarasota County, 22. Punta Gorda, Charlotte County, middle Irvingtonian:
23. McLeod Limerock Mine, Levy County, late Irvingtonian: 24. Coleman 2A, Sumter County.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 61
species, which excludes several small, biochronologically significant faunas.
These faunas will be briefly discussed after the summaries of the larger sites.
Figure 5 is a correlation chart showing the relative stratigraphic position and age
of the major Florida Blancan and Irvingtonian faunas discussed in the text.
DuBar (1958) initiated the use of terrestrial vertebrate fossils for the age
determination of Plio-Pleistocene shell beds in Florida; however, he was hindered
by the limited diversity of the vertebrate samples available to him. Furthermore,
35 years ago paleontologists had only a limited understanding of Late Cenozoic
AGE NALMA
RANCHO-
LABREAN
1
0.79 Z
0
I-
cc
1.u4
Iul I ___
*1 DEVILS DEN. REDDes. MELmURNE
NORTH
FLORIDA
DI EVILS DEN. REDDCK MELBOURNE
ARREDONDO 2A. ICHETUCKNEE RIV.
IDAYTONA BEACH
HAILE 7A, BA WILLISTON 3A 3B
S COLEMAN2A
1.0
HAILE 21A
CRYSTAL RIVER
POWER PLANT
HALE 16A
SINGLIS1A
HAILE 7C
HALE 15A
SANTA FE RIV. 1
SOUTH
FLORIDA
ISEMINLE FIELD.VEBO
CUTLER. MONKEY JUNGLE I
I BRADENTON 1
APOLLO BEACH
LEISEY SHELL PIT. PUNTA GORDA
POOL BRANCH, PAYNE CREEK
RIGBY SHELL PIT
DESOTO SHELL PIT
FORSBERG SHELL PIT
MACASPHALT SHELL PIT
ACLINE SHELL PIT. KISSIMMEE
RIVER, BRIGHTON CANAL
ST. PETE. TIMES, LEHIGH ACRES
Figure 5. Correlation chart showing relative position and age of the Florida late Pliocene and Pleistocene
vertebrate faunas discussed in text The faunas are divided into those from northern and southern peninsular
Florida, with the arbitrary boundary between the two regions placed at 280N latitude. The faunas listed
within a box are approximately equivalent in age, but their order in the box does not represent stratigraphic
superposition. Vertical size of boxes expresses the range of possible ages of the sites enclosed in the box.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
mammalian chronology in Florida and elsewhere. Webb (1974a) summarized the
chronology of Florida late Pliocene and Pleistocene mammals and provided faunal
lists for the two Blancan and four Irvingtonian faunas known from the state at that
time. In the past decade, large and taxonomically diverse samples of Blancan and
Irvingtonian vertebrate fossils have been collected from Florida in direct
stratigraphic context with well known marine units (see Table 4) including:
Pinecrest Beds (Macasphalt Shell Pit, Morgan and Ridgway 1987; Hulbert 1988a;
Emslie 1992a,b; this paper), Caloosahatchee Formation (De Soto Shell Pit, this
paper), and Bermont Formation (Leisey and Rigby Shell Pits, Hulbert and Morgan
1989; Webb et al. 1989; papers in this volume). Furthermore, the Late Cenozoic
mammalian biochronology of North America has vastly improved over the past 20
years, thanks to the studies of Skinner and Hibbard (1972), Webb (1974a), Lindsay
et al. (1975), Kurten and Anderson (1980), Repenning (1980 1987), Lundelius et
al. (1987), and many others.
BLANCAN
There is a 2-million-year gap in the Florida terrestrial vertebrate record
between the well known early Pliocene (late Hemphillian) faunas of the Bone
Valley Formation (5.2 to 4.5 Ma) and various late Pliocene (late Blancan) faunas
(2.5-2.0 Ma).. Although early Blancan terrestrial faunas are unknown from
Florida (Morgan and Ridgway 1987), the marine Bee Ridge Fauna from Sarasota
County in southwestern Florida (Morgan 1994) appears to be between 3.5 and 3.0
Ma in age (=Blancan III or IV of Repenning 1987). The Bee Ridge Fauna consists
exclusively of cetaceans, pinnipeds, and other marine vertebrates that are not
comparable with continental faunas of early Blancan age from western North
America. The oldest faunas considered in our study are late Blancan in age
(Blancan V of Repenning 1987). The two most widely cited reviews of Florida
Pliocene and Pleistocene faunas list only two late Blancan vertebrate faunas from
the state, Haile 15A and Santa Fe River 1 (Webb 1974a; Kurten and Anderson
1980). Reasonably complete mammalian faunal lists have been published for both
of these sites (Webb 1974a; Robertson 1976). Morgan and Ridgway (1987)
described the late Blancan St. Petersburg Times LF from Pinellas County along the
central Gulf Coast. They also discussed several other smaller Blancan faunas and
presented a brief review of Florida Blancan sites. Numerous recent publications
have discussed the late Blancan vertebrate fauna from the Macasphalt Shell Pit in
Sarasota County (Morgan and Ridgway 1987; Hulbert 1988a; Jones et al. 1991;
Emslie 1992a,b).
Table 4. Florida Blancan and Irvingtonian vertebrate faunas collected in stratigraphic superposition with or in association with marine geologic units.
Formation Age Site Remarks
Pinecrest Beds late Pliocene Macasphalt Shell Pit Collected in place from Unit 4 of Petuch (1982) within
Tamiami Formation (2.5-2.0 Ma) Pinecrest Beds
Pinecrest Beds late Pliocene Kissimmee River Not collected in place, associated molluscan fauna typical
Tamiami Formation (2.5-2.0 Ma) of Pinecrest Beds
Pinecrest Beds late Pliocene St Petersburg Times Collected in place immediately above Pinecrest Beds
Tamiami Formation (2.5-2.0 Ma)
Pinecrest Beds late Pliocene Lehigh Acres Not collected in place, associated molluscan fauna typical
Tamiami Formation (2.5-2.0 Ma) of Pinecrest Beds
Pinecrest Beds late Pliocene Brighton Canal Not collected in place, associated molluscan fauna typical
Tamiami Formation (2.5-2.0 Ma) of Pinecrest Beds
Pinecrest Beds late Pliocene Acline Shell Pit Collected in place from Pinecrest Beds
Tamiami Formation (2.5-2.0 Ma)
Caloosahatchee latest Pliocene De Soto Shell Pit Collected in place within Caloosahatchee Fm.
Formation (2.0-1.6 Ma)
Caloosahatchee latest Pliocene Forsberg Shell Pit' Collected in place within or underlying Caloosahatchee Fm.
Formation (2.0-1.6 Ma)
Bermont Formation early Pleistocene Leisey Shell Pit Collected in place within Bermont Fm.
(1.6-1.0 Ma)
Bermont Formation early Pleistocene Crystal River Collected in place within Bermont Fm.
(1.6-1.0 Ma) Power Plant
Bermont Formation early Pleistocene Rigby Shell Pit Collected in place within Bermont Fm.
(1.6-1.0 Ma)
Bermont Formation early Pleistocene Punta Gorda LF2 Collected in place above Bermont Fm.
(1.6-1.0 Ma) of Webb (1974a)
IThe Forsberg Shell Pit and Punta Gorda locality of Waldrop and Wilson (1990) are the same site. However, this is a different locality from the Punta Gorda Local Fauna of Webb (1974a) and Kurten and
Anderson (1980).
2 = Gomphothere Site of Brooks (1968).
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
Santa Fe River 1.--The Santa Fe sites are river bottom deposits in northern
peninsular Florida collected along an approximately 10 km stretch of the Santa Fe
River, which forms the border between Columbia County on the north and
Gilchrist County on the south (Fig. 4, site 1). The Santa Fe River 1 LF, including
sites 1, IA, and 1B, was collected from a bend in the river less than 0.5 km in
length. The bottom of the Santa Fe River in this region produces a mixture of
Blancan and Rancholabrean vertebrates. It is not difficult to separate the faunas of
these two ages because there is very little overlap between them with regard to
chronologically significant mammals. Although the Santa Fe 1B site was
originally thought to be a purely Blancan assemblage (Webb 1974a), a re-
examination of the material from this locality reveals the presence of several
Rancholabrean taxa, including Megalonyx jeffersonii, Holemsina septentrionalis,
Castoroides ohioensis,. and Mammuthus columbi.
The Santa Fe River 1 site has produced one of the most diverse faunas of
large mammals known from the late Blancan of Florida (Table 2), including at
least 22 species. Blancan indicators include Borophagus diversidens, Canis
lepophagus, Nannippus peninsulatus, Equus (Dolichohippus), Platygonus
bicalcaratus, Hemiauchenia blancoensis, and Rhynchotherium praecursor. The
co-occurrence of these taxa with the Neotropical immigrants Dasypus bellus (small
Blancan form), Holmesina floridanus (small Blancan form), Glyptotherium
arizonae, "Glossotherium" chapadmalense, and Titanis walleri indicates a post-
interchange late Blancan fauna. Five species from the Santa Fe River 1 LF are
characteristic of Florida late Blancan and earliest Irvingtonian faunas:
Glyptotherium arizonae, the dwarf Florida form of Megalonyx leptostomus,
Chasmaporthetes ossifragus, Capromeryx arizonensis, and Titanis walleri. Santa
Fe 1 is one of four Florida late Blancan localities in which Smilodon is present
(Berta 1987), a genus that does not appear until the early Irvingtonian in the
western United States (Lundelius et al. 1987).
General discussions of the Santa Fe River fauna can be found in Webb
(1974a), Robertson (1976), Kurt6n and Anderson (1980), and Morgan and
Ridgway (1987). Taxonomic references to Blancan vertebrate taxa from the Santa
Fe River 1 LF include Weaver and Robertson (1967) on the turtle Trachemys
platymarginata; Brodkorb (1963) on Titanis walleri (type locality); Gillette and
Ray (1981) on Glyptotherium arizonae; Robertson (1976), Edmund (1987), and
Hulbert and Morgan (1993) on Holmesina floridanus; Kurten (1965) and Berta
(1987) on Smilodon gracilis; Berta (1981) on Chasmaporthetes ossifragus; and
MacFadden and Waldrop (1980) on Nannippus peninsulatus.
Blancan mammals also have been reported from three additional Santa Fe
River sites. MacFadden and Waldrop (1980) recorded Nannippus phlegon (=N.
peninsulatus) from Santa Fe River 4A, a fauna that also includes Equus
(Dolichohippus), Capromeryx arizonensis, and Platygonus bicalcaratus. In their
review of the Blancan mustelid Trigonictis, Ray et al. (1981) described and figured
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 65
specimens of T. macrodon from Santa Fe River 8A. Berta (1981) reported and
figured a maxilla of the hyaenid Chasmaporthetes ossifragus from Santa Fe River
15.
Haile 15A.-The Haile 15A LF was deposited in a fissure filling in the late
Eocene Crystal River Formation near Newberry, Alachua County (Fig. 4, site 2).
This site is located about 30 m above sea level and about 80 km inland from the
Gulf of Mexico. Along with abundant freshwater and terrestrial vertebrates, the
Haile 15A LF contains a substantial fauna of estuarine and nearshore marine fishes
which led Robertson (1976) to propose that it was deposited during a period of
high sea level. However, Opdyke et al. (1984) provided data suggesting that the
limestone karst region of northern peninsular Florida may have undergone
epeirogenic uplift during the Pleistocene. Despite the possibility of some isostatic
uplift in response to karstification, it still seems highly probable that the Haile 15A
site was deposited during a period when sea level was higher than present.
Biochronological data from the vertebrate fauna suggests an age between 2.5 and
2.0 Ma for Haile 15A. Evidence from both eustatic sea level curves (Haq et al.
1987) and the oxygen isotope record (Shackleton and Opdyke 1977) indicates that
this 0.5 million year time interval was a period of generally low worldwide sea
levels. However, there were several glacial-interglacial cycles during the late
Pliocene, with periods of normal or higher than modern sea level at approximately
2.3 and 2.1 Ma (Shackleton and Opdyke 1977). Which of these presumed late
Pliocene high sea level stands best approximates the age of Haile 15A, and other
Florida late Blancan high sea level faunas discussed below, is problematic at the
present time.
The Haile 15A LF has 23 species of mammals, including four taxa
characteristic of western Blancan faunas, Sigmodon medius, Nannippus
peninsulatus, Equus (Dolichohippus), and the otter Satherium piscinarium. Haile
15A also records the association of typical Blancan mammals with South
American immigrants, thereby indicating a post-interchange late Blancan fauna.
Neotropical immigrants present at Haile 15A include Dasypus bellus (small
Blancan form), Holmesina floridanus (type locality), "Glossotherium"
chapadmalense, and Hydrochaeris holmesi. Haile 15A also records late
occurrences of two species known from the late Hemphillian (earliest Pliocene)
Palmetto Fauna in central Florida, the flying squirrel Cryptopterus webbi (type
locality; Pratt pers. comm.) and the hipparionine horse Cormohipparion emsliei
(Hulbert 1988a). Haile 15A is one of the earliest occurrences of Smilodon gracilis
(see Berta 1987).
Robertson (1976) monographed the Haile 15A mammalian fauna and
Campbell (1976) reviewed the avifauna. Additional taxonomic references on this
fauna include Weaver and Robertson (1967) on Trachemys platymarginata (type
locality), Edmund (1987) and Hulbert and Morgan (1993) on Holmesina
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
floridanus, Berta (1987) on Smilodon gracilis, Martin (1979) on Sigmodon medius,
Kinsey (1974) on Mylohyus floridanus (type locality), and Webb (1974b) on
Hemiauchenia macrocephala.
Macasphalt Shell Pit.--The Macasphalt Shell Pit LF was collected from a
commercial shell mine operated by the APAC (formerly Macasphalt) Corporation,
located 8 km east of Sarasota, Sarasota County (Fig. 4, site 6). The fossil
vertebrates from the Macasphalt Shell Pit are derived from a dark brown to
blackish sandy organic layer up to 1 m in thickness (unit 4 or "black layer" of
Petuch 1982). The vertebrate-bearing unit and the marine shell beds that occur
both below and above it are referred to the Pinecrest Beds (Petuch 1982; Stanley
1986), or the Pinecrest Sand Member of the Tamiami Formation of some authors
(Hunter 1968; DuBar 1974). Waldrop and Wilson (1990) recently renamed this
unit the Fruitville Formation. Measured stratigraphic sections at the Macasphalt
Shell Pit place the vertebrate-bearing unit from 5-8 m below the present ground
surface. This is near or slightly above present sea level based on the current
surface elevation of 7-8 m. Unit 4 contains a rich nearshore marine and estuarine
vertebrate and invertebrate fauna providing strong evidence that, like Haile 15A,
the Macasphalt Shell Pit LF was deposited during a period of relatively high sea
level.
The extraordinarily rich marine molluscan fauna from the Pinecrest Beds
exposed in the Macasphalt (=APAC) Shell Pit has prompted numerous studies by
invertebrate paleontologists (e.g. Petuch 1982; Stanley 1986; Lyons 1991; Scott
and Allmon [eds.] 1992). However, there is by no means complete agreement on
the age of the richly fossiliferous marine shell beds directly underlying and
overlying the primary bone-bearing horizon. Stanley (1986) regarded these beds to
be early Pliocene in age, Lyons (1991) considered them to be middle Pliocene (3.5-
3.0 Ma), while others have suggested a late Pliocene age (Ward 1992; Zullo 1992).
A recent paper on the integrated geochronology of the Macasphalt Shell Pit (Jones
et al. 1991), including analyses of mammalian biochronology, ostracode
biostratigraphy, paleomagentic polarity, and strontium isotope stratigraphy, placed
the age of the vertebrate-bearing unit 4 at Macasphalt between 2.5 and 2.0 Ma.
The Macasphalt vertebrate fauna, composed of well over 100 species,
contains a mixture of marine, estuarine, freshwater, and terrestrial taxa, with the
brackish and freshwater component predominating. The mammalian fauna
consists of 25 species, including many characteristic Blancan forms: dwarf
Megalonyx leptostomus, Trigonictis macrodon, Sigmodon medius, Nannippus
peninsulatus, Equus (Dolichohippus), Platygonus bicalcaratus, Hemiauchenia
blancoensis, and Rhynchotherium cf. R. praecursor. The association of these
species, in particular N. peninsulatus, in the Macasphalt Shell Pit LF with a
diverse suite of Neotropical immigrants, including Dasypus bellus (small Blancan
form), Holmesina floridanus (small Blancan form), "Glossotherium"
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 67
chadpadmalense, and Neochoerus dichroplax, is typical of all well-sampled
Florida late Blancan faunas (Morgan and Ridgway 1987).
Morgan and Ridgway (1987) summarized the Macasphalt Shell Pit LF and
provided a list of the diagnostic mammals. Macasphalt is the type locality of the
hipparionine horse Cormohipparion emsliei described by Hulbert (1988a). Prior to
its discovery at Macasphalt, the youngest record of Cormohipparion was from the
late Hemphillian Bone Valley Formation of central Florida. Morgan (1991)
reported the molossid bat Tadarida cf. T. brasiliensis from this site. Jones et al.
(1991) briefly mentioned the Macasphalt mammalian fauna in their
geochronological discussion of the APAC Shell Pit. Emslie (1992a,b) described
the rich avifauna of over 40 taxa from the Macasphalt Shell Pit.
Kissimmee River.--The Kissimmee River LF is derived primarily from UF
sites Kissimmee River 5 and 6 located south of Ft. Basinger in Okeechobee County
(Fig. 4, site 4). Vertebrate fossils were collected from the Kissimmee River basin
in the 1960s and 1970s during and after a project conducted by the U. S. Army
Corps of Engineers to channelize the Kissimmee River. The extensive spoil piles
produced by the dredging of the river channel have long been known to produce a
rich Pliocene molluscan fauna representative of the Pinecrest Beds. Certain
Pinecrest localities along the Kissimmee River have produced a fairly diverse
Blancan vertebrate fauna as well. Although most of the Kissimmee River LF was
collected from spoil piles, primarily by the late Howard Converse, the fossils seem
to constitute a uniform late Blancan fauna that compares closely to other Florida
samples of this age. The close association of this vertebrate fauna with marine
molluscs of the Pinecrest Beds is similar to the situation encountered in the
Macasphalt Shell Pit, St. Petersburg Times, Brighton Canal, Lehigh Acres, and
Acline Shell Pit sites. The Kissimmee River LF consists of a mixture of terrestrial,
freshwater, and marine taxa, suggesting that the site formed in a shallow marine
depositional environment. The exact elevation of the vertebrate-bearing unit is
unknown; however, the current surface elevation of about 10 m in this region
strongly suggests that the fauna was deposited during a period of high sea level.
There are no published references to the Kissimmee River LF, which consists
of at least 24 species of mammals. Characteristic Blancan mammals from the
Kissimmee River LF include the dwarf Megalonyx leptostomus, Nannippus
peninsulatus, Equus (Dolichohippus), and Platygonus bicalcaratus. Seven species
of Neotropical immigrants are present in this fauna, the capybara Neochoerus
dichroplax and six xenarthrans, Dasypus bellus (small Blancan form), Holmesina
floridanus (small Blancan form), Glyptotherium arizonae, Pachyarmatherium
leiseyi, "Glossotherium" chapadmalense, and Eremotherium sp. The oldest
previous record of Eremotherium was from the earliest Irvingtonian Inglis 1A LF
(Webb 1974a; Webb and Wilkins 1984). There are now four late Blancan records
of Eremotherium, including the Kissimmee River, Brighton Canal, and Lehigh
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
Acres faunas in southern Florida, and the extensive sample from the latest Blancan
Haile 7C LF in northern peninsular Florida. The Kissimmee fossils of
Pachyarmatherium leiseyi constitute the oldest record of this species in Florida.
All other Florida occurrences of P. leiseyi are from early Irvingtonian faunas. The
two osteoderms of this species from the Kissimmee River 6 site were collected from
spoil piles, as was the remainder of the fauna, and thus there is a possibility of
faunal mixing. However, scutes of P. leiseyi are also known from a Blancan fauna
in South Carolina in association with Nannippus (J. Knight, pers. comm.). The
association of typical Blancan mammals with a large suite of Neotropical
immigrants in the Kissimmee River fauna places the age of this site as post-
Interchange between 2.5 and 2.0 Ma.
Haile 7C.--Vertebrate fossils from the recently discovered Haile 7C LF are
preserved in clays and fine-grained sands deposited in a solution feature in the
marine Eocene Crystal River Formation in a commercial limestone mine near
Newberry, Alachua County (Fig. 4, site 11). The site probably represents either a
sinkhole pond or spring and is numerically dominated by freshwater taxa,
including anurans, salamanders, the turtles Trachemys platymarginata (=T.
idahoensis of Jackson 1988) and Chelydra n. sp., and the alligator Alligator
mississippiensis. Many of the turtles are represented by complete shells, often
including the articulated skulls and limbs. Likewise, several of the mammals from
Haile 7C are known from associated or articulated skeletons. The Haile 7C LF
includes 16 taxa of mammals. The large mammal fauna is dominated by
Eremotherium, Holmesina, and a new species of Tapirus (Hulbert in prep.). Small
rodents, carnivores, and most ungulates are very rare in the site. Only two papers
have discussed the Haile 7C LF (Hulbert et al. 1989; Hulbert and Morgan 1993).
The age of Haile 7C is still somewhat uncertain because of the limited
diversity of the mammalian fauna. The mammals present are indicative of either a
latest Blancan or earliest Irvingtonian age, probably dating between 2.2 and 1.7
Ma. A late Blancan age assignment is difficult to substantiate because there are no
strictly Blancan mammals present in the fauna. Although the Haile 15A LF is
located within 1 km of Haile 7C, the vertebrate faunas from these two sites differ
considerably. Haile 7C not only lacks the diagnostic Blancan genera found at
Haile 15A, but it also contains no evidence of the marine taxa present in the latter
fauna, indicating that Haile 7C was not deposited during the same high sea level
stand.
Haile 7C is clearly younger than 2.5 Ma, as it has six genera of Neotropical
immigrants, Dasypus, Holmesina, Glyptotherium, Eremotherium, Neochoerus, and
Erethizon. Eremotherium and Erethizon were previously unknown from the
Blancan of Florida, although we also report Eremotherium from three additional
late Blancan faunas in the southern half of the state. Erethizon occurs in several
late Blancan faunas in the western United States (Harrison 1976; Frazier 1981).
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 69
Although turtles are not often regarded as particularly useful index fossils, the
large emydid Trachemys platymarginata is common in most Florida late Blancan
faunas. It is replaced by the smaller T. scripta in the early Irvingtonian. The Haile
7C Trachemys is essentially identical to the type series of T. platymarginata from
Haile 15A. Furthermore, Haile 7C and Haile 15A are the only two known
localities for a large undescribed species of Chelydra.
Lundelius et al. (1987) noted that there is not a sharp faunal break between
the Blancan and Irvingtonian, but rather they documented a gradual transition
between these two NALMA that occurred between about 2.0 and 1.7 Ma. Thus, it
is not surprising that some faunas from this time period are difficult to place in one
or the other of these two ages. The absence of typical Blancan genera, such as
Borophagus, Nannippus, and Equus (Dolichohippus) from Haile 7C argues for an
early Irvingtonian age; however, carnivores and horses are virtually absent in this
fauna. Perhaps the majority of Florida late Blancan faunas occur in the time
period immediately following the onset of the Great American Interchange
between 2.5 and 2.2 Ma. If so, there might be a significant period of time, perhaps
as long as half a million years between about 2.2 and 1.7 Ma, after the extinction
of Nannippus and most other typical Blancan taxa, but before the appearance of
most diagnostic Irvingtonian taxa. We suggest that the Haile 7C LF dates to the
very end of the Blancan during the early portion of the transitional period between
the Blancan and Irvingtonian NALMA. Regardless of its placement in the land
mammal biochronology, Haile 7C is clearly latest Pliocene in age (younger than
2.5 Ma and older than 1.6 Ma). The Borchers LF in Kansas is a well known fauna
that also appears to fall in this transitional time period, and has likewise been
regarded as either late Blancan or early Irvingtonian (Lundelius et al. 1987).
Other Florida late Blancan faunas.--Among the Florida Blancan localities
reviewed by Morgan and Ridgway (1987) are several important sites, including St.
Petersburg Times, Bass Point Waterway, El Jobean, and Acline Shell Pit, not listed
in Table 2 because they have fewer than ten diagnostic species of mammals. We
briefly discuss several of these smaller sites, as well as the previously unpublished
Brighton Canal and Lehigh Acres local faunas. Most of these sites possess critical
Blancan taxa or occur in stratigraphic superposition with well known marine units.
The St. Petersburg Times Site is located about 10 km inland from the Gulf of
Mexico in downtown St. Petersburg, Pinellas County (Fig. 4, site 3). The
vertebrate fossils constituting the St. Petersburg Times LF were recovered from a
black sandy organic layer near the bottom of a borrow pit. Underlying the bone-
bearing layer was a semi-indurated marine shell bed tentatively referred to the
Pinecrest Beds based on its molluscan fauna. The vertebrate unit was about 12-15
m below the surface, and thus was near present sea level based on current surface
elevations. The stratigraphic sequence and the absence of marine taxa in the St.
Petersburg Times Site both suggest deposition during a period of low sea level.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 37, PT. I, NO. 1
The vertebrate fauna from the St. Petersburg Times Site consists of 30 species and
is dominated by freshwater forms, especially sirens, water snakes, turtles, and
aquatic birds. Nine species of terrestrial mammals occur in the St. Petersburg
Times LF (Morgan and Ridgway 1987), four of which are characteristic of Florida
late Blancan faunas, Holmesina floridanus, Glyptotherium arizonae, small
Megalonyx leptostomus, and Nannippus peninsulatus. Morgan and Ridgway
(1987) also reviewed the lower vertebrates in the St. Petersburg Times LF. Becker
(1987) and Emslie (1992a) discussed the avifauna.
The Lehigh Acres LF is here designated for a vertebrate fauna collected by
Thomas Missimer from the Lehigh Corporation Pits south of Alva, Lee County
(Fig. 4, site 10). The fossils were derived from a semi-indurated calcareous marl
containing a mixture of marine and freshwater molluscs characteristic of the
Pinecrest Beds. Although collected primarily from spoil piles, there is no
indication that the fauna is mixed. The Lehigh Acres LF is composed of 22 species
of vertebrates, including nearshore marine, freshwater, and terrestrial species.
There are nine species of terrestrial mammals in the fauna, including four typical
Blancan taxa, "Glossotherium" chapadmalense, Nannippus peninsulatus, Equus
(Dolichohippus), and Platygonus bicalcaratus. The association of Nannippus with
"Glossotherium" and Eremotherium characterizes a post-Interchange late Blancan
fauna. The Eremotherium from Lehigh Acres represents one of the few Blancan
records for this genus.
The Brighton Canal LF is a small, previously undescribed late Blancan
vertebrate fauna collected by Muriel Hunter from spoil piles along the Brighton
Canal near Brighton, Highlands County (Fig. 4, site 5). The predominant
stratigraphic unit in this region of the canal is the Pinecrest Beds, based on the
molluscan fauna collected from the same spoil piles that produced the vertebrates.
The Brighton Canal LF includes nine species of terrestrial mammals, four of which
are characteristic of Florida late Blancan faunas, Glyptotherium arizonae,
Nannippus peninsulatus, Equus (Dolichohippus), and Mylohyus floridanus. The
South American immigrant Eremotherium is also present.
The Bass Point Waterway 1 LF, Sarasota County (Fig. 4, site 7), includes
Holmesina floridanus, Smilodon gracilis, Nannippus peninsulatus, and Equus
(Dolichohippus). The nearby El Jobean LF in Charlotte County (Fig. 4, site 8) also
contains Holmesina floridanus, Smilodon gracilis, and Nannippus peninsulatus.
These two sites were first mentioned by Churcher (1984) is his review of
Ischyrosmilus gracilis (=Smilodon gracilis following Berta 1987) from Florida.
The association of Nannippus and Holmesina is diagnostic of most Florida post-
Interchange late Blancan faunas. The Bass Point Waterway and El Jobean sites are
among the earliest records for Smilodon gracilis.
The Acline Shell Pit, located near Acline in Charlotte County (Fig. 4, site 9),
has produced a small Blancan fauna from marine shell beds referred to the
Pinecrest Beds (Olsson and Petit 1964). The most diagnostic mammals in the
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 71
Acline fauna are "Glossotherium" chapadmalense and Cuvieronius tropicus.
Morgan and Ridgway (1987) mentioned four additional late Blancan vertebrate
faunas from southwestern Florida not discussed here, including Sommers Pit and
Northport in Sarasota County, Port Charlotte in Charlotte County, and Mule Pen
Quarry in Collier County.
IRVINGTONIAN
Webb (1974a) and Kurtdn and Anderson (1980).listed only four Irvingtonian
faunas from Florida: Inglis 1A, Punta Gorda, Pool Branch, and Coleman 2A.
Inglis 1A (Klein 1971; Webb 1974a; Webb and Wilkins 1984) and Coleman 2A
(Martin 1974) both have diverse mammalian faunas composed of more than 40
species, whereas Punta Gorda and Pool Branch have much smaller faunas
consisting primarily of large mammals. A fifth Florida Irvingtonian site, Haile
16A, has been mentioned numerous times in taxonomic papers, but no published
mammalian faunal list is available.
Earliest Irvingtonian faunas (2.0-1.6 Ma), including Inglis IA and De Soto
Shell Pit, contain many taxa typical of late Blancan faunas and lack Mammuthus
which does not arrive in North America until after 1.6 Ma. Late early Irvingtonian
faunas (1.6-1.0), such as Leisey, usually have Mammuthus hayi, lack most Blancan
holdovers, and possess a number of taxa also typical of middle Irvingtonian faunas.
We use the term early Irvingtonian in the context of Lundelius et al. (1987) only
when discussing faunas of both the earliest and late early Irvingtonian (i.e., 2.0 to
1.0 Ma). The middle Irvingtonian encompasses the interval from 1.0 to 0.6 Ma,
and the late Irvingtonian is from 0.6 to 0.3 Ma. The reasons for locating the
boundaries at these dates are presented above.
Inglis 1A.--The Inglis IA LF was collected from a solution cavity in the
Eocene Inglis Formation along the now-defunct Cross Florida Barge Canal in
Citrus County (Fig. 4, site 12). The Inglis 1A site occurs in a karst solution feature
that is as much as 5 m below present mean sea level. However, the fauna totally
lacks marine forms, suggesting that it was deposited during a period of low sea
level. Richmond and Fullerton (1986) recorded a glacial interval from 2.01 to 1.87
Ma (corresponding to marine oxygen isotope stage 40); this age is in agreement
with that provided by vertebrate biochronology (see below). Correlation of Inglis
1A with a glacial period is supported by certain taxa in the vertebrate fauna that
suggest cooler and/or more arid conditions (Carr 1980; Meylan 1982; Webb and
Wilkins 1984; Morgan 1991).
Our list of mammals from Inglis IA (Table 2) includes several additions and
corrections to the published mammalian fauna for this site (Webb 1974a; Webb
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
and Wilkins 1984). The Inglis mammal fauna consists of 53 species, with the best
represented forms being xenarthrans, carnivores, ungulates, and small mammals.
Inglis IA possesses several genera listed by Lundelius et al. (1987) as
characteristic of the Irvingtonian, including Smilodon, Lepus, and Equus (s.s.), and
lacks Borophagus, Nannippus, Equus (Dolichohippus), Rhynchotherium, and other
genera restricted to the Blancan. However, Inglis IA does share a number of taxa
with Florida late Blancan faunas, including Glyptotherium arizonae, the dwarf
Florida form of Megalonyx leptostomus, Trigonictis macrodon, Chasmaporthetes
ossifragus, Geomys propinetis, Sylvilagus webbi, Capromeryx arizonensis, and the
phororhacid bird Titanis walleri. The presence of both Blancan holdovers and
characteristic Irvingtonian taxa, as well as the absence of Mammuthus, indicates an
earliest Irvingtonian age (2.0-1.6 Ma) for Inglis 1A. The mammals from Inglis 1A
compare closely to the fauna from Curtis Ranch in Arizona (Lindsay and Tessman
1974). Diagnostic mammals shared by these two faunas include Glyptotherium
arizonae, Canis edwardii, Sigmodon curtisi, Ondatra idahoensis, and Capromeryx
arizonensis.
The geology and stratigraphy of the Inglis deposit were discussed by Klein
(1971), who also reviewed the carnivores and ungulates. The birds (Carr 1980)
and squamate reptiles (Meylan 1982) of Inglis 1A have been studied in detail.
There are numerous taxonomic references on mammals from the Inglis 1A LF,
including Gillette and Ray (1981) on Glyptotherium arizonae, McDonald (1977)
on Megalonyx leptostomus, Morgan et al. (1988) on Desmodus archaeodaptes,
Morgan (1991) on the chiropteran fauna, Ray et al. (1981) on Trigonictis
macrodon, Berta (1981) on Chasmaporthetes ossifragus, Berta (1987) on Smilodon
gracilis, Van Valkenburgh et al. (1990) on Miracinonyx inexpectatus, Frazier
(1981) on Erethizon kleini (type locality), Ahearn (1981) on Hydrochaeris holmesi,
Wilkins (1984) on Geomys propinetis (type locality), Martin (1979) on Sigmodon
curtisi, and White (1991) on Sylvilagus webbi (type locality).
De Soto Shell Pit.--The De Soto Shell Pit LF is here designated for the
vertebrate fauna collected from organic layers in the Caloosahatchee Formation
exposed in three commercial shell mines south of Arcadia in De Soto County (Fig.
4, site 13). This is the first published reference to the De Soto Shell Pit LF. The
mammalian fauna from the De Soto Shell Pit is composed of 32 species (see Table
2), 25 of which are in common with Inglis lA. The biochronologically significant
species of mammals shared by the De Soto and Inglis faunas are: Glyptotherium
arizonae, small Megalonyx leptostomus, Canis edwardii, Trigonictis macrodon,
Chasmaporthetes ossifragus, Geomys propinetis, Ondatra idahoensis, Sigmodon
curtisi, Capromeryx arizonensis, an undescribed dwarf species of Hemiauchenia,
and an undescribed species of Tapirus. Like Inglis, the De Soto fauna lacks genera
restricted to the Blancan, but has a number of taxa common to both Florida late
Blancan and earliest Irvingtonian faunas, has Equus (s.s.) rather than Equus
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 73
(Dolichohippus), and lacks Mammuthus. The common De Soto emydid turtle is
Trachemys platymarginata, a species limited to the late Blancan and earliest
Irvingtonian in Florida. All of these factors taken together strongly indicate an
earliest Irvingtonian age. The earliest occurrence of the arvicoline rodent
Atopomys appears to be in the De Soto Shell Pit LF. This genus is also known
from the late early Irvingtonian Haile 16A LF. Elsewhere in North America
Atopomys is not known prior to the middle Irvingtonian (Winkler and Grady
1990).
The De Soto Shell Pit LF contains a number of nearshore marine taxa,
including sharks, bony fishes, and the monk seal Monachus, indicating that the site
probably formed near sea level. Therefore, the Inglis IA and De Soto Shell Pit
local faunas are not quite the same age. Inglis was deposited below present sea
level during a glacial interval, while De Soto was deposited about 5-10 m above
modern sea level during an interglacial. Based on the oxygen isotope record
(Shackleton and Opdyke 1977), the only high sea level stand during the latest
Pliocene occurred about 1.8 Ma. Independent evidence for the age of the De Soto
Shell Pit LF comes from the rich marine molluscan fauna of the Caloosahatchee
Formation that occurs in direct stratigraphic superposition both below and above
the units containing the vertebrate fauna. Lyons (1991) considered the entire
molluscan fauna of the Caloosahatchee Formation to be late Pliocene in age (older
than 1.8 Ma). Corals collected near "the top of the Caloosahatchee" Formation
along the Caloosahatchee River have yielded Helium/Uranium dates of 1.89 to 1.78
Ma (Bender 1973). These dates are consistent with the earliest Irvingtonian (2.0-
1.6 Ma) age suggested by the De Soto Shell Pit vertebrate fauna.
Haile 16A.--The Haile 16A LF is located in the Haile Quarry complex
northeast of Newberry, Alachua County (Fig. 4, site 15). The fossils were derived
from massive dark, silty clays filling a large fissure formed in marine limestones
referred to the Eocene Crystal River Formation. The depth, areal extent, and
stratigraphy of the Haile 16A locality could not be determined since the site was
destroyed by mining operations before it could be properly excavated. Although all
fossils from this site were obtained by collecting and screenwashing spoil piles,
there is no indication that the fauna is mixed. Like many other northern Florida
karst deposits, Haile 16A has an unbalanced mammal fauna. The most abundant
large mammals are the ground sloths Eremotherium, Megalonyx, and
Paramylodon, and the tremarctine bear Arctodus pristinus. Ungulates are
uncommon in this fauna and proboscideans are absent. The microvertebrate fauna
from Haile 16A is extremely abundant and taxonomically diverse, including large
samples of insectivores, bats, and rodents, as well as birds, snakes, lizards, and
frogs.
No faunal list from Haile 16A has been published, but at least 33 species of
mammals are known to be present (Table 2). Because the small mammals from
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
this site have not been completely studied, the actual number of species present is
probably somewhat higher. Haile 16A has been regarded as middle Irvingtonian
by most previous authors (Frazier 1981; Morgan et al. 1988; Winkler and Grady
1990; Morgan 1991), although Lundelius et al. (1987) questionably placed this
fauna in the late Irvingtonian. A detailed analysis of the mammalian fauna from
Haile 16A suggests that a late early Irvingtonian age is more likely. The
Holmesina from Haile 16A is similar in size to individuals of H. floridanus from
Inglis 1A and Leisey, and is distinctly smaller than specimens referred to H.
septentrionalis from the middle Irvingtonian McLeod Limerock Mine (Hulbert and
Morgan 1993). The largest available sample of the newly described armadillo
Pachyarmatherium (Downing and White this volume) is actually from Haile 16A
not Leisey. It appears to be restricted to late Blancan and early Irvingtonian faunas
in Florida. The cotton rat Sigmodon libitinus is known from Haile 16A (type
locality), as well as the early Irvingtonian Leisey Shell Pit and Payne Creek Mine
local faunas. Other biochronologically diagnostic species shared by Haile 16A and
Leisey Shell Pit include: Megalonyx wheatleyi; Erethizon dorsatum; undescribed
species of Podomys, Pedomys, and Synaptomys; Tapirus haysii; Equus "fraternus;"
and Palaeolama mirifica. All of these species are absent from Florida earliest
Irvingtonian faunas, although several do occur in middle Irvingtonian and younger
sites. Although its biochronological significance is unknown, the only known
Florida occurrence of the zapodid rodent Zapus is from Haile 16A.
Three species of mammals from Haile 16A indicate that this site is slightly
older than Leisey: Trigonictis cf. T. cookii, Geomys propinetis, and Sylvilagus
webbi. The mustelid Trigonictis has been considered a Blancan indicator in
western faunas (e.g. Kurtdn and Anderson 1980). However, the presence of T.
cookii at Haile 16A and T. macrodon in the Inglis IA and De Soto Shell Pit local
faunas (Ray et al. 1981) confirms that this genus survived into the early
Irvingtonian in eastern North America. With the exception of Haile 16A,
Sylvilagus webbi and Geomys propinetis are both restricted to Florida late Blancan
and earliest Irvingtonian faunas. Winkler and Grady (1990) regarded the
arvicoline rodent Atopomys as characteristic of middle Irvingtonian faunas. The
presence of Atopomys in the early Irvingtonian Haile 16A and De Soto Shell Pit
local faunas strongly indicates that this rare genus evolved considerably earlier
than previously thought. Although Haile 16A shares several taxa with Florida late
Blancan and earliest Irvingtonian faunas, on the whole this fauna has more species
in common with Leisey and other late early Irvingtonian sites. The Haile 16A LF
probably derives from the early half of the late early Irvingtonian (1.6 to 1.3 Ma),
and is intermediate in age between earliest Irvingtonian faunas such as Inglis 1A
and late early Irvingtonian faunas like Leisey.
Many taxonomic publications have mentioned mammals from Haile 16A
including Edmund (1987) and Hulbert and Morgan (1993) on Holmesina,
Downing and White (this volume) on Pachyarmatherium, McDonald (1977) on
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 75
Megalonyx wheatleyi, Morgan et al. (1988) and Morgan (1991) on Desmodus
archaeodaptes, Emslie (this volume) on Arctodus pristinus, Ray et al. (1981) on
Trigonictis cf. T. cookii, Wilkins (1984) on Geomys propinetis, Frazier (1981) on
Erethizon dorsatum, Martin (1979) on Sigmodon libitinus, Winkler and Grady
(1990) on Atopomys salvelinus, and Hulbert (this volume) on Tapirus haysii and
Equus "fraternus."
Pool Branch.--The Pool Branch LF was collected from the banks of Pool
Branch, a tributary of the Peace River in Polk County (Fig. 4, site 21). The Pool
Branch mammalian fauna was first listed by Webb (1974a) and later updated by
McDonald (1985). Pool Branch was one of only four Irvingtonian faunas
discussed by Webb (1974a) and Kurten and Anderson (1980). Papers mentioning
taxa from this fauna include Webb (1974b) on Hemiauchenia macrocephala,
McDonald (1985) on Nothrotheriops, and Hulbert (this volume) on Equus and
Tapirus.
Twelve species of mammals from Pool Branch are listed in Table 2.
Carnivores are absent and small mammals are represented only by two partial
rodent teeth, both referable to the arvicoline Ondatra annectens. This species of
muskrat also occurs at Leisey and is characteristic of early and middle Irvingtonian
faunas. The giant tapir, Tapirus haysii, is restricted to late early and middle
Irvingtonian faunas in Florida (Hulbert this volume). McDonald (1985) reported
and figured a number of postcranial elements of the ground sloth Nothrotheriops
from Pool Branch, the first record of this genus from eastern North America. A
much larger sample of this sloth, now referred to N. texanus, has been recovered
from the Leisey Shell Pit LF (McDonald this volume). Another similarity between
Leisey and Pool Branch is the presence of an undescribed species of hemionine
horse, Equus (Hemionus) sp. (Hulbert this volume). All 12 species of mammals
listed from Pool Branch in Table 2 also occur at Leisey strongly indicating a late
early Irvingtonian age.
Payne Creek Mine.--An interesting and mostly unstudied vertebrate fauna
has been collected from the Payne Creek Mine, located in Polk County about 10
km southwest of Pool Branch (Fig. 4, site 20). Steadman (1984) reported 10 avian
taxa from the Payne Creek Mine LF, including an extralimital record of the least
grebe, Podiceps dominicus. The mammalian fauna from the Payne Creek Mine
consists of at least 22 species, including several typical early Irvingtonian
xenarthrans and rodents (Table 2). There are four diagnostic species of
xenarthrans from the Payne Creek Mine, Holmesina floridanus,
Pachyarmatherium leiseyi, Megalonyx wheatleyi, and Eremotherium n. sp. The
stage of evolution of the H. floridanus osteoderms from Payne Creek is suggestive
of an early Irvingtonian age. The new species of Eremotherium is characteristic of
Florida late Blancan and early Irvingtonian faunas (De Iuliis and Cartelle in prep).
Megalonyx wheatleyi appears to be restricted to late early and middle Irvingtonian
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
faunas. This fauna also contains the typical Florida Irvingtonian tremarctine bear,
Arctodus pristinus. The Payne Creek Mine LF has a diverse assemblage of small
mammals, including the cotton rat Sigmodon libitinus and two species of
arvicolines, the muskrat Ondatra annectens and an undescribed species of the vole
Pedomys (see Morgan and White this volume). All three of these species also are
present in the Leisey Shell Pit and Haile 16A local faunas. The similarity of the
Payne Creek Mine fauna to that of Leisey strongly indicates a late early
Irvingtonian age (probably between 1.3 and 1.0 Ma).
Crystal River Power Plant.--The Crystal River Power Plant LF was
discovered during the construction of the Crystal River Nuclear Power Plant,
located just inland from the Gulf of Mexico about 6 km west of Red Level in Citrus
County (Fig. 4, site 17). Vertebrate fossils were encountered about 10 m below
present sea level beneath an extensive marine shell bed. The marine molluscan
fauna from this shell bed is very similar to that from Leisey, and is probably
equivalent to the Bermont Formation. The vertebrate fauna contains a large
sample of freshwater fish, but no marine forms are present. The absence of marine
vertebrates, coupled with the fact that the site was discovered well below present
sea level, provides strong evidence that the Crystal River Power Plant LF was
deposited during a glacial interval.
The mammalian fauna from the Crystal River Power Plant LF is composed of
16 species (Table 2). The presence of Lutra and Castoroides confirms a post-
Blancan age for this site as both genera first appear in the Irvingtonian. Species
characteristic of Florida Irvingtonian faunas include Arctodus pristinus, a large
species of Procyon, Tapirus haysii, and Equus "leidyi." Several taxa from Crystal
River further restrict the age of this fauna within the Irvingtonian. The presence of
Smilodon gracilis and Tapirus haysii rules out a late Irvingtonian age. One of the
most diagnostic mammals in the Crystal River Power Plant LF is the canid Canis
edwardii, a species restricted to the early Irvingtonian. The stage of evolution
represented by the small sample of Holmesina osteoderms from Crystal River is
also consistent with an early Irvingtonian age (Hulbert and Morgan 1993).
Megalonyx wheatleyi is now known from four Florida late early Irvingtonian
faunas, including Crystal River. Every species of mammal listed in Table 2 from
from Crystal River occurs at Leisey as well, suggesting that these two faunas are
very similar in age.
Rigby Shell Pit.--The Rigby Shell Pit LF was collected from a commercial
shell mine near Nokomis, Sarasota County (Fig. 4, site 22). The vertebrate fossils
were derived from a unit within marine shell beds of the Bermont Formation and
consist of a mixture of marine, freshwater, and terrestrial species. The most
abundant large vertebrates in the fauna are Hemiauchenia macrocephala and
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 77
Equus "leidyi." The Rigby Shell Pit fauna was briefly mentioned by Hulbert and
Morgan (1989), but no published faunal list is available.
All 12 species of mammals identified from Rigby are also known from Leisey,
five of which are indicative of an Irvingtonian age. Arctodus pristinus and Equus
"leidyi" are typical of many Florida Irvingtonian faunas, but do not provide a more
precise age refinement. The most age diagnostic mammal from Rigby is Canis
edwardii, which is restricted to early Irvingtonian sites in Florida. A distal
humerus of Holmesinafloridanus from Rigby is distinctly larger than comparable
specimens from Leisey, suggesting that the Rigby Shell Pit may be slightly younger
(Hulbert and Morgan 1993). Except for the larger size of Holmesina, the
mammalian fauna from Rigby is indistinguishable from that of the Leisey Shell Pit
and is probably late early Irvingtonian in age as well.
Haile 21A.-The Haile 21A LF is located in the Haile Quarry complex
northeast of Newberry in Alachua County (Fig. 4, site 16). The site consisted of
clays, sands, and limestone breccias filling a karst solution cavity in the marine
late Eocene Crystal River Formation. This locality has been completely destroyed
by mining operations. Morgan et al. (1988) and Morgan (1991) briefly discussed
the Haile 21A site and its vertebrate fauna. The mammalian fauna from Haile 21A
consists of 20 species, but is overwhelmingly dominated by the large peccary,
Platygonus vetus (Wright this volume). Haile 21A is the type locality of the
primitive vampire bat, Desmodus archaeodaptes (Morgan et al. 1988), and also
has a large sample of the vespertilionid bat Myotis austroriparius. In contrast to
most other north Florida Irvingtonian karst deposits, small mammals are scarce in
the Haile 21A LF, with the exception of bats.
Tapirus haysii, Equus "leidyi," Platygonus vetus, and a large species of
Procyon occur in the Haile 21A LF, all of which are limited to the Irvingtonian in
Florida. Furthermore, Tapirus haysii, has not yet been recorded from either
earliest Irvingtonian or late Irvingtonian faunas (Hulbert this volume), thereby
limiting the age of Haile 21A to late early or middle Irvingtonian. The presence of
Canis edwardii in the Haile 21A LF (Berta this volume) strongly suggests an early
Irvingtonian age. Haile 21A is the only other Florida fauna besides Leisey in
which both C. edwardii and the larger C. armbrusteri are present. The association
of these two canids seems to be typical only of late early Irvingtonian faunas, at
least in Florida.
Specimens of Smilodon gracilis from Haile 21A are within the size range of
the Leisey sample, but are smaller than specimens of S. gracilis from the middle
Irvingtonian McLeod Limerock Mine (Berta 1987; this volume). The skeleton of a
very large and apparently undescribed species of the sabercat Homotherium was
recovered from Haile 21A. This large Homotherium, although very rare
everywhere except Haile 21A, has also been recorded from three other early
Irvingtonian faunas, Leisey, Inglis 1A, and Haile 16A. Besides Haile 21A, the
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
only other localities for Desmodus archaeodaptes are Inglis lA and Haile 16A
(Morgan et al. 1988; Morgan 1991). Haile 21A shares many diagnostic
mammalian taxa with Leisey and likewise probably correlates with the latter part
of the early Irvingtonian (1.6 and 1.0 Ma).
McLeod Limerock Mine.--The McLeod LF was recovered from the McLeod
Limerock Mine located 3.3 km north of Williston, Levy County (Fig. 4, site 24).
McLeod was excavated in 1941 by a Frick field party led by Ted Galusha, and the
material from this site is now housed in the American Museum of Natural History.
The fossils were recovered from a fine to coarse grayish clayey sand deposited in a
solution feature in the marine Eocene Crystal River Formation. This site has long
since been destroyed by mining operations (Frazier 1977).
No mammalian faunal list has been published for the McLeod LF. Fourteen
diagnostic species of mammals known to be present in the McLeod LF are listed in
Table 2, although this list is probably incomplete. The occurrence of Smilodon
gracilis at McLeod indicates that this site is middle Irvingtonian or older (Berta
1987). The larger species S. fatalis (=S. populator) appears in the late
Irvingtonian. S. gracilis specimens from McLeod are larger and more advanced
than early Irvingtonian samples of this sabercat from Inglis 1A and Leisey Shell
Pit, suggesting a younger age for McLeod (Berta 1987). The presence of Tapirus
haysii also indicates a pre-late Irvingtonian age for McLeod, as this species is
restricted to late early and middle Irvingtonian faunas in Florida (Hulbert this
volume). The presence of Canis armbrusteri at McLeod rules out an earliest
Irvingtonian age, as the oldest Florida records of this wolf is from the late early
Irvingtonian Leisey Shell Pit. This canid survives until the late Irvingtonian where
it is well represented at Coleman 2A (Martin 1974; Berta this volume).
Megalonyx wheatleyi, represented at McLeod by a nearly complete skeleton
(McDonald 1977), is most typical of middle Irvingtonian faunas, although it does
occur in several Florida late early Irvingtonian sites.
One of the more diagnostic species present at McLeod is the extinct round-
tailed muskrat, Neofiber leonardi. This species has been reported principally from
late Irvingtonian faunas correlative with Coleman 2A (see below), including
Rezabek (type locality) and Kanopolis in Kansas and Slaton in Texas (Frazier
1977). McLeod represents one of the earliest records of the large pampathere,
Holmesina septentrionalis, a species that first appears in the middle Irvingtonian
and survives until the end of the Pleistocene (Hulbert and Morgan 1993). One
species from McLeod, Neofiber leonardi, appears to be restricted to the late
Irvingtonian elsewhere, whereas two other species, Smilodon gracilis and Tapirus
haysii, are otherwise unknown after the middle Irvingtonian. The Concurrent
Range Zone of these three species in Florida constrains the age of McLeod to either
late middle Irvingtonian or early late Irvingtonian, probably between 0.7 and 0.5
Ma.
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 79
Studies on mammals from the McLeod LF include Frazier (1977) on Neofiber
leonardi, McDonald (1977) on Megalonyx wheatleyi, Edmund (1987) and Hulbert
and Morgan (1993) on Holmesina septentrionalis, Berta (1987) on Smilodon
gracilis, Berta (this volume) on Canis armbrusteri, Seymour (1993) on Panthera
onca, and Emslie (this volume) on Arctodus pristinus.
Coleman 2A.--The Coleman 2A LF was collected from sands and clays
deposited in a karst solution feature in the marine late Eocene Crystal River
Formation in Sumter County in the north-central portion of the peninsula (Fig. 4,
site 25). Like most other northern Florida karst deposits, the Coleman 2A Site was
discovered in a commercial limestone mine and was subsequently destroyed by
mining operations. Prior to Leisey, the Coleman 2A LF was the only Irvingtonian
mammalian fauna from Florida to be studied comprehensively (Martin 1974).
Martin (1974:90) noted that the Coleman 2A LF, "...stands near the
Irvingtonian-Rancholabrean boundary." He considered this fauna to be latest
Irvingtonian in age because it lacks Bison and includes several taxa of mammals
unknown from Florida Rancholabrean faunas. Coleman 2A is the only well known
Florida vertebrate fauna that is late Irvingtonian in age. At least three species of
mammals from Coleman 2A indicate an Irvingtonian age, Arctodus pristinus,
Canis armbrusteri, and Platygonus cumberlandensis. A. pristinus occurs in most
well sampled Florida Irvingtonian faunas, but is replaced in the Rancholabrean by
the smaller tremarctine bear Tremarctos floridanus. C. armbrusteri ranges from
the late early Irvingtonian through the late Irvingtonian, but is replaced in the early
Rancholabrean by C. dirus. The large peccary P. cumberlandensis is known
primarily from middle and late Irvingtonian faunas, although Martin (1974) also
reported this species the early Rancholabrean Haile 7A LF. Wright (this volume)
refers Florida early Irvingtonian Platygonus to the species, P. vetus, whereas
Rancholabrean faunas are generally characterized by the smaller P. compressus.
Coleman 2A represents the oldest known record of two Rancholabrean (and
living) species, Didelphis virginiana and Neofiber alleni. All other North
American fossil occurrences of D. virginiana are from the Rancholabrean. The
extinct muskrat N. leonardi occurs in several western faunas considered to be
correlatives of Coleman 2A (Frazier 1977). The presence of Didelphis and N.
alleni in Coleman 2A indicate a latest Irvingtonian age for this fauna. The tapir at
Coleman 2A is Tapirus veroensis, the Rancholabrean species, and not T. haysii
which is characteristic of late early and middle Irvingtonian faunas in Florida.
Another link to the Rancholabrean is the cotton rat Sigmodon bakeri, originally
described from Coleman 2A, but subsequently recognized from at least three
Florida early Rancholabrean sites (Martin 1974). The large vole Pitymys aratai is
presently known only from Coleman 2A (Martin 1974). Coleman 2A represents
the oldest record of the Florida mouse, Podomys floridanus. The association of
taxa restricted to the Irvingtonian (Arctodus pristinus and Canis armbrusteri) with
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
species typical of Rancholabrean faunas supports Martin's (1974) contention that
Coleman 2A is latest Irvingtonian in age, probably between 0.5 and 0.3 Ma.
In addition to Martin's (1974) review of the entire mammalian fauna, other
studies on mammals from Coleman 2A include Webb (1974b) on Palaeolama,
Martin (1979) on Sigmodon bakeri (type locality), Frazier (1981) on Erethizon
dorsatum, Wilkins (1984) on Geomys pinetis, Wilkins (1985) on Thomomys
orientalis, Edmund (1987) on Holmesina septentrionalis, Seymour (1993) on
Panthera onca, Emslie (this volume) on Arctodus pristinus, and Berta (this
volume) on Canis armbrusteri. Ritchie (1980) reviewed the avifauna from
Coleman 2A.
Other Florida Irvingtonian faunas.--Three additional Florida Irvingtonian
faunas not included in Table 2 are worthy of mention. The Punta Gorda LF (Fig.
4, site 23) was collected in the banks of Alligator Creek near Punta Gorda,
Charlotte County (Webb 1974a). The fossils were derived from a 1 m thick clay
unit overlying marine shell beds tentatively referred to the Bermont Formation.
Brooks (1968) provided a stratigraphic section for the Punta Gorda LF
(="gomphothere site"). Punta Gorda was one of the first Irvingtonian sites
reported from Florida (Webb 1974a), and was one of only four Irvingtonian faunas
from the state mentioned by Kurten and Anderson (1980). Webb (1974a) listed the
mammalian fauna from Punta Gorda which is composed of only nine species. The
most abundant mammals in the Punta Gorda LF are the proboscideans Mammuthus
hayi and Cuvieronius tropicus. The mammoth from this site is very similar to M.
hayi reported from Leisey (Webb and Dudley this volume). Although the presence
of Cuvieronius at Punta Gorda was originally thought to indicate an early
Pleistocene age, this last North American gomphothere is now known to have
survived into the early Rancholabrean in Florida. Other Irvingtonian taxa present
at Punta Gorda include Equus "leidyi" and Holmesinafloridanus. The age of the
Punta Gorda LF is not entirely clear because of the depauperate fauna, particularly
the lack of carnivores and rodents, but there are several constraints on its age. The
presence of mammoth confirms that this site is younger than 1.6 Ma, thus ruling
out an earliest Irvingtonian age. A late early Irvingtonian age for Punta Gorda is
most probable owing to the similarity between the mammoths from this site and
Leisey.
A mixed assemblage of Irvingtonian and Rancholabrean vertebrates has been
recovered from Apollo Beach in Hillsborough County, just north of the Leisey
Shell Pit (Fig. 4, site 18). Mammals in the Apollo Beach fauna that are shared with
Leisey include Castoroides, Neochoerus, Erethizon dorsatum, Arctodus pristinus,
Tapirus haysii, and an undescribed species of Equus (Hemionus). The Apollo
Beach LF is probably late early Irvingtonian in age.
Waldrop and Wilson (1990) reported Canis edwardii and Chasmaporthetes
ossifragus from a unit underlying the Caloosahatchee Formation in the Forsberg
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 81
Shell Pit, near Punta Gorda in Charlotte County (Fig. 4, site 14). These same two
carnivores also occur together in the Inglis 1A and De Soto Shell Pit faunas.
Chasmaporthetes is known from both late Blancan and earliest Irvingtonian faunas
in Florida, whereas Canis edwardii is restricted to the early Irvingtonian. Three
additional mammalian taxa, Pachyarmatherium leiseyi, an intermediate-sized
Holmesina floridanus, and Smilodon gracilis, have been collected recently from
within shell beds of the Caloosahatchee Formation at the Forsberg Shell Pit. This
represents the first well-documented earliest Irvingtonian record ofP. leiseyi. The
stage of evolution of the Holmesina from the Forsberg Shell Pit is typical of other
early Irvingtonian faunas such as Inglis 1A, Haile 16A, and Leisey Shell Pit.
Smilodon gracilis is found from the late Blancan through the middle Irvingtonian
in Florida. Most of the mammalian taxa from the Forsberg Shell Pit LF are
indicative of an early Irvingtonian age. The Concurrent Range Zone of Canis
edwardii and Chasmaporthetes ossifragus further constrains the age as earliest
Irvingtonian, between 2.0 and 1.6 Ma.
RANCHOLABREAN
Webb (1974a) provided mammalian faunal lists for more than 30
Rancholabrean sites from Florida and Kurt6n and Anderson (1980) discussed many
of these same sites as well. Rancholabrean vertebrate faunas are not treated
separately in our discussion or in Table 2. We have combined a number of the best
known sites to compile composite faunal lists for both the early Rancholabrean and
late Rancholabrean (Table 2). The composite Rancholabrean faunal lists are
important to our discussion because they extend the chronological distribution of
late Pliocene and early to middle Pleistocene mammals to include the late
Pleistocene and Holocene. We briefly summarize our criteria for recognizing
Rancholabrean faunas in Florida, and for distinguishing between the early and late
Rancholabrean.
The first appearance of Bison in North America at about 300 ka is generally
used to define the base of the Rancholabrean NALMA (Savage 1951; Lundelius et
al. 1987). The boundary between the early and late Rancholabrean corresponds to
the beginning of the last or Sangamonian interglacial at about 130 ka. Although
the presence of Bison is convenient for defining the Rancholabrean, this genus is
often absent in Florida Rancholabrean faunas. Therefore, it has been necessary to
establish other faunal criteria for recognizing Rancholabrean faunas.
Didelphis virginiana was long to thought to be restricted to Rancholabrean
and modern faunas. However, Martin (1974) reported Didelphis from the latest
Irvingtonian Coleman 2A fauna. Both the glyptodont Glyptotherium floridanum
and the ground sloth Megalonyx jeffersonii are known only from Rancholabrean
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
faunas in Florida (McDonald 1977; Kurt6n and Anderson 1980; Gillette and Ray
1981). The dire wolf, Canis dirus, is restricted to Rancholabrean faunas in North
America (Kurtdn and Anderson 1980). C. dirus is one of the most common large
carnivores in Florida Rancholabrean faunas. The Florida cave bear, Tremarctos
floridanus, also is found only in Rancholabrean faunas in Florida. The larger
tremarctine, Arctodus pristinus, is the characteristic bear of late Blancan and
Irvingtonian faunas in the state. Although the largest and best known North
American species of Smilodon, S. fatalis (=S. populator of Berta 1985), first
appears in the late Irvingtonian (Kurtdn and Anderson 1980), it is not recorded
from Florida until the early Rancholabrean. The rice rat, Oryzomys palustris, is
unknown in Florida or elsewhere in the United States prior to the Rancholabrean.
Neotropical oryzomyine rodents ultimately were derived from North America
during the Pliocene (Baskin 1986), but the genus Oryzomys apparently evolved
either in South America or tropical North America and then reinvaded temperate
North America in the late Pleistocene. Didelphis virginiana, Oryzomys palustris,
and Dasypus novemcinctus were the last Neotropical mammals to reach the United
States as participants in the Great American Interchange.
The presence of the giant bison, Bison latifrons, is often used to identify early
Rancholabrean faunas (Robertson 1974; Kurten and Anderson 1980). Despite the
supposed occurrence of B. latifrons in several Wisconsinan faunas in western
North America, the two richest samples of this species in Florida, Bradenton LF
and Haile 8A (Robertson 1974), are in faunas known to be early Rancholabrean in
age based on other taxa. The gomphothere Cuvieronius tropicus also appears to
have gone extinct in Florida during the early Rancholabrean, with the youngest
record from the early Rancholabrean Daytona Beach LF. Although there are no
published late Rancholabrean records of the giant ground sloth Eremotherium from
Florida, several fragamentary specimens of E. mirabile have been identified
recently from the late Rancholabrean Vero and Oklawaha River faunas. The vole,
Pitymys hibbardi, originally described from the early Rancholabrean Williston 3A
LF (Holman 1959) and since identified from the Bradenton and Haile 7A faunas
(Martin 1974), appears to be restricted to the early Rancholabrean in Florida. The
extinct cotton rat, Sigmodon bakeri, described from the late Irvingtonian Coleman
2A LF (Martin 1974), is known from the same three early Rancholabrean localities
as is P. hibbardi.
The late Rancholabrean comprises the time interval from about 130 to 10 ka,
including the last or Sangamonian interglacial (ca. 130 to 120 ka) and the last or
Wisconsinan glacial (120 to 10 ka). The latter half of the late Rancholabrean (<50
ka) is within the range of radiocarbon dating, and as a result a very detailed
chronology is known for this time period. Because of its proximity to the Recent,
there are far more late Rancholabrean vertebrate faunas known in North America
than from any other age. This is also true for Florida where there are more than
100 late Rancholabrean faunas. The composite list of Florida late Rancholabrean
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 83
mammals in Table 2 is derived from the faunal lists of the eight most prolific
faunas of this age faunass listed in footnote to Table 2). See Webb (1974a) and
Webb and Wilkins (1984) for individual faunal lists for the best known Florida late
Rancholabrean mammalian faunas.
Several species of Florida Pleistocene mammals are restricted to late
Rancholabrean faunas. The American lion Panthera atrox (=P. leo atrox of
Kurtdn and Anderson 1980) is a Rancholabrean immigrant from the Old World.
In Florida, P. atrox is known only from late Rancholabrean faunas. Florida's only
surviving large cat, the puma or Florida panther, Puma concolor, first appears in
the state during the late Rancholabrean, as does Bison antiquus (Robertson 1974).
Four species of rodents, Sciurus niger, Pitymys pinetorum, Microtus
pennsylvanicus, and Ondatra zibethicus, are restricted to Florida late
Rancholabrean faunas, although the latter two species occur in older faunas
elsewhere (Martin 1974; Kurt6n and Anderson 1980). The extant cotton rat
Sigmodon hispidus is typical of Florida late Rancholabrean faunas (Martin 1974
1979), although the earliest appearance of this species is in the late early
Rancholabrean Daytona Beach and Haile 8A local faunas. The end of the
Pleistocene is marked by the appearance of humans about 12 ka and the subsequent
extinction of the mammalian megafauna by about 10 ka, as well as the return to
interglacial conditions with a rise in sea level to its present position.
SUMMARY
Vertebrate fossils of four different ages have been recovered from the Leisey
Shell Pit in southwestern Hillsborough County, Florida. In ascending
stratigraphic/chronologic order these are: Arcadia Formation (late early or early
middle Miocene); Bone Valley Formation or stratigraphic equivalent (late
Miocene); Bermont Formation (early Pleistocene); and Fort Thompson Formation
(late Pleistocene). The Bermont Formation at Leisey consists of massive, sandy,
marine shell beds, with occasional lenses of dark, organic-rich silt and fine-grained
sand. These lenses produce the Leisey Shell Pit Local Fauna, a diverse, late early
Irvingtonian (1.6-1.0 Ma) vertebrate assemblage composed of over 200 taxa.
The age of the Leisey Shell Pit LF has been determined primarily by
vertebrate biochronology, with corroboration from paleomagnetic polarity and
strontium isotope chronology. Many types of biochronologic information are used,
including first occurrences of immigrant taxa; first occurrences of species that
evolved in situ; concurrent or overlapping range zones; stage of evolution in
morphologically-changing lineages; and last occurrences of key taxa that
presumably represent extinction. The latter must be used with caution as there are
many other reasons a taxon may be absent from a fauna. Although broadly similar
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. I
to other regions of North America, fossil vertebrates from Florida include some
endemic elements and a diverse fauna of Neotropical immigrants, as well as
differences in timing of first and last occurrences.
Dating marine shell beds using interbedded terrestrial vertebrate faunas has
long been applied in Florida, but not always successfully due to inadequate faunas
and the limited understanding of mammalian biochronology in the state prior to
the last 20 years. Both of these problems have been rectified with recent
discoveries of rich land mammal faunas from southern Florida, particularly the late
Blancan Macasphalt Shell Pit LF, the earliest Irvingtonian De Soto Shell Pit LF,
and of course, Leisey Shell Pit. Our work suggests that the Pinecrest Beds are late
Pliocene (2.5-2.0 Ma), the Caloosahatchee Formation is latest Pliocene (2.0-1.6
Ma), and the Bermont Formation is early Pleistocene (1.6-1.0 Ma). Previous
interpretations of the age of the Bermont ranged from late Pliocene to middle
Pleistocene. Results from vertebrate biochronology are consistent with those from
paleomagnetic polarity (reversed, Matuyama Chron) and broadly concordant with
strontium isotope chronology. However, the numerical dates from the strontium
isotope ratios are older than those based on vertebrate biochronology, ranging into
the late Pliocene. Combining age criteria from all of these sources, as well as sea-
level evidence, results in a most likely chronologic range for the Leisey Shell Pit
LF of 1.5 to 1.1 Ma, or late early Irvingtonian.
Among vertebrate faunas outside of Florida, the Leisey Shell Pit LF shares
the most taxa with the early Irvingtonian Gilliland LF in Texas. The correlative
Holloman LF of Oklahoma is also very similar. Leisey is older than the two best
known middle Irvingtonian faunas from eastern North America, Cumberland
Cave, Maryland and Port Kennedy Cave, Pennsylvania.
Biochronologic analysis of Florida vertebrate faunas results in the recognition
of eight distinct assemblages from the late Pliocene through the end of the
Pleistocene. These predominantly mammalian assemblages should be useful for
correlation within Florida and elsewhere in the southeastern United States. Age
ranges for the eight Florida vertebrate assemblages listed below are approximate,
but should be accurate to within about 0.2 Ma.
1. Late Blancan (2.5-2.0 Ma). Characterized by the limited occurrence
of "Glossotherium" chapadmalense, Borophagus diversidens, Canis
lepophagus, Sigmodon medius, Nannippus peninsulatus, Equus
(Dolichohippus) sp., Platygonus bicalcaratus, Hemiauchenia blancoensis,
and Rhynchotherium praecursor. First occurrence of Neotropical
immigrant genera that participated in the Great American Interchange,
including Dasypus, Holmesina, Glyptotherium, Pachyarmatherium,
"Glossotherium," Eremotherium, and Titanis. Only Florida Plio-
Pleistocene vertebrate assemblage characterized by the overlapping range
zone of Nannippus and these Neotropical immigrants. Best exemplified
MORGAN & HULBERT: OVERVIEW OF THE LEISEY SHELL PIT LOCAL FAUNA 85
by the Santa Fe River 1, Macasphalt Shell Pit, Haile 15A, and Kissimmee
River local faunas.
2. Latest Blancan (ca. 2.0 Ma). This is probably the most poorly defined
of the Florida Plio-Pleistocene vertebrate assemblages, as the single fauna
representing this time interval has yet to be fully described. Characterized
by the first well-documented occurrences of new species of Eremotherium
and Tapirus; by the first occurrence ofErethizon and Neotoma; and by the
last occurrence of a large, undescribed species of Chelydra. The
Holmesinafloridanus sample from this assemblage is intermediate in size
between the very small form from the late Blancan and the larger form
from the earliest Irvingtonian. Only known by the Haile 7C LF.
3. Earliest Irvingtonian (2.0-1.6 Ma). Characterized by the limited
occurrence of Erethizon kleini, Sigmodon curtisi, Ondatra idahoensis, and
an undescribed dwarf species of Hemiauchenia; and by the first
occurrence of Paramylodon harlani, Canis edwardii, Lepus sp., Equus
"leidyi," and Mammut americanum. Taxa shared with the late Blancan
assemblage but absent in succeeding faunas include Megalonyx
leptostomus, Trigonictis macrodon, Chasmaporthetes ossifragus, and
Capromeryx arizonensis, as well as the emydid turtle Trachemys
platymarginata and the giant phororhacid bird Titanis walleri. Best
exemplified by the Inglis 1A and De Soto Shell Pit local faunas.
4. Late early Irvingtonian (1.6-1.0 Ma). Characterized by the limited
occurrence of Nothrotheriops texanus, Ondatra annectens, and
Mammuthus hayi; the first occurrences of Megalonyx wheatleyi, Canis
armbrusteri, Lutra canadensis, Castoroides, Erethizon dorsatum, Geomys
pinetis, Podomys, Sigmodon libitinus, Pedomys, Synaptomys, Sylvilagus
floridanus, Sylvilagus palustris, Tapirus haysii, Palaeolama mirifica, and
the emydid turtle Trachemys scripta; and the last occurrence of
Glyptotherium arizonae, Pachyarmatherium leiseyi, Canis edwardii,
Trigonictis, Geomys propinetis, and Sylvilagus webbi. Best exemplified
by the Haile 16A, Leisey Shell Pit, Pool Branch, Payne Creek Mine,
Crystal River Power Plant, and Haile 21A local faunas.
5. Middle Irvingtonian (1.0-0.6 Ma). Characterized by the presence of
Neofiber leonardi and an advanced grade ofSmilodon gracilis; by the first
occurrence of Holmesina septentrionalis, Panthera onca, and Sigmodon
bakeri; and by the last occurrence of Megalonyx wheatleyi, Smilodon
gracilis, and Tapirus haysii. Only known by the McLeod LF which
probably falls late in the interval.
6. Late Irvingtonian (0.6-0.3 Ma). Characterized by the limited
occurrence of Urocyon minicephalus and Pitymys aratai; by the first
occurrence of Didelphis virginiana, Procyon lotor, Thomomys orientalis,
Podomys floridanus, Neofiber alleni, and Tapirus veroensis; and by the
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, PT. I, NO. 1
last occurrence of Arctodus pristinus and Canis armbrusteri. Represented
only by the Coleman 2A LF which is very late in the interval.
7. Early Rancholabrean (0.3 Ma-130 ka). Characterized by the limited
occurrence of Bison latifrons and Pitymys hibbardi; by the first occurrence
of Glyptotherium floridanum, Megalonyx jeffersonii, Canis dirus,
Tremarctos floridanus, Smilodon fatalis, Oryzomys palustris, Sigmodon
hispidus, Synaptomys australis, Platygonus compressus, and Mammuthus
columbi; and by the last occurrence of Cuvieronius and Sigmodon bakeri.
Best exemplified by the Bradenton, Daytona Beach, Haile 7A, Oldsmar,
and Williston 3A local faunas.
8. Late Rancholabrean (130-10 ka). Characterized by the limited
occurrence of Panthera atrox, Dinobastis serus, and Bison antiquus; by
the first occurrence of Sciurus niger, Pitymys pinetorum, Microtus
pennsylvanicus, Ondatra zibethicus, and Puma concolor; and by the last
local occurrence of numerous genera at the end of the interval. These
include Holmesina, Glyptotherium, Eremotherium, Megalonyx,
Paramylodon, Desmodus, Conepatus, Smilodon, Panthera, Tremarctos,
Castoroides, Neochoerus, Mylohyus, Platygonus, Hemiauchenia,
Palaeolama, Equus, Tapirus, Mammut, and Mammuthus. Among the
more than 100 late Rancholabrean faunas from Florida the best known are
Arredondo 2A, Cutler Hammock, Devils Den, Ichetucknee River,
Melbourne, Monkey Jungle Hammock, Reddick 1, Seminole Field, and
Vero.
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