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Group Title: Bulletin of the Florida Museum of Natural History
Title: Fossil history of the panther (Puma concolor) and the cheetah-like cat (Miracinonyx inexpectatus) in Florida
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 Material Information
Title: Fossil history of the panther (Puma concolor) and the cheetah-like cat (Miracinonyx inexpectatus) in Florida
Series Title: Bulletin - Florida Museum of Natural History ; volume 40, number 2
Physical Description: p. 177-219 : ill., maps ; 23 cm.
Language: English
Creator: Morgan, Gary
Seymour, Kevin L.
Donor: unknown ( endowment )
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1997
Copyright Date: 1997
 Subjects
Subject: Florida panther   ( lcsh )
Animals, Fossil -- Florida   ( lcsh )
Paleontology -- Pleistocene   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Includes bibliographical references (p. 217-219).
Language: Abstract also in Spanish.
General Note: Cover title.
Statement of Responsibility: Gary S. Morgan and Kevin L. Seymour.
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Bibliographic ID: UF00095787
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 37997832
issn - 0071-6154 ;

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    Copyright
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    Abstract
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Full Text



BULLIITI


of the



SFLORIDA
MUSEUM OF
NATURAL HISTORY

FOSSIL HISTORY OF THE PANTHER (PUMA
CONCOLOR) AND THE CHEETAH-LIKE CAT
(MIRACINONYX INEXPECTATUS)
IN FLORIDA

Gary S. Morgan and Kevin L. Seymour


Volume 40. No. 2 pp. 177-219


UNIVERSITY OF FLORIDA


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ISSN: 0071-6154


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FOSSIL HISTORY OF THE PANTHER (PUMA CONCOLOR)
AND THE CHEETAH-LIKE CAT (MIRACINONYX
INEXPECTATUS) IN FLORIDA


Gary S. Morgan' and Kevin L. Seymour2




ABSTRACT

Fossils of the Florida panther or puma (Puma concolor) are reported from 15 late Pleistocene
(Rancholabrean) sites and one Holocene archaeological site on the Florida peninsula (11 of these are
unpublished records). This large cat was widely distributed in Florida during the late Pleistocene, from as
far north as Columbia County near the Georgia state line, throughout the northern and central portions of the
state, along both the Atlantic and Gulfcoasts, and as far south as Dade County at the southernmost tip of the
peninsula. P. concolor is unknown from late Pleistocene faunas in the Florida panhandle. The living
Florida panther, P. concolor coryi, is now restricted to southernmost peninsular Florida in Glades, Hendry,
Collier, and Dade counties, although it was fairly common farther north along the central Atlantic coast until
the late nineteenth century. All of the P. concolor fossils reported here compare closely in size and
morphology to Recent skulls and postcranial skeletons ofP. concolor from southern Florida. The associated
mammalian faunas from the Florida sites containing P. concolor indicate that this species is restricted to the
late Rancholabrean d Recent (between 130 ka and the present).
The cheetah-like cat, or puma-like cat, Miracinonyx is identified from eight late Pliocene and
Pleistocene sites in Florida, only two of which were published previously. M. inexpectatus occurs in four
Florida late Pliocene and early Pleistocene sites, including the late Blancan Santa Fe River 2A and Northport
faunas and the early Irvingtonian Inglis IA and Leisey Shell Pit local faunas. Three associated metacarpals
from the Rancholabrean Lecanto 2A local fauna are tentatively identified as M. inexpectatus, representing
one of the youngest records of this species. Fossils identifiable only as Miracinonyx sp. are recorded from
the late Irvingtonian Coleman 2A local fauna, from a Plio-Pleistocene locality in Port Charlotte and from the
Pleistocene Cardinale site, an underwater fossil site in the Gulf of Mexico located 25 km offshore.
Miracinonyx appears to be closely related to the genus Puma and is distinguished from P. concolor by its
somewhat larger size and conspicuous elongation ofthe limbs and metapodials.

RESUME

Los fosiles de Ia pantera o puma de la Florida (Puma concolor ) son reportados de 15 sitios del
Pleistoceno tardio (Rancholabreano) y de un sitio arqueol6gico del Holoceno en la peninsula de la Florida.
Este gato de gran tamanfo estaba ampliamente distribuido en la Florida durante el Pleistoceno tardio, desde
tan al norte como el condado de Columbia pr6ximo a la line estatal de Georgia, a trav6z de las porciones
nortelas y centrales del estado, a lo largo de ambas costas a atllntica y la del Golfo, y tan al sur como en el
condado de Dade en la punta mAs surefia de la peninsula. P. concolor es desconocido de la fauna del


SNew Mexico Museum ofNatual History, 1801 Mountin Road NW, Albuquerquqe, NM 87104.
SDepartment of Palaeobiology, Royal Ontrio Museum, 100 Queen's Prk, Toronto. OntMio, Caud MSS 2C6.

Morgan, G. S., and K. L. Seymour. 1997. Fossil history ofthe panther (Puma concolor) and the cheetah-
like cat (Miracinonyx inexpectatus) in Florida. Bull. Florida Mus. Nat. Hist 40(2):177-219.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Pleistoceno tardio en la region noroeste de la Florida. La painter de la Florida que actualmente existed, P.
concolor coryi, esta ahora restringida a la parte mis surefla de la peninsula de la Florida en los condados de
Glades, Hendry, Collier, y Dade, aunque dta era regularmente comun mis al norte a lo largo de la costa
central atlintica hasta finales del siglo diecinueve. Todos los fosiles de P. concolor reportados aqui se
asemejan bastante en tamalio y morfologa a los craneos y esqueletos postcraneales de P. concolor del
Reciente en el sur de la Florida. Las faunas de mamlferos asociadas a los sitios de la Florida que contienen
P. concolor indican que sdta especie esta restringida al Rancholabreano tardio y al Reciente centree 130 kilo
afios y el presente.
El gato semejante al cheetah, o semejante al puma, Miracinonyx es identificado en ocho sitios del
Plioceno tardio y del Pleistoceno en la Florida, solo dos de estos sitios fueron reportados previamente. M.
inexpectatus ocurre n cuatro sitios del Plioceno tardlo y del Pleistoceno temprano on la Florida, incluyendo
las faunas locales del Santa Fe River 2A y Northport del Blancan tardio y del Inglis IA y Leisey Shell Pit del
Irvingtoniano temprano. Los tres huesos asociados del metacarpo de la fauna local del Lecanto 2A del
Rancholabreano son tentativamente identificados como M. inexpectatus representando uno de los mis
recientes registros de esta especie. Los fosiles identificados unicamente como Miracinonyx sp. son
registrados en la fauna local de Coleman 2A del Irvingtoniano tardio, on una localidad en Port Charlotte del
Plio-Pleistocene y n el sitio Cardinale del Pleistoceno, que es un sitio de fosiles bajo el agua on el Golfo de
M6xico localizado a 25 Km mar afuera Miracinonys aparenta star cercanamente relacionado con el
genero Puma y se distingue de P. concolor por su aproximado gran tamano y su conspicua elongaci6n de las
extremidades y de los metapodios.

ACKNOWLEDGMENTS

Many individuals have collected fossils of Puma concolor and Miracinonyx from sites throughout
Florida and generously donated their finds to the Florida Museum of Natural History (FLMNH). The
largest fossil sample ofP. concolor from Florida is from the Cutler Hammock site, which was excavated by
archaeologists from the Metro-Dade Archaeology Division and the Dade County Archaeological
Conservancy under the direction of Robert Carr. Access to the Cutler Hammock site was granted by the
Deering family, who donated the entire Cutler collection of vertebrate fossils to the FLMNH. Lewis D.
Ober, William G. Weaver, and several of their students from Miami-Dade Community College excavated
the Monkey Jungle Hammock Site and deposited the fossils in the FLMNH. Frank Dumond, general
manager of Monkey Jungle, kindly permitted access to the property. Erika Simons and Laurie Wilkins
helped collect panther fossils from the Wekiva River. Arthur Poyer collected a maxilla of P. concolor from
Orange Lake 2A. Robin Brown donated a panther mandible from the Peace River to the FLMNH collection.
Thomas Cardinale brought the underwater site in the Gulf of Mexico west of St. Petersburg (now named the
Cardinale Site in his honor) to the attention of FLMNH paleontologists and donated the fossils from this site,
including a Miracinonyx radius. Steven Beck contributed a large collection of vertebrate fossils from the
Lecanto 2A site to the FLMNH, including the metacarpals of Miracinonyx inexpectatus described here.
Ralph (Tony) Estevez donated a metatarsal of M. inexpctatus from the Leisey Shell Pit to the FLMNH.
We are grateful to Erika Simons who took the photographs in Figures 5-7 and helped produce all the
photographic plates. Marc Frank provided information on fossil felid specimens in the FLMNH Vertebrate
Paleontology Collection. We thank Laurie Wilkins and Charles Woods of the FLMNH Mammalogy
Collection for allowing us to examine their excellent collection of Recent skulls and postcranial skeletons of
P. concolor coryi. John Eisenberg, Richard Franz, and two anonymous reviewers made numerous helpful
comments on the manuscript We are particularly grateful to John Eisenberg for providing funds from the
Ordway endowment for the publication of this paper.

INTRODUCTION

Puma (=Felis) concolor, the puma or Florida panther, previously was
identified from six Pleistocene fossil sites in Florida (Webb, 1974a; Kurt6n, 1976;
McDonald, 1990). This meager record is surprising considering the large number
of Pleistocene vertebrate faunas known from Florida (Webb, 1974a; Kurten and






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 179


Anderson, 1980), and the fact that southern peninsular Florida is the only region
east of the Mississippi River where P. concolor still survives. Fossils of P.
concolor reported here add 11 new localities from the state. The number of
Florida sites known to contain P. concolor is now 16, which also takes into
account one previous record that is shown below to be erroneous. Of these sites, 15
are late Pleistocene (late Rancholabrean) in age and one is a Holocene
archaeological site. We also identify the cheetah-like or puma-like cat,
Miracinonyx, a probable fossil relative of P. concolor, from eight Florida sites
ranging in age from late Pliocene (late Blancan and earliest Irvingtonian) to
Rancholabrean.
Simpson (1929) first reported a puma-like cat from the Pleistocene of Florida.
He referred an upper carnassial from Seminole Field in Pinellas County to Felis cf.
F. inexpectata (Cope). It should be noted here that the generic names for the large
cats have changed significantly over the years. We repeat the generic names as
they were used by the original authors to avoid confusion in this brief historical
review. Discussion under the Systematic Paleontology section clarifies our opinion
on the generic designations of the Florida fossil cat species reviewed here. Ray
(1958) identified a radius, astragalus, six phalanges, and two fragmentary
metapodials from Melbourne in Brevard County as Felis cf. F. (Puma) inexpectata.
Ray noted that the Melbourne fossils could not be clearly separated from
comparable elements of extant Felis concolor. The next mention of fossil panther
from Florida was by Kurt&n (1965, fig. 9) who referred a single astragalus from
Reddick 1A in Marion County to Felis concolor. Kurt6n regarded the Reddick
astragalus as indistinguishable in size and morphological characters from the
living puma.
Webb (1974a) listed Felis concolor from the Santa Fe 2A locality, a mixed
late Blancan and Rancholabrean assemblage. This record apparently was based on
a metacarpal that is here re-identified as a Blancan specimen of Miracinonyr
inexpectatus (see discussion below), as no other fossils of F. concolor are present
in the Florida Museum of Natural History collections from Santa Fe 2A. Kurt6n
(1976) reviewed the fossil history of pumas (Fells concolor) and puma-like cats (F.
inexpectata) in North America. He referred specimens from Blancan and
Irvingtonian sites to F. inexpectata and Rancholabrean fossils to the living species
F. concolor. Kurten distinguished F. inexpectata from F. concolor by its larger
size and more elongated postcranial elements. Kurten (1976) reported fossils ofF.
concolor from four Florida Rancholabrean sites, three listed above, including
Seminole Field, Melbourne, and Reddick lA, and one that represented a new
record, a maxillary fragment with P3-P4 from Devil's Den in Levy County. The
fossils from Seminole Field and Melbourne, identified as F. concolor by Kurt6n
(1976), were the same specimens earlier referred to Felis cf. F. inexpectata by
Simpson (1929) and Ray (1958), respectively.
Van Valkenburgh et al. (1990) transferred F. inexpectata to the genus
Miracinonyx, along with the closely related species M. trumani, and also suggested







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


that M. inexpectatus might be near the ancestry of the puma. They referred several
specimens from the earliest Irvingtonian (latest Pliocene) Inglis IA site in Citrus
County, Florida, to M. inexpectatus. McDonald (1990) listed F. concolor from the
Rancholabrean Warm Mineral Springs site located in Sarasota County in the
southwestern portion of the peninsula. Berta (1995) identified a metatarsal of M.
inexpectatus from the early Irvingtonian (early Pleistocene) Leisey Shell Pit
located along the eastern shore of Tampa Bay in Hillsborough County.
Prior to 1980 there were few Recent specimens of the Florida panther, Puma
concolor coryi, in museum collections. The comparatively large sample of Florida
panther specimens now available in the Mammalogy Collection of the Florida
Museum of Natural History permits a considerably more detailed analysis of
individual, sexual, and geographical variation in this subspecies than was
previously possible. Unfortunately, this large sample of museum specimens has
come at great cost to the dwindling population of the Florida panther, since the
majority of these endangered cats were killed by vehicles as they attempted to cross
busy South Florida highways. Wilkins et al. (1997) provides a detailed analysis of
individual and geographical variation in the pelage and crania of extant P.
concolor coryi.

METHODS AND ABBREVIATIONS

We have examined and confirmed the identification of all Puma concolor
fossils known from Florida, with one exception. P. concolor fossils from eleven of
these sites were previously unreported. The single panther fossil we were not able
to examine was an astragalus (UF 8895) from the late Rancholabrean Reddick IA
site, identified and figured by Kurtdn (1965, fig. 9B). Several specimens from
Reddick 1A in the James Gut Collection, including this P. concolor astragalus,
were catalogued into the UF Vertebrate Paleontology Collection but were never
received by the museum. Sometime prior to Gut's death, Kurtdn obviously had the
opportunity to examine fossils in Gut's private collection.
Dental terminology is standard: incisors (I/i), canines (C/c), premolars (P/p)
and molars (M/m). Upper teeth are indicated by upper case letters (e.g. P4 is the
upper fourth premolar) and lower teeth are indicated lower case letters (e.g. ml is
the lower first molar). For comparative purposes, we examined and measured 20
skulls (12 males, 8 females) and 18 postcranial skeletons (10 males, 8 females) of
Recent adult Puma concolor coryi from southern peninsular Florida, primarily
Collier and Dade counties.
Detailed map data, field notes, and other information on most Florida sites
containing fossils of Puma concolor and Miracinonyx are on file in the Vertebrate
Paleontology Collection of the Florida Museum of Natural History. P. concolor
and Miracinonyx fossils from Florida are housed in the following museums (with
acronyms): Field Museum of Natural History, Chicago (FMNH), Florida Museum
of Natural History, University of Florida, Gainesville (UF), American Museum of






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 181


Natural History, New York (AMNH), Museum of Comparative Zoology, Harvard
University, Cambridge (MCZ), and Royal Ontario Museum, Toronto (ROM).


SYSTEMATIC PALEONTOLOGY
Class Mammalia Linnaeus, 1758
Order Carnivora Bowdich, 1821
Family Felidae Gray, 1821
Puma concolor (Linnaeus, 1771)

Material Examined

All fossils of Puma concolor identified from Florida are listed below. The
fossils are listed separately for each site with their catalogue numbers and a brief
description of the element. Each specimen was directly compared with the large
Recent sample of P. concolor skulls, teeth, and postcranial bones from Florida to
confirm its identification. The features illustrated in Seymour (1983) and other
publications listed below were used where possible.

Late Pleistocene (late Rancholabrean)

Ichetucknee River, Columbia County: UF 45420, left metacarpal II.
Santa Fe River 1, Columbia/Gilchrist County line: UF 124614, right mandible with
ml.
Devil's Den, Levy County: UF 9256, partial left maxilla with P3-P4; UF 133802,
left ml.
Wekiva River, Levy County: UF 124387, right metacarpal II; UF 124388, right
metacarpal IV; UF 124389, left metacarpal V; UF 124386, left proximal
femur, UF 124390, right metatarsal Il; UF 124391, left proximal metatarsal
IV.
Orange Lake 2A, Marion County: UF 134300, partial right maxilla with P3-M1.
Reddick IA, Marion County (from Kurtdn, 1965: fig. 9): UF 8895, left astragalus.
Silver Springs, Marion County: UF 51221, left ulna.
Rock Springs, Orange County: UF 8954, phalanx.
Melbourne, Brevard County: MCZ 17791, right radius, right astragalus, right
distal metatarsal II, and at least two phalanges.
Seminole Field, Pinellas County: AMNH 23540, incomplete left P3; AMNH
23541, right P4.
Leisey Shell Pit 2, Hillsborough County: UF 125192, juvenile left metacarpal IV.
Peace River, Hardee County: UF 45949, left mandible with broken canine and
worn p3-p4; UF 93415, left p4.
Warm Mineral Springs, Sarasota County: UF 23870, left distal femur.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Cutler Hammock, Dade County: UF 172944, right P4; UF 124376, left mandible
with p3-ml; UF 93415, left p4; UF 124377, left humerus; UF 124378, right
distal humerus; UF 124379, left metacarpal H; UF 124380, left proximal
metacarpal III; UF 124382, right proximal metacarpal V; UF 124381, left
proximal metacarpal V; UF 128051, proximal phalanx of right manus, digit
V; UF 124384, left astragalus; UF 124382, right proximal metatarsal V; UF
124383, left proximal metatarsal V.
Monkey Jungle Hammock, Dade County: UF 45393, right P4; UF 45392, left ml;
UF 24520, partial right p3; UF 124385, left metacarpal V.

Holocene

Granada Site, Dade County: UF zooarchaeology collection (uncatalogued), distal
epiphysis of metapodial and phalanx.

Distribution

All of the fossil sites in Florida that have produced specimens of Puma
concolor are indicated on the map in Figure 1. The 15 late Pleistocene sites and
one Holocene archaeological site are distributed throughout peninsular Florida,
from Columbia County in the northern part of the state not far from the Georgia
line to Dade County at the extreme southeastern tip of the peninsula. Seven sites
are located in Columbia, Levy, and Marion counties in northern Florida, two are in
the central part of the state (one in Orange County and one near the central
Atlantic Coast in Brevard County), three are in southwestern Florida along the
Gulf Coast (one each in Pinellas, Hillsborough, and Sarasota counties), one is in
south-central Florida in Hardee County, and three sites are located in southernmost
Florida in Dade County. Fossils of P. concolor currently are unknown from the
Florida panhandle. Fewer late Pleistocene fossil sites are known from the
panhandle than from the peninsula, but at least five late Rancholabrean river-
bottom/spring deposits in this region have diverse mammalian faunas, including
the Aucilla River in Jefferson and Taylor counties, Wacissa River in Jefferson
County, Wakulla Springs and Wakulla River in Wakulla County, St. Marks River
in Leon and Wakulla counties, and Chipola River in Jackson County (Webb,
1974a).
There are three major types of Rancholabrean fossil deposits in Florida that
have produced fossils of Puma concolor. River or spring deposits comprise eight
of the fossil sites from which panthers are known, sinkhole/cave/fissure deposits
account for four panther records, and sites deposited in coastal rivers or lagoons
constitute the remaining three sites. Three of the underwater sites, Devil's Den,
Rock Springs, and Warm Mineral Springs, probably were dry caves or sinkholes in
the late Pleistocene, and thus could be classified in either of the first two locality
types. Most of the late Pleistocene sites known from Florida fall into one of these







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY


.pO


Figure 1. Late Pleistocene (Rancholabrean ) and Holocene sites from Florida containing Puma concolor.
Rancholabrean: 1. Ichetucknee River, Columbia County, 2. Santa Fe River 1, Columbia/Gilchrist County
line; 3. Devil's Den, Levy County, 4. Wekiva River, Levy County, 5. Orange Lake 2A, Marion County, 6.
Reddick IA, Marion County, 7. Silver Springs, Marion County, 8. Rock Springs, Orange County, 9.
Melbourne, Brevard County 10. Seminole Field, Pinellas County, 11. Leisey Shell Pit 2, Hillsborough
County, 12. Peace River, Hardee County, 13. Warm Mineral Springs, Sarasota County, 14. Cutler
Hammock, Dade County, 15. Monkey Jungle Hammock, Dade County. Holocene: 16. Granada Site, Dade
County.




three categories (Webb, 1974a), and thus the type of site probably has little affect
on the presence or absence of P. concolor fossils. This is not unexpected
considering that living Florida panthers and western pumas/mountain lions have
very large home ranges and, as a consequence, regularly travel through a number
of habitats over a wide geographical area.






BULETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Wilkins et al. (1997) map the distribution of Recent specimens of Puma
concolor coryi collected in the Florida peninsula over the past century. Florida
panthers are now restricted to the southern third of the peninsula in Glades,
Hendry, Collier, and Dade counties; however, in the late 19th century a number of
specimens were collected along the central Atlantic coast in Volusia, Brevard, and
Indian River counties. P. concolor probably was found throughout the Florida
peninsula prior to the arrival of Europeans. The fossil record clearly establishes
that P. concolor was widely distributed in the Florida peninsula in the late
Pleistocene.

Taxonomy

Until recently most members of the family Felidae were placed in the genus
Felis. Mammalogists generally have used the binomen Fells concolor for the
puma/mountain lion/cougar and the subspecies name F. concolor coryi for the
Florida panther. Over the past decade or so felids have been split into a number of
genera on the basis of their phylogenetic relationships. As a consequence, felid
systematists now prefer to use the genus Puma for the puma or Florida panther
(Glass and Martin, 1978; Kral and Zima, 1980; Collier and O'Brien, 1985;
Herrington, 1986; Wayne et al., 1989; Van Valkenburgh et al., 1990; Salles, 1992;
Janczewski et al., 1995), and we follow this convention in the present paper. We
do note, however, as did Salles (1992:4), that at this point in the understanding of
felid phylogenetics, the choice of generic rank for felids is still largely arbitrary.

Diagnosis

The puma is different from all other living cats because of its combination of
large size, unicolor coat as an adult, relatively long tail, and small head. These
external features are, however, little help to the paleontological diagnosis of this
species or its relatives. Werdelin (1983) used a correspondence analysis to show
that the puma has the cranial proportions of a small cat, while attaining the large
body size of a pantherine cat. He also pointed out that the puma differs from other
cat species in its short, broad teeth, especially the p4 and ml, and relatively small
canines compared with the pantherines. When compared to the jaguar, in
particular, the puma has a relatively small canine, larger p3, and longer ml
(Werdelin 1983:385). Comparisons with small cats, such as the jaguarundi
(Herpailurusyagouaroundi), show the puma to have relatively round canines and a
long P3 (Glass and Martin, 1978). These two authors also found that living pumas
have a relatively smaller p4, larger ml, larger canines, smaller P3, and less
reduced protocone on P4, when compared to a possible fossil relative, such as
Miracinonyx trumani. Other osteological features that can be used to help
differentiate puma skulls from jaguar skulls were described by Hoffstetter (1949),
whereas postcranial features differentiating puma bones from jaguar bones were






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 185


elucidated by Seymour (1983). We utilized these two works for the identification of
isolated feline fossils.
In summary, there are few if any unique osteological features within this
group of cats. Identification of fossil material is difficult, with direct comparison
preferred in all cases. Furthermore, there were several other large felids present in
the Pleistocene of Florida whose bones might be confused with puma fossils. Van
Valkenburgh et al. (1990) and Seymour (1993) noted that several of these felids, in
particular Smilodon gracilis/S. fatalis, Miracinonyx inexpectatus, and Panthera
onca often occur together in sites, so that the problem of identification of isolated
remains is not imaginary. Papers illustrating many bones of these species are as
follows: Berta (1987) for Smilodon gracilis, Rawn-Schatzinger (1992) for
Homotherium serum, Merriam and Stock (1932) for Panthera atrox and Smilodon
fatalis, Van Valkenburgh et al. (1990) for Miracinonyx inexpectatus and M.
trumani, and Seymour (1983) for Panthera onca. Although these publications are
excellent resources, they tend to illustrate the bones of one or two species and
generally lack differential diagnoses which might be used to identify isolated feline
fossils.

Descriptions and Comparisons

The fossils listed above under Material Examined are very similar in size and
morphology to bones and teeth of living Puma concolor from Florida and are
confidently referred to that species. Some confusion existed in the past concerning
the few previously published fossil records of P. concolor from Florida. Kurtdn
(1965, 1976) rectified this situation by demonstrating that some of the large felid
fossils from Melbourne and Seminole Field were conspecific with P. concolor. We
found the P. concolor samples from both Melbourne and Seminole Field to be a
mixture of species. Included within the Melbourne material (MCZ 17791) is a
right radius and right astragalus of P. concolor. One distal metapodial (a right
metatarsal II) most likely represents P. concolor, but the other (a left metatarsal V)
represents a canid. Of the six phalanges, two are middle phalanges and four are
proximal phalanges. The middle phalanges are diagnostically felid-shaped, but we
were unable to identify them to species. The smallest and one of the largest of the
four proximal phalanges most likely represent P. concolor, but the other two large
phalanges are heavier and probably belong to Panthera onca, the jaguar. The
Seminole Field material as identified above differs somewhat from Kurtdn (1965,
1976). We agree that AMNH 23541 represents a right P4 of P. concolor.
However, there are five isolated teeth catalogued under AMNH 23540, only one of
which represents P. concolor (an incomplete left P3); the other teeth represent
primarily jaguar. Kurt6n (1965, 1976) identified an astragalus of Puma concolor
from Reddick. The Reddick astragalus was not available for study (see more
detailed discussion above in Methods section), but its identification has not been
questioned.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Table 1. Measurements of the upper dentition of Pleistocene and Recent Puma concolor from Florida. The
mean (x) and observed range (O.R.) are provided for the Recent sample, which consists of 20 individuals (12
males, 8 females) from Collier and Dade counties in southern Florida.

P3 P4
L W L W

Recent
x 14.2 8.7 21.7 11.6
O.R. 12.5- 8.0- 19.2- 10.4-
15.3 10.1 23.3 12.8

Devils Den
UF 9256 14.3 8.7 23.1 11.6

Orange Lake 2A
UF 134300 15.1 8.2 23.9 11.8

Cutler Hammock
UF 172944 21.5 10.5

Monkey Jungle Hammock
UF 45393 22.4 11.0

Seminole Field
AMNH 23540 8.5 -
AMNH 23541 21.4 10.6



Measurements of the upper dentition (P3 and P4) of Puma concolor fossils
from Florida are provided in Table 1. This table also includes comparative dental
measurements for 20 adult skulls (12 males, 8 females) of extant P. concolor from
southern Florida. Kurtdn (1973a) presented comparative measurements of the P4
and ml of Recent P. concolor from throughout the species' wide geographical
range (from western Canada to southern Argentina). Although several of the
measurements for fossils of P. concolor from Florida are slightly outside the size
range of Recent Florida panthers, these measurements fit well within the size range
of the species as a whole.
The fossil sample of Puma concolor from Florida includes two maxillary
fragments with P3-P4, one is the specimen from Devil's Den (UF 9256) reported
by Kurten (1976) and the second is a recently collected fossil from Orange Lake
2A (UF 134300). Both of these maxillae are illustrated in Figure 2. There are also
three isolated P4s, one each from Monkey Jungle Hammock (UF 45393; see Figs.
2E, F), Cutler Hammock (UF 172944), and Seminole Field (AMNH 23541), as
well as a partial P3 from Seminole Field (AMNH 23540). The measurements in
Table 1 reveal that, with one minor exception, all of the fossils are within the size
range of the modern sample of P. concolor from southern Florida. The P4 from





MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY


*a


41


Figure 2. Maxillae and upper dentition of Rancholabrean (late Pleistocene) Puma concolor from Florida.
A. lateral view and B. occlusal view, partial right maxilla with P3-M1, UF 134300, Orange Lake 2A,
Marion County, C. lateral view and D. occlusal view, partial left maxilla with P3-P4, UF 9256, Devil's
Den, Levy County, E. lateral view and F. occlusal view, right P4, UF 45393, Monkey Jungle Hammock,
Dade County. Scale bar equals 2 cm.


'iYxv


~P~ F







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Table 2. Measurements ofthe lower dentition of Pleistocene and RecentPuma concolor and Plio-Pleistocene
Miracinonyx from Florida. Comparative measurements are also provided for Miracinonyx inexpectatus
from Hamilton Cave, West Virginia and M. trumani from Natural Trap Cave, Wyoming (from Van
Valkenburgh et al., 1990). The mean (x) and observed range (O.R.) are provided for the Recent P. concolor
sample, which consists of 20 individuals (12 males 8 females) from Collier and Dade counties in southern
Florida. The measurements for M. trumani are also mean values, but have different sample sizes for
individual teeth (see footnotes)


p3 p4 ml
L W L W L W


Puma concolor (Recent)
x
O.R.


Puma concolor (fossil)
Cutler
UF 124376


Devil's Den
UF 133802

Monkey Jungle
UF 45392


Miracinonyx inexpectalus
Inglis 1A
UF 21604

Northport
ROM 29000
ROM 32286
Hamilton Cave
USNM 401092

Miracinonyx trumani
Natural Trap Cave
x


16.4 8.3
14.3- 7.6-
18.4 8.9


11.1 7.1 14.9


8.2 18.6 8.4


- 18.1 8.9


- 17.7 7.9



12.6 7.2 17.1 9.0 19.0 9.6


8.6 19.2
- -17

7.0 18.3



6.7' 16.0


9.5
8.5 -


8.2 20.1


7.72 16.1 7.92


'Men of9 individuals (fiom Van Valk h a,1990, table 2).
2 Mean of lOindividual (from Van Valmbughet 1990,tabe 2).





MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY


jj/ D
C

Figure 3. Mandible and lower dentition of Rancholabrean (late Pleistocene) Puma concolor from Florida.
A. lateral view and B. occlusal view, left mandible with p3-ml, UF 124376, Cutler Hammock, Dade
County, C. lateral view and D. occlusal view, left ml, UF 133802, Devil's Den, Levy County. Scale bar
equals 2 cm.


Orange Lake 2A is slightly longer anteroposteriorly (23.9 mm) than the largest
Florida P. concolor measured (23.3), but the Orange Lake fossil is otherwise very
similar to the Recent sample. No differences in dental morphology were observed
between the Pleistocene and Recent samples of P. concolor.
Table 2 compares measurements of the lower dentition and mandible of fossil
and living Puma concolor from Florida. Measurements are given for only three
fossils, a well-preserved left mandible with p3-ml from Cutler Hammock (UF
124376) and isolated left mis from Devil's Den (UF 133802) and Monkey Jungle
(UF 45392). The Cutler mandible and Devil's Den ml are illustrated in Figure 3.
Mandibles from Santa Fe River 1 and Peace River are either damaged or the teeth


r7J
b^ite^






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


are too worn to provide accurate measurements. As with the upper dentition, the
measurements of the fossils closely match those of the Recent sample. The p3
from the Cutler mandible is slightly broader (7.1 mm) than the largest p3 (6.9 mm)
in the extant P. concolor sample, and the ml from the same Cutler specimen is
slightly longer in the anteroposterior dimension (18.6 mm) than the ml in any
Recent Florida panther mandible measured (18.4 mm). The overall morphology of
the fossil mandibles and lower dentition closely matches the Recent sample of P.
concolor.
Measurements of limb bones of Recent and fossil Puma concolor are included
in Table 3. A complete humerus from Cutler Hammock (UF 124377) is well
within the Recent sample in all measurements. A radius from Melbourne (MCZ
17791) is slightly smaller in several dimensions, including proximal breadth and
depth, than the Recent P. concolor sample but is otherwise very similar to the
radius of a small female Florida panther. An ulna from Silver Springs (UF 51221)
is within the range of extant P. concolor ulnae in all measurements except for
having a slightly broader shaft. A proximal femur from the Wekiva River (UF
124386) is at the small end of the observed range of measurements of Recent P.
concolor femora, mirroring the situation in several metacarpals from the Wekiva
River discussed below. A distal femur from Warm Mineral Springs (UF 23870) is
very slightly broader (52.3 mm) than the largest extant panther femur measured
(UF 52.2 mm), but the fossil shows no other significant differences from the
Recent sample.
Table 4 provides comparative measurements of metacarpals and metatarsals
of Recent and Pleistocene Puma concolor from Florida. With the exception of two
specimens from the Wekiva River, the measurements of the fossil metapodials are
within the range of variation in the extant sample. The Wekiva sample includes a
metacarpal HI (UF 124387), metacarpal IV (UF 124388), and metacarpal V (UF
124389). These specimens were given separate catalogue numbers because they
were not directly associated in the field, but the metacarpal m and IV probably
belong to the same individual, as they are smaller than the smallest Recent
specimens measured in total length and in several other dimensions (Table 4).
Except for their somewhat smaller size, these specimens are similar in morphology
to P. concolor and probably represent a small female individual of that species.

Biochronology

All Florida fossil sites that have produced Puma concolor fossils are
Rancholabrean in age. The first appearance of the genus Bison in North America
generally is used to define the beginning of the Rancholabrean Land Mammal Age
(Savage, 1951; Lundelius et al., 1987). Surprisingly, the age of the earliest North
American arrival of Bison, and thus the lower boundary of the Rancholabrean, is
not well established, with dates ranging from about 300 to 500 ka. We follow







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY


Table 3. Measurements of limb bones of Pleistocene and Recent Puma concolor and Plio-Pleistocene
Miracinonyx from Florida. Comparative measurements also are provided for Miracinonyx inexpectatus
from Hamilton Cave, West Virginia and M. trumani from Natural Trap Cave, Wyoming (from Van
Valkenburgh et al., 1990) The mean (x) observed range (O.R.) and sample size (N) are provided for
measurements of the Recent sample ofP. concolor, which consists of between 15 and 18 individuals from
Collier and Dade counties in southern Florida. The measurements forM. trumani are also mean values, but
have different sample sizes for each limb element (see footnotes).


total proximal proximal shaft distal
length width depth' width width


Humers
Puma concolor (Recent)
x
O.R.

N
Puma concolor (fossil)
Cutler
UF 124377
UF 124378
Miracinonyx inexpectatus
Inglis
UF 95766
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave3
Radius
Puma concolor (Recent)
x
O.R.

N
Puma concolor (fossil)
Melbourne
MCZ 17791
Miracinonyx inexpectatus
Inglis
UF 45346
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave'

Ulna
Puma concolor (Recent)
x
O.R.


225.1 43.4 56.7 19.0 50.9
204- 38.1- 50.5- 15.9- 45.7-
245 47.8 63.2 22.2 57.5
16 16 16 16 16


227 43.2 57.5


18.9 47.5
- 49.8


264 53.5 71.6 22.2 62.1


269

236.9


- 21.5 59.5

- 18.1 52.7


194.1 23.1 17.0 17.4 37.3
181- 20.5- 15.2- 15.1- 32.9-
210 25.3 18.9 19.7 41.7
18 18 18 18 16


186 20.2


232 27.4

245

232.9


14.2 18.1 31.6


19.4 23.2 42.5

- 42.7

36.4


233.9 14.7 27.3 20.8 -
215- 12.9- 23.4- 17.0- -
255 17.1 31.2 25.6 -
15 15 15 15 -







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Table 3 Continued


total proximal proimal shaft distal
length width depth width width


distal
depth


Puma concolor (fossil)
Silver Springs
UF51221

Femur
Puma concolor (Recent)
x
O.R.

N
Puma concolor (fossil)
Wekiva River
UF 124386
Warm Mineral Springs
UF 23870
Miracinonyx inexpectatus
Inglis
UF 45353
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Caves

Tibia
Puma concolor (Recent)
x
O.R.

N
Miracinonyx inexpectatus
Inglis
UF 45350
UF 45351
UF 45354
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave6


241 17.0 28.1 26.8



269.3 51.1 25.3 19.3 47.9
252- 45.5- 22.8- 16.1- 43.3-
290 56.3 27.7 21.7 52.2
18 17 17 17 16


46.1 22.0 -

52.3


293 57.9 28.0 21.1 54.5

334 27.3 59.8


291.7


- 22.2 55.1


249.9 50.0 51.7 18.1 36.5
234- 44.9- 45.7- 15.5- 33.3-
265 55.2 56.9 20.5 39.8
18 18 18 1 8 18


58.4 64.5 26.3 -
61.9 63.9 27.6 -
42.0

311 68.0 41.9


289.2 58.8


- 42.6


Proxim depth of una is anteropotor dephroimdepofolm .th of feuri nteropoteior deph of fnom l hed.
SShft width of ulna is maximum anteropostmior depth of shaft
Mean of 8 individuals (from Van Valkenbunh d a 1990, able 3).
SMean of 10-1 individuals (from Van Vlkenbuh t a. 1990, table 3).
Mean of individuals (from Van Va nburgh et al 1990. table 3).
Mean of5 individuals (from Van Valkburgh ct al 1990. table 3).







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 193





Table 4. Measurements of metacarpals and metatarsals of Pleistocene and Recent Puma concolor and Plio-
Pleistocene Miracinonyx from Florida. Comparative measurements are also provided for Miracinonyx
inexpectatus from Hamilton Cave, West Virginia andM. trumani from Natural Trap Cave, Wyoming (from
Van Valkenburgh et al., 1990). The mean (x) and observed range (O.R.) are provided for measurements on
the Recent sample ofP. concolor, which consists of 18 individuals (10 males, 8 females) from Collier and
Dade counties in southern Florida. The measurements for M. trumani are also mean values, but have
different sample sizes for each metapodial (see footnotes).


total proximal proximal shaft distal
length width depth width width


Metacarpal H
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Cutler
UF 124379
Miracinonyx inexpectatus
Inglis IA
UF 45464
Hamilton Cave
USNM 401092
Miracinonyx cf. M. inexpectatus
Lecanto 2A
UF 128349
Miracinonyx trumani
Natural Trap Cave'

Metacarpal IH
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Cutler
UF 124380
Ichetucknee
UF 45420
Wekiva River
UF 124387
Miracinonyx inexpectatus
Hamilton Cave
USNM 401092
Miracinonyx cf. M. inexpectatus
Lecanto 2A
UF 128350
Miracinonyx trumani
Natural Trap Cave2


70.3 12.5 17.1


13.1 19.4


7.7 14.2


9.2 14.9

8.5 14.6


- 15.3

10.5 16.8


75.4


105.3


105.3

95.3


19.1 15.8


7.9 12.4


12.4 17.4







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Table 4 Continued


total proximal proximal shaft distal
length width depth width width


Metaarpal IV
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Wekiva River
UF 124388
Miracinonyx inexpectatus
Santa Fe 2A
UF 45382
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave'

Metacarpal V
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Cutler
UF 124381
Monkey Jungle
UF 124385
Wekiva River
UF 124389
Miracinonyx inexpectatus
Hamilton Cave
USNM 401092
Miracinonyx cf. M. inexpectatus
Lecanto 2A
UF 128351
Miracinonyx trumani
Natural Trap Cave4

Metataraul
Puma concolor (Recent)
x
O.R.

Miracinonyx inexpectatus
Inglis IA
UF 45471
Leisey Shell Pit
UF 124172
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave'


72.2 10.1 12.0


96.9

100.4

93.2


13.1 15.6


7.4 11.3


8.8 14.6


- 12.8

;4.5 12.7


8.3 13.2


9.7 16.1


10.4 15.6

9.2 13.9


11.7 17.6


11.2 16.2








MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 195



Table 4 Continued


total proximal proximal shaft distal
length width depth width width


Metatrsal III
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Wekiva River
UF 124390

Metatarsal IV
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Wekiva
UF 124391
Miracinonyx inexpectatus
Inglis 1A
UF 45470
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave'

Metatarsal V
Puma concolor (Recent)
x
O.R.

Puma concolor (fossil)
Cutler
UF 124382
UF 124383
Miracinonyx inexpectatus
Hamilton Cave
USNM 401092
Miracinonyx trumani
Natural Trap Cave7
Miracinonyx sp.
Port Charlotte
FMNH PM 39892


10.4 13.9


102.8
96.1-
110.0


12.2 14.4


129.5

129.5

121.3


18.7 20.5 11.7

14.1 20.9 13.3

- 12.0


-17.1 12.3 8.2
-15.4 12.1 8.5


117.3

110.7


108.3


16.7


9.8 17.1

9.8 14.9


7.9 14.9


Mean of8 individuals (from Van Vanbugh et al. 1990, tab 5).
Mean of 12individmuas (from Van Valibe h et aL 1990 table 5).
Mean of 10-12 individuals (from Van Valkenbgh at l 1990. tble 5)
SMean of 10 individual (from Van Valknbuh e al 1990, tble 5)
SMean of 10 individuals (from Vn Vaenbuh et al 1990 table 5)
Mean of 7 individual (from Van Valnbunrgh t i. 1990. tabl 5)
' Meanof6 individual (from Van Valkmbugh et aL 1990 tble5)






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Lundelius et al. (1987) and Morgan and Hulbert (1995) who placed this boundary
at about 300 ka.
The presence of Bison is convenient for determining that a particular fauna is
Rancholabrean in age; however, this genus is often absent in Florida faunas known
to be Rancholabrean based on other criteria. Morgan and Hulbert (1995) identified
additional species of mammals that are restricted to Florida Rancholabrean faunas.
Several of these species (e.g. Tremarctos floridanus and Smilodon fatalis) are
known from older pre-Rancholabrean faunas elsewhere in North America. Among
xenarthrans, a glyptodont (Glyptotheriumfloridanum) and Jefferson's ground sloth
(Megalonyx jeffersonii) occur in Florida only during the Rancholabrean (Kurtdn
and Anderson, 1980; Gillette and Ray, 1981). Large carnivores restricted to
Florida Rancholabrean faunas (Kurten and Anderson, 1980) include dire wolf
(Canis dirus), Florida cave bear (Tremarctosfloridanus), American lion (Panthera
atrox=P. leo atrox of Kurt6n and Anderson, 1980), and sabertooth cat (Smilodon
fatalis). The living black bear (Ursus americanus) does not appear in Florida until
the Rancholabrean. Three extinct species of rodents, giant beaver (Castoroides
ohioensis), giant capybara (Neochoerus pinckneyi), and a large bog lemming
(Synaptomys australis), are found in Florida only in Rancholabrean faunas. At
least six living species of rodents make their first appearance in the state during the
Rancholabrean (Morgan and Hulbert, 1995), including fox squirrel (Sciurus niger),
rice rat (Oryzomys palustris), cotton rat (Sigmodon hispidus), meadow vole
(Microtus pennsylvanicus), pine vole (Pitymys pinetorum), and muskrat (Ondatra
zibethicus). The extinct tapir, Tapirus veroensis, is typical of Florida
Rancholabrean faunas, although this species does occur in two Florida latest
Irvingtonian faunas, Coleman 2A in Sumter County and Sebastian Canal in
Brevard County (Hulbert, 1995; Morgan and Portell, 1996).
The Rancholabrean Land Mammal Age is further subdivided into the early
Rancholabrean (300 to 130 ka) and late Rancholabrean (130 to 11 ka), with the
boundary between the two subages corresponding to the beginning of the last or
Sangamonian interglacial at about 130 ka (see more detailed discussion in Morgan
and Hulbert, 1995). The beginning of the last interglacial (130 ka) is also the
boundary between the middle and late Pleistocene. Fewer than 10 Florida faunas
are definitely early Rancholabrean in age, whereas several hundred late
Rancholabrean faunas are known from throughout the state. However, some early
Rancholabrean faunas probably have gone unrecognized because only two species
of mammals are known to be restricted to faunas of this age in Florida (Morgan
and Hulbert, 1995), the giant bison (Bison latifrons) and an extinct pine vole
(Pitymys hibbardi). One other large mammal, Cuvier's gomphothere (Cuvieronius
tropicus), had its last occurrence in Florida during the early Rancholabrean.
Late Rancholabrean faunas are considerably more numerous in Florida than
are early Rancholabrean faunas, probably because the younger Rancholabrean
faunas are closer in time to the present (and are thus more likely to have been
preserved) and are more easily dated, both biochronologically and by the






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 197


radiocarbon ('4C) method. Faunas from the last half of the late Rancholabrean (i.e.
faunas younger than 50,000 years BP) can be radiocarbon dated. Surprisingly few
Florida late Rancholabrean sites have been radiocarbon dated, primarily because of
the rarity of wood and charcoal in these sites. We hope that the absolute ages of
more Florida late Rancholabrean sites will be determined in the near future
through direct radiocarbon dating of bone collagen of extinct Pleistocene
megafauna using accelerator mass spectrometry (AMS) (Stafford et al., 1991).
For the great majority of Florida late Rancholabrean sites that have not been
radiocarbon dated, biochronology remains the most reliable method for estimating
their age. At least 10 species of mammals are restricted to late Rancholabrean
faunas in Florida. Three of these are extinct species and seven are living species
that make their first appearance in the state during the late Rancholabrean. Two
members of the extinct Pleistocene megafauna, Panthera atrox and the extinct
bison, Bison antiquus, are found in Florida only during the late Rancholabrean.
The large extinct vampire bat, Desmodus stock (D. magnus, originally described
from Reddick, is a synonym) is rare in Florida, but the few sites in the state from
which it is known are late Rancholabrean in age (Morgan, 1991). Five living
species of rodents, Sciurus niger, Sigmodon hispidus, Pitymys pinetorum, Microtus
pennsylvanicus, and Ondatra zibethicus, first appear in Florida during the late
Rancholabrean, although the latter two species occur in somewhat older faunas
elsewhere (Kurtdn and Anderson, 1980). The black bear (Ursus americanus) is
unknown in Florida faunas prior to the late Rancholabrean, despite its nearly
statewide distribution at present.
Puma concolor first appeared in Florida during the Rancholabrean and still
survives in the southernmost part of the peninsula, albeit in ever-decreasing
numbers. Furthermore, all of the 15 Rancholabrean sites from which P. concolor
is known can be shown to be late Rancholabrean in age, based on either
radiocarbon dates or the presence of one or more of the species of mammals
discussed in the previous paragraph. The Florida panther is not known to occur in
any undoubted early Rancholabrean faunas in Florida, and therefore appears to be
a good biochronological indicator for late Rancholabrean faunas in the state.

Florida Fossil Sites Containing Puma concolor

The map of Florida in Figure 1 shows the locations of the 15 Pleistocene
fossil sites and one Holocene archaeological site in the state from which Puma
concolor is known. Each site has been assigned a number that corresponds to the
numbers in Figure 1. The general location and a brief description are given for
each locality, as well as information on the age and associated mammalian fauna.

1. Ichetucknee River
The Ichetucknee River is a well known late Rancholabrean vertebrate
fauna collected from the bottom of a spring-fed river located 8 km northwest






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


of Fort White, Columbia County, northern Florida (Webb, 1974a; Kurt6n and
Anderson, 1980). No radiocarbon dates are available for the Ichetucknee
River fauna, but a late Rancholabrean age is indicated by the presence of
Panthera a(rox and Bison antiquus, as well as the extant rodents Sigmodon
hispidus, Ondatra zibethicus, and Pitymys pinetorum. Other mammals from
the Ichetucknee River that are characteristic of Florida Rancholabrean faunas
include Canis dirus, Tremarctos floridanus, Smilodon fatalis, Oryzomys
palustris, Synaptomys australis, and Tapirus veroensis.

2. Santa Fe River 1
The Santa Fe River 1 fauna was recovered from the bottom of the Santa
Fe River in the vicinity of Ginnie Spring and Devil's Ear Spring in northern
Florida. In this region the Santa Fe River forms the boundary between
Columbia County on the north bank and Gilchrist County on the south bank.
This site produces a mixed fauna of Blancan and Rancholabrean vertebrates.
The mammals from these two ages are very different at the species level, and
consequently the two faunas are not difficult to separate. There is also a
difference in preservation between the Blancan and Rancholabrean fossils.
The panther mandible reported here is most similar in preservation to the
Rancholabrean fossils from the Santa Fe 1 fauna. Morgan and Ridgway
(1987) discussed the Santa Fe 1 Blancan fauna, and Morgan and Hulbert
(1995) provided a current faunal list. Rancholabrean mammals from Santa
Fe 1 include Canis dirus, Castoroides ohioensis, Ondatra zibethicus, Tapirus
veroensis, and Bison sp.

3. Devil's Den
Devil's Den is a water-filled sinkhole over 20 m in depth located 3 km
northwest of Williston in Levy County. This sinkhole presumably was dry or
nearly so during the late Wisconsinan low sea level stand, at which time
regional water tables were very low as well. The Devil's Den mammalian
fauna was reviewed by Martin and Webb (1974), and Kurt6n (1966) discussed
the large sample of Florida cave bear (Tremarctos floridanus) known from
this site. A radiocarbon date suggesting an early Holocene age of 7000 to
8000 yBP for the Devil's Den fauna (Martin and Webb, 1974) is probably
spurious, as this site is almost certainly late Pleistocene (late Rancholabrean)
in age. Rancholabrean indicators from Devil's Den include Megalonyx
jeffersonii, Canis dirus, Tremarctos floridanus, Smilodon fatalis, Oryzomys
palustris, and Synaptomys australis. Six mammals from Devil's Den, Ursus
americanus, Sciurus niger, Sigmodon hispidus, Microtus pennsylvanicus,
Ondatra zibethicus, and Pitymys pinetorum, are extant species that make
their first appearance in Florida during the late Rancholabrean.






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 199


4. Wekiva River
The Wekiva River fauna was collected from the bottom of the Wekiva
River, 7 km northeast of Gulf Hammock in Levy County, north-central
Florida. This fauna, which includes the extinct giant land tortoise
Geochelone crassiscutata and 17 species of mammals, has not been published
upon previously. A Rancholabrean age is indicated by the presence of Canis
dirus and Bison sp.

5. Orange Lake 2A
The Orange Lake 2A site was discovered in an abandoned limestone
quarry near the town of Orange Lake, Marion County, north-central Florida.
The vertebrate fossils are derived from clays and sands deposited in a narrow
vertical fissure formed in Eocene marine limestone. This unpublished fauna
is dominated by microvertebrates, but there are enough large mammals
present to establish its late Pleistocene age. Diagnostic Rancholabrean
mammals from Orange Lake 2A include Megalonyxjeffersonii, Canis dirus,
Tremarctos floridanus, Sigmodon hispidus, Pitymys pinetorum, and
Synaptomys australis.

6. Reddick 1A
The Reddick IA local fauna is located in an abandoned limestone quarry
about 1 km southeast of Reddick, Marion County, north-central Florida. The
vertebrate fossils are derived from fissure and cave fillings formed in Eocene
marine limestone. Reddick is one of the best known and richest
Rancholabrean vertebrate assemblages from Florida, with more than 50
species of mammals reported (Gut and Ray, 1963; Webb, 1974a; Kurt6n and
Anderson, 1980). The age of the Reddick fauna is Rancholabrean based on
the presence of Megalonyx jeffersonii, Glyptotherium floridanum, Desmodus
stock, Canis dirus, Tremarctos floridanus, Oryzomys palustris, Sigmodon
hispidus, Pitymys pinetorum, Synaptomys australis, Tapirus veroensis, and
Bison sp. Among these species, the extinct vampire, Desmodus stock and
the rodents Sigmodon hispidus and Pitymys pinetorum further indicate a late
Rancholabrean age.

7. Silver Springs
Silver Springs, located 10 km northeast of Ocala, Marion County, north-
central peninsular Florida, is one of the largest freshwater springs in North
America. Fossil vertebrates have been known from Silver Springs and the
Silver Springs Run for some time, but the fauna has not been reviewed.
Hoffman (1983) described a mammoth kill site in the Silver Springs Run,
several kilometers east and downstream from the main spring. Hoffman did
not provide a faunal list for this site, but he did mention the presence of Bison
and a "large cat," in addition to Mammuthus columbi. He did not give the






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


scientific name or identify the skeletal element of the "large cat," but
apparently it is not the same specimen as the Puma concolor ulna reported
here (UF 51221). Hoffman (1983) obtained a radiocarbon date of 9840 yBP
on collagen from a mammoth bone from this site. Despite the early Holocene
radiocarbon date from Silver Springs, a late Rancholabrean age seems more
likely as recent compilations of radiocarbon dates strongly indicate that no
member of the North American Pleistocene megafauna survived after 11,000
yBP (Meltzer and Mead, 1983). The presence of Megalonyx jeffersonii,
Neochoerus pinckneyi, Tapirus veroensis, and Bison sp. confirms a
Rancholabrean age for the Silver Springs fauna.

8. Rock Springs
Rock Springs is located 10 km north of Apopka, Orange County, central
Florida. The vertebrate fauna was collected from Rock Springs, an
underwater artesian spring, and from Rock Springs Run, a small stream that
flows from the spring. Rock Springs has a rich bat fauna (Morgan, 1991)
suggesting that the site was not completely underwater in the late Pleistocene.
Wilkins (1983) reviewed the mammalian fauna from Rock Springs, including
Canis dirus, Tremarctos floridanus, Ursus americanus, Tapirus veroensis,
and Bison sp, all of which are indicative of Florida Rancholabrean faunas.

9. Melbourne
The Melbourne local fauna was collected in the first half of this century,
primarily from a golf course 5 km west of Melbourne, Brevard County, on
Florida's central Atlantic Coast. The rich mammalian fauna from
Melbourne, consisting of 49 species, has been reviewed by several authors
(Gazin, 1950; Ray, 1958; Kurten and Anderson, 1980). The vertebrate fossils
from Melbourne occur above the marine late Pleistocene Anastasia
Formation, which was deposited during the last (Sangamonian) interglacial,
thus indicating a late Rancholabrean age (younger than 120 ka). A
Rancholabrean age for the Melbourne fauna is supported by the presence of
Megalonyx jeffersonii, Glyptotherium floridanum, Canis dirus, Tremarctos
floridanus, Smilodon fatalis, Castoroides ohioensis, Neochoerus pinckneyi,
Sigmodon hispidus, Oryzomys palustris, Synaptomys australis, Tapirus
veroensis, and Bison sp.

10. Seminole Field
The Seminole Field site is located in the town of Seminole, west of St.
Petersburg, Pinellas County, on the central Gulf Coast of Florida. Seminole
Field has a rich mammalian fauna consisting of 46 species (Simpson, 1929).
The vertebrate fossils from Seminole Field were found above a marine shell
bed of the Fort Thompson Formation that was deposited during the last
interglacial, thus suggesting a late Rancholabrean age for the Seminole Field







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 201


local fauna. The mammalian fauna from Seminole Field is similar to those
from Melbourne and Vero and, like those two faunas, is late Rancholabrean
in age (Simpson, 1929; Webb, 1974a; Kurten and Anderson, 1980).
Rancholabrean indicators in the Seminole Field fauna include Megalonyx
jeffersonii, Glyptotherium floridanum, Canis dirus, Tremarctos floridanus,
Smilodon fatalis, Sigmodon hispidus, Oryzomys palustris, Synaptomys
australis, Tapirus veroensis, and Bison sp.

11. Leisey Shell Pit 2
The Leisey Shell Pits are a series of commercial shell pits located 7 km
southwest of Ruskin along the eastern shore of Tampa Bay, Hillsborough
County, west-central Florida. The classic Leisey Shell Pit localities are
mostly early Irvingtonian in age; however, Rancholabrean mammal fossils
were recovered from spoil piles in Leisey Pit 2 (Morgan and Hulbert, 1995).
In addition to the single Puma concolor fossil, other Rancholabrean
mammals from Leisey Shell Pit 2 include Sigmodon hispidus, Tapirus
veroensis, and Bison sp.

12. Peace River
Although fossils from the Peace River in south-central Florida were
among the first Pleistocene mammals reported from the state (Leidy, 1889a),
little has been written about this important fauna since and no current faunal
list is available. The Peace River is over 100 km long and vertebrate fossils
are known from about half its length, particularly in Hardee and De Soto
counties. Both of the Puma concolor fossils from the Peace River reported
here are from Hardee County; one was found near Zolfo Springs and the
second came from near Gardner, about 10 km farther south. Rancholabrean
species found in the Peace River in the general vicinity of the two P. concolor
fossils include Glyptotherium floridanum, Tapirus veroensis, and Bison
antiquus.

13. Warm Mineral Springs
Warm Mineral Springs is a deep, water-filled sinkhole located near
Northport, Sarasota County, southwestern Florida, about 20 km inland from
the Gulf of Mexico. The sinkhole reaches about 70 m in depth, but the
vertebrate fossils were recovered from a ledge about 13 m deep. This ledge
was subaerially exposed during the late Pleistocene when sea levels and water
tables were considerably lower than at present. Remains of extinct
Pleistocene mammals were recovered from the 13 m ledge beneath a layer of
organic material, which was radiocarbon dated at 9 to 11 ka (Clausen et al.,
1975; McDonald, 1990). McDonald (1990) reported 20 species of mammals
from Warm Mineral Springs, including Puma concolor. Only two of these
species are extinct, Megalonyxjeffersonii and Smilodonfatalis, both of which






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


are restricted to Rancholabrean faunas in Florida. Two species of extant
rodents identified from Warm Mineral Springs, Sigmodon hispidus and
Pitymyspinetorum, are unknown in the state prior to the late Rancholabrean.

14. Cutler Hammock
The Cutler Hammock site is located in a sinkhole in a tropical hardwood
hammock 4 km east of Perrine, Dade County, in southernmost peninsular
Florida. This site is less than 5 m above sea level and only about 0.3 km
inland from the Atlantic Ocean. The Cutler Hammock sinkhole formed in
the oolitic facies of the marine late Pleistocene Miami Limestone, which was
deposited during the last interglacial high sea level stand and has been dated
between 140 and 110 ka by the uranium series method (Osmond et al., 1965).
The Cutler Hammock fauna is therefore younger than the last interglacial and
is probably very late Pleistocene (late Rancholabrean) in age, between 20 and
11 ka. Unfortunately, the bones from this site are too highly leached to
provide accurate radiocarbon dates (T. W. Stafford, pers. comm.). To date,
47 species of mammals have been identified from the Cutler Hammock local
fauna, including 16 species of extinct Pleistocene megafauna (Emslie and
Morgan, 1995; Morgan, in press). A taphonomic study of the Cutler
Hammock fauna indicates that many of the bones in the site, particularly
those of Mylohyus nasutus and juvenile Equus, were accumulated at a
carnivore den, probably by the dire wolf, Canis dirus (Emslie and Morgan,
1995). A Rancholabrean age for Cutler Hammock is supported by the
presence of Canis dirus, Tremarctos floridanus, Smilodon fatalis, Oryzomys
palustris, and Bison sp. The presence of Panthera atrox, Sciurus niger,
Sigmodon hispidus, and Pitymys pinetorum at Cutler Hammock restricts the
age of this site to late Rancholabrean.

15. Monkey Jungle Hammock
Monkey Jungle Hammock is located on the property of the Monkey
Jungle tourist attraction, 5 km west of Goulds and 12 km southwest of Cutler
Hammock in Dade County. The Monkey Jungle Hammock and Cutler
Hammock sites are the two southernmost Rancholabrean vertebrate faunas in
the continental United States. These two sites formed under similar
depositional conditions, are close in age, and have a large number of species
in common. The fossils at Monkey Jungle also were found in a sinkhole in a
tropical hardwood hammock developed in the marine late Pleistocene Miami
Limestone. Both the Monkey Jungle and Cutler sites formed during periods
of much lower sea level and correspondingly lower water tables. A very late
Pleistocene (late Rancholabrean) age for these two sites is most likely, despite
the lack of radiocarbon dates. The Monkey Jungle Hammock local fauna has
been discussed several times (Martin, 1977; Ober, 1978; Morgan, 1985;
Morgan 1991). Morgan (in press) provides a current mammalian faunal list






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 203


numbering 41 species, including nine members of the extinct Pleistocene
megafauna. Like Cutler Hammock, the presence of Panthera atrox, Sciurus
niger, Sigmodon hispidus, and Pitymys pinetorum in the Monkey Jungle
Hammock fauna indicates a late Rancholabrean age. Other Rancholabrean
species from Monkey Jungle are Canis dirus, Tremarctos floridanus, and
Oryzomys palustris.

16. Granada Site
The Granada Site (archaeology site number 8Dall) from Miami in
Dade County along the Atlantic Coast of southeastern peninsular Florida is
the only archaeological site in the state from which Puma concolor has been
identified (Wing and Reitz, 1982; Wing, pers. comm.). The Granada Site is
within the historical geographic range of P. concolor coryi.

Miracinonyx inexpectatus (Cope, 1895)

Material Examined

Late Pliocene (late Blancan)

Santa Fe River 2A, Columbia/Gilchrist County line: UF 45382, right metacarpal
IV.
Northport, Sarasota County: ROM 29000, left mandible fragment with p3 and p4;
ROM 32286, incomplete p4.

Latest Pliocene/early Pleistocene (early Irvingtonian)

Inglis IA, Citrus County: UF 21604, left mandible with p3-ml; UF 95766, right
humerus; UF 45347, right humerus missing proximal end; UF 45346, right
radius; UF 45344, left proximal ulna; UF 45464, left metacarpal II; UF
45462, right metacarpal V; UF 45465, 45467, two scapholunars; UF 45353,
left femur; UF 45352, left distal femur, UF 45350, right proximal tibia; UF
45354, right distal tibia; UF 45351, left proximal tibia; UF 45471, left
metatarsal II; UF 45470, left metatarsal IV; UF 18114, 45463, 45469 three
left astragali; UF 45451 left navicular, UF 45452, right cuboid; UF 45468,
left cuboid; UF 45466, right ectocuneiform.
Leisey Shell Pit, Hillsborough County: UF 124172, right metatarsal II.


Taxonomy

Schultz et al. (1985) noted that fragmentary felid specimens are notoriously
difficult to identify to species, and we would add that even relatively complete but






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


isolated specimens should be identified with caution. Van Valkenburgh et al.
(1990) also warned against using functional characters in an analysis of
relationships between carnivores that have evolved in similar directions. This is
particularly true for two closely related genera such as Miracinonyx and Puma.
Although Van Valkenburgh et al. (1990) comparatively illustrated many elements
of Puma with both species of Miracinonyx, their paper was primarily a
phylogenetic analysis and did not provide much data on the differential
identification of individual skeletal elements of these two genera. Taking into
account the problems in identifying isolated felid fossils discussed above, we
tentatively identify new specimens of Miracinonyx here using the limited material
available for direct comparison. In general, postcranial elements of Miracinonyx,
which constitute the majority of the sample described below, are noticeably
elongated compared to the same elements in Puma concolor.
Several fossils of Puma concolor from Florida originally were identified as
Felis inexpectata, specifically specimens from the Rancholabrean Seminole Field
(Simpson, 1929) and Melbourne (Ray, 1958) sites. In his review of North
American fossil pumas, Kurten (1976) referred the specimens from these two sites
to P. concolor, and we concur with his identifications. Kurt6n recognized F.
inexpectata as a larger, longer-limbed, puma-like cat that was restricted to Blancan
and Irvingtonian faunas. Adams (1979) named Miracinonyx as a North American
subgenus of the Old World cheetah Acinonyx, including the Blancan and
Irvingtonian species Acinonyx (Miracinonyx) studeri (=F. inexpectata) and the
Rancholabrean species A. (M.) trumani. Van Valkenburgh et al. (1990) reviewed
the systematics of the North American cheetah-like cats. They elevated
Miracinonyx to the generic level, including the species M. inexpectatus and M.
trumani. These authors also established that Felis inexpectata (Cope, 1895) is the
senior synonym of F. studeri Savage, 1960, in agreement with the conclusion of
Kurten (1976). The Florida fossil record of cheetah-like cats now referred to the
genus Miracinonyx is briefly reviewed here because of the possible ancestral
relationship of M. inexpectatus to Puma concolor (see Van Valkenburgh et al.
1990).

Descriptions and Comparisons

Fossils from four Florida sites are here referred to Miracinonyx inexpectatus
(see map in Figure 4). A right metacarpal IV from the Santa Fe River 2A locality,
a mixed late Blancan and Rancholabrean assemblage, probably was the basis for
the listing of Puma concolor from this site by Webb (1974a). This specimen is
much larger and more elongated than any metacarpal IV of P. concolor from
Florida (see Table 4) and is re-identified as M. inexpectatus. The metacarpal of
Miracinonyx from Santa Fe 2A is almost certainly late Blancan in age. Van
Valkenburgh et al. (1990) referred a mandible and a metatarsal IV from the early
Irvingtonian Inglis 1A local fauna, Citrus County, Florida, to M. inexpectatus.







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY


a.0 .
es 9


Figure 4. Late Pliocene and Pleistocene sites f m Florida containing fossils ofMiracinonyx. Miracinonyx
inexpectatus: 1. Santa Fe River 2A, Columbia/Oilchrist County line, late Blancan; 2. Northport, Sarasota
County, late Blancan; 3. Inglis 1A, Citrus County, early Irvingtonian; 4. Leisey Shell Pit 3, Hillsborough
County, early Irvingtonian. Miracinonyx f. M. inexpectatus: 5. Lecanto 2A. Citrus County,
Rancholabrean; Miracinonyx sp.: 6. Port Charlotte. Charlotte County, Blancan?; 7. Coleman 2A, Sumter
County, late Irvingtonian; 8. Cardinale Site, Gulf of Mexico, 25 km west of Pinellas County,
Rancholabrean?.



Webb and Wilkins (1984) had previously listed Fells cf. F. inexpectata from Inglis
IA. We have identified a number of additional elements ofM. inexpectatus from
Inglis (see list under Material Examined). A more thorough examination of the
substantial felid sample from Inglis IA, which includes Miracinonyx, Smilodon,
Homotherium, and at least one other smaller cat, would almost surely reveal
additional specimens of M. inexpectatus, particularly carpals, tarsals, and
phalanges. Berta (1995) recently identified a right metatarsal II from the early






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Irvingtonian Leisey Shell Pit local fauna in Hillsborough County as M.
inexpectatus.
Measurements of the lower teeth from the Inglis lA mandible (UF 21604;
Fig. 5) referred to Miracinonyx inexpectatus are presented in Table 2.
Comparative dental measurements of M. inexpectatus from the middle
Irvingtonian Hamilton Cave in West Virginia and M. trumani from the late
Rancholabrean Natural Trap Cave in Wyoming are also provided in Table 2 (from
Van Valkenburgh et al., 1990). The Inglis teeth are somewhat smaller than the
teeth from Hamilton Cave but are otherwise very similar. Measurements of the
Inglis teeth fit within the range of variation of the larger sample of Blancan and
Irvingtonian M inexpectatus reported by Kurtdn (1976, table IV).
Two specimens from Northport, Sarasota County, may be referred to
Miracinonyx, probably M. inexpectatus. A left mandible fragment with p3 and p4
(ROM 29000) shows the very short diastema of a large short-faced feline (see
measurements in Table 2). The p4 looks like a large puma, but we cannot match


~ 7rrs~~- .4


Figure 5. Left mandible with p3-ml, UF 21604, of Miracinonyx inexpectatus from Inglis IA, Citrus
County, Florida, earliest Irvingtonian: A, Lateral view, B, occlusal view. Scale bar equals 2 cm.


~ler





MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 207


the morphology of the p3 with any particular species because of an expanded
posterior portion. Nevertheless, at this point we do not think it can represent any
other feline. An incomplete p4 (ROM 32286) appears to be too large to represent
P. concolor and so probably also represents Miracinonyx. The teeth of the larger
Northport specimen (ROM 29000) are slightly larger than those of the Hamilton
Cave M. inexpectatus, whereas the smaller Northport specimen (ROM 32286) is
more similar in size to the comparable tooth (p4) from Inglis. The age of the
Northport faunas is not well known, but they appear to have a mixture of late
Blancan and Rancholabrean elements. The two Northport Miracinonyx specimens
are probably late Blancan. Morgan and Ridgway (1987) reported a maxillary
fragment with P4 of the Blancan canid Borophagus diversidens from Northport.
The sample of postcranial elements referred to Miracinonyx inexpectatus
from Inglis 1A includes three complete limb bones (humerus, radius, and femur)
and several partial limbs, as well as four complete metapodials. Measurements of
limb bones ofM. inexpectatus from Inglis IA are presented in Table 3, along with
comparative measurements of Miracinonyx from Hamilton Cave and Natural Trap
Cave (from Van Valkenburgh et al., 1990). The Inglis humerus (UF 95766)
compares closely in length with the humerus of the Hamilton Cave M.
inexpectatus; however, both the radius (UF 45346) and femur (UF 45353) of
Miracinonyx from Inglis are close to the mean size of specimens of M. trumani
from Natural Trap Cave and are shorter than comparable elements from Hamilton
Cave. Despite their similarity in size to limb elements ofM. trumani from Natural
Trap Cave, the radius and femur from Inglis are here referred to M. inexpectatus,
as are the complete humerus and several partial tibiae. The Inglis limb bones are
within the known range of variation of M. inexpectatus limbs based on
measurements of that species from various Blancan and Irvingtonian sites
elsewhere in North America (Kurtdn, 1976, tables V, VI, VIII, and IX).
Measurements of metacarpals and metatarsals of Miracinonyx inexpectatus
from Inglis 1A, Santa Fe 2A, and Leisey Shell Pit are compared to measurements
ofM. inexpectatus from Hamilton Cave and M. trumani from Natural Trap Cave in
Table 4 (from Van Valkenburgh et al., 1990). Table 4 includes measurements of
five metapodials of M. inexpectatus from Florida: a right metacarpal IV (UF
45382) from Santa Fe 2A; a left metacarpal II (UF 45464), left metatarsal II (UF
45471), and left metatarsal IV (UF 45470) from Inglis IA; and a right metatarsal
II (UF 124172) from Leisey Shell Pit. Overall, the Inglis and Santa Fe metapodials
are somewhat shorter than the comparable elements from Hamilton Cave, whereas
the Leisey metatarsal II is slightly longer than the metatarsal II from Hamilton
Cave. These measurements may reflect an increase in size through time as Santa
Fe 2A (late Blancan) and Inglis 1A (earliest Irvingtonian) are 0.5-1.0 Ma older
than Leisey Shell Pit (late early Irvingtonian) and Hamilton Cave (middle
Irvingtonian) (Morgan and Hulbert, 1995). The Florida Blancan and Irvingtonian
metapodials here referred to M. inexpectatus are all considerably longer than the





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


41t

A
B C


D
Figure 6. Metatarsals in anterior view of Recent male Puma concolor (A, C) from Volusia County,
Florida, and the extinct cheetah-like cat Miracinonyx inexpectatus (B, D) from Inglis 1A, Citrus County,
Florida, earliest Irvingtonian. A Left metatarsal II, Recent P. concolor, UF 24042, Volusia County, B.
Left metatarsal II, M. Inexpectatus, UF 45471, Inglis IA; C. Left metatarsal IV, Recent P. concolor, UF
24042, Volusia County, D. Lft metatarsal IV, M. Inexpectatus, UF 45470, Inglis IA. Scale bar equals
2 cm.

comparable elements in Recent Puma concolor (see Fig. 6; also see comparative
measurements of P. concolor in Table 4).






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 209


Miracinonyx cf. M. inexpectatus

Material Examined

middle to late Pleistocene (Rancholabrean)

Lecanto 2A, Citrus County: UF 128349, right metacarpal I; UF 128350, right
metacarpal Il; UF 128351, right metacarpal V.

Descriptions and Comparisons

Several possibly associated metacarpals, including metacarpal H, H, and V
(UF 128349, 128350, 128351, respectively) from the Rancholabrean Lecanto 2A
local fauna in Citrus County, Florida, are tentatively referred to Miracinonyx
inexpectatus (see Fig. 7). Measurements of the three metacarpals of Miracinonyx
from Lecanto 2A are compared to measurements of metacarpals ofM. inexpectatus
from the middle Irvingtonian Hamilton Cave, West Virginia, and metacarpals of
M. trumani from the late Rancholabrean Natural Trap Cave, Wyoming, in Table 4
(from Van Valkenburgh et al., 1990). The metacarpal H and metacarpal V from
Lecanto are intermediate in size between the comparable metacarpals from
Hamilton Cave and Natural Trap Cave, although they are both closer in size to the
Hamilton Cave M. inexpectatus. The metacarpal Il from Lecanto is exactly the
same length (105.3 mm) as the metacarpal II from Hamilton Cave, and is
considerably longer than the sample of metacarpal ifls from Natural Trap Cave.
The Lecanto metacarpals are tentatively referred to M. inexpectatus based on their
similarity in size and morphology to that species. There are other unstudied fossils
of Miracinonyx from Lecanto 2A, but these specimens consist of rather
undiagnostic carpals, tarsals, and phalanges.

Biochronology

The age of the Lecanto 2A local fauna is of interest because almost all
previously reported specimens of Miracinonyx inexpectatus are from late Blancan
and Irvingtonian sites. Morgan (1991) briefly reviewed the mammalian fauna
from Lecanto 2A, which is Rancholabrean in age based on the presence of Canis
dirus, Tremarctos floridanus, Oryzomys palustris, and Sigmodon hispidus. More
detailed study of the small mammal fauna from Lecanto 2A is required to
determine if the site is early or late Rancholabrean.
The only other possible Rancholabrean record of Miracinonyx inexpectatus is
an isolated P4 from Cavetown, Maryland. Kurten (1976) originally identified this
specimen as Fells (=Puma) concolor, although he noted that the tooth was actually
in the size range ofM. inexpectatus. Van Valkenburgh et al. (1990) referred the
Cavetown P4 to M. inexpectatus. Kurtdn (1976) regarded the Cavetown site to be






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


E

A F
B C -


D


Figure 7. Metacarpals in anterior view of Recent male Puma concolor (A, C, E) from Volusia County,
Florida, and the extinct cheetah-like cat Miracinonyx inexpectatus (B, D, F) from Lecanto 2A, Citrus
County, Florida, Rancholabrean. A. Right meacarpal II, Recent P. concolor, UF 24042, Volusia County,
B. Right metacarpal II, Miracinonyx cf. M. inexpectatus, UF 128349, Lecanto 2A C. Right metacarpal
III, Recent P. concolor, UF 24042, Volusia County; D. Right metacarpal III, Miracinonyx cf. M.
inexpectatus, UF 128350, Lecanto 2A; E. right metacarpal V, Recent P. concolor, UF 24042, Volusia
County; F. Right metacarpal V, Miracinonyx cf. M. inexpectatus, UF 128351, Lecanto 2A. UF 128349-
128351 are probably from a single individual but were catalogued separately, because their association in
the field could not be confined. Scale bar equals 2 cm.


of uncertain age, but probably Rancholabrean, whereas Kurtdn and Anderson
(1980) considered this same fauna to be Wisconsinan (=late Rancholabrean).
There is a substantial gap in time, spanning the late Irvingtonian and much of
the Rancholabrean, between the youngest well-dated specimens of Miracinonyx
inexpectatus from the middle Irvingtonian (Conard Fissure, Arkansas;
Cumberland Cave, Maryland; Port Kenndey Cave, Pennsylvania; and Hamilton
Cave, West Virginia) and the earliest definite specimens of M. trumani from the
very late Rancholabrean (Crypt Cave, Nevada, and Natural Trap Cave, Wyoming).
Miracinonyx underwent significant morphological change during the
approximately 0.5 Ma time period when there is a very poor record of this genus in


210






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 211


North America (Van Valkenburgh et al., 1990). Therefore, specimens of
Miracinonyx from sites of intermediate age, such as the late Irvingtonian Coleman
2A (see below) and the Rancholabrean Lecanto 2A local faunas, are very important
to understanding the evolutionary history of the genus. Unfortunately, there is no
possibility of collecting additional material of Miracinonyx from Coleman 2A or
Lecanto 2A, as both fossil sites have been destroyed by limestone mining
operations.

Miracinonyx sp.

Material Examined

Blancan?

Courtland Waterway, Port Charlotte, Charlotte County: FMNH PM 39892, left
metatarsal V.

middle Pleistocene (late Irvingtonian)

Coleman 2A, Sumter County: UF 45363, incomplete left scapholunar.

Rancholabrean?

Cardinale Site, Gulf of Mexico, 25 km west of St. Petersburg, Pinellas County: UF
69800, partial right radius lacking proximal and distal ends.

Descriptions and Comparisons

Overall, the Coleman 2A scapholunar (UF 45363) compares closely with
Puma concolor (and not Panthera onca or Smilodon gracilis); however, the facet
for the articulation of the uniform is angled more ventrally. Both species of
Miracinonyx have this facet angled more ventrally (Van Valkenburgh et al., 1990,
fig. 9), whereas this facet in Puma concolor is almost vertical. Therefore, we
tentatively identify the Coleman scapholunar as Miracinonyx sp. The diverse
mammalian fauna from Coleman 2A is late Irvingtonian in age (Martin, 1974;
Morgan and Hulbert, 1995).
A partial right radius lacking both the proximal and distal ends (UF 69800)
from the underwater Cardinale Site off the west coast of Florida in the Gulf of
Mexico is tentatively referred to Miracinonyx. The shaft of this specimen is
considerably more elongated than in any radius of Recent Puma concolor from
Florida examined, but is similar to a complete radius of M. inexpectatus from
Inglis IA (UF 45346). Because UF 69800 is incomplete and the site from which it
was recovered lacks age-diagnostic mammals, we are only able to identify this






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


specimen as Miracinonyx sp. The fossils from the Cardinale Site were collected
from the bottom of the Gulf of Mexico at a depth of about 20 m approximately 25
km west of St. Petersburg in Pinellas County. The fauna represents a mixture of
Pliocene marine vertebrates and Pleistocene terrestrial mammals. Fossils of these
two different ages presumably were derived from superposed strata that became
mixed on the ocean floor. The Pliocene taxa include the extinct great white shark
Carchardon megalodon, the marlin Istiophorus sp., a mysticete whale, and the
monachine phocid Callophoca obscura. The partial Miracinonyx radius, four
Equus cheekteeth, and a fragment of proboscidean tusk constitute the Pleistocene
fauna. Although the Equus teeth belong to an advanced species of the genus
typical of the Irvingtonian and Rancholabrean, they do not provide a more specific
age refinement for this fauna. The presence of terrestrial mammals on the
continental shelf in water 20 m deep obviously indicates that these animals
inhabited this region during a period of much lower sea level. It is tempting to
suggest that this fauna lived on the continental shelf during the last glacial interval
(Wisconsinan, between 120 and 10 ka) when sea levels were as much as 100 m
lower than present; however, as many as 20 glacial/interglacial cycles may be
represented in Florida during the late Pliocene and Pleistocene (Morgan and
Hulbert, 1995).
A left metatarsal V (FMNH PM 39892) from Port Charlotte in Charlotte
County, referred to Puma concolor by Seymour (1983), is much larger and more
elongate than the living Florida panther. The measurements of this specimen
compare most closely to those of Miracinonyx trumani from Natural Trap Cave
(Table 4). We hesitate to refer this single metatarsal to M. trumani because this
species currently is known only from several very late Rancholabrean sites in the
western United States (Crypt Cave, Nevada, and Natural Trap Cave, Wyoming),
and we have not had the opportunity to directly compare the Florida specimen with
material from these two localities. Furthermore, several Florida specimens referred
to M. inexpectatus from the late Blancan and early Irvingtonian (including both
limb bones and metapodials, see Tables 3 and 4) are smaller than comparable
fossils ofM. inexpectatus from middle Irvingtonian sites, such as Hamilton Cave,
and are actually closer in size to specimens of M. trumani. The Port Charlotte
fossil is of unknown stratigraphic provenience, although vertebrate faunas from
this vicinity range in age from late Blancan to Rancholabrean.

DISCUSSION

Puma concolor is now known from 15 late Pleistocene localities in Florida
(previously there were only five valid fossil records of the Florida panther from the
state), whereas the Miracinonyx records from Florida have quadrupled from two to
eight. This dramatic increase in the number of fossil sites for Puma concolor and
Miracinonyx reflects a trend in recent studies of Florida's rich late Pliocene and
Pleistocene mammalian faunas in which the published fossil record of a particular







MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 213


species is not necessarily an accurate reflection of its actual distribution or
abundance. Perhaps the most obvious reason for this trend has been a very active
field program over the past 15 years by vertebrate paleontologists from the Florida
Museum of Natural History (FLMNH), concentrating on Florida late Pliocene and
Pleistocene faunas. Furthermore, over this same time period there also has been a
comprehensive curatorial effort to identify and catalog fossils in the FLMNH
vertebrate paleontology collection (e.g. more than 150,000 specimens have been
catalogued since 1980, including both recently collected fossils and uncatalogued
material that had resided in the collection for as much as 30 or 40 years). Through
these combined efforts a vast new wealth of fossils from Florida Plio-Pleistocene
sites has accumulated over the past two decades; however, research and publication
of these fossils has not kept pace with field and curatorial activities.
The most recent comprehensive review of Florida Pleistocene mammals
(Webb, 1974a) is now more than 20 years out of date. Webb (1974a, table 2.1)
listed the mammalian species from most of the late Pliocene and Pleistocene faunas
known from Florida up until the early 1970s, including two Blancan sites, four
Irvingtonian sites, and 35 Rancholabrean sites. Hulbert's (1992) list of the fossil
vertebrates of Florida updated the overall fauna and taxonomy of Plio-Pleistocene
mammals from the state. Several papers have provided current mammalian faunal
lists for Florida late Blancan and Irvingtonian sites (Morgan and Ridgway, 1987;
Morgan and Hulbert, 1995), but very few Rancholabrean faunas from the state
have been studied recently. Based on data from the FLMNH vertebrate
paleontology collection and locality files, as well as information summarized in
several recent publications (e.g., Morgan and Hulbert, 1995), there are now 22
Blancan sites, 19 Irvingtonian sites, and well over 100 Rancholabrean sites (a very
conservative estimate) known from Florida.
The fossil record of large felids provides a good example of the gap between
the published record of Florida Pleistocene mammals and the actual fossil record as
preserved in museum collections, particularly the vertebrate paleontology
collection at the Florida Museum of Natural History. Kurtdn's (1965) review of the
Pleistocene Felidae of Florida summarized nearly 80 years of research on Florida's
rich fauna of large fossil cats, beginning with Joseph Leidy's (1889b) description
of a skull of Machairodus floridanus (=Smilodon fatalis) from a Rancholabrean
fissure fill near Ocala in Marion County. Kurtdn identified four species of large
felids from Florida Pleistocene faunas (number of sites he reported for each species
in parentheses): American lion, Panthera atrox (2); jaguar, Panthera onca (11);
puma, Puma concolor (3); and sabertooth cat, Smilodon fatalis (10). All of the
fossils studied by Kurt6n (1965) were Rancholabrean in age. In a review of North
American Pleistocene jaguars, Kurtdn (1973b) reported P. onca from the late
Irvingtonian Coleman 2A site in Sumter County and added one new
Rancholabrean record of the jaguar from the Waccasassa River in Levy County.
Webb (1974a) added one new site record for Panthera onca, and Webb (1974b)
provided four new site records for Smilodon floridanus (=S. fatalis), bringing to 14







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


the total number of Florida Pleistocene sites from which each of these large cats
had been identified. Webb (1974a, b) also reported the gracile sabertooth,
Smilodon gracilis, from two late Blancan sites and one early Irvingtonian site from
Florida, and the scimitar cat, Homotherium, from the early Irvingtonian Inglis IA
site. Waldrop (1974) reported the only valid Florida record of the scimitar cat,
Homotherium serum, from the late Rancholabrean Reddick 1A local fauna in
Marion County.
Over the past two decades most contributions on Florida fossil felids have
concentrated on late Blancan and Irvingtonian species. Churcher (1984) reported
Ischyrosmilus gracilis (=Smilodon gracilis) from the late Blancan Bass Point
Waterway 1 and El Jobean sites in southwestern Florida. Berta (1987, 1995)
described samples of Smilodon gracilis from the early Irvingtonian Inglis IA in
Citrus County and Leisey Shell Pit in Hillsborough County and the middle
Irvingtonian McLeod Limerock Mine in Levy County. Samples from three Florida
early Irvingtonian sites (Haile 16A, Haile 21A, and Leisey Shell Pit) suggest that a
large, undescribed species of Homotherium inhabited Florida during the
Irvingtonian (Berta, 1995). Van Valkenburgh et al. (1990) reported Miracinonyx
inexpectatus from the early Irvingtonian Inglis IA fauna, the first record of a
cheetah-like cat from Florida. Berta (1995) later added a second record of M.
inexpectatus from the Leisey Shell Pit. Seymour (1993) described a skeleton of
Panthera onca from the middle Irvingtonian McLeod Limerock Mine, the oldest
record of jaguar from Florida, and also reported a large sample of P. onca from the
late Irvingtonian Coleman 2A site.
Morgan and Hulbert (1995) listed the mammal species from five late Blancan
and 11 Irvingtonian sites from Florida, including four species of large felids
(number of Florida sites from which each species has been identified in
parentheses): Smilodon gracilis (11); undescribed species of Homotherium (7);
Miracinonyx inexpectatus (6), and Panthera onca (2). Combining published data
(in particular, Kurtdn, 1965, and Webb, 1974a) and unpublished specimens in the
FLMNH vertebrate paleontology collection, there are six species of large felids
known from Florida Rancholabrean sites (number of Florida sites from which each
species has been identified in parentheses-previously published records listed first,
followed by total number of records including unpublished specimens):
Miracinonyx inexpectatus (0, 1), Panthera atrox (2, 12), Panthera onca (13, 37)
Puma concolor (5, 15), Homotherium serum (1, 1), and Smilodon fatalis (14, 26).
Excluding the two Rancholabrean felids that are extremely rare in Florida
(Miracinonyx inexpectatus and Homotherium serum), the number of sites from
which the four remaining species have been identified has increased dramatically
over the past 20 years, ranging from nearly doubling in the sabertooth cat to an
increase by a factor of three in the jaguar and puma to a sixfold increase in the
American lion. These new data present a considerably more accurate picture of the
Pleistocene distribution and abundance of Florida's large cat fauna than can be
obtained from the published literature.






MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 215


The preceding brief summary of large felids from Florida late Pliocene and
Pleistocene vertebrate faunas establishes their current status (number of species
present, number of Florida sites for each species, and updated taxonomy), and
allows for a preliminary analysis of their biochronology, biogeography, and
possible ecological interrelationships. During the late Blancan and Irvingtonian
there were four species of large felids in Florida. No more than three species occur
in a single site, in part resulting from the differing biochronologic ranges of the
species. Miracinonyx inexpectatus is the only one of these four cats that occurs
from the late Blancan through the late Irvingtonian. Smilodon gracilis ranges
from the late Blancan to the middle Irvingtonian. The large undescribed
Homotherium is well represented only in the early Irvingtonian, although it is
tentatively identified from two late Blancan faunas as well. Panthera onca first
appears in the middle Irvingtonian and still survives in Middle and South America,
although it disappeared from Florida at the end of the Rancholabrean.
The late Blancan Santa Fe River fauna and the early Irvingtonian Inglis IA
and Leisey Shell Pit local faunas all record the association of Smilodon gracilis,
Homotherium, and Miracinonyx inexpectatus. If the fossil record for the late
Blancan and early Irvingtonian (late Pliocene through early Pleistocene, from
about 2.5 to 1.0 Ma) in Florida is considered to be fairly complete, and the
presence of more than 40 faunas of this age would indicate that it is, then these
three species could be considered the typical large felid fauna during this time
period. This fauna, consisting of a very large sabertooth cat, a smaller gracile
sabertooth cat, and a cheetah-like cat, compares reasonably well in body size to a
typical fauna of large African felids that includes lion, leopard, and cheetah. The
similarity to the Recent African large carnivore fauna is reinforced by the presence
in two of the three Florida faunas (Santa Fe and Inglis) of the hyaena
Chasmaporthetes ossifragus (see Berta, 1981). Van Valkenburgh et al. (1990)
discussed a fairly widespread association of large felids from North American
middle Irvingtonian faunas (Conard Fissure, Hamilton Cave, Port Kennedy Cave)
that, like the Florida faunas, includes M. inexpectatus and S. gracilis, but
substitutes Panthera onca for Homotherium. P. onca also replaces Homotherium
in Florida middle Irvingtonian (McLeod) and late Irvingtonian (Coleman 2A)
faunas.
The Florida Rancholabrean large felid fauna is composed of six species, of
which only four, Panthera atrox, Panthera onca, Puma concolor, and Smilodon
fatalis, are widespread in fossil sites in the state. The two rare species,
Homotherium serum and Miracinonyx inexpectatus, each have been identified
from only one Rancholabrean fauna in Florida. The cheetah-like cat is also known
from several localities of uncertain age that may be late Pleistocene. Of the four
common large cats, S. fatalis and P. onca occur in both early and late
Rancholabrean faunas, whereas P. atrox and P. concolor are restricted to the late
Rancholabrean. Two of the Rancholabrean felids, the jaguar and puma, are still
extant. The remaining four species are extinct, although some workers (e.g.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(2)


Kurtdn and Anderson, 1980) consider P. atrox to be an extinct subspecies of the
lion, P. leo.
Four species of large felids are present in four Florida Rancholabrean sites:
Ichetucknee River, Santa Fe River, and Reddick 1 in the northern part of the state,
and Cutler Hammock at the extreme southern tip of the peninsula. Ichetucknee
River, Santa Fe River, and Cutler Hammock all have a large cat fauna composed of
Panthera atrox, Panthera onca, Puma concolor, and Smilodon fatalis, whereas
Reddick has P. onca, P. concolor, and S. fatalis, as well as the very rare scimitar
cat, Homotherium serum. Four additional Florida sites contain three large felids.
The most common association, including P. onca, P. concolor, and S. fatalis,
occurs at Devil's Den, Melbourne, and Seminole Field. The large felid fauna from
Monkey Jungle Hammock is composed of P. atrox, P. onca, and P. concolor.
Three of the four common large cats in Florida Rancholabrean faunas, American
lion, jaguar, and puma, compare reasonably well in body size with a typical
African felid fauna composed of lion, leopard, and cheetah, respectively. There is
no ecological counterpart of the sabertooth cat in living African faunas, although
its large body size and saber-like canines have led most paleontologists to propose
that they preyed upon large, thick-skinned mammals, such as mammoths,
mastodonts, and ground sloths (Kurtdn and Anderson, 1980).
One of the most interesting associations of large carnivores in Florida
Rancholabrean faunas is that of Puma concolor and Panthera onca, the only two
surviving members of the North American late Pleistocene large cat fauna. The
puma and jaguar occur together in 10 of the 15 Florida Rancholabrean sites from
which P. concolor is known. Although only the puma or Florida panther is still
found in Florida, the jaguar is far more common in the fossil record having been
identified from 2 Irvingtonian and 37 Rancholabrean sites in the state. P. concolor
is the only large cat that inhabited much of temperate North America prior to the
arrival of Europeans about 500 years ago. The puma has been extirpated from
large areas of its former range, particularly in eastern North America, and now
occurs only in southernmost peninsular Florida, the western United States and
western Canada, and it is also widespread in the Neotropical Region from Mexico
to the southern tip of South America. Throughout the tropical portion of its range
in Middle and South America, P. concolor coexists with P. onca; however, pumas
generally favor drier, more upland habitats, whereas jaguars more commonly occur
in wetter areas, such as in forests and along rivers (Schaller and Crawhaw, 1980;
Seymour, 1989).
Jaguars now are found primarily in tropical forests from Mexico south to
Brazil, although their historical range extended northward into the deserts of
southern Arizona, New Mexico, and Texas (Seymour, 1989). Moreover, Panthera
onca was widely distributed across the south temperate region of North America
(south of 400 North latitude) in the late Rancholabrean (Seymour, 1989). The
reason for the extinction, or more properly extirpation, of the jaguar from Florida
and elsewhere in temperate North America is not known. It is generally assumed








MORGAN & SEYMOUR: P. CONCOLOR & M. INEXPECTATUS FOSSIL HISTORY 217



that most of the large carnivores that went extinct at the end of the Pleistocene
(e.g., Canis dirus, Tremarctos floridanus, Smilodon fatalis, and Panthera atrox,
among others) disappeared because of the extinction of the large herbivores upon
which they depended for food (Kurt6n and Anderson, 1980). The considerable
reduction in the jaguar's range after the end of the Pleistocene is likely related to
the extinction of many of the species it preyed upon, probably including peccaries
(Mylohyus nasutus and Platygonus compressus), capybaras (Hydrochaeris holmesi
and Neochoerus pinckneyi), armadillos (Dasypus bellus), and tapirs (Tapirus
veroensis). These North American species are all now extinct, although living
species of peccaries, capybaras, armadillos, and tapirs still inhabit the Neotropics
in Middle and South America where they comprise some of the favored prey of
jaguars (Emmons, 1987; Seymour, 1989).

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