|
of the
FLORIDA
MUSEUM OF
NATURAL HISTORY
THE FLORIDA PANTHER Puma concolor coryi: A
MORPHOLOGICAL INVESTIGATION OF THE
SUBSPECIES WITH A COMPARISON TO OTHER
NORTH AND SOUTH AMERICAN COUGARS
Laurie Wilkins, Julio M. Arias-Reveron, Bradley
Stith, Melody E. Roelke, and Robert C. Belden
Volume 40 No. 3. on. 221-269 1997
UNIVERSITY OF FLORIDA
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THE FLORIDA PANTHER Puma concolor coryi:
A MORPHOLOGICAL INVESTIGATION OF THE
SUBSPECIES WITH A COMPARISON TO OTHER NORTH
AND SOUTH AMERICAN COUGARS
Laurie Wilkins1, Julio M. Arias-Reveron2, Bradley M. Stith3,
Melody E. Roelke4, and Robert C. Belden5
ABSTRACT
The endangered Florida panther, Puma concolor coryi, has been the subject of ecological and
biomedical research, but little work has been done on the morphology of the subspecies. Interest in the
morphologic characters that describe the Florida population has increased with the discovery of panthers
outside their known range of southwestern Florida, the acquisition of many more specimens than were
available to earlier researchers, and the first adult specimens and live captures from the Everglades. The
necessity to define morphologic features of Florida panthers also had law enforcement implications.
Characters previously used to describe Puma concolor coryi were quantified and re-evaluated using
statistical methods. All historic and recent specimens from the southeastern U.S. (n=79) were examined for
pelage color, cranial profile and proportions, and other morphological traits. These specimens were
compared to a sample of North American and South American specimens. The characters measured provide
a basis on which to describe the Florida population and discriminate between it and other subspecies.
Specimens of panthers inhabiting the Everglades region differ from the balance of the Florida
population in cranial morphology and other features, a result that is consistent with recent genetic research.
There is no evidence to support a permanent geographic or ecological separation of Florida panthers into two
populations. The best explanation for the observed morphological differences, consistent with historical
information, is that the Everglades panthers are descendants of captive-released animals and may be hybrids.
The extent to which other Florida panthers may have been affected by this possible hybridization cannot be
detected with the current analytical methods. Most specimens recovered in the last 20 years from southwest
Florida exhibit the classic P. concolor coryi morphologic features.
SFlorida Museurn of Natural History, Uiversity of Florida (UF), Gainesvile FL 32611-7800, U.SA.
SInstituto Tecnologio de Costa Rica Centro de Investiacion y Desarrolo on Agiicultra Sostenible pr el Topico Humedo (CIDASTH),
Apartado 223-4400, Ciudad Quesada San Calo, Costa Rica (fonned3y University of Florida IFAS Citrus Research and Education Center,
700 Experiment Station Road, Lake Alfred FL 33850, U.SA.
' Depatment of Wildlife Ecology and Conservtion, University of Floridra Gainesvill FL 32611, U.SA
4 Genetics Laboatory, National Cancer Institute, Frederick MD 21702, U.SA. (fonerdy with the Florida Game and Fresh Water
Connission).
SFlorida Game and Freshwater Fish Commission, 4005 South Main Street, Gainesvie FL 32601, U.SA
Wilkins, L, J. M. Arias-Reveron, B. M. Stith, M. E. Roelke, R. C. Belden. 1997. The Florida panther
Puma concolor coryi: A morphological investigation of the subspecies with a comparison to other North
and South American cougars. Bull. Florida Mus. Nat. Hist. 40(3):221-269.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
RESUME
La pantera de Florida, Puma concolor coryi, amenazada de extinsi6n, ha sido objeto de studios
ecol6gicos y biom6dicos, pero poco se ha hecho acerca de la morfologia de sus subspecies. El interest en los
caractres morfol6gicos que described la poblaci6n de Florida se ha incrementado con el descubrimiento de
panteras fuera de su rango conocido en el sudoeste de Florida, la adquicisi6n de mas especimenes (que antes
no estaban disponibles a previous investigadores) y los pimeros especimenes y captures vivas provenientes de
los Everglades. La necesidad de definir caracteristicas subespecificas de las panteras de Florida tiene
tambien implicancias legales.
Los caracteres previamente utilizados para describir Puma concolor coryi fueron cuantificados y
re-evaluados usando m6todos estadisticos. En todos los especimenes historicos y recientes del sudeste de los
E.E.U.U. (n=79) se examinaron el color de pelo, perfil cranial, proporciones y otros rasgos morfol6gicos.
Estos especimenes, fueron comparados con una muestra representative de Norte y Sudamerica. Los
caracteres medidos proporcionaron la base para la descripci6n de la poblaci6n de Florida y para su
descriminaci6n dentro y entire otras subespecies.
Los especimenes de pantera que viven en la region de los Everglades difieren del resto de la
poblaci6n de Florida en su morfologia cranial y otras caracteristicas, hallazgo consistent con recientes
investigaciones gen6ticas. No existen evidencias que soporten una separacin geogrifica o ecol6gica de la
pantera de Florida en dos poblaciones. La mejor explicaci6n de las diferencias morfol6gicas observadas,
consistent con la informaci6n hist6rica, es que las panteras de los Everglades son descendientes de animals
de cautiverio liberados y que por lo tanto serian hbridos. Con este m6todo analitico no se puede detactar
hasta que punto otras panteras de Florida podrian haber sido afectados por esta hibridaci6n. La mayoria de
los especimenes provenientes del sudoeste de Florida recuperados en los 6ltimos veinte alfos, exhiben los
clasicos rasgos morfol6gicos deP. concolor coryi.
TABLE OF CONTENTS
Introduction ........................................... ............................................... .............................. 223
A know ledg ents........................................................................................ .............................................. 224
A abbreviations ...................................................... ................................................................................. 225
Taxonomic Synonymy of Puma concolor coryi...................................... 225
Historical Distribution............................ ........................... ............................ 226
M methods .................................... ...................................................................................................... 227
Pelage Color....................................... ................................................... .......................................... 230
Pelage Characters and Kinked Tail................................. ..................................................................... 235
Cranial Profile........ .................. ...................................... ..................................... . 240
Cranial Proportions.......................................................................................................................... 247
Discussion and Conclusions................................ ................................................................................. 250
Literature Cited ................................................................................ ..................................... ............ 254
Appendices ...................... .... .................................... .......................... 257
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
INTRODUCTION
The puma (Puma concolor) has the most extensive distribution of all
American carnivores (Cabrera and Yepes 1960). At one time, the range of species
covered almost the entire North and South American continents from northern
British Columbia to Patagonia, and it was found in virtually every habitat from
high mountains to tropical swamps (Young 1946). Consistent with this broad
distribution, the species exhibits considerable geographic variation, and 30
subspecies have been recognized (Goldman 1946). The Florida panther (P. c.
coryi) once ranged the southeastern states from Louisiana throughout the lower
Mississippi River Valley east through the southeastern states. Historically, its
distribution was continuous and intergraded with other populations to the north
and west (Goldman 1946). It has been isolated for at least the past 100 years in the
wild lands of south Florida (Bangs 1899) as human settlement patterns caused the
decimation of adjacent cougar populations. Current population estimates vary
between 30 and 80 individuals in the Big Cypress and Everglades ecosystems
(Belden 1986a, Maehr 1997).
This study reviews the morphological characters of the Florida panther (Puma
concolor coryi). It examines color, cranial morphology and pelage features of the
subspecies in the context of the geographic variation expressed by the species
throughout its range. The Florida panther was discovered and named a subspecies
by Charles Cory (1896). Since that time, there have been only a few published
accounts that provide descriptive information (Bangs 1898, 1899, Nelson and
Goldman 1929, Goldman 1946, Layne and McCauley 1977, Lazell 1981, Belden
1986b). Specimens of cougars from the southeast have always been rare. At the
time of Goldman's comprehensive taxonomic review (1946), only 17 P. c. coryi
museum specimens were available, including three from Louisiana and 14 from
Florida. Layne and McCauley (1977) published weights and measurements for an
additional 15 individuals from Louisiana, Arkansas, and Florida; however, only
five had been preserved as museum specimens. The need to describe the panther
in Florida with a suitable suite of morphological characters has become more
important in the last 15 years with (1) the recovery of panthers from areas outside
their known range in Florida, (2) the probability of escaped or released captive
cats of other subspecies into Florida environment, and (3) problems of verification
in law enforcement issues.
In 1986, cats in the Everglades were captured for the first time with the
initiation of a radio-telemetry study (Smith and Bass 1994). It was noted that the
Everglades cats differed from the panthers in the Big Cypress in size and overall
appearance and in the absence of the two physical traits that had been documented
in the Big Cypress population; namely, the kinked tail and a mid-dorsal cowlick, or
whorl (Belden 1986a, Roelke 1990, Wilkins and Belden, unpubl. data). Genetic
studies revealed that free-ranging panthers in Florida consist of two genetically
distinct stocks that had evolved separately (O'Brien et al. 1990, Roelke et al. 1993).
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Further, the two genotypes are strongly partitioned geographically into the
southeastern (Everglades) and southwestern (Big Cypress) populations. The
presence or absence of the kinked tail followed a similar pattern, being present in
most Big Cypress cats and absent in the Everglades population. O'Brien and his
colleagues (1990) suggested the source of the Everglades genotype may have been
from seven captive cats released into the Everglades National Park between 1957
and 1967 (archives, National Park Service, Washington DC), from the Piper
collection of Everglades Wonder Gardens (Vanas 1976).
A total of 45 panther deaths were documented in Florida between 1972 and
1990, most the result of road mortality, illegal kills, and intraspecific aggression
(Roelke 1990). This new material, preserved as specimens in the collections of the
Florida Museum of Natural History (FLMNH), provided an opportunity to review
physical traits described by earlier investigators. The objectives were to (1)
identify and quantify the traits that best describe the Florida population; (2)
compare the recent specimens to historic museum specimens (pre-1950) to
determine what changes, if any, have occurred over time as a result of isolation or
small population numbers; and (3) discern what morphological differences exist
within the Florida population that might correspond to the reported genetic
differences. In addition to the main objectives, we hoped to resolve some
troublesome identifications associated with cats killed outside the current known
range of the panther, which were thought to be captive released individuals.
ACKNOWLEDGEMENTS
This project was funded in part by the National Park Service (NPS) and the Florida Game and Fresh
Water Fish Commission (GFC). GFC also provided invaluable logistical support with the loan of a State
vehicle, arranged by James Brady, for the transport of the hundreds of pounds of equipment necessary to
conduct the color analysis.
We are grateful to the many museum curators and collection staff who allowed one of us (Iw) to
examine specimens under their care, especially. C. Smart and T. Daeschler (Academy of Natural Sciences,
Philadelphia), G. Musser (American Museum of Natural History) D. Holmes (Arkansas Museum of Science
and History, Little Rock), P. Jenkins (British Museum of Natural History) Collier County (Florida)
Historical Society, J. Bayless (Everglades Regional Collection Center, National Park Service, Homestead
FL) B. Patterson and R. Izor (Field Museum of Natural History) M. Hafner (Museum of Zoology,
Louisiana State University, Baton Rouge) M. Rutzmoser, J. Chupasko, and M. Massaro (Museum of
Comparative Zoology, Harvard University) M. Douglas (Oklahoma State University) M. Carleton and L
Gordon (National Museum of Natural History, Washington) P. Landers (Zoological Museum, University
Wisconsin), and numerous others who provided courteous assistance with loas, correspondence and
telephone inquiries. Access to skin collections was made more difficult because of the need to also
accommodate a 100 Ib spectrophotometer. We thank all those who facilitated this arduous task, especially
Linda Gordan (National Museum of Natural History) and Timothy McCarthy (then of the American
Museum of Natural History). Francisco Bisbal of Venezuela and Andr6s Novaro of Chile, students at
University of Florida, examined specimens in South American Museums while conducting their own
research. Jay Sylvester of the Milton Roy Corporation made the pelage color study possible by his generous
loan ofthe color spectrophotometer.
Henry Setzer originally thought of the contour gauge as a measuring tool for the profile measurement;
he also assisted by examining specimens for me while on museum tours of his own. Linda Chandler, Laurie
Walz, and Wendy Zomlefer, produced the graphics. Oron L Bass, Jr. (National Park Service) provided
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
information concerning the panthers in the Everglades region. Technical advice was provided by Clarence
Abercrombie, Richard Hulbert, Stephen Linda, Rodrigo Medellin and Timothy OBrien, all previously or
currently at the University of Florida. Reviews by Jacqueline Belwood, Michael Kennedy, and Melvin
Sunquist greatly improved the manuscript. Constructive comments were also provided by Oron Bass,
Deborah Jansen, Thomas Logan, and William Robertson. Rhoda J. Bryant and Dianna Carver (Florida
Museum of Natural History) assisted in manuscript preparation. Ideas, insight, and encouragement were
offered by John F. Eisenberg
ABBREVIATIONS
Class designations for populations of southeastern U.S. cougars (P. concolor coryi), see
methods for explanation:
HIST historic museum specimens from Florida, including type specimens (pre-
1950)
RECENT recent museum specimens and living animals from southwestern Florida
(non-Everglades cats), since 1950
GLADES Everglades individuals (southeast Florida)
PIPER captive specimens from the Piper collection (Everlgades Wonder
Gardens)
TEST Test animals with no data or of uncertain identity
ARUND historic museum specimens from Louisiana (formerly P. c. arundivaga)
ARK/LA Louisiana and Arkansas specimens acquired within the last 30 years,
identification to be confirmed.
TAXONOMIC SYNONOMY OF PUMA CONCOLOR CORYI (BANGS)
Felis concolor floridana Cory, 1896. Hunting and Fishing in Florida, p. 109.
(name preoccupied by Felis floridana Desmarest, 1820 (=Lynx floridanus
Rafinesque, 1817). Type from Alapata Flats, north of Lake Okeechobee,
Florida, FMNH 9255, skin only, M adult, Field Museum of Natural History,
Chicago. NOTE: Bangs (1898, p. 234) reported the type locality to be north
of Lake Okeechobee and east of Kissimee River based on conversation with
Mr. Cory.
Felis coryi Bangs, 1899. Proc. Biol. Soc. Wash. 13:15, Jan. 31 (renaming ofF. c.
floridana Cory). Type from "wilderness back of Sebastian, Florida, MCZ
7742, skin and skull, old M adult, Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts. Collected by F. R. Hunter January 1,
1898.
Felis arundivaga Hollister, 1911. Proc. Biol. Soc. Washington. 24:176, June 16.
Type from 12 miles southwest of Vidalia, Concordia Parish, Louisiana,
USNM 137122, skin and skull, M adult, United States National Museum
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
(Biological Surveys Collection), Washington, D.C. Collected by B. V. Lilly,
June 17, 1905.
Felis concolor coryi (Bangs 1899) Nelson and Goldman, 1929. J. Mamm.,
10(4):347. Same as Felis coryi Bangs 1899 (see above).
Systematists prefer the use of Puma instead of Felis Jardine (1834). We
follow this convention according to Wilson and Reeder 1993.
Nelson and Goldman (1929) synonomized Felis arundivaga Hollister 1911,
the canebrake puma, with Felis concolor coryi, the Florida puma. According to
them, the Louisiana pumas did not appear to be satisfactorily separable from the
Florida race, although the skull of type specimen and one other slightly exceed
those from Florida in length.
HISTORICAL AND RECENT DISTRIBUTION
The former range of P. c. coryi was thought to be the austroriparian zones in
eastern Texas or western Louisiana and the lower Mississippi River Valley east
through the southeastern states in general, intergrading to the north with P. c.
couguar, and to the west and northwest with P. c. stanleyana and P. c. hippolestes
(Goldman 1946). Archeological and early records of pumas exist for Alabama
(1880), Arkansas (1821-1920), Georgia (1773-1920), Louisiana (1819-1943), and
Mississippi (1758-1880) (Young 1946, pp. 12, 14, 19, 23, and 26). In recent years
there have been panther reports over much of the historical range (reviewed in
Layne and McCauley 1976) and include sightings or specimens from Alabama,
Arkansas, Louisiana, Mississippi, and Tennessee (Goertz and Abegg 1966, Eaton
1971, Jenkins 1971, Noble 1971, Wolfe 1971, Sealander and Gipson 1973, Lowery
1974, Lowman 1975, Morine 1976). Only seven specimens outside of Florida
could be located: Lousiana (4)-the type specimen (USNM 137122, 1905) from
Concordia Parish, two specimens from Prairie mer Rouge (1800s), and one from
Caddo Parish (1965); Arkansas (2)-Ashley County (1969) and Logan County
(1975); South Carolina (l)-with conflicting data suggesting it was either from
Oregon or South Carolina (records of ANSP) (Appendix 3).
Archeological and paleontological evidence exists for pumas in the
southeastern United States outside of Florida (Parmalee 1960, Kurten 1965, Reitz
and Roe, Univ. Georgia, pers. comm.). Within Florida, Pleistocene fossil have
been recovered throughout the peninsula from Dade County in the south to
Columbia County near the Georgia border (Webb 1974, Kurten 1976, Morgan
1997).
The Florida panther was believed to be extinct in north Florida by the turn of
the century (Bangs 1898) although northern sightings and actual shootings have
been reported since that time (Tinsley 1970, Layne and McCauley 1976, Belden
1986a, and Konecny and Eisenberg 1984). Historic and recent specimens from
Florida (Fig. 1) are restricted to the southern part of the state. The northern-most
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
localities represented in Florida are those of the type specimen locality "wilderness
west of Sebastian" (Bangs 1898), in the vicinity of the current boundary between
Indian River and Brevard counties, New Smyrna in 1859, and the recent find of a
skeleton in Farmton Wildlife Management Area, Volusia Co. (Belden et al. 1988).
In 1983 an illegal kill was made in Palm Beach County, an area not known to
be inhabited by panthers. Eventually, the skull and partial skeleton of that animal
was recovered, but the identity of the cat as a true Florida panther could not be
resolved (Abercrombie 1984, Belden 1986b). A second puma, also of questionable
identity, was killed in Palm Beach County in 1984. In 1988, a young puma was hit
by a car in Jefferson County in North Florida, well outside the current range of
panthers (Roelke 1988). It was later learned that the animal was captive bred, not
of Florida stock, and intentionally released by the owner. There are over 1000
cougars in captivity in the State of Florida (B. Cook, Law Enforcement Division,
Florida Game and Fresh Water Fish Commission, pers. comm.). The need to
develop a means to identify the Florida panther, particularly in law enforcement
issues, was the genesis of this study.
METHODS
Museum specimens of P. concolor coryi were examined and compared to
specimens of puma throughout its range in North and South America. The sample
from the southeastern United States consisted of 72 specimens from Florida dating
back to the mid-1800s and the only seven specimens known from outside of Florida
(Fig. 1, Appen. 2), including one from South Carolina with mixed data. The data
set from Florida contains all specimens, including those of uncertain identity.
In addition to traditional skull measurements, techniques were developed to
quantify non-linear characters of color and cranial profile. The latter was intended
to measure the distinctive nasal contour seen in Florida panthers, the "roman
nose," noted by Goldman (1946). Color was measured in museum skins using a
color spectrophotometer, and they were examined for the presence of a mid-dorsal
cowlick.
Multivariate techniques were employed to evaluate the possible
morphological boundaries of populations (subspecies) and variation within the
Florida population. Principal component analysis (PCA) and Canonical
discriminant analysis (CDA) are multivariate techniques of data reduction that aid
in detecting patterns in the data (measurements of characters) and relationships
between and within classes or groups of individuals (other taxonomic units,
OTUs). With PCA, the sample is not subdivided a prior into discrete groups, and
the characters are unweighted. The principal components (axes) are
representations of the variables (measures), each of which vary in their relative
contribution ("factor loading") to each PC axis. PCA is not designed to
discriminate between groups, merely to aid in distinguishing trends in the data.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
SOUTHEASTERN SPECIMEN LOCALITIES
Total specimens: 72
o recent
* historic (pre-1950)
x questionable origin
4oA
Figure 1. Geographic distribution of Puma concolor coryi specimens examined (dots) from the southeastern
United States. Three groups of specimens are represented: recent specimens since 1950; historical
specimens, including those collected by Charles Cory and Outram Bangs in the late 1800s; and recent
specimens whose identity was uncertain. The most northern records include a single specimen from 1859,
locality listed only as New Smyrna, and the skeleton of a cougar discovered in 1987 in the Farmpton
Wildlife Management Area, Volusia County.
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
CDA is a powerful procedure that maximizes intergroup differences to
portray the relationships of the groups more clearly. CDA utilizes OTUs that have
been divided into groups on the basis of an a priori classification. It may be used
to assign group membership to new specimens, or to describe group differences
and relationships. MANOVA emphasizes the testing of similarity/difference
among centroids of the a priori groups and, in that respect is closely related to
discriminant analysis. MANOVA tests the hypothesis that all a priori groups have
the same multi-dimensional mean centroidd) for the variables measured.
In order to examine the morphometric relationships within the Florida
population, where a priori classification was required, the Florida and southeast
specimens were assigned to the following classes: Florida historic (=HIST, pre-
1950 Florida specimens); Florida recent (=RECENT, non-Everglades Florida cats
since 1950); historic Louisiana specimens (=ARUND); Everglades cats
gladesES); animals from Florida of questionable identity, or specimens with no
data (=TEST), Piper captive cats from Everglades Wonder Gardens (=PIPER); and
the southeastern cats from Arkansas and Louisiana (=ARK/LA) that were collected
between 1965 and 1975 long after cougars were thought to survive in Louisiana
(Goertz and Abegg 1966, Sealander and Gipson 1973). The TEST animals are two
females shot in Palm Beach County outside the known range of the Florida
population, a specimen in the Everglades Regional Collection Center with no data,
the skull of a male cougar found in Volusia County, a male from South Carolina
with mixed data, and a specimen from a private collection recently donated to the
Florida Museum of Natural History with no data. These labels are used throughout
the following discussion. At one time, the canebrake puma from Louisiana was
considered a separate subspecies P. c. arundivaga (Hollister 1911), but Nelson and
Goldman (1929) synonomized it with P. c. coryi since they were unable to find any
distinctive characters to separate it.
Only adults were included in the study. Cougars are sexually dimorphic, with
males being larger than females (Goldman 1946, Kurtdn 1973, Anderson 1983,
Maehr and Moore 1992, Gay and Best 1995). This sexual variation dictates
separate analyses by sex, at least for variables associated with skull measures,
thereby reducing the effective sample size for each subspecies. Skins of juveniles
and those that were notably faded as a result of continuous exposure to light were
eliminated from the color analysis. All data sets were tested for normality prior to
analyses using the Shapiro-Wilk statistic (W). With one exception (cranial
profile), multivariate analyses were done with the Statistical Analysis System (SAS
Institute Inc. 1985). Group sample sizes varied with each statistical procedure,
depending upon the availability and condition of specimens. Many skulls were
damaged or did not have skins, so it was impossible to combine variables, since
complete specimens (skins, or undamaged skulls) were often unavailable. Also,
some characters were qualitative and others quantitative. Therefore, each
character analysis was conducted independently. Because sample sizes and
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
analyses varied, methods and results are combined for each of the four characters
discussed: pelage color, pelage traits, cranial profile, and cranial variation.
PELAGE COLOR
Methods
Cory (1896) reported the pelage of the Florida panther as being "more rufous
or reddish brown" than more northern cats. Goldman (1946), using color
standards developed by Ridgeway (1912), described them as 'tawny' heavily mixed
with black in the mid-line becoming cinnamon-buffy or dull 'clay color' on the
sides of the neck. Florida cats are considered dark, but no melanistic cougars have
ever been authenticated. Goldman (1946, p. 235) described a distinct facial pattern
as: face in general greyish-brown, the blackish areas at the base of vibrissae
prominent, ears black externally. There are subtle color differences between
populations in facial patterns and in color shifts along the flanks, inside of the legs,
and the underside of the neck and chest; however, these were not incorporated as
variables in the quantitative analyses.
A spectrophotometer (Color Scan by Milton Roy Company, Analytical
Products Division, Rochester, New York) was used to measure the color of 282
museum pelts representing 13 subspecies from North and South America. The
instrument measures spectral variables that correspond to dominant wavelength (or
hue), saturation (or chroma), and lightness. In addition, it evaluates each sample
(reading) with respect to its position on a red-green scale and a yellow-blue scale
(from I to 100), with low values toward green and blue and high values toward red
and yellow, respectively. Measurements for dominant wavelength and saturation
can be expressed in several ways, but those used in this study are trichromatic
coefficients for x (=dominant wavelength) and y (=saturation). Each is calculated
from the percentage of the three primary colors required to match the sample being
measured. Lightness (luminous reflection) is expressed as a number on a relative
scale from I to 100.
Seven readings were taken on each skin: three mid-dorsal, three lateral, and
one belly (Fig. 2). Dorsal values represent the darker midline exhibited by many
individuals. Lateral color measurements were taken at the hip, ribs and shoulder.
These represent the predominant color of each pelt. The mid-ventral belly
measurement was eventually discarded because of the tremendous variability
shown by the values as a result of dark basal underfur showing through the lighter
guard hairs of the belly fur. High correlation coefficients (>0.9) for each of the two
back measurements and each of the three lateral measurements allowed reduction
of the dorsal and lateral color variables to one value for each. The final data set
consisted of eight variables: three values each (hue, saturation, lightness) for a
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
FLORIDA PANTHER COLOR MEASURES
Figure 2. Spectrophotometer readings taken for seven regions of the pelt: three dorsal, three lateral, and a
single belly measure. Dorsal values represent the darker midline; lateral measures were taken randomly
within the defined regions and represent the predominant color exhibited by each animal. Only the mid-
dorsal and mid-lateral measurements (*) were used in the subsequent analyses.
single mid-back and a single mid-lateral measure plus a value for degree of red and
one for the degree of yellow for the mid-lateral measure. These eight remaining
variables were subjected to PCA as an exploratory method to determine if the
considerable within- and between-subspecies variation observed formed a pattern
that would warrant further analysis. MANOVA tested the hypothesis that no
overall subspecies differences existed between the means of the Florida population
and those of other North and South American populations.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Results
Color variables were normally distributed (p<0.05 test for normality) in those
subspecies groups represented by large sample sizes, and approached normality in
less well represented groups. No observable color differences could be detected
between males and females, or between historic and recent coryi so these classes
were combined in subsequent procedures.
PCA and MANOVA produced similar results. There was considerable
overlap among both North and South American subspecies as would be expected
given the variation present in the species overall (not shown). However,
examination of the principal components when P. c. coryi is compared to selected
North American and South American subspecies separately reveals patterns that
correspond to the qualitative descriptions given by Goldman (1946). P. c. coryi is
darker than western and northern inland populations from North America (Fig.
3a). There is virtually no difference in color measures between P. c. coryi and
coastal populations from Oregon and Washington (P. c. oregonensis and P. c.
olympus) (Fig. 3b). This also was noted by Goldman (1946, p. 237): "In dark
general color tones coryi approaches the geographically distinct olympus...." P. c.
coryi is less red than tropical subspecies from Guatemala, Costa Rica, Panama,
Venezuela, and Brazil (P. c. mayensis, P. c. costaricensis, P. c. concolor) (Fig.
3c), although the latter cannot themselves be separated from one another.
MANOVA showed significant differences between coryi and most other North
American subspecies, including P. c. hippolestes (p=0.002) and P. c. stanleyana
(p=0.0001), two subspecies with which it presumably intergraded in the past
(Table 1), but it could not be distinguished from northwest coastal populations of
P. c. californica, P. c. oregonensis or P. c. olympus based on color variables
(p>0.1). Among the South American subspecies, the differences in the means of
color variables for P. c. coryi are significant for all subspecies tested except P. c.
araucanus from Chile (p=0.62) and P. c. bangsi from Colombia (p=0.3) (Table 2)
The two GLADES cats, the female cougar from Corbett Management Area
(TEST), and one PIPER clustered within the spread of P. c. coryi values in PCA;
one PIPER is outside the range of variation expressed by P. c. coryi (Figs. 3a, b).
As with the PCA, no significant differences were detected between the means of P.
c. coryi and the Everglades cats or the Corbett female with MANOVA, but P. c.
coryi was shown to be significantly different than the PIPER (p=0.02) (Table 1.)
With the exception of one captive, the PIPER and GLADES cats and test animals
clustered within the range of variation ofP.c. coryi (Fig. 3a, b; Table 1). However,
the colors of these test animals differed in ways that were not measured with the
spectrophotometer.
Table 1. Probability ( 0.05) of difference between the means of color variables for selected North American Puma concolor subspecies, including
corvi and Everglades cats. Variables include measures of hue, saturation, lightness degree of redness, and degree of yellowness for mid-dorsal an
mid-lateral region of skin (probability of no overall species effect 0.0001 MANOVA). Not all subspecies were tested.
coryi ever captive test azteca calif hippo kaibab mayen missoul olm reon stanley
(n=24) (n=2) (n=2) (n=l) (n=36) (n=23) (n= 1) (n=22) (n=4) (n=16) (n= ) (n=-8) (n=23)
coryi .580 .019' .426 .0001* .422 .002* .0001* .003* .0008* .384 .152 .0001*
everglades .624 .326 .032* .424 .196 .125 .185 .322 .917 .870 .045* >
captive .000*1 .0001* .0008* .0002* .018* .012* .002*
hipolestes .560 .0077* .127 .0005* .0003* -
kibabensis .003* .021* .0001* .0001' -
mayensis .238 .0016* .034* -
missoulensis .0007* .029* -
olympus .271 -
azteca .275
californica .0001*
IUF2 male shot by hunter, Combett Munagemnt Are. 0
*r3gnrHt KOp<0.05.
Table 2. Probability ( 0.05) of difference between the means of color variables for selected South American Puma concolor subspecies,
including cory and Everglades cats. Variables include measures of hue saturation, lightness degree of redness, and degree of yellowness
for mid-dorsal and mid-lateral region of skin (probability of overall species effect was 0.0001 MANOVA).
coryi acrocod araucan bangsi borben concolor costrac incarum osgoodi pearsoni
(n=24) (n= 4) (n=9) (n) (15) (n=16) (n=8) (n=6) (n=4) (n=4)
coryi 1.000 .0001* .6205 .2953 .0001* .0001* .0001' .0027* .0107* .0001*
acrocodia .0053* .0856 .0001* .0036* .0380* .0203 .0762 .0001*
araucanus .9680 .0001* .0005* .0041* .0239 .0470* .0003*
bangsi .0003* .0083* .0466* .0977 .1924 .0218*
borbensis .0745 .3084 .0046* .0650 .0001*
concolor .7781 .0615 .2442 .0001*
Significant atp<0.05.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
FLORIDA PANTHER COLOR MEASURES
COMPARED WITH OTHER PUMA
POPULATIONS
a) North America
4, redder -- '
-4 -2 0 2 4
FIRST PRINCIPAL COMPONENT
b) North West Coast
2
\2
-2
I\2 11
Shredder I1.
-4 -2 0 2 4
FIRST PRINCIPAL COMPONENT
Legend on following page.
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
FLORIDA PANTHER COLOR MEASURES
COMPARED WITH OTHER PUMA
POPULATIONS
C) CentraSouth America
334
Z, 4
X 2
1 2 4
S-2
1 4
4
Zi 2
0
-4 N redder --.
-4 -2 0 2 4
FIRST PRINCIPAL COMPONENT
Figure 3. Plots of the first two canonical variables representing color for cougars from North America based
on eight color measures. Numbers represent groupings (subspecies) or individuals as follow: (a) 1, coryi
(Florida); 2, missoulensis (Montana, North Dakota, Canada); 3, hippolestes (Colorado, Wyoming); 4,
kaibobensis (Arizona, Utah, Nevada); 5, mayensis (southern Mexico, Guatemala); E, Everglades. Note
that coryi is darker than western and northern inland populations, and darker but not as red as populations
from southern Mexico and Guatemala. (b) 1, coryi; 2, combined oregonensis and olympus (Washington,
Oregon, Olympic Peninsula); P, Piper cats from Everglades Wonder Gardens; T, test animal (Corbett
female). There is virtually no difference between Puma concolor coryi and animals inhabiting the northwest
coast (c) 1, coryi (Florida); 2, costaricensis (Costa Rica, Panama); 3, osgoodi (Bolivia); 4, concolor
(Brasil, Venezuela). Note that coryi is as dark as but not as red as tropical subspecies from Panama,
Venezuela, and Brasil, although the latter can not be separated from each other. Cougars from mountainous
regions of Bolivia are somewhat lighter and less red than the other three groups.
For example, one of the PIPER cats had dorsal and lateral coloration similar to
Florida cats, but graded to a bright reddish color along the flank, whereas coryi
grades to a dull clay color.
PELAGE CHARACTERS AND KINKED TAIL
Methods
Two other pelage features, white flecks and a mid-dorsal whorl, have
frequently been observed in the Florida panthers. A third unusual trait is that of
the kinked tail, a skeletal feature that is visible externally.
Flecks.- In most Florida cats, the head, neck, and shoulders are irregularly
flecked with white hairs (Goldman 1946), a feature noted by Bangs (1899) as "little
bunches of white hairs, scattered here and there." A certain amount of white
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
flecking may be seen on pumas from any part of their range, but it is much more
prevalent in the Florida subspecies (Goldman 1946). The density of flecks on any
particular animal is correlated with age (Roelke and Wilkins, unpubl. data). They
consist of only a few isolated white hairs or small patches on very young animals,
but old animals have liberal flecking that extends along the back almost to the hip
(Fig 4a). It is generally believed that flecking is caused by ticks, and there are
seven species that comprise the ectoparasite fauna of cougars in Florida. Heavy
infestations of ticks, especially Ixodes scapularis, are associated with open wounds
and scars on both live and dead animals (Forrester et al. 1985). Of 318 museum
skins examined, only four (two from Brasil, one from Panama, one from Peru)
showed the dense flecking seen on Florida cats, although light flecking could
occasionally be seen on cougars from throughout their range. This may reflect a
sampling error, or unusually high densities of Ixodes ticks in Florida compared to
other localities. However, it is also possible that panthers in Florida may be more
sensitive to the bite of the Ixoides tick. Whatever the reason, flecking is
consistently more prevalent in the Florida population. As an environmentally
induced color change and not a genetically inherited trait, it is not considered a
true morphologic character. However, it is useful in recognizing cats from Florida.
Whorl/crooked tail.- The whorl, or cowlick, is a structural reversal of hairs
that occurs mid-back and/or at base of the neck. The mid-dorsal whorl can be an
abbreviated narrow ridge of only four centimeters, but it is more often a
pronounced oblong or tear drop shape up to 30 cm in length (Fig. 4b). The whorl
at the base of the neck is chevron-shaped and may be up to 10 cm long (not
shown); it is quite distinct from the usual swirl that is caused by the change in
direction of hairs in this region. Whorls occur in both sexes and are present at
birth, as seen in four fetuses recovered from a car-struck female. Florida animals
frequently display the mid-dorsal whorl, sometimes the neck whorl, and in a few
instances both appear in the same animal. The mid-dorsal whorl was not
mentioned by early describers although it was distinctly present in many of the
specimens they examined.
The kinked or crooked tail is the result of a modification of the distal caudal
vertebrae. Often the third vertebra from the end is shortened and curved, resulting
in a 90-degree bend in the tail (Fig.4c). The last tail vertebra often is truncated
and also sometimes curved, resulting in a double kink. The kink is palpable
through the skin and is often visible as a curl at the tip of the tail in living animals.
The whorl and kinked tail are considered to be genetic markers of the Florida
subspecies (O'Brien et al. 1990). The two characters are not linked, as animals
occasionally will exhibit one trait and not the other. Skeletons were often not
preserved in collections, so the frequency of this trait in older museum specimens
could not be determined. However, it is visible in two published photographs
panthers shot in the 1940's (Hamilton and Whitaker 1979, pg. 307; Tinsley 1970,
p. 23).
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
FLORIDA PANTHER PELAGE/SKELETAL TRAITS
FLECKS, WHORLS AND KINKED TAILS
Figure 4. Pelage traits of the Florida panther. (a) flecks shown on an old male where flecking is extensive.
(b) mid-dorsal cowlick or whorl. (c) kinked tail, a modification of bones of the tip of the tail, is seen as a curl
in the tail of the living animal. Photographs by R. Belden (top) and M. Roelke (bottom).
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
PRESENCE OF WHORLS IN FLORIDA AND
OTHER POPULATIONS OF COUGARS
' Loclities vgue subspecies nne
Figure 5. Darkened areas represent geographical populations of cougars (subspecies). Numbers represent
number of skins examined. Number of specimens displaying mid-dorsal whorl and frequency (%) shown
adjacent to population.
The frequency of the whorl in Puma concolor was determined by examining
648 skins in museum collections, representing 15 North American and 14 South
American subspecies, including the panthers from Florida (n=49). Sample sizes
for each population varied. In addition to museum skins, live animals (23 Florida,
23 Texas, 50 Colorado) were examined for whorls and kinked tails. At the time
the museum skins were examined, the neck whorl had not been discovered, so no
quantitative data are available for any population outside of Florida.
WILKINS ET AL.: FLORIDA PANTHER MORPHOLOGY
Results
Whorl.- The mid-dorsal whorl was found on skins of cougars throughout
their range, but in very low frequencies compared to its presence in Florida
animals (Fig. 5, 6). It is expressed in six North American subspecies (8 of 456
specimens, or 1.75%) and four South American subspecies (21 of 145, or 14.4%).
Among the South American forms, it is more prevalent in subspecies from Chile
and Argentina (20 of 74, or 27%) (Table 3, Fig. 5). The trait was present in 12 of
16 historic specimens from Florida, dating back to the late 1800s, but was absent in
4 of 6 type series taken by Bangs in the wilderness west of Sebastian. Among all
historic and recent specimens and live captures in Florida, it is more prevalent in
the cats from southwestern Florida (92.6%) than in the cats from the Everglades
and Palm Beach County, Florida (22.2%) (Table 4). By the mid-1980s when this
study was initiated, virtually all cats in the Big Cypress exhibited the whorl,
compared to only 2 of 8 cats from the recent Everglades population. It is
noteworthy that the only skin known of the eastern puma, P. c. couguar (dated
1847 from Greenwich, Rhode Island, in the collections of the Museum of
Comparative Zoology [MCZ 42598] Harvard University) did possess a mid-dorsal
whorl, whereas the type specimen of P. c. arundivaga from Louisiana did not.
Kinked tail.- There is no information available on the occurrence of the
kinked tail in other subspecies or in historic P. c. coryi because post-cranial
skeletons were not preserved. In Colorado, 2 of approximately 50 animals live-
captured had kinked tails, but none of the recent Texas cougars (n=23) did (R
Armstrong,). As with the whorl, kinked tails occur in very high frequencies
Table 3. Number and frequency of whorl in Florida specimens compared to combined North and South
American subspecies.
Total Total % Frequency
Locality Examined Whorl of Occurrence
P. concolor coryi' 72 57 79.2
Total North America(excluding coryi) 456 8 1.75
Total South America 145 21 14.40
Combined ssp. from Chile2
and Argentina 74 20 27.00
coryi includes all Florida specimens and living animals examined, captive specimens included.
Combined total of Chilean and Argentinean specimens are also presented to illustrate the high
frequency of the whor in this subspecies complex (consisting of four subspecies: araucans,
pearoni, puma, palagonica)
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
FREQUENCY OF WHORLS AND
KINKED TAILS
SN = 61 N =49
= whorl
--I kinked tail
N = 145
N = 456
Florida, N. Ame
(excluding
Everglades)
PUMAS
erica
S. America
Figure 6. High frequency of whoris/kinked tails in Big Cypress compared to Everglades and puma
populations in North and South America. Everglades includes two animals from West Palm Beach, only one
of which had a skin.
in the recent specimens and live captures from Florida, and are more prevalent in
southwestern Florida (87.8%) than in southeastern Florida (10%) animals (Table
4, Fig. 6).
CRANIAL PROFILE
Methods
According to Goldman (1946), the distinguishing features of the skull of P. c.
coryi are a broad, flat frontal region, the result of remarkably high-arched nasal
bones. He specifically mentions the outline of the nasals "rising to form a distinct
convexity..."(p. 236) (Fig. 7a), a trait that has become known as the roman nose.
80-
60-
40-
20 -
0 J
N=9
N 10
Everglades,
Florida
WILKINS ET AL.: FLORIDA PANTHER MORPHOLOGY
Table 4. Frequency of occurrence of whorl and kinked tail in specimens and live captures of Florida
cougars.
Whorl % Kink %
Locality n Present Occurrence n Present Occurrence
Southwest Florida' 54 50 92.6 49 433 87.8
Historical Southeast2 9 5 55.5 post-cranial skeletons not available
Recent Southeast3 9 2 22.2 10 1 10.0
Total 72 59
'Al specimens from Big Cypress and other regions west of Shark Slough.
2Specimens taken by Bongo and Cory in the late 1800s
'Includes Everglades and two individuals from Palm Beach County, one of which had a skin.
The cranial profile was duplicated with a carpenter's contour gauge. The gauge
was placed 1/8" to the left of the midline of the skull. The tip of the nasals and the
point at which the contour gauge intersected the temporal line (ridge of bone that
curves forward from the saggital crest towards the post-orbital process) provided
two consistent reference points (Fig. 7a). When the images produced by the
contour were rotated with reference to a horizontal line (Fig. 7b) and
superimposed, the distinctive inflated nasal region of P. concolor coryi becomes
apparent (Fig. 7c). A total of 338 specimens representing 29 subspecies were
measured. Some taxa are represented by a single or few specimens. Each contour
was digitized using the mensuration program Sigma Scan. The images were
interpolated to increments of 0.05 inches. The contours were normalized on both
the X and Y axes; along the X axis to eliminate the variation due to size alone, and
along the Y axis to define the highest point on the curve (highest point is Y=l)
(Fig. 8) In the final data set, each contour measurement consisted of 20 values,
each value representing an increment of 0.05 inches along the profile.
The highest point of the crania of most subspecies is the frontal region, the
nasals gradually sloping from there. In the skulls of most P. c. coryi the frontal
region is flat relative to the highly arched nasals, so the inflated nasal region
becomes the highest point on the P. c. coryi skulls. This is shown in the
comparison of a normalized profile of a Florida panther skull compared to one
from Colorado (Fig. 8). The point on the X axis where Y=1.0 (the highest point)
then becomes a measure of the degree of inflation at the anterior portion of the
cranium. The closer that high point is to X=0, the greater the inflation of the
nasals. The means, standard deviation, and minimum-maximum values for the
highpoint were calculated and compared. The contour values were not normally
distributed. For this reason, and because some classes contained small samples,
the Mann-Whitney-Wilcoxan 2-tailed non-parametric test was used to test for
differences in the means between (1) males and females, (2) historic P. c. coryi and
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
MEASUREMENT OF CRANIAL PROFILE
a) Region of Cranium Measured
flattened
inflated frontal
nasal region
region
b) Profiles of Four Specimens
c) Profiles Shown in Overlay
Figure 7. Measurement of cranial profile. (a) outline of Florida panther skull illustrating inflated nasals
relative to flattened frontal region and region measured with contour gage placed 1/8" to the left of the
midline of the skull; (b) actual contours (from top to bottom) of male coryi, compared to cats from New
Mexico, Colorado, and Texas; (c) contours overlaid, shaded area is the difference between coryi and others.
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
SKULL PROFILES
0 0.2 0.4 0.6 0.8
NORMALIZED PROFILE
Figure 8. Images of profiles interpolated to increments of 0.05 inches and normalized on both the X and Y
axes. The point on the X axis where Y=1.0 (the highest point) becomes a measure of the degree of inflation
of the anterior portion of the cranium. The closer the high point is to X=0, the greater the inflation of the
nasals. Shown is the comparison of a normalized profile of a Florida panther skull compared to that of a
Colorado animal. This illustrates the inflated anterior portion of the skull of the Florida panther compared to
the same area that is more sloping in the Colorado individual.
other pre-defined classes of Florida specimens, and (3) historic P. c. coryi and
other subspecies.
Results
Hypothesis of no difference between males and females could not be rejected
at alpha = 0.05, so sexes were combined in this analysis. When HIST P. c. coryi
specimens are compared to other subspecies, the differences are significant in 15 of
27 subspecies (Table 5a). Notable differences include the North American
subspecies P. c. azteca from Arizona, New Mexico (n=55, p=0.028), P. c.
californica from California (n=30, p=0.0), P. c. hippolestes from Colorado (n=24,
p=0.0) and P. c. stanleyana from Texas (n=28, p=0.0001). No significant
differences were detected between HIST P. c. coryi and North American subspecies
P. c. cougar of eastern U.S.(n=4), P. c. olympus (n=l) and P. c. oregonensis
(n=24), the latter two from northwestern U.S.. Significant differences were
recorded between HIST P. c. coryi and three GLADES cats, two PIPER from
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Table 5a. Mean, standard deviation, and minimum-maximum values for contour measurements of
subspecies (n = 286), excluding coryi. Figure in last column is probability of differences between means of
coryi (historic only n = 18) and other subspecies (p < 0.05 Mann-Whitney-Wilcoxan non-parametric 2-tailed
test).
Standard
Subspecies n Mean deviation min max Z-score Probability
4 .525 .0289 .500-.550 2.297
6 .567 .0983 .400-.650 -
55 .493 .1029 .350-.700 2.9799
3 .533 .0289 .500-.550 2.139
8 .475 .0886 .350-.600 -1.8576
1 .600 .0000 .600-.600 -1.5842
2 .525 .0354 .500-.550 1.6914
30 .547 .0890 .350-.700 -4.1807
1 .450 .0000 .450- .450 0.5940
4 .462 .0479 .400-.500 1.4415
5 .600 .0866 .450 -.650 -3.0032
4 .438 .0750 .350 -.500 0.77924
2 .525 .0354 .500-.550 1.6914
24 .546 .0920 .350-.700 -4.0068
6 .533 .0753 .400-.600 2.7126
22 .539 .0950 .300-.650 -3.6935
6 .500 .0837 .400-.600 2.2621
14 .561 .0789 .350-.650 -3.7672
1 .550 .0000 .550 -.550 1.3862
24 .460 .1021 .350-.600 -1.6904
11 .514 .1206 .300-.650 2.2949
2 .650 .0707 .600-.700 -2.2944
11 .568 .1055 .400-.750 3.5264
2 .525 .0354 .550-.550 -1.6914
1 .350 .0000 .350-.350 -0.71326
4 .450 .0913 .350-.550 -0.95976
28 .536 .0941 .350-.650 3.8647
7 .650 .0500 .600-.750 -3.8613
286
0.0216 *
-
0.0288 *
0.0325 *
0.0632
0.1131
0.0908
0.0000 ****
0.5524
0.1494
0.0027 **
0.4359
0.0908
0.0000 ****
0.0067 **
0.0002 **
0.0237 *
0.0002 ***
0.1657
0.0910
0.0217 *
0.0218 *
0.0004 ***
0.0908
0.4757
0.3372
0.0001 ***
0.0001 ***
See table Sb for anndivaga (Lou na) scores.
*p <0.05
** <0.01
*** < 0.001
****p < 0.0001
Everglades Wonder Gardens, three historic specimens of Louisiana, and the recent
kills from Arkansas and Louisiana (p<0.05). No significant differences can be
reported between HIST and RECENT P. c. coryi or for four TEST specimens from
Florida; namely, the no data specimen from ENP, two Palm Beach females, and
the skull found recently in Volusia County (Table 5b).
The consistently high profile values exhibited by historic P. c. coryi, followed
closely by recent P. c. coryi, illustrate that the inflated nasal region in P. c. coryi
araucanus
arundivaga1
azteca
bangsi
borbensis
brown
cabrarae
californica
capricornesis
concolor
costaricensis
cougar
green
hippolestes
incarum
kaibabensis
mayensis
missoulensis
olympus
oregonensis
osgoodi
patagonica
pearsoni
puma
schoregori
sOderstrami
stanleyana
vancouverensis
Total
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
has the lowest mean value of all groups measured, despite the overlap recorded
(Fig. 9). The wide range of values for most subspecies suggests that the failure of
the analyses to discriminate between P. c. coryi and some of the other groups may
be due to small sample sizes.
Everglades cats differed significantly from the HIST cats regardless of small
sample size. The two cats from the Everglades do in fact have small rounded
crania with flat nasals, more similar to cats from South America. In this case, the
flattened nasals represent a real difference from those of both HIST and RECENT
P. c. coryi.
The failure of the analyses to detect the similarity of the ARUND historic
from Louisiana to HIST coryi highlights one limitation of the technique; namely
the profile scores for three ARUND historical range from 0.4 (inflated nasals) to
0.65 (inflated frontals). The score 0.65 is that of the male type specimen from
Louisiana (=arundivaga), that has, in addition to the inflated nasals, an enlarged
frontal region that obscured the inflated nasals in the normalized profile. The
technique was unable to distinguish between actual and relative differences in the
conformation of the skull.
Table 5b. Mean, standard deviation and minimum-maximum values for contour measurements of coryi
classes (n = 49). Figure in last column is probability of differences between the means of coryi (historic n =
18) vs coryi (recent), Everglades, captive, Test 1, Test 2, Test 3, arundivaga (historic) and arundivaga test
(recent). (p<0.05 Mann-Whitney-Wilcoxan 2-tailed non-parametric test).
Standard
coryi class n Mean deviation min max Z-score Probability
Recent 18 .455 .1041 .300-.600 1.442 0.1493
Piper 3 .567 .0289 .550-.600 2.4553 0.0141*
Everglades 3 .550 .1000 .450 -.650 2.0794 0.0376*
Historic 18 .411 .0832 .350-.600
Test 11 1 .550 .0000 .550-.550 1.3862 0.1657
Test 22 2 .500 .0707 .450-.550 1.4177 0.1563
Test 33 1 .350 .0000 .350 -.350 -0.71326 0.4757
Louisiana (historic) 3 .400 .1323 .400-.650 2.0275 0.0426*
Louisiana/Arkansas (recent) 2 .583 .0763 .500- .650 2.4037 0.0162*
Total 52
2 Test I GLADES 7040 no data speimen from ENP.
3Test2-UF 19077 Canal Pomt, PiBeah Co.; UF 23985 Corbett Management are Palm Beach Co.
Test 3 UF 24042 Skeeton found in Voli Co.
*p<0.05
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 40(3)
POPULATIONS
araucanus
arundivaga
azteca
bangsi
brown
cabrarae
californica
capricornensis
concolor
Everglades coryi -
Historic coryi
Recent coyi -
costaricensis
cougar
discolor
green -
hippolestes
incarum
kaibabensis
mayensis
missoulensis
olympus
oregonensis
osgoodi-
patagonica -
pearson-
puma
schoregori
soderstromi
stanleyana
vancouverensis
0.25
SKULL PROFILES
0.35
0.55 0.65
PROFILE HIGHPOINT LOCATION
Figure 9. Means and standard deviations for value of X when Y=1.0 (the high point) for all populations
measured. Historic coryi shows the greatest inflation of the anterior skull, followed by recent coryi. The
extinct P. c. couguar, the eastern cougar, also shows this tendency, as does oregonensis, cougars from the
northwestern United States.
SI I I I
IIC---------
I LII
SI I I
I I-
I I I
S I i
I I ..I-
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
CRANIAL PROPORTIONS
Methods
Cranial measures provide a suitable method for the study of geographic
variation. According to Goldman (1946), the skull of the Florida panther differs
from that of western subspecies in a number of proportions. Earlier work showed
P. c. coryi could be correctly classified with a high level of confidence when
compared with other North American subspecies using cranial measurements and
statistical methods (Abercrombie 1984, Belden 1986b). Eighteen cranial
measurements were taken on adult specimens of P. c. coryi (n=55), including
historic specimens from Louisiana (n=3) and other North American subspecies
(n=183). Lack of sufficient samples of all South American forms and some North
American subspecies limited the cranial analysis to the use of six subspecies: P. c.
azteca, P. c. californica, P. c. coryi, P. c. hippolestes, P. c. kaibabensis, P. c.
oregonensis. There were too few specimens available of the now-extinct P. c.
cougar, from eastern U.S. to include in this analysis.
Unfortunately, many P. c. coryi skulls were damaged, the result of having
been shot in the head (older specimens) or hit by cars (recent specimens), resulting
in numerous missing values. In all multivariate SAS statistical procedures,
observations with missing variables will be eliminated from the analysis.
Therefore, even with a sufficient sample of specimens, elimination of damaged
individuals and separate analyses by sex would preclude statistical methodology.
Missing values were replaced with mean values for a particular variable which was
calculated using existing values within the appropriate class (i.e. historic males,
historic females, recent males, etc.). Specimens of unknown sex were classified
with discriminant-function analysis also using specimens of known sex from the
appropriate class (as above) to create a calibration data set. These procedures were
necessary only for the Florida data set, since complete skulls of known-sex
individuals from other subspecies groups were selected for measurement. Skulls of
adult males are not only larger, but more angular and massive. Females are
smaller and have a more smoothly rounded brain case and lesser development of
sagittal and lambdoid crests (Goldman 1946). The two sexes might, therefore, be
described by a different set of variables. Stepwise discriminant analysis selected 11
variables as important for discriminating females and 16 for males.
Geographic variation was explored using PCA. Because no discrete clusters
were formed, subsequent CDA was conducted to maximize intergroup differences.
Presented here is only one of several analyses conducted, including only three
subspecies: Florida and Louisiana specimens (P. c. coryi), southwestern U.S. (P. c.
azteca), and Texas (P. c. stanleyana), those groups inhabiting the southern part of
the U.S. This was thought to be an appropriate strategy considering the disjunct
distribution of P. c. coryi, the considerable variation exhibited by the subspecies in
North America, and the limited scope of this study. Also, PCA suggested that
clinal variation might be a confounding factor that could not be resolved with the
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
present data set. The objective was to determine if P. c. coryi could be
discriminated from populations that it most closely resembles and to which it is
closest geographically.
Discriminant function analysis examined possible misclassification of
specimens (p<0.05), using those specimens with reliable data as the base
calibration group. Suspected hybrids (cats from the Everglades), others lacking
data or with uncertain data, and those animals that could not be reliably identified
as Florida panthers were treated as a test group.
Results
The three groups produced only two canonical variables, which together
explained 100% of the variation. Canonical variable I (CV1) explained 81% of the
variation for females, and 80% for males, with 19% and 20% respectively being
explained by CV2. The measurements contributing to CV1 for females were total
length, zygomatic breadth, pterygoid width, and upper carnassial crown length,
while width of palate pterygoid width, condyle width, and maxillary tooth row
were major contributing variables to CV2. For males total length, zygomatic
breadth, condylobasal length, and post-orbital process breadth contributed to CVI,
and zygomatic breadth, maxillary tooth row, and palatal width at canines
contributed most to CV2 (Table 6). The plots of the canonical variables showed
groups much more discrete than with other analyses (Fig. 10). As expected, HIST
and RECENT P. c. coryi overlapped considerably. A single female Florida cougar
was within the range of variation expressed by the Texas group. Test animals fell
either on the periphery or outside the range of P. c. coryi, as did the two females
from the Everglades. The only test animal that could confidently be assigned to P.
c. coryi was the skull found at the Frampton Wildlife Refuge in Volusia County in
1987.
The results of the discriminant analysis follow the CDA plots in that animals
outside the range of variation of P. c..coryi were reclassified (Table 7). However,
because only three groups were represented, the individual being reclassified was
placed in the group it most closely resembled. This confounds the interpretation,
but is nevertheless instructive. Cats killed within the last 20 years in Louisiana
and Arkansas were reclassified into the Texas subspecies P. c. stanleyana and are
probably not relicts from the original population inhabiting those states but either
new introductions from the west or escaped captive individuals. The two
Everglades cats, the Canal Point cat and one Piper captive were reclassified as P. c.
azteca. The skeleton found in Volusia County was reclassified as P. c. coryi. The
captive Piper male from Everglades Wonder Gardens, which although well outside
the range of variation of P. c. coryi, was more similar to it than to any of the other
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
FLORIDA PANTHER CRANIAL MEASURES
COMPARED TO OTHER NORTH AMERICAN
POPULATIONS
MALES
-4 0 4
FIRST CANONICAL VARIABLE
FEMALES
-4 0 4
FIRST CANONICAL VARIABLE
Figure 10. Plots of the first two canonical variables for skull and dental measurements: 1 = Florida historic,
2 = Florida recent, 3 = Louisiana historic, 4 = Texas, 5 = Arizona and New Mexico, E = Everglades, P =
Piper (captive), T = Test (of uncertain origin). One observation hidden in each graph.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Table 6. Details of canonical correlation analysis for three subspecies ofPuma concolor (coryi, stanleyana,
azteca). Coefficients having dominant loadings on a standardized variable are shown in bold type and
discussed in the text
Females Males
Canonical variable Canonical variable
1 2 1 2
Canonical correlation .89 .68 .87 .66
Eigenvalue 3.70 .87 3.07 .76
Proportion of variance explained .81 .19 .80 .20
Cumulative proportion explained .81 1.00 .80 1.00
Standardized canonical coefficients
Total length -0.84 -0.59 -1.15 -0.07
Condylobasal length 1.85 0.05
Zygomatic breadth 0.87 -0.33 1.18 -1.80
Cranium height -0.67 0.44
Width of palate 0.50 1.08 0.44 -0.11
Palatal length -0.22 -0.80
Maxillary tooth row -0.49 134 0.09 1.14
Mandibular tooth row 0.62 -0.54
Mastoid breadth -0.26 0.27 -0.52 -0.04
Post-orbital process breadth -1.61 -0.01
Postorbital constriction 0.56 -0.11 0.55 -0.16
Width at canines -0.30 136
Pterygoid width 0.72 -0.88 -0.01 -0.15
Condyle width 0.15 -0.70 -0.11 0.08
Upper carnassial crown length -0.74 0.00 -
Upper carnassial crown width -0.50 -0.54
groups so was reclassified as P. c. coryi. It is interesting to note that the historic
Louisiana males remained within the P. c. coryi sample, however the Louisiana
female did not. Both Everglades cats were reclassified.
DISCUSSION AND CONCLUSIONS
Small sample sizes and problematic techniques limit the interpretive value of
some of the results. Had it been possible to combine the characters measured,
include a broader representation of populations, and have more robust samples, a
more cogent picture of the relationship between Florida and other populations
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
Table 7. Reclassification ofspecimens ofadult male and female Florida cougars according to discriminant
function analyses. Specimens of uncertain identity were tested against a calibrated data set ofPuma c. coryi
(n=45), P. c. azteca (n=29), and P. c. stanleyana (n=21), separate analysis by sex, equal prior probability.
Individual Reclassified Posterior
Reclassified Identity To Class Probability
MALES
LSU 11363 ARK/LA stanleyana 0.40
LSU 17032 ARK/LA stanleyana 0.97
MSH 1379 ARK/LA stanleyana 0.88
UF 12462 Captive coryi 1.00
ANS 2241 So.Carolina azteca 0.99
UF 24042 Volisia Co. coryi 0.99
FEMALES
EVER 7040 ENP specimen stanleyana 0.41
UF 19077 Canal Point azteca 0.78
UF 23985 Corbett coryi 0.77
UF 24557 Everglades(#27) azteca 0.97
UF 24563 Everglades (#15) azteca 0.98
might have emerged. Within the limited scope of the study, however, some
conclusions might be drawn.
The geographic races of the cougar, like those of other animals are based on a
combination of characters, including size, color, and cranial and dental measures
that prevail in areas over which environmental conditions tend to be uniform
(Goldman 1946). Puma concolor coryi, best known from the Florida population,
appears to be well defined based on pelage markings, color, and the cranial profile.
None of these characters is unique in itself; however, in combination, they provide
a basis to describe the Florida population, whether or not one accepts the concept
of a subspecies. Goldman (1946) placed heavy emphasis on the inflated nasals as a
distinguishing feature of the Florida panther. Quantitative measures of this trait
reinforced Goldman's view of its importance in identifying individuals belonging
to the Florida population. Of North American subspecies examined, the Florida
population most closely resembles cats from the northwest coast in details of color
and cranial profile. This may reflect a similarity of some environmental parameter
that prevails over the two geographic areas (for example, high humidity levels) and
deserves further investigation.
Two Piper cats from the Everglades Wonder Gardens exhibit significant
differences when compared to both the historic and recent specimens of P. c. coryi
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
based on cranial measures and skull contour. The baseline color of one of the
Everglades cats was within range of variation of Florida panthers, but qualitative
details of the pelage markings would preclude this assignment. These two animals
had not been included in the genetic study, but came from the same breeding
compound that produced the captive animals released into the Everglades.
Cats of questionable origin or suspect data defied classification. They
clustered apart from or on the periphery of the Florida sample. The one exception
to this is the skull recovered from Volusia County that could most confidently be
assigned to the Florida population. Individuals containing a hybrid strain, with the
exception of the Everglades cats, could not be detected within this study, unless it
is the single individual that was killed in Canal Point in Palm Beach County,
which was not included with genetics study.
The two cats from the Everglades differ significantly from the rest of the
Florida cats in cranial profile and in cranial morphology, in addition to the absence
of the kinked tail and mid-dorsal whorl. These results are consistent with those of
the genetic studies that show the distribution of genetic markers to be strongly
partitioned between Big Cypress and Everglades ecosystems. The morphologic
differences between the Big Cypress and Everglades cats cannot be explained by
geographic separation within Florida either now or in the distant past. This would
tend to support the view put forth by O'Brien et al. (1990) that the cats inhabiting
the Everglades are descendants of the captive and probably hybrid cats that were
released into the Everglades.
The results of genetic work conducted by O'Brien et al (1990) suggest that
little genetic mixing has taken place between the Everlades and Big Cypress
populations, yet no permanent physiographic barrier exists between the Everglades
and Big Cypress systems. Virtually no differences could be detected between the
historic P. c. coryi morphotype and the recent Big Cypress panthers. The only
indication of change is the slightly diminished arched nasal profile in the recent
cats. This could be the result of dilution of the character through hybridization, but
it may also be a sampling error, or genetic drift as a result of small population size.
Considering that perhaps 30 years has elapsed since captive cats were released into
the Everglades, the consistent dichotomy between the Everglades cats and Big
Cypress cats is open to conjecture.
The Everglades is a vast and complex system, but short term cycles of
drought and excessive rainfall do effect water levels in some regions (Duever et al
1994). Shark River Slough, a deep marsh in the western Everglades, when flooded
might produce a cyclical but effective barrier limiting dispersal of animals from
either side. Radio-collared animals in the Everglades were not documented
crossing the Slough until 1989, when a severe drought caused lowered water levels
(Bass and Maehr 1991), and then only three of ten monitored animals ever crossed
from the Everglades side (Bass 1997). Dispersal around the Slough to the north
would be limited by development and habitat fragmentation, while the
WILKINS ET AL.: FLORIDA PANTHER MORPHOLOGY
Caloosahatchee River is an effective barrier between north and south in the
southwestern part of the state (Maehr 1997).
Distribution of panthers in south Florida is limited by the availability of
suitable habitat and prey (Smith and Bass 1994). Between 1986 and 1991, radio
telemetry studies revealed that female panthers in the Everglades utilized home
ranges entirely east of the Slough, as did their female offspring (Bass 1997). In
southwest Florida, females remained in or near the home ranges of their probable
mothers (Maehr et al 1991). Young female recruitment into natal home range is a
pattern repeated in other cougar and felid populations. Due to high home range
stability for adult males, vacant home ranges for young dispersing males are
limited, and mortality is greatest in subadult and non-resident males (Maehr et al.
1991). High rates of aggressive encounters between males of Florida panthers
compared to other cougar populations, and the highest mortality factor of male
cougars in Florida (Dunbar 1997), may be a further indication of limited dispersal
opportunity as a function of suitable habitat.
A combination of factors that include cyclically high water levels in Shark
River Slough, limited habitat availability, and the dynamics of puma behavior
could explain the high degree of isolation that existed between the two populations
of cougars in south Florida and that would have resulted in a reduction in gene
flow and shared morphologic traits. With this recent extended dry period recorded
in Florida, the effects of introgression may become more conspicuous as animals
cross more freely between the two refuges.
Frequencies of whorls and kinked tails have changed over time. Four of six
cats collected and studied by Outram Bangs between 1896 and 1898 did not have a
mid-dorsal whorl. Among recent panthers of known origin from the Big Cypress
Swamp between 1972 and 1987, all had kinked tails (n=30), and 25 of 27 (92%)
had the back whorl. In 1986-1987, six panthers were captured in the Everglades
National Park (ENP); none had a kinked-tail and only one had the whorl. In 1988
the first BCS animal with a straight-tail and whorl was documented (#25 rogue).
By early 1990, four straight-tailed panthers (three of which had whorls) were
documented in the BCS and the first cat from ENP (#39) with both the whorl and
the kink was captured in ENP (Roelke 1990). This suggests there has been more
mixing between the two areas in recent years.
The presence of a mid-dorsal whorl and kinked-tail clearly identifies a cat as
being from the native Florida population, regardless of the genetic interpretation.
The expression of these traits in the Florida populations in high frequencies has
been considered a morphologic indicator of inbreeding and reduced levels of
genetic variability. Of six North American subspecies of cougar, Roelke et al.
(1993) demonstrated that the authentic Florida panther (excluding Everglades cats)
exhibits less variation than any other puma subspecies with the fewest polymorphic
loci (P) 4.9%, and low heterozygosity (H) with 1.8%, compared to (P) of 27% and
(H) of 1.8-6.7% for other subspecies. This is nearly as low as the level of allozyme
variation reported in the cheetah. This condition has undoubtedly resulted from
Appendix 1 Continued.
Catalog No. Sex Age COND'-4 BW1' NW"'5 F1"' K1'' Date Comments
UF 10425
UF 10460
UF 10481
UF 10573
UF 11852
UF 11915
UF 11927
UF 12004
UF 12462
UF 12823
UF 12827
UF 14390
UF 14699
UF 16374
UF 18798
UF 18944
UF 19077
UF 19090
UF 19096
UF 20957
UF 20958
UF 20777
UF 20973
UF 22409
UF 22529
UF 23985
UF 23986
UF 24042
UF 24096
M F !ALC P A A P 1981 Fetus G-81-19
M F !ALC P A A P 1981 Fetus G-81-19
F F !ALC P A A P 1981 Fetus G81-19
F F !ALC P A A P 1981 Fetus G81-19
F 4 S,SK,SKL A A A A 1979 Captive Everglades Wonder Gdns
F 1+ !S,SK,SKL P P 2 P 1979 G-80-4
M 2 S,SK,SKL P P 2 P 1980 G-80-15
M U !S P A 1 NA 1940s Rug
M 3 S,SK,SKL A A 1 A 1981 Captive Everglades Wonder Gdns
U 2 !SK NA NA NA NA 1940s? Collected by R. Allen
U 1 !SK NA NA NA NA 1940s? Collected by R. Allen
M 3 S,SK,SKL P A 2 P 1978 G78-65
(F) 3 PS,SK,PSKL P A 3 NA 1940s? Collected by R. Allen
M 3 SK,SKL A A P P 1982 FGC #6 "blunt trauma", Skin examined
by R. Belden in life
F 3 S,SK,SKL R A P P 1983 FGC #3 Skin mounted in Tallahassee
M 2? !S,SK,SKL R A P P 1983 Palmdale, Glades Co., skull fragmentary,skin not
tanned; cowlick could be "coaxed"
(F) 2 SK,SKL NA NA NA A 1983 Palm Beach Co. illegal kill
F 4 SK,PSKL P A P P 1983 FGC #5 skin examined by R. Belden
M 4 S,SK,SKL P A 5 P 1983 FGC #1
(F) 1+ SK,SKL NA NA NA P 1985 "Bones"; found by film crew G85-BNZ
F 3 !SK,SKL NA NA NA P? 1985 G84-26 "squash", specimen mutilated,no record
of pelage characters; no kink, but abnormal
blunt tail tip
M 5 PS,SK,SKL P A 5 P 1984 FGC #2 nasal region destroyed, skin not tanned.
F 1+ SK,PSKL NA NA NA NA 1985 Butchered female, tail missing G85-C-S
M 4 S,SK,SKL P A 4 P 1985 FGC #7 "raccoon"
M 4 S,SK,SKL P A 5 P 1985 FGC #4 "tattoo"
F 3 S,SK,SKL A A 2 A 1984 Corbett Management Area
M 2 PS,SK,PSKL P A P P 1987 FGC #10 killed by FGC #12
(M) 3 SK,PSKL NA NA NA NA 1987 Volusia Co. tip of tail missing
M 3 !S,SK,SKL P A 3 P 1988 GFC #13 "big Al", skull
fragmented
Appendix 1 Continued.
22*
Catalog No.1 Sex2 Age3 COND' BW1,' NW1.' F'l K1's Date Comments
UF 24097*** M 1 !S,SK,SKL A A A A 1988 G88-16 Jefferson Co.
UF 24160 (M) ? !SKL NA NA NA P 1979 Gannet Strand
UF 24267 F 5 S,SK,SKL P A 5 P 1988 FOC #8
UF 24268 F 3 S,SK,SKL P A 4 P 1986 FGC #PCO59
UF 24314 M 4 S,SK,SKL P A 4 P 1988 GFC #20
UF 24315 M 3 S,SK,SKL P A 2 A 1988 GFC #25 "rogue"
UF 24316 M 2 SK,SKL P A P P 1988 GFC #24 Highlands Co.; examined in life by
M. Roelke
UF 24561 M 2 S,SK,SKL P A 2 P 1989 89-64 Corkscrew Sanctuary
UF 24557 F 3 S,SK,SKL A A 2 A 1989 GFC #27 Everglades
UF 24563 F 4 PS,SK,SKL A A P A 1988 GFC #15 Everglades, skin in poor condition
J Billie M 3 S,SK P P 4 P7 1983 Illegal kill; kink discernible in skin; specimen
examined at time of trial
UF 24595 M !S,SK,SKL P A 3 P 1989 GFC #33 rabies
UF 24611 M !S,SKSKL P A 3 P 1990 GFC #35 subadult
UF 24621 M !S,SK,SKL P A 3 P 1990 GFC #30 killed by another male
Total specimens examined = 72: 49 Skins (2 partial); 60 Skulls; 37 Skeletons (6 partial); 4 Fluid preserved
USNM U.S. NtionaMueum; MCZ Museum of Coampaive Zooloy MCZB denotes Bp' collection; FMNH Field Musmea of Naural istoy, ANSP Philadelphia Acadmy Sciene; UWZ
Univesity of Wisconn; EVER Everglades Ntioal Prk CCM Colier Coty Mumem (Florid); U Florida Mue of Naturl ltoy, University ofFlorids; T-type specimen; COND-Codliin;
BW-BackWhorl; NW-NockWhorl; F-Flocb; K-Kink.
SM-nmu; F-fmce; ()n me or femae bied d size of adult idividl U camtbe determind.
Alp cl based a fsion of skull futures: 1 -juvenile or subdult, all utues open, size small; 2-younm adult, aus distit ad e fsion, sase nal; 3-adult, partial fusion of ;
4-imture dult, most or all sutue fused; 5-old, sue idlistinct; F-ftus; + indicates inermdiate between classes.
s Type of pncrtion: S-ski, SK-skull, SKL-pot cranial saleton, P-partial, ALC-alcohol.
SOccurrece of pelas a seletal fetma: P-preas, A-habrn, R-unlimesary (see tet), NA- not applicable whn skin orpost-crmnlsBlaeteis not available for inpetion.
Category of flecb: 1- <5; 2-5-20; 3-20-50, 4- >50< 100; 5-> 100; A-absees, P-proesu, ibt not evaluhaed due to poor condition or abm aki vals e stind
SUSNM19483 deleed fiom tdy; phased from Wads Scitific n Miami, noted speimoe iporer a suipplier; no fthr da
SFMNH91621 no data, bt believed to be Cory secirn FMNH14901 previous mountedad thought discarded; based on oolor ad polap a s.
SSpecimn not believed to be Florida pater based color; captive released ito north Florlda (nfomatioa from Florida Gams & FiahCeaminsiao); skulmlanot iachded inalyas because of yo
aubadult
I Skulls included in crani amlyi because miuia, juvenile, framed or not yet promised;
Appendix 2
Specimens of southeastern Puma concolor, outside of Florida, examined
Back Neck
Catalog No.1 Sex2 Age3 Condition4 Whorl5 Whorl5 Flecks6 Kink5 Data Comments
USNM 137122T M 3 S,SK A A 3 NA 1905 LA,Vidalia; type specimen of
Fconcolor arundivaga
(Hollister)
USNM 1157 (F) 2 SK NA NA NA NA 1800s LA, Prairie mer Rouge
USNM 1158 (M) 3 SK NA NA NA NA 1800s LA, Prairie mer Rouge
LSU 17032 M 3 SK,SKL NA NA NA NA 1969 AR, Ashley Co.; tip oftail
missing, near LA border
LSU 11363 M 2 S,SKSKL A A A A 1965 LA, LaCaddo Parish, near
Keithville; skin mtd.; pelage
features per M.Haffner LSU;
no kink apparent
MSH 1379 M 3 SK NA NA NA NA 1975 AR, Logan Co.
ANSP 2241 (M) 2 SK NA NA NA NA 1800s SC, data questionable
LUSNM U.S. National Museum ofNatural History, LSU Louisiana State Univenity, MSH Akansas Museum of Science and Histoy
NSP Academy of Natural Sciences of Philadelphia
3. e; F-famale; () male or female based on size of adult individual
' Definitions same as Appendix 1.
WILKINS ET AL. FLORIDA PANTHER MORPHOLOGY
Appendix 3
Live animals examined for kinked tail and whorl (1980-1990)'
Number Locality Whod Kink Comments
Yes Yes
R Yes
Yes Yes
Yes Yes
No No
R Yes/No
Yes Yes
Yes Yes
Yes Yes
No No
No No
No No
R Yes
Yes Yes
R Yes
Yes Yes
Yes Yes
Yes Yes
Yes No
No No
Yes No
Yes Yes
R No
Shot in foot
White Oak
parent of GFC #19
parent of GFC #19
mother of #16 & #21
modified kink3
"Jumbo"
"Jumbette"
Price cat
offspring of#14
offspring of#15
from #15 "Annie"
Whorl single ridge
Whorl single ridge
First to show no
kink or whorl
First to show both
kink and whorl
GFC #9
Big Guy
GFC #11
GFC #12
GFC #14
GFC #16
GFC #17
GFC #18
GFC #19
GFC #21
GFC #22
GFC #23
GFC #26
GFC #28
GFC #29
GFC #31
GFC #32
GFC #34
GFC #36
GFC #37
GFC #38
GFC #39
GFC #40
Total
Big Cypress
Total 17
Whorl absent 1(6%)
Rudimentary 4(23.5%)
Whorl present 12(70%)
No kink 4(23.5%)
Kink 13(76.5%)
Everglades
Total 6
Whorl absent 4(66.6%)
Rudimentary 1(16.6%)
Whorl present 1(16.6%)
No kink 5(83.3%)
Kink 1(16.6%)
23
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Everglades
Everglades
Big Cypress
Big Cypress
Big Cypress
Everglades
Everglades
Everglades
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Big Cypress
Everglades
Big Cypress
pture data fom IR Belden and M. RoelkeO Game and Flmida Game and Freshwater Commission
R=namentsy, approximately 5cm o less single ridge
mod kinkpresent at first capture, but absent at later capture. Kink ofa different shape (sideways) than those observed in Big Cypress
cats; no explanation, unless due to an injury informationn fnm D. Janzen M. Roelke pers. comm)
APPENDIX 4
Color variables for Puma concolor (coryi). List of all measures for mid-back and mid lateral: V1, V10 = hue; V2,
V11 saturation; V3, V12 = lightness; V19 = degree of red; V20 = degree of yellow for lateral.
Catalogue No. Class VI V2 V3 V10 Vll V12 V19 V20
US 528190 coryi 0.42140 0.38172 28.93000 0.38615 0.37570 48.93000 7.65000 18.04000
US 265596 coryi 0.42266 0.38209 27.43000 0.38656 0.37471 42.42000 7.20000 16.09000
MC 19855 coryi 0.41196 0.37800 27.48000 0.39150 0.37678 45.19000 8.20000 17.89000
MC 86992 coryi 0.41173 0.37938 29.16000 0.48422 0.38006 36.43000 8.84000 17.31000
MC 85489 coryi 0.41435 0.37986 28.37000 0.40158 0.38140 36.29000 8.03000 17.21000
MC 87743 coryi 0.40450 0.37274 25.60000 0.40477 0.38116 34.79000 8.45000 17.02000
MC 87742 coryi 0.41749 0.37673 29.49000 0.400071 0.38057 34.64000 7.78000 16.43000
MC 5650 coryi 0.40565 0.37195 24.84000 0.40755 0.38325 35.58000 8.69000 17.96000
AM 144512 coryi 0.42419 0.38392 30.36000 0.38884 0.37659 46.57000 7.80000 17.92000
FM 14902 coryi 0.42302 0.38768 30.64000 0.40038 0.38427 44.51000 8.35000 20.34000
FM 14900 coryi 0.34076 0.38914 29.58000 0.40016 0.38631 42.84000 7.61000 20.15000
FM 1255T coryi 0.42736 0.38531 29.25000 0.39062 0.38079 42.87000 6.77000 17.89000
UF 9789 coryi 0.41559 0.38415 32.93000 0.38820 0.37634 45.63000 7.59000 17.52000 Z
UF 22529 coryi 0.42239 9,37523 31.04000 0.38673 0.37541 47.14000 7.65000 17.56000
UF 14390 coryi 0.42094 0.38448 30.39000 0.39548 0.38022 44.94000 8.26000 19.02000
UF 11927 coryi 0.41989 0.38490 8.15000 0.38722 0.37608 48.14000 7.72000 18.04000
UF 12004 coryi 0.43287 0.39457 37.25000 0.39287 0.38354 51.85000 7.66000 21.56000
UF 19096 coryi 0.41287 0.37981 30.26000 0.38384 0.37280 47.39000 7.63000 16.72000 7
UF 11915 coryi 0.41325 0.37931 30.99000 0.38688 0.37343 44.37000 7.82000 16.41000 0
UF 10424 coryi 0.42330 0.38549 32.86000 0.38488 0.37401 47.97000 7.65000 17.26000 ^
UF 22409 coryi 0.41857 0.38027 29.72000 0.48283 0.38073 46.22000 7.62000 20.50000 <
UF 24096 coryi 0.42447 0.38372 29.79000 0.38624 0.37444 47.51000 7.82000 17.40000 0
,az
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
the loss of habitat and more than 150 years of persecution. Examination of historic
specimens of the late 1800s shows that the whorl was not fixed in the population
by the late 1800s, as it was missing from four of the original six Bangs specimens,
but the frequency was nevertheless high (present in 5 of 9 specimens, or 55%,
from the late 1800s).
The process leading to lowered levels of variability and expression of unusual
traits might have begun much earlier and be explained in part by the peninsular
nature of Florida in which genetic exchange with more northern populations was
restricted on three sides. The Florida land mass receded to an even narrower
peninsula than it is today when sea levels rose to their present level 8000 BP
(Watts and Hansen 1988). Furthermore, climatic changes accompanied by a shift
to less productive soils, and a vegetation community dominated by pine flatwoods
by 5000 BP might further limit population numbers. Together these would have
created conditions that would limit dispersal and intergradation with more
northern forms. This view gains some support in the high frequency of the whorl
in cats from Chile and Argentina (27% overall, but higher within individual
populations) and the peninsular nature of those two countries (Eisenberg and
Redford 1982).
The Florida panther exhibits a combination of unique and shared characters
that are measurable and quantifiable. Further, the morphotype remains relatively
unchanged from the early historic specimens of the late 1800s in spite of a possible
introgression with another form. This does not suggest, however, that managed
outbreeding is undesirable. The loss of genetic variability and associated problems
of lowered reproductive potential and immunological deficiency (Roelke et al.
1993) that threaten this population, in addition to loss of habitat, requires
immediate implementation of management decisions, and habitat preservation.
LITERATURE CITED
Abercrombie, A. 1984. Subspecific identity of Felis concolor UF 19077 by discriminant analysis. Report
to Game and Fresh Water Fish Commission.
Anderson, A. E. 1983. A critical review of literature on puma (Fells concolor). Colorado Div. Wildl.,
Spec. Rept. 54:1-91
Bangs, 0. 1898. The land mammals of peninsular Florida and the coast region of Georgia. Proc. Boston
Soc. Nat. Hist., 28(7):157-235.
1899. The Florida puma. Proc. Biol. Soc. Wash., 13:15-17.
Bass, O. L, Jr. 1997. Ecology and population dynamics of the Florida Panther in Everglades National
Park. Pp. 82-97 in D. B. Jordan, ed. Proceedings of the Florida Panther Conference, 1-3 November
1994, Ft Myers, Florida.
, and D. S. Maehr. 1991. Do recent panther deaths in Everglades National Park suggest an ephemeral
population? Res. Explor. 7(4):427.
Belden, R. C. 1986a. Florida panther recovery plan implementation-a 1983 progress report. Pages 159-
172 in S. D. Miller and D. D. Everett, eds. Cats of the world: biology, conservation and
management. Proc. 2nd international cat symposium, Caesar Kleberg Wildl. Res. Inst.,
Kingsville,TX, and the Natl. Wildl. Fed., Washington, D.C. 501 pp.
1986b. "Florida panther characteristics", Recovery plan implementation, annual performance
report. Study No. E-1-10 I-E-5a, Florida Game and Fresh Water Fish Comm. 13 pp.
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
W. B. Frankenberger, R. T. McBride, S. T. Schwikert. 1988. Panther habitat use in southern
Florida. J. Wildl. Manage. 52(4):660-663.
Cabrera, A. and J. Yepes. 1960. Mamnifferos Sud Americanos, 2nd., 2 vols. Buenos Aires: Ediar.
Cory, C. B. 1896. Hunting and fishing in Florida. Estes and Lauriat, Boston. 305 pp.
Duever, J. J., J. F. Meeder, L C. Meeder, and J. M. McCollom. 1994. The climate of South Florida and its
role in shaping the Everglades ecosystem. Pp. 225-248 in S. M. Davis and J. C. Ogden, eds.
Everglades the ecosystem and its restoration. St. Lucie Press, Delray Beach, Florida.
Dunbar, M. R. 1997. Florida panther biomedical investigations. Pp. 342-393 in D. B. Jordan, ed.
Proceedings of the Florida Panther Conference, 1-3 November 1994, Ft. Myers, Florida.
Eaton, R. L 1971. Florida panther. Natl. Parks and Cons. Mag., 45(12):18-20.
Eisenberg, J. F. and K. H. Redford. Comparative niche structure and evolution of mammals of the Nearctic
and southern South America. Pages 77-86 in M. A. Mares and H. H. Genoways, eds. Mammalian
biology in South America. Special publication series, Vol. 6. Pymatuning Laboratory of Ecology,
University of Pittsburgh, Pittsburgh, PA.
Forrester, D. J., J. A. Conti, and R. C. Belden. 1985. Parasites of the Florida panther (Felis concolor
coryi). Proc. Helminthol. Soc. Wash. 52(1):95-97.
Gay, S. W. and T. L Best 1995. Geographic variation in sexual dimorphism of the Puma (Puma
concolor) in North and South America. The Southwestern Nat. 40(2):148-159.
Goertz, J. W. and R. Abegg. 1966. Pumas in Louisiana. J. Mamm., 47:727.
Goldman, E. A. 1946. Classification of the races of the puma. Pages 175-302 in S. P. Young and E. A.
Goldman, eds. The puma, mysterious American cat. American Wildlife Institute, Washington, D. C.
Reprinted (1964) by Dover Publ., Inc., New York. 358 pp.
Hamilton, W. J., Jr. 1943. The mammals of eastern United States; an account of recent land mammals
occurring east of the Mississippi. Comstock Press, Ithaca, N.Y. 432 pp.
__and J. O. Whitaker, Jr. 1979. Mammals of Eastern U.S., second edition Cornell Univ. Press, Ithaca,
N.Y.
Hollister, N. 1911. The Louisiana puma. Proc. Biol. Soc. Wash. 24:175-177.
Jenkins, J. H. 1971. The status and management of the bobcat and cougar in the southeastern states. Pp.
87-91 in S. E. Jorgensen and L D. Mech, eds. Proceedings of a symposium on the native cats of
North America; their status and management USDI, Fish and Wildl. Serv., Bur. of Sport Fisheries
and Wildl. Region 3. 139 pp.
Konecny, M. and J. F. Eisenberg. 1984. On the captive breeding and reintroduction of the Florida panther
into suitable habitats. Report prepared for the Florida Game and Fresh Water Fish Commission.
Kurt6n, B. 1965.The Pleistocene Felidae of Florida. Bull. Florida State Mus. 9(6):215-273.
1976. Fossil puma (Mammalia: Felidae) in North America. Netherlands J. Zool. 26(4):502-534.
Layne, J. M. and M. N. McCauley. 1977. Biological overview of the Florida panther. Pages 5-45 in P. C.
H. Pritchard, ed. Proceedings of the Florida panther conference Florida Audubon Society in
cooperation with the Florida Game and Fresh Water Fish Commission. 121pp.
Lazell, J. D. Jr. 1981. Diagnosis and identification of the races ofFelis concolor in eastern North America,
a preliminary report. U. S. Fish and Wildlife Service, 1 November. 22 pp. mimeographed.
Lowery, G. H., Jr. 1974. The mammals of Louisiana and its adjacent waters. Louisiana State Univ. Press,
Baton Rouge. 565 pp.
Lowman, G. E. 1975. A survey of endangered, threatened, rare, status undetermined, peripheral, and
unique mammals of the southeastern national forests and grasslands. U. S. Dept Agric. Forest Serv.,
Southern Region. 132 pp.
Maehr, D. S. 1997. The comparative ecology of bobcat, black bear, and Florida panther in south Florida.
Bull. Florida Mus. Nat. Hist. 40(1):1-176.
E. D. Land, and J. C. Roof. 1991. Social ecology of Florida panthers. Res. Explor. 7(4):414-431.
and C. T. Moore. 1992. Models of mass growth for three North American cougar populations. J.
Wildl. Mgmt. 56:700-707.
Morgan, G.. 1997. Fossil history of the panther (Puma concolor) and the cheetah-like cat (Miracinonyx
inexpectatus) in Florida. Bull. Florida Mus. Nat. Hist. 40(2):177-219.
Morine, D. E. 1976. Preservingthe Pascagoula. Nat Conservancy News 26(4):12-16.
Nelson, E. W. and E. A Goldman. 1929. List of the pumas, with three described as new. J. Mamm.,
10:345-350.
Noble, R. E. 1971. A recent record of the puma (Felis concolor) in Arkansas. Southwestern Nat, 16:209.
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
O'Brien, S. J., M. E. Roelke, N. Nuki, K. Richards, W. Johnson, W. L Franklin, A. Anderson, O. L Bass, R.
C. Belden, and J. S. Martenson. 1990. Genetic introgression with the Florida panther (Felis
concolor coryi). Nat. Geo. Res. 6(4):485-494.
Pamnalee, P, W. 1960. Animal remains from the Etowah Site, Mound C, Barrow Co., Georgia. Florida
Anthrop. Vol. XIII nos. 2-3.
Ridgeway, R. 1912. Color standards and color nomenclature. Published by Ridgeway, Washington, D.C.
Roelke, M. E. 1990. Florida panther biomedical investigation 1 July 1986 30 June 1990. Study No.
7506, Florida Game and Fresh Water Fish Comm. 199 pp.
_, J. S. Martenson, and S. J. O'Brien. 1993. The consequences of demographic reduction and genetic
depletion in the endangered Florida panther. Current Biology, 3(6):340-349.
Sealander, J. A. and P. S. Gipson. 1973. Status of the mountain lion in Arkansas. Proc. Ark. Acad. Sci.
27:38-40.
Smith, T. R., and O. R. Bass. 1994. Landscape, white-tailed deer, and the distribution of Florida panthers in
Everglades National Park. Pp. 693-708 in S. M. Davis and J. C. Ogden, eds. Everglades the
ecosystem and its restoration. St. Lucie Press, Delray Beach, Florida.
Tinsley, J. B. 1970. The Florida panther. Great Outdoors Publishing Co., St. Petersburg. 60 pp.
Vanas, J. 1976. The Florida panther in the Big Cypress Swamp and the role of Everglades Wonder Gardens
in past and future captive breeding programs. Pages 109-111 in P. C. H. Pritchard, ed. Proceedings
of the Florida panther conference. Florida Audubon Society in cooperation with the Florida Game
and Fresh Water Fish Commission. 121pp.
Watts, W. A., and B. C. S. Hansen. 1988. Environments of Florida in the late Wisconsin and Holocene. Pp.
307-323 in B. A Purdy, ed. Wet Site Archaeology. The Texford Press, Inc., Caldwell, New Jersey.
Webb, S. D. (ed.). 1974. Pleistocene mammals ofFlorida. Univ. Presses ofFla., Gainesville. 270 pp.
Wilson, D. E. and D. M. Reeder, eds. 1993. Mammal species of the World. Smithsonian Institution Press,
Washington, D. C. p. 296.
Wolfe, J. L 1971. Mississippi land mammals. Distribution, identification, ecological notes. Miss. Mus.
Sci., Miss. Game and Feish Comm., 44 pp.
Young, S. P. 1946. History, life habits, economic status, and control. Pages 1-173 in S. P. Young and E.
A. Goldman, eds. The puma, mysterious American cat. American Wildlife Institute, Wash. D.C..
Reprinted by Dover Publications, Inc., New York (1964).
APPENDIX 1
Florida specimens of Puma concolor examined
Catalog No.I Sex Age3 COND'4 BW1' NW1,5 F'6 K1.3 Date
U U S,PSK
(F) 2 SK
M 4 S,SK
M 4 S
A A 0 NA
NA NA NA NA
P A 5 NA
P P 5 NA
USNM 19483*
USNM 145264
USNM 265596
USNM 528190
resident
MCZB 5489
MCZB 5650
MCZB 6992
MCZB 7742T
MCZB 7743
MCZB 7744
MCZ 31781
MCZ 10703
MCZ 19855
MCZ 54423
FMNH 1255T
FMNH 14900
FMNH 14902
FMNH 50058
FMNH 91621**
ANSP 15270
ANSP 16705
ANSP 16706
AMNH 100222
AMNH 144512
UWZ
EVER 7040
CCM
UF 9789
UF 10424
M U !S,SKL P
F 2 S,SKSKL A
A 4 NA
P 3 P
1859
1940
1975?
1896
1896
1897
1898
1898
1898
1934
1922
1973
1895
1896?
1896?
1939
1896?
1933
1935
1935
1946
1900
Comments
Purchase, Wards Scientific
New Smyrna
Purchase from E.Ross Allen
Confiscated at tanner 1977; belonged to Florida
Type F.concolor coryi (Bangs)
near Everglades (city?) O
"E.Florida", Wyman Collection
Everglades, NE Flamingo
Type F.concolorfloridana (Cory)
Marked M, but probable F (Cory)
Cory specimen
No data, probable Cory specimen
F 4 S,SK A P 5 NA
F 3 S,SK A A 3 NA
M 4 S,SK A A 3 NA
M 3 S,SK P P 3 NA
F 4 S,SK R A 4 NA
(M) 1 !S,SK A A 1 NA
(M) 3 SK NA NA NA NA
(M) 3 SK NA NA NA NA
F 4 S,SK P A 4 NA
(F) 1 SK,PSKL NA NA NA NA
F U !S P A 4 NA
F 2 S,SK P A 3 NA
(M) 3 S,SK P A 4 NA
F 2 SK NA NA NA NA
U U !S P A 3 NA
(M) 4 SK NA NA NA NA
(F) 4 SK NA NA NA NA
(F) 3 SK NA NA NA NA
(F) 3 SK NA NA NA NA
M 2 S,SK P A 3 NA
(F) 3 SK NA NA NA NA
(F) 3 SK NA NA NA NA
U U !S P A 3 ?
No data
Mtd. specimen, Earl Brown Coll.,
kink present in photo (Kelly's)
1972 Glades Co., tail missing
1981 w/4 fetuses G-81-19
Appendix 4 continued.
Catalogue No. Class VI V2 V3 VIO V11 V12 V19 V20
UF 24268 coryl 0.42702 0.38487 31.33000 0.38978 0.37504 45.20000 8.22000 17.33000
UF 24267 coryi 0.41416 0.38203 33.32000 0.38336 0.37185 46.25000 7.62000 16.16000
UF 24314 coryl 0.41583 0.38259 34.83000 0.38149 0.37211 50.83000 7.62000 17.16000
FM 91621 coryi 0.43707 0.38591 27.75000 0.40735 0.38497 45.28000 6.22000 21.75000
UF 11852 captive 0.42582 0.38027 5.91000 0.39663 0.37803 39.25000 8.20000 16.98000
UF 12462 captive 0.43884 0.38656 30.88000 0.38979 0.37478 45.71000 8.35000 17.42000
UF 23985 corbett 0.42496 0.38235 28.52000 0.39149 0.37716 42.88000 7.80000 17.29000
UF 24557 everglades 0.42117 0.38359 26.59000 0.39562 0.37716 42.88000 7.80000 17.29000
UF 24563 everglades 0.38239 0.36295 26.15000 0.38391 0.37007 33.10000 6.84000 14.81000
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
APPENDIX 5
Sample data set for contour measurement for the type specimen of P. concolor
coryi. X and Y are the coordinates along the anterior portion of the nasal
region; XN and YN are the normalized values. The position on the X axis for
which YN=1.000 is the highpoint and marked in bold type.
Catalog Sex Status X- Y XN YN
MCZB 7742 M HIST 0.43750 0.07176 0.05000 0.09289
MCZB 7742 M HIST 0.87500 0.13107 1.00000 0.16966
MCZB 7742 M HIST 1.31250 0.21641 0.15000 0.28012
MCZB 7742 M HIST 1.75000 0.33917 0.20000 0.43902
MCZB 7742 M HIST 2.18750 0.51020 0.25000 0.66040
MCZB 7742 M HIST 2.62500 0.68299 0.30000 0.88406
MCZB 7742 M HIST 3.06250 0.76805 0.35000 0.99416
MCZB 7742 M HIST 3.50000 0.77256 0.40000 1.00000
MCZB 7742 M HIST 3.93750 0.69983 0.45000 0.90586
MCZB 7742 M HIST 4.37500 0.63546 0.50000 0.82254
MCZB 7742 M HIST 4.81250 0.56039 0.55000 0.72537
MCZB 7742 M HIST 5.25000 0.52478 0.60000 0.67927
MCZB 7742 M HIST 5.68750 0.49976 0.65000 0.64689
MCZB 7742 M HIST 6.12500 0.46806 0.70000 0.60586
MCZB 7742 M HIST 6.56250 0.43376 0.75000 0.56146
MCZB 7742 M HIST 7.00000 0.37115 0.80000 0.48042
MCZB 7742 M HIST 7.43750 0.31526 0.85000 0.40807
MCZB 7742 M HIST 7.87500 0.21586 0.90000 0.27941
MCZB 7742 M HIST 8.31250 0.11414 0.95000 0.14774
MCZB 7742 M HIST 8.75000 0.00259 1.00000 0.00335
WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY
APPENDIX 6
Measure of highpoint of contour. Value of XN when YN=1.0 is the highpoint
measure for all P concolor coryi (including P. c. arundivaga)
Catalog Subspecies Sex StatusI XN YN
US 1157
US 1158
LSU 11363
LSU 17032
MSH 51375
US 137122
UF 11852
UF 12462
UF 24097
MCZ 54423
UF 24557
UF 24563
AM 100222
ANS 16705
ANS 16706
FM 50058
MCZ 19855
MCZ B5489
MCZ B7743
UF 14699
US 145264
AM 144512
ANS 15270
FM 14902
MCZ 10703
MCZ 31781
MCZB 5650
MCZB 6992
MCZB 7742
US 265596
UF 12827
UF 10424
UF 18798
UF 19090
UF 20973
BILLIE
UF 11927
UF 14390
UF 19096
UF 22409
UF 22529
UF 23986
UF 24096
UF 24268
UF 24314
arundivaga
arundivaga
arundivaga?
arundivaga?
arundivaga?
arundivaga
coryi
coryi
coryi
coryi
coryi
coryi
coryl
coryl
coryi
coryl
coryl
corj'
coryi
coryl
coryl
corya
coryi
cory'
coryl
coryi
coryl
cory,
coryl
coryi
cory'
coryi
coryi
coryw
coryi
corli
coryi
coro
coryl
coryl
coryi
cory
cory,
HIST
HIST
TEST
TEST
TEST
TYPE
CAPTIVE
CAPTIVE
CAPTIVE-J*
GLADES-J*
GLADES
GLADES
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST
HIST/TYPE
HIST
HIST-J*
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
RECENT
0.600
0.650
0.650
0.500
0.600
0.400
0.550
0.600
0.550
0.650
0.450
0.550
0.400
0.550
0.350
0.500
0.500
0.350
0.350
0.500
0.350
0.350
0.600
0.350
0.350
0.350
0.450
0.350
0.400
0.350
0.400
0.300
0.450
0.450
0.600
0.500
0.400
0.400
0.350
0.350
0.550
0.600
0.400
0.550
0.600
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Appendix 6 continued
UF 24315 coryi M RECENT 0.600 1.000
UF 24316 coryi M RECENT 0.400 1.000
UF 24561 coryi M RECENT 0.350 1.000
US 528190 coryi M RECENT 0.350 1.000
UF 11915 coryi F RECENT-J* 0.500 1.000
UF 20957 coryl F RECENT-J* 0.600 1.000
GLADES 7040 coryi F TESTI 0.550 1.000
UF 19077 coryi F TEST2 0.550 1.000
UF 23985 coryi F TEST2 0.450 1.000
UF 24042 coryi M TEST3 0.350 1.000
Sjuvenles and esubedult aintted from study
SStus historic, recent, captive, svegldes. test (those nimals of questionable identity);
wpe*-type specimen for Pubpecied
WILKINS ET AL.: FLORIDA PANTHER MORPHOLOGY
Appendix 7
Skull measurements used in the cranial analysis of Puma concolor coryi.
VARIABLES.
1. Total Length (TL) Anterior tips of premaxillae to posterior point in median line over the foramen
magnum.
2. Condylobasal length (CBL) Anterior tips of premaxillae to posterior pane of occipital condyles.
3. Zygomatic breadth (ZYB) Greatest distance between outside borders of zygomata.
4. Cranium height (CRH) Vertical distance from lower border of palatines to height of frontals, at
vertical plane of postorbital processes.
5. Width of palate (PAW)- Greatest width of palate between outside margins of carnassial
alveoli.
6. Palatal length (PAL) Anterior tips of premaxillae to posterior edge of palate.
7. Maxillary tooth row (MXTR) Anterior alveolus of canine to posterior alveolus of carnassial.
8. Mandlbular tooth row (MDTR)- Anterior alveolus of canine to posterior alveolus of carnassial.
9. Mastoid breadth (MASB) Greatest posterior width of skull including outside margins of the
mastoids.
10. Post-orbital process breadth (POPB) Greatest width between the outer points ofpostorbital
processes.
11. Postorbital constriction (POC) Least distance posterior to the postorbital processes.
12. Width at canines (WC) Width at outside margins of canine aveoli.
13. Interorbital breadth (IOB) Least distance between orbits.
14. Pterygold width (PW) Least distance at outer edges ofpterygoids.
15. Condyle width (CW) Greatest width of occupital condyles.
16. Upper canassial crown length (UCL) Antero-posterior length of crown at cingulum.
17. Upper carnassial crown width (UCW) Greatest width of carnassial.
18. Lower carnassial crown length (LCL) Antero-posterior length of crown at cingulum.
267
BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3)
Appendix 8.
Number of Puma concolor specimens examined and number possessing
a mid-dorsal whor
Whorl
Subspecies N Locality N F(%) Source of data
NORTH AMERICA
arundlvaga 2
azteca 128
brownie 4
californica 45
coryi* 48
coryi.)**
cougar
hippolestes
improcera 2
kaibabensis 33
mayensis 4
missoulensis 16
olympus 18
oregonensis 58
stanleyana 117
vancouverensis 2
SOUTH AMERICA
araucanus 11
anthonyi
acrocodia
bangsi
borbensis
(incl. discolor)
concolor
costaricensis
green
incarum
osgoodi
patagonica
pearsoni
puma
soderstromi
undetermined***
TOTAL
1
4
7
Louisiana Arkansas
Arizona New Mexico,
Mexico
Arizona California
California Oregon
Florida
unknown origin (incl. captive)
Rhode Island
Colorado Montana Utah
Wyoming British Colombia
Baja, California, Mexico
Arizona Utah Nevada
Mexico Belize
Montana
Washington
Oregon Washington
Texas
Vancouver Island
Chile
Venezuela
Brazil (Matto Grosso)
Colombia Venezuela
15 Brazil Peru
21 Venezuela Brazil Guyana
Surinam
9 Costa Rica Panama Venezuela
1 Brazil
8 Peru
4 Bolivia Brazil
6 Argentina
7 Argentina
6 Chile Argentina
1 Venezuela
44 Argentina
648
USNM, LSU'
4 3.1 USNM, AMNH, FMNH
0 USNM, MVZ
9 USNM, MCZ
39 81.5 USNM, AMNH,
FMNH, ANSP, UF
1 2.5 USNM, FMNH, UF
1 100.0 MCZ
0 USNM, FMNH
0 MCZ, MVZ'
1 3.0 USNM, AMNH
0 USNM, FMNH, MCZ
0 USNM
1 5.5 USNM
0 USNM
1 1.0 USNM, UF, MR"
0 USNM, MVZ
3 27.2 USNM, FMNH,
AMNH, MR"
0 AMNH'
0 USNM, AMNH, FMNH
0 FMNH, AMNH, MCZ,
EBRG
0 USNM, AMNH
1 4.8 USNM, FMNH, MCZ,
EBRG'
0 USNM, AMNH, MCZ
0 USNM
0 USNM, FMNH, AMNH
0 USNM, FMNH
0 USNM, MR4
3 42.8 USNM, FMNH,
CMNH
4 66.6 USNM, AMNH,
FMNH, MR
0 EBRGO
10 22.0 BMNH, MCNBR'
69
268
WILKINS ET AL.: FLORIDA PANTHER MORPHOLOGY
Apendix 8 Continued
*ory -inchudes only museum spec ns; whod counted even ifdifficult to detect, (.e. dimenstary, N-3); seedcussion in tt
regarding presence in live animals
*coryl() includes purchase (USNM1943); known aptives (UF1852 UF12462 Everglades Wonder Grdes and suspected Cory
specimen (FMNH91621 with no data)
*undetennined subspecies, all specimens fro Argntina
1USNM U.S. National Museum; AMNH American Museum of Natul Histoy; FMNH Fied Museum of Natural Histy, MCZ Museum of
Comparative Zoology UP Florida Museum ofNatural History, BMNH British Museum ofNatund History
2LSU Louisiana State University (M. Hafmer, pers. comm.)
3MVZ Museum of Vertebrte Zoology (W. Liicker, Jr. pers. comm)
4MR Meody Rodkc (pe. comm.), based on examination ofliving animals
5AMNH American Museum of Natural History, type specimen (M. Lawrence, pes. comm.)
6EBRG Venezuela Estacion Biologica de Rancho Grande (F.Bisbal pers.comm.)
7CMNH Colorado Museum of Naturl History (C.Chase, pes. comm.)
SMCNBR Museo de Ciencias Naturaes Bernardino Revadavia" (A. Novaro, per. comm.)
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