-1 1 ==8-- - ---- 4- _ ./- 1 - 1 - 1* 1, -5---- ME<~I11 i ]1 ~ -4*,Il, 2- ~ -4 1 -'- 1-1 ' :4-11 -=a '' r j ..4 41 - 161 9.1 _ h, 4 It, ~r* -.4-,41 -, |- n 1 -1'-1 1 '741 , 0. *1 ,£tr-=-- f.-f=cr I j~!r- - A--7- 1 C._, ___J -ER-=a 4_-·31;E·-Ze_ ~-i!;3& I 1 5 .t - - -- of fhe 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 34 3, pp. 211-269 1997 El . -3-*. '- 1 - ./ ··· 11'1 " ... itn /- .1 -1 *21· L X -=-1 ' . 4-- , '3-- f wk . -4 1 - 1--fpz=- 1- .* _--*14; _ 4- - 4-1 - -1 4 1 4 5.-4=1=1; - UNIVERSiTY OF FLORIDA GAINESVILLE Numbers of the BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISrORY m published at irregular intervals. Volumes contain about 300 pages and are not necessarily completed in any one calendar year. JOHN F. EISENBERG, EDITOR RICHARD FRANZ, CO-EDITOR RHODA J. BRYANT, A£ANAGING EDITOR Communications concerning purchase or exchange of the publications and all manuscripts should be addressed to: Managing Editor, Bulletin; Florida Museum of Natural History, University of Florida P, 0. Box 117800, Gainesville FL 32611-7800; U.S.A This journal is printed on recycled paper. ISSN: 0071-6154 CODEN: BF 5BA5 Publication date: December 31, 1997 Price: $ 4.50 THE FLORmA PANTHER kma concolor cogi: A MORPHOLOGICAL INVESTIGATION OF THE SUBSPECIES WITH A COMPARISON TO OTHER NORTH AND SOUTH AMERICAN COUGARS Laurie Wilkinsi, Julio M. Arias-Reveron2, Bradley M. Stith~, Melody E. Roelkei, and Robert C. Beldens ABSTRACT The endangered Florida panther, Puma concolor cogi, 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 cogi 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 hits. These specimens were compared to a sample ofNorth 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 inforination, 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 cory, morphologic features. 1 Florida Museum of-Natural History, University of Florida (UF), Gainesville FL 32611-MOO, USA 2 Instituto Tecnologico de Costa Rica, Centro de Investigacion y Desarrodo en Agricult rra Sostenible para el Tropico Humedo (CIDAS% Apartado 223-4400, Ciudad Quesada, San Cdos, Costa Rica (fonnerly: Univemity of Florida, IFAS Ci~us Research and Education Center. 700 Experiment Station Road, Lake Alfred FL 33830, U.SA 3 Department of Wildtife Ecology and Conservation, University of Florida, Gainesville FL 32611, U.SA 4 Genetics Laboratory, Na8onal Cancer Institute, Fredmick MD 21702, U.SA (formedy with the Flotida Game and Fresh Water Comrnission). 6 Florida Game and Freshwater Fish Commissionl 4005 South Main Street Gainesville FL 32601, U.SA Wilkins, L,J. M. Arias-Reveron, B. M. Stith. M. E. Roelke, E C. Belden. 1997. The Florida panther Puma concolor cogi: A morphological investigation ofthe subspecies with a comparison to other North and South American cougars. Bull. Florida Mus. Nat. Hist 40(3):221-269. 222 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3) RESUMEN La pantera de Florida, Puma concolor cool, amenazada dc extinsi64 ha sido objeto de estudios ecoldgicos y biomudicos, pew poco se ha hecho acerca de la morfologia dc mis subespecies. El interes en los caracteres morfolagicos que describen la poblaci6n de Florida se ha incrementado con el des¢ubrimiento de panteras fuera de su rango conocido en el sudoeste de Florida, la adquicisidn de mas especimenes (que antcs no estaban disponibles a previos investigadores) y los primeros especimenes y capturas vivas provenientes de los Everglades. La necesidad de definir cancteristicas subespecificas de las panteras de Florida tiene tambien implicancias legales Los caractercs previamente utilizados para describir Puma concolor cogi fueron cuantificados y re-evaluados usando m6todos estadisticos. En todos los especimenes historicas y recientes del sudeste de los E.E.U.U. (n=79) se examinaron el color de pelo, perfil cranial, proporciones y otros rasgos morfoldgica Estos especimenes, fueron comparados con una muestra representativa de Norte y Sudamerica. Los caracteres medidos proporcionaron la base para la descripci6n de la poblaci6n de Florida y para su descriminaci6n dentro y entre otras subespecies. 1~ especimenes de pantera que viven en la regidn de los Everglades difieren del resto de la poblaci6n de Florida en su morfologia cranial y otras caracteristical hallazgo consistente con recientes investigaciones genaticas. No existen evidencias que soporten una separacian geogrAfica o ecoldgica de la pantera de Florida en dos poblaciones. La mejor explicaci6n de las diferencias morfo16gicas observadas. consistente con la informaci6n histbrica, es que las panteras de los Everglades son descendientes de animales de cautiverio liberados y que por 10 tanto scrian hibridos. Con este maodo 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 ditimos veinte aitos, exhiben los clasicos rasgos morfol6gicos de P. concolor coryi TABLE OF CONTENTS Introdur,inn 223 Acknowledgments 224 Abbreviatinne 225 TaxonomicSynonymy ofPuma concolor coryi 225 Historical Distribution. 226 Methods 227 Pelage r.ir- 230 Pelage Characters and Kit)ked Tail 235 Cranial Profile 240 Cranial Proporti,-9 247 Discussion and Conclusion. 230 Literature Cited 254 Appenli;re« 257 WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY 223 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 OP. 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 19864 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). 224 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, amnged by James Brady, for the transport of the hundreds of pounds of equipment necessary to co.,dirt the color analysis We are grateful to the many museum curators and collection staff who allowed one of us (lw) to examine specimens under their care, especially. C. Sma,t and T. Daeschler (Academy of Natural Sciences Philadelphia), G. Musser (American Museum of Natural Hi~vy), 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 FLA B. Patterson and R. Izor (Field Museum of Natural History), M. Haher (Museum of Zoology, Louisiana State University, Baton Rouge), M. Rutzmoscr, 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 loana~ 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 Andrds Novam 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 of the color grectrophotometer. 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 Wak and Wendy Zomlefer. produced the graphics. Oron L Bass. Jr. (National Park Service) provided WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY 225 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. Revima by Jacqueline Belwood, Michael Kennedy, and Melvin Sunquist greatly improved the manuscript Consinictive comments were also provided by Oron Bass Deborah Jamen, Thomas Ing,4 and William Robertson. Rhoda J. Bryant and Dianna Camer (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 cogi), sce 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-Evergl*les cats), since 1950 GLADES Everglacles 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 ~ANGS) Felis concolor «#oridana Cory, 1896. Hunting and Fishing in Florida, p, 109. (name preoccupied by Felis flori(lam: Desmarest 1%20 (=Lynx jloridanus Rafinesque, 1817). Type from Alapata Flats, north of Lake Okeechobee, Florida, FMNH 9255, skin only, M adult, Field Museum of Natural HiStOfy, 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 cogi Bangs, 1899. Proc. Biol. Soc. Wash. 13:15, Jan. 31 (renaming of F. 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. 1 Hunter January 1, 1898. Felis arundivaga Hollistcr, 1911. Proc. Biol. Soc. Washington. 24:176, June 16. Type from 12 miles southwest of Vidalig Concordia Parish, Louisiana USNM 137122, skin and skull, M adult, United States National Museum 226 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 FWis cogi 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 cogi, 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. cogi 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. stanteyana 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 (18005), and one from Caddo Parish (1965); Aikansas (2)-Ashley County (1969) and Logan County (1975); South Carolina (1)-with conflicting data suggesting it was either from Oregon or South Carolina (records ofANSP) (Appendix 3). Archeological and paleontological evidence exists for pumas in the southeastern United States outside of Florida (Parmalee 1960, Kurt6n 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, Kurt6n 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 19864 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 227 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 inh,hited 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, pen. comm.), The need to develop a means to identify the Florida panther, particularly in law enforcement issues, was the genesis of this stu*. METHODS Museum specimens of P. concolor cool were examined and compared to specimens of puma throughout its range in North and South America. The sample from the soatheastern 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 priori into discrete groups, and the characters are unweighted. The principal components (axes) are representations of the variables (measures), each of which vafy 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. 228 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3) SOUTHEASTERN SPECIMEN LOCALITIES N Carli 0 Lake ern Hemando Orange Pasco ~~~ Polk ~ ,$'00 0 Mandee Hardee 9 Highlands - De Soto Martin Charlo#e C~lades 0 0 Lee _-_-_· __Hendry ·-- --- -6- Palm Beach~ift 6-5. *2 _-Z *i -««-' NTotal specimens: 72 \€i.«0 o recent 0 historic (pre-1950) w questionable origin , 50: . - 0 0 4 Figure 1. Geographic distribution of Puma concolor coo, specimens examined (dots) from the southeastern United States. Bree groups of specimens are represented: recent specimens since 1950; historical specimens including those collected by Charles Cory and Outram Bangs in the late 18005; 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 229 CDA is a powerful procedure that maxinlizes 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 discrimin,nt onglysis. MANOVA tests the hypothesis that all o priori groups have the same multi-dimensional mean (centroid) 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 (=GLADES); 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 Everglacles 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. coo,i 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, Kurt6n 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 ofjuveniles 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 230 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 mid4ine 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 1 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 1 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 231 FLORIDA PANTHER COLOR MEASURES 214 5 5 --0-- g =7 .25 E ~36 g Figure 2. Spectrophotometer readings taken for seven regions of the pelt: three dorsal, three lateral, and a single belly measure. Dorsal values represent the cia,ker midline; lateral measures were taken randomly within the defined regions and represent the 1 ' ' ant 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. 232 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 cogi 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 cogi is darker than western and northern inland populations from North America (Fig. 3a). There is virtually no difference in color measures between P. c. cogi and coastal populations from Oregon and Washington (P. c. oregonensis and P. c. 0/ympus) (Fig. 3b). This also was noted by Goldman (1946, p. 237): "In dark general color tones cogi approaches the geographically distinct olympus...." P. c. coryi is less red than tropical subspecies from Guatemala Costa Rica, Panama Venezuela, and BrazA CP. 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 hippo/estes (p=0.002) and P. c. stanteyana (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 vadables (p>0.1). Among the South American subspecies, the differences in the means of color variables for P. c. coo,i 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 Cornett 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. 34 b). As with the PCA, no significant differences were detected between the means of P. c cogi and the Everglades cats or the Corbett female with MANOVA, but P. c. cogi 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 of P.c. cogi (Fig. 34 b; Table 1). However, the colors of these test animals differed in ways that were not measured with the spectrophotometer. Table 1. Probability (E 0.05) of difference between the means of color variables for selected North American Puma concolor subspecies, including cool and Everglades cats. Variables include measures of hue, saturation, lightness degree ofredness, and degree of yellowness for mid-dorsal and mid-lateral region ofskin (probability ofno overall species effect 0.0001 - MANOVA). Not all subspecies were tested. W ILKIN S ET A L: FLO R ID A P R M O R PH O LO G Y 233 coryi ever captive testl azteca calif hippo kaibab mayen missoul olymp oregon stanley (n=24) (n=2) (n=2) (n=l) Or•36) (n=23) (n=ll) (n=22) (n=4) (n=16) (n=12) (n=18) (n=23) coryt . 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* hippolestes . 360 .0077* . 127 .0005* .0003* - kamabensis .003* .021 * .0001* .0001 * -- mayensis .238 .0016* .034* -- missoulensis .0007* .029* - olympus .271 -- azteca .275 californica .0001 * 1 VF D985 female shot by himter, Corbett Management Area * signincant *<0.05. Table 2. Probability (5 0.05) of difference between the means of color variables for selected South American Puma concolor subspecies including coryi and Everglades cats. Variables include measures of hue saturation„ lightness degree of redness, and degree of yellowness for mid-dorsal and mid-lateral region ofskin (probability ofoverall species e#ect was 0.0001 - MANOVA). coryi acrocod araucan bangsi borben concolor costrac incarum osgoodi pearsoni (n=24) (n=4) (n=9) (n=4) (n=15) (n=16) (n=8) (n=6) (n=4) (n=4) coryi 1 .000 .0001* .6205 .2933 . 0001 * .0001* .0001 * .0027* .0107* .0001 * acrocodia .0033* .0856 .0001 * .0036* .0380* .0203 .0762 .0001 * araucanus .9680 0001 * .0003* .0041 * .0239 .0470* .0003* bangsi 0003* .0083* .0466* .0977 . 1924 .0218* borbensis .0745 .3084 .0046* .0650 .0001 * concolor .7781 .0615 .2442 .0001 * * signiticant *<0.05. 234 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 40(3) FLORIDA PANTHER COLOR MEASURES COMPARED WITH OTHER PUMA POPULATIONS 4 - a) North America 2. 4 434 335 S E C O N D P R IN C IP A L C O M P O N E N T S E C O N D P R IN C IP A L C O M P O N E N T O- 2 J 1 01 11 1 1 1 -2. 4 - da rk er 1 -4. redder -+ 4 -2 0 2 4 FIRST PRINCIPAL COMPONENT b) North West Coast 4- /2//1 2\22 /42/2. 2/ 42 // 21 3 \1/ 21 2 12\ / ~ redder ~ \11 / 1 4 -2 6 2 4 FIRST PRINCIPAL COMPONENT Legend on following page. WILKINS ET At-: FLORIDA PANTHER MORPHOLOGY 235 FLORIDA PANTHER COLOR MEASURES COMPARED WITH OTHER PUMA POPULATIONS C) CentraUSouth America 2 3 2- \ :r \1 0- # 1 j/ h34 , SE C O N D P R IN C IP A L C O M P O N E N T 4 -2 - V ' // - d ar ke r -4 4- redder - 6;4 FIRST PRINCIPAL COMPONENT Figure 3. Plots ofthe 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, cogi (Florida); 2, missoutensis (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, coo,i; 2, combined oregonensis and otympus (Washington, Oregon, Olympic Peninsula); P, Piper cats from Everglades Wonder Gardens; T, test animal (Corbett female). There is virtually no difference between Puma concolor cocyi and animals inhabiting the northwest coast (c) 1, coryi (Florida); 2, costancensis (Costa Rica Panama); 3, osgood, (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 similat 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 236 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 (Golanun 1946). The density of flecks on any particular animal is correlated with age (Roelke and Wilking, unpubl. Atn). They consist of only a few isolated white hairs or small patches on very young animals, but old animals have liberal necking 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 whort, 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 237 FLORIDA PANTHER PELAGE/SKELETAL TRAITS FLECKS, WHORLS AND KINKED TAILS 2 im.V-"- *li: 2 .''i 38.1 ,I Figure 4. Pelage hits ofthe 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 ofthe living animal. Photographs by R Belden (top) and M. Roelke (bottom) 238 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL- 40(3) PRESENCE OF WHORLS IN FLORIDA AND OTHER POPULATIONS OF COUGARS f 58 0 km 50 Ll-LLU 2 16 QBangs 18 & Cory 9 5.5% 21 5~% (5)(1) 33 005(1 128 117 31 1% " 79% (57) 4) (1) Lake V Okeecho e 4 173 ~~ 81~ 1 Shark Total N. America Cypress Sloug 8 of 456 = 1.75% 7 -21 4.8% (1) swal* Miami 1 1 92.6%(50) 7 BE¥ ades Total S. America 15 25%(2 121 of 145 = 14.4% 8 44 / SOUTH FLORIDA 66.6% (4) 6 Southern cone: 272% (3) 11 20 of 74 = 27.0% 7 42.8% (3) ? 22% (10)1 6 1 Localities vague, subspecies undetermined. 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 1he 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 239 Results Whorl.- The mid-dorsal whorI 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 18005, 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 Evergla(les 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. cocyi 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. concdor coy# 72 57 79.2 Total North America(excluding cogn) 456 8 1.75 Total South America 145 21 14.40 Combined ssp. from Chile2 and Argentina 74 20 27.00 rmy; includes all Florida specimens and living animals examinei captive specimens included. Combined total of Chilean and Agentinean specimens are also presented to illustrate the high frequency of the whorl in this subspecies complex (consisting of four subspecies: waucam,3, pearloni, pum., polagonica) 240 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3) FREQUENCY OF WHORLS AND KINKED TAILS 100 N = 61 N = 49 80 - % OC CU RE NC E ~ whorl E-7 kinked tail 60 - 40 - N = 145 9 milir 9 ill Everglades, Florida, N. America S. America Florida (excluding Everglades) PUMAS Figure 6. High frequency of whorts/kinked tails in Big C~ress compared to Everglades and puma populations in North and South America Everglades includes two animals from West Palm Beach, only one ofwhich 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. WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY 241 Table 4. Frequency ofoccurrence ofwhorl and kinkedtail in specimens and live captures of Florida cougars. Whorl % Kink % ~cality n Present Occurrence n Present Occurrence Southwest Florida' 54 50 92.6 49 433 87.8 Historical Southeast2 9 5 55.3 post-cranial skeletons not available Recent Southeast1 9 2 22.2 10 1 10.0 Total 72 59 'All specimens from Big Cypress and other regions west of Shark Slough. ,Specimens taken by Bongo and Cory in the late 18005 'Includes Eveglades and two individuals from Palm Beach County. one ofwhich had a skin. The cranial profile was duplicated with a carpenter's contour gauge. The gauge was placed 1/8" to the left ofthe 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=1) (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 242 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL- 40(3) MEASUREMENT OF CRANIAL PROFILE a) Region of Cranium Measured flattenedinflated frontalnasal regionregion "' 111 e 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 nattened #ontal 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 cory, and others. WILKINS ET AL: FLORIDA PANTHER MORPHOLOGY 243 SKULL PROFILES 1.0 - P. C. COTyi P. c. hippolestes A< 0.8- bc• YONG K 0.6- . H E IG H T t. \ n-0,0. a4 - 1 . 0i. d' 1 ,82 - J J V ¥ . . . 1 0 0.2 0.4 a6 0.8 1.0 NORMALIZED PROFILE Figure 8. Images of profiles interpolated to increments ofO.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 ofthe degree of inflation ofthe anterior portion ofthe cranium. The closer the high point is to X=0, the greater the inflation of the nasals. Shown is the comparison ofa normalized profile of a Florida panther skull compared to that ofa 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. cogi 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. coo'i 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 califomica from California (n=30, p=O.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. cogi and North American subspecies P. c. cougar of eastern U.S.(n=4), P. C. olyinpus (n=l) and P. c, oregonensis (n=24), the latter two from northwestern U.S.. Significant differences were recorded between HIST P. c. cogi and three GLADES cats, two PIPER from 244 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 40(3) Table 55 Mean, standard deviation, and '' ' i values for contour measurements of subspecies (n = 2861 excluding coiyl. Figure in last column is probability of differences between means of cogi (historic only n = 18) and other subspecies (p E 0.05 Mann-Whitney-Wilcoxan non-parametric 2-tailed test) Standard Subspecies n Mean deviation min - max Z-score Probability araucanus 4 .525 .0289 .500 -.550 2.297 0.0216 * arundn,agal 6 .567 .0983 .400 -.650 - - azteca 55 .493 .1029 .350 -.700 2.9799 0.0288 * bangli 3 .533 .0289 .500 -.550 2. 139 0.0325 * borbensis 8 .473 .0886 .350 -.600 -1 .8576 0.0632 browni 1 .600 .0000 .600 -.600 -1.5842 0.1131 cabrarae 2 .525 .0354 .500 -.550 1 .6914 0.0908 californica 30 .541 .0890 .350 -.700 -4. 1807 0.0000 **** capricornesis 1 .450 .0000 .450 -.450 0.5940 0. 5524 concolor 4 .462 .0479 .400 -.500 1 .4413 0. 1494 costaricensis 5 .600 .0866 .450 -.650 -3 .0032 0.0027 cougar 4 .438 .0730 .350 -.500 0.77924 0.4359 greeni 2 . 525 .0354 .500 - .550 1 .6914 0.0908 hippolestes 24 .546 .0920 .350 -.700 4.0068 0.0000 **** incarum 6 .533 .0753 .400 -.600 2.7126 0.0067 - katbobensis 22 .539 .0950 .300 -.650 -3.6935 0.0002 - mayensis 6 .500 .0837 .400 -.600 2.2621 0.0237 * missoutensis 14 .561 .0789 .350 -.650 -3 .7672 0.0002 *** olympus 1 . 550 .0000 .550 -.550 1 .3862 0. 1657 oregonensis 24 .460 .1021 .350 -.600 -1.6904 0.0910 osgoodi 11 .514 .1206 .300 -.650 2.2949 0.0217 * patagonica 2 .650 .0707 .600 -.700 -2.2944 0.0218 * pearsom 11 .568 .1055 .400 -.750 3.5264 0.0004 *** puma 2 .525 .0354 .530 - . 550 - 1 .6914 0.0908 schoregori 1 .350 .0000 .330 -.350 -0.71326 0.4757 soderstrdmi 4 .450 .0913 .350 - .550 -0.95976 0.3372 stanleyana 28 .536 .0941 .350 - .650 3.8647 0.0001 *** vancouverensis 7 .650 .0300 .600 - .750 -3 . 8613 0.0001 *** Total 286 ' fp*~ 0 for anmdimga O.ouijana) scores. ...~%:ojl .... 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 H[ST 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. cogi, followed closely by recent P. c. co