Puma concolor

27. Puma Puma concolor French: Couguar / German: Puma / Spanish: Puma Other common names: Cougar, Mountain Lion, Catamount, Panther; Florida Panther (coryi) Taxonomy. Felis concolor Linnaeus, 1771, Cayenne region, French Guiana. In the Western Hemisphere the Puma ranges from Patagonia to northern British Columbia, a span of about 100 degrees of latitude. Recent analyses based upon mitochondrial gene sequences suggest the North American Puma derived from a small number of founders about 10,000 years ago and that sufficient genomic differentiation exists to support the recognition of two subspecies in the Western Hemisphere. Regardless, a taxonomic revision 1s needed for this species and its many recognized races. Earlier taxonomic work based upon phenotypic characters formed the basis for the recognition and classification ofat least thirty subspecies. Subspecies and Distribution. P. c. concolor Linnaeus, 1771 — E Venezuela through the Guianas to lower Amazonian Brazil. P. c. acrocodia Goldman, 1943 — C Brazil (Matto Grosso), SE Bolivia, and the Chaco of Paraguay and Argentina. P. c. anthony: Nelson & Goldman, 1931 — S Venezuela and adjacent N Brazil. P. c. araucanus Osgood, 1943 — S Chile and S Argentina. P.c. azteca Merriam, 1901 — SW USA (Arizona and New Mexico); NW Mexico. P. c. bangsi Merriam, 1901 — Andean Colombia. P. c. borbensis Nelson & Goldman, 1933 — Amazonian Brazil, Colombia, Ecuador, and Peru. P. c. browni Merriam, 1903 — NW Mexico (N Baja California). P. c. cabrerae Pocock, 1940 — NW Argentina. P. c. californica May, 1896 — SW USA (California). P. c. capricornensis Goldman, 1946 — SE Brazil and NE Argentina. P. c. cory: Bangs, 1899 — SE USA (Florida). P. c. costaricensis Merriam, 1901 — Nicaragua through Panama. P. c. cougar Kerr, 1792 — NE USA. P. c. greeni Nelson & Goldman, 1931 — E Brazil. P. c. hippolestes Merriam, 1897 — C USA. P. c. improcera Phillips, 1912 — NW Mexico (S Baja California). P. c. incarum Nelson & Goldman, 1929 — Andean Peru. P.c. kaibabensis Nelson & Goldman, 1931 — W USA (Nevada & Utah). P. c. mayensis Nelson & Goldman, 1929 — S Mexico through El Salvador. P. c. missoulensis Goldman, 1943 — NW Canada through NW USA (Idaho & Montana). P. c. oregonensis Rafinesque, 1832 — NW USA (Oregon and Washington). P. c. osgoodi Nelson & Goldman, 1929 — C Bolivia. P. c. patagonica Merriam, 1901 — S Argentina (E side of Lago Pueyrredon). P. c. pearsoni Thomas, 1901 — Patagonian Argentina and Chile. P. c. puma Molina, 1782 — C Chile and adjacent Argentina. P. c. schorgeri Jackson, 1955 — Midwestern USA P. c. soderstromi Lonnberg, 1913 — Andean Ecuador. P. c. stanleyana Goldman, 1936 — S USA (Texas) and adjacent Mexico. P. c. vancouverensis Nelson & Goldman, 1932 — SW Canada (Vancouver I). Descriptive notes. Head-body 86-155 cm, tail 60-97 cm. Average weight ranges from 53-72 kg for adult males, and 34-48 kg for adult females. An exceptionally large male weighed 120 kg. Mass at birth is about 0-6 kg. Growth rates are similar for both males and females. Puma are large, slender cats, tawny above and whitish below. Melanism has been reported rarely in South America, but not in North America. Kittens are buffcolored, with rows of irregularly-shaped black spots until 9-12 months old. Thetail of the adult is a long and sweeping “|” that is tipped in dark brown to black hair. Puma have proportionally the longest rear legs in the Felidae. Hair length, color, and texture vary geographically throughout the species’ range in the Western Hemisphere. Individuals from colder, higher altitudes tend to have thicker and longer hair than those from more tropical climates. Dentition follows the typical felid pattern of prominent canines, modest incisors, and sharp carnassial cheek teeth for shearing tendons and bones. Puma exhibit a range of vocalizations, which are infrequently heard but may be associated with mating behavior. These include bird-like chirps that appear to be used by females to communicate with kittens, and the more stereotypical scream, which is likely related to mating behavior. Unlike the great cats, the Puma is unable to roar, a product of reduced larynx and hyoid apparatus. The prevalence of a crooked tail, atrial septal defects, a dorsal whorl of hair, and other anomalies in the Florida subspecies (coryi)is likely a product of reduced population size and several generations of inbreeding. Interestingly, all of these anomalies were absent from young born of matings between Texas female Pumas and Florida Panther males. Seven Texas female Pumas were inserted into the Florida Panther population as part of a genetic augmentation effort in 1995; five of these females were known to have reared at least two litters each. Habitat. The Pumais usually associated with remote, rugged terrain where there is cover for stalking and ambush-hunting, secure places to establish natal dens, and at least one species of abundant deersized prey. It has been suggested that viable populations of Puma are impossible without at least one species of deer-sized prey. Its extensive distribution throughout the Western Hemisphere, from sea level to more than 4000 m elevation, suggests a tolerance of environmental conditions that is rare among mammals. Habitat use can be highly seasonal where prey species such as Elk migrate altitudinally in response to snowfall, or can be annually static, for example, in subtropical southern Florida, where prey species have stable annual home ranges. Specific habitat preferences are as variable as the regions in which the Puma lives, and range from mixed conifer and curlleaf mountain mahogany vegetation in rugged topography in Wyoming to subtropical hardwood hammocks, pine flatwoods, and palm forests in southern Florida. Female Puma locate secretive natal dens in boulder piles, dense vegetation, or other natural structures that provide some protection from the elements and reduce detection by potential predators. In Florida these sites tended to be at least one km from a paved road, and were usually located within dense thickets of saw palmetto (Serenoa repens), which provided vertical cover, horizontal cover, and reduced temperatures compared to outside air. Where the species is dependent upon forest, occupied areas tend to be at least 20,000 ha without major roads. The species increasingly is found in landscape patches that have been fragmented by expanding human activity and infrastructure such as highways, ranches, produce farms, human settlements, and extractive industries. In these areas, remnant landscape connections and restored habitat corridors can be important demographic linkages. In Florida and California dispersing individuals are tolerant of habitat that may include canals, highways, relatively open terrain, and other features that are usually recognized as barriers by resident adults. Food and Feeding. The Puma’s extensive distribution is reflected in a diverse list of prey. In general, Puma from temperate climates eat larger prey than Puma from tropical climates, and solitary adults tend to eat larger prey than females with kittens. The smaller prey size of Puma in the tropics may be the result of niche separation with the larger, socially dominantJaguar. Puma are ambush hunters; they utilize cover to closely approach potential prey before an attack is made. Common prey in North America include Elk, White-tailed Deer, Mule Deer, Wild Boar, Collared Peccary, porcupines, rabbits, and hares. In South America common prey include vizcacha, Guanaco (Lama guanicoe), brocket deer, Pampas Deer (Ozotoceros bezoarticus), pudu (Pudu), agouti, armadillo, and porcupine. Domestic stock is occasionally taken throughout the species’ range, and exotic species such as Wild Boar and European Hare can be important dietary components even where larger prey are available. In marginal habitat such as in the Everglades of southern Florida, atypical prey such as alligator (Alligator mississipiensis) and North American River Otter may be taken. This is the result of inherently low populations of White-tailed Deer in the Everglades region. Puma routinely remove the entrails of large prey before caching and burying their kills with leaves and other debris for future use. The use of such kills may last from one to 27 days. Humans can alter the abundance and spatial use patterns of Puma and their prey through hunting and otheractivities. Historically, Puma predation on humans has been rare, although in recent years there has been an apparent increase in encounters, especially involving juvenile or underweight cats. Many of these attacks have occurred in the urbanwilderness interface where the likelihood of an encounter increases with presence of humans in occupied peripheral Puma habitat. Activity patterns. Puma are primarily crepuscular and nocturnal and do the bulk of traveling and hunting at night. This pattern appears related to the activity of their prey and the concealment offered by darkness. In Florida, activity peaks occurred from 01:00-07:00 h and 18:00-22:00 h. Females with kittens tended to leave their natal dens to hunt at 22:00 h and return to them at 08:00 h. Activity patterns of females at natal dens were similar to those of solitary adults, but exhibited less difference between activity peaks and nadirs. Movements, Home range and Social organization. Puma exist in a system of dynamic land tenure: resident adults have prior rights to home ranges, and residency is dependent upon the death or departure of other residents. Annual home range size varies from 50 to more than 1000 km? and appears to be related to primary productivity and prey abundance. Movement distances greater than 20 km in a 24hour period have been recorded. Male home ranges are generally more than twice as large as female home ranges, and tend to incorporate as many females as possible within a territory boundary that is not continually defended. Males tend to use their larger home ranges evenly across seasons, whereas female movements are related to reproductive condition and the presence of kittens. Females restrict their movements to the immediate vicinity of natal dens for about two months, after which time kittens join their mothers at kills. Aggressive encounters between males appear to be contests for breeding rights and often result in the death or injury of at least one of the combatants. The basic social unit ofthis solitary speciesis the female-kitten group, which lasts for 10-24 months. Independence of offspring may be related to onset of estrus in the female and subsequent sexual encounters with adult males. In Florida, independence ofjuveniles was preceded by encounters between the mother and an adult male in 63% of 27 documented instances. Females are philopatric and tend to establish home ranges adjacentto those of their mothers. In Florida, dispersal distances ranged from six to 32 km in females and from 24 to 224 km in males. In Wyoming, a male dispersed 274 km. Females are capable of long-distance dispersal, but seldom do so. Translocated individuals have been known to return to capture locations from more than 470 km away. Breeding. Puma reproduction can occur year-round,varies regionally, and, if seasonal, appears to be linked to the reproductive patterns of primary prey. For example, in Florida, reproduction has occurred in every month, but most litters are born in the spring, after the pulse of White-tailed Deer fawn production. Gestation varies from 92-96 days,following a bout of copulations that maylast several days. Adult males may be aspermatic for portions of the year. In Florida, despite poor sperm quality, reduced genetic variability seems insufficient to limit population growth. Litter size averages two and ranges from one to four. Earliest documentation offirst reproduction in the species is 18 months for females. Males generally begin breeding after three years, although sexual maturity is likely attained at or before two years. Status and Conservation. The Puma is listed on Appendix II of CITES with the subspecies coryi, costaricensis, and cougar listed on Appendix I. Classified as Least Concern on The IUCN Red List. In North America Puma inhabit less than 50% of their original range, primarily due to the loss of habitat and intensive human activities. A similar pattern likely pertains in Central and South America where deforestation is occurring at a rapid pace. In North America legal status varies: the Puma is an unregulated predator in Texas; a hunted species with distinct seasons in Nevada, Montana, and Alberta, Canada; a non-hunted species in California and Manitoba, Canada. The Puma is fully protected in Florida. Races cougar and schorgeri are probably extinct whereas coryz, costaricensis, and brownii are listed as threatened or endangered by the US Fish and Wildlife Service. In South America Puma hunting is prohibited in Argentina, Brazil, Bolivia, Chile, Colombia, Costa Rica, French Guiana, Guatemala, Honduras, Nicaragua, Panama, Paraguay, Surinam, Venezuela, and Uruguay; it is unprotected in Ecuador, El Salvador, and Guyana. The Puma can be hunted in Mexico and Peru. Small population size has resulted in reduced genetic variability and maladaptive physiological traits such as atrial septal defects in coryi. However,there is no evidence that such phenotypic characters have negatively affected demographics in the population. Efforts to save the Florida Puma include experimental translocations and genetic introgression via the transport of female Puma from Texas. The Puma, like other large carnivores, is an effective conservation flagship because it requires large areas, exhibits top-down regulation, and can be popular with the public. A landscape approach to the conservation of this species will be important in maintaining both large and small populations because the species occurs at inherently low densities. Successful management and recovery strategies for Puma will require local and regional planning to balance expanding human populations and infrastructure development with habitat restoration. Land management policies that enhance prey populations and offer the Puma interconnected reserve systems are needed. Efforts to create blueprints for such large-scale conservation include Paseo Pantera (Wildlife Conservation Society) in Central and South America and the Florida Ecological Network. Bibliography. Ackerman et al. (1986), Anderson (1983), Ashman et al. (1983), Azevedo (2008), Banfield (1974), Barone et al. (1994), Beier (1993, 1995), Belden & Hagedorn (1993), Creel (2006), Culver, Hedrick et al. (2008), Culver, Johnson et al. (2000), Cunningham et al. (1999), Dalrymple & Bass (1996), Etling (2001), Fuller, K.S. et al. (1987), Gay & Best (1995), Gonyea (1976), Hoctoret al. (2000), Holt (1994), Iriarte et al. (1990), Kelly et al. (2008), Kilgo et al. (1998), Kurtén (1973), Laing & Lindzey (1993), Logan & Irwin (1985), Logan & Sweanor (2001), Logan et al. (1986), Maehr (1990, 1997), Maehr & Caddick (1995), Maehr & Cox (1995), Maehr & Moore (1992), Maehr, Belden et al. (1990), Maehr, Land & Roof (1991), Maehr, Land, Roof & McCown (1989), Maehr, Land, Shindle et al. (2002), Maehr, Roof et al. (1989), Mclvor et al. (1995), Murphy et al. (1999), Pierce et al. (2000), Pimm et al. (2006), Rabb (1959), Romo (1995), Rosas-Rosas et al. (2003), Ruth et al. (1998), Scognamillo et al. (2003), Seidensticker et al. (1973), Sunquist & Sunquist (2002), Thompson & Stewart (1994), Toweill et al. (1988), Van Dyke et al. (1986), Wilson, D.E. & Reeder (2005), Wilson, P. (1984), Yanez et al. (1986), Young & Goldman (1946).Published as part of Don E. Wilson & Russell A. Mittermeier, 2009, Felidae, pp. 54-168 in Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona :Lynx Edicions on pages 156-158, DOI: 10.5281/zenodo.637689