key: cord-0040111-sxqbpk3i
authors: Stein, Andrew B.; Hayssen, Virginia
title: Panthera pardus (Carnivora: Felidae)
date: 2013-06-12
journal: nan
DOI: 10.1644/900.1
sha: 2439338da09def95d08768cecfdc0d2d98bc2197
doc_id: 40111
cord_uid: sxqbpk3i

Panthera pardus (leopard; Linnaeus, 1758) is the smallest of the 4 large felids in the genus Panthera. A solitary and adaptable species, P. pardus is the widest ranging of all wild felids, inhabiting rain forests, mountains, semiarid environments, and suburban areas throughout sub-Saharan Africa, the Middle East, and South Asia to the Russian Far East. Despite this distribution, P. pardus is listed as “Near Threatened” by the International Union for Conservation of Nature and Natural Resources and several Asian subspecies are listed as endangered. P. pardus primarily feeds on small to medium-sized ungulates, but has a varied diet including fish, reptiles, birds, and small mammals.

NOMENCLATURAL NOTES. A thorough discussion of the type locality is in Pocock (1930a) . The species name pardus is from the Greek pardos for male panther. Fitzinger (1868) provides pre-Linnean synonyms. The name pernigrajperniger is often attributed to Hodgson, but was published by Gray (1863) when cataloging Hodgson's specimens. Thus, following Ellerman and Morrison-Scott (1951) , we have attributed the name to Gray. An alternative attribution would be Hodgson in Gray (1863) . Other common names are graupanther (Brehm 1863) ; leopard, panther, panthere (Pocock 1930a (Pocock , 1930b ; chui, nsowa, steppenleopard, tui (Matschie 1895) ; ihwanqana, ingwe, nkoe, nkwe, nkwe, nngwe, yingwe (Roberts 1951); chita, harvard, ingwi, inkwi, ngo, nyalugwi, shabel, siveri, tijger (Ward 1910) ; chui, damissa, dumok, erith, lagho bagh, malilda, nimr, ol-owarukeri, osebo, terdwa (Dollman and Burlace 1928) ; and bars (Heptner and Sludskii 1992) . " [T] he terms 'panther' and 'leopard' are used somewhat indiscriminately and inconsistently as synonyms. Leopard is the name by which the animal is commonly known amongst English-speaking people in Europe, America and Africa; but ... panther ... is more usually employed by Indian sportsmen" (Pocock 1930a:64) . The generic synonymy for Panthera includes Felis and Leopardus as well as Jaguarius, Leo, Pardus, and Tigris (Mazak 1981 ).

Panthera pardus (Fig. 1) is a large spotted cat distinguished from other members of the genus Panthera by its distribution throughout sub-Saharan Africa and Asia and its characteristic dark, rosette spots. In Africa, P. pardus pardus may be confused with cheetahs (Acinonyx jubatus), which also have been called "hunting leopards" in Asia (Sterndale 1884) , but cheetahs are taller and more slender with circular, solid spots, unlike the robust P. pardus with rosettes. Cheetahs also have a diagnostic tearmark from the inside of the eye to the outside of the mouth, absent in P. pardus (Krausman and Morales 2005) .

The coat pattern of P. pardus and the jaguar (Panthera onca) is similar, but P. onca often has a small spot within the rosette pattern, whereas P. pardus often does not. P. pardus and P. onca are not sympatric because P. onca inhabits Central and South America and P. pardus occurs in Africa, the Middle East, and Asia. P. pardus is marginally smaller than P. onca (length of head and body, 1.0-1.5 m, 1.1-1.8 m; mass, 30-90 kg, 61-132 kg, respectively), but P. pardus has a somewhat longer tail (0.8-1.0 m, 0.5-0.8 m, respectively- Nelson and Goldman 1933; Roberts 1951; Hall and Kelson 1959; Rosevear 1974; Eisenberg and Redford 1999; Stein 2008) . P. pardus has a more slender head and smaller foot pads compared to the more robust skull and more evenly rounded foot pads of P. onca (Nelson and Goldman 1933) .

Individual measurements can vary by geographic region with smaller individuals typically coming from Cape Province, South Africa, where adult male Panthera pardus pardus have a mean mass of 31 kg (range, 20-45 kg), whereas the more typical masses for an adult male P. pardus are reported from Zimbabwe as 60 kg (range, 52-71 kg) with individuals up to 90 kg on occasion (Kitchener 1991) . Ranges of mean body mass (kg) for 34 females and 47 males from India, the Ivory Coast, Namibia, and South Africa were 21.2-54.0 and 30.9-62.6, respectively (Robinette 1963; Smithers 1983; Grimbeek 1991; Bailey 1993; Jenny 1996; Marker and Dickman 2005; Stein 2008 ). Rariges of mass (kg) without stomach contents for 9 females and 8 males were 26.8-36.4 and 33.6-60.0, respectively, in Zambia (Wilson 1968 ). Body mass of adults from the former Soviet Union ranged from 32 to 60 kg (Heptner and Sludskii 1992) .

Mean length of head and body for 2 females and 2 males from Namibia were 106.5 and 132.0 em, respectively (Stein 2008) . Ranges of mean total length (em) for 20 females and 23 males from South Africa and Namibia were 185. 0-198.4 and 210.0-217.5, respectively (Grimbeek 1991; Bailey 1993; Marker and Dickman 2005; Stein 2008 ). Measurements (range; em) of P. p. orientalis in the former Soviet Union (unknown sex and samples size) were: length of head and body, 120-171; length of tail, 75-102; length of hind foot, 24-26; length of ear, 7.5-8.0; height at shoulder, 50-78 (Heptner and Sludskii 1992) . Measurements for 2 female P. p. nimr from the Judean Desert (mm) were: total length, 1,684-1,920; length of head and body, 930-1,130; length of tail, 754-790; length of hind leg from the hip to foot pad, 223-230; length of ear, 59-64; width of front interdigital foot pad, 52-52; width of hind interdigital foot pad, 48-49; mass, 22-23.5 kg (IIani 1981) . Mean and range of head and body lengths and length of tail (em) for 10 P. p. nimr of unknown sex from Jordan, Sinai, Iraq, Oman, and Saudi Arabia were 197.11,160.0-226.1; 78.12, 66.0-94.0 (Harrison and Bates 1991) . Ear length of 1 animal was 44 mm (Harrison and Bates 1991) .

Mean cranial measurements (em; n in parentheses) for males and females, respectively, from Namibia were: greatest length, 17 (2), 15.6 (3); greatest width, 15.6 (2), 11.3 (3); upper canine length, 3.6 em (3), 2.7 (5-Stein 2008; Fig. 2 ). Cranial measurements (mm, -indicates missing measurement) for 2 males and 2 females, respectively, from Gabon were: greatest length, 226.5, -, 171.5, -; condylobasal length, 202.0, -, 158.5, 166.0; zygomatic width, 148.8, 147.0, 106.7, 114.5; skull width, 91.7, -, 68.7, 73.1; I-M distance, 87.8, 83.0, 64.5, 67.0; C-M distance, 71.6, 68.1, 55.2, 57.0; palatal length, 101.0, 99.8, 75.7, 76.0; mandibular length, 149.8, 145.5, 114.2, 121.5 (Dobroruka and van Bree 1965) . Mean cranial measurements (mm; range, n) for male and female P. p. delacori, respectively, from Southeast Asia were: greatest skull length, 212, 207-218, 4; 188, 180-196, 6; condylobasal length, 195, 186-202, 4; 172, 165-178, 5; zygomatic width, 139, 134-148,4; 125, 116-130,6; greatest skull width, 69.0, 67-71, 2; 65,65,1; interorbital width, 37.6, 36-41,3; 31,31,1; postorbital width, 42.6, 42-43,3; 37, 37, 1 (Dobroruka 1963) . Mean cranial measurements (mm; range, n) for animals of unknown sex from Jordan, Sinai, Iraq, Oman, and Saudi Arabia were: greatest skull length, 193, 166-213, 6; condylobasal length, 177, 151-194, 6; zygomatic width, 121, 105-132, 7; mandibular length, 130, 111-142, 7 (Harrison and Bates 1991) . Cranial measurements (mm; range) for males and females, respectively, from the former Soviet Union (no sample size) were: greatest skull length, 193-256, 180-218; condylobasal length, 1'86-224, 170-188; zygomatic width, 123-172, 116-135; muzzle width above canines, 53-65,47-53; length of upper toothrow, 65-79, 60-68 (Heptner and Sludskii 1992) . Canines average 34.4 mm in length (Christiansen 2007) . The mandibular symphysis is not fused (Kalita et al. 2001) .

Fur color varies from a pale yellow to rich ochre to black. Fur is generally soft and thick, with individuals living in colder climates having longer fur than those from warmer habitats (Turnbull-Kemp 1967) . Fur along the ventral portion of the torso is generally lightly colored and long, regardless of the region. Fur is short and uniform in color on the rostrum and becomes spotted with solid dots along the muzzle and forehead. Whisker spots can be used for individual identification (Pennycuick and Rudnai 1970) .

The solid spots become irregular, rosette patterns along the neck and shoulders extending along the back and midsection to the rump and tail. Large, irregular spots are present along the limbs from the elbow and knee to the feet and along the ventral side of the torso. One of us (ABS) observed variation in eye color from golden yellow to a pale blue.

Panthera pardus is the most widely distributed wild cat species in the world (Fig. 3) , with a range extending from the Cape of Good Hope, South Africa, through the Middle East and Southeast Asia to the Amur Peninsula in the Russian Far East (Nowell and Jackson 1996) . The population of P. p. pardus is distributed throughout sub-Saharan regions except for the Skeleton Coast of Namibia and most of South Africa outside of the Limpopo region, Eastern and Western Cape provinces. P. p. adersi is extinct (Goldman and Walsh 2002) . In the Middle East, P. p. nimr occurs in Israel on the Arabian Peninsula (EI-Mashjary 1995; AI-Johany 2007). P. p. ciscaucasica persists in protected areas and mountainous areas of the Russian North Caucasus, Georgia, Armenia, Azerbaijan, Nagorno-Karabakh Republic, Turkey, Turkmenistan, Afghanistan, Pakistan, and Iran in southwestern Asia (Henschel et al. 2008) . P. p. fusca is pervasive throughout the protected area system within India. P. p. kotiya is present on Sri Lanka. P. p. delacouri is increasingly rare in China and Southeast Asia outside of protected areas. In northern China, P. p. japonensis also is rare. P. p. melas is still found in Malaysia, but not on the islands of Borneo or Sumatra (Nowell and Jackson 1996; Meijaard 2004) .

Panthera pardus pardus is primarily distributed throughout sub-Saharan Africa with smaller isolated populations in the Atlas Mountains of Morocco, Ahaggar in southeastern Algeria, the eastern desert of Egypt, and Niger (Ray et al. 2005) . The remaining populations of P. p. pardus are in West Africa extending from Nigeria to Senegal. The eastern and southern African population is more contiguous, although increasingly patchy, extending east to west from Somalia to Gabon and eastern Nigeria and south of the Sahel from Sudan to South Africa with an isolated population in the western Cape region (Nowell and Jackson 1996; Ray et al. 2005 ).

The origin of the modern Panthera pardus is between 470,000 and 850,000 years ago in Africa. Modern P. pardus migrated to Asia more recently, between 170,000 and 300,000 years ago (Uphyrkina et al. 2001 ). The earliest documented ancestral fossils of P. pardus were from Laetoli, Tanzania, along with lion (Panthera leo) fossils dated at approximately 3.5 million years ago (Turner and Anton Henschel et al. 2008 ). Subspecies are: 1, P. p. ciscaucasica; 2, P. p. delacouri; 3, P. p. fusca; 4, P. p. japonensis; 5, P. p. kotiya; 6, P. p. melas; 7, P. p. nimr; 8, P. p. orientalis; 9, P. p. pardus. 1997 ). The oldest fossil within Asia is from the Inain Siwaliks at approximately 2 million years ago (Hemmer 1976 ). This Asiatic P. pardus was similar in appearance to P. onca and the now extinct P. gombazogensis (Kitchener 1991) .

One of us (ABS) has observed that large male Panthera pardus may develop a dewlap extending from the lower jaw to the chest. P. pardus has large and well-defined musculature on the forelimbs, shoulders, and neck. P. pardus can drag a carcass more than double its body mass using its mouth and hoist the carcass into trees (Scheepers and Gilchrist 1991) . A 39-kg P. pardus had a forelimb length of 57 em, a hind-limb length of 76 em, and an intergirdle distance of 74 em (Day and Jayne 2007) . In length, a humerus was 21 em and a femur was 23 em (Ray et al. 1996; . Limb bone mean lengths and circumferences (mm) for 5 P. pardus were: humerus, 211. 2, 90.1; radius, 263.6, 60.9; femur, 353.4, 87.0; tibia, 302.3, 83.4 (Christiansen 1999) . Meniscal ossicles are present (Walker et al. 2002) . Dental formula is i 3/3, c 1/1, p 3/2, m 1/1, total 30 (Stander 1997) .

Hair of P. p. nimr is about 1.5-2.0 em in length on dorsum and 4-5 em in length on ventrum (Borner 1977) . Hair on the distal part of the tail is longer than on the proximal region, thus the last one-third of the tail appears to have a larger diameter (Borner 1977) . Hair density is about 3,000 hairs/em.' on dorsum with 1 guard hair for every 4 underfur hairs (Heptner and Sludskii 1992) . Yellow guard hairs are "30 mm long and 121 microns thick but the black hairs [are] 40 mm long and 96 microns thick" (Heptner and Sludskii 1992:204) . Similarly, yellow underfur is 20 mm long (21 urn thick) and black underfur is 24 mm long (32 urn thick-Heptner and Sludskii 1992). Dark, smokey-gray dorsal underfur in a winter coat is up to 19 mm long (Pocock 1934) . Winter hair (45-50 mm) is longer than summer hair (20-25 mm -Pocock 1934; Heptner and Sludskii 1992) .

A 94-cm-Iong adult female had the following measurements (cm or g): height at withers, 67; height at hind limb, 69; length of head, 27; width of head, 15.5; mass of head, 2,000; mass of skull and mandible, 550; orbital cavity, 6 by 5; eyeball mass, 45; eyeball size, 3.4 by 3.2 by 3.3; cornea lateromedial by dorsoventral diameters, 2.2 by 1.8; mass of larynx, 105; thyroid size, 6.0 by 1.6 by 0.6; tracheal length, 26 (with 43 tracheal rings); lung mass with trachea, 405; heart mass, 175; heart circumference, 22.5; esophageal length, 42; mass of stomach, 250; lesser curvature of stomach, 24.5; greater curvature of stomach, 44; length of small intestine, 106; length of large intestine, 75.2; cecal length, 3.2; rectal length, 40; mass of intestines, 650; mass of spleen, 80; size of spleen, 21.5 by 4.5 by 1.8; mass of 5-lobed liver, 525; length of hepatic duct, 13.7 (Archana et al. 2006 ). An adult female had 13 ribs, a left lung with 2 lobes, a right lung with 5 lobes, and a gall bladder (Archana et al. 2006) .

Panthera pardus has a digitigrade foot structure with the forefeet having 5 toes and the hind feet having 4. The 1st toe, set on the inside of the foot above the wrist, is only used when bringing down prey. The feet of adult male P. pardus are 70-90 mm in length and width, forming a near circular track for the forefoot (Stuart and Stuart 1994) . The hind foot is often slightly longer than it is wide, but conforms to these measurements. The female feet are similar but are 55-70 mm in length and are 5-10 mm more slender (Stuart and Stuart 1994) . The metacarpal-phalanx ratio is 1.87 (Iwanluk et al. 2001) . Maximal speeds of 60 km/h, horizontal leaps of 6 m, and vertical leaps of 3 m are reported in the secondary literature (Nowak 1999) .

Typically, feces of P. pardus are a cylindrical, sausage shape with a diameter between 20 and 30 mm (Stuart and Stuart 1994) . Feces of P. pardus are primarily made up of undigested hair and bone (Keogh 1983) .

Mean measurements (em or g) of a pair of kidneys from an adult female were: length, 7.12; width, 5.12; thickness, 3.47; cortical thickness, 0.71; paracortical thickness, 0.96; medullary thickness, 1.08; mass, 158 (Sarma et al. 2004) . A neonate had a cardiac foramen ovale (Macdonald and Johnstone 1995) .

Acoustical compliances (em:' of air) were: middle-ear cavity, 4.3; tympanic membrane-ossicular, 2.15; total middle ear, 1.41 (Huang et al. 2000) . Minimum frequency for acoustic reflectance was 0.77 kHz with an admittancenotch frequency of 3.12 kHz (Huang et al. 2000) . Regarding the larynx, the longitudinal length of the vocal fold (at rima glottis) is 20 mm and the distance from the base of the cricoid cartilage to the superior border of the vocal fold is 50 mm (Hast 1989 ).

Ranges of heart rate (beats/min), respiration (breaths/ min), and rectal temperature (OC) of immobilized P. pardus after 20 and 70 min, respectively, were: 70-88, 55-95; 6-28, 11-30; 39.1-40.6, 37.8-38.8 (Belsare and Athreya 2010) . Hemoglobin of P. pardus has 2 forms, 80% of which is the major form (Abbasi and Braunitzer 1985; Ahmed et al. 1988 ). Mean hematological values were: blood clotting time, 5.29 min; hemoglobin 12.3-14.83 g/dl; hematocrit, 0.37-0.47; packed cell volume, 39-43%; red blood cells, 6.95-10.26 X 10 6/JlI; mean corpuscular volume, 45.5-61.6 um'; mean corpuscular hemoglobin, 14.0-21.4 pg (Hawkey and Hart 1986; Jain 1986; Pospisil et al. 1987; Singh et al. 1999a ). The differential counts (%) from 10.3-15.0 X 10 6/ml leukocytes were segmented neutrophils, 61-84; banded neutrophils, 4; basophils, 0-1; eosinophils, .0-11; monocytes, 1-3; lymphocytes, 9-23 (Hawkey and Hart 1986; Jain 1986; Pospisil et al. 1987) . Serum calcium and inorganic phosphorus (mg/dl) were 10.97 and 3.63, respectively, from 1 female (Singh et al. 1999b) . Plasma protein and fibrinogen (g/dl) were 8.1 and 0.1, respectively (Jain 1986 (Crissey et al. 2003) .

In females, baseline serum progesterone is 1.6 ng/ml and increases to 13-98 ng/ml during the luteal phase (Schmidt et al. 1988 ); fecal progesterone is 706-732 pg/g (De Haas van Dorsser et al. 2007) . Progesterone stays at basal levels when animals are isolated (Schmidt et al. 1988 ). Basal concentrations (ng/ml) of luteinizing hormone and follicle-stimulating hormone in males and females, respectively, were: 1.9, 1.8; 28.7, 27.5 (Brown et al. 1989) . Basal serum estrogen in females was 8.8 pg/ml (Brown et al. 1988 ) and peak fecal estrogen (1,433 ng/g) was twice basal levels (De Haas van Dorsser et al. 2007 ). The peak urinary relaxin concentration during 2 pregnancies was 3.6-4.6 ng/mg creatinine (De Haas van Dorsser et al. 2006) . Sperm density, number of motile sperm, and number of normal sperm, respectively, were: 51.6-55.8 X 10 6/ml, 57%, and 72% (Jayaprakash et al. 2001; De Haas van Dorsser and Strick 2005) . Total sperm length, head length, and head width (urn) are: 54.6, 4.2, and 2.5, respectively (De Haas van Dorsser and Strick 2005) . Males over 8 years and under 3 years had lower sperm counts than males of intermediate ages and sperm counts were lower in summer than in winter (De Haas van Dorsser and Strick 2005) .

Scats have the following bile acids: deoxycholic, chenodeoxycholic, and dehydrocholic (Khorozyan et al. 2007 ). Urine contains cauxin (McLean et al. 2007 ). Marking fluid contains 1.15 mg/ml lipids as well as the following: acids: acetic, butyric, heptanoic, hexanoic, isoheptanoic, isohexanoic, isooctanoic, isovaleric, octanoic, nonanoic, propionic, and valerie; neutral compounds: 2-acetyl-lpyrroline, acetaldehyde, and acetone; and basic compounds: cadaverine, dimethylamine , ethylenediamine, phenylethylamine, putrescine, and trimethylamine (Poddar-Sarkar and Brahmachary 2004).

Mating occurs mid-January-mid-February in Iran (Farhadinia et al. 2009 ), January-February in Amur, and November-December in Nepal (Hayssen et al. 1993) . Births occur February-March in India and Nepal, April-May in Amur, in the spring and early summer in Pakistan, during the rainy season in Angola, at the start of the rainy and start of the dry seasons in Zaire, and year-round in South Africa (Hayssen et al. 1993) .

The mean length of estrus is 5-13 days; mean cycle length is 20-55 days and the follicular phase is 18-23 days (Hayssen et al. 1993; Cunningham and Gross 2000; De Haas van Dorsser et al. 2007 ). Gestation is 88-112 days (Acharjyo and Patnaik 1985; Hayssen et al. 1993; Cunningham and Gross 2000; De Haas van Dorsser et al. 2006 , 2007 . Lactation is 114-130 days with den emergence at 42 days (Hayssen et al. 1993 ) and independence at 13 months (Sunquist 1983; Le Roux and Skinner 1989) . Lactating females may leave cubs alone for up to 36 h (Seidensticker 1977) . Interbirth interval is 3.5-45 months with most intervals 8-12 months (Acharjyo and Patnaik 1985; Hayssen et al. 1993) .

Females have 4 mammae. Litter size is 1-6, with a mode of 2, and litters of 5 or 6 are rare (Eaton 1977; Acharjyo and Patnaik 1985; Hayssen et al. 1993; Kumar and Luna 2005) . Birth mass from secondary sources is 43-60 g (Kingdon 1977; Smithers 1983 ) but is 280-1,000 g from primary sources (Desai 1975; Acharjyo and Patnaik 1985; Shukla et al. 2003) . Neonates have. closed eyes, short fur, and pink skin on nose tip, paws, and perineal area (Desai 1975) . From tip to tip neonates averaged 43.8 em (Acharjyo and Patnaik 1985) . Sex ratio at birth was 49 males to 41 females (Acharjyo and Patnaik 1985) . Eyes open at 4-9 days, incisors erupt at 21-29 days, canines erupt at 30 days, and molars start to appear at 52 days (Desai 1975; Cunningham and Gross 2000; Shukla et al. 2003) .

Females 1st mate at 23-32 months with a 1st birth from 27 to 52 months, whereas males can 1st sire young at 1.5 years (Hayssen et al. 1993 ). In Kruger National Park an average of 28% of adult females produced young each year (Bailey 1993) . Infanticide may occur when territorial males are removed before cubs reach independence (Ilani 1990; Bailey 1993) . Fully developed dentition is present at 2 years of age (Stander 1997) . Incisors and canines show wear before premolars and molars (Stander 1997) . A table of age and tooth wear from 8 months until 10 years is available (Stander 1997) . Males have more enamel flaking and canine fractures than do females (Stander 1997 (Miquelle and Murzin 2003) . Smaller populations are present in Thailand and Malaysia (Grassman 1999) . In Rajaji National Park, India, the population of P. pardus was estimated at 14.99 individuals/LOu km 2 (Harihar et al. 2009 ). The stronghold of P. p. pardus is in Africa, where large, continuous populations still exist (Henschel et al. 2008) . Density of P. pardus ranges from 2.49 to 11.11 individuals/Inn km 2 in South Africa (Balme et al. 2010) and is 3.6 individuals/l Ofl km 2 in north-central Namibia (Stein et al. 2011) . Although a radiocollared female lived over 10 years in Thailand, at least 27 animals in zoos have lived 20-27 years (Grassman and Larney 2002; Weigl 2005) .

Space use.-Panthera pardus occupies a variety of habitats where competitors are present, prey sizes vary, and cover is variable. Considering all of these influences on movements and habitat use of P. pardus, generally home ranges of P. pardus are largest where prey availability is relatively low, although ranges are smallest where prey availability is high and cover is available. In semiarid and arid environments with low prey density, ranges of P. pardus are the largest recorded, including the Kalahari Desert (male home range X == 2,182 km 2- Bothma and Le Riche 1984) and the mountainous areas of Cape Province, South Africa (X == 388 km 2- Norton and Lawson 1985) . The ranges of P. pardus in northeastern Namibia were relatively large (X == 451 km 2 for males and 188 km 2 for females) in Kaudam National Park (Stander et al. 1997a ). In north-central Namibia P. pardus had medium to large ranges; a male's range was 108 km 2 and 2 female ranges averaged 50 km 2 with the availability of desert warthogs (Phacochoerus aethiopicus) and greater kudu (Tragelaphus strepsiceros [currently Strepsiceros strepsiceros]- Stein et al. 2011) . In rocky areas of eastern Botswana, ranges of P. pardus were 32.9 km 2 for females in Botswana (Steyn and Funston 2009) and 40-69 km 2 for males in the Cedarberg Wilderness Area, Cape Province, South Africa (Norton and Henley 1987) . Ranges of P. pardus ranges within rain-forested areas vary from medium sized such as 86 km 2 for males and 25 km 2 for females in Tai National Park, Ivory Coast (Jenny 1996) , to small range sizes of 32-46 km 2 for males and 14-26 km 2 for females in Huai Kha Kaeng National Park, Thailand (Rabinowitz 1989; Simcharoen et al. 2008) . In Nepal, 2 males had home ranges of 47 and 48 krrr', whereas a female had a range of 17 km 2 (Odden and Wegge 2005) . Ranges of P. p. pardus are smallest in forested and rocky areas such as Kruger National Park, where prey includes impala (Aepyceros melampus; X == 38 km 2 for males and 15 km 2 for females -Bailey 1993) and the Lolldaiga Hills, Kenya (X == 33 km 2 for males and 14 km 2 for females-Mizutani and Jewell 1998), but 1 female in the Serengeti, Tanzania, had a home range of 15.9 km 2 (Bertram 1982) . However, the smallest ranges were in Sri Lanka (8-10 km 2-Eisenberg and Lockhart 1972).

Individual P. pardus that have larger ranges tend to have areas of overlap with neighbors, yet core areas or territories are exclusively maintained (Bothma and Le Riche 1984; Steyn and Funston 2009) . Females typically share portions of their territories with their female offspring (Bailey 1993; Steyn and Funston 2009) .

Panthera pardus feeding on large prey items may remain in a single location for several days (Bothma and Le Riche 1984; Bailey 1993 ), but will typically move through its entire home range over a period of7-10 days (Mizutani and Jewell 1998) . Human disturbances may influence the range use and activity patterns of P. pardus (Marker and Dickman 2005) .

Typically P. pardus is nocturnal with peak activity during the hours of dawn and dusk (Eisenberg and Lockhart 1972; Chambers et al. 1984; Bailey 1993) or diurnal with peak activity during late morning and late afternoon-early evening (Norton and Henley 1987) . The homing instinct is strong in P. pardus (Stander et al. 1997b) , making translocation an impractical solution to conflict with people. Radiotagged P. pardus returned to their original range in proportion to the distance from the release site (Stander et al. 1997b) .

Juveniles remain with their mother 12-18 months (Bailey 1993) . Young males disperse, whereas young females often take over part of their mother's range (Bailey 1993) .

Diet.-Diet selection of Panthera pardus is primarily driven by opportunity to catch and maintain possession of its prey. Although it prefers prey within the range of 10-40 kg, in the absence of larger competitors, it may feed on larger prey (Hayward et al. 2006; Stein 2008) . The costs of attempting to kill larger prey (> 150 kg) may restrict the diet of P. pardus, although it has been recorded feeding on prey in this size range (Scheepers and Gilchrist 1991) . As well, P. pardus can persist on a variety of smaller prey in environments of lean resources or an absence of larger prey (Stuart and Stuart 1993; Hayward et al. 2006 ). Most (69%) kills by P. pardus in southern India were < 50 kg (Johnsingh 1992) . In Africa, P. p. pardus feeds on diverse species depending on ungulate species available. Primary prey are impala in Kruger National Park, South Africa, and Rhodes Matopos National Park, Zimbabwe (Smith 1978; Bailey 1993) ; impala, springbok (Antidorcas marsupialis), and small antelope in the Kalahari, South Africa (Mills 1990; Owen Smith and Mills 2008) ; impala, bush duiker (Sylvicapra grimmia), nyala (Tragelaphus angasii), red duiker (Cephalophus natalensis), southern reedbuck (Redunca arundinum), and desert warthog in the Phinda-Mkhuze complex, South Africa (Balme et al. 2010 ); rodents, bush-pig (Potamochoerus larvatus), and red-flanked duiker (Cephalophus rufilatus) in Lope National Park, Gabon (Henschel et al. 2005) ; rock hyrax (Procavia johnstoni [currently P. capensis]) and groove-toothed rat (Otomys) on Mt. Kenya (Roedel et al. 2004) ; chital (Axis axis) in Nepal (Odden et al. 2010) and India (Arivazhagan et al. 2007 ); cattle (Bos taurus), northern plains gray langur (Presby tis entellus [currently Semnopithecus entellus]), goral (Naemorhedus goral), and dogs (Canis lupusfamiliaris) in the Himalayas (Mukherjee and Mishra 2001) ; bezoar goats (Capra aegagrus) in Armenia (Khorozyan and Malkhasyan 2003) ; tufted deer (Elaphodus cephalophus) and bamboo rats (Rhizomys sinense) in the Wolong Reserve, China (Johnson et al. 1993) ; and sambar (Cervus unicolor [currently Rusa unicolor] ), muntjac (Muntiacus), Gee's golden langur (Trachypithecus geei), goral, and livestock in Bhutan (Wang and Macdonald 2009 ). In mountainous and semiarid areas, P. pardus preys upon small prey such as rock hyrax, bush duiker, and crested porcupine (Hystrix cristata- Bothma and Le Riche 1984; Norton et al. 1986; Stuart and Stuart 1993; Stander et al. 1997a ). In the absence of larger predators P. pardus may prey on slightly larger prey such as greater kudu (Karanth and Sunquist 1995; Stein 2008 greater cane rat (Thryonomys swinderianus), and striped ground squirrel (Xerus erythropus); and miscellaneous: domestic dog, African civet (Civettictis civetta), common genet (Genetta genetta), common dwarf mongoose (Helogale parvula), scrub hare (Lepus saxatilis), tree pangolin (Manis tricusp is), ground pangolin (M. temminckii), long-tailed pangolin (M. tetradactyla), banded mongoose (Mungos mungo), aardvark (Orycteropus ajer) , birds, reptiles, fish, dung beetles (Fey 1964; Pienaar 1969; Eisenberg and Lockhart 1972; Le Roux and Skinner 1989; Edgaonkar and Chellam 1998; Zuberbuhler 2001; Roedel et al. 2004; D' Amour et al. 2006; Bodendorfer et al. 2006; Odden and Wegge 2009) , and perhaps the Indian giant squirrel (Ratuja indica-Mehta 1997) . The probability of a kill is greatest in areas with intermediate cover (Balme et al. 2007 ). P. pardus will eat grass (Isachne beuttneri and Streptogyna crinite-Hoppe-Dominik 1988) and drink water every 2.7 days in the Kalahari, South Africa (Bothma 2005) .

Cannibalism can occur (Pienaar 1969; Bodendorfer et al. 2006; Steyn and Funston 2006) . P. pardus, in general, does not target domestic stock or humans, but particular individuals may develop the habit of raiding livestock or human settlements (Sterndale 1884; Corbett 1947; Turnbull-Kemp 1967; Mizutani 1999) . A P. pardus killed 51 sheep and lambs in a single event (Stuart 1986 ). In Kashmir, 48.5% of 35 attacks by P. pardus on humans were fatal (Nabi et al. 2009 ), whereas in Uganda, 32.5% of 114 attacks were fatal (Treves and Naughton-Treves 1999) .

Diseases and parasites.-External parasites include flies: . Lipoptena chalcomelaena and Wohlfahrtia magnifica; ticks:

Amblyomma hebraium, A. nuttali, A. thaolloni, A. variegatum, Haemaphysalis aciculifer, H. bispinosa, H. concinna, H. den tipalp is, H. elliptica, H. hystricis, H. konigsbergeri, H. leachi, H. papuana, H. parmata, Ixodes cavipalpus, I. cumulatimpuctatus, I. moreli, I. muniensis, I. oldi, I. pilosus, I. rasus, I. vanidicus (Turnbull-Kemp 1967; Bailey 1993; Rosen et al. 1998; Apanaskevich et al. 2007) . Mange increased during the early dry season (Bailey 1993) .

Endoparasites include acanthocephalans: Acanthocephala, Cucullanorhynchus constrictruncatus; cestodes: Hymenolepididae, Mesocestoides, Pseudophyllidae, Spirometra, and Taeniidae; protozoa: Babesia, Coccidia, Cryptosporidium, Giardia, Sarcocystis, (Strauss and Sivanandam 1966; Turnbull-Kemp 1967; Somvanshi et al. 1987; Bailey 1993; Pythal et al. 1993; Patton and Rabinowitz 1994; Tehsin 1996; Upadhye and Dhoot 2000; Dhoot et al. 2002; Penzhorn et al. 2002; Gawande et al. 2007; Amin et al. 2008; Huttner et al. 2009; Mowlavi et al. 2009; Fayer 2010) .

Some P. pardus were seropositive for feline immunodeficiency virus (Troyer et al. 2005 ) and for type 2 feline coronavirus (Kennedy et al. 2002) . The yeast M alassezia symphodialis was isolated from the ear canal of 2 P. pardus (Coutinho et al. 2006) . The bacteria Salmonella enteritidis and S. typhimurium occur in P. pardus (Babu et al. 1993 ). Pyometra in a captive, 14-year-old female was treated by ovariohysterectomy and systemic antibiotics (McCain et al. 2009 ). Adenocarcinoma (Ranganath et al. 2008) , uterine leiomyoma (Siegal-Willott et al. 2005) , histiocytoma (Nath et al. 2006) , lymphosarcoma (Sujatha et al. 2005) , squamus cell carcinoma (Sabapara et al. 2003) , avian influenza H5N1 (Keawcharoen et al. 2004) , bovine tuberculosis (Renwick et al. 2006) , Clostridium perjringens enterotoxicosis (Neiffer 2001) , degenerative spinal disease (Kolmstetter et al. 2000) , feline enteritis (Singh et al. 1983) , hepatitis (Gupta 1978) , hiatal hernia (Kearns et al. 2000) , neoplasia (Owston et al. 2008) , pulmonary anthracosis (Sujatha et al. 2007) , and rabies (Jayakumar et al. 1989 ) occur in P. pardus.

Interspecific interactions.-Although they coexist, tigers (Panthera tigris) restrict the distribution of P. pardus in Nepal (Seidensticker 1976; Odden et al. 2010 ); 5 P. pardus were killed by P. tigris in Nepal (McDougal 1988) . In Rajaji National Park in India, with the displacement of human communities and the increased population, densities of P. pardus declined sharply and diet of P. pardus showed a pronounced shift (Harihar et al. 2011) . Diet of P. pardus can be 'an indicator of the presence of intraguild competitors, where dietary shifts signaled the decline of a P. tigris population in India (Ramakrishnan et al. 1999) . In Cameroon evidence of changes in the population of P. pardus were not always tied to the presence of one larger competitor, however, because reduced lion (Panthera leo) numbers did not signal an increase in the population of P. pardus where spotted hyenas (Crocuta crocuta) were still present (Croes et al. 2011) . Dietary overlap of P. pardus with cheetahs (Acinonyx jubatus) is 68.7%, with wild dogs (Lycaon pictus) is 65.7%, and with P. leo is 39.1% (Hayward and Kerley 2008) . P. leo, spotted hyenas, wild dogs, and P. tigris will opportunistically kill P. pardus or their cubs, just as P. pardus will kill the unprotected cubs of intraguild members. These larger predators also take kills from P. pardus (Schaller 1972; Mills 1990; Creel and Creel 2002) . P. pardus tends to select smaller prey when inhabiting areas with larger competitors Sunquist 1995, 2000) . In Bandipur, India, P. pardus accounted for 15% of 379 kills, whereas dholes (Cuon a/pinus) took 80%, and P. tigris took 5% (Johnsingh 1983) . Human agropastoralists regularly scavenge the prey of P. pardus (Treves and Naughton-Treves 1999) .

Panthera pardus can be anesthetized with a mixture of tiletamine hydrochloride and zolazepam hydrochloride at dosages of 4-5 mg/kg (Swanepoel et al. 2010) or a xylazine hydrochloride (1.4 mg/kgj-ketamine hydrochloride (5 tag] kg) mixture (Belsare and Athreya 2010) . When mass is not estimated an initial dose of 50 mg xylazine-150 mg ketamine can be supplemented with 50-75 mg of ketamine only (Belsare and Athreya 2010) . Odden and Wegge (2005) also suggests using 3.6-5.9 mg/kg ofketamine and 0.07-0.12 mg/ kg of medetomidine. Other drugs and dosages are: diazepam (0.17-0.18 mg/kg), ketamine (6-12 mg/kg), xylazine (0.5-2.0 mg/kg), and telazol (2-7 mg/kg-e- Sabapara 1995) .

Novel odors have short-term (3-h) effects on behavior of captive P. pardus (Yu et al. 2009 ). P. pardus in structurally enriched enclosures is more active than those in unenriched enclosures (Mallapur et al. 2002) .

Reproductive behavior.-Adult Panthera pardus are solitary with the exception of females rearing cubs and during mating when males and females associatefor several days before separating again (Eisenberg and Lockhart 1972) .

Females attract mates through the release of scent marks and vocalizations that attract a male, which associates with her for 1-4 days (Bailey 1993) . In mating, the male mounts the female and holds the skin on the nape of her neck (De Haas van Dorsser et al. 2007) . Males often leap off the female as she aggressively snarls and occasionally strikes at the male (Seidensticker 1977) . The female rolls "on her back in front of the male and presents herself. She sits with her forelimbs extended fully on the ground, her hind limbs remaining half bent" (Desai 1975,:297) . Insertion occurs 4-8 s after mounting and coitus is 10-50 s; 5-60 copulations occurred during a 9-h period in peak estrus (Desai 1975) , whereas 13 copulations occurred during a 1.5-h period, with a mean duration of copulation of 3 s and an average interval between copulations of 6.5 min (Laman and Knott 1997) . A female associated with a male 11 days after her cubs were killed (Bailey 1993) .

Communication.-"Cubs emit low cries when hungry or uncomfortable" (Desai 1975:299) . Guttural sounds accompany coitus and at the peak of copulation both males and females make a high-pitched sound (Desai 1975) . "A leopard call consists of a repeated pattern of strokes sounding much like the sawing of wood" (Eisenberg and Lockhart 1972:71) . Average number of strokes per call is 13-16 with a range of 2-30 with an intercall interval of6 min and a range of 1-10 min (Ulmer 1966; Eisenberg and Lockhart 1972) . Duetting may occur (Eisenberg and Lockhart 1972) . Both males and females roar (Ulmer 1966) . Marking behavior includes tree scratching and soil scrapes (Eisenberg and Lockhart 1972) . Scent marking, roaring, and conspicuous behaviors maintain spacing distances (Muckenhirn and Eisenberg 1971) .

Miscellaneous behavior.-In Kenya and South Africa, 66% of activity of Panthera pardus is nocturnal (Hayward and Slotow 2009 ). In the rain forests of West Africa, P. pardus is diurnal with strong individual prey preferences (Jenny and Zuberbuhler 2005) . In Oman, P. p. nimr was most active at 0200-0700 h and least active at 1200-1500 h (Spalton et al. 2006) .

Panthera pardus attacks its prey in a variety of ways, but primarily stalks to within a short distance of its target before pouncing (Stander et al. 1997a) . Females with cubs increase foraging efficiency by killing smaller prey (Bothma and Coertze 2004a) . P. pardus kills smaller prey by biting the nape of the neck or 'puncturing the skull with its canines, whereas larger prey are bitten on the throat, avoiding the horns and antlers of antelope and deer. In the Kalahari, P. pardus exhibited a flexible hunting strategy that did not regularly include the typical stalk-ehase-kill sequence, but rather involved longer stalking periods at further distances related to the target prey species and reduced available cover (Bothma and Le Riche 1989) .

After making a kill P. pardus will either eat a small prey item immediately or cache the kill for feeding in safety (Bothma and Le Riche 1984; Bailey 1993) . P. pardus may drag its kills several hundred meters to specific types of trees or bushes of a prescribed height, trunk thickness, and foliage density (Bothma and Le Riche 1984; Bailey 1993) . In Kruger National Park, P. pardus hoisted 84% of its kills into trees (Bailey 1993) . In the Kalahari Desert, Tsavo National Park, Kenya, or the commercial farmlands of north-central Namibia, where larger predators are less common, P. pardus cached its kills under bushes (Hamilton 1976; Bothma and Le Riche 1984; Stein 2008) . In northern Botswana, male P. pardus tended to take prey to trees more than females (Stein et al. 2010) . P. pardus also will cache kills in caves (de Ruiter and Berger 2001) .

In northeastern Namibia, P. pardus staggered its activities in different parts of its shared range when another animal was occupying the area (Stander et al. 1997a) . P. pardus fights on rare occasions, usually when a newcomer challenges the resident animal (Corbett 1947; Hamilton 1976; Bailey 1993) .

In the Kalahari, adult males scent marked 2.3 times more than adult females without cubs and 5.9 times more often than females with cubs (Bothma and Coertze 2004b) . The frequency of scent marking increased during courting and bouts of mating. Tree scratching also is used to mark territory, but less frequently than scent marking (Bothma and Coertze 2004b) . Scrapes in Iran had a mean length of 39.3 cm and a mean width of 22.7 em (Ghoddousi et al. 2008) . P. pardus prefers specific tree species for scratching, for example Acacia erioloba in the Kalahari or water berry (Syzygium corda tum) in the Soutpansberg mountains of South Africa, but may not scratch trees in other parts of its range (Hamilton 1976; Stuart and Stuart 1994; Bothma and Le Riche 1995) .

Diploid chromosome number (2n) is 38 with a fundamental number (FN) of 72 and includes 5 metacentric, 7 submetacentric, 4 acrocentric, and 2 telocentric pairs (Hsu et al. 1963; Tanomtong et al. 2008) . The X chromosome is a small submetacentric and the Y chromosome is the smallest metacentric (Tanomtong et al. 2008) .

The most genetically diverse (expected heterozygosity, 0.77-0.80) population of Panthera pardus pardus is found in sub-Saharan Africa, whereas the lowest genetic variation (expected heterozygosity, 0.340-0.356) occurs in the isolated Amur peninsula, P. p. orientalis, of the Russian Far East (Spong et al. 2000; Uphyrkina et al. 2001 Uphyrkina et al. , 2002 . For P. p. kotiya, percent polymorphism and percent average heterozygosity for wild-caught, captive-born, and melanistic P. pardus, respectively, were 4.0, 1.2; 4.0, 1.4; 4.0, 2.0 (Miththapala et al. 1991) . Two of 12 loci were polymorphic in P. pardus (Newman et al. 1985) . Inbreeding coefficients for captive P. pardus ranged from 0 to 0.5 (Shoemaker and Wharton 1984) . V-chromosome, mitochondrial, and autosomal DNA suggest that P. leo and P. pardus are sister taxa (Davis et al. 2010) .

Melanism is inherited as a recessive trait (nonagouti) of the agouti locus in P. pardus (Robinson 1969 (Robinson , 1970 Roychoudhury and Acharjyo 1984) and is not the result of the 2-base pair deletion in the ASIP gene or either of 2 "inframe" deletions in the Me lR gene (Eizirik et al. 2003 ). Albino P. pardus have been seen (Divyabhanusinh 1993) . Hybrids between P. pardus and P. leo, P. onca, P. tigris, and cougars (Puma concolor) have been reported (Gray 1971) .

Microsatellite DNA from scats can individually identify P. pardus (Perez et al. 2006; Mondol et al. 2009 ), as can spot patterns (Miththapala et al. 1989) . Artificial insemination can be successful (Dresser et al. 1982) . Cytochrome b from fecal genetic material can distinguish sympatric tigers and P. pardus (Nagata et al. 2005 (Ray et al. 2005) . The range of P. pardus has been drastically reduced worldwide and reduced approximately 37% throughout Africa (Ray et al. 2005) . As of 1964, only 10-15 P. p. orientalis were estimated to live in the former Union of Soviet Socialist Republics (Bannikov 1964) . Conservation efforts and the regulation of the distribution of trophy hunting permits can reduce mortality (Balme et al. 2009 ). Efforts to distribute financial benefits from trophy hunting and photographic tourism may also mitigate conflicts with farmers (Stein et al. 2010 ).

On the International Union for Conservation of Nature and Natural Resources (IUCN) Red List of Threatened Species, P. p. melas, P. p. orientalis, and P. p. nimr are "Critically Endangered" (Ario et al. 2008; Jackson and Nowell 2008; Mallon et al. 2008 ), whereas P. p. kotyi and P. p. ciscaucasica are "Endangered" (Khorozyan 2008; Kittle and Watson 2008) . In 2011, the total captive population of P. p. nimr consisted of 42 males, 32 females, and 3 unsexed individuals derived from 14 founders (Budd and Leus 2011) .

The earliest writings of Panthera pardus were recorded in Sumeria dating back to 3100 BC (Turnbull-Kemp 1967) . In Sumerian culture, the god Nin-urta used cyclone winds as a weapon, in the form of the P. pardus-headed Shargaz. The Chinese believe P. pardus to be 1 of 4 beasts of power (Turnbull-Kemp 1967) . P. pardus is present on ancient Egyptian hieroglyphics and drawings (Budge 1978) . Ancient Rome used P. pardus for gladiatorial fighting.

.In East Africa, killing a P. pardus can assist a young Maasai male achieve warrior status (Hazzah et al. 2009 ). Skins of P. pardus are used in the ceremonial dress of cultures throughout Africa, and consuming meat or genitalia of P. pardus is thought to transfer the power and stealth of the P. pardus (Turnbull-Kemp 1967) .

review of previous versions of this manuscript by T. K. Fuller, S. DeStefano, and L. L. Sievert.

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The spotted ambassadors of a vanishing world

Opinion 417. Rejection for nomenclatural purposes of volume 3 (zoologie) of the work by Lorenz Oken entitled Okens Lehrbuch der Naturgeschichte published in 1815-1816. Opinions and Declarations Rendered by the International Commission on Zoological Nomenclature

Are long digits correlated with high forepaw dexterity? A comparative test in terrestrial carnivores (Carnivora)

Panthera pardus ssp. orientalis. International Union for Conservation of Nature and Natural Resources 2010. International Union for Conservation of Nature and Natural Resources Red list of threatened species

Schalm's veterinary hematology. Lea and Febiger

Rabies in a wild leopard, Felis (Panthera) pardus

Semen characteristics of the captive Indian leopard, Panthera pardus

Spatial organization of leopard (Panthera pardus) in Tai National Park, Ivory Coast: is rain forest habitat a tropical haven

Hunting behaviour in West African forest leopards

Large mammalian prey-predators in Bandipur

Prey selection in three large sympatric carnivores in Bandipur

Food habits of Asiatic leopards (Panthera pardus fusea) in Wolong Reserve

Anatomy of the mandible of Indian leopard

Prey selection by tigers, leopards, and dhole in tropical forests

Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon a/pinus) in Nagarahole

Hiatal hernia and diaphragmatic eventration in a leopard (Panthera pardus)

Avian influenza H5N1 in tigers and leopards

Detection of feline coronavirus in captive Felidae in the USA

A photographic reference system of the microstructure of the hair of southern African bovids

Panthera pardus ssp. saxicolor. International Union for Conservation of Nature and Natural Resources 2010. International Union for Conservation of Nature and Natural Resources Red list of threatened species

Using thin-layer chromatography of fecal bile acids to study the leopard (Panthera pardus ciscaucasica) population

Ecology of the leopard (Panthera pardus) in Khosrov Reserve, Armenia: implications for conservation. Scientific Reports, Societa Zoologica "La Torbiera

IlIA: carnivores

Natural history of wild cats

Panthera pardus ssp. kotiya. International Union for Conservation of Nature and Natural Resources 2010. International Union for Conservation of Nature and Natural Resources Red list of threatened species

Degenerative spinal disease in large felids

Retrospective study on reproductive patterns of large wild cats in captivity

An observation of leopard (Panthera pardus Linnaeus) mating behaviour in Serengeti National Park

A note on the ecology of the leopard (Panthera pardus Linnaeus) in the Londolozi Game Reserve

Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata

Mammals collected in central

A hand-book to the Carnivora. Part 1. Cats, civets, and mungooses

Notes on two African mammals

The game animals of Africa

Harmsworth's natural history. A complete survey of the animal kingdom with thousands of photographs from life and an unrivalled series of colour plates

Ward's records of big game with their distribution, characteristics, dimensions, weights, and horn and tusk measurements

Comparative anatomy of the cardiac foramen ovale in cats (Felidae), dogs (Canidae), bears (Ursidae) and hyaenas (Hyaenidae)

Enclosure design and space utilization by Indian leopards (Panthera pardus) in four zoos in southern India

Panthera pardus ssp. nimr, International Union for Conservation of Nature and Natural Resources 2010. International Union for Conservation of Nature and Natural Resources Red list of threatened species

Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian leopards

Die geographische Verbreitung der Katzen und ihre Verwandtschaft untereinander

Uber chinesische Saugetier, besonders aus den Sammlungen de Herrn Wilhelm Filchner

Pyometra in captive large felids: a review of eleven cases

Leopard and tiger intereactions at Royal Chitwan National Park

Characterization of cauxin in the urine of domestic and big cats

Leopard (Panthera pardus) attempting to prey on Indian giant squirrel (Ratufa indica centralis)

Biogeographic history of the Javan leopard Panthera pardus based on a craniometric analysis

Zoologische Annalen, Erster Band Von Jahre 1793

The Kalahari hyena: comparative behavioral ecology of two species

Zoologie et Paleontologie, Comprenant L'Anatomie, la Physiologie, la Classification et l

Spatial distribution of far eastern leopard in Southwest Primorski Krai, and recommendations for their conservation. Wildlife Conservation Society, World Wildlife Fund and Tigris Foundation

Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation

Identification of individual leopards (Panthera pardus kotiya) using spot pattern variation

Genetic variation in Sri Lankan leopards

Impact of leopards on a working ranch in Laikipia Kenya

Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya

A panel of microsatellites to individually identify leopards and its application to leopard monitoring in human dominated landscapes

Trichinella britovi in a leopard (Panthera pardus saxicolor) in Iran

Home ranges and predation of the Ceylon leopard (Panthera pardus fusca). Pp. 142-176 in The world's cats

Predation by leopard Panthera pardus in Majhatal Harsang wildlife sanctuary, western Himalya

Comparison of injury pattern in victims of bear (Ursus thibetanus) and leopard (Panthera pardus) attacks. A study from a tertiary care center in Kashmir

Fecal genetic analysis using PCF-RFLP of cytochrome b to identify sympatric carnivores, the tiger Panthera tigris and the leopard Panthera pardus, in far eastern Russia

A case of histiocytoma in a leopard Panthera pardus

Clostridium perfringens enterotoxicosis in two Amur leopards (Panthera pardus orientalis)

Revision of the jaguars

Die von mir in den Jahren 1892-95 in Ost und Central-Africa, speciell in den Massai-Landern und den Landern am Victoria nyansa gesammelten und beobachteten Saugethiere

Biochemical genetic variation in eight endangered or threatened felid species

Home range and movements of male leopards in the Cedarberg Wilderness Area, Cape Province

Radio tracking of leopards and caracals in the Stellenbosch area, Cape Province

Prey of leopards in four mountainous areas of the south-western Cape Province

Walker's mammals of the world

Wild cats: status, survey and conservation action plan. International Union for Conservation of Nature and Natural Resources/Species Survival Commission Cat Specialist Group

Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park

Kill rates and food consumption of leopards in Bardia National Park

Do tigers displace leopards? If so, why?

Lehrbuch der naturgeschichte

Predator-prey size relationships in an African large-mammal food web

Neoplasia in felids at the Knoxville Zoological Gardens

Parasites of wild Felidae in Thailand: a coprological survey

A method of identifying individual lions Panthera leo with an analysis of the reliability of identification

Seroprevalence of Toxoplasma gondii in free-ranging lion and leopard populations in southern Africa

Critically endangered Arabian leopard Panthera pardus nimr in Israel: estimating population parameters using molecular scatology

Predator-prey relationships amongst the larger mammals of the Kruger National Park

Notes upon some African species of the genus Felis, based upon specimens recently exhibited in the Society's gardens

Descriptions of two subspecies of leopards

The panthers and ounces of Asia

The panthers and ounces of Asia. Part II. The panthers of Kashmir, India, and Ceylon

Proceedings of the Zoological Society of

The leopards of Africa

Putative chemical signals of leopard

Basic haematological values in carnivores. II. The felidae

Death of a wild Indian leopard Panthera pardus fusca (Meyer) due to parasitism with the lung fluke Paragonimus . westermanii (Kerbert, 1878) and the hookworm Galoncus perniciosus (Linstow, 1885)

The density and behaviour of large cats in a dry tropical forest mosaic in Huai Kha Khaeng Wildlife Sanctuary

Tiger decline caused by the reduction of large ungulate prey: evidence from a study of leopard diets in southern India

Adenocarcinoma in a leopard (Panthera pardus): a case report

Setting conservation and research priorities for larger African carnivores

Anatomy of the humerus of leopard (Panthera pardus)

Anatomical study of the femur (os femoris) of leopard (Panthera pardus)

Bovine tuberculosis in southern African wildlife: a multi-species hostpathogen system

The mammals of South Africa. Trustees of "The Mammals of South Africa

Weights of some of the larger mammals of northern Rhodesia

The breeding of spotted and black leopards

Inheritance of the black form of the leopard, Panthera pardus

Notes on the feeding habits of the leopard in the alpine zone of Mount Kenya

Dual infestation of a leopard by Wohlfahrtia magnifica and Lipoptena chalcomelaena

The carnivores of West Africa

Genetics of cat colour in the leopard Panthera pardus

Chemical restraint and sedation of leopards (Panthera pardus)

Squamous cell carcinoma of lungs in an Indian leopard Panthera pardus

Macromorphological study on the kidney of leopard (Panthera pardus)

Leopardus pardus tullianus Valenciennes

Opredelitel' mlekopitayushchikh Rossiiskoi Imperlii. [Guide to the mammals of Imperial Russia

The Serengeti lion: a study of predator-prey relations

Leopard predation on giraffe calves in the Etosha National Park

Boekdrukkerij van de Gebroeders Nys, voor Rekening van de Koninklijke Akademie voor Zee-en Landmagt

Serum concentrations of oestradiol and progesterone, and sexual behaviour during the normal oestrous cycle in the leopard (Panthera pardus)

Die Saugthiere in Abbildugen nach der Natur mit Beschreibungen

Die Saugthiere in Abbildungen nach der Natur mit Beschreibungen 1776-1778. Wolfgang Walther, Erlangen

Report on additions to the Society's menagerie in February 1878

On the ecological separation between tigers and leopards

Notes on early maternal behavior of the leopard

An analysis of inbreeding within leopards in captivity

Relationship of age with body weight in orphaned leopard cubs

Chronic obstipation in a leopard (Panthera pardus) caused by intrapelvic uterine leiomyoma compression of the distal colon

Home range size and daytime habitat selection of leopards in Huai Kha Khaeng Wildlife Sanctuary

Infectious feline enteritis in panther cubs

Hematology of tigers (Panthera tigris), leopards (Panthera pardus), and clouded leopards (Neofelis nebulosa) in captivity

Serum calcium and inorganic phosphorus in tigers (Panthera tigris) and leopard (Panthera pardus) kept in captivity

Movement patterns and feeding behavior of the leopard in the Rhodes Matopos National Park

The mammals of the southern African subregion

Sarcocystis in a leopard (Panthera pardus)

Critically endangered Arabian leopards Panthera pardus nimr persist in the Jabal Samhan Nature Reserve

High genetic variation in leopards indicates large and long-term stable effective population size

Field age determination of leopards by tooth wear

sic] Kaqece, and II. [sic] Ghau. 1997a. The ecology of asociality in Namibian leopards

Nonconsumptive utilization of leopards: community conservation and ecotourism in practice

Ecology and conservation of the leopard (Panthera pardus) in north cenrtal Namibia

Farm management and economic analyses of leopard conservation in north-central Namibia

Leopard population and home range estimates in north-central Namibia

Mammalia of India and Ceylon

A case of cannibalism in leopards

Land-use and socio-spatial organization of female leopards in a semi-arid wooded savanna

A double infection of filariasis in a black panther (Panthera pardus) from Pahang

The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province

Prey of leopards in the western Soutpansberg

A field guide to the tracks and signs of southern and East African wildlife

Pulmonary anthracosis in large wild felids

Lymphosarcoma in a leopard. A case report

Dispersal of three radiotagged leopards

Factors affecting location failure of GPS collars fitted to African leopards (Panthera pardus)

Catalogue of the mammals of China (south of the river Yangtsze) and of the island of Formosa

Cy~ogenetic study of the leopard, Panthera pardus (Carnivora, Felidae) by conventional staining, G-banding and high-resolution staining technique

Panther, Panthera pardus (Linnaeus) with guinea worm infection

with descriptions of allied species from other localities

The mammals of the tenth edition of Linnaeus: an attempt to fix the types of the genera and the exact bases and localities of the species

The Duke of Bedford's zoological exploration of eastern Asia. XIV. On mammals from southern Shen-si, central China

Risk and opportunity for humans coexisting with large carnivores

Catalogus mammalium tam viventium quam fossilium. Quinquennale supplementum anno 1904. R. Friedlander and Sohn

Seroprevalence and genomic divergence of circulating strains of feline immunodeficiency virus among Felidae and Hyaenidae species

The leopard

The big cats and their fossil relatives

Voices of the Felidae

Sudden death of a leopard (Panthera pardus) due to babesiosis

Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus

Conservation genetics of the far eastern leopard (Panthera pardus orientalis)

Sur une novelle espece de panthere

Schreber's Die Saugthiere in Abbildugen nach der Natur mit Beschreibungen

Meniscal ossicles in large non-domestic cats

Feeding habits and niche partitioning in a predator guild composed of tigers, leopards and dholes in a temperate ecosystem in central Bhutan

Records of big game with their distribution, characteristics, dimensions, weights, and horn and tusk measurements

Das Fellmuster der wildebenden Katzenarten und der Hauskatze in Vergleichender und Stammesgeschicher Hinsicht

Longevity of mammals in captivity; from the living collections of the world. A list of mammalian longevity in captivity

Weights of some mammals from eastern Zambia

Effects of odors on behaviors of captive Amur leopards Panthera pardus orientalis

Predator-specific alarm calls in Campbell's monkeys, Cercopithecus campbelli

Persische panther

Weitere Mitteilungen fiber Persische Panther

We thank K. Doyle, curator of the mammal collection at the University of Massachusetts, Amherst, for the skull of Panthera pardus skull from their collection. We also thank F. Belcharid, C. Burton, A. Sanei, and C. Stuart for information related to the current range of P. pardus. J. Edwards helped with the map. We appreciate the critical

Associate editor of this account was DAVID ZEGERS~Editor was MEREDITH J. HAMILTON.