Panthera leo (Linnaeus, 1758)

Don E. Wilson & Russell A. Mittermeier, 2009, Felidae, Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona: Lynx Edicions, pp. 54-168 : 137-138

6.

Lion

Panthera leo

French: Lion / German: Lowe / Spanish: Leon

Taxonomy. Felis leo Linnaeus, 1758 ,

Morocco, North Africa.

Proposed races massaica, somaliensis and roosevelt: are included in nubica. Nominal race leo (Linnaeus, 1758) from north-west Africa and race melanochaita (C. E. H. Smith, 1858) from Cape region, South Africa, are extinct. Six extant subspecies currently recognized.

Subspecies and Distribution.

Pl. azandica]. A. Allen, 1924 — NE Zaire.

P. l. bleyenberghi Lonnberg, 1914 — S Zaire, Zambia, and Angola.

P. l. kruger: Roberts, 1929 — NW (Kalahari), N, and SE South Africa.

P.l. nubica de Blainville, 1843 — NE and E Africa.

P.l. persica Meyer, 1826 — from Iraq to C India in the 19" century; now restricted to the Gir Forest, India.

P. l. senegalensis Meyer, 1826 — West Africa E to the Central African Republic.

Descriptive notes. Head-body 172-250 cm (males), 158-192 cm (females), tail 61-100 cm. Shoulder height 107-123 cm; weight 190 (150-225 kg; record 272 kg) for males, and 126 (122-192 kg) for females. The Lions of the Asian subspecies persica are in general smaller. Brown amber to clear cream-colored iris. Eyes edged with white underneath. Coat of adults is generally a uniform sandy or tawny on the upperparts and flanks and white on the underparts. Some adults retain the rosettes and spots characteristic of young lions to a lesser or greater degree, even into their later years. Backs of rounded ears are black,tail is just over half the length of head and body, with well-developed tuft of long tawny to black hairs on the tip concealing a horny spur. Melanistic forms are extremely rare, very pale individuals known from Kaokoland, Namibia, and south-west Botswana. Some nearly white individuals, not albinos, from Timbavati, north-east South Africa. Hair on face, upper body parts, flanks and tail is short, on underparts softer and longer. Adult males have a mane of long hair (up to 16 cm) on the sides of the face and on top of the head, extending onto the shoulder, around the neck, and for a short distance along the spine. In sub-adults mane color is sandy, yellowish, or tawny, but may change to black with advancing age. Adult Lions lacking manes occur in the Tsavo region, Tanzania, and elsewhere uncommonly. The mane serves as a sexual signal to females and distinguishes the male at great distances. It is also an indicator of individual fitness: mane development is strongly influenced by testosterone. Some adult males have tufts of long hair on the elbows and a long band of hair extending from the mane over the chest to the anterior part of the abdomen. In the extinct North African nominate subspecies this covered the whole belly. Long whitish whiskers are arranged in parallel rows on the sides of the upperlip, each arising from a black spot, but with the top row of spots without whiskers. Recognition of individuals from this pattern of spots is possible. The massive skull is relatively flat on top. The distinguishing morphological character of the remaining Asian Lions, rarely seen in African Lions, is a longitudinal fold of skin along their bellies. Male Asiatic Lions have only moderate mane growth on top of the head, so that their ears are alwaysvisible.

Habitat. Lions have a wide habitat tolerance. They are not found in dense lowland forests of West Africa and the Congo Basin, but will penetrate deep into desert, especially along avenues of watercourses. They are common in semi-desert areas. They will drink when water is available, but are not dependent on it, getting their moisture requirements from their prey and even plants (tsamma melon in the Kalahari Desert). Optimal habitat appears to be open woodlands and thick bush, scrub, and grass complexes, providing ample supply of food in the form of medium to large-sized ungulates, some shade for resting in the heat of the day, and the barest of cover to facilitate stalking of prey. Range up to 4000 m in Ethiopia. The Lion formerly ranged from North Africa through south-west Asia, where it disappeared from most countries within the last 150 years, west into Europe, where it became extinct almost 2000 years ago, and east into India. Lions survived in the central Saharan Desert on the edge of the Air Mountains until about 60 years ago.

Food and Feeding. Mediumto large-sized ungulates make up the bulk of their diet, but Lions are generalists, taking a large variety of vertebrates from mice to young rhinos, Hippos (Hippopotamus amphibius), and African Elephants (Loxodonta africana), birds up to the size of an ostrich (Struthio camelus), as well as reptiles,fish, and even insects. Major large ungulate species include a large variety of bovids like African Buffalo, Blue Wildebeest, Hartebeest, Waterbuck, Kob ( Kobus kob), Sable Antelope, Gemsbok , eland, Impala, Springbok, as well as Giraffe (Giraffa camelopardalis), warthog, and zebra in Africa, and deer as well as a high percentage oflivestock in Asia. Individual differences in prey selection and killing techniques are discernible in different prides in the same area, indicating that learning plays a strong role in the Lion’s hunting behavior; i.e. hunting of Brown Fur Seals (Arctocephalus pusillus) on the Namibian coast. In the Serengeti 30% of stalks were successful when members of the pride took part, against 17-19% when only a single individual was involved. Lions do not take account of wind direction when stalking prey; however, they will often try to cut prey off by running ahead of it. Speeds of 45-60 km/h may be achieved in a rush for up to a few hundred meters (usually not more than 100-200 m). The attack is delivered to the rump or shoulders, with the weight of the Lions often bringing the prey to the ground. Sometimes the prey’s neck is broken in falling. As soon as the prey is down it is seized by the throat or muzzle to effect strangulation. The Lions feed at the site of the kill or drag it to the nearest cover. The belly is ripped open and the stomach and intestines pulled out. Lions, especially males, frequently scavenge, acquiring up to 53% of food items in the open plains of the Serengeti this way. This figure decreases to 5% in more arid environments, where prey occurs at lower density. However, in the woodlands of Kruger National Park, male Lions acquire most of their food by hunting rather than scavenging. In Kruger the main prey of male Lions was African Buffalo; females fed on the most abundant medium-sized ungulates, such as Blue Wildebeest and Burchell’s Zebra. Lions will watch vultures descending on kills of other predators. Protection of kills against Spotted Hyenas,is greatly enhanced by the presence of a male. Females may lose up to 20% of their kills to Spotted Hyenas and a further 17% to unrelated Lions. In the absence of their normal prey, Lions can cause grave losses to cattle and small stock, or may even become “man-eaters”. The individuals involved are not always old and decrepit.

Activity patterns. Lions usually hunt at night, but there are exceptions. Major hunting peaks usually occur between 02:00 h and 04:00 h, but in Etosha, most hunts occurred from 21:30-22:30 h. In Botswana’s Chobe National Park, Lions seem to be most successful on moonless nights. Hunting levels may be increased during storms, when noise, wind, and waving vegetation make it difficult for prey to detect Lions. They will sleep and rest, usually in compact groups, for long periods when satiated. They avoid exerting themselves during the heat of the day. Although a terrestrial species, they are good climbers and will sometimes drape themselves along branches of trees to take advantage of cool breezes and avoid flies or dangerous animals. Resting periods are interrupted by short periods of intense activity when hunting or exhibiting aggression. Their roar can, under optimal conditions, be heard from as far as 5 km away, appears to serve to demarcate territory, and may be individually distinctive. Prides often indulge in communal roaring. They also use scats, urine spraying, and scrapes for demarcation.

Movements, Home range and Social organization. Mostly sedentary but some populations follow migratory prey seasonally. Some individuals nomadic, especially males and subadults. In the arid savanna of Etosha National Park Lions move a mean 13-2 km in 24 hours; in the S Kalahari an average 15-2 km (range 11-34 km). In contrast, Serengeti Lions averaged 4-5-6-5 km in 24 hours. Pride home range size varies from 25-226 km?, but can be considerably larger in low prey density areas, e.g., Etosha National Park, with up to 2075 km ®. In some areas there is an overlap of home ranges but in others ranges are largely exclusive, and actively defended, meriting the term territories. Whether ranges are defended appears to be dependent on the local movement of prey species. Average emigration/mortality of pride Lions in Etosha National Park was 17%. Core unit of the Lions’ matriarchial society is the pride, which consists of a group of related females, none dominant, and their cubs. Pride size, measured by the number of adult females, varies from a minimum average of 2-2 in very arid environments to an average of 4-6, up to eight in year-round high prey-density areas like the Ngorongoro Crater, Tanzania. Maximum pride size was 39. Subspecies persica has on average smaller prides of 2-5 females. Prides are “fission-fusion” social units. Membership is stable, but pride members are often scattered in small subgroups throughout the pride’s range, and each member spends considerable time alone. Female pride members display several cooperative behaviors unique among felids, giving birth in synchrony, raising cubs communally, with allosuckling of cubs and cooperative hunting. Males, for the short time they are living together with females, concentrate their energy on defending their tenure over the pride(s). In Etosha males only participated in 4% of 461 hunting opportunities, in the Serengeti in only 3%, but more recent studies show that males elsewhere are efficient hunters, tackling especially large and dangerous prey like Buffalo. Prides often divide into smaller sub-groups when foraging. In some regions there is a complex division of labor among lionesses. Some, often larger and heavier individuals, repeatedly play the role of “center”, and others the role of “wing”, chasing the prey towards the “center”. A single male or coalition of up to seven males holds tenure over one or more prides, and effectively excludes strange males from siring cubs with pride females. The average tenure is only 2-3 years. Only in exceptional cases do male Lions breed with related pride females. Coalition males in preand post-tenure periods hunt and scavenge cooperatively. Larger coalitions of 4-6 males can maintain tenure more than twice as long as 1-2 males. Coalitions of more than four males are always related, but pairs frequently consist of unrelated males. Despite maternal defense, infanticide is common when males take over a new pride. Most females with dependent cubs lose their cubs within a month of takeover, and pregnant females lose the cubs shortly after giving birth. In this way, males assure paternity during their short reproductive lifetime. Females show a burst of heightened sexual activity for about three months following a takeover, attracting other males and encouraging competition. This ensures that the fittest coalition is able to gain tenure. Females remain infertile during this “testing” phase, and afterwards, when tenure has stabilized, tend to breed in synchrony. Litters born synchronously have a higher survival rate and tend to show a sex ratio biased towards males. Sociality in Lions may have evolved to increase hunting success on very large prey, defense of young and kills, maintenance of long term territories, and insurance against individual injury or incapacity.

Breeding. Largely aseasonal, but with birth peaks in certain regions. Courtship initiated by either member of pair; pair remain in close association during the mating period. The male follows the female at all times and rests with her. Female invites copulation by lordosis. Other males of the coalition are tolerated close by, and sometimes there are even multiple-mate copulations, but strange lions are driven off. At least one member of male coalitions larger than two fails to breed successfully, but through kin selection, related non-breeding helpers still benefit. Pair may copulate every 15 minutes, with copulation lasting up to one minute. Mating lasts over a period of several hours and also through the night. Between copulations the pair will lie down next to each other or walk together until the next mating. During copulation the female purrs loudly and towards the end the male may gently neckbite the female. The majority of matings do not result in pregnancy; of 14 observed mating periods only four resulted in fertilization. Lions are polyestrous, estrus lasting 4-16 days, interestrus interval varying from a few days to over a year, on average 16 days, gestation mean of 110 days (range 100-114). Litter size averages 2-5-3, range 1-6, but 89% is 1-4. The mean interbirth interval is 20 months (range 11-25) if previous litter survives to maturity, or 4-6 months if previous litter was lost. For parturition females leave their pride for 4-8 weeks. Mass at birth is about 1-5 kg. Females rejoin the pride with their 4-8 week-old cubs; any female with milk will allow suckling. Cubs will suckle regularly for 6-7 months, suckling less after seven months and discontining by twelve months. They remain with their mothers for 21-30 months and grow rapidly for the first three years. Cub mortality is high, with a 40-50% survival rate, depending on food supply and pride takeover. Young Lions learn to hunt by watching the adults in the pride, and leave their natal pride at the age of 24-42 months. Males reach sexual maturity at 26 months, but only get the opportunity to mate when they are about five years old, and usually only during their pride tenure. Female become pregnantfor the first time at about 43 months of age and continue to breed until they are about 15 years old, producing a litter every two years.

Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List. There are no sound estimates of the total number of Lions in Africa. Most recent data range from 16,500 to 30,000. They occur throughout most of Africa, but are increasingly restricted to protected areas. They are of great social-economic value for trophy hunting, game viewing, and negatively as stock-raiders. Man eating Lions are also a problem, especially in Tanzania. Recent increase in reintroduction of Lions into smaller fenced reserves. Hunting restricted to “problem” animals in most places, in some countries prohibited but in others no legal protection. Generally considered problem animals, whose existence is at odds with human settlement and cattle cultures. Scavenging makes them vulnerable to poisoned carcasses. Some populations already isolated, Namibia —Etosha National Park c. 300, Zimbabwe Hwange National Park complex 500. Subspecies persica up from less than 20 in Gir forest around 1900 to about 300 in 1995. Lions are sparsely distributed in West and Central Africa. Densities vary from 0-17 adults/subadults/ 100 km? in Savuti, Chobe National Park, Botswana, to 1-5-2/ 100 km? in Ngalagadi-Transfrontier Park ( Botswana / South Africa) and Etosha National Park, Namibia; to 3-10 and up to 18/ 100 km ® in East and South African protected areas; and 14/ 100 km? in Gir National Park, India. Highest known density is in Maasai Mara National Reserve, Kenya with 30/ 100 km? Translocation of Asian lions to a second protected area outside of Gir National Park, which is at carrying capacity,is planned. Recent canine distemper outbreak in Serengeti-Mara ecosystem killed 33% of an estimated population of 3000 lions. Other threatening diseases include bovine tuberculosis, and FIV. Lions are heavily parasitized by all manner of external and internal parasites.

Bibliography. Barnett et al. (2006), Bauer & Van der Merwe (2004), Berry (1981), Bertram (1975a, 1975b), Bosman & Hall-Martin (1997), Bothma (1998), Bothma & Walker (1999), Bridgeford (1985), Bryden (1978), Caraco & Wolf (1975), Chellam (1987, 1993), Chellam & Jonsingh (1993), Cooper, J. (1942), Cooper, S.M. (1991), Cowie (1966), Eloff (1964, 1973a, 1973b), Fagotto (1985), Funston & Mills (1997), Funston et al. (2001), Grant etal. (2005), Grinnell & Mc-Comb (2001), Guggisberg (1975), Hanby et al. (1995), Hunter (1999), Hunter et al. (2007), Jackson (1995), Jonsingh & Chellam (1991), Joslin (1973), Kingdon (1971-1982), Kruuk & Turner (1967), Makacha & Schaller (1969), McBride (1990), Mills (1990), Mills & Shenk (1992), Mills et al. (1989), Mitchell et al. (1965), Moser (2008), Munson et al. (1995), Nowell & Jackson (1996), Ogutu & Dublin (2002, 2004), Orford et al. (1988), Owens & Owens (1984), Packer & Kock (1995), Packer & Pusey (1983, 1987, 1997), Packer, lkanda et al. (2006), Packer, Scheel & Pusey (1990), Patterson, B.D., Kasiki et al. (2004), Patterson, B.D., Kays et al. (2006), Patterson, G. (1988), Pienaar (1969), Pusey & Packer (1987), Rashid (1991), Rodgers (1974), Rosevear (1974), Rudnai (1973a, 1973b, 1974), Ruggiero (1991), Schaller (1972), Scheel & Packer (1991), Schenkel (1966), Skinner & Smithers (1990), Smuts (1979, 1982), Smuts, Hanks & Whyte (1978), Smuts, Robinson & Whyte (1980), Spong (2002), Stander (1991, 1992a, 1992b, 1997), Van Orsdol (1982, 1984), Verberne & Leyhausen (1976), Walker (1994), West & Packer (2002), Woodroffe & Frank (2005).

Figures

6. Lion (Panthera leo), 7. Jaguar (Panthera onca)

Subspecies and Distribution. Pl. azandica]. A. Allen, 1924 — NE Zaire . P. l. bleyenberghi Lonnberg, 1914 — S Zaire , Zambia , and Angola . P. l. kruger: Roberts, 1929 — NW (Kalahari), N, and SE South Africa . P.l. nubica de Blainville, 1843 — NE and E Africa. P.l. persica Meyer, 1826 — from Iraq to C India in the 19" century; now restricted to the Gir Forest, India . P. l. senegalensis Meyer, 1826 — West Africa E to the Central African Republic .