Amur leopard

The Amur leopard (Panthera pardus orientalis) is a leopard subspecies native to the Primorye region of southeastern Russia and northern China. It is listed as Critically Endangered on the IUCN Red List. In 2007, only 19–26 wild leopards were estimated to survive in southeastern Russia and northeastern China.[1] It was considered as one of the rarest cats on Earth.[3]

Amur leopard
A captive Amur leopard at the Colchester Zoo, England
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. pardus
Subspecies:
P. p. orientalis
Trinomial name
Panthera pardus orientalis
(Schlegel, 1857)
Historic and present distribution of the subspecies (barring northern China to the west of Manchuria)
Synonyms[2]

P. p. japonensis (Gray, 1862)

As of 2015, fewer than 60 individuals were estimated to survive in Russia and China.[4] Camera-trapping surveys conducted between 2014 and 2015 revealed 92 individuals in an 8,398 km2 (3,242 sq mi) large transboundary area along the Russian-Chinese border.[5] In 2019, it was reported that the population is close to 90 leopards.[6]

Results of genetic research indicate that the Amur leopard is genetically close to leopards in northern China and Korea, suggesting that the leopard population in this region became fragmented in the early 20th century.[7] The North Chinese leopard was formerly recognised as a distinct subspecies P. p. japonensis, but was subsumed under the Amur leopard in 2017.[2]

Taxonomic history

In 1857, Hermann Schlegel described a leopard skin from Korea under the scientific name Felis orientalis.[8] Since Schlegel's description, several naturalists and curators of natural history museums described zoological specimens of leopards from the Russian Far East and China:

In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed P. p. japonensis to P. p. orientalis. The remaining are not considered valid subspecies.[2]

Genetic research

Phylogenetic analysis of leopard samples from Primorsky Krai and North Korea revealed that they cannot be distinguished. It is considered probable that the population became fragmented less than a century ago.[7] Phylogenetic analysis of an old leopard skin from South Korea indicates that it belongs to the Amur leopard subspecies.[17]

The complete mitochondrial genome of a wild male leopard specimen from Shaanxi Province in central China has been amplified and is 16,966 base pairs long.[18]

Characteristics
Close-up of a leopard at Ménagerie du Jardin des Plantes

The Amur leopard differs from other leopard subspecies by its thick fur that is pale cream-colored, particularly in winter. Rosettes on the flanks are 5 cm × 5 cm (2.0 in × 2.0 in) and widely spaced, up to 2.5 cm (0.98 in), with thick, unbroken rings and darkened centers.[16] Its fur is fairly soft with long and dense hair. The length of hair on the back is 20–25 mm (0.79–0.98 in) in summer and up to 70 mm (2.8 in) in winter. The winter coat varies from fairly light yellow to dense yellowish-red with a golden tinge or rusty-reddish-yellow. In summer, the fur is brighter with more vivid coloration pattern. It is rather small in body size, with males larger than females. Males measure from 107–136 cm (42–54 in) with an 82–90 cm (32–35 in) long tail, a shoulder height of 64–78 cm (25–31 in), and a weight of 32.2–48 kg (71–106 lb). Females weigh from 25–42.5 kg (55–94 lb).[19]

The North Chinese leopard was first described on the basis of a single tanned skin which was fulvous above, and pale beneath, with large, roundish, oblong black spots on the back and limbs, and small black spots on the head. The spots on the back, shoulders and sides formed a ring around a central fulvous spot. The black spots on the nape were elongated, and large ones on the chest formed a necklace. The tail was spotted and had four black rings at the tip.[9]

Distribution and habitat

In the Russian Far East, the Amur leopard inhabits an area of about 7,000 km2 (2,700 sq mi) today.[20] It is well adapted to a cold climate and snow.[7] Leopards cross between Russia, China, and North Korea across the Tumen River despite a high and long wire fence marking the boundary.[21]

The first camera trap image of an Amur leopard in northeastern China was taken in 2010 in Hunchun National Nature Reserve located in the Changbai Mountains in Jilin and Heilongjiang Provinces.[22] This habitat consists of broadleaved conifer and Korean pine forests at altitudes of 600–1,200 m (2,000–3,900 ft), where annual average temperature is about 1.5 °C (34.7 °F).[23] In this area, leopards were repeatedly photographed by camera traps set up between January 2013 and July 2014 covering up to 4,858 km2 (1,876 sq mi).[24][25][26]

Elsewhere in China, leopard range is fragmented with small populations occurring foremost in isolated reserves. In Shanxi Province, leopards were recorded in 16 protected areas during camera trapping surveys between 2007 and 2014. In Shaanxi Province, leopards were recorded in six nature reserves, including Foping National Nature Reserve.[27]

Historical range

Leopard fossils from the Pleistocene have been excavated in Japan, but the species has not been identified with certainty.[28]

Historic records from before 1930 indicate that the Amur leopard occurred in Eastern Siberia, in Northern China near Beijing and in the mountains to the northwest, and in Korea.[10][29] Its range once extended throughout Manchuria in northeastern China, including Jilin and Heilongjiang provinces, and throughout the Korean Peninsula. In Russia, its range was dramatically reduced during the 1970s to about 20% of its former range. The northern boundary of its occurrence commenced on the coast of the Sea of Japan at 44°N and ran south at a distance of 15–30 km (9.3–18.6 mi) from the coast to 43°10'N. There its range turned steeply westward, north of the Suchan River basin, then north to encompass the source of the Ussuri River and two right bank tributaries in the upper reaches of the Ussuri, and westward toward the bank of Khanka Lake. In the 1950s, leopards were observed 50 km (31 mi) north of Vladivostok and in Kedrovaya Pad Nature Reserve. The association of the leopard with mountains is fairly definite, and to snow-free south-facing rocky slopes in winter. The species is confined more to places where wild sika deer live or where deer husbandry is practiced.[19]

Leopards were extirpated on the Korean Peninsula during the period of Korea under Japanese rule.[30] At least 624 leopards were killed during the Japanese occupation between 1910 and 1945. In South Korea, the last known leopard was captured in 1970.[17] The Amur leopard is considered locally extinct.[1]

In China, Amur leopards occurred in the Lesser Khingan, Changbai Mountains and Wanda Mountains until the 1970s. In the following decades, the range decreased to a few areas in Jilin and Heilongjiang Provinces.[31] Today, only small and isolated populations remain in China.[18]

Ecology and behavior
Skeleton at the Museum of Osteology

Amur leopards are solitary, unless females have offspring.[19] Records from camera-traps indicate that they are more active during the day than at night and during twilight, both in the summer and winter seasons. This activity pattern coincides with activity of prey species such as Siberian roe deer, sika deer and wild boar.[26]

They are extremely conservative in their choice of territory. An individual's territory is usually located in a river basin which generally extends to the natural topographical borders of the area. The territory of two individuals overlaps sometimes, but only slightly. Depending on sex, age and family size, the size of an individual's territory varies from 5,000–30,000 ha (19–116 sq mi). They use the same hunting trails, migration routes and even rest places over the course of many years.[32] Leopards are resident at places where wild animals are abundant, and follow herds of ungulates. In the Ussuri region the main prey of leopards are Siberian roe deer (Capreolus pygargus) and Manchurian sika deer (Cervus nippon mantchuricus), Manchurian wapiti (Cervus canadensis xanthopygus), Siberian musk deer (Moschus moschiferus), Amur moose (Alces alces cameloides) and Ussuri wild boar (Sus scrofa ussuricus). They also catch hare (Lepus), Eurasian badger (Meles meles), fowl, and mice. In Kedrovaya Pad Nature Reserve roe deer is their main prey year-round, but they also prey on young Asian black bear if they are less than two years old.[19] When density of ungulates is low, leopards have large home ranges of up to 100 km2 (39 sq mi).[33]

During a study of radio-collared Amur leopards in the early 1990s, a territorial dispute between two males at a deer farm was documented, suggesting that Amur leopards favour such farms for hunting.[34] Female leopards with cubs are often found in the proximity of deer farms. The large number of domesticated deer is a reliable food source in difficult times.[35] In 2011, an adult Amur leopard female was radio-collared in the vicinity of the Land of the Leopard National Park in the Khasansky region of Primorskii Krai. During three years of tracking, she used a home range of 161.7 km2 (62.4 sq mi) with a core area of 23.3 km2 (9.0 sq mi). During estrus, she moved in a core area of 52.9 km2 (20.4 sq mi). After giving birth in late June, she reduced her movements to an area of about 3 km2 (1.2 sq mi) for a month, in which she shifted her cubs three times. From autumn onwards, she gradually increased her home range. When the cubs were more than one year old, the family moved together in the initial home range of 161.7 km2 (62.4 sq mi).[36]

Amur leopards share habitat with Siberian tigers, but in the Changbai Mountains have been recorded more often in higher altitudes and far from settlements and roads than tigers.[26]

Reproduction
Cub in Cottbus Zoo

Amur leopards become sexually mature at the age of 2–3 years. They are able to reproduce up to 10–15 years of age. Estrus lasts 12–18 days, and in exceptional cases up to 25 days. Gestation lasts 90–105 days, and usually between 92 and 95 days. A newborn cub weighs 500–700 g (1.1–1.5 lb). The young open their eyes on the 7th–10th day and begin to crawl on the 12th–15th day. By the second month they emerge from their dens and also begin to eat meat. Cubs are weaned when three months old, and then learn to hunt. Lactation continues for five or six months. Cubs reach independence at the approximate age of two to three years. They stay with their mother until they are around 18 months to two years old.[36] Juveniles sometimes stay with their mother until she comes into estrus again. Until the 1970s, cubs were seen in Kedrovaya Pad Nature Reserve and in northeastern China most often between the end of March and May. Litters comprised two to three cubs. In captivity some individuals have lived for 21 years.[19]

During a population census in 1997, four females found with young had only one cub each. Results of radio telemetry studies confirmed that young stay with their mother for two years. In Kedrovaya Pad Nature Reserve, the young of two different litters were observed with their mothers at the same time.[32]

Threats

The Amur leopard is threatened by poaching, poaching of prey species, habitat loss and deforestation or exploitation of forests. Its natural habitat is threatened by forest fires and construction of new roads.[1] Due to the small number of reproducing Amur leopards in the wild, the gene pool is so reduced that the population is at risk from inbreeding depression.[37] In 2015, a wild Amur leopard was found with canine distemper virus in Primorskii Krai. The small population is possibly exposed to domestic or wild disease carriers and transmitters.[38]

Tigers can eliminate leopards if densities of large and medium-sized prey species are low. Competition between these predators supposedly decreases in summer, when small prey species are more available. In winter, conditions are less favorable for tigers and the extent of trophic niche overlap with that of leopards probably reaches its peak.[33]

Poaching

Poaching of leopards is a main threat to their survival. There are rumours but no evidence that Chinese traders buy leopard skins; no skins were confiscated at borders to China. In 14 months from February 2002 to April 2003, seven skins or part of skins were confiscated, six in Russia and one in China. Leopards are most often killed by local Russians from small villages in and around the leopard habitat. Most of these villagers hunt entirely illegally; they have no licenses for hunting nor for their guns, and they are not members of one of the local hunting leases.[35]

In 1999, skins of poached leopards were offered for $500–1,000 near a protected area in Russia.[39]

Forest degradation

Human induced fires are a main threat to the survival of the leopard. Setting fire to fields is a habit of rural farmers who start them for a particular purpose such as improving fertility for livestock grazing, killing ticks and other insects, making scrap metals visible so that they can be easily collected, culling vegetation along train tracks, and stimulating fern growth. Young ferns are sold in shops, served in restaurants and also exported to China as a popular dish. Surveys using satellite images and GIS techniques revealed that on average 19% of south-west Primorye burns annually, and a total of 46% burned at least once in six years. Due to a long and frequent fire history, much of the land in south-west Primorye has been converted to permanent grasslands. These frequent fires cause degradation of suitable leopard habitat into unsuitable habitat. Repeated fires have created open "savannah" landscapes with grass, oak bushes and isolated trees that leopards seem to avoid, again probably because of low ungulate densities.[35]

Large deer farms extended over thousands of hectares in leopard habitat. Deer farming used to be a large-scale business; the velvet of deer antlers was sold to Asian pharmacies.[34] The number of deer farms has decreased considerably since the late 1990s.[35]

Development projects

A number of plans for economic activities in south-west Primorye were developed that posed a serious threat to the leopard's survival. A plan to build an oil pipeline from central Siberia through Primorye to the coast of the Sea of Japan has been shelved. A plan for an open pit coal mine in the heart of the leopard range was not carried out following pressure from environmentalists and the Ministry of Natural Resources. The strategic location of south-west Primorye, close to the main population centres of Primorsky Krai, the Japanese Sea and the borders of Korea and China, makes it more attractive for economic activities including transport, industries, tourism and development of infrastructure. Logging is not a major threat; the use of the road network established for the transport of logs from forests increases anthropogenic pressures in unprotected leopard habitat.[35]

Inbreeding

An acute problem is potential inbreeding. The remaining population could disappear as a result of genetic degeneration, even without direct human influence. The levels of diversity are remarkably low, indicative of a history of inbreeding in the population for several generations. Such levels of genetic reduction have been associated with severe reproductive and congenital abnormalities that impede the health, survival and reproduction of some but not all genetically diminished small populations. Cub survival has been declining from 1.9 cubs per one female in 1973 to 1.7 in 1984 and 1.0 in 1991. Besides a decline in natural replacement, there is a high probability of mortality for all age groups as a result of certain diseases or direct human impact.[40]

Results of genetic analyses imply that the Amur leopard population lost genetic diversity over a short period of time.[7]

Conservation

The Amur leopard is listed in CITES Appendix I. It is stated to need better protection from illegal trade in skins and bones.[1]

In 2001, a meeting was held in Vladivostok with the aim of devising and planning management recommendations and activities needed to ensure the recovery and continued survival of the wild Amur leopard population in range countries. Chinese participants announced the creation of a new protected area in Jilin Province, the Hunchun Nature Reserve.[41] Since 2014, Russian and Chinese biologists collaborate in transboundary monitoring of the Amur leopard population.[5]

The Amur Leopard and Tiger Alliance (ALTA) is an initiative of Russian and western conservation organisations to conserve the Amur leopard and tiger, and secure a future for both species in the Russian Far East and Northeast China. ALTA operates across Northeast Asia under the guiding principle that only co-operative, co-ordinated conservation actions from all interested parties can save these endangered species from extinction. ALTA works in close co-operation with local, regional, and federal government and non-government organisations to protect the region's biological wealth through conservation, sustainable development, and local community involvement. The Phoenix Fund and the Wildlife Conservation Society provide a local framework for implementing ALTA projects, working closely with many Russian and Chinese agencies. With regard to conservation of leopards, ALTA aims at retaining a leopard population of 35 adult females (100 total) in south-west Primorye and the Jilin-Heilongjiang border region; and creating a second population of 20 adult females (60 total) in the former range of the leopard. Conservation projects for the leopard include:[35]

  • four antipoaching teams with a total of 15 members in the leopard's range
  • a special task force of local police and anti-poaching teams led by the Khasan prosecutor
  • monitoring of the leopard population through snow track counts and camera trap counts
  • monitoring and analysis of the impact of fires on leopard habitat and the effectiveness of firefighting
  • habitat assessment with geographic information system (GIS) techniques: assessment of the role of habitat quality, land ownership, land use, protection status, settlements, deer farms, roads, and human settlements with use of monitoring data and satellite images
  • development of land-use plans that take in account future needs of leopards
  • support for protected areas in the leopard range
  • compensation of livestock kills by leopards and tigers
  • a comprehensive education program for school children and students in the leopard range
  • support for hunting leases, and an ungulate recovery program
  • media campaign to create awareness about the leopard's plight
  • support and technical assistance for the new Hunchun Nature Reserve in China that borders on the leopard range in Russia

An oil pipeline planned to be built through leopard habitat was rerouted, following a campaign by conservationists.[42]

Reintroduction into the wild

Since 1996, the idea of reintroducing leopards in the south of Sikhote-Alin has been discussed by ALTA members.[43] During a workshop in 2001, the outlines and principles of a plan for the development of a second population of the leopard in the Russian Far East was prepared. For reintroduction to be successful, one fundamental question needs to be answered: Why did leopards disappear from the southern Sikhote-Alin in the middle of the 20th century? It was recommended to assess reasons for localized extinctions, obtain support of local people, increase prey in areas proposed for reintroduction, ensure that conditions exist conducive for reintroduction in the selected area, and ensure survival of the existing population. There are two sources of leopards for reintroduction: leopards born and raised in zoos and leopards raised in a special reintroduction center passed through a rehabilitation program for life in the wild.[40]

If this reintroduction is to succeed, it is clear that the design of the breeding and release centre, and the management of the leopards in it, must focus strongly on overcoming the difficulties imposed by the captive origin of the cats. Three necessary behaviours should be acquired prior to release: hunting and killing of live natural prey; avoidance of humans and avoidance of tigers.[44]

In March 2009, the Minister of Natural Resources of Russia during his meeting with Vladimir Putin reassured that the ministry is planning to introduce new "imported" leopards into the area and creating suitable and safe habitat for them. The government already allocated all required funds for the project.[45]

Contiguous patches of potential leopard habitat as potential reintroduction sites were identified in the southern Sikhote-Alin. Three coastal potential habitat patches could harbour approximately 72 adult leopards.[20]

In captivity
In Tierpark Hellabrunn
At Audubon Zoo

A captive breeding programme for the Amur leopard was established in 1961 from nine wild-born founders.[46] A molecular genetic survey revealed that at least two founders of the captive pedigree had genetic information that is not consistent with any wildborn Amur leopard.[37] Both the American and European zoo populations include contribution of genes from a male founder that was not an Amur leopard. It has been the strategy of the European Endangered Species Programme to minimize his contribution and maintain genetic diversity of the captive population.[44]

As of December 2011, 173 captive Amur leopards were held in zoos worldwide. Within the EESP, 54 male, 40 female and 7 unsexed individuals are kept. In American and Canadian zoos, another 31 males and 41 females are kept within the Population Management Program.[47]

Naming and etymology

The names 'Amurland leopard' and 'Amur leopard' were coined by Pocock in 1930, when he compared leopard specimens in the collection of the Natural History Museum, London. In particular, he referred to a leopard skin from the Amur Bay as 'Amur leopard'.[16] Since at least 1985, this name has been used for the leopard population in Eastern Siberia and for the captive population in zoos worldwide.[48][46]

The Amur leopard is also known as the "Siberian leopard",[49] "Far Eastern leopard",[50][37][43] and "Korean leopard".[17]

In culture
Further information: Tigers in Korean culture
Amur leopard and Korean magpie depicted in Korean folk painting from the late Joseon Dynasty

The Amur leopard were depicted in many Korean folk paintings in Joseon Dynasty.

In media

The Animal Planet documentary The Last Leopard (2008) is about the plight of Amur leopards in Russia. The television series "Wild Russia" showed a glimpse into the life of leopards. A female leopard and her cub were featured on Planet Earth episodes "Seasonal Forests".[3]

See also

Leopard subspecies: African leopard  · Arabian leopard  · Anatolian leopard  · Persian leopard  · Indian leopard  · Indochinese leopard  · Javan leopard  · Sri Lankan leopard  · Panthera pardus spelaea

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