Thered panda (Ailurus fulgens), also known as thelesser panda, is a smallmammal native to theeastern Himalayas andsouthwestern China. It has dense reddish-brown fur with a black belly and legs, white-lined ears, a mostly white muzzle and a ringed tail. Its head-to-body length is 51–63.5 cm (20.1–25.0 in) with a 28–48.5 cm (11.0–19.1 in) tail, and it weighs between 3.2 and 15 kg (7.1 and 33.1 lb). It is well adapted to climbing due to its flexible joints and curved semi-retractile claws.
The red panda was formallydescribed in 1825. The two recognised subspecies, the Himalayan and the Chinese red panda,genetically diverged about 250,000 years ago. The red panda's place on theevolutionary tree has been debated, but modern genetic evidence places it in close affinity withraccoons,weasels, andskunks. It is not closely related to thegiant panda, which is abear, though both possess elongated wrist bones or "false thumbs" used for graspingbamboo. The evolutionarylineage of the red panda (Ailuridae) stretches back around25 to 18 million years ago, as indicated by extinct fossil relatives found in Eurasia and North America.
The origin of the namepanda is uncertain, but one of the most likely theories is that it derived from theNepali word "ponya".[4] The wordपञ्जाpajā orपौँजाpañjā means "ball of the foot" and "claws".[5] The Nepali words "nigalya ponya" has been translated as "bamboo-footed" and is thought to be the red panda's Nepali name; in English, it was simply calledpanda, and was the only animal known under this name for more than 40 years; it became known as thered panda orlesser panda to distinguish it from thegiant panda, which was formally described and named in 1869.[4]
The modern red panda is theonly recognised species in the genusAilurus. It is traditionally divided into twosubspecies: the Himalayan red panda (A. f. fulgens) and the Chinese red panda (A. f. styani). The Himalayan subspecies has a straighter profile, a lighter coloured forehead andochre-tipped hairs on the lower back and rump. The Chinese subspecies has a more curved forehead and sloping snout, a darker coat with a less white face and more contrast between the tail rings.[3]
In 2020, results of a genetic analysis of red panda samples showed that the red panda populations in the Himalayas and China were separated about 250,000 years ago. The researchers suggested that the two subspecies should be treated as distinct species. Red pandas in southeastern Tibet and northern Myanmar were found to be part ofstyani, while those of southern Tibet were offulgens in the strict sense.[12]DNA sequencing of 132 red panda faecal samples collected inNortheast India and China also showed two distinct clusters indicating that theSiang River constitutes the boundary between the Himalayan and Chinese red pandas.[13] They probably diverged due toglaciation events on the southernTibetan Plateau in thePleistocene.[14]
Phylogeny
The placement of the red panda on theevolutionary tree has been debated. In the early 20th century, various scientists placed it in thefamilyProcyonidae withraccoons and their allies. At the time, most prominent biologists also considered the red panda to be related to the giant panda, which would eventually be found to be abear. A 1982 study examined the similarities and differences in the skull between the red panda and the giant panda, other bears and procyonids, and placed the species in its own familyAiluridae. The author of the study considered the red panda to be more closely related to bears.[3] A 1995mitochondrial DNA analysis revealed that the red panda has close affinities with procyonids.[15] Further genetic studies in 2005, 2018 and 2021 have placed the red panda within thecladeMusteloidea, which also includes Procyonidae,Mustelidae (weasels and relatives) andMephitidae (skunks and relatives).[16][17][18]
Phylogenetic relationship of the red panda based on analysis of
Reconstructed skull and head ofSimocyon, a large carnivorous early relative of the modern red panda
The family Ailuridae appears to have evolved in Europe in either theLate Oligocene orEarly Miocene, about25 to 18 million years ago. The earliest memberAmphictis is known from its 10 cm (4 in) skull and may have been around the same size as the modern species. Itsdentition consists of sharp premolars andcarnassials (P4 and m1) andmolars adapted for grinding (M1, M2 and m2), suggesting that it had a generalisedcarnivorous diet. Its placement within Ailuridae is based on the grooves on the side of itscanine teeth. Other early orbasal ailurids includeAlopecocyon andSimocyon, whose fossils have been found throughout Eurasia and North America dating from theMiddle Miocene, the latter of which survived into theEarly Pliocene. Both have similar teeth toAmphictis and thus had a similar diet.[19] Thepuma-sizedSimocyon was likely a tree-climber and shared a "false thumb"—an extended wrist bone—with the modern species, suggesting the appendage was an adaptation toarboreal locomotion and not to feed onbamboo.[19][20]
Later and moreadvanced ailurids are classified in the subfamily Ailurinae and are known as the "true" red pandas. These animals were smaller and more adapted for anomnivorous orherbivorous diet. The earliest known true panda isMagerictis from the Middle Miocene of Spain and known only from one tooth, a lower second molar. The tooth shows both ancestral and new characteristics having a relatively low and simplecrown but also a lengthened crushing surface with developedtooth cusps like later species.[21] Later ailurines includePristinailurus bristoli which lived in eastern North America from the late Miocene to the Early Pliocene[21][22] and species of the genusParailurus which first appear in Early Pliocene Europe, spreading across Eurasia into North America.[21][23] These animals are classified as asister taxon to the lineage of the modern red panda. In contrast to the herbivorous modern species, these ancient pandas were likely omnivores, with highly cusped molars and sharppremolars.[21][22][24]
The earliest fossil record of the modern genusAilurus dates no earlier than thePleistocene and appears to have been limited to Asia. The modern red panda's lineage became adapted for a specialised bamboo diet, having molar-like premolars and more elevated cusps.[21] The false thumb would secondarily gain a function in feeding.[19][20]
Genomics
Analysis of 53 red panda samples from Sichuan and Yunnan showed a high level ofgenetic diversity.[25] The fullgenome of the red panda was sequenced in 2017. Researchers have compared it to the genome of the giant panda to learn the genetics ofconvergent evolution, as both species have false thumbs and are adapted for a specialised bamboo diet despite having the digestive system of a carnivore. Both pandas show modifications to certain limb development genes (DYNC2H1 andPCNT), which may play roles in the development of the thumbs.[26] In switching from a carnivorous to a herbivorous diet, both species have reactivated taste receptor genes used for detectingbitterness, though the specific genes are different.[27]
Description
Red panda skull
Red panda face
The red panda'scoat is mainly red or orange-brown with a black belly and legs. The muzzle, cheeks, brows and inner ear margins are mostly white while the bushy tail has red andbuff ring patterns and a dark brown tip.[28][29][30] The colouration appears to serve ascamouflage in habitat with redmoss and whitelichen-covered trees. Theguard hairs are longer and rougher while the denseundercoat is fluffier with shorter hairs.[29] The guard hairs on the back have a circular cross-section and are 47–56 mm (1.9–2.2 in) long. It has moderately longwhiskers around the mouth, lower jaw and chin. The hair on the soles of the paws allows the animal to walk in snow.[28]
The red panda has a relatively small head, though proportionally larger than in similarly sized raccoons, with a reduced snout and triangular ears, and nearly evenly lengthed limbs.[28][29] It has a head-body length of 51–63.5 cm (20.1–25.0 in) with a 28–48.5 cm (11.0–19.1 in) tail. The Himalayan red panda is recorded to weigh 3.2–9.4 kg (7.1–20.7 lb), while the Chinese red panda weighs 4–15 kg (8.8–33.1 lb) for females and 4.2–13.4 kg (9.3–29.5 lb) for males.[28] It has five curveddigits on each foot, each with curved semi-retractile claws that aid in climbing.[29] Thepelvis and hindlimbs have flexible joints, adaptations for an arboreal quadrupedal lifestyle.[31] While notprehensile, the tail helps the animal balance while climbing.[29]
The forepaws possess a "false thumb", which is an extension of a wrist bone, the radialsesamoid found in many carnivorans. This thumb allows the animal to grip onto bamboo stalks and both the digits and wrist bones are highly flexible. The red panda shares this feature with the giant panda, which has a larger sesamoid that is more compressed at the sides. In addition, the red panda's sesamoid has a more sunken tip while the giant panda's curves in the middle. These features give the giant panda more developed dexterity.[32]
The red panda's skull is wide, and its lower jaw is robust.[28][29] However, because it eats leaves and stems, which are not as tough, it has smaller chewing muscles than the giant panda. The digestive system of the red panda is only 4.2 times its body length, with a simple stomach, no noticeable divide between theileum andcolon, and nocaecum.[28]
Both sexes have pairedanal glands that emit a secretion consisting of long-chainfatty acids,cholesterol,squalene and2-Piperidinone; the latter is the most odoriferous compound and is perceived by humans as having an ammoniacal or pepper-like odour.[33]
The red panda inhabitsNepal, the states ofSikkim,West Bengal andArunachal Pradesh in India,Bhutan, southernTibet, northernMyanmar and China'sSichuan andYunnan provinces.[1] The global potential habitat of the red panda has been estimated to comprise 47,100 km2 (18,200 sq mi) at most; this habitat is located in thetemperate climate zone of theHimalayas with a mean annual temperature range of 18–24 °C (64–75 °F).[34] Throughout this range, it has been recorded at elevations of 2,000–4,300 m (6,600–14,100 ft).[35][36][37][38][39]
The red panda prefersmicrohabitats within 70–240 m (230–790 ft) of water sources.[50][51][52][53] Fallen logs andtree stumps are important habitat features, as they facilitate access to bamboo leaves.[54] Red pandas have been recorded to use steep slopes of more than 20° and stumps exceeding a diameter of 30 cm (12 in).[50][52] Red pandas observed inPhrumsengla National Park used foremost easterly and southerly slopes with a mean slope of 34° and acanopy cover of 66 per cent that were overgrown with bamboo about 23 m (75 ft) in height.[51] InDafengding Nature Reserve, it prefers steep south-facing slopes in winter and inhabits forests with bamboo 1.5–2.5 m (4 ft 11 in – 8 ft 2 in) tall.[55] InGaoligongshan National Nature Reserve, it inhabits mixed coniferous forest with a dense canopy cover of more than 75 per cent, steep slopes and a density of at least 70 bamboo plants/m2 (6.5 bamboo plants/sq ft).[56] In some parts of China, the red panda coexists with the giant panda. InFengtongzhai andYele National Nature Reserves, red panda microhabitat is characterised by steep slopes with lots of bamboostems, shrubs, fallen logs and stumps, whereas the giant panda prefers gentler slopes with taller but lesser amounts of bamboo and less habitat features overall. Suchniche separation lessens competition between the two bamboo-eating species.[50][54]
Behaviour and ecology
Red panda sleeping in a tree
The red panda is difficult to observe in the wild,[57] and most studies on its behaviour have taken place in captivity.[58] The red panda appears to be bothnocturnal andcrepuscular,sleeping intermittently at night. It typically rests or sleeps in trees or other elevated spaces, stretched out prone on a branch with legs dangling when it is hot, and curled up with its hindlimb over the face when it is cold. It is adapted for climbing and descends to the ground head-first with the hindfeet holding on to the middle of the tree trunk. It moves quickly on the ground by trotting or bounding.[29]
Social spacing
Adult pandas are generallysolitary andterritorial. Individuals mark theirhome range or territorial boundaries with urine,faeces and secretions from theanal and surrounding glands. Scent-marking is usually done on the ground, with males marking more often and for longer periods.[29] In China'sWolong National Nature Reserve, the home range of a radio-collared female was 0.94 km2 (0.36 sq mi), while that of a male was 1.11 km2 (0.43 sq mi).[59] A one-year-long monitoring study of ten red pandas in eastern Nepal showed that the four males had median home ranges of 1.73 km2 (0.67 sq mi) and the six females of 0.94 km2 (0.36 sq mi) within a forest cover of at least 19.2 ha (47 acres). The females travelled 419–841 m (1,375–2,759 ft) per day and the males 660–1,473 m (2,165–4,833 ft). In the mating season from January to March, adults travelled a mean of 795 m (2,608 ft) and subadults a mean of 861 m (2,825 ft).[41] They all had larger home ranges in areas with low forest cover and reduced their activity in areas that were disturbed by people, livestock and dogs.[60]
Diet and feeding
The red panda is largely herbivorous and feeds primarily on bamboo, mainly the generaPhyllostachys,Sinarundinaria,Thamnocalamus andChimonobambusa.[61] It also feeds on fruits, blossoms,acorns, eggs, birds and small mammals. Bamboo leaves may be the most abundant food item year-round and the only food they can access during winter.[62] In Wolong National Nature Reserve, leaves of the bamboo speciesBashania fangiana were found in nearly 94 per cent of analysed droppings, and itsshoots were found in 59 per cent of the droppings found in June.[59]
The red panda grabs food with one of its front paws and usually eats sitting down or standing. When foraging for bamboo, it grabs the plant by the stem and pulls it down towards its jaws. It bites the leaves with the side of thecheek teeth and then shears, chews and swallows. Smaller food like blossoms, berries and small leaves are eaten differently, being clipped by the incisors.[29] Having thegastrointestinal tract of a carnivore, the red panda cannot properly digest bamboo, which passes through its gut in two to four hours. Hence, it must consume large amounts of the most nutritious plant matter. It eats over 1.5 kg (3 lb 5 oz) of fresh leaves or 4 kg (9 lb) of fresh shoots in a day with crude proteins and fats being the most easily digested. Digestion is highest in summer and fall but lowest in winter, and is easier for shoots than leaves.[68] The red panda'smetabolic rate is comparable to other mammals of its size, despite its poor diet.[69] The red panda digests almost a third ofdry matter, which is more efficient than the giant panda digesting 17 per cent.[68] Microbes in the gut may aid in its processing of bamboo; themicrobiota community in the red panda is less diverse than in other mammals.[70]
Communication
Sounds of red panda twittering
At least seven different vocalisations have been recorded from the red panda, comprising growls, barks, squeals, hoots, bleats, grunts and twitters. Growling, barking, grunting and squealing are produced during fights and aggressive chasing. Hooting is made in response to being approached by another individual. Bleating is associated with scent-marking and sniffing. Males may bleat during mating, while females twitter.[71] During bothplay fighting and aggressive fighting, individuals curve their backs and tails while slowly moving their heads up and down. They then turn their heads while jaw-clapping, move their heads laterally and lift a forepaw to strike. They stand on their hind legs, raise the forelimbs above the head and then pounce. Two red pandas may "stare" at each other from a distance.[29]
Reproduction and parenting
Two cubs perched in a tree in Langtang National Park, NepalRed panda tending its cub
Red pandas arelong-day breeders, reproducing after thewinter solstice as daylight grows longer. Mating thus takes place from January to March, with births occurring from May to August. Reproduction is delayed by six months for captive pandas in the southern hemisphere.Oestrous lasts a day, and females can enter oestrous multiple times a season, but it is not known how long the intervals between each cycle last.[72]
As the reproductive season begins, males and females interact more, and will rest, move, and feed near each other. An oestrous female will spend more time marking and males will inspect her anogenital region. Receptive females make tail-flicks and position themselves in alordosis pose, with the front lowered to the ground and the spine curved.Copulation involves the male mounting the female from behind and on top, though face-to-face matings as well as belly-to-back matings while lying on the sides also occur. The male will grab the female by the sides with his front paws instead of biting her neck. Intromission is 2–25 minutes long, and the couplegroom each other between each bout.[72]
Gestation lasts about 131 days.[73] Prior to giving birth, the female selects a denning site, such as a tree, log or stump hollow or rock crevice, and builds a nest using material from nearby, such as twigs, sticks, branches, bark bits, leaves, grass and moss.[57]Litters typically consist of one to four cubs that are born fully furred but blind. They are entirely dependent on their mother for the first three to four months until they first leave the nest. They nurse for their first five months.[73] The bond between mother and offspring lasts until the next mating season. Cubs are fully grown at around 12 months and at around 18 months they reachsexual maturity.[29] Two radio-collared cubs in eastern Nepal separated from their mothers at the age of 7–8 months and left their birth areas three weeks later. They reached new home ranges within 26–42 days and became residents after exploring them for 42–44 days.[41]
The red panda is primarily threatened by thedestruction andfragmentation of its habitat, the causes of which include increasing human population,deforestation, the unlawful taking ofnon-wood forest material and disturbances by herders andlivestock.[1] Trampling by livestock inhibits bamboo growth,[74] andclearcutting decreases the ability of some bamboo species to regenerate.[83] The cut lumber stock in Sichuan alone reached 2,661,000 m3 (94,000,000 cu ft) in 1958–1960, and around 3,597.9 km2 (1,389.2 sq mi) of red panda habitat were logged between the mid-1970s and late 1990s.[48] Throughout Nepal, the red panda habitat outside protected areas is negatively affected by solid waste, livestock trails and herding stations, and people collecting firewood andmedicinal plants.[43][84] Threats identified in Nepal'sLamjung District include grazing by livestock during seasonaltranshumance, human-made forest fires and the collection of bamboo as cattlefodder in winter.[85] Vehicular traffic is a significant barrier to red panda movement between habitat patches.[60]
Poaching is also a major threat.[1] In Nepal, 121 red panda skins wereconfiscated between 2008 and 2018.Traps meant for other wildlife have been recorded killing red pandas.[86] In Myanmar, the red panda is threatened by hunting using guns and traps; since roads to the border with China were built starting in the early 2000s, red panda skins and live animals have been traded and smuggled across the border.[39] In southwestern China, the red panda is hunted for its fur, especially for the highly valued bushy tails, from which hats are produced. The red panda population in China has been reported to have decreased by 40 per cent over the last 50 years, and the population in western Himalayan areas are considered to be smaller.[48] Between 2005 and 2017, 35 live and seven dead red pandas were confiscated in Sichuan, and several traders were sentenced to 3–12 years of imprisonment. A month-long survey of 65 shops in nine Chinese counties in the spring of 2017 revealed only one in Yunnan offered hats made of red panda skins, and red panda tails were offered in an online forum.[87]
Conservation
The red panda is listed inCITES Appendix I and protected in all range countries; hunting is illegal. It has been listed asEndangered on theIUCN Red List since 2008 because the global population is estimated at 10,000 individuals, with a decreasing population trend. A large extent of its habitat is part of protected areas.[1]
A red pandaanti-poaching unit and community-based monitoring have been established in Langtang National Park. Members of Community Forest User Groups also protect and monitor red panda habitats in other parts of Nepal.[91] Community outreach programs have been initiated in eastern Nepal using information boards, radio broadcasting and the annual International Red Panda Day in September; several schools endorsed a red panda conservation manual as part of theircurricula.[92]
Since 2010,community-based conservation programmes have been initiated in 10 districts in Nepal that aim to help villagers reduce their dependence on natural resources through improved herding and food processing practices and alternative income possibilities. The Nepali government ratified a five-year Red Panda Conservation Action Plan in 2019.[93] From 2016 to 2019, 35 ha (86 acres) of high-elevationrangeland inMerak, Bhutan, was restored and fenced in cooperation with 120 herder families to protect the red panda forest habitat and improve communal land.[94] Villagers in Arunachal Pradesh established twocommunity conservation areas to protect the red panda habitat from disturbance and exploitation of forest resources.[46] China has initiated several projects to protect its environment and wildlife, includingGrain for Green, The Natural Forest Protection Project and the National Wildlife/Natural Reserve Construction Project. For the last project, the red panda is not listed as a key species for protection but may benefit from the protection of the giant panda andgolden snub-nosed monkey, with which it overlaps in range.[95]
TheLondon Zoo received two red pandas in 1869 and 1876, the first of which was caught inDarjeeling. TheCalcutta Zoo received a live red panda in 1877, thePhiladelphia Zoo in 1906, andArtis andCologne Zoos in 1908. In 1908, the first captive red panda cubs were born in an Indian zoo. In 1940, theSan Diego Zoo imported four red pandas from India that had been caught in Nepal; their first litter was born in 1941. Cubs that were born later were sent to other zoos; by 1969, about 250 red pandas had been exhibited in zoos.[96] TheTaronga Conservation Society started keeping red pandas in 1977.[97]
In 1978, abreed registry, theInternational Red Panda Studbook, was set up, followed by the Red PandaEuropean Endangered Species Programme in 1985. Members of international zoos ratified a global master plan for thecaptive breeding of the red panda in 1993. By late 2015, 219 red pandas lived in 42zoos in Japan.[98] ThePadmaja Naidu Himalayan Zoological Park participates in the Red PandaSpecies Survival Plan and kept about 25 red pandas by 2016.[99] By the end of 2019, 182 European zoos kept 407 red pandas.[100] Regional captive breeding programmes have also been established in North American, Australasian and South African zoos.[4]
Cultural significance
Red panda on a 2009 stamp from India
The red panda's role in the culture and folklore of local people is limited. A drawing of a red panda exists on a 13th-century Chinese scroll.[101] In Nepal'sTaplejung District, red panda claws are used for treatingepilepsy; its skin is used in rituals for treating sick people, making hats,scarecrows and decorating houses.[86] In western Nepal,Magarshamans use its skin and fur in their ritual dresses and believe that it protects against evil spirits. People in central Bhutan consider red pandas to bereincarnations ofBuddhist monks. Some tribal people in northeast India and theYi people believe that it brings good luck to wear red panda tails or hats made of its fur.[101] In China, the fur is used for local cultural ceremonies. At weddings, the bridegroom traditionally carries the hide. Hats made of red panda tails are also used by local newlyweds as a "good-luck charm".[48]
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