The koala (Phascolarctos cinereus, or, inaccurately, koala bear[a]) is an arboreal herbivorous marsupial native to Australia. It is the only extant representative of the family Phascolarctidae and its closest living relatives are the wombats, which comprise the family Vombatidae. The koala is found in coastal areas of the mainland's eastern and southern regions, inhabiting Queensland, New South Wales, Victoria, and South Australia. It is easily recognisable by its stout, tailless body and large head with round, fluffy ears and large, spoon-shaped nose. The koala has a body length of 60–85 cm (24–33 in) and weighs 4–15 kg (9–33 lb). Pelage colour ranges from silver grey to chocolate brown. Koalas from the northern populations are typically smaller and lighter in colour than their counterparts further south. These populations possibly are separate subspecies, but this is disputed.
Koalas typically inhabit open eucalypt woodlands, and the leaves of these trees make up most of their diet. Because this eucalypt diet has limited nutritional and caloric content, koalas are largely sedentary and sleep up to 20 hours a day. They are asocial animals, and bonding exists only between mothers and dependent offspring. Adult males communicate with loud bellows that intimidate rivals and attract mates. Males mark their presence with secretions from scent glands located on their chests. Being marsupials, koalas give birth to underdeveloped young that crawl into their mothers' pouches, where they stay for the first six to seven months of their lives. These young koalas, known as joeys, are fully weaned around a year old. Koalas have few natural predators and parasites, but are threatened by various pathogens, such as Chlamydiaceae bacteria and the koala retrovirus, as well as by bushfires and droughts.
Koalas were hunted by Indigenous Australians and depicted in myths and cave art for millennia. The first recorded encounter between a European and a koala was in 1798, and an image of the animal was published in 1810 by naturalist George Perry. Botanist Robert Brown wrote the first detailed scientific description of the koala in 1814, although his work remained unpublished for 180 years. Popular artist John Gould illustrated and described the koala, introducing the species to the general British public. Further details about the animal's biology were revealed in the 19th century by several English scientists. Because of its distinctive appearance, the koala is recognised worldwide as a symbol of Australia. Koalas are listed as Vulnerable by the International Union for Conservation of Nature. The animal was hunted heavily in the early 20th century for its fur, and large-scale cullings in Queensland resulted in a public outcry that initiated a movement to protect the species. Sanctuaries were established, and translocation efforts moved to new regions koalas whose habitat had become fragmented or reduced. The biggest threat to their existence is habitat destruction caused by agriculture and urbanisation.
Etymology
The word koala comes from the Dharug gula, meaning no water.[4] Although the vowel 'u' was originally written in the English orthography as "oo" (in spellings such as coola or koolah), it was changed to "oa", possibly in error.[5] Because of the koala's supposed resemblance to a bear, it was often miscalled the koala bear, particularly by early settlers.[6] The generic name, Phascolarctos, is derived from the Greek words phaskolos "pouch" and arktos "bear". The specific name, cinereus, is Latin for "ash coloured".[7]
Taxonomy and evolution
The koala was given its generic name Phascolarctos in 1816 by French zoologist Henri Marie Ducrotay de Blainville,[9] who would not give it a specific name until further review. In 1819, German zoologist Georg August Goldfuss gave it the binomial Lipurus cinereus. Because Phascolarctos was published first, according to the International Code of Zoological Nomenclature, it has priority as the official name of the genus.[10] French naturalist Anselme Gaëtan Desmarest proposed the name Phascolartos fuscus in 1820, suggesting that the brown-coloured versions were a different species than the grey ones. Other names suggested by European authors included Marodactylus cinereus by Goldfuss in 1820, P. flindersii by René Primevère Lesson in 1827, and P. koala by John Edward Gray in 1827.[2]
The koala is classified with wombats (family Vombatidae) and several extinct families (including marsupial tapirs, marsupial lions and giant wombats) in the suborder Vombatiformes within the order Diprotodontia.[11] The Vombatiformes are a sister group to a clade that includes macropods (kangaroos and wallabies) and possums.[12] The ancestors of vombatiforms were likely arboreal,[8] and the koala's lineage was possibly the first to branch off around 40 million years ago during the Eocene
The modern koala is the only extant member of Phascolarctidae, a family that once included several genera and species. During the Oligocene and Miocene, koalas lived in rainforests and had less specialised diets.[14] Some species, such as the Riversleigh rainforest koala (Nimiokoala greystanesi) and some species of Perikoala, were around the same size as the modern koala, while others, such as species of Litokoala, were one-half to two-thirds its size.[15] Like the modern species, prehistoric koalas had well developed ear structures which suggests that long-distance vocalising and sedentism developed early.[14] During the Miocene, the Australian continent began drying out, leading to the decline of rainforests and the spread of open Eucalyptus woodlands. The genus Phascolarctos split from Litokoala in the late Miocene[14][16] and had several adaptations that allowed it to live on a specialised eucalyptus diet: a shifting of the palate towards the front of the skull; larger molars and premolars; smaller pterygoid fossa;[14] and a larger gap between the molar and the incisor teeth.[17]
During the Pliocene and Pleistocene, when Australia experienced changes in climate and vegetation, koala species grew larger.[15] P. cinereus may have emerged as a dwarf form of the giant koala (P. stirtoni). The reduction in the size of large mammals has been seen as a common phenomenon worldwide during the late Pleistocene, and several Australian mammals, such as the agile wallaby, are traditionally believed to have resulted from this dwarfing. A 2008 study questions this hypothesis, noting that P. cinereus and P. stirtoni were sympatric during the middle to late Pleistocene, and possibly as early as the Pliocene.[18] The fossil record of the modern koala extends back at least to the middle Pleistocene.[19]
Genetics and variations
Traditionally, three distinct subspecies have been recognised: the Queensland koala (P. c. adustus, Thomas 1923), the New South Wales koala (P. c. cinereus, Goldfuss 1817), and the Victorian koala (P. c. victor, Troughton 1935). These forms are distinguished by pelage colour and thickness, body size, and skull shape. The Queensland koala is the smallest of the three, with shorter, silver fur and a shorter skull. The Victorian koala is the largest, with shaggier, brown fur and a wider skull.[20][21] The boundaries of these variations are based on state borders, and their status as subspecies is disputed. A 1999 genetic study suggests that the variations represent differentiated populations with limited gene flow between them, and that the three subspecies comprise a single evolutionarily significant unit.[21] Other studies have found that koala populations have high levels of inbreeding and low genetic variation.[22][23] Such low genetic diversity may have been a characteristic of koala populations since the late Pleistocene.[24] Rivers and roads have been shown to limit gene flow and contribute to the genetic differentiation of southeast Queensland populations.[25] In April 2013, scientists from the Australian Museum and Queensland University of Technology announced they had fully sequenced the koala genome.[26]
Koalas typically inhabit open eucalypt woodlands, and the leaves of these trees make up most of their diet. Because this eucalypt diet has limited nutritional and caloric content, koalas are largely sedentary and sleep up to 20 hours a day. They are asocial animals, and bonding exists only between mothers and dependent offspring. Adult males communicate with loud bellows that intimidate rivals and attract mates. Males mark their presence with secretions from scent glands located on their chests. Being marsupials, koalas give birth to underdeveloped young that crawl into their mothers' pouches, where they stay for the first six to seven months of their lives. These young koalas, known as joeys, are fully weaned around a year old. Koalas have few natural predators and parasites, but are threatened by various pathogens, such as Chlamydiaceae bacteria and the koala retrovirus, as well as by bushfires and droughts.
Koalas were hunted by Indigenous Australians and depicted in myths and cave art for millennia. The first recorded encounter between a European and a koala was in 1798, and an image of the animal was published in 1810 by naturalist George Perry. Botanist Robert Brown wrote the first detailed scientific description of the koala in 1814, although his work remained unpublished for 180 years. Popular artist John Gould illustrated and described the koala, introducing the species to the general British public. Further details about the animal's biology were revealed in the 19th century by several English scientists. Because of its distinctive appearance, the koala is recognised worldwide as a symbol of Australia. Koalas are listed as Vulnerable by the International Union for Conservation of Nature. The animal was hunted heavily in the early 20th century for its fur, and large-scale cullings in Queensland resulted in a public outcry that initiated a movement to protect the species. Sanctuaries were established, and translocation efforts moved to new regions koalas whose habitat had become fragmented or reduced. The biggest threat to their existence is habitat destruction caused by agriculture and urbanisation.
Etymology
The word koala comes from the Dharug gula, meaning no water.[4] Although the vowel 'u' was originally written in the English orthography as "oo" (in spellings such as coola or koolah), it was changed to "oa", possibly in error.[5] Because of the koala's supposed resemblance to a bear, it was often miscalled the koala bear, particularly by early settlers.[6] The generic name, Phascolarctos, is derived from the Greek words phaskolos "pouch" and arktos "bear". The specific name, cinereus, is Latin for "ash coloured".[7]
Taxonomy and evolution
The koala was given its generic name Phascolarctos in 1816 by French zoologist Henri Marie Ducrotay de Blainville,[9] who would not give it a specific name until further review. In 1819, German zoologist Georg August Goldfuss gave it the binomial Lipurus cinereus. Because Phascolarctos was published first, according to the International Code of Zoological Nomenclature, it has priority as the official name of the genus.[10] French naturalist Anselme Gaëtan Desmarest proposed the name Phascolartos fuscus in 1820, suggesting that the brown-coloured versions were a different species than the grey ones. Other names suggested by European authors included Marodactylus cinereus by Goldfuss in 1820, P. flindersii by René Primevère Lesson in 1827, and P. koala by John Edward Gray in 1827.[2]
The koala is classified with wombats (family Vombatidae) and several extinct families (including marsupial tapirs, marsupial lions and giant wombats) in the suborder Vombatiformes within the order Diprotodontia.[11] The Vombatiformes are a sister group to a clade that includes macropods (kangaroos and wallabies) and possums.[12] The ancestors of vombatiforms were likely arboreal,[8] and the koala's lineage was possibly the first to branch off around 40 million years ago during the Eocene
The modern koala is the only extant member of Phascolarctidae, a family that once included several genera and species. During the Oligocene and Miocene, koalas lived in rainforests and had less specialised diets.[14] Some species, such as the Riversleigh rainforest koala (Nimiokoala greystanesi) and some species of Perikoala, were around the same size as the modern koala, while others, such as species of Litokoala, were one-half to two-thirds its size.[15] Like the modern species, prehistoric koalas had well developed ear structures which suggests that long-distance vocalising and sedentism developed early.[14] During the Miocene, the Australian continent began drying out, leading to the decline of rainforests and the spread of open Eucalyptus woodlands. The genus Phascolarctos split from Litokoala in the late Miocene[14][16] and had several adaptations that allowed it to live on a specialised eucalyptus diet: a shifting of the palate towards the front of the skull; larger molars and premolars; smaller pterygoid fossa;[14] and a larger gap between the molar and the incisor teeth.[17]
During the Pliocene and Pleistocene, when Australia experienced changes in climate and vegetation, koala species grew larger.[15] P. cinereus may have emerged as a dwarf form of the giant koala (P. stirtoni). The reduction in the size of large mammals has been seen as a common phenomenon worldwide during the late Pleistocene, and several Australian mammals, such as the agile wallaby, are traditionally believed to have resulted from this dwarfing. A 2008 study questions this hypothesis, noting that P. cinereus and P. stirtoni were sympatric during the middle to late Pleistocene, and possibly as early as the Pliocene.[18] The fossil record of the modern koala extends back at least to the middle Pleistocene.[19]
Genetics and variations
Traditionally, three distinct subspecies have been recognised: the Queensland koala (P. c. adustus, Thomas 1923), the New South Wales koala (P. c. cinereus, Goldfuss 1817), and the Victorian koala (P. c. victor, Troughton 1935). These forms are distinguished by pelage colour and thickness, body size, and skull shape. The Queensland koala is the smallest of the three, with shorter, silver fur and a shorter skull. The Victorian koala is the largest, with shaggier, brown fur and a wider skull.[20][21] The boundaries of these variations are based on state borders, and their status as subspecies is disputed. A 1999 genetic study suggests that the variations represent differentiated populations with limited gene flow between them, and that the three subspecies comprise a single evolutionarily significant unit.[21] Other studies have found that koala populations have high levels of inbreeding and low genetic variation.[22][23] Such low genetic diversity may have been a characteristic of koala populations since the late Pleistocene.[24] Rivers and roads have been shown to limit gene flow and contribute to the genetic differentiation of southeast Queensland populations.[25] In April 2013, scientists from the Australian Museum and Queensland University of Technology announced they had fully sequenced the koala genome.[26]
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