Sloths [1] Temporal range: Early Oligocene to Holocene | |
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Bradypus variegatus , a three-toed sloth | |
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Choloepus hoffmanni , a two-toed sloth | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Superorder: | Xenarthra |
Order: | Pilosa |
Suborder: | Folivora Delsuc, Catzeflis, Stanhope, and Douzery, 2001 [2] |
Families | |
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Red: two-toed sloth,Blue: three-toed sloth,Purple: both two-toed sloth and three-toed sloth | |
Synonyms | |
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Sloths are a Neotropical group of xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their lives hanging upside down in the trees of the tropical rainforests of South America and Central America. Sloths are considered to be most closely related to anteaters, together making up the xenarthran order Pilosa.
There are six extant sloth species in two genera – Bradypus (three–toed sloths) and Choloepus (two–toed sloths). Despite this traditional naming, all sloths have three toes on each rear limb-- although two-toed sloths have only two digits on each forelimb. [3] The two groups of sloths are from different, distantly related families, and are thought to have evolved their morphology via parallel evolution from terrestrial ancestors. Besides the extant species, many species of ground sloths ranging up to the size of elephants (like Megatherium ) inhabited both North and South America during the Pleistocene Epoch. However, they became extinct during the Quaternary extinction event around 12,000 years ago, along with most large animals across the Americas. The extinction correlates in time with the arrival of humans, but climate change has also been suggested to have contributed. Members of an endemic radiation of Caribbean sloths also formerly lived in the Greater Antilles but became extinct after humans settled the archipelago in the mid-Holocene, around 6,000 years ago.
Sloths are so named because of their very low metabolism and deliberate movements. Sloth, related to slow, literally means "laziness," and their common names in several other languages (e.g. German : Faultier, French : paresseux, Spanish : perezoso, Romanian: leneș, Finnish: laiskiainen) also mean "lazy" or similar. Their slowness permits their low-energy diet of leaves and avoids detection by predatory hawks and cats that hunt by sight. [3] Sloths are almost helpless on the ground but are able to swim. [4] The shaggy coat has grooved hair that is host to symbiotic green algae which camouflage the animal in the trees and provide it nutrients. The algae also nourish sloth moths, some species of which exist solely on sloths. [5]
Sloths belong to the superorder Xenarthra, a group of placental mammals believed to have evolved in the continent of South America around 60 million years ago. [6] One study found that xenarthrans broke off from other placental mammals around 100 million years ago. [7] Anteaters and armadillos are also included among Xenarthra. The earliest xenarthrans were arboreal herbivores with sturdy vertebral columns, fused pelvises, stubby teeth, and small brains. Sloths are in the taxonomic suborder Folivora [2] of the order Pilosa. These names are from the Latin 'leaf eater' and 'hairy', respectively. Pilosa is one of the smallest of the orders of the mammal class; its only other suborder contains the anteaters.
The Folivora are divided into at least eight families, only two of which have living species; the remainder are entirely extinct (†): [8]
The common ancestor of the two existing sloth genera dates to about 28 million years ago, [8] with similarities between the two- and three- toed sloths an example of convergent evolution to an arboreal lifestyle, "one of the most striking examples of convergent evolution known among mammals". [13] The ancient Xenarthra included a significantly greater variety of species, with a wider distribution, than those of today. Ancient sloths were mostly terrestrial, and some reached sizes that rival those of elephants, as was the case for Megatherium . [4]
Sloths arose in South America during a long period of isolation and eventually spread to a number of the Caribbean islands as well as North America. It is thought that swimming led to oceanic dispersal of pilosans to the Greater Antilles by the Oligocene, and that the megalonychid Pliometanastes and the mylodontid Thinobadistes were able to colonise North America about 9 million years ago, well before the formation of the Isthmus of Panama. The latter development, about 3 million years ago, allowed megatheriids and nothrotheriids to also invade North America as part of the Great American Interchange. Additionally, the nothrotheriid Thalassocnus of the west coast of South America became adapted to a semiaquatic and, eventually, perhaps fully aquatic marine lifestyle. [14] In Peru and Chile, Thalassocnus entered the coastal habitat beginning in the late Miocene. They presumably waded and paddled in the water for short period, but over a span of 4 million years, they eventually evolved into swimming creatures, becoming specialist bottom feeders of seagrasses, similar to the extant sirenians. [15]
Both types of extant tree sloth tend to occupy the same forests; in most areas, a particular species of the somewhat smaller and generally slower-moving three-toed sloth (Bradypus) and a single species of the two-toed type will jointly predominate. Based on morphological comparisons, it was thought the two-toed sloths nested phylogenetically within one of the divisions of the extinct Greater Antilles sloths. [16] Though data has been collected on over 33 different species of sloths by analyzing bone structures, many of the relationships between clades on a phylogenetic tree were unclear. [17] Much of the morphological evidence collected to support the hypothesis of diphyly has been based on the structure of the inner ear. [18]
Recently obtained molecular data from collagen [8] and mitochondrial DNA sequences [19] fall in line with the diphyly (convergent evolution) hypothesis but have overturned some of the other conclusions obtained from morphology. These investigations consistently place two-toed sloths close to mylodontids and three-toed sloths within Megatherioidea, close to Megalonyx , megatheriids and nothrotheriids. They make the previously recognized family Megalonychidae polyphyletic, with both two-toed sloths and Greater Antilles sloths being moved away from Megalonyx. Greater Antilles sloths are now placed in a separate, basal branch of the sloth evolutionary tree. [8] [19]
The following sloth family phylogenetic tree is based on collagen and mitochondrial DNA sequence data. [8]
Folivora |
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The marine sloths of South America's Pacific coast became extinct at the end of the Pliocene following the closing of the Central American Seaway; the closing caused a cooling trend in the coastal waters which killed off much of the area's seagrass (and which would have also made thermoregulation difficult for the sloths, with their slow metabolism). [20]
Ground sloths disappeared from both North and South America shortly after the appearance of humans about 11,000 years ago. Evidence suggests human hunting contributed to the extinction of the American megafauna. Ground sloth remains found in both North and South America indicate that they were killed, cooked, and eaten by humans. [4] Climate change that came with the end of the last ice age may have also played a role, although previous similar glacial retreats were not associated with similar extinction rates.
Megalocnus and some other Caribbean sloths survived until about 5,000 years ago, long after ground sloths had died out on the mainland, but then went extinct when humans finally colonized the Greater Antilles. [21]
Sloths can be 60 to 80 cm (24 to 31 in) long and, depending on the species, weigh from 3.6 to 7.7 kg (7.9 to 17.0 lb). Two-toed sloths are slightly larger than three-toed sloths. [22] Sloths have long limbs and rounded heads with tiny ears. Three-toed sloths also have stubby tails about 5 to 6 cm (2.0 to 2.4 in) long.
Sloths are unusual among mammals in not having seven cervical vertebrae. Two-toed sloths have five to seven, while three-toed sloths have eight or nine. The other mammals not having seven are the manatees, with six. [23]
Sloths have colour vision but have poor visual acuity. They also have poor hearing. Thus, they rely on their sense of smell and touch to find food. [24]
Sloths have very low metabolic rates (less than half of that expected for a mammal of their size), and low body temperatures: 30 to 34 °C (86 to 93 °F) when active, and still lower when resting. Sloths are heterothermic, meaning their body temperature may vary according to the environment, normally ranging from 25 to 35 °C (77 to 95 °F), but able to drop to as low as 20 °C (68 °F), inducing torpor. [24]
The outer hairs of sloth fur grow in a direction opposite from that of other mammals. In most mammals, hairs grow toward the extremities, but because sloths spend so much time with their limbs above their bodies, their hairs grow away from the extremities to provide protection from the elements while they hang upside down. In most conditions, the fur hosts symbiotic algae, which provide camouflage [25] from predatory jaguars, ocelots, [26] and harpy eagles. [27] Because of the algae, sloth fur is a small ecosystem of its own, hosting many species of commensal and parasitic arthropods. [28] There are a large number of arthropods associated with sloths. These include biting and blood-sucking flies such as mosquitoes and sandflies, triatomine bugs, lice, ticks and mites. Sloths have a highly specific community of commensal beetles, mites and moths. [29] The species of sloths recorded to host arthropods include [29] the pale-throated three-toed sloth, the brown-throated three-toed sloth, and Linnaeus's two-toed sloth. Sloths benefit from their relationship with moths because the moths are responsible for fertilizing algae on the sloth, which provides them with nutrients. [30]
Their limbs are adapted for hanging and grasping, not for supporting their weight. Muscle mass makes up only 25 to 30 percent of their total body weight. Most other mammals have a muscle mass that makes up 40 to 45 percent of their total body weight. [31] Their specialised hands and feet have long, curved claws to allow them to hang upside down from branches without effort, [32] and are used to drag themselves along the ground, since they cannot walk. On three-toed sloths, the arms are 50 percent longer than the legs. [24]
Sloths move only when necessary and even then, very slowly. They usually move at an average speed of 4 metres (13 ft) per minute but can move at a marginally higher speed of 4.5 metres (15 ft) per minute if they are in immediate danger from a predator. While they sometimes sit on top of branches, they usually eat, sleep, and even give birth hanging from branches. They sometimes remain hanging from branches even after death. On the ground, the maximum speed of sloths is 3 metres (9.8 ft) per minute. Two-toed sloths are generally better able than three-toed sloths to disperse between clumps of trees on the ground. [33]
Sloths are surprisingly strong swimmers and can reach speeds of 13.5 metres (44 ft) per minute. [34] They use their long arms to paddle through the water and can cross rivers and swim between islands. [35] Sloths can reduce their already slow metabolism even further and slow their heart rate to less than a third of normal, allowing them to hold their breath underwater for up to 40 minutes. [36]
Wild brown-throated three-toed sloths sleep on average 9.6 hours a day. [37] Two-toed sloths are nocturnal. [38] Three-toed sloths are mostly nocturnal but can be active in the day. They spend 90 per cent of their time motionless. [24]
Sloths are solitary animals that rarely interact with one another except during breeding season, [39] though female sloths do sometimes congregate, more so than do males. [40]
Sloths descend about once every eight days to defecate on the ground. The reason and mechanism behind this behavior have long been debated among scientists. There are at least five hypotheses: 1) fertilize trees when feces are deposited at the base of the tree; [41] 2) cover feces and avoid predation; [42] [43] [44] 3) chemical communication between individuals; [45] 4) pick up trace nutrients in their claws, that are then ingested; [46] and 5) favor a mutualistic relationship with populations of fur moths. [44] [46] More recently, a new hypothesis has emerged, which presents evidence against the previous ones and proposes that all current sloths are descendants from species that defecated on the ground, and there simply has not been enough selective pressure to abandon this behavior, since cases of predation during defecation are actually very rare. [47]
Baby sloths learn what to eat by licking the lips of their mother. [48] All sloths eat the leaves of Cecropia .
Two-toed sloths are omnivorous, with a diverse diet of insects, carrion, fruits, leaves and small lizards, ranging over up to 140 hectares (350 acres). Three-toed sloths, on the other hand, are almost entirely herbivorous (plant eaters), with a limited diet of leaves from only a few trees, [39] and no other mammal digests its food as slowly.
They have made adaptations to arboreal browsing. Leaves, their main food source, provide very little energy or nutrients, and do not digest easily, so sloths have large, slow-acting, multi-chambered stomachs in which symbiotic bacteria break down the tough leaves. [39] As much as two-thirds of a well-fed sloth's body weight consists of the contents of its stomach, and the digestive process can take a month or more to complete.
Three-toed sloths go to the ground to urinate and defecate about once a week, digging a hole and covering it afterwards. They go to the same spot each time and are vulnerable to predation while doing so. Considering the large energy expenditure and dangers involved in the journey to the ground, this behaviour has been described as a mystery. [49] [50] [51] Recent research shows that moths, which live in the sloth's fur, lay eggs in the sloth's feces. When they hatch, the larvae feed on the feces, and when mature fly up onto the sloth above. These moths may have a symbiotic relationship with sloths, as they live in the fur and promote growth of algae, which the sloths eat. [5] Individual sloths tend to spend the bulk of their time feeding on a single "modal" tree; by burying their excreta near the trunk of that tree, they may also help nourish it. [52]
The pale- and brown-throated three-toed sloths mate seasonally, while the maned three-toed sloth breeds at any time of the year. The reproduction of pygmy three-toed sloths is currently unknown. Litters are of one newborn only, after six months' gestation for three-toed, and 12 months for two-toed. Newborns stay with their mother for about five months. In some cases, young sloths die from a fall indirectly because the mothers prove unwilling to leave the safety of the trees to retrieve the young. [53] Females normally bear one baby every year, but sometimes sloths' low level of movement actually keeps females from finding males for longer than one year. [54] Sloths are not particularly sexually dimorphic and several zoos have received sloths of the wrong sex. [55] [56]
The average lifespan of two-toed sloths in the wild is currently unknown due to a lack of full-lifespan studies in a natural environment. [57] Median life expectancy in human care is about 16 years, with one individual at the Smithsonian Institution's National Zoo reaching an age of 49 years before her death. [58]
Although habitat is limited to the tropical rainforests of Central and South America, in that environment sloths are successful. On Barro Colorado Island in Panama, sloths have been estimated to constitute 70% of the biomass of arboreal mammals. [59] Four of the six living species are currently rated "least concern"; the maned three-toed sloth (Bradypus torquatus), which inhabits Brazil's dwindling Atlantic Forest, is classified as "vulnerable", [60] while the island-dwelling pygmy three-toed sloth (B. pygmaeus) is critically endangered. Sloths' lower metabolism confines them to the tropics, and they adopt thermoregulation behaviors of cold-blooded animals such as sunning themselves. [61]
The majority of recorded sloth deaths in Costa Rica are due to contact with electrical lines and poachers. Their claws also provide another, unexpected deterrent to human hunters; when hanging upside-down in a tree, they are held in place by the claws themselves and often do not fall down even if shot from below.
Sloths are victims of animal trafficking where they are sold as pets. However, they generally make very poor pets, as they have such a specialized ecology. [62]
The Sloth Institute Costa Rica is known for caring, rehabilitating and releasing sloths back into the wild. [63] Also in Costa Rica, the Aviarios Sloth Sanctuary cares for sloths. It has rehabilitated and returned about 130 individuals to the wild. [64] However, a report in May 2016 featured two former veterinarians from the facility who were critical of the sanctuary's efforts, accusing it of mistreating the animals. [65]
Xenarthra is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Extinct xenarthrans include the glyptodonts, pampatheres and ground sloths. Xenarthrans originated in South America during the late Paleocene about 60 million years ago. They evolved and diversified extensively in South America during the continent's long period of isolation in the early to mid Cenozoic Era. They spread to the Antilles by the early Miocene and, starting about 3 million years ago, spread to Central and North America as part of the Great American Interchange. Nearly all of the formerly abundant megafaunal xenarthrans became extinct at the end of the Pleistocene.
Megatherium is an extinct genus of ground sloths endemic to South America that lived from the Early Pliocene through the end of the Pleistocene. It is best known for the elephant-sized type species Megatherium americanum, native to the Pampas through southern Bolivia during the Pleistocene. Various other species belonging to the subgenus Pseudomegatherium ranging in size comparable to considerably smaller than M. americanum are known from the Andean region.
Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. Ground sloths varied widely in size, with the largest, belonging to genera Megatherium, Lestodon, and Eremotherium, being around the size of elephants, with body masses of around 4 tonnes. Ground sloths are a paraphyletic group, as living tree sloths are thought to have evolved from ground sloth ancestors.
Megatheriidae is a family of extinct ground sloths that lived from approximately 23 mya—11,000 years ago.
Mylodontidae is a family of extinct South American and North American ground sloths within the suborder Folivora of order Pilosa, living from around 23 million years ago (Mya) to 11,000 years ago. This family is most closely related to another family of extinct ground sloths, Scelidotheriidae, as well as to the extant arboreal two-toed sloths, family Choloepodidae; together these make up the superfamily Mylodontoidea. Phylogenetic analyses based on morphology uncovered the relationship between Mylodontidae and Scelidotheriidae; in fact, the latter was for a time considered a subfamily of mylodontids. However, molecular sequence comparisons were needed for the correct placement of Choloepodidae. These studies have been carried out using mitochondrial DNA sequences as well as with collagen amino acid sequences. The latter results indicate that Choloepodidae is closer to Mylodontidae than Scelidotheriidae is. The only other living sloth family, Bradypodidae, belongs to a different sloth radiation, Megatherioidea.
Scelidotheriidae is a family of extinct ground sloths within the order Pilosa, suborder Folivora and superfamily Mylodontoidea, related to the other extinct mylodontoid family, Mylodontidae, as well as to the living two-toed sloth family Choloepodidae. The only other extant family of the suborder Folivora is the distantly related Bradypodidae. Erected as the family Scelidotheriidae by Ameghino in 1889, the taxon was demoted to a subfamily of Mylodontidae by Gaudin in 1995. However, recent collagen sequence data indicates the group is less closely related to Mylodon and Lestodon than Choloepus is, and thus it has been elevated back to full family status by Presslee et al. (2019).
The maned sloth is a three-toed sloth that is native to South America. It is one of four species of three-toed sloths belonging to the suborder Xenarthra and are placental mammals. They are endemic to the Atlantic coastal rainforest of southeastern and northeastern Brazil, located in the states of Espírito Santo, Rio de Janeiro and Bahia. Each of the individuals within the species are genetically distinct with different genetic makeup.The maned sloth is listen under Vulnerable (VU) according to the IUCN Red List and have a decreasing population trend.
The brown-throated sloth is a species of three-toed sloth found in the Neotropical realm of Central and South America.
Linnaeus's two-toed sloth, also known as the southern two-toed sloth, unau, or Linne's two-toed sloth is a species of sloth from South America, found in Venezuela, Suriname, Guyana, French Guiana, Colombia, Ecuador, Peru, and Brazil north of the Amazon River. There is now evidence suggesting the species' range expands into Bolivia. It's the largest extant sloth species.
The order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes anteaters and sloths. The name comes from the Latin word for "hairy".
Choloepus is a genus of xenarthran mammals of Central and South America within the monotypic family Choloepodidae, consisting of two-toed sloths, sometimes also called two-fingered sloths. The two species of Choloepus, Linnaeus's two-toed sloth and Hoffmann's two-toed sloth, were formerly believed on the basis of morphological studies to be the only surviving members of the sloth family Megalonychidae, but have now been shown by molecular results to be closest to extinct ground sloths of the family Mylodontidae.
Megalonychidae is an extinct family of sloths including the extinct Megalonyx. Megalonychids first appeared in the early Oligocene, about 35 million years (Ma) ago, in southern Argentina (Patagonia). There is, however, one possible find dating to the Eocene, about 40 Ma ago, on Seymour Island in Antarctica. They first reached North America by island-hopping across the Central American Seaway, about 9 million years ago, prior to formation of the Isthmus of Panama about 2.7 million years ago. Some megalonychid lineages increased in size as time passed. The first species of these were small and may have been partly tree-dwelling, whereas the Pliocene species were already approximately half the size of the huge Late Pleistocene Megalonyx jeffersonii from the last ice age.
The three-toed or three-fingered sloths are arboreal neotropical mammals. They are the only members of the genus Bradypus and the family Bradypodidae. The five living species of three-toed sloths are the brown-throated sloth, the maned sloth, the pale-throated sloth, the southern maned sloth, and the pygmy three-toed sloth. In complete contrast to past morphological studies, which tended to place Bradypus as the sister group to all other folivorans, molecular studies place them nested within the sloth superfamily Megatherioidea, making them the only surviving members of that radiation.
The pygmy three-toed sloth, also known as the monk sloth or dwarf sloth, is a species of sloth in the family Bradypodidae. The species is endemic to Isla Escudo de Veraguas, a small island off the Caribbean coast of Panama. The species was first described by Robert P. Anderson of the University of Kansas and Charles O. Handley Jr., of the Smithsonian Institution in 2001. The pygmy three-toed sloth is significantly smaller than the other three members of its genus, but otherwise resembles the brown-throated three-toed sloth. According to Anderson and Handley Jr., the head-and-body length is between 48 and 53 centimetres, and the body mass ranges from 2.5 to 3.5 kg.
Lestodon is an extinct genus of giant ground sloth native to South America during the Pleistocene epoch. Its fossil remains have been primarily been found in the Pampas and adjacent regions. The largest member of the family Mylodontidae, It is estimated to have weighed 4,100 kilograms. It was a herbivore and primarily fed on the grasses and low-growing plants.
The mammalian order Pilosa, which includes the sloths and anteaters, includes various species from the Caribbean region. Many species of sloths are known from the Greater Antilles, all of which became extinct over the last millennia, but some sloths and anteaters survive on islands closer to the mainland.
Ahytherium is an extinct genus of megalonychid sloth that lived during the Pleistocene of what is now Brazil. It contains a single species, A. aureum.
Nothrotheriidae is a family of extinct ground sloths that lived from approximately 17.5 mya—10,000 years ago, existing for approximately 17.49 million years. Previously placed within the tribe Nothrotheriini or subfamily Nothrotheriinae within Megatheriidae, they are now usually placed in their own family, Nothrotheriidae. Nothrotheriids appeared in the Burdigalian, some 19.8 million years ago, in South America. The group includes the comparatively slightly built Nothrotheriops, which reached a length of about 2.75 metres (9.0 ft). While nothrotheriids were small compared to some of their megatheriid relatives, their claws provided an effective defense against predators, like those of larger anteaters today.
Lakukullus is an extinct genus of nothrotheriid ground sloths that lived during the Middle Miocene around 13.8 to 11.8 million years ago of what is now Bolivia.