Two-toed sloth
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''Choloepus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of xenarthran mammals of Central and South America within the monotypic
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Choloepodidae, consisting of two-toed sloths, sometimes also called two-fingered sloths. The two species of ''Choloepus'' (which means "lame foot"), Linnaeus's two-toed sloth (''Choloepus didactylus'') and
Hoffmann's two-toed sloth Hoffmann's two-toed sloth (''Choloepus hoffmanni''), also known as the northern two-toed sloth is a species of sloth from Central and South America. It is a solitary, largely nocturnal and arboreal animal, found in mature and secondary rainf ...
(''Choloepus hoffmanni''), were formerly believed on the basis of morphological studies to be the only surviving members of the
sloth Sloths are a group of Neotropical 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 l ...
family Megalonychidae, but have now been shown by molecular results to be closest to extinct ground sloths of the family Mylodontidae.


Extant species


Evolution

A study of retrovirus and mitochondrial DNA suggests that ''C. didactylus'' and ''C. hoffmani'' diverged 6 to 7 million years ago. Furthermore, based on
cytochrome c oxidase subunit I Cytochrome c oxidase I (COX1) also known as mitochondrially encoded cytochrome c oxidase I (MT-CO1) is a protein that in humans is encoded by the ''MT-CO1'' gene. In other eukaryotes, the gene is called ''COX1'', ''CO1'', or ''COI''. Cytoch ...
sequences, a similar divergence date ( years ago) between the two populations of ''C. hofmanni'' separated by the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
has been reported. Their ancestors evolved with marine vertebrae, the three toed-sloth and the manatee are the only other mammals with similar vertebrae.


Relation to the three-toed sloth

Both types of 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. Although similar in overall appearance, the relationship between the two genera is not close. Recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analyses support analysis of morphological data from the 1970s and 1980s, indicating the two genera are not closely related and adapted to their arboreal lifestyles independently. It was unclear from this work from which ground-dwelling sloth taxa the three-toed sloths evolved. Based on the morphological comparisons, it was thought the two-toed sloths nested phylogenetically within one of the divisions of Caribbean sloths. 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. Much of the morphological evidence to support the hypothesis of diphyly has been based on the structure of the inner ear. Most morphological studies have concluded that
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
is the mechanism that resulted in today’s two genera of tree sloths. This means that the extant genera evolved analogous traits, such as locomotion methods, size, habitat, and many other traits independently from one another as opposed to from their last common ancestor. This makes tree sloths “one of the most striking examples of convergent evolution known among mammals”. Recently obtained molecular data from collagen and mitochondrial DNA sequences 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 ''Megalonyx'' ( Greek, "large claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America during the Pliocene and Pleistocene epochs. It became extinct during the Quaternary extinction event at the end ...
'', megatheriids and
nothrotheriid Nothrotheriidae is a family of extinct ground sloths that lived from approximately 17.5 mya—10,000 years ago, existing for approximately . Previously placed within the tribe Nothrotheriini or subfamily Nothrotheriinae within Megatheriidae, they ...
s. They make the previously recognized family Megalonychidae polyphyletic, with both two-toed sloths and the Caribbean sloths being moved out of that family and away from ''Megalonyx''. Caribbean sloths are placed in a separate, basal branch of the sloth evolutionary tree. The following sloth family phylogenetic tree is based on collagen and mitochondrial DNA sequence data (see Fig. 4 of Presslee ''et al''., 2019).


Characteristics

The name "two-toed sloth" was intended to describe an anatomical difference between the genera ''Choloepus'' and ''Bradypus'', but does so in a potentially misleading way. Members of ''Choloepus'' have two digits on their forelimbs (the thoracic limbs) and three digits on their hindlimbs (the pelvic limbs), while members of ''Bradypus'' have three digits on all limbs. Although the term "two-fingered" sloth is arguably less misleading, the shorter "two-toed" is much more widely used. Members of ''Choloepus'' are larger than three-toed sloths, having a body length of , and weighing . Other distinguishing features include a more prominent snout, longer fur, and the absence of a
tail The tail is the section at the rear end of certain kinds of animals’ bodies; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammal ...
.


Behaviour

Two-toed sloths spend most of their lives hanging upside down from trees. They cannot walk, so they pull hand-over-hand to move around, which is at an extremely slow rate. Being predominantly nocturnal, their fur, which grows greenish algae to blend in, is their main source of protection. Their body temperatures depend at least partially on the ambient temperature; they cannot shiver to keep warm, as other mammals do, because of their unusually low metabolic rates and reduced musculature. Two-toed sloths also differ from three-toed sloths in their climbing behavior, preferring to descend head first.


Lifecycle

Two-toed sloths have a gestation period of six months to a year, depending on the species. The mother gives birth to a single young, while hanging upside down. The young are born with claws, and are weaned after about a month, although they will remain with the mother for several more months, and do not reach sexual maturity until the age of three years, in the case of females, or four to five years, in the case of males. During natal dispersion, two-toed sloths prefer tropical forests over other types of habitat, often using riparian forest buffers to disperse. Although they also occupy shade-grown cacao plantations, they avoid open pastures.


Feeding

They eat primarily leaves, but also shoots,
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
s, nuts,
berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, ras ...
,
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, e ...
, some native
flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
, and even some small vertebrates. In addition, when they cannot find food, they have been known to eat the algae that grow on their fur for nutrients. They have large, four-chambered stomachs, which help to ferment the large amount of plant matter they eat. Food can take up to a month to digest due to their slow metabolism. Depending on when in the excretion cycle a sloth is weighed, urine and feces may account for up to 30% of the animal’s body weight, which averages about 6 kg (13 lb). They get most of their fluids from water in the leaves that they eat but sloths have also been observed drinking directly from rivers.


Dentition and skeleton

Two-toed sloths have a reduced, ever growing dentition, with no incisors or true
canine Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
s, which overall lacks homology with the dental formula of other mammals. Their first tooth is very canine-like in shape and is referred to as a caniniform. It is separated from the other teeth, or molariforms, by a
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
. The molariforms are used specifically for grinding and are mortar and pestle-like in appearance and function. Thus, they can grind food for easier digestibility, which takes the majority of their energy. The dental formula of two-toed sloths is: (unau) Two-toed sloths are unusual among mammals in possessing as few as five
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
, which may be due to mutations in the homeotic genes. All other mammals have seven cervical vertebrae, other than the three-toed sloth and the manatee.


Notes


References

* (1758): Systema naturae perregna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, syonymis, locis. Laurentii : Salvi, 824 pp.


External links


National Geographic: Two-toed sloth profileBristol Zoo: Two-toed slothNational Zoo: Two-toed sloth
{{Taxonbar, from=Q511079 Sloths Mammals of Colombia Taxa named by Carl Linnaeus