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Arboreal locomotion is the
locomotion Locomotion means the act or ability of something to transport or move itself from place to place. Locomotion may refer to: Motion * Motion (physics) * Robot locomotion, of man-made devices By environment * Aquatic locomotion * Flight * Locomoti ...
of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose numerous mechanical challenges to animals moving through them and lead to a variety of anatomical, behavioral and ecological consequences as well as variations throughout different species.Cartmill, M. (1985). Climbing. In ''Functional Vertebrate Morphology'', eds. M. Hildebrand D. M. Bramble K. F. Liem and D. B. Wake, pp. 73–88. Cambridge: Belknap Press. Furthermore, many of these same principles may be applied to climbing without trees, such as on rock piles or mountains. Some animals are exclusively arboreal in habitat, such as the
tree snail Tree snail is a common name that is applied to various kinds of tropical air-breathing land snails, pulmonate gastropod mollusks that have shells, and that live in trees, in other words, are exclusively arboreal in habitat. Some other species of a ...
.


Biomechanics

Arboreal habitats pose numerous mechanical challenges to animals moving in them, which have been solved in diverse ways. These challenges include moving on narrow branches, moving up and down inclines, balancing, crossing gaps, and dealing with obstructions.


Diameter

Moving along narrow surfaces, such as a branch of a tree, can create special difficulties to animals who are not adapted to deal with balancing on small diameter substrates. During locomotion on the ground, the location of the
center of mass In physics, the center of mass of a distribution of mass in space (sometimes referred to as the balance point) is the unique point where the weighted relative position of the distributed mass sums to zero. This is the point to which a force may ...
may swing from side to side. But during arboreal locomotion, this would result in the center of mass moving beyond the edge of the branch, resulting in a tendency to topple over and fall. Not only do some arboreal animals have to be able to move on branches of varying diameter, but they also have to eat on these branches, resulting in the need for the ability to balance while using their hands to feed themselves. This resulted in various types of grasping such as pedal grasping in order to clamp themselves onto small branches for better balance.


Incline

Branches are frequently oriented at an angle to gravity in arboreal habitats, including being vertical, which poses special problems. As an animal moves up an inclined branch, it must fight the force of gravity to raise its body, making the movement more difficult. To get past this difficulty many animals have to grasp the substrate with all four limbs and increase the frequency of their
gait Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency. Di ...
sequence. Conversely, as the animal descends, it must also fight gravity to control its descent and prevent falling. Descent can be particularly problematic for many animals, and highly arboreal species often have specialized methods for controlling their descent. One way animals prevent falling while descending is to increase the amount of contact their limbs are making with the substrate to increase friction and braking power.


Balance

Due to the height of many branches and the potentially disastrous consequences of a fall, balance is of primary importance to arboreal animals. On horizontal and gently sloped branches, the primary problem is tipping to the side due to the narrow base of support. The narrower the branch, the greater the difficulty in balancing a given animal faces. On steep and vertical branches, tipping becomes less of an issue, and pitching backwards or slipping downwards becomes the most likely failure. In this case, large-diameter branches pose a greater challenge since the animal cannot place its forelimbs closer to the center of the branch than its hindlimbs.


Crossing gaps

Some arboreal animals need to be able to move from tree to tree in order to find food and shelter. To be able to get from tree to tree, animals have evolved various adaptations. In some areas trees are close together and can be crossed by simple
brachiation Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb ...
. In other areas trees are not close together and animals need to have specific adaptations to jump far distances or glide.


Obstructions

Arboreal habitats often contain many obstructions, both in the form of branches emerging from the one being moved on and other branches impinging on the space the animal needs to move through. These obstructions may impede locomotion, or may be used as additional contact points to enhance it. While obstructions tend to impede limbed animals, they benefit snakes by providing anchor points.


Anatomical specializations

Arboreal organisms display many specializations for dealing with the mechanical challenges of moving through their habitats.


Limb length

Arboreal animals frequently have elongated limbs that help them cross gaps, reach fruit or other resources, test the firmness of support ahead, and in some cases, to
brachiate Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb. ...
. However, some species of lizard have reduced limb size that helps them avoid limb movement being obstructed by impinging branches.


Prehensile tails

Many arboreal species, such as tree porcupines,
green tree python The green tree python (''Morelia viridis'') is a species of snake in the family Pythonidae. The species is native to New Guinea, some islands in Indonesia, and the Cape York Peninsula in Australia. First described by Hermann Schlegel in 1872, it ...
s,
emerald tree boa The emerald tree boa (''Corallus caninus'')Mehrtens JM. 1987. ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . is a boa species found in the rainforests of South America. Since 2009 the species ''Corallus batesii' ...
s,
chameleon Chameleons or chamaeleons (family Chamaeleonidae) are a distinctive and highly specialized clade of Old World lizards with 202 species described as of June 2015. The members of this family are best known for their distinct range of colors, bein ...
s,
silky anteater The silky anteater, also known as the pygmy anteater, has traditionally been considered a single species of anteater, ''Cyclopes didactylus'', in the genus ''Cyclopes'', the only living genus in the family Cyclopedidae. Found in southern Mexico, ...
s,
spider monkey Spider monkeys are New World monkeys belonging to the genus ''Ateles'', part of the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil. The g ...
s, and
possums Possum may refer to: Animals * Phalangeriformes, or possums, any of a number of arboreal marsupial species native to Australia, New Guinea, and Sulawesi ** Common brushtail possum (''Trichosurus vulpecula''), a common possum in Australian urban a ...
, use
prehensile tail A prehensile tail is the tail of an animal that has adapted to grasp or hold objects. Fully prehensile tails can be used to hold and manipulate objects, and in particular to aid arboreal creatures in finding and eating food in the trees. If the ta ...
s to grasp branches. In the
spider monkey Spider monkeys are New World monkeys belonging to the genus ''Ateles'', part of the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil. The g ...
and
crested gecko The crested gecko or eyelash gecko (''Correlophus ciliatus'') is a species of gecko native to southern New Caledonia. In 1866, the crested gecko was species description, described by a French zoologist named Alphonse Guichenot. This species was t ...
, the tip of the tail has either a bare patch or adhesive pad, which provide increased friction.


Clawed

Claws can be used to interact with rough substrates and re-orient the direction of forces the animal applies. This is what allows
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s to climb tree trunks that are so large as to be essentially flat, from the perspective of such a small animal. However, claws can interfere with an animal's ability to grasp very small branches, as they may wrap too far around and prick the animal's own paw.


Adhesion

Adhesion is an alternative to claws, which works best on smooth surfaces. Wet adhesion is common in
tree frog A tree frog (or treefrog) is any species of frog that spends a major portion of its lifespan in trees, known as an arboreal state. Several lineages of frogs among the Neobatrachia have given rise to treefrogs, although they are not closely relat ...
s and
arboreal salamander The arboreal salamander (''Aneides lugubris'') is a species of climbing salamander. An insectivore, it is native to California and Baja California, where it is primarily associated with oak and sycamore woodlands, and thick chaparral. Descrip ...
s, and functions either by suction or by capillary adhesion. Dry adhesion is best typified by the specialized toes of
gecko Geckos are small, mostly carnivorous lizards that have a wide distribution, found on every continent except Antarctica. Belonging to the infraorder Gekkota, geckos are found in warm climates throughout the world. They range from . Geckos ar ...
s, which use
van der Waals force In molecular physics, the van der Waals force is a distance-dependent interaction between atoms or molecules. Unlike ionic or covalent bonds, these attractions do not result from a chemical electronic bond; they are comparatively weak and th ...
s to adhere to many substrates, even glass.


Gripping

Frictional gripping is used by primates, relying upon hairless fingertips. Squeezing the branch between the fingertips generates a frictional force that holds the animal's hand to the branch. However, this type of grip depends upon the angle of the frictional force, thus upon the diameter of the branch, with larger branches resulting in reduced gripping ability. Animals other than primates that use gripping in climbing include the chameleon, which has mitten-like grasping feet, and many birds that grip branches in perching or moving about.


Reversible feet

To control descent, especially down large diameter branches, some arboreal animals such as
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s have evolved highly mobile ankle joints that permit rotating the foot into a 'reversed' posture. This allows the claws to hook into the rough surface of the bark, opposing the force of gravity.


Low center of mass

Many arboreal species lower their center of mass to reduce pitching and toppling movement when climbing. This may be accomplished by postural changes, altered body proportions, or smaller size.


Small size

Small size provides many advantages to arboreal species: such as increasing the relative size of branches to the animal, lower center of mass, increased stability, lower mass (allowing movement on smaller branches), and the ability to move through more cluttered habitat. Size relating to weight affects gliding animals such as the reduced weight per snout-vent length for 'flying' frogs.


Hanging under perches

Some species of
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
,
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
, and all species of
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 li ...
achieve passive stability by hanging beneath the branch. Both pitching and tipping become irrelevant, as the only method of failure would be losing their grip.


Behavioral specializations

Arboreal species have behaviors specialized for moving in their habitats, most prominently in terms of posture and gait. Specifically, arboreal mammals take longer steps, extend their limbs further forwards and backwards during a step, adopt a more 'crouched' posture to lower their center of mass, and use a diagonal sequence
gait Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency. Di ...
.


Ecological consequences

Arboreal locomotion allows animals access to different resources, depending upon their abilities. Larger species may be restricted to larger-diameter branches that can support their weight, while smaller species may avoid competition by moving in the narrower branches.


Climbing without trees

Many animals climb in other habitats, such as in rock piles or mountains, and in those habitats, many of the same principles apply due to inclines, narrow ledges, and balance issues. However, less research has been conducted on the specific demands of locomotion in these habitats. Perhaps the most exceptional of the animals that move on steep or even near vertical rock faces by careful balancing and leaping are the various types of mountain dwelling caprid such as the
Barbary sheep The Barbary sheep (''Ammotragus lervia''), also known as aoudad (pronounced ɑʊdæd is a species of caprine native to rocky mountains in North Africa. While this is the only species in genus ''Ammotragus'', six subspecies have been describ ...
,
markhor The markhor (''Capra falconeri'') is a large ''Capra'' species native to Central Asia, the Karakoram, and the Himalayas. It is listed on the IUCN Red List as Near Threatened since 2015. The markhor is the national animal of Pakistan, where it ...
,
yak The domestic yak (''Bos grunniens''), also known as the Tartary ox, grunting ox or hairy cattle, is a species of long-haired domesticated cattle found throughout the Himalayan region of the Indian subcontinent, the Tibetan Plateau, Kachin Sta ...
,
ibex An ibex (plural ibex, ibexes or ibices) is any of several species of wild goat (genus ''Capra''), distinguished by the male's large recurved horns, which are transversely ridged in front. Ibex are found in Eurasia, North Africa and East Africa. ...
,
tahr Tahrs ( , ) or tehrs ( ) are large artiodactyl ungulates related to goats and sheep. There are three species, all native to Asia. Previously thought to be closely related to each other and placed in a single genus, ''Hemitragus'', genetic stud ...
,
rocky mountain goat The mountain goat (''Oreamnos americanus''), also known as the Rocky Mountain goat, is a hoofed mammal endemic to mountainous areas of western North America. A subalpine to alpine species, it is a sure-footed climber commonly seen on cliffs an ...
, and
chamois The chamois (''Rupicapra rupicapra'') or Alpine chamois is a species of goat-antelope native to mountains in Europe, from west to east, including the Alps, the Dinarides, the Tatra and the Carpathian Mountains, the Balkan Mountains, the Ril ...
. Their adaptations may include a soft rubbery pad between their hooves for grip, hooves with sharp keratin rims for lodging in small footholds, and prominent dew claws. The
snow leopard The snow leopard (''Panthera uncia''), also known as the ounce, is a Felidae, felid in the genus ''Panthera'' native to the mountain ranges of Central Asia, Central and South Asia. It is listed as Vulnerable species, Vulnerable on the IUCN Red ...
, being a predator of such mountain caprids, also has spectacular balance and leaping abilities; being able to leap up to ≈17m (~50 ft). Other balancers and leapers include the
mountain zebra The mountain zebra (''Equus zebra'') is a zebra species in the family Equidae, native to southwestern Africa. There are two subspecies, the Cape mountain zebra (''E. z. zebra'') found in South Africa and Hartmann's mountain zebra (''E. z. hartman ...
,
mountain tapir The mountain tapir, also known as the Andean tapir or woolly tapir (''Tapirus pinchaque'') is the smallest of the four widely recognized species of tapir. It is the only one to live outside of tropical rainforests in the wild. It is most easily d ...
, and
hyrax Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es.


Brachiation

Brachiation is a specialized form of arboreal locomotion, used by primates to move very rapidly while hanging beneath branches. Arguably the epitome of arboreal locomotion, it involves swinging with the arms from one handhold to another. Only a few species are brachiators, and all of these are primates; it is a major means of locomotion among
spider monkey Spider monkeys are New World monkeys belonging to the genus ''Ateles'', part of the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil. The g ...
s and
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India ...
s, and is occasionally used by female
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s. Gibbons are the experts of this mode of locomotion, swinging from branch to branch distances of up to 15 m (50 ft), and traveling at speeds of as much as .


Gliding and parachuting

To bridge gaps between trees, many animals such as the flying squirrel have adapted membranes, such as
patagia The patagium (plural: patagia) is a membranous Animal body, body part that assists an animal in obtaining lift (force), lift when gliding flight, gliding or flight. The structure is found in extant taxon, extant and extinct groups of flying and g ...
for
gliding flight Gliding flight is heavier-than-air flight without the use of thrust; the term volplaning also refers to this mode of flight in animals. It is employed by gliding animals and by aircraft such as gliders. This mode of flight involves flying a s ...
. Some animals can slow their descent in the air using a method known as parachuting, such as ''
Rhacophorus ''Rhacophorus'' is a genus of frogs in the shrub frog family (biology), family (Rhacophoridae) and the related Hylidae make up the true tree frogs. They are found in India, Japan, Madagascar, Africa, and Southeast Asia. Over 40 species are curren ...
'' (a "
flying frog A flying frog (also called a gliding frog) is a frog that has the ability to achieve gliding flight. This means it can descend at an angle less than 45° relative to the horizontal. Other nonflying arboreal frogs can also descend, but only at angle ...
" species) that has adapted toe membranes allowing it to fall more slowly after leaping from trees.


Limbless climbing

Many species of snake are highly arboreal, and some have evolved specialized musculature for this habitat. While moving in arboreal habitats, snakes move slowly along bare branches using a specialized form of concertina locomotion, but when secondary branches emerge from the branch being moved on, snakes use
lateral undulation Undulatory locomotion is the type of motion characterized by wave-like movement patterns that act to propel an animal forward. Examples of this type of gait include crawling in snakes, or swimming in the lamprey. Although this is typically the ...
, a much faster mode. As a result, snakes perform best on small perches in cluttered environments, while limbed organisms seem to do best on large perches in uncluttered environments.


Arboreal animals

Many species of animals are arboreal, far too many to list individually. This list is of prominently or predominantly arboreal species and higher taxa.


Evolutionary history

The earliest known climbing
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
is the
varanopid Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an ''Archaeothyris''-like synapsid in the Late Carbonifero ...
amniote ''
Eoscansor ''Eoscansor'' (EE-oh-SKAN-sor; meaning "dawn climber") is a species of small varanopid amniote that lives from the upper Pennsylvanian subperiod in northern New Mexico, United States 305 million years ago. The species ''Eoscansor cobrensis'' was ...
'' from the Late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
( Pennsylvanian) of North America which is clearly specialised with adaptations for grasping, likely onto tree trunks. ''
Suminia ''Suminia'' is an extinct genus of basal anomodont that lived during the Tatarian age of the late Permian, spanning approximately from 268-252 Ma.Rybczynski N. 2000. Cranial anatomy and phylogenetic position of Suminia getmanovi, a basal anomodon ...
'', a
anomodont Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
from Russia dating to the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
, about 260 million years ago, was also likely a specialised climber.


See also

*
Suspensory behavior Suspensory behaviour is a form of arboreal locomotion or a feeding behavior that involves hanging or suspension of the body below or among tree branches. This behavior enables faster travel while reducing path lengths to cover more ground when trave ...
*
Cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often u ...
*
Fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric eviden ...
*


References


Sources

* * Cartmill, M. (1974). Pads and claws in arboreal locomotion. In Primate Locomotion, (ed. F. A. J. Jenkins), pp. 45–83. New York: Academic Press. * * * * {{locomotion Habitat Trees