Arboreal locomotion is the locomotion 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. [1] 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 tree snails.
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. [1]
Moving along narrow surfaces, such as a branch of a tree, can create special difficulties for 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 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. [2]
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 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. [3]
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. [1] 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.[ citation needed ]
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. In other areas, trees are not close together and animals need to have specific adaptations to jump far distances or glide. [4]
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, [5] [6] they benefit snakes by providing anchor points. [7] [8] [9]
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Arboreal organisms display many specializations for dealing with the mechanical challenges of moving through their habitats. [1]
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. [1] However, some species of lizard have reduced limb size that helps them avoid limb movement being obstructed by impinging branches.
Many arboreal species, such as howler monkeys, green tree pythons, emerald tree boas, chameleons, silky anteaters, spider monkeys, and possums, use prehensile tails to grasp branches. In the spider monkey and crested gecko, the tip of the tail has either a bare patch or adhesive pad, which provides increased friction.
Claws can be used to interact with rough substrates and re-orient the direction of the force the animal applies. This is what allows squirrels 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 is an alternative to claws, which works best on smooth surfaces. Wet adhesion is common in tree frogs and arboreal salamanders, and functions either by suction or by capillary adhesion. Dry adhesion is best typified by the specialized toes of geckos, which use van der Waals forces to adhere to many substrates, even glass.
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.
To control descent, especially down large diameter branches, some arboreal animals such as squirrels 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.
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 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. [1] Size relating to weight affects gliding animals such as the reduced weight per snout-vent length for 'flying' frogs. [10]
Some species of primate, bat, and all species of sloth achieve passive stability by hanging beneath the branch. [1] Both pitching and tipping become irrelevant, as the only method of failure would be losing their grip.
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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.[ citation needed ]
Brachiation is a specialized form of arboreal locomotion, used by primates to move very rapidly while hanging beneath branches. [11] 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 monkeys and gibbons, and is occasionally used by female orangutans. 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 56 km/h (35 mph).[ citation needed ]
To bridge gaps between trees, many animals such as the flying squirrel have adapted membranes, such as patagia for gliding flight. Some animals can slow their descent in the air using a method known as parachuting, such as Rhacophorus (a "flying frog" species) that has adapted toe membranes allowing it to fall more slowly after leaping from trees. [12]
Many species of snake are highly arboreal, and some have evolved specialized musculature for this habitat. [13] While moving in arboreal habitats, snakes move slowly along bare branches using a specialized form of concertina locomotion, [14] but when secondary branches emerge from the branch being moved on, snakes use lateral undulation, a much faster mode. [15] 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. [15]
The earliest known climbing tetrapod is the varanopid amniote Eoscansor from the Late Carboniferous (Pennsylvanian) of North America which is clearly specialised with adaptations for grasping, likely onto tree trunks. [16] Suminia , a anomodont synapsid from Russia dating to the Late Permian, about 260 million years ago, was also likely a specialised climber. [17]
Bipedalism is a form of terrestrial locomotion where a tetrapod moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped, meaning 'two feet'. Types of bipedal movement include walking or running and hopping.
Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes. Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joints than their lizard ancestors, enabling them to swallow prey much larger than their heads. To accommodate their narrow bodies, snakes' paired organs appear one in front of the other instead of side by side, and most have only one functional lung. Some species retain a pelvic girdle with a pair of vestigial claws on either side of the cloaca. Lizards have independently evolved elongate bodies without limbs or with greatly reduced limbs at least twenty-five times via convergent evolution, leading to many lineages of legless lizards. These resemble snakes, but several common groups of legless lizards have eyelids and external ears, which snakes lack, although this rule is not universal.
Brachiation, 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. This form of locomotion is the primary means of locomotion for the small gibbons and siamangs of southeast Asia. Gibbons in particular use brachiation for as much as 80% of their locomotor activities. Some New World monkeys, such as spider monkeys and muriquis, were initially classified as semibrachiators and move through the trees with a combination of leaping and brachiation. Some New World species also practice suspensory behaviors by using their prehensile tail, which acts as a fifth grasping hand. Evidence has shown that the extinct ape Proconsul from the Miocene of East Africa developed an early form of suspensory behaviour, and was therefore referred to as a probrachiator.
Rectilinear locomotion or rectilinear progression is a mode of locomotion most often associated with snakes. In particular, it is associated with heavy-bodied species such as terrestrial African adders, pythons and boas; however, most snakes are capable of it. It is one of at least five forms of locomotion used by snakes, the others being lateral undulation, sidewinding, concertina movement, and slide-pushing. Unlike all other modes of snake locomotion, which include the snake bending its body, the snake flexes its body only when turning in rectilinear locomotion.
Concertina movement is the movement occurring in snakes and other legless organisms that consists of gripping or anchoring with portions of the body while pulling or pushing other sections in the direction of movement.
Animal locomotion, in ethology, is any of a variety of methods that animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flying, hopping, soaring and gliding. There are also many animal species that depend on their environment for transportation, a type of mobility called passive locomotion, e.g., sailing, kiting (spiders), rolling or riding other animals (phoresis).
Horsfeld's tarsier is the only species of tarsier in the genus Cephalopachus. Named by American naturalist Thomas Horsfield, it is also referred to as western tarsier. The species occurs on Borneo, Sumatra and nearby islands and is, like other members of the group, entirely nocturnal.
Knuckle-walking is a form of quadrupedal walking in which the forelimbs hold the fingers in a partially flexed posture that allows body weight to press down on the ground through the knuckles. Gorillas and chimpanzees use this style of locomotion, as do anteaters and platypuses.
Chrysopelea, more commonly known as the flying snake or gliding snake is a genus that belongs to the family Colubridae. Flying snakes are mildly venomous, though the venom is dangerous only to their small prey. Their range is in Southeast Asia, southernmost China, India, and Sri Lanka.
Orthograde is a term derived from Greek ὀρθός, orthos + Latin gradi that describes a manner of walking which is upright, with the independent motion of limbs. Both New and Old World monkeys are primarily arboreal, and they have a tendency to walk with their limbs swinging in parallel to one another. This differs from the manner of walking demonstrated by the apes.
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 or if it can keep a constant speed for a long distance. "Cursorial" is often used to categorize a certain locomotor mode, which is helpful for biologists who examine behaviors of different animals and the way they move in their environment. Cursorial adaptations can be identified by morphological characteristics, physiological characteristics, maximum speed, and how often running is used in life. There is much debate over how to define a cursorial animal specifically. The most accepted definitions include that a cursorial organism could be considered adapted to long-distance running at high speeds or has the ability to accelerate quickly over short distances. Among vertebrates, animals under 1 kg of mass are rarely considered cursorial, and cursorial behaviors and morphology are thought to only occur at relatively large body masses in mammals. There are a few mammals that have been termed "micro-cursors" that are less than 1 kg in mass and have the ability to run faster than other small animals of similar sizes.
A number of animals are capable of aerial locomotion, either by powered flight or by gliding. This trait has appeared by evolution many times, without any single common ancestor. Flight has evolved at least four times in separate animals: insects, pterosaurs, birds, and bats. Gliding has evolved on many more occasions. Usually the development is to aid canopy animals in getting from tree to tree, although there are other possibilities. Gliding, in particular, has evolved among rainforest animals, especially in the rainforests in Asia where the trees are tall and widely spaced. Several species of aquatic animals, and a few amphibians and reptiles have also evolved this gliding flight ability, typically as a means of evading predators.
A facultative biped is an animal that is capable of walking or running on two legs (bipedal), as a response to exceptional circumstances (facultative), while normally walking or running on four limbs or more. In contrast, obligate bipedalism is where walking or running on two legs is the primary method of locomotion. Facultative bipedalism has been observed in several families of lizards and multiple species of primates, including sifakas, capuchin monkeys, baboons, gibbons, gorillas, bonobos and chimpanzees. Several dinosaur and other prehistoric archosaur species are facultative bipeds, most notably ornithopods and marginocephalians, with some recorded examples within sauropodomorpha. Different facultatively bipedal species employ different types of bipedalism corresponding to the varying reasons they have for engaging in facultative bipedalism. In primates, bipedalism is often associated with food gathering and transport. In lizards, it has been debated whether bipedal locomotion is an advantage for speed and energy conservation or whether it is governed solely by the mechanics of the acceleration and lizard's center of mass. Facultative bipedalism is often divided into high-speed (lizards) and low-speed (gibbons), but some species cannot be easily categorized into one of these two. Facultative bipedalism has also been observed in cockroaches and some desert rodents.
Around 350 BCE, Aristotle and other philosophers of the time attempted to explain the aerodynamics of avian flight. Even after the discovery of the ancestral bird Archaeopteryx which lived over 150 million years ago, debates still persist regarding the evolution of flight. There are three leading hypotheses pertaining to avian flight: Pouncing Proavis model, Cursorial model, and Arboreal model.
A limb is a jointed, muscled appendage of a tetrapod vertebrate animal used for weight-bearing, terrestrial locomotion and physical interaction with other objects. The distalmost portion of a limb is known as its extremity. The limbs' bony endoskeleton, known as the appendicular skeleton, is homologous among all tetrapods, who use their limbs for walking, running and jumping, swimming, climbing, grasping, touching and striking.
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 travelling, searching for food and avoiding predators. Different types of suspensory behaviour include brachiation, climbing, and bridging. These mechanisms allow larger species to distribute their weight among smaller branches rather than balancing above these weak supports. Primates and sloths are most commonly seen using these behaviours, however, other animals such as bats may be seen hanging below surfaces to obtain food or when resting.
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 type of gait utilized by limbless animals, some creatures with limbs, such as the salamander, forgo use of their legs in certain environments and exhibit undulatory locomotion. In robotics this movement strategy is studied in order to create novel robotic devices capable of traversing a variety of environments.
Vertical clinging and leaping (VCL) is a type of arboreal locomotion seen most commonly among the strepsirrhine primates and haplorrhine tarsiers. The animal begins at rest with its torso upright and elbows fixed, with both hands clinging to a vertical support, such as the side of a tree or bamboo stalk. To move from one support to another, it pushes off from one vertical support with its hindlimbs, landing on another vertical support after an extended period of free flight. Vertical clinging and leaping primates have evolved a specialized anatomy to compensate for the physical implications of this form of locomotion. These key morphological specializations have been identified in prosimian fossils from as early as the Eocene.
A sucker in zoology is a specialised attachment organ of an animal. It acts as an adhesion device in parasitic worms, several flatworms, cephalopods, certain fishes, amphibians, and bats. It is a muscular structure for suction on a host or substrate. In parasitic annelids, flatworms and roundworms, suckers are the organs of attachment to the host tissues. In tapeworms and flukes, they are a parasitic adaptation for attachment on the internal tissues of the host, such as intestines and blood vessels. In roundworms and flatworms they serve as attachment between individuals particularly during mating. In annelids, a sucker can be both a functional mouth and a locomotory organ. The structure and number of suckers are often used as basic taxonomic diagnosis between different species, since they are unique in each species. In tapeworms there are two distinct classes of suckers, namely "bothridia" for true suckers, and "bothria" for false suckers. In digeneal flukes there are usually an oral sucker at the mouth and a ventral sucker posterior to the mouth. Roundworms have their sucker just in front of the anus; hence it is often called a pre-anal sucker.
The webbed foot is a specialized limb with interdigital membranes (webbings) that aids in aquatic locomotion, present in a variety of tetrapod vertebrates. This adaptation is primarily found in semiaquatic species, and has convergently evolved many times across vertebrate taxa.