Vertical clinging and leaping

Last updated
Galago leaping Bushbaby Leaping captured in Multiflash.jpg
Galago leaping

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. [1] Vertical clinging and leaping primates have evolved a specialized anatomy to compensate for the physical implications of this form of locomotion. [2] [3] These key morphological specializations have been identified in prosimian fossils from as early as the Eocene. [4] [5] [6]

Contents

Vertical clinging and leaping primates

Indri indri clinging to a tree Indri andasibe mantadia Nationalpark 2019-10-16 3.jpg
Indri indri clinging to a tree
Propithecus verreauxi leaping from tree Sifaka leaping.jpg
Propithecus verreauxi leaping from tree
P. verreauxi employs a similar form of locomotion on the ground Verreaux's Sifaka GIF.GIF
P. verreauxi employs a similar form of locomotion on the ground

Vertical clinging and leaping has only been observed in primates, and primarily in prosimians. [5] A few anthropoids have been observed to engage in this behavior, such as Pithecia, [7] and are similar in morphological adaptations to prosimians who rely on VCL.

Vertical clinging and leaping strepsirrhine taxa include: [5]

Vertical clinging and leaping haplorrhine taxa include: [5]

Variations in posture

Variations in mid-flight posture Types of Vertical Clinging Leaps, Hunt et al 1996.png
Variations in mid-flight posture

Variations in mid-flight posture

The variations in vertical leaping can be classified into three types based on differences in mid-flight posture: [1]

(A) During the stretched-out vertical cling leap, the femur is extended. (ex. Indriidae)
(B) During the curled-up vertical cling leap, the torso is sub-orthograde with all limbs positioned in front of the body. (ex. Galago )
(C) During the limbs-down vertical cling leap, the torso is pronograde, with all limbs positioned underneath the body. (ex. Cheirogaleus )

Variations in clinging posture

There is an array of clinging postures that primates use while foraging and resting in trees. These postures include sitting, bipedal standing, squatting, forelimb suspension, forelimb-hindlimb suspension and more. Stabilization is key to how a primate positions themselves while clinging. The placement of the torso depends on the weight distribution between the hands, feet, and tail. [1] Vertical clinging and leaping primates may also use their tails to grip adjacent branches in order to stabilize their positional modes.

Morphological specializations

Locomotion and movement are the main contributing factors to primate's body shape and structure, thus the anatomy of vertical climbers and leapers is highly specialized to enable them to move effectively within their arboreal habitat. Features found in the hips, knees, feet, hands, arms, and tails are specialized to facilitate climbing and leaping in primates, and aids in other postures such as tail suspensions and foot hanging. With the features described, these primates can move through trees efficiently and obtain food easily. Body size also has a direct correlation to how fast and how far a primate can leap. [2] There are other musculoskeletal specializations that differentiate VCL dependent primates, such as their hip joints and muscles. Leaping primates have more proximally positioned lesser and third trochanters, and have relatively large muscles for hip extensors, knee extensors or ankle plantar flexors. [10] This is an adaptation that has come from the increased need for hind limb propulsion.

The evolution of key features of VCL primate anatomy can be explained by analyzing how they leap. Longer limbs allow for enhanced locomotive abilities. Leaping requires primates to achieve a certain height and distance and then land on a vertical support. Longer hindlimbs therefore allow more time and distance for acceleration and takeoff, so longer and stronger femurs evolved because they increase the distance over which acceleration occurs. [3] Additionally, during landing, longer limbs help with the deceleration of the body, since hindlimb length is directly proportional to the time required for landing. Therefore, longer limbs provide more time for deceleration and can prevent injury after landing with high velocity. [2]

Launch point considerations

Launch point branch diameter

Launch angles vary depending on height and range needs Launch angles.png
Launch angles vary depending on height and range needs

Before leaping, primates must fundamentally consider the distance they are traveling and the stability in their launch point and landing point. A large branch is preferable to a small branch as a launch point, as the wider diameter provides a sturdier base for the jump and also reduces the energy expended during takeoff. In fact, tarsiers have been observed climb to lower heights in order to leap from and land on larger diameter branches. However, this also limits the height of the leap, as the primate must be able to land on a branch that is large enough to stabilize the animal and also minimize the energy necessary for maintaining balance during landing. [11]

Launch point angle

Launch angle does not appear to be species-specific, and the range of angles varies between 30 and 70 degrees. The optimum ballistic trajectory angle would be 45 degrees, [11] but often the animal will need to cover more horizontal than vertical distance, or vice versa. In these cases, a trajectory other than 45 degrees would be more beneficial to give more height or more range (see photo), so vertical clinging and leaping animals adjust their launch angles accordingly to compensate for this. [12]

Fossil evidence

Locomotor adaptations in Eocene prosimians

Ida, an early Eocene primate fossil who provided the first evidence of vertical clinging and leaping Ida, One of the Oldest and Most Complete Primate Fossils from the early Eocene period.jpg
Ida, an early Eocene primate fossil who provided the first evidence of vertical clinging and leaping

There is evidence of vertical clinging and leaping adaptations in the postcranial skeleton of early Eocene fossils of prosimians. Lacking any common quadrupedal features, the postcranial bones and fossils of the Eocene prosimians most closely resemble the specializations of vertical clingers and leapers. Common morphological elements identified in these early fossils include a high phalangeal index, low intermembral index, femurs with cylindrical head and high, narrow patellar grooves, and fused fibula and tibia. [4] The high phalangeal index is the elongation of the phalangeal finger bones, which helps vertical clingers who require increased prehensile power in their hands to support their large bulk. The low intermebral index is the total length of the humerus and radius in percentage of total length of the femur and tibia. The features of the femur is part of an evolution of elongated femurs that helps the primates' extreme range of flexion and extension, which allows them to perform their leaps. Thus, VCL seems to be the only known locomotor adaptation in the Eocene primates, meaning it is one of, if not the earliest, locomotor adaptations. [5] [2]

Early Eocene primate fossil: Ida

The earliest evidence of VCL can be found in an early Eocene adolescent primate, named Ida. She exhibited legs that were longer than her arms, suggesting that leaping was a key aspect of her locomotion. Her death is hypothesized to be linked to her vicinity to the Messel Pit, near Frankfurt, Germany. The hot magma from the earth mixing with the underground water tables led to explosions that released toxic gasses. Ida was found with a broken wrist, and it is believed that due to this, she could not leap or cling to the higher tree branches, and therefore had to stay lower to the ground where she came into contact with the toxic gasses and died. [6]

Related Research Articles

Tarsier Family of dry nosed primates

Tarsiers are haplorrhine primates of the family Tarsiidae, which is itself the lone extant family within the infraorder Tarsiiformes. Although the group was once more widespread, all of its species living today are found in the islands of Southeast Asia, specifically the Philippines, Malaysia, Indonesia, and Brunei.

Strepsirrhini Suborder of primates which includes lemurs, galagos, pottos and lorises

Strepsirrhini or Strepsirhini is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Collectively they are referred to as strepsirrhines. Also belonging to the suborder are the extinct adapiform primates which thrived during the Eocene in Europe, North America, and Asia, but disappeared from most of the Northern Hemisphere as the climate cooled. Adapiforms are sometimes referred to as being "lemur-like", although the diversity of both lemurs and adapiforms does not support this comparison.

Brachiation Form of arboreal locomotion involving swinging by the arm

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 Milocene of East Africa developed an early form of suspensory behaviour, and was therefore referred to as a probrachiator.

Forelimb

A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. In bipedal animals with an upright posture, the term upper limb is often used.

Horsfields tarsier Species of primate

Horsfield's tarsier, also known as the western tarsier, is the only species of tarsier in the genus Cephalopachus. It occurs on Borneo, Sumatra and nearby islands and is, like other members of the group, entirely nocturnal.

Knuckle-walking

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.

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.

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 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 is 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.

<i>Akidolestes</i> extinct genus of mammals

Akidolestes is an extinct genus of mammals from the family spalacotheriid.

<i>Lagerpeton</i> genus of lagerpetid from the Late Triassic period

Lagerpeton is a genus of lagerpetid avemetatarsalian, comprising a single species, L. chanarensis. First described from the Chañares Formation of Argentina by A. S. Romer in 1971, Lagerpeton's anatomy is somewhat incompletely known, with fossil specimens accounting for the pelvic girdle, hindlimbs, posterior presacral, sacral and anterior caudal vertebrae. Skull and shoulder material has also been described.

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. 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.

<i>Necrolemur</i> Extinct genus of primates

Necrolemur is a small bodied omomyid with body mass estimations ranging from 114 grams to 346 grams. Necrolemur’s teeth feature broad basins and blunt cusps, suggesting their diet consisted of mostly soft fruit, though examination of microwear patterns suggests that populations from lower latitudes also consumed insects and gums. While they do not sport a true tooth comb like modern lemurs, microwear patterns on their lower incisors suggest they engaged in fur grooming behaviors. Like tarsiers, Necrolemur had large, front-facing, close set eyes and were likely nocturnal. Analysis of cranial and postcranial fossils by paleontologists suggest members of the family Omomyidae, including the genus Necrolemur, possessed highly specialized adaptations for leaping.

Shoshonius is an extinct genus of omomyid primate that lived during the Eocene. Specimens identified as Shoshonius have been found exclusively in central Wyoming and the genus currently includes two species, Shoshonius cooperi, described by Granger in 1910, and Shoshonius bowni, described by Honey in 1990.

Arboreal locomotion Movement of animals through trees

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. Furthermore, many of these same principles may be applied to climbing without trees, such as on rock piles or mountains.

The evolution of human bipedalism, which began in primates about four million years ago, or as early as seven million years ago with Sahelanthropus, or about 12 million years ago with Danuvius guggenmosi, has led to morphological alterations to the human skeleton including changes to the arrangement and size of the bones of the foot, hip size and shape, knee size, leg length, and the shape and orientation of the vertebral column. The evolutionary factors that produced these changes have been the subject of several theories.

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.

Adapis is an extinct adapiform primate from the Eocene of Europe. While this genus has traditionally contained five species, recent research has recognized at least six morphotypes that may represent distinct species. Adapis holds the title of the first Eocene primate ever discovered. In 1821, Georges Cuvier, who is considered to be the founding father of paleontology, discovered Adapis in fissure fillings outside of Paris, France. Given it's timing and appearance in the fossil record, Cuvier did not recognize the primate affinities of Adapis and first described it as a small extinct pachyderm; only later in the 19th century was Adapis identified as a primate.

<i>Afrotarsius</i> Extinct genus of primates

Afrotarsius is a primate found in the Paleogene of Africa.

<i>Archicebus</i> Genus of fossil primates that lived in the early Eocene forests (~55 million years ago

Archicebus is a genus of fossil primates that lived in the early Eocene forests of what is now Jingzhou in the Hubei Province in central China, discovered in 2003. The only known species, A. achilles, was a small primate, estimated to weigh approximately 20–30 grams (0.71–1.06 oz), and is the only known member of the family Archicebidae. When discovered, it was the oldest fossil haplorhine primate skeleton found, appearing to be most closely related to tarsiers and the fossil omomyids, although A. achilles is suggested to have been diurnal whereas tarsiers are nocturnal. Resembling tarsiers and simians, it was a haplorhine primate, and it also may have resembled the last common ancestor of all haplorhines as well as the last common ancestor of all primates. Its discovery further supports the hypothesis that primates originated in Asia, not in Africa.

Danuvius guggenmosi is an extinct species of great ape that lived 11.6 million years ago during the Middle–Late Miocene in southern Germany. It is the sole member of the genus Danuvius. The area at this time was probably a woodland with a seasonal climate. A male specimen was estimated to have weighed about 31 kg (68 lb), and two females 17 and 19 kg. Both genus and species were described in November 2019.

References

  1. 1 2 3 Hunt, Kevin D.; Cant, John G. H.; Gebo, Daniel L.; Rose, Michael D.; Walker, Suzanne E.; Youlatos, Dionisios (1996). "Standardized descriptions of primate locomotor and postural modes". Primates. 37 (4): 363–387. doi:10.1007/bf02381373. ISSN   0032-8332. S2CID   37235291.
  2. 1 2 3 4 Demes, B.; Jungers, W.L.; Fleagle, J.G.; Wunderlich, R.E.; Richmond, B.G.; Lemelin, P. (1996). "Body size and leaping kinematics in Malagasy vertical clingers and leapers" (PDF). Journal of Human Evolution. 31 (4): 367–388. doi:10.1006/jhev.1996.0066. ISSN   0047-2484. S2CID   84810171. Archived from the original (PDF) on 2020-06-21.
  3. 1 2 Ryan, T.M.; Ketcham, R.A. (2005). "Angular orientation of trabecular bone in the femoral head and its relationship to hip joint loads in leaping primates". Journal of Morphology. 265 (3): 249–263. doi:10.1002/jmor.10315. PMID   15690365. S2CID   28975464.
  4. 1 2 Ni, X.; Gebo, D.L.; Dagosto, M.; Meng, J.; Tafforeau, P.; Flynn, J.J.; Beard, K.C. (2013). "The oldest known primate skeleton and early haplorhine evolution". Nature. 498 (7452): 60–64. Bibcode:2013Natur.498...60N. doi:10.1038/nature12200. PMID   23739424. S2CID   4321956.
  5. 1 2 3 4 5 Napier, J.R.; Walker, A.C. (1967). "Vertical Clinging and Leaping – A Newly Recognized Category of Locomotor Behaviour of Primates". Folia Primatologica. 6 (3–4): 204–219. doi:10.1159/000155079. PMID   6070682.
  6. 1 2 "Who Was Ida?". National Geographic. 2011-10-24. Retrieved 2018-11-02.
  7. 1 2 3 Walker, S.E. (2005). "Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela". American Journal of Primatology. 66 (4): 369–387. doi:10.1002/ajp.20162. PMID   16104032. S2CID   15006017.
  8. Kinzey, W.G.; Rosenberger, A.L.; Ramirez, M. (1975). "Vertical clinging and leaping in a neotropical anthropoid". Nature. 255 (5506): 327–328. Bibcode:1975Natur.255..327K. doi:10.1038/255327a0. PMID   805378. S2CID   35759389.
  9. 1 2 Garber, P.A. (1992). "Vertical clinging, small body size, and the evolution of feeding adaptations in the Callitrichinae". American Journal of Physical Anthropology. 88 (4): 469–482. doi:10.1002/ajpa.1330880404. PMID   1503119.
  10. Baker, Jeremy J.; Searight, Katherine J.; Stump, Madeliene Atzeva; Kehrer, Matthew B.; Shanafelt, Colleen; Graham, Eric; Smith, Timothy D. (2011-07-19). "Hip Anatomy and Ontogeny of Lower Limb Musculature in Three Species of Nonhuman Primates". Anatomy Research International. 2011: 580864. doi: 10.1155/2011/580864 . ISSN   2090-2743. PMC   3335645 . PMID   22567295.
  11. 1 2 Crompton, R.H.; Blanchard, M.L.; Coward, S.; Alexander, R.M.; Thorpe, S.K. (2010-07-01). "Vertical Clinging and Leaping Revisited: Locomotion and Habitat Use in the Western Tarsier, Tarsius bancanus Explored Via Loglinear Modeling". International Journal of Primatology. 31 (6): 958–979. doi:10.1007/s10764-010-9420-8. S2CID   45884124.
  12. Sussman, R.W. (2003). "Chapter 3: Lorisiformes". Primate Ecology and Social Structure. Pearson Custom Publishing. p. 78. ISBN   978-0-536-74363-3.