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The arthropod leg is a form of jointed of , usually used for . Many of the terms used for arthropod leg segments (called podomeres) are of origin, and may be confused with terms for bones: coxa (meaning hip, : coxae), trochanter, femur (: femora), tibia (: tibiae), tarsus (: tarsi), ischium (: ischia), metatarsus, carpus, dactylus (meaning ), patella (: patellae).

Homologies of leg segments between groups are difficult to prove and are the source of much argument. Some authors posit up to eleven segments per leg for the most recent common ancestor of arthropods but modern arthropods have eight or fewer. It has been argued that the ancestral leg need not have been so complex, and that other events, such as successive loss of function of a , could result in parallel gains of leg segments.

In arthropods, each of the leg segments articulates with the next segment in a and may only bend in one plane. This means that a greater number of segments is required to achieve the same kinds of movements that are possible in vertebrate animals, which have rotational ball-and-socket joints at the base of the fore and hind limbs.

(2009). 9781444309225, John Wiley & Sons. .


Biramous and uniramous
The appendages of arthropods may be either biramous or uniramous. A uniramous limb comprises a single series of segments attached end-to-end. A biramous limb, however, branches into two, and each branch consists of a series of segments attached end-to-end.

The external branch (ramus) of the appendages of is known as the exopod or exopodite, while the internal branch is known as the endopod or endopodite. Other structures aside from the latter two are termed exites (outer structures) and endites (inner structures). Exopodites can be easily distinguished from exites by the possession of internal musculature. The exopodites can sometimes be missing in some crustacean groups ( and ), and they are completely absent in insects.

The legs of and are uniramous. In crustaceans, the first antennae are uniramous, but the second antennae are biramous, as are the legs in most species.

For a time, possession of uniramous limbs was believed to be a shared, , so uniramous arthropods were grouped into a taxon called . It is now believed that several groups of arthropods evolved uniramous limbs independently from ancestors with biramous limbs, so this taxon is no longer used.


Chelicerata
legs differ from those of insects by the addition of two segments on either side of the tibia, the patella between the femur and the tibia, and the metatarsus (sometimes called basitarsus) between the tibia and the tarsus (sometimes called telotarsus), making a total of seven segments.

The tarsus of spiders has claws at the end as well as a hook that helps with web-spinning. Spider legs can also serve sensory functions, with hairs that serve as touch receptors, as well as an organ on the tarsus that serves as a humidity receptor, known as the tarsal organ.

The situation is identical in , but with the addition of a pre-tarsus beyond the tarsus. The claws of the scorpion are not truly legs, but are , a different kind of that is also found in spiders and is specialised for predation and mating.

In , there are no metatarsi or pretarsi, leaving six segments per leg.


Crustacea
The legs of are divided primitively into seven segments, which do not follow the naming system used in the other groups. They are: coxa, basis, ischium, merus, carpus, propodus, and dactylus. In some groups, some of the limb segments may be fused together. The claw (chela) of a lobster or crab is formed by the articulation of the dactylus against an outgrowth of the propodus. Crustacean limbs also differ in being biramous, whereas all other extant arthropods have uniramous limbs.


Myriapoda
(, and their relatives) have seven-segmented walking legs, comprising coxa, trochanter, prefemur, femur, tibia, tarsus, and a tarsal claw. Myriapod legs show a variety of modifications in different groups. In all centipedes, the first pair of legs is modified into a pair of venomous fangs called forcipules. In most millipedes, one or two pairs of walking legs in adult males are modified into sperm-transferring structures called gonopods. In some millipedes, the first leg pair in males may be reduced to tiny hooks or stubs, while in others the first pair may be enlarged.


Insects
Insects and their relatives are hexapods, having six legs, connected to the thorax, each with five components. In order from the body they are the coxa, trochanter, femur, tibia, and tarsus. Each is a single segment, except the tarsus which can be from three to seven segments, each referred to as a tarsomere.

Except in species in which legs have been lost or become vestigial through evolutionary adaptation, adult insects have six legs, one pair attached to each of the three segments of the thorax. They have paired appendages on some other segments, in particular, mouthparts, antennae and , all of which are derived from paired legs on each segment of some common ancestor.

Some insects do however have extra walking legs on their abdominal segments; these extra legs are called . They are found most frequently on the larvae of moths and sawflies. Prolegs do not have the same structure as modern adult insect legs, and there has been a great deal of debate as to whether they are homologous with them.

(1977). 9780412613906, Springer.
Current evidence suggests that they are indeed homologous up to a very primitive stage in their embryological development, but that their emergence in modern insects was not homologous between the and . Such concepts are pervasive in current interpretations of phylogeny.

In general, the legs of larval insects, particularly in the , vary more than in the adults. As mentioned, some have prolegs as well as "true" thoracic legs. Some have no externally visible legs at all (though they have internal rudiments that emerge as adult legs at the final ). Examples include the maggots of or grubs of . In contrast, the larvae of other , such as the and have thoracic legs, but no prolegs. Some insects that exhibit hypermetamorphosis begin their metamorphosis as , specialised, active, legged larvae, but they end their larval stage as legless maggots, for example the .

Among the , the legs of larvae tend to resemble those of the adults in general, except in adaptations to their respective modes of life. For example, the legs of most immature are adapted to scuttling beneath underwater stones and the like, whereas the adults have more gracile legs that are less of a burden during flight. Again, the young of the are called "crawlers" and they crawl around looking for a good place to feed, where they settle down and stay for life. Their later have no functional legs in most species. Among the , the legs of immature specimens are in effect smaller versions of the adult legs.


Fundamental morphology of insect legs
A representative insect leg, such as that of a or , has the following parts, in sequence from most proximal to most distal: coxa, trochanter, femur, tibia, tarsus, and pretarsus.

Associated with the leg itself there are various around its base. Their functions are and have to do with how the leg attaches to the main exoskeleton of the insect. Such sclerites differ considerably between unrelated insects.


Coxa
The coxa is the proximal segment and functional base of the leg. It articulates with the pleuron and associated sclerites of its thoracic segment, and in some species it articulates with the edge of the sternite as well. The homologies of the various basal sclerites are open to debate. Some authorities suggest that they derive from an ancestral subcoxa. In many species, the coxa has two lobes where it articulates with the pleuron. The posterior lobe is the meron which is usually the larger part of the coxa. A meron is well developed in Periplaneta, the Isoptera, Neuroptera and Lepidoptera.


Trochanter
The trochanter articulates with the coxa but usually is attached rigidly to the femur. In some insects, its appearance may be confusing; for example it has two subsegments in the Odonata. In parasitic Hymenoptera, the base of the femur has the appearance of a second trochanter.


Femur
In most insects, the femur is the largest region of the leg; it is especially conspicuous in many insects with saltatorial legs because the typical leaping mechanism is to straighten the joint between the femur and the tibia, and the femur contains the necessary massive musculature.


Tibia
The tibia is the fourth section of the typical insect leg. As a rule, the tibia of an insect is slender in comparison to the femur, but it generally is at least as long and often longer. Near the distal end, there is generally a tibial spur, often two or more. In the , the tibia of the foreleg bears a large apical spur that fits over a semicircular gap in the first segment of the tarsus. The gap is lined with comb-like bristles, and the insect cleans its antennae by drawing them through.


Tarsus
The ancestral tarsus was a single segment and in the extant , and certain insect larvae the tarsus also is single-segmented. Most modern insects have tarsi divided into subsegments (tarsomeres), usually about five. The actual number varies with the , which may be useful for diagnostic purposes. For example, the characteristically have 5-segmented fore- and mid-tarsi, but 4-segmented hind tarsi, whereas the have four tarsomeres on each tarsus.

The distal segment of the typical insect leg is the pretarsus. In the , Protura and many insect larvae, the pretarsus is a single claw. On the pretarsus most insects have a pair of claws ( ungues, singular unguis). Between the ungues, a median unguitractor plate supports the pretarsus. The plate is attached to the of the flexor muscle of the ungues. In the , the parempodia are a symmetrical pair of structures arising from the outside (distal) surface of the unguitractor plate between the claws. It is present in many Hemiptera and almost all . Usually, the parempodia are bristly (setiform), but in a few species they are fleshy.

(1995). 9780801420665, Cornell University Press.
Sometimes the parempodia are reduced in size so as to almost disappear. Above the unguitractor plate, the pretarsus expands forward into a median lobe, the arolium.

Webspinners () have an enlarged basal tarsomere on each of the front legs, containing the -producing glands.

Under their pretarsi, members of the generally have paired lobes or pulvilli, meaning "little cushions". There is a single pulvillus below each unguis. The pulvilli often have an arolium between them or otherwise a median bristle or empodium, meaning the meeting place of the pulvilli. On the underside of the tarsal segments, there frequently are pulvillus-like organs or plantulae. The arolium, plantulae and pulvilli are adhesive organs enabling their possessors to climb smooth or steep surfaces. They all are outgrowths of the exoskeleton and their cavities contain blood. Their structures are covered with tubular tenent hairs, the apices of which are moistened by a glandular secretion. The organs are adapted to apply the hairs closely to a smooth surface so that adhesion occurs through surface molecular forces.Stanislav N Gorb. "Biological attachment devices: exploring nature's diversity for biomimetics Phil. Trans. R. Soc. A 2008; 366(1870): 1557-1574 doi:10.1098/rsta.2007.2172 1471-2962

Insects control the ungues through muscle tension on a long tendon, the "retractor unguis" or "long tendon". In insect models of locomotion and motor control, such as Drosophila (), (), or stick insects (), the long tendon courses through the tarsus and tibia before reaching the femur. Tension on the long tendon is controlled by two muscles, one in the femur and one in the tibia, which can operate differently depending on how the leg is bent. Tension on the long tendon controls the claw, but also bends the tarsus and likely affects its stiffness during walking.


Variations in functional anatomy of insect legs
The typical thoracic leg of an adult insect is adapted for running (), rather than for digging, leaping, swimming, predation, or other similar activities. The legs of most are good examples. However, there are many specialized adaptations, including:

  • The forelegs of (Gryllotalpidae) and some scarab beetle () are adapted to burrowing in earth ().
  • The forelegs of mantidflies (), (Mantodea), and ambush bugs () are adapted to seizing and holding prey in one way, while those of Gyrinidae are long and adapted for grasping food or prey in quite a different way.
  • The forelegs of some butterflies, such as many , are reduced so greatly that only two pairs of functional walking legs remain.
  • In most grasshoppers and crickets (), the hind legs are ; they have heavily bipinnately muscled femora and straight, long tibiae adapted to leaping and to some extent to defence by kicking. (Alticini) also have powerful hind femora that enable them to leap spectacularly.
  • Other beetles with spectacularly muscular hind femora may not be saltatorial at all, but very clumsy; for example, particular species of (Bruchinae) use their swollen hind legs for forcing their way out of the hard-shelled seeds of plants such as in which they grew to adulthood.
  • The legs of the , the and , are adapted for seizing prey that the insects feed on while flying or while sitting still on a plant; they are nearly incapable of using them for walking.
  • The majority of aquatic insects use their legs only for swimming (natatorial), though many species of immature insects swim by other means such as by wriggling, undulating, or expelling water in jets.


Evolution and homology of arthropod legs
The embryonic body segments () of different taxa have diverged from a simple body plan with many similar appendages which are serially homologous, into a variety of body plans with fewer segments equipped with specialised appendages.
(2025). 9780470515662, John Wiley. .
The homologies between these have been discovered by comparing in evolutionary developmental biology.

1antennae (jaws and fangs)antennaeantennae1st antennae
21st legs--2nd antennae
32nd legs1st legsmandiblesmandiblesmandibles (jaws)
43rd legs2nd legs1st maxillae1st maxillae1st maxillae
54th legs3rd legs2nd maxillae2nd maxillae2nd maxillae
65th legs4th legscollum (no legs)1st legs1st legs
76th legs-1st legs2nd legs2nd legs
87th legs-2nd legs3rd legs3rd legs
98th legs-3rd legs-4th legs
109th legs-4th legs-5th legs


See also

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