Arthropod Coxa
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The arthropod leg is a form of jointed
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
of
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s, usually used for
walking Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an 'inverted pendulum' gait in which the body vaults o ...
. Many of the terms used for arthropod leg segments (called podomeres) are of
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
origin, and may be confused with terms for bones: ''coxa'' (meaning
hip In vertebrate anatomy, hip (or "coxa"Latin ''coxa'' was used by Celsus in the sense "hip", but by Pliny the Elder in the sense "hip bone" (Diab, p 77) in medical terminology) refers to either an anatomical region or a joint. The hip region is ...
, plural ''coxae''), ''trochanter'', ''femur'' (plural ''femora''), ''tibia'' (plural ''tibiae''), ''tarsus'' (plural ''tarsi''), ''ischium'' (plural ''ischia''), ''metatarsus'', ''carpus'', ''dactylus'' (meaning
finger A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers ( Pentadactyly). Chambers ...
), ''patella'' (plural ''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 In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
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 ''Hox''-gene, could result in
parallel Parallel is a geometric term of location which may refer to: Computing * Parallel algorithm * Parallel computing * Parallel metaheuristic * Parallel (software), a UNIX utility for running programs in parallel * Parallel Sysplex, a cluster of ...
gains of leg segments. In arthropods, each of the leg segments articulates with the next segment in a
hinge joint A hinge joint (ginglymus or ginglymoid) is a bone joint in which the articular surfaces are molded to each other in such a manner as to permit motion only in one plane. According to one classification system they are said to be uniaxial (having ...
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 joint The ball-and-socket joint (or spheroid joint) is a type of synovial joint in which the ball-shaped surface of one rounded bone fits into the cup-like depression of another bone. The distal bone is capable of motion around an indefinite number of ...
s at the base of the fore and hind limbs.


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
crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
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 (
amphipod Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descr ...
s and
isopod Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
s), and they are completely absent in insects. The legs of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and
myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
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, derived character, so uniramous arthropods were grouped into a taxon called
Uniramia Uniramia (''uni'' – one, ''ramus'' – branch, i.e. single-branches) is a group within the arthropods. In the past this group included the Onychophora, which are now considered a separate category. The group is currently used in a narrower sen ...
. 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

Arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroon ...
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 have 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
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always end ...
s, but with the addition of a pre-tarsus beyond the tarsus. The claws of the scorpion are not truly legs, but are
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") and ...
s, a different kind of
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
that is also found in spiders and is specialised for predation and mating. In ''
Limulus ''Limulus'' is a genus of horseshoe crab, with one extant species, the Atlantic horseshoe crab (''Limulus polyphemus''). One fossil species is currently assigned to the genus though several other species have been named, which have since been as ...
'', there are no metatarsi or pretarsi, leaving six segments per leg.


Crustacea

The legs of
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s 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 Chela may refer to: * ''Chela'' (fish), a genus of small minnow-type fish in the Cyprinid family * Chela (organ), a pincer-like organ terminating certain limbs of some arthropods such as crabs * Chela (meteorite), a meteorite fall of 1988 in Tanz ...
) 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

Myriapod Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
s (
millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
s,
centipede Centipedes (from New Latin , "hundred", and Latin , " foot") are predatory arthropods belonging to the class Chilopoda (Ancient Greek , ''kheilos'', lip, and New Latin suffix , "foot", describing the forcipules) of the subphylum Myriapoda, an ...
s 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 Gonopods are specialized appendages of various arthropods used in reproduction or egg-laying. In males, they facilitate the transfer of sperm from male to female during mating, and thus are a type of intromittent organ. In crustaceans and milli ...
. 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 The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
, 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 cerci, all of which are derived from paired legs on each segment of some
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
. Some
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l insects do however have extra walking legs on their abdominal segments; these extra legs are called
proleg A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few other types of in ...
s. 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. 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
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
and
Symphyta Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
. Such concepts are pervasive in current interpretations of phylogeny. In general the legs of larval insects, particularly in the
Endopterygota Endopterygota (from Ancient Greek ''endon'' 'inner' + ''pterón'' 'wing' + New Latin ''-ota'' 'having'), also known as Holometabola, is a superorder of insects within the infraclass Neoptera that go through distinctive larval, pupal, and adult s ...
, 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
ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
). Examples include the maggots of
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
or grubs of
weevils Weevils are beetles belonging to the superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and herbivorous. Approximately 97,000 species of weevils are known. They belong to several families, ...
. In contrast, the larvae of other
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
, such as the
Scarabaeidae The family Scarabaeidae, as currently defined, consists of over 30,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several sub ...
and
Dytiscidae The Dytiscidae – based on the Greek ''dytikos'' (δυτικός), "able to dive" – are the predaceous diving beetles, a family of water beetles. They occur in virtually any freshwater habitat around the world, but a few species live a ...
have thoracic legs, but no prolegs. Some insects that exhibit
hypermetamorphosis Hypermetamorphosis, or heteromorphosis,P.J. Gullan & P.S. Cranston. 2010. The Insects: An Outline of Entomology, 4th Edition. Wiley-Blackwell. is a term used in entomology that refers to a class of variants of holometabolism, that is to say, com ...
begin their metamorphosis as
planidia A planidium is a specialized form of insect larva seen in the first-instar of a few families of insects that have parasitoidal ways of life. They are usually flattened, highly Sclerotization, sclerotized (hardened), and quite mobile. The functi ...
, specialised, active, legged larvae, but they end their larval stage as legless maggots, for example the
Acroceridae The Acroceridae are a small family of odd-looking flies. They have a hump-backed appearance with a strikingly small head, generally with a long proboscis for accessing nectar. They are rare and not widely known. The most frequently applied common ...
. Among the
Exopterygota The Exopterygota (Ancient Greek ''ἔξω'' (éxō, “outside”) + ''πτερόν'' (pterón, “wing”) + New Latin ''-ota'' (“having”)), also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infracla ...
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 Ephemeroptera 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
Coccoidea Scale insects are small insects of the Order (biology), order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient g ...
are called "crawlers" and they crawl around looking for a good place to feed, where they settle down and stay for life. Their later
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s have no functional legs in most species. Among the
Apterygota The name Apterygota is sometimes applied to a subclass of small, agile insects, distinguished from other insects by their lack of wings in the present and in their evolutionary history; notable examples are the silverfish, the firebrat, and 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
housefly The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It is believed to have evolved in the Cenozoic Era, possibly in the Middle East, and has spread all over the world as a commensal of humans. It is the most common fl ...
or
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
, has the following parts, in sequence from most
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
to most
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
: *coxa *trochanter *femur *tibia *tarsus *pretarsus. Associated with the leg itself there are various
sclerite A sclerite (Greek , ', meaning "hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly ...
s around its base. Their functions are
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
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 This glossary of entomology describes terms used in the formal study of insect species by entomologists. A–C A synthetic chlorinated hydrocarbon insecticide, toxic to vertebrates. Though i ...
legs because the typical leaping mechanism is to straighten the joint between the femur and the tibia, and the femur contains the necessary massive
bipennate Muscle architecture is the physical arrangement of muscle fibers at the macroscopic level that determines a muscle’s mechanical function. There are several different muscle architecture types including: parallel, pennate and hydrostats. Force pr ...
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
Apocrita Apocrita is a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" ( petiole) formed ...
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
Protura The Protura, or proturans, and sometimes nicknamed coneheads, are very small (0.6-1.5mm long), soil-dwelling animals, so inconspicuous they were not noticed until the 20th century. The Protura constitute an order of hexapods that were previously ...
,
Diplura The order Diplura ("two-pronged bristletails") is one of three orders of non-insect hexapods within the class Entognatha (alongside Collembola (springtails) and Protura). The name "diplura", or "two tails", refers to the characteristic pair of ...
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
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, which may be useful for diagnostic purposes. For example, the
Pterogeniidae Pterogeniidae is a family of beetles belonging to the supefamily Tenebrionoidea. They are found in South Asia, Southeast Asia, China, New Guinea and the Solomon Islands. Adults and larvae have been found associated with the fruiting bodies of pol ...
characteristically have 5-segmented fore- and mid-tarsi, but 4-segmented hind tarsi, whereas the Cerylonidae have four tarsomeres on each tarsus. The distal segment of the typical insect leg is the pretarsus. In the
Collembola Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
, 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
apodeme An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
of the flexor muscle of the ungues. In the
Neoptera Neoptera (Ancient Greek ''néos'' (“new”) + ''pterón'' (“wing”)) is a classification group that includes most orders of the winged insects, specifically those that can flex their wings over their abdomens. This is in contrast with the mo ...
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
Heteroptera The Heteroptera are a group of about 40,000 species of insects in the order Hemiptera. They are sometimes called "true bugs", though that name more commonly refers to the Hemiptera as a whole. "Typical bugs" might be used as a more unequivocal al ...
. Usually the parempodia are bristly (setiform), but in a few species they are fleshy. 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 (
Embioptera The order Embioptera, commonly known as webspinners or footspinners, are a small group of mostly tropical and subtropical insects, classified under the subclass Pterygota. The order has also been called Embiodea or Embiidina. More than 400 ...
) have an enlarged basal tarsomere on each of the front legs, containing the
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
-producing glands. Under their pretarsi, members of the
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
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. 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 ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
'' (
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
),
locusts Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumst ...
(
Acrididae The AcrididaeMacLeay WS (1821) ''Horae Entomologicae or Essays on the Annulose Animals'' 2 are the predominant family of grasshoppers, comprising some 10,000 of the 11,000 species of the entire suborder Caelifera. The Acrididae are best known bec ...
), or stick insects ( Phasmatodea), 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
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
es are good examples. However, there are many specialized adaptations, including: *The forelegs of the
Gryllotalpidae Mole crickets are members of the insect family Gryllotalpidae, in the order Orthoptera (grasshoppers, locusts, and crickets). Mole crickets are cylindrical-bodied, fossorial insects about long as adults, with small eyes and shovel-like fore ...
and some
Scarabaeidae The family Scarabaeidae, as currently defined, consists of over 30,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several sub ...
are adapted to burrowing in earth. *The forelegs of the
Mantispidae Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in ...
,
Mantodea Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
, and
Phymatinae Insects in the subfamily Phymatinae are commonly called ambush bugs after their habit of lying in wait for prey, relying on their superb camouflage. Armed with raptorial forelegs, ambush bugs routinely capture prey ten or more times their own s ...
are adapted to seizing and holding prey in one way, while those of the
Gyrinidae The whirligig beetles are water beetles, comprising the family Gyrinidae that usually swim on the surface of the water if undisturbed, though they swim underwater when threatened. They get their common name from their habit of swimming rapidly i ...
are long and adapted for grasping food or prey in quite a different way. *The forelegs of some butterflies, such as many
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
, are reduced so greatly that only two pairs of functional walking legs remain. *In most
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassho ...
the hind legs are
saltatorial This glossary of entomology describes terms used in the formal study of insect species by entomologists. A–C A synthetic chlorinated hydrocarbon insecticide, toxic to vertebrates. Though i ...
; they have heavily bipinnately muscled femora and straight, long tibiae adapted to leaping and to some extent to defence by kicking.
Flea beetle The flea beetle is a small, jumping beetle of the leaf beetle family (Chrysomelidae), that makes up the tribe Alticini which is part of the subfamily Galerucinae. Historically the flea beetles were classified as their own subfamily. Though m ...
s 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 The bean weevils or seed beetles are a subfamily (Bruchinae) of beetles, now placed in the family Chrysomelidae, though they have historically been treated as a separate family. They are granivores, and typically infest various kinds of seeds or ...
use their swollen hind legs for forcing their way out of the hard-shelled seeds of plants such as '' Erythrina'' in which they grew to adulthood. *The legs of the
Odonata Odonata is an order of flying insects that includes the dragonflies and damselflies. Members of the group first appeared during the Triassic, though members of their total group, Odonatoptera, first appeared in Late Carboniferous. The two comm ...
, the
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
and
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies, which constitute the other odonatan suborder, Anisoptera, but are smaller and have slimmer bodies. Most species fold the wings along ...
, 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, 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 (
somite The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryonic stage of somitogenesis, along the head-to-tail axis in segmented animals. In vertebrates, somites subdivide in ...
s) of different
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s 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. The homologies between these have been discovered by comparing
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s in
evolutionary developmental biology Evolutionary developmental biology (informally, evo-devo) is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved. The field grew from 19th-century beginni ...
.


Leg modification classifications

*
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 ...
legs are modified for
digging Digging, also referred to as excavation, is the process of using some implement such as claws, hands, manual tools or heavy equipment, to remove material from a solid surface, usually soil, sand or rock on the surface of Earth. Digging is actu ...
and
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of sh ...
ing. For example:
mole cricket Mole crickets are members of the insect family Gryllotalpidae, in the order Orthoptera (grasshoppers, locusts, and crickets). Mole crickets are cylindrical-bodied, fossorial insects about long as adults, with small eyes and shovel-like fore ...
s have fossorial forelegs *
Saltatorial This glossary of entomology describes terms used in the formal study of insect species by entomologists. A–C A synthetic chlorinated hydrocarbon insecticide, toxic to vertebrates. Though i ...
legs are modified for
jumping Jumping or leaping is a form of locomotion or movement in which an organism or non-living (e.g., robotic) mechanical system propels itself through the air along a ballistic trajectory. Jumping can be distinguished from running, galloping and o ...
and leaping. For example: many
orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassho ...
ns have saltatory hindlegs * Natatorial legs are modified for
swimming Swimming is the self-propulsion of a person through water, or other liquid, usually for recreation, sport, exercise, or survival. Locomotion is achieved through coordinated movement of the limbs and the body to achieve hydrodynamic thrust that r ...
. For example:
whirligig beetle The whirligig beetles are water beetles, comprising the family Gyrinidae that usually swim on the surface of the water if undisturbed, though they swim underwater when threatened. They get their common name from their habit of swimming rapidly i ...
s have natatorial mesothoracic and metathoracic legs *
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 ...
legs are modified for
running Running is a method of terrestrial locomotion allowing humans and other animals to move rapidly on foot. Running is a type of gait characterized by an aerial phase in which all feet are above the ground (though there are exceptions). This is ...
. For example: lots of
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
es have very sensitive cursorial legs *
Raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...
legs are modified for
grasp A grasp is an act of taking, holding or seizing firmly with (or as if with) the hand. An example of a grasp is the handshake, wherein two people grasp one of each other's like hands. In zoology particularly, prehensility is the quality of an ap ...
ing, usually a hunting adaptation. For example:
Mantids Mantidae is one of the largest families in the order of praying mantises, based on the type species '' Mantis religiosa''; however, most genera are tropical or subtropical. Historically, this was the only family in the order, and many referen ...
use their raptorial forelegs to hunt


References

{{Locomotion Arthropod morphology Animal locomotion Spider anatomy Trilobite anatomy