Structure

In the classic textbook interpretation, the insectan leg has six well-sclerotized segments, arranged proximal to distal: the coxa, trochanter, femur, tibia, tarsus, and pretarsus. A more fundamental and complete segmentation scheme, which facilitates the recognition of leg and leg-derived homologies among all arthropods, involves 10 or 11 segments. These segments include the epicoxa (debatably present as the wing articulation and a fused portion of the tergum), subcoxa (absorbed into the pleuron), coxa, trochanter, prefemur (typically fused with the trochanter), femur, patella (fused with the tibia), tibia, basitarsus, eutarsus (often subdivided), and pretarsus. A more modern interpretation of the free leg of extant insects, therefore, depicts it as consisting of seven to eight distinct segments, which are the classical six plus a basitarsus and a prefemur, in some insects.

Each segment in the insectan leg, unless secondarily lost or fused, is independently movable by muscles inserted on its base. Thus, subdivisions of the eutarsus, marked by flexible cuticle but without corresponding internal muscles, are not true segments; these subdivisions are referred to as tarsomeres. The areas of flexion between segments are joints, and the well-sclerotized contact points in the joints are the condyles. The various joints contribute to the mechanical efficiency of the leg. The articulation between the coxa and the body, for example, allows the leg to move forward and rearward, whereas that between the coxa and the trochanter allows the leg to be lifted at the end of the backstroke and depressed at the beginning of the backstroke.

Leg joints are of two types. Monocondylic joints have a single point of articulation, somewhat like a ball-and-socket joint, and usually are situated dorsally. They allow considerable freedom of movement and are characteristic of the legs of larval insects. Dicondylic joints consist of an anterior and a posterior condyle, or a dorsal and ventral condyle in the case of the trochanterofemoral joint. They typically limit movement to that of a hinge. Adult legs usually have dicondylic joints, although the tibiotarsal joint is often monocondylic.

The coxa (plural coxae) is typically short and rather stout, although it varies in shape among taxa. It is set in a coxal cavity and articulates with the thorax at the coxal process of the pleural sulcus (groove). Quite often, it also articulates with the thoracic trochantin and sternum, somewhat restricting its movement. To withstand the forces of movement, the coxa is strengthened by a ringlike basicostal sulcus that sets off a basal sclerite, the basicoxite. Internally, the basicostal sulcus is expressed as a ridge, the basicosta, that provides for muscle attachment. Posterior to the point of articulation, the basicoxite is called the meron and in insects such as adult Neuroptera and Lepidoptera, it can be quite large. In higher Diptera, the meron is detached from the coxa and forms a plate in the mesothoracic pleuron. In some insects, an additional external groove, the coxal sulcus, divides the coxa lengthwise.

The trochanter is small and freely movable in a vertical direction on the coxa, but it is often rather fixed to the base of the femur. In the larvae and adults of numerous fossil insects and a few extant taxa, such as Odonata, two trochanteral segments are present, the distal one being the prefemur.

The femur (plural femora) is usually the largest and strongest segment of the leg. Its size is related to the mass of the tibial extensor muscles within it, varying from a small, thick segment in larval insects to the enormous segment in the hind leg of jumping Orthoptera. The femur often is equipped with spines and other cuticular modifications, especially in predatory insects.

The tibia (plural tibiae) typically is long and slender in adult insects. Proximally, it is bent slightly toward the femur, allowing the shaft of the tibia to be flexed close against the femur for more locomotory power in insects such as grasshoppers. It often bears spines for grooming or for engaging the substrate to aid in locomotion. Many insects also have apical or subapical movable spurs on their tibiae.

The tarsus (plural tarsi) is a simple, undivided segment in holometabolous larvae and basal hexapods such as Protura and some Collembola. In collembolans, the tarsus and tibia are fused into a single tibiotarsus. In most insects, a separate segment, the basitarsus, is present and the eutarsus is subdivided into two to four sections or tarsomeres. The ventral surface of the basitarsus and eutarsus often bears pads called tarsal pulvilli that aid movement on smooth surfaces and are especially well developed in some Orthoptera. The basitarsus and eutarsus generally are well endowed with sensory hairs and chemoreceptors. The ventral surface often has a secretory epithelium that produces a wax, possibly for waterproofing, inhibiting the uptake of undesirable water-soluble compounds, or preventing entrapment in surface films.

The pretarsus, also called the acropod or posttarsus, arises from the distal end of the eutarsus. In the Protura, Collembola, and larvae of many holometabolous insects, the pretarsus is a simple, clawlike segment. Typically, however, the pretarsus consists of a membranous base, a pair of hollow claws (ungues), and various sclerites and lobes. A sclerotized unguitractor plate articulates with the eutarsus into which the plate is partly invaginated. The muscles that flex the claws are inserted on a process of this plate. The claws articulate with the unguifer, a median process at the distal end of the eutarsus. A saclike, hollow lobe, the arolium, arises between the claws. In adult Diptera, other than crane flies, an arolium is absent. Instead, a padlike lobe called the pulvillus (plural pulvilli) arises from an auxiliary plate (auxilia) beneath the base of each claw, while an unpaired, lobelike or bristlelike process, the empodium, stems medially from the unguitractor plate. The pretarsal pads and lobes are covered with adhesive setae (tenent hairs) that allow the insect to climb and hold onto smooth surfaces.

Variation in structure, and hence function, can be found among the three pairs of legs within an individual, as well as between larvae and adults, between males and females, and among taxa. The thoracic legs of many larval insects are serially uniform or, sometimes, lacking. The legs of adults often vary in structure among the three pairs, although even adults of some insects (e.g., some female Coccidae, Psychidae, and Strepsiptera) are devoid of legs. The variation among the three pairs of legs in adults often is associated with acquisition of food, courtship, and mating. Developmental variation occurs in holometabolous insects, which have simple, rather generalized legs in the larvae and more specialized legs in the adults. Among the hemimetabolous groups, many Hemiptera-Heteroptera gain a tarsomere in the final molt.

Sexual dimorphism in leg structure is particularly prevalent. The reduced forelegs of nymphalid butterflies have short tarsomeres in females but lack all segments beyond the tibia in males. The forelegs of male Ephemeroptera are typically elongated to grasp the female. The slender, elongate legs of crane flies are even longer in males than in females for species in which a guarding male stands over the female during oviposition; in some crane flies the distalmost tarsomere is prehensile in males and used for holding females. The hind femora of males of many Coreidae are enlarged for intrasexual fighting. The pulvilli beneath the claws of some flies, such as Tachinidae, are considerably larger in males than in females.

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