The thorax represents the second tagma of the insect body. The thorax evolved early in the phylogenetic history of insects. In most Paleozoic insects the thorax is well developed and differentiated from the head and abdomen, and the three distinct tagmata probably developed during the Devonian.

FIGURE 6 Mandible articulation. Top: lateral view; bottom: posterior view (Orthoptera: Stenoplematidae).

In terms of insect phylogeny, the thorax of Apterygota is strikingly different in shape compared with the head or abdomen. Of modern apterygotes only the Collembola display taxa in which thoracic tagmatization and segmentation are not obvious.

Apparently, the primary, functional role of the thorax has always been locomotion, since the primary modifications of the thorax have been for locomotion (first walking, and then flight). Modification for locomotion probably developed before other morphological adaptations, such as metamorphosis. Diverse independent and interdependent mechanisms for locomotion have evolved throughout the Insecta, including walking, flight, and jumping. Active participation in flight by insects is unique among invertebrates.

Anatomy of the Thorax

The cervix is the connection between the head (occipital foramen) and the anteriormost part of the thorax (pronotum) (Fig. 2). Typically, the area between the head and pronotum is membranous. The ground plan for the insect cervix contains two cervical sclerites on each side of the head that articulate with an occipital condyle of the head and the prothoracic episternum. Musculature attached to these sclerites increases or decreases the angle between the sclerites, and creates limited mobility of the head.

The thorax of modern insects consists of three segments termed the prothorax, mesothorax, and metathorax. The last two collectively are called the pterothorax (Greek, ptero = wing or feather) because extant insects bear wings on these segments only. The individual dorsal sclerites or terga of the thoracic segments are also known as nota (Greek, notos = back; sing, notum). The nota of Apterygota and many immature insects are similar to the terga of the abdomen with typical secondary segmentation. The nota of each thoracic segment are serially distinguished as the pronotum, mesonotum, and metanotum.

The size and shape of the prothorax are highly variable. The prothorax may be a large plate as in Orthoptera, Hemiptera, and Coleoptera, or reduced in size forming a narrow band between the head and mesothorax as in Hymenoptera. The prothorax is usually separated or free from the mesothorax. The sclerites are separated by a membrane that may be large and conspicuous in more primitive holometabolous insects such as Neuroptera and Coleoptera, or reduced in size in more highly evolved holometabolous insects such as Diptera and Hymenoptera.

The pterothorax includes the thoracic segments immediately posteriad of the prothorax. In winged insects the relationship between thoracic segments involved in flight can be complicated. In contrast, the thorax of larval insects and most wingless insects is relatively simple. The mesothorax and metathorax of these insects are separated by membrane. Adult winged insects show a mesothorax and metathorax that are consolidated (i.e., more or less united) to form a functional unit modified for flight.

The development of the pterothoracic segments varies among winged insects. When both pairs of wings participate equally in flight, the two thoracic segments are about the same size. This condition is seen in Odonata, some Lepidoptera, and some Neuroptera. When one pair of wings is dominant in flight, the corresponding thoracic segment is commensurately larger and modified for flight, whereas the other thoracic segment is reduced in size. This condition is seen in Diptera and Hymenoptera, where the forewing is large and dominant in flight. The reverse condition is seen in the Coleoptera, where the hind wing is large and dominant in flight.

In more closely related insect groups, such as families within an order, that are primitively wingless or in which wings have been secondarily lost in modern or extant species, many modifications to the thorax occur. Many wingless forms can be attributed to environmental factors that promote or maintain flightlessness. For instance, island-dwelling insects are commonly short winged (brachypterous), or wingless, whereas their continental relatives are winged, presumably because for island species, flight increases the likelihood of being carried aloft, moved out to sea, and subsequently lost to the reproductive effort of the population. The anatomical consequences of flightlessness can be predictable; in the Hymenoptera, short wings bring a disproportionate enlargement of the pronotum and reduction in size of the mesonotum and metanotum.

Sutures and Sclerites of Wing-Bearing Segments

The wing-bearing segments of the thorax are subdivided into a myriad of sclerites that are bounded by sutures and

Scutellum Posterior wing process Antecostal sulcus Postnotum Wing stub Pleural wing process Phragma Subalare Pleural sulcus Epimeron Coxal opening Postcoxal bridge Spinasternum Sternellum

FIGURE 7 Diagram of the pterygote pterothorax.

membranous areas. These sutures and sclerites are the product of repeated modification of the thorax in response to various demands placed on the insect body by the environment. Similar modifications have occurred independently in many groups of insects; some modifications are unique. Generalizations are difficult to make, given the large number of sutures and sclerites, coupled with the number of insects that there are to consider.

Dorsal Aspect

The nota of the pterothorax are further subdivided into the prescutum, scutum, and scutellum; again, serially distinguished as mesoscutum and mesoscutellum, and metascutum and metascutellum (Fig. 7). Additionally, there are sclerites anterior and posterior to the notum, as discussed shortly.

The prescutum is the anterior portion of the scutum, laterally bearing prealar bridges separated by the prescutal suture from the mesoscutum. The scutum is the largest dorsal sclerite of the notum and is bounded posteriorly by the scutoscutellar suture, which divides the notum into the scutum and scutellum. The scutellum is generally smaller than the scutum. In Heteroptera it is a small triangular sclerite between the bases of the hemelytra. In Coleoptera the scutellum is the small triangular sclerite between the bases of the elytra. In Diptera and Hymenoptera the scutellum is relatively large, forming a subhemispherical sclerite, sometimes projecting posteriad. The posteriormost sclerite of the notum is the postnotum, separated from the notum by secondary segmentation. In some insects there is a postscutellum (metanotal acrotergite) that forms the posteriormost thoracic sclerite of the metanotum, or the posteriormost sclerite of the thorax. In Diptera the postscutellum appears as a transverse bulge below the scutellum.

The acrotergite and postnotum deserve further explanation. Again, the anteriormost sclerite is an acrotergite, the anterior precostal part of the notal plate. The postnotum is an intersegmental sclerite associated with the notum of the preceding segment. The postnotum bears the antecosta, a marginal ridge on the inner surface of the notal sclerite

FIGURE 7 Diagram of the pterygote pterothorax.

corresponding to the primary intersegmental fold. The postnotum also usually bears a pair of internal projecting phragmata. The antecostal suture divides the acrotergite from the antecosta, the internal ridge marking the original intersegmental boundary. Thus, when the antecosta and acrotergite are developed into larger plates and are associated with the notum anterior to them, they are referred to as a postnotum. The final structure associated with the dorsal aspect of the pterothorax is the alinotum (Greek, ala = wing; notos = back; pl., alinota). The alinotum is the wing-bearing sclerite of the pterothorax.

Wing Articulation

The thoracic components necessary for wing movement include the prealar bridge, anterior notal wing process, and posterior notal wing process. The components of the wing itself that articulate with the thoracic components are the humeral and axillary sclerites; they form the part of the wing closest to the body and are not treated in this article.

The prealar bridge is a heavily sclerotized and rigid supporting sclerite between the unsclerotized membrane of the pterothorax and the pleuron; it supports the notum above the thoracic pleura. The prealar bridge is comprosed of cuticular extensions from the anterior part of the prescutum and antecosta. The anterior notal wing process is the anterior lobe of the lateral margin of the alinotum supporting the first axillary sclerite (Fig. 7). The posterior notal wing process is a posterior lobe of the lateral margin of the alinotum that supports the third axillary sclerite of the wing base (Fig. 7).

Lateral Aspect

The pleuron (Greek = side; pl., pleura) is a general term associated with the lateral aspect of the thorax. Adults, nymphs, and active larvae all display extensive sclerotization of the pleural area. Sclerites forming this part of the body wall are derived from the precoxa, subscoxa, or supracoxal arch of the subcoxa.

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