orthopteran

/awr thop"teuhr euhn/, adj.

1. orthopterous.

n.

2. an orthopterous insect.

[1895-1900; < NL Orthopter(a) (see ORTHOPTERON) + -AN]

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▪ insect

Introduction

  broadly, any member of one of four insect orders. Orthopteran has come to be regarded as the common name for these related groups, which exhibit considerable morphological, physiological, and paleontological diversity. Although sometimes the insects are combined into the order Orthoptera, generally several orders are implied in the term orthopteran.

 Among the orthopterans, cockroaches (cockroach) and mantids (mantid) are placed in the order Dictyoptera, although they are sometimes placed in Blattodea and Mantodea, respectively, which may be considered as separate orders or as suborders of Dictyoptera. The grylloblattids (order Grylloblattodea (ice bug)) and walking sticks (walkingstick) (order Phasmida) are given ordinal rank also. On the other hand, members of the suborders Ensifera (katydids (katydid), crickets (cricket), and camel crickets) and Caelifera (pygmy sand crickets (pygmy sand cricket), grasshoppers (grasshopper), and locusts (locust)) are considered to comprise the order Orthoptera. For completeness of discussion, all of these groups, handled here as four separate orders, are included in this article.

 Orthopterans, abundant in tropical regions throughout the world in both numbers of species and individuals, are common in the summer months in temperate regions, when their relatively large size and chirping sounds attract considerable attention. Zoologists have long been interested in cockroaches, one of the oldest insect groups known. Most of the 24,000 species of orthopterans are plant feeders, with mouthparts adapted for chewing. Locusts, known as pests since biblical times, are very destructive to agricultural products.

General features

      Orthopterans may be bizarre in appearance, unusually large in size, or show peculiar behaviour. They range in size from a few millimetres to more than 30 centimetres. Some tropical walking sticks resembling tree twigs are more than 30 centimetres (11.8 inches) long, and others, much smaller, resemble leaves of plants. The size range of present-day cockroaches is typical of the diversity of body size among orthopterans: tiny flightless cockroaches (Attaphila), living as commensals in the nests of ants, are only two millimetres long when mature, whereas a species of Megaloblatta found in South America reaches 10 centimetres in length with a wing span of almost 19 centimetres.

      Approximately 24,000 species of orthopterans have been identified. Throughout the U.S. there are about 1,300 species; not all of them inhabit any one place. There are more species (e.g., 282 in Arizona) in southern and southwestern sections of the U.S. than in the North (e.g., 103 in all the New England states). Of the 600 species found in Europe, Great Britain has only 35, including four established, introduced species (adventives).

      The largest families of orthopterans are worldwide in range, although all have decreased numbers of species in cold temperate zones. Few mantids or walking sticks, for example, occur outside tropical or subtropical areas. There are about 20 mantid species and 27 walking stick species in the southern regions of the U.S., compared with the 400 mantid species and 600 walking stick species that have been identified in Central and South America. A few northern groups include the grylloblattids and several genera of grasshoppers.

Importance

      Among the orthopterans are many species that are either harmful to agricultural products or are pests (pest). Grasshoppers are capable of causing widespread devastation of the agricultural crops grown in many countries throughout the world. In cattle-growing regions there often is competition between grasshoppers and livestock for available forage. Mormon crickets (a common name for species of the genus Anabrus that originated during the early years of the Mormon settlement in Utah) are major pests of both crops and open rangeland in the western part of the U.S. during seasons that are favourable for their development. Cockroaches, known throughout the world as domestic pests, are a frequent nuisance, especially in warm-temperate to tropical areas. Although cockroaches occasionally carry organisms such as bacteria or parasites that produce intestinal diseases, they are more generally considered to be mechanical carriers of contaminating filth.

      Mantids, predators on other insects, have become adapted to resemble the flowers, tree trunks, or grass stems on which they await their prey. Crickets, katydids, and grasshoppers are known for the songs they produce using stridulatory mechanisms, and research concerned with song production is an active field. The biology of migratory (migration) grasshoppers or locusts involves hormones that promote transformation of nonmigratory, solitary, shorthorned grasshoppers into gregarious hordes of locusts capable of causing great destruction. This transformation has been studied in attempts to control these pests.

Natural history

life cycle

General features

 Since orthopterans undergo simple metamorphosis and have externally developing wings, they are known as hemimetabolous insects. The grylloblattids are wingless, and all large orthopteran groups contain a few wingless species, even though the basic structure of the orthopteran thorax proves their relationship to winged insects. A typical orthopteran life cycle has three stages: egg, nymph, and adult. Usually eggs are deposited outside the body on the ground or on vegetation; however, in some cases (e.g., viviparous cockroaches), eggs are incubated in a brood chamber within the body of the female, and nymphs are born alive. Nymphs resemble adults except for their smaller size and lack of development of reproductive organs and wings; there is no pupa, or resting stage. In most orthopteran groups, the hatching insect that wriggles from the egg is not a fully formed nymph with freely moving legs; actually it is little more than an active embryo and is still enclosed in a thin membrane. This stage is called a vermiform larva; shedding of the enclosing membrane occurs at the intermediate molt. The shapeless skins shed by young grasshoppers crawling from egg pods or by mantids leaving an egg case are examples of such exuviae (cast skins).

      The number of nymphal stages between the intermediate molt and adulthood varies from about 4 to 13. Generalizations are approximately as follows: cockroaches, 5–13; mantids, 4–9; grylloblattids, about 8; crickets and katydids, 5–9; walking sticks, 4–6; and grasshoppers, 4–9, most often 5–7.

egg laying

      Egg-laying habits are distinctive in many orthopteran species. Among cockroaches, only one family (Blaberidae) is viviparous; the other four families contain species that have well formed egg cases (oothecae). Among these families, some species carry the oothecae protruding from the body until hatching time is near; others, however, deposit their egg cases within several days of formation. Usually oothecae contain from a few to more than 30 eggs arranged in two rows. Along one edge of the ootheca is a seam that bears a keel, or ridge; the shape of the ridge varies in species that carry the ootheca externally. Minute openings from the base of the keel to the interior of the ootheca are known to be a ventilating device in some species. The ootheca is first carried in the body with the keel uppermost; in certain groups, however, it is rotated prior to deposition so that the keel is on one side. Details concerning positioning of the ootheca and other aspects of egg laying were not correlated with behaviour patterns or group affinities until a basis for understanding their significance was established.

      Mantids lay eggs in clusters of less than 10 to more than 300. Usually they are laid in regular layers surrounded by a viscous quick-drying liquid that provides a light but tough protective covering. The egg masses of most mantids have a distinctive shape and size. Although most mantid species attach oothecae to vegetation, these egg cases may be attached to rocks in some environments or placed in grooves in the sand and covered over in the desert.

      Grylloblattid eggs are laid in damp moss, decaying logs, or in pockets between broken rocks and wet soil. The eggs, about three millimetres long and usually black in colour, are laid loosely or inserted into the hatching site with the ovipositor.

      Of the walking stick species studied, most have eggs that look like small seeds and are dropped loosely on the ground. At least one species, however, attaches its eggs to foliage, and a large, heavily bodied species of the southeastern U.S. (Anisomorpha buprestoides) scratches a depression in sandy soil with its front and middle legs, deposits eggs in it, and covers them with sand.

      Crickets and katydids utilize their ovipositors to insert eggs into soil or plant material. The eggs of tree crickets (Oecanthus), for example, are inserted in rows in the canes of blackberries and various other stems; the eggs of field crickets (Gryllus) are laid in soil; the flattened eggs of certain katydids (Scudderia) are forced between the upper and lower epidermis layers at the margin of tree leaves; and the eggs of other katydids (Microcentrum) are laid in overlapping rows lengthwise on twigs of trees.

      Most grasshoppers lay their eggs in soil; a few drill holes in dead wood or place their eggs at the bases of grass clumps or on the surfaces of leaves. Before laying the eggs, the female manipulates the valves of her ovipositor to make a hole in suitable soil (the type varies with the species). During the digging process, the abdomen is greatly extended, and the female manipulates the ovipositor valves to open and close and rotate on the long axis. Then she deposits several dozen eggs in the hole. The eggs are surrounded by a mucilaginous mass (the egg pod) that dries in a cylindrical shape. The number of egg pods laid by one female and the number of eggs in each pod vary according to the species and local conditions. The egg pods are laid over an interval of several weeks.

      A few orthopteran species have females only; therefore, reproduction occurs without fertilization ( parthenogenesis). Only rarely are species that normally reproduce bisexually parthenogenetic; and when parthenogenesis does occur in bisexual species, it is usually only partially successful because the few nymphs that do hatch often are deformed and fail to reach maturity. In the laboratory, however, there have been a few cases in which several generations have been produced parthenogenetically, proving that there is an inherent capability in these bisexual groups for reproduction without males.

Growth and life span

 Increases in the number of antennal segments occur during the development of some orthopterans. The German cockroach, Blattella germanica, has been studied in detail in this regard, and it has been shown that a newly hatched nymph has about 24 segments in each antenna. Each of the succeeding nymphal stages (there are usually six in this species) shows an increase in the number of segments until, by the time the adult stage is reached, the average number is 94. The two basal segments do not divide; the third segment, as well as certain other ones, is a growth centre that divides during molting. It is customary for grasshoppers to have 20–30 antennal segments when mature; this is about twice the number present in the first-stage nymph.

      The life span of orthopterans depends in part upon whether or not there are long periods during winter or at other times (e.g., a dry season) when the insects are quiescent. Some species habitually spend several months resting during unfavourable periods; such species have one generation per year, and the life span of an individual is approximately one year. The portion of the life span spent as an adult varies, but is likely to be about one or two months. In some species egg maturation is delayed several months after the final molt, and the females do not lay the eggs until they have matured. In other species eggs are not hatched until one or more years after they are laid; therefore, more than one winter or dry season is passed in the egg stage, and a single life cycle can occupy two or more full years. For instance, a walking stick commonly found in the U.S., Diapheromera femorata, often has some eggs that hatch the year following deposition and others that hatch after two winters have been passed amid dead leaves on the ground. There are some orthopterans that develop very slowly, and their life cycles require several years for completion; an example is a North American cockroach (Cryptocercus punctulatus) that inhabits rotting logs, feeds on decaying wood, and attains maturity after six or seven years. Grylloblattids live for five to seven years. The time required for domestic cockroaches to reach maturity varies with species and environmental conditions. The German cockroach completes nymphal growth in about 95 days, but the American cockroach (Periplaneta americana) needs about 225 days. Similarly, adults live from a month or two to several years, depending on conditions and species.

Reproduction

General features

      Typically each female has paired ovaries consisting of tubes in which eggs develop, moving posteriorly into a single oviduct as they “ripen.” The oviduct leads to a vagina and then to the exterior where there is either a simple or specialized ovipositor consisting of paired appendages called ovipositor valves. Attached to the oviduct or vagina is a sac (called the spermatheca) for storage of male sperm; as eggs move down the oviduct, they are fertilized by the sperm. The typical male contains paired testes that produce vast numbers of slender active sperm; these are stored in enlargements of the tubes leading posteriorly from the testes. Accessory gland secretions provide not only the medium for carrying the sperm but also a material that solidifies to form a thin-walled sac, or reservoir, containing some sperm and fluid. This reservoir, called the spermatophore, is almost universally found among orthopterans.

      Grasshopper spermatophores consist of a bladderlike reservoir and a spermatophore tube. The spermatophore is formed during the first two minutes or so of copulation, after which the tube extends from the male genital organs to the spermathecal duct of the female. The spermathecal duct opens at the base of the ovipositor valves. Sperm pass to the female during copulation; after sperm transfer is complete, parts of the spermatophore may remain attached to both male and female. In some orthopterans, particularly crickets and katydids, the entire spermatophore is attached to the female.

courtship behaviour

      All orthopteran groups have species that show definite courtship behavior prior to actual mating. Male cockroaches are attracted particularly by females that are virgin and in a receptive condition. Such females frequently secrete pheromones (pheromone). Pheromones, chemical substances secreted by certain insects, influence the behaviour of other individuals of the same species. Antennae of males of a domestic roach, Periplaneta americana, have specialized sense organs that detect the odour of female P. americana pheromones; upon detection of the odour, the male initiates searching movements, first with the antennae, then with the palpi. Finally, the male, with folded wings raised and fluttering, actively searches out the female. If the female is still receptive when he finds her, the male protrudes his posterior abdominal segments, pushes under the end of the female, and grasps the terminal ventral segments of the female with his genital hooks; then he expels the spermatophore, which becomes attached to the spermathecal opening of the female. The entire process lasts up to an hour.

      Among the grasshoppers, species with coloured hind wings and the habit of making sounds during flight use hovering and other special flight patterns to attract the attention of females. Crickets and katydids have the most dramatic courtship displays because “songs” enter into the precopulatory behaviour. Females of some species are receptive only to the specific song of a male of the same species; in others, however, mating calls are not necessary, and a female will mate with a male who is unable to sing because his wings have been removed. Here, as in grasshoppers, a variety of mating positions are assumed.

      A striking sequel to mating occurs frequently in mantids when the female eats the male. There is a popular opinion that mantid males always are eaten, but many escape under natural conditions. But in the close confines of a small cage cannibalism of the male is more common.

Ecology

      In a broad sense, ecology represents the sum total of interrelations between organisms and their environment. In the case of orthopterans, the basic requirements of food and moisture; shelter, including protection from weather and from enemies; favourable habitats, involving special niches such as caves or deserts; as well as preferred seasons and conditions conducive to successful reproduction, are involved.

Food

      Mantids are the only orthopteran group that feeds almost entirely on insects, but some members of primarily plant-feeding groups also capture and devour insects. For instance, tree crickets (Oecanthus) regularly eat flowers, leaves, and other plant parts, in addition to many aphids and other weak insects. Some katydids are active predators of insects. Most cockroaches are typical scavengers, but some are specialized feeders. Cryptocercus, for example, digests cellulose in decaying logs with the aid of symbiotic protozoans in its intestines. All walking sticks and a majority of grasshoppers are plant feeders. Although many grasshoppers feed on a wide variety of plant species, some are restricted to a single plant species or one group of plants. Some orthopterans consume only certain parts of plants; for example, coneheaded katydids of the genus Neoconocephalus feed mainly on seeds of grasses. Plant preferences among some leaf-eating orthopterans change with the seasons of the year; in other leaf-eating groups feeding habits are dependent on the stage of the life cycle—i.e., nymphs do not eat the same plants as adults of the same species. Moisture requirements of orthopterans vary, as evidenced by the habitats they occupy. Some can absorb water from a drop on the cuticle (skin); others obtain it from water vapour in the air if the relative humidity is sufficiently high.

Habitat

      Shelter utilized by orthopterans ranges from general hiding places amid living plant foliage or dead leaves on the ground to special structures such as subterranean galleries in soil or the recesses in caves occupied by various crickets. Some Gryllacrididae are leaf rollers and produce silk to maintain the rolled shape of their hiding places. The loose bark of trees and logs and the water-filled leaf bases of bromeliads often shelter certain genera of cockroaches, some of which are semi-aquatic in their habits. In Africa a few cockroaches (Cyrtotria), of elongated and cylindrical body shape, are adapted to enter round holes in hollow plant stems where they sometimes live. With the exception of cockroaches, most conspicuous orthopterans are active by day (diurnal), although there are nocturnal species in every group. Although grylloblattids are essentially nocturnal, they are sometimes active on cloudy days or in winter. The majority of crickets and katydids are nocturnal, as are many walking sticks and some mantids; however, many mantids prey on insects that visit flowers by day.

      The degree of moisture, types of vegetation, and altitude above sea level influence the location of orthopteran communities. Grasshoppers breed in the Himalayan Mountains at altitudes as high as 6,000 metres (about 18,000 feet), and each mountain altitudinal zone has distinctive species; fully winged, actively flying species are usually not restricted to a single zone but are found in adjacent ones. On the other hand, at high altitudes there are proportionately greater numbers of grasshoppers with short, nonfunctional wings or none at all.

      Caves (cave) are a special habitat occupied by orthopterans on all continents. The long-horned grasshoppers and the crickets are the principal orthopteran representatives; nearly 200 species of these two groups have been found in caves. In addition, more than 30 cockroach species inhabit caves; and a third group, the grylloblattids, has at least one cavernicolous species in Japan and three in the U.S. Some of these orthopterans inhabit lava tubes and fissures resulting from past volcanic action. Air currents and high humidity in these tubes and fissures produce an “ice cave” condition. In the U.S., the grylloblattids and a few dozen cave crickets (Gryllacrididae) are the principal cavernicoles. It is noteworthy that a bone from a bison skeleton, found in a French cave in the 1920s, bore a prehistoric carving that depicted Troglophilus, a European cave cricket.

      Usually the orthopteran species found on a given continent are distinct from those of other continents, especially if the land masses are well separated. For example, there are about 2,000 species and 500 genera of grasshoppers in Africa; although several of the genera are found in North America, none of the species is. Some species in North Africa, however, also inhabit southern Europe and western Asia. Explanations for distinct continental species are found not only in the far, overwater distances involved but also in the long periods of isolation that have occurred and in the different conditions that have prevailed in past geologic periods.

      Oceanic islands have been populated, in part, by species that were transported by hurricanes, floating debris, birds, and in recent centuries by human activities. The 85 species of Hawaiian orthopterans include four distinct genera of crickets and katydids comprising about 45 of the 85 species. Evidently some of these evolved following the establishment of a few parent species. The remainder are believed to have been established as a result of the activities of man. Since cockroaches are scavengers, they are often found where man is found; i.e., in his buildings and campsites. Early man probably spread cockroaches as he moved about seeking food. Modern commerce has been even more helpful to these unwelcome travellers. An analysis of the distribution of 11 domiciliary species found in the U.S. and related species found elsewhere suggested that five reached America in shipping from West Africa; another African species might not have reached America directly; two probably came from Europe; two might have come from the Orient; and one was native to the West Indies. Thus man has played an important role in spreading cockroaches throughout the world.

      The familiar, large, black field crickets of the U.S. are good examples of ecological differences among similar species. There are six native species in the eastern U.S.; several others occur in the western states, and at least two introduced species have become established in the Gulf States. Using the general appearance of dead specimens, the native eastern species are very similar and difficult to distinguish. For many years, the taxonomy of these species was unsettled. Entomologists using behaviour rather than morphology as a major taxonomic criterion have found that five of the six species have distinct songs; that four of them overwinter as nymphs, the other two chiefly as eggs; and that, to a considerable extent, the habitat preferences are different—i.e., one species lives in abandoned fields, another in leaf litter of open woodland. Laboratory attempts to crossbreed males and females of different species have been unsuccessful; the pair either failed to copulate or, if they did, produced unhealthy hybrids. Ecological differences also are important for other groups of closely related orthopterans.

Form and function

Adaptive features

Movement

      Orthopterans exhibit various adaptations (adaptation) for movement; some are present in an entire family or suborder, others are peculiar to certain genera. The head of mantids is borne by the prothorax in such a way that it is easily turned to face in different directions. Since the mantid diet consists almost entirely of insects, vision is critically important and is unusually well developed. The best known orthopterans with specialized front legs (leg) are mantids; the principal leg segments are hinged and spined for seizing and holding prey. Some Orthoptera, especially certain groups of Tettigonioidea, also have front legs with long spines that enable them to hold other insects, although the hinging is not comparable to that in mantids.

      Although some cockroaches burrow in soil, sand, or decomposing wood, the principal burrowers are found among the Orthoptera. In both groups the legs, especially the front tibiae, are short and strong, with heavy spurs. Mole crickets, false mole crickets, and sand crickets are accomplished burrowers. Small tunnels serve as shelter and as egg-laying locations, and roots or tubers encountered while burrowing are sometimes used as food. Several genera of camel crickets (Gryllacrididae) in the southwestern U.S. have conspicuous, sometimes basket-shaped, clusters of spurs on the hind legs. They often live on sand dunes and burrow chiefly for shelter. A few desert-living grasshoppers, some in the southwestern U.S., others in Africa, exhibit what has been called “self-burial.” Instead of making an elongated cylindrical burrow, the grasshopper rests on the surface of the sand or moves forward and backward, manoeuvring its legs until it has submerged itself and covered its body with sand. The apparent purpose is protection.

      Hind legs of Orthoptera, though useful in walking, are used primarily for leaping. Particularly important are the large muscle in the femur, the hinged attachment of tibia to femur, and the tendon extending within the leg from the femur to the end of the tarsus. In a few semi-aquatic Orthoptera, the hind tibia is broadened as a paddle or equipped with fringed spurs to permit effective swimming strokes in water. The ability to run swiftly is common among cockroaches and some mantids. Cockroaches escape enemies by running; mantids utilize their running ability both to escape predators and to catch prey. Some mantids that live on the ground, in deserts, or on tree trunks in the tropics are active runners; however, the majority of mantids stalk their prey slowly or wait quietly until an unsuspecting insect moves nearby.

      Body shape is important to many orthopterans, either allowing them to live in places where adequate shelter from weather and enemies is provided or affording them concealment through camouflage. Most cockroaches have flat bodies that enable them to hide beneath stones, under other objects on the ground, or under the loose bark of logs. Examples of orthopterans whose camouflages resemble parts of plants are members of the Phasmida; some of them resemble leaves, others look like twigs or rough pieces of small limbs from trees. Several katydids and grasshoppers resemble leaves; some are green or brown, others have spots that resemble leaves affected by plant diseases. There are some slender grasshoppers that live among grasses, where they conceal themselves by clinging lengthwise to stems and remaining motionless or by quickly sidling around behind stems.

Camouflage

      There are two basic types of insect colours (coloration). Structural colours occur when irregular cuticle or scale surfaces break up and reflect certain wave lengths of light. Metallic lustres of some orthopterans (e.g., silvery patches on some grasshoppers) are examples. Most orthopteran colours are due to pigments; often they are located in the cuticle, but sometimes they occur in some deeper body layer. The pigments may be naturally occurring ones or, like melanin, dependent on an oxidation process or a hormonal balance that influences metabolism; these latter pigments are present in varying amounts in different individuals of the same species.

      Among some orthopterans, especially grasshoppers, body colours tend to simulate the colour of the habitat background. This is particularly true of species inhabiting rocky or sandy environments. In some cases, colour changes occur rapidly; this was demonstrated by certain light gray African grasshoppers that became black after being caged a few days on dark burnt-over ground. In other cases more time is required. Colour changes usually involve the effect of bright light on integumentary pigments. Among some orthopterans, however, light must enter the eyes, and a rhythm related to some nervous-endocrine mechanism is apparently involved.

      An unusual and rapid colour change occurs in an Australian alpine grasshopper (Kosciuscola tristis), which lives at above 5,000 feet elevation. The adult male, bright greenish blue on the upper part of its body at temperatures above 25° C (77° F), is dull and blackish below 15° C (59° F). At intermediate temperatures, correspondingly intermediate shades of colour occur. Detailed experiments by Australian entomologists prove that temperature, not light intensity, relative humidity, or degree of crowding is the controlling factor. The epidermal cells of the integument contain brown and blue granules; at warmer temperatures on sunny days the blue granules, in a discrete layer uppermost in the epidermal cells, are near the surface of the integument. At night or on cloudy days, the brown granules migrate from the bottom of the epidermal cells and change places with the blue granules. Thirty minutes is sufficient time for a colour change to take place.

Defense

      There are no known stinging orthopterans but many have chemical mechanisms in the form of glands that produce irritating fluids or repugnant odours. The disagreeable smell of some cockroaches, especially when disturbed, is well known. Examples are several species of Eurycotis in Florida and tropical America; both sexes have a large gland in the hind part of the abdomen between the sixth and seventh segments. An acidic, milky fluid consisting of several chemical constituents is emitted either as an oozing liquid or as a three-foot spray. Another cockroach (Diploptera) has a defense gland that ejects a mixture of quinones from the second abdominal spiracles. Ants, beetles, and other predators become confused and avoid these cockroaches when they release their secretions; however, certain mantid predators are not affected.

      Man may handle most walking sticks safely, but a large, heavily bodied species in the southeastern U.S. (Anisomorpha buprestoides) sometimes forcibly ejects a milky fluid that is extremely irritating if introduced into the human eye. This species has a pair of circular pores on the thorax leading to reservoirs of the fluid; each reservoir has circular muscles that permit ejection of fluid without the general body contraction characteristic of some grasshoppers. When handled, most grasshoppers and some other orthopterans regurgitate from the mouth a brown fluid that superficially resembles molasses. Release of the fluid from the forward part of the alimentary canal is triggered by a response of the nervous system to pressure on certain parts of the body, especially the sides of the thorax or the femurs. Some grasshoppers have other defense mechanisms (e.g., some exude fluid through spiracles or from special glands opening on the body or even leg joints). Sometimes hissing sounds and blowing of bubbles from spiracles accompany secretion.

Physiology and biochemistry

Body composition

 Several grasshopper species have been analyzed chemically. They consist of (by dry weight) roughly 50 percent to 75 percent crude protein, 4 percent to 18 percent fats, 4 percent to 16 percent carbohydrates, and 3 percent to 19 percent ash.

      The tough and usually hard outer body wall (exoskeleton) of orthopterans is called the integument or cuticle; (cuticle) its most important component is chitin, a stable polysaccharide chemically similar to plant cellulose. Chitin makes the cuticle strong and flexible but does not provide rigidity. sclerotin, the horny substance of the cuticle formed by a tanning-like process involving protein produced in the exoskeleton, is found in hard body plates (sclerites), leg spurs, and sharp tubercles; sclerotin is responsible for the rigidity of these structures. A heavily “sclerotized” cuticle is one that is hard and usually dark-coloured.

Hormones (hormone)

      The importance of hormones in the biology of orthopterans has been revealed by research. Together with the related pheromones, which tend to coordinate individuals within the population of a species instead of regulating function within an individual, hormones are important in many activities of orthopterans related to mating and reproduction. Other activities involving hormones in grasshoppers include control of fat accumulation in metabolism, control of peristalsis in the malpighian tubules (excretory organs attached to the posterior part of the alimentary canal), secretion of an enzyme at hatching time for dissolving the cuticle that encloses the embryo, and control of the number of molts in nymphal growth.

      Detailed studies on the reproduction of cockroaches have disclosed an interrelated series of neurological and glandular functions that combine to control mating and egg production. Frequently, dorsal abdominal glands of the male aid in attracting the female to a mating position. In several cases, once a female has mated and an ootheca is being carried, mechanical pressure of the ootheca causes a stimulation to be transmitted to glandular bodies closely associated with the cerebral ganglia and called corpora allata; this in turn inhibits development of additional eggs in the ovarioles until laying and subsequent removal of pressure occur. In other cases, virgin females are receptive to mating just when yolk deposition is occurring in the first oocytes of developing eggs (egg). Following mating, the mechanical stimulation of the inserted spermatophore inhibits further attraction of the female to the male abdominal glands until after the first group of eggs is deposited.

 Locust is a common name for several species of short-horned grasshoppers that often increase suddenly in numbers and undertake mass (animal behaviour) migration. A locust has both solitary and gregarious phases. Gregarious locusts outnumber solitary ones, migrate both as nymphs and adults, and travel in swarms. Swarming adults are tremendously destructive to crops. Typically, gregarious locusts have darker bodies and longer wings compared with solitary forms. Colour changes in adults are correlated with maturation of reproductive organs.

      Hormones and pheromones are involved in many stages of locust development. Solitary locusts can transform into gregarious ones as a result of hormonal changes induced by crowding. The presence of mature male locusts under conditions of crowding stimulates a maturation hormone that causes females to mature rapidly. Head glands in the female are stimulated to release another hormone that speeds egg maturation. A favourable season followed by an unfavourable one may cause gregarious locusts to develop. In a favourable season with enough food, the population of solitary locusts increases. If the next season is a poor one, the solitary locusts are forced to crowd together where food is available. Crowding exposes the females to male secretions, females and their eggs respond by maturing rapidly, a population explosion occurs, and a locust horde results. In Schistocerca gregaria, the attainment of reproductive activity is sometimes synchronized with environmental contact with certain aromatic shrubs that produce terpenoids in season.

sound production and hearing

      Some orthopterans make conspicuous sounds, while others produce sounds that are outside the range of human hearing. In both cases sound production is important to behaviour necessary for success of the species concerned. Except for Grylloblattodea, in which sound production is unknown, all major groups of orthopterans produce some sort of sound, though sound production is widespread only in crickets, katydids, and grasshoppers.

      The stridulatory mechanism of grasshoppers involves moving the hindleg across the folded front wing (tegmen). Serrations, or pegs, which vary in shape, number, and location among different species, are located on the inner surface of the femur and rub across special raised veins of the tegmen, creating a characteristic lisp; sometimes the serrations are on the tegminal veins. In the hindwings of other grasshoppers are stiff veins that make a crackling sound (crepitation) in flight.

      Among male crickets and katydids, a front wing with an enlarged transverse vein near its base bears teeth that rasp when shuffled across a scraper on the other front wing. The row of teeth is called the file, and the membrane to which it is attached vibrates when the teeth move over the scraper. During stridulation the tegmina are lifted at an angle of 15° to 40° to the surface of the abdomen, then rapidly opened and closed (shuffled); sound is produced during the closure.

      The best-known auditory organs of orthopterans, the tympanic organs on each side of the abdomen, are found in both sexes of grasshoppers and on the front tibiae of most crickets and katydids. There are auditory nerves running from special cells beneath a tympanic membrane (a thin area of cuticle, backed by an air sac and free to vibrate) to a ganglion of the central nervous system. In addition to these evident tympanic structures, other less evident auditory organs occur in the orthopterans. Many orthopterans, however, have no conspicuous tympana and are entirely dependent for sound reception on sensory hairs located on cerci, the head, other parts of the body, and an auditory organ called Johnston's organ, which is widespread in the second segment of the antenna.

Evolution and paleontology

      Cockroaches are the most abundant and the earliest fossilized orthopterans found; fossils have been discovered at various localities in North America, Europe, and northern Asia. Several hundred Carboniferous Period (about 280,000,000 to 345,000,000 years old) and at least a hundred Triassic cockroach species (about 190,000,000 to 225,000,000 years old) have been described; they differ from present Blattaria chiefly in details of wing venation. In addition, some early species had long ovipositors, unknown in recent species. Only a few fossil mantids are known, the oldest in Baltic amber of the Oligocene (about 26,000,000 to 38,000,000 years ago). Some phasmatid-like Jurassic species (about 136,000,000 to 190,000,000 years old) are believed to be primitive walking sticks.

      Although fewer in number than the Blattaria, fossil Orthoptera have contributed to orthopteran classification. Ensifera occur from the Triassic to the present; apparently Ensifera and Caelifera separated as distinct evolutionary lines as early as the Carboniferous. The earliest fossils in the acrididoid line had long antennae. Shorter antennae, reduction of tarsal segments to three, and reduction in length of the ovipositor occurred by the Eocene Period (about 38,000,000 to 54,000,000 years ago).

      By the late 19th century, all principal groups of orthopterans except Grylloblattodea were represented in collections; however, the order Orthoptera, broader in scope than it is at the present time, included earwigs and other groups. Gryllacridids were not placed in a separate family, and Phasmida were considered a family closely related to Mantidae because both are walking rather than jumping in habit. By the 20th century, however, basic morphological studies, as well as extensive reports on fossils, contributed new insights into the fundamental relationships of the major groups. The Grylloblattids were first reported in 1914, and numerous publications since then have analyzed their phylogenetic significance. In the late 1930s extensive studies of fossils were correlated with important work on current species, especially concerning the Orthoptera (restricted sense). Meanwhile, comparative studies of wing venation, the proventriculus, reproductive organs, and behaviour have steadily advanced the knowledge of group relationships. Additional details and supporting data, for example, were given in an extensive phylogenetic study in 1968. The rank of a suborder (Acridomorpha) for grasshoppers alone (Eumastacoidea through Acridoidea of this article) has not been evaluated sufficiently. Many of the earliest fossil orthopterans were different enough from any present ones to justify the recognition of separate though extinct families.

Classification

Distinguishing taxonomic features

      Among the distinctive features of orthopterans are their wings (wing), which, when present, usually number four. The two forewings, generally long and narrow, are many-veined and somewhat thickened. Among the Orthoptera, Dictyoptera, and Phasmida the forewings, hardened and of a leathery consistency, are known as tegmina. The hindwings, broad with many veins, usually are folded fanlike beneath the forewings when at rest. The females have ovipositors. Some are concealed by ventral abdominal segments; others are as long as the body. Orthopterans have mandibulate mouthparts adapted for chewing and undergo simple metamorphosis.

      The classification of orthopterans into orders and families is based chiefly on comparative morphology, on indications of ancestry derived from fossils, and on relationships suggested by patterns of behaviour and the physiology of body systems. Similar anatomical structures that have been significant in deducing orthopteran relationships include: tarsal segments; hindlegs; wings; stridulatory and auditory organs; head capsule; thoracic sclerites; ovipositor; male genitalia; and proventriculus. In recent years correlations between behaviour and mating and reproduction, especially in cockroaches and crickets, have been used widely to support classification of families and subfamilies as well as distinctions between species.

 In the grylloblattids, cockroaches, and mantids, the cerci are segmented; segments are lacking in katydids, crickets, grasshoppers, and walking sticks. The external male genitalia are sometimes concealed; in Grylloblattodea (grylloblattids), Dictyoptera (cockroaches and mantids), and Phasmida (walking sticks), the genitalia are asymmetric; in the Orthoptera (katydids, crickets, grasshoppers, and locusts) they are symmetrical. The Orthoptera have the femur of the hindleg enlarged for jumping; other groups have hindlegs similar in size to the middle legs. In mantids the front legs are modified for seizing prey, in mole crickets for digging. The tarsus consists of five segments in grylloblattids, mantids, and cockroaches; there are usually five segments in walking sticks, usually only three or four in Orthoptera. The antennal segments vary from fewer than seven to many long and setaceous ones.

Annotated classification

Orthopteran

 Common name for several orders of related insects; wings, when present, number 4; chewing mouthparts; mostly plant feeders; size range from 2 mm to 30 cm; more than 24,000 species; worldwide distribution.

      Order Dictyoptera

 Hindlegs similar to middle ones, adapted for running; tarsi 5-segmented; wing pads not reversed during nymphal stages (front wings remain above hindwings); antennae usually filiform, with more than 30 segments; cerci many-segmented; without auditory tympanum.

      Suborder Blattaria (cockroaches)

 Head usually concealed from above by shieldlike pronotum; two ocelli represented by pale areas (fenestrae); front legs adapted for running; proventriculus (gizzard) heavily armed on inner lining, with longitudinal folds between the teeth; members from Carboniferous to present; worldwide distribution; sizes 2 to 100 mm, average about 15 mm; more than 3,000 species.

      Family Cryptocercidae

 Wingless; blackish; cerci (sensory appendages) concealed by apical segments of abdomen; eyes small; size usually at least 15 mm; found in China, Manchuria, U.S.; 3 species.

      Family Blattidae

 Middle and hindfemurs with numerous strong spines similarly arranged on both ventral margins; ovipositor a plowlike (vavular) modification of apical ventral segment of abdomen; male subgenital plate symmetrical with widely spaced styli; size range about 10 to 35 mm; worldwide distribution; about 550 species.

      Family Polyphagidae

 Middle and hindfemurs mostly with few strong spines ventrally; spines on front and back ventral margins arranged differently; anal area of folded hindwing not plaited fanlike; postclypeus usually enlarged; range in size about 3 to 35 mm; worldwide distribution; about 200 species.

      Family Blattellidae

 Variable, mostly small species; mostly with numerous similar spines on ventral margins of middle and hindfemurs; ranging in size from 3 to 100 mm, with an average of about 12 mm; worldwide distribution; about 1,700 species.

      Family Blaberidae

 Mostly large species; spines of middle and hindfemurs variable; male subgenital plate asymmetrical; viviparous or ovoviparous; from 10 to 60 mm in size; worldwide in distribution; about 650 species.

      Suborder Mantodea (mantids)

 Head usually conspicuous anterior to a narrow pronotum, seldom concealed by a broad pronotum; 3 distinct ocelli; front legs adapted for seizing prey; proventriculus not heavily armed, inner lining with fine anastomosing ridges between teeth of moderate size.

      Family Mantidae

 Oligocene to present (fossils rare); size from 20 to 155 mm, average about 60 mm, worldwide distribution; about 2,000 species.

      Order Grylloblattodea (Notoptera)

 Legs similar, adapted for running; 5-segmented tarsi; wingless; eyes small or absent; no ocelli; antennae fairly long, filiform, about 25–40 segments; female with well-developed ovipositor, resembling Tettigoniidae (see below under Orthoptera); cerci long, with 8 to 9 segments.

      Family Grylloblattidae (ice bugs (ice bug))

 Recent (no fossils); size about 20–25 mm; found in northwestern North America, eastern Siberia, Japan; about 15 species.

      Order Phasmida (Cheleutoptera or Phasmatoptera)

 Legs similar, adapted for walking; tarsi nearly always 5-segmented; often wingless; when winged, tegmina often shorter than wings; wing pads not reversed as nymphs; ovipositor short, often concealed; male genitalia asymmetrical; cerci short, unsegmented; no tympanum or conspicuous stridulatory organs; usually very slender, elongated, sometimes broadened, even leaflike.

      Family Phasmatidae (walking sticks, stick insects)

 Variable, rarely less than an inch long; a broadly conceived family; Triassic to present; 15 to 325 mm in size with average about 70 mm; worldwide distribution centred in warm countries; about 2,000 species.

      Family Phylliidae (leaf insects)

 Depressed, leaflike, winged; tegmina short in males, covering most of abdomen in females; antennae long, pubescent in males, very short in females; all members recent; about 60 mm in size; Indo-Malayan distribution from Ceylon to Fiji; about 25 species.

      Order Orthoptera

 Hindlegs almost always enlarged and adapted for jumping; tarsi usually 3- or 4-segmented, occasionally fewer than 3 or as many as 5 segments; front wings more or less thickened; wing pads reversed (hindwing partly covering front wing) during late nymphal stages; cerci nearly always unsegmented; specialized auditory and stridulatory organs often present.

      Suborder Ensifera

 Antennae usually long, with more than 30 segments; auditory organs, if present, consist of tympanum at base of front tibia; many species with stridulatory mechanism at base of tegmen; ovipositor usually present as rounded or flattened bladelike, or elongated cylindrical spearlike, structure.

      Superfamily Prophalangopsoidea

 Tarsi compressed or cylindrical, 3- or 4-segmented; tympanum present; simple stridulatory vein on each male tegmen.

      Family Prophalangopsidae

 Triassic to present. 25–47 mm in size; recent species found in India and northwestern North America; 3 species.

      Superfamily Tettigonioidea

 Tarsi usually depressed, 4-segmented, at least on middle and hind legs; tympanum present; stridulatory organ specialized, with left tegmen uppermost and scraper (at edge of lower tegmen) often better developed on right tegmen.

      Family Tettigoniidae (katydids, bush crickets)

 Jurassic to present; size range from 15 to 120 mm, average about 35 mm; distribution worldwide; about 3,000 species.

      Family Phasmodidae (phasmodids)

 Very slender, resembling Phasmatidae (see above); head prognathous (i.e., mouth directed forward instead of downward); hind femurs not enlarged; all members recent; found in Australia; size range from 25 to 85 mm; 2 species.

      Superfamily Gryllacridoidea

 Tarsi depressed, usually 4-segmented; front wings without stridulatory mechanism; usually no tympanum; ovipositor usually flattened, bladelike; 12 to 50 mm in size with average 25 mm.

      Family Stenopelmatidae (stenopelmatids, Jerusalem crickets)

 Heavy body, often wingless, with front legs usually strongly armed for digging; ovipositor inconspicuous; represented from Miocene to present; members found in southeastern Asia, Southern Africa, North America, and Central America; about 35 species.

      Family Gryllacrididae (gryllacridids)

 Usually wingless or with vestigial wings; found from Jurassic to present; chiefly tropical, most nocturnal, in trees and shrubs; 1 species in U.S.; about 550 species.

      Family Schizodactylidae (schizodactylids)

 Winged or wingless; when winged, wing tips very long, rolled in a coil at rest; tarsal segments 2 and 3 with conspicuous lateral lobes; all members recent, found in southern Asia, South Africa; about 8 species.

      Family Rhaphidophoridae (camel crickets and allies)

 Wingless; usually of humpback appearance; tarsi compressed; legs often very long and slender; found from Oligocene to present; worldwide distribution; about 300 species.

      Family Lezinidae

 Wingless; cerci with apical segmentation; found in Asia and Africa; about 10 species.

      Family Henicidae

 Usually wingless; front tibia with 1 or more dorsal spurs; front coxa usually with spine; represented in Old and New World tropics; about 150 species.

      Superfamily Grylloidea

 Tarsi 3-segmented; tegmina of males usually highly developed for stridulation; tympanum usually at base of front tibia; ovipositor most often round in cross section, spearlike, other times somewhat flattened or reduced; 2 to 50 mm in size with average 18 mm.

      Family Gryllidae (crickets)

 Front legs usually not for digging; ovipositor usually elongated; species from Triassic to present; worldwide distribution; about 2,400 species.

      Family Gryllotalpidae (mole crickets)

 Front legs highly modified for digging, with tibiae expanded and with fingerlike dactyls; ovipositor vestigial; represented from Oligocene to present; distribution worldwide; about 65 species.

      Suborder Caelifera

 Antennae short, with less than 30 segments; auditory organs, if present, at base of abdomen in form of tympanum; ovipositor usually consists of paired valvular appendages adapted for digging.

      Superfamily Tridactyloidea

 Tarsi 1- or 2-segmented; antennae short, with 12 or fewer segments.

      Family Tridactylidae (pygmy sand crickets)

 Front tarsi 2-segmented; hind tibiae with unsegmented tarsus, no “claws,” 2 long “paddles”; usually winged; 3 to 18 mm in size; worldwide distribution; about 60 species.

      Family Cylindrachaetidae (false mole crickets)

 Body elongated, cylindrical, wingless; antennae 7- to 8-segmented; eyes reduced; ocelli absent; front tibia with 4 or 5 dactyls; cerci very short. All members recent; inhabit Australia and southern South America; size 40–60 mm; about 8 species.

      Superfamily Tetrigoidea

 Pronotum strongly elongated, extending backward over most or sometimes all of abdomen; tegmina short, scalelike; hind wings usually fully developed; no tympanum; front and middle tarsi 2-segmented; hind tarsus 3-segmented.

      Family Tetrigidae (pygmy grasshoppers or grouse locusts)

 Miocene to present; 8–25 mm in size with average about 14 mm; worldwide distribution; about 700 species.

      Superfamily Eumastacoidea

 Abdominal spiracles situated in lateral membrane between terga and sternums; basal abdominal terga without auditory tympanum; basal segment of hind tarsus with serrated margins or at least with an external tubercle (all U.S. species wingless).

      Family Eumastiacidae (eumastacids or monkey grasshoppers)

 Sides of abdomen without stridulatory organ; antennae relatively short; frontal costa forked below median ocellus; represented from Miocene to present; worldwide in warm countries; about 200 species.

      Family Tanaoceridae (tanaocerids)

 Specialized stridulatory organ on side of 3rd abdominal tergum, rubbed by hind femur; antennae very long; frontal costa not forked; found in southwestern U.S.; 3 species.

      Superfamily Pneumoroidea

 Body robust; male fully winged, female short winged; hind legs short; tympanum absent; male with stridulatory ridges on side of 3rd abdominal tergum.

      Family Pneumoridae (pneumorids)

 All members recent; restricted to southern Africa; 11 to 107 mm in size; about 18 species.

      Superfamily Proscopioidea

 Body sticklike, superficially like Phasmatidae (see above under Order Phasmida); head elongate, conical, usually oblique or vertical; wings rarely present, tympanum absent; no stridulatory organ.

      Family Proscopiidae (proscopiids)

 All representatives recent; from Costa Rica to southern South America; size 30 to 180 mm with average about 60 mm; about 100 species.

      Superfamily Acridoidea

 Typically robust, with both pairs of wings and tympanum present, although extremely variable; pronotum not or briefly prolonged over abdomen; abdominal spiracles located in terga; tarsi 3-segmented, basal segment without serrated margins; size 8 to 120 mm with average about 28 mm.

      Family Acrididae (grasshoppers)

 Many of unusual appearance; stridulatory mechanism often present, but not with organ on abdomen; vertex without a definite longitudinal furrow; tympanum normally present (absent in some, especially wingless genera); ocelli small; hind tibiae usually not expanded apically for swimming; Eocene to present; worldwide distribution; more than 5,000 species.

      Family Pamphagidae (pamphagids)

 Usually large; wings variable; many wingless bulky species in arid regions; vertex of head with longitudinal furrow; lower basal lobe of hind femur longer than upper lobe; members recent; in Africa and southern parts of Europe and Asia; about 300 species.

      Family Pyrgomorphidae (pyrgomorphids)

 Head conical; longitudinal furrow of vertex present; lower basal lobe of femur longer than upper lobe; tympanum usually present; no stridulatory mechanism; recent; worldwide in tropical and subtropical areas, unknown in U.S.; about 350 species.

      Each of the following 7 families contains only a few species: Pauliniidae, Xyronotidae, Ommexechidae, Trigonopterygidae, Charilaidae, Lathiceridae, Lentulidae. The Xyronotidae includes 1 Mexican species only; the other families are not represented in North America.

Critical appraisal

      The arrangement of orders, suborders, superfamilies, and families presented above is a consensus of current opinion; however, some entomologists recognize different relationships and additional families too minor or too little understood to be included here. In most large groups, there are a few peculiar species that vary slightly from the characteristics described. Although an attempt was made in this classification to indicate the earliest geologic period in which major groups were found, available information is not always clear as to whether the insect found belongs to a modern family or an ancestral one with more primitive features and a different name. The known species shown for each group represent only living ones that have been named, not fossils. In some cases, modern catalogs and monographs can be used to obtain accurate counts; for other cases, however, estimates are given.

      Several other orders of insects are orthopteroid in their general relationships. Among them, the Dermaptera (earwigs (earwig)) differ from orthopterans by having short leathery front wings devoid of veins, hind wings with veins radiating from a central point midway of the anterior margin, and most cerci modified into pincer-like structures. Isoptera (termites, sometimes placed in Dictyoptera) have similar front and hind wings; although in many ways they resemble some cockroaches, they differ in their elaborate caste system and their habits (i.e., living in complex colonies consisting of reproductive individuals, sterile workers, and soldiers). Zorapterans (zorapteran) show some morphological relationship to cockroaches but have two-segmented tarsi, peculiar wing venation, a primitive caste system, and other differences. Embiopterans (webspinner) (web spinners) are also orthopteroid in basic morphology, but are notably distinct from orthopterans by the much enlarged silk-producing basal segment of the front tarsus. Plecoptera (stonefly) (stoneflies) are also orthopteroid, but their front and hindwings are of a similar texture (unlike orthopterans), and their immature stages are specialized for an aquatic life.

Ashley B. Gurney

Additional Reading

R.D. Alexander, “Acoustical Communication in Arthropods,” A. Rev. Ent., 12:495–526 (1967), with many orthopteran examples; “Life Cycle Origins, Speciation, and Related Phenomena in Crickets,” Qt. Rev. Biol., 43:1–41 (1968), a valuable summary of modern advances in cricket biology; and with D. Otte, “The Evolution of Genitalia and Mating Behavior in Crickets (Gryllidae) and Other Orthoptera,” Misc. Publs. Mus. Zool. Univ. Mich., 133:5–62 (1967), well illustrated; E.D. Ball et al., “The Grasshoppers and Other Orthoptera of Arizona,” Tech. Bull. Univ. Ariz., 93:257–373 (1942), well illustrated; W.S. Blatchley, Orthoptera of Northeastern America (1920), an important guide, some parts out of date; I.J. Cantrall, “The Ecology of the Orthoptera and Dermaptera of the George Reserve, Michigan,” Misc. Publs. Mus. Zool. Univ. Mich., 54:1–182 (1943), a survey of 78 species of orthopterans; R.F. Chapman, The Insects: Structure and Function, 3rd ed. (1982), with many orthopteran examples; L. Chopard, “Dictyoptères,” “Notoptères,” “Chéleutoptères,” and “Orthoptères,” in Traité de zoologie, vol. 9 (1949), a general treatise on Orthoptera; V.M. Dirsh, “A Preliminary Revision of the Families and Subfamilies of Acridoidea,” Bull. Br. Mus. Nat. Hist. (Ent.), 10:351–419 (1961), the current classification of grasshoppers; “The Post-Embryonic Ontogeny of Acridimorpha (Orthoptera),” EOS, Madr., 43:413–514 (1968), basic information on growth changes in grasshoppers; S.K. Gangwere, “A Monograph on Food Selection in Orthoptera,” Trans. Am. Ent. Soc., 87:67–230 (1961), a summary of techniques for studying food preferences of orthoptera and a review of information on the food eaten by various orthopteran groups, chiefly U.S. species; A.B. Gurney, “Praying Mantids of the United States: Native and Introduced,” A. Rep. Smithson. Instn. for 1950, pp. 339–362 (1951), a semipopular review with photos of several species; Daniel Otte, The North American Grasshoppers, 3 vol. (1981– ), a revision of the taxonomic order. D.M. Guthrie and A.R. Tindall, The Biology of the Cockroach (1968), emphasis on the physiology of cockroach species commonly used as laboratory animals; P.T. Haskell, Insect Sounds (1961), many references to orthoptera; M. Hebard, “The Dermaptera and Orthoptera of Illinois,” Bull. Ill. St. Nat. Hist. Surv., 20:125–279 (1934), keys to species, illustrated; J.R. Helfer, How to Know the Grasshoppers, Cockroaches and Their Allies (1963, reissued 1987), a semipopular handbook; A.D. Imms, Imms' General Textbook of Entomology, 10th ed. rev. by O.W. Richards and R.G. Davies, 2 vol. (1977), an important but detailed text; F.A. McKittrick, “Evolutionary Studies of Cockroaches,” Mem. Cornell Univ. Agric. Exp. Stn. 389 (1964), classification of cockroaches; K. Princis, “Blattariae,” in Orthopterorum Catalogus, 8 pt. (1962–71), a catalog of living cockroaches, of use to researchers in systematics and distribution; D.R. Ragge, Grasshoppers, Crickets and Cockroaches of the British Isles (1965), a well-illustrated standard work; J.A.G. Rehn, “Man's Uninvited Fellow Traveler, the Cockroach,” Sci. Mon., 61: 265–276 (1945), popular and factual; J.A.G. Rehn and H.J. Grant, A Monograph of the Orthoptera of North America (North of Mexico), vol. 1 (1961), detailed and authoritative; L.M. Roth, “Oothecae of the Blattaria,” A. Ent. Soc. Am., 61:83–111 (1968), an illustrated review; “The Evolution of Male Tergal Glands in the Blattaria,” A. Ent. Soc. Am., 62:176–208 (1969), a review.“Evolution and Taxonomic Significance of Reproduction in Blattaria,” A. Rev. Ent., 15:75–96 (1970), a summary of reproductive habits; with T. Eisner, “Chemical Defenses of Arthropods,” A. Rev. Ent., 7:107–136 (1962), concerned with secretory glands that produce repugnant substances; and with E.R. Willis, The Biotic Associations of Cockroaches (1960), a documented summary; A.G. Sharov, in Trans. Paleont. Inst., Moscow, 118:1–208 (1968; Eng. trans., “Phylogeny of the Orthopteroidea,” pp. 1–251, 1971), evolutionary development; B.P. Uvarov, Grasshoppers and Locusts, vol. 1 (1966), the first of a 2-volume book on this subject; T.J. Walker, “Cryptic Species Among Sound-Producing Ensiferan Orthoptera (Gryllidae and Tettigoniidae),” Q. Rev. Biol., 39:345–355 (1964), emphasis on the large number of orthoptera species reproductively isolated by inconspicuous calling songs; F.E. Zeuner, Fossil Orthoptera Ensifera, 2 vol. (1939), a review with excellent illustrations.Ashley B. Gurney

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