Crickets (also known as "true crickets"), of the family Gryllidae, are insects related to bush crickets, and, more distantly, to grasshoppers. The Gryllidae have mainly cylindrical bodies, round heads, and long antennae. Behind the head is a smooth, robust pronotum. The abdomen ends in a pair of long cerci (spikes); females have a long, cylindrical ovipositor. The hind legs have enlarged femora (thighs), providing power for jumping. The front wings are adapted as tough, leathery elytra (wing covers), and some crickets chirp by rubbing parts of these together. The hind wings are membranous and folded when not in use for flight; many species, however, are flightless. The largest members of the family are the bull crickets, Brachytrupes, which are up to 5 cm (2 in) long.
More than 900 species of crickets are described; the Gryllidae are distributed all around the world except at latitudes 55° or higher, with the greatest diversity being in the tropics. They occur in varied habitats from grassland, bushes, and forests to marshes, beaches, and caves. Crickets are mainly nocturnal, and are best known for the loud, persistent, chirping song of males trying to attract females, although some species are mute. The singing species have good hearing, via the tympana (eardrums) on the tibiae of the front legs.
Crickets often appear as characters in literature. The Talking Cricket features in Carlo Collodi's 1883 children's book, The Adventures of Pinocchio, and in films based on the book. The eponymous insect is central to Charles Dickens's 1845 The Cricket on the Hearth, as is the chirping insect in George Selden's 1960 The Cricket in Times Square. Crickets are celebrated in poems by William Wordsworth, John Keats, and Du Fu. They are kept as pets in countries from China to Europe, sometimes for cricket fighting. Crickets are efficient at converting their food into body mass, making them a candidate for food production. They are used as food in Southeast Asia, where they are sold deep-fried in markets as snacks. They are also used to feed carnivorous pets and zoo animals. In Brazilian folklore, crickets feature as omens of various events.
| Scientific classification | |
|---|---|
| Kingdom: | Animalia |
| Clade: | Euarthropoda |
| Class: | Insecta |
| Order: | Orthoptera |
| Suborder: | Ensifera |
| Superfamily: | Grylloidea |
| Family: | Gryllidae Laicharting, 1781 |
| Subfamilies | |
| See Taxonomy section | |
| Synonyms | |
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Description
Crickets are small to medium-sized insects with mostly cylindrical, somewhat vertically flattened bodies. The head is spherical with long slender antennae arising from cone-shaped scapes (first segments) and just behind these are two large compound eyes. On the forehead are three ocelli (simple eyes). The pronotum (first thoracic segment) is trapezoidal in shape, robust, and well-sclerotinized. It is smooth and has neither dorsal or lateral keels (ridges).At the tip of the abdomen is a pair of long cerci (paired appendages on rearmost segment), and in females, the ovipositor is cylindrical, long and narrow, smooth and shiny. The femora (third segments) of the back pair of legs are greatly enlarged for jumping. The tibiae (fourth segments) of the hind legs are armed with a number of moveable spurs, the arrangement of which is characteristic of each species. The tibiae of the front legs bear one or more tympani which are used for the reception of sound.
The wings lie flat on the body and are very variable in size between species, being reduced in size in some crickets and missing in others. The fore wings are elytra made of tough chitin, acting as a protective shield for the soft parts of the body and in males, bear the stridulatory organs for the production of sound. The hind pair is membranous, folding fan-wise under the fore wings. In many species, the wings are not adapted for flight.
The largest members of the family are the 5 cm (2 in)-long bull crickets (Brachytrupes) which excavate burrows a metre or more deep. The tree crickets (Oecanthinae) are delicate white or pale green insects with transparent fore wings, while the field crickets (Gryllinae) are robust brown or black insects.
Distribution and habitat
Crickets have a cosmopolitan distribution, being found in all parts of the world with the exception of cold regions at latitudes higher than about 55° North and South. They have colonised many large and small islands, sometimes flying over the sea to reach these locations, or perhaps conveyed on floating timber or by human activity. The greatest diversity occurs in tropical locations, such as in Malaysia, where 88 species were heard chirping from a single location near Kuala Lumpur. A greater number than this could have been present because some species are mute.Crickets are found in many habitats. Members of several subfamilies are found in the upper tree canopy, in bushes, and among grasses and herbs. They also occur on the ground and in caves, and some are subterranean, excavating shallow or deep burrows. Some make galleries in rotting wood, and certain beach-dwelling species can run and jump over the surface of pools.
Biology
Defence
Crickets are relatively defenceless, soft-bodied insects. Most species are nocturnal and spend the day hidden in cracks, under bark, inside curling leaves, under stones or fallen logs, in leaf litter, or in the cracks in the ground that develop in dry weather. Some excavate their own shallow holes in rotting wood or underground and fold in their antennae to conceal their presence. Some of these burrows are temporary shelters, used for a single day, but others serve as more permanent residences and places for mating and laying eggs. Crickets burrow by loosening the soil with the mandibles and then carrying it with the limbs, flicking it backwards with the hind legs or pushing it with the head.Other defensive strategies are the use of camouflage, fleeing, and aggression. Some species have adopted colourings, shapes, and patterns that make it difficult for predators that hunt by sight to detect them. They tend to be dull shades of brown, grey, and green that blend into their background, and desert species tend to be pale. Some species can fly, but the mode of flight tends to be clumsy, so the most usual response to danger is to scuttle away to find a hiding place.
Chirping
Most male crickets make a loud chirping sound by stridulation. The stridulatory organ is located on the tegmen, or fore wing, which is leathery in texture. A large vein runs along the centre of each tegmen, with comb-like serrations on its edge forming a file-like structure, and at the rear edge of the tegmen is a scraper. The tegmina are held at an angle to the body and rhythmically raised and lowered which causes the scraper on one wing to rasp on the file on the other. The central part of the tegmen contains the "harp", an area of thick, sclerotinized membrane which resonates and amplifies the volume of sound, as does the pocket of air between the tegmina and the body wall. Most female crickets lack the necessary adaptations to stridulate, so make no sound.Several types of cricket songs are in the repertoire of some species. The calling song attracts females and repels other males, and is fairly loud. The courting song is used when a female cricket is near and encourages her to mate with the caller. A triumphal song is produced for a brief period after a successful mating, and may reinforce the mating bond to encourage the female to lay some eggs rather than find another male. An aggressive song is triggered by contact chemoreceptors on the antennae that detect the presence of another male cricket.
Crickets chirp at different rates depending on their species and the temperature of their environment. Most species chirp at higher rates the higher the temperature is (about 62 chirps a minute at 13 °C (55 °F) in one common species; each species has its own rate). The relationship between temperature and the rate of chirping is known as Dolbear's law. According to this law, counting the number of chirps produced in 14 seconds by the snowy tree cricket, common in the United States, and adding 40 will approximate the temperature in degrees Fahrenheit.
In 1975, Dr. William H. Cade discovered that the parasitic tachinid fly Ormia ochracea is attracted to the song of the cricket, and uses it to locate the male to deposit her larvae on him. It was the first known example of a natural enemy that locates its host or prey using the mating signal. Since then, many species of crickets have been found to be carrying the same parasitic fly, or related species. In response to this selective pressure, a mutation leaving males unable to chirp was observed amongst a population of field crickets on the Hawaiian island of Kauai, enabling these crickets to elude their parasitoid predators. A different mutation with the same effect was also discovered on the neighboring island of Oahu (ca. 100 miles (160 km) away).
Flight
Some species, such as the ground crickets (Nemobiinae), are wingless; others have small fore wings and no hind wings (Copholandrevus), others lack hind wings and have shortened fore wings in females only, while others are macropterous, with the hind wings longer than the fore wings. In Teleogryllus, the proportion of macropterous individuals varies from very low to 100%. Probably, most species with hind wings longer than fore wings engage in flight.Some species, such as Gryllus assimilis, take off, fly, and land efficiently and well, while other species are clumsy fliers. In some species, the hind wings are shed, leaving wing stumps, usually after dispersal of the insect by flight. In other species, they may be pulled off and consumed by the cricket itself or by another individual, probably providing a nutritional boost.
Gryllus firmus exhibits wing polymorphism; some individuals have fully functional, long hind wings and others have short wings and cannot fly. The short-winged females have smaller flight muscles, greater ovarian development, and produce more eggs, so the polymorphism adapts the cricket for either dispersal or reproduction. In some long-winged individuals, the flight muscles deteriorate during adulthood and the insect's reproductive capabilities improve.
Diet
Captive crickets are omnivorous; when deprived of their natural diet, they accept a wide range of organic foodstuffs. Some species are completely herbivorous, feeding on flowers, fruit, and leaves, with ground-based species consuming seedlings, grasses, pieces of leaf, and the shoots of young plants. Others are more predatory and include in their diet invertebrate eggs, larvae, pupae, moulting insects, scale insects, and aphids. Many are scavengers and consume various organic remains, decaying plants, seedlings, and fungi. In captivity, many species have been successfully reared on a diet of ground, commercial dry dog food, supplemented with lettuce and aphids.Crickets have relatively powerful jaws, and several species have been known to bite humans.
Reproduction and lifecycle
Male crickets establish their dominance over each other by aggression. They start by lashing each other with their antennae and flaring their mandibles. Unless one retreats at this stage, they resort to grappling, at the same time each emitting calls that are quite unlike those uttered in other circumstances. When one achieves dominance, it sings loudly, while the loser remains silent.Females are generally attracted to males by their calls, though in nonstridulatory species, some other mechanism must be involved. After the pair has made antennal contact, a courtship period may occur during which the character of the call changes. The female mounts the male and a single spermatophore is transferred to the external genitalia of the female. Sperm flows from this into the female's oviduct over a period of a few minutes or up to an hour, depending on species. After copulation, the female may remove or eat the spermatophore; males may attempt to prevent this with various ritualised behaviours. The female may mate on several occasions with different males.
Most crickets lay their eggs in the soil or inside the stems of plants, and to do this, female crickets have a long, needle-like or sabre-like egg-laying organ called an ovipositor. Some ground-dwelling species have dispensed with this, either depositing their eggs in an underground chamber or pushing them into the wall of a burrow. The short-tailed cricket (Anurogryllus) excavates a burrow with chambers and a defecating area, lays its eggs in a pile on a chamber floor, and after the eggs have hatched, feeds the juveniles for about a month.
Crickets are hemimetabolic insects, whose lifecycle consists of an egg stage, a larval or nymph stage that increasingly resembles the adult form as the nymph grows, and an adult stage. The egg hatches into a nymph about the size of a fruit fly. This passes through about 10 larval stages, and with each successive moult, it becomes more like an adult. After the final moult, the genitalia and wings are fully developed, but a period of maturation is needed before the cricket is ready to breed.
The species Gryllus bimaculatus exhibits the reproductive pattern of polyandry. This sexual selective pattern increases the overall fitness of the species and promotes genetic variation. The females select and mate with multiple viable sperm donors and exhibit a distinct preference for novel mates.
Inbreeding avoidance
Female Teleogryllus oceanicus crickets from natural populations mate polyandrously and store sperm from multiple males. Female crickets exert a postcopulatory fertilization bias in favour of unrelated males to avoid the genetic consequences of inbreeding. Fertilization bias depends on the control of sperm transport to the sperm storage organs. The inhibition of sperm storage by female crickets can act as a form of cryptic female choice to avoid the severe negative effects of inbreeding.In controlled-breeding experiments with the cricket Gryllus firmus, seven inbred lines were tested, and during 14 generations of brother–sister matings, nymphal weight and early fecundity declined substantially. This observed inbreeding depression appeared to be due to an increased frequency of homozygous combinations of deleterious recessive alleles generated by the inbreeding (thereby decreasing fitness). These results support the general idea that the principal benefit of outcrossing is the masking of deleterious recessive alleles by wild-type alleles.
Phylogeny and taxonomy
The phylogenetic relationships of the Gryllidae, summarized by Darryl Gwynne in 1995 from his own work (using mainly anatomical characteristics) and that of earlier authors,[a] are shown in the following cladogram, with the Orthoptera divided into two main groups, Ensifera (crickets sensu lato) and Caelifera (grasshoppers). Fossil Ensifera are found from the late Carboniferous period (300 Mya) onwards, and the true crickets, Gryllidae, from the Triassic period (250 to 200 Mya).A phylogenetic study by Jost & Shaw in 2006 using sequences from 18S, 28S, and 16S rRNA supported the monophyly of Ensifera. Most ensiferan families were also found to be monophyletic, and the superfamily Gryllacridoidea was found to include Stenopelmatidae, Anostostomatidae, Gryllacrididae and Lezina. Schizodactylidae and Grylloidea were shown to be sister taxa, and Rhaphidophoridae and Tettigoniidae were found to be more closely related to Grylloidea than had previously been thought. The authors stated that "a high degree of conflict exists between the molecular and morphological data, possibly indicating that much homoplasy is present in Ensifera, particularly in acoustic structures." They considered that tegmen stridulation and tibial tympanae are ancestral to Ensifera and have been lost on multiple occasions, especially within the Gryllidae.
