Grasshoppers are a group of insects belonging to the suborder Caelifera. They are amongst what are possibly the most ancient living groups of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago.
Grasshoppers are typically ground-dwelling insects with powerful hind legs which allow them to escape from threats by leaping vigorously. Their front legs are shorter and used for grasping food. As hemimetabolous insects, they do not undergo complete metamorphosis; they hatch from an egg into a nymph or "hopper" which undergoes five moults, becoming more similar to the adult insect at each developmental stage.[1] The grasshopper hears through the tympanal organ which can be found in the first segment of the abdomen attached to the thorax; while its sense of vision is in the compound eyes, a change in light intensity is perceived in the simple eyes (ocelli). At high population densities and under certain environmental conditions, some grasshopper species can change colour and behavior and form swarms. Under these circumstances, they are known as locusts.
Grasshoppers are plant-eaters, with a few species at times becoming serious pests of cereals, vegetables and pasture, especially when they swarm in the millions as locusts and destroy crops over wide areas. They protect themselves from predators by camouflage; when detected, many species attempt to startle the predator with a brilliantly coloured wing flash while jumping and (if adult) launching themselves into the air, usually flying for only a short distance. Other species such as the rainbow grasshopper have warning coloration which deters predators. Grasshoppers are affected by parasites and various diseases, and many predatory creatures feed on both nymphs and adults. The eggs are subject to attack by parasitoids and predators. Grasshoppers are diurnal insects, meaning they are most active during the day time.
Grasshoppers have had a long relationship with humans. Swarms of locusts can have devastating effects and cause famine, having done so since Biblical times.[2] Even in smaller numbers, the insects can be serious pests. They are used as food in countries such as Mexico and Indonesia. They feature in art, symbolism and literature. The study of grasshopper species is called acridology.
Grasshoppers belong to the suborder Caelifera. Although "grasshopper" has been used as a common name for the suborder in general,[3][4][5] modern sources restrict it to the more "evolved" families.[6] They may be placed in the infraorder Acrididea[7] and have been referred to as "short-horned grasshoppers" in older texts[8] to distinguish them from the also-obsolete term "long-horned grasshoppers" (now bush-crickets or katydids) with their much longer antennae. The phylogeny of the Caelifera, based on mitochondrial ribosomal RNA of thirty-two taxa in six out of seven superfamilies, is shown as a cladogram. The Ensifera (crickets, etc.), Caelifera and all the superfamilies of grasshoppers except Pamphagoidea appear to be monophyletic.[9][10]
In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the Permo-Triassic boundary;[11] the earliest insects that are certainly Caeliferans are in the extinct families Locustopseidae and Locustavidae from the early Triassic, roughly 250 million years ago. The group diversified during the Triassic and have remained important plant-eaters from that time to now. The first modern families such as the Eumastacidae, Tetrigidae and Tridactylidae appeared in the Cretaceous, though some insects that might belong to the last two of these groups are found in the early Jurassic.[12][13] Morphological classification is difficult because many taxa have converged towards a common habitat type; recent taxonomists have concentrated on the internal genitalia, especially those of the male. This information is not available from fossil specimens, and the palaeontological taxonomy is founded principally on the venation of the hindwings.[14]
The Caelifera includes some 2,400 valid genera and about 11,000 known species. Many undescribed species probably exist, especially in tropical wet forests. The Caelifera have a predominantly tropical distribution with fewer species known from temperate zones, but most of the superfamilies have representatives worldwide. They are almost exclusively herbivorous and are probably the oldest living group of chewing herbivorous insects.[14]
The most diverse superfamily is the Acridoidea, with around 8,000 species. The two main families in this are the Acrididae (grasshoppers and locusts) with a worldwide distribution, and the Romaleidae (lubber grasshoppers), found chiefly in the New World. The Ommexechidae and Tristiridae are South American, and the Lentulidae, Lithidiidae and Pamphagidae are mainly African. The Pauliniids are nocturnal and can swim or skate on water, and the Lentulids are wingless.[12] Pneumoridae are native to Africa, particularly southern Africa, and are distinguished by the inflated abdomens of the males.[15]
Grasshoppers have the typical insect body plan of head, thorax, and abdomen. The head is held vertically at an angle to the body, with the mouth at the bottom. The head bears a large pair of compound eyes which give all-round vision, three simple eyes which can detect light and dark, and a pair of thread-like antennae that are sensitive to touch and smell. The downward-directed mouthparts are modified for chewing and there are two sensory palps in front of the jaws.[16]
The thorax and abdomen are segmented and have a rigid cuticle made up of overlapping plates composed of chitin. The three fused thoracic segments bear three pairs of legs and two pairs of wings. The forewings, known as tegmina, are narrow and leathery while the hindwings are large and membranous, the veins providing strength. The legs are terminated by claws for gripping. The hind leg is particularly powerful. The legs of these species are so powerful that they can jump quite a long distance. they also use this to flee from danger.[17] The femur is robust and has several ridges where different surfaces join and the inner ridges bear stridulatory pegs in some species. The posterior edge of the tibia bears a double row of spines and there are a pair of articulated spurs near its lower end. The interior of the thorax houses the muscles that control the wings and legs.[16]
The abdomen has eleven segments, the first of which is fused to the thorax and contains the tympanal organ and hearing system. Segments two to eight are ring-shaped and joined by flexible membranes. Segments nine to eleven are reduced in size; segment nine bears a pair of cerci and segments ten and eleven have the reproductive organs. Female grasshoppers are normally larger than males, with short ovipositors.[16] The name of the suborder "Caelifera" comes from the Latin and means chisel-bearing, referring to the shape of the ovipositor.[18]
The grasshopper's auditory organs are located on its abdomen, rather than on its head. These organs consist of a pair of membranes, each positioned on either side of the first abdominal segment and tucked under the wings. Known as tympanal organs, these simple eardrums vibrate in response to sound waves, enabling the grasshopper to hear the songs of other grasshoppers.[19]
Those species that make easily heard noises usually do so by rubbing a row of pegs on the hind legs against the edges of the forewings (stridulation). These sounds are produced mainly by males to attract females, though in some species the females also stridulate.[20]
Grasshoppers may be confused with crickets, but they differ in many aspects; these include the number of segments in their antennae and the structure of the ovipositor, as well as the location of the tympanal organ and the methods by which sound is produced.[21] Ensiferans have antennae that can be much longer than the body and have at least 20–24 segments, while caeliferans have fewer segments in their shorter, stouter antennae.[20]
Most grasshoppers are polyphagous, eating vegetation from multiple plant sources,[22] but some are omnivorous and also eat animal tissue and animal faeces.[23] In general their preference is for grasses, including many cereals grown as crops.[24] The digestive system is typical of insects, with Malpighian tubules discharging into the midgut. Carbohydrates are digested mainly in the crop, while proteins are digested in the ceca of the midgut. Saliva is abundant but largely free of enzymes, helping to move food and Malpighian secretions along the gut. Some grasshoppers possess cellulase, which by softening plant cell walls makes plant cell contents accessible to other digestive enzymes.[25] Grasshoppers can also be cannibalistic when swarming.[26][27][28]
Grasshoppers have a typical insect nervous system, and have an extensive set of external sense organs. On the side of the head are a pair of large compo