Beetles are one of the most diverse groups of insects. Their order, Coleoptera (meaning "sheathed wing"), has more species in it than in any other order in the animal kingdom. Forty percent of all described insect species are beetles (about 350,000 species), and new species are regularly discovered. Approximately one out of every five animals on our planet is a beetle (20% of all animal life). Estimates put the total number of species, described and undescribed, at between 5 and 8 million. This is why when J. B. S. Haldane, a Scottish geneticist, was asked what his studies of nature revealed about God, he replied, "An inordinate fondness for beetles".
Beetles can be found in almost all habitats, but are not known to occur in the sea or in the polar regions. They have a major impact on the ecosystem in three ways: feeding on plants and fungi, breaking down animal and plant debris, and eating other invertebrates. Certain species are agricultural pests, such as the Colorado potato beetle, Leptinotarsa decemlineata, while others are important controls of agricultural pests. For example, lady beetles (family Coccinellidae) consume aphids, fruit flies, thrips, and other plant-sucking insects that damage crops.
Beetles are generally characterised by a particularly hard exoskeleton and hard forewings (elytra). The beetle's exoskeleton is made up of numerous plates called sclerites, separated by thin sutures. This design creates the armoured defences of the beetle while maintaining flexibility. The elytra are not used for flight, but tend to cover the hind part of the body and protect the second pair of wings (alae). Elytra must generally be raised in order to move the hind flight wings. A beetle's flight wings are crossed with veins and, after landing, are folded, often along these veins, and stored below the elytra. In some cases the ability to fly has been lost, most notably in the ground beetles (family Carabidae) and the true weevils(family Curculionidae), but also in some desert and cave-dwelling species of other families. Many of these species have the two elytra fused together, forming a solid shield over the abdomen. In a few families both the ability to fly and the elytra have been lost, with the best known example being the glowworms of the family Phengodidae, in which the females are larviform throughout their lives.
Beetles have mouthparts similar to those of grasshoppers. Of these parts, the most commonly known are likely the mandibles, which appear as large pincers on the front of some beetles. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see Predation, below). Two pairs of finger-like appendages are found around the mouth in most beetles, serving to move food into the mouth. These are the maxillary and labial palpi.
The eyes are compound, and may display some remarkable adaptability, as in the case of whirligig beetles (family Gyrinidae), in which the eyes are split to allow a view both above and below the waterline. Other species also have divided eyes (some Cerambycidae and Curculionidae), while many beetles have eyes that are notched to some degree. A few beetle genera also possess ocelli, which are small, simple eyes usually situated farther back on the head (on the vertex).
Beetle antennae are primarily organs of smell, but may also be used to physically feel out a beetle's environment. Further, they may be used in some families during mating, or among a few beetles for defence. Antennae vary greatly in form within the Coleoptera, but are often similar within any given family. In some cases males and females of the same species will have different antennal forms. Antennae may be filiform, clavate, flabellate or genticulate.
The legs, which are multi-segmented, end in two to five small segments called tarsi, which are vaguely comparable to feet. Like many other insect orders beetles bear claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs may be variously modified and adapted for other uses. Among aquatic families (Dytiscidae, Haliplidae, many Hydrophilidae, and others) the legs, most notably the hind pair, are modified for swimming and often bear rows of long hairs to aid this purpose. Other beetles have fossorial legs that are widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (family Histeridae). The hind legs of some beetles, such as flea beetles (within Chrysomelidae) and flea weevils (within Curculionidae), are enlarged and designed for jumping.
Oxygen is obtained via a tracheal system. Air enters a series of tubes along the body through openings called spiracles, and is then taken into increasingly finer fibres. Pumping movements of the body force the air through the system.
Beetles have hemolymph instead of blood, and the open circulatory system of the beetle is powered by a tube-like heart attached to the top inside of the thorax.
Various techniques are employed for retaining both air and water supplies. For example, predaceous diving beetles (family Dytiscidae) employ a technique of retaining air, when diving, between the abdomen and the elytra.
Beetles are endopterygotes with complete metamorphosis.
The eggs of beetles are very small. Some may be brightly coloured. A single female may lay from several dozen to several thousand eggs, placing them in clumps.
The larva of a beetle is called a grub, and often represents the principal feeding stage of the life-cycle. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles and lady bird beetles, while others feed within their food sources. The larval period varies between species but can be as long as several years. All beetle larvae go through several instars, which are the developmental stages between each moult. In many species the larvae simply increase in size with each successive instar. In some cases, however, more dramatic changes occur. Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the planidium) is highly mobile in order to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as hypermetamorphosis; examples include the blister beetles (family Meloidae) and some rove beetles, particularly those of the genus Aleochara.
As with Lepidoptera, beetle larvae pupate for a period of time, and from the pupa emerges a fully formed, sexually mature adult beetle, or imago. Adults have an extremely variable lifespan, from weeks to years, depending on the species.
Conflict can play a part in the mating rituals of species such as burying beetles (genus Nicrophorus) where conflicts between males and females rage until only one of each is left, thus ensuring reproduction by the strongest and fittest. Many beetles are territorial and will fiercely defend their small patch of territory from intruding males.
Pairing is generally short but in some cases will last for several hours. During pairing sperm cells are transferred to the female to fertilise the egg.
There are other notable ways of caring for the eggs and young, such as those employed by leaf rollers, who bite sections of leaf causing it to curl inwards and then lay the eggs, thus protected, inside.
A number of longhorn beetles (family Cerambycidae) bear a striking resemblance to wasps. This defense, known as mimicry, can be found to a lesser extent in other beetle families, such as the scarab beetles.
Many species, including lady beetles and blister beetles, can secrete poisonous substances to make them unpalatable. These same species often exhibit aposematism, where bright or contrasting color patterns warn away potential predators.
Large ground beetles will tend to go on the attack, using their strong mandibles to forcibly persuade a predator to seek out easier prey.
The four extant suborders of beetle are these:
These suborders diverged in the Permian and Triassic. Their phylogenetic relationship is uncertain, with the most popular hypothesis being that Polyphaga and Myxophaga are most closely related, with Adephaga an outgroup to those two, and Archostemata an outgroup to the other three.
The large number of beetle species poses special problems for classification, with some families consisting of thousands of species and needing further division into subfamilies and tribes.
See the article subgroups of the order Coleoptera for a complete list of families and * for a complete list of World families and subfamilies.
Many agricultural, forestry, and household pests are represented by the order. These include:
Some farmers introduce beetle banks to foster and provide cover for beneficial beetles.
Several species of the dung beetles, most notably Scarabaeus sacer (often referred to as "scarab"), enjoyed a sacred status among the ancient Egyptians, as the creatures were likened to the god Khepri. Some scholars suggest that the people's practice of making mummies was inspired by the brooding process of the beetle.
Many thousands of amulets and stamp seals have been excavated that depict the scarab. In many artifacts, the scarab is depicted pushing the sun along its course in the sky. During and following the New Kingdom, scarab amulets were often placed over the heart of the mummified deceased. The amulets were often inscribed with a spell from the Book of the Dead which entreated the heart, "Do not stand as a witness against me."
Coleopterists have formed organizations to facilitate the study of beetles. Among these is The Coleopterists Society, an international organization based in the United States.
Research in this field is often published in peer-reviewed journals specific to the field of coleopterology, though journals dealing with general entomology also publish many papers on various aspects of beetle biology. Some of the journals specific to beetle research are:
There is a thriving industry in the collection of beetle specimens for amateur and professional collectors. Some countries have established laws governing or prohibiting the collection of certain rare (and often much sought after) species.
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