Beetle

Etymology

Coleoptera comes from the Greek koleopteros, given to the group by Aristotle for their elytra, hardened shield-like forewings, from koleos "sheath", and pteron, "wing". The English name "beetle" comes from the Old English word bitela, little biter, related to bītan (to bite), leading to Middle English betylle. Another Old English name for beetle is ceafor, chafer, used in names such as cockchafer, from the Proto-Germanic *kabraz- (compare German Käfer).

Distribution and diversity

Beetles are by far the largest order of insects: the roughly 400,000 species make up about 40% of all insect species so far described, and about 25% of all animals. In 2015, Nigel Stork and colleagues provided four independent estimates of the total number of beetle species, giving a mean estimate of some 1.5 million with a "surprisingly narrow range" spanning all four estimates of 0.9 to 2.1 million beetle species. The four estimates made use of host-specificity relationships (1.5 to 1.9 million), ratios with other taxa (0.9 to 1.2 million), plant:beetle ratios (1.2 to 1.3), and body size by year of description (1.7 to 2.1 million).

Beetles are found in nearly all natural habitats, including freshwater and marine habitats, everywhere vegetative foliage is found, from trees and their bark to flowers, leaves, and underground near roots - even inside plants in galls, in every plant tissue, including dead or decaying ones.

The heaviest beetle, indeed the heaviest insect stage, is the larva of the goliath beetle, Goliathus goliatus, which can attain a mass of at least 115 g (4.1 oz) and a length of 11.5 cm (4.5 in). Adult male goliath beetles are the heaviest beetle in its adult stage, weighing 70–100 g (2.5–3.5 oz) and measuring up to 11 cm (4.3 in). Adult elephant beetles, Megasoma elephas and Megasoma actaeon often reach 50 g (1.8 oz) and 10 cm (3.9 in).

The longest beetle is the Hercules beetle Dynastes hercules, with a maximum overall length of at least 16.7 cm (6.6 in) including the very long pronotal horn. The smallest recorded beetle and the smallest free-living insect (as of 2015), is the featherwing beetle Scydosella musawasensis which may measure as little as 0.325 mm in length.

Permian

The oldest known insect that unequivocally resembles a Coleopteran is from the Lower Permian (270 mya), though it has 13-segmented antennae, elytra with more fully developed venation and more irregular longitudinal ribbing, and abdomen and ovipositor extending beyond the apex of the elytra. In the Permian–Triassic extinction event at the end of the Permian, some 30% of all insect species became extinct, so the fossil record of insects only includes beetles from the Lower Triassic (220 mya). Around this time, during the Late Triassic, fungus-feeding species such as Cupedidae appear in the fossil record. In the stages of the Upper Triassic, alga-feeding insects such as Triaplidae and Hydrophilidae begin to appear, alongside predatory water beetles. The first weevils, including the Obrienidae, appear alongside the first rove beetles (Staphylinidae), which closely resemble recent species. Some entomologists are sceptical that such early insects are so closely related to present-day species, arguing that this is extremely unlikely; for example, the structure of the metepisternum suggests that the Obrienidae could be Archostemata, not weevils at all, despite fossils with weevil-like snouts.

In 2009, a fossil beetle was described from the Pennsylvanian of Mazon Creek, Illinois, pushing the origin of the beetles to an earlier date, 318 to 299 million years ago. Fossils from this time have been found in Asia and Europe, for instance in the red slate fossil beds of Niedermoschel near Mainz, Germany. Further fossils have been found in Obora, Czech Republic and Tshekarda in the Ural mountains, Russia. However, there are only a few fossils from North America before the middle Permian, although both Asia and North America had been united to Euramerica. The first discoveries from North America made in the Wellington formation of Oklahoma were published in 2005 and 2008.

As a consequence of the Permian–Triassic extinction event, the fossil record of insects is scant, including beetles from the Lower Triassic. However, there are a few exceptions, such as in Eastern Europe. At the Babiy Kamen site in the Kuznetsk Basin, numerous beetle fossils were discovered, including entire specimens of the infraorders Archostemata (e.g. Ademosynidae, Schizocoleidae), Adephaga (e.g., Triaplidae, Trachypachidae) and Polyphaga (e.g. Hydrophilidae, Byrrhidae, Elateroidea). However, species from the families Cupedidae and Schizophoroidae are not present at this site, whereas they dominate at other fossil sites from the Lower Triassic. Further records are known from Khey-Yaga, Russia, in the Korotaikha Basin.

Jurassic

During the Jurassic (210 to 145 million years ago), there was a dramatic increase in the diversity of beetle families, including the development and growth of carnivorous and herbivorous species. The Chrysomeloidea diversified around the same time, feeding on a wide array of plant hosts from cycads and conifers to angiosperms. Close to the Upper Jurassic, the Cupedidae decreased, but the diversity of the early plant-eating species increased. Most recent plant-eating beetles feed on flowering plants or angiosperms, whose success contributed to a doubling of plant-eating species during the Middle Jurassic. However, the increase of the number of beetle families during the Cretaceous does not correlate with the increase of the number of angiosperm species. Around the same time, numerous primitive weevils (e.g. Curculionoidea) and click beetles (e.g. Elateroidea) appeared. The first jewel beetles (e.g. Buprestidae) are present, but they remained rare until the Cretaceous. The first scarab beetles were not coprophagous but presumably fed on rotting wood with the help of fungus; they are an early example of a mutualistic relationship.

There are more than 150 important fossil sites from the Jurassic, the majority in Eastern Europe and North Asia. Outstanding sites include Solnhofen in Upper Bavaria, Germany, Karatau in South Kazakhstan, the Yixian formation in Liaoning, North China, as well as the Jiulongshan formation and further fossil sites in Mongolia. In North America there are only a few sites with fossil records of insects from the Jurassic, namely the shell limestone deposits in the Hartford basin, the Deerfield basin and the Newark basin.

Cretaceous

The Cretaceous saw the fragmenting of the southern landmass, with the opening of the southern Atlantic Ocean and the isolation of New Zealand, while South America, Antarctica, and Australia grew more distant. The diversity of Cupedidae and Archostemata decreased considerably. Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute into different patterns; the Carabidae predominantly occurred in the warm regions, while the Staphylinidae and click beetles (Elateridae) preferred temperate climates. Likewise, predatory species of Cleroidea and Cucujoidea hunted their prey under the bark of trees together with the jewel beetles (Buprestidae). The diversity of jewel beetles increased rapidly, as they were the primary consumers of wood, while longhorn beetles (Cerambycidae) were rather rare: their diversity increased only towards the end of the Upper Cretaceous. The first coprophagous beetles are from the Upper Cretaceous, they may have lived on the excrement of herbivorous dinosaurs; The first species where both larvae and adults are adapted to an aquatic lifestyle are found. whirligig beetles (Gyrinidae) were moderately diverse, although other early beetles (e.g. Dytiscidae) were less, with the most widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae.

Many fossil sites worldwide contain beetles from the Cretaceous. Most are in Europe and Asia and belong to the temperate climate zone during the Cretaceous. Lower Cretaceous sites include the Crato fossil beds in the Araripe basin in the Ceará, North Brazil, as well as overlying Santana formation; the latter was near the equator at that time. In Spain, important sites are near Montsec and Las Hoyas. In Australia, the Koonwarra fossil beds of the Korumburra group, South Gippsland, Victoria, are noteworthy. Major sites from the Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia.

Cenozoic

Today'S beetle species developed in the Paleogene and Neogene. During this time, the continents began to be located closer to where they are today. Around 5 million years ago, the land bridge between South America and North America was formed, and the faunal exchange between Asia and North America began. Though many recent genera and species already existed during the Miocene, their distribution differed considerably from today's.

Phylogeny

The very large number of beetle species poses special problems for classification. Some families contain tens of thousands of species, and need to be divided into subfamilies and tribes. This immense number led the evolutionary biologist J. B. S. Haldane to quip, when some theologians asked him what could be inferred about the mind of the Creator from the works of His Creation, "An inordinate fondness for beetles". Polyphaga is the largest suborder, containing more than 300,000 described species in more than 170 families, including rove beetles (Staphylinidae), scarab beetles (Scarabaeidae), blister beetles (Meloidae), stag beetles (Lucanidae) and true weevils (Curculionidae). These beetles can be identified by the presence of cervical sclerites (hardened parts of the head used as points of attachment for muscles) absent in the other suborders. Adephaga contains about 10 families of largely predatory beetles, includes Ground beetles (Carabidae), water beetles (Dytiscidae) and whirligig beetles (Gyrinidae). In these insects, the testes are tubular and the first abdominal sternum (a plate of the exoskeleton) is divided by the hind coxae (the basal joints of the beetle's legs). Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and the telephone-pole beetle. Myxophaga contains about 65 described species in four families, mostly very small, including Hydroscaphidae and the genus Sphaerius.

The consistency of beetle morphology, in particular their possession of elytra, has long suggested that Coleoptera is monophyletic, though there have been doubts about the arrangement of the suborders, namely the Adephaga, Archostemata, Myxophaga and Polyphaga within that clade. The twisted-wing parasites, Strepsiptera, are the sister group to the beetles, having split from them in the Early Permian.

Molecular phylogenetic analysis confirms that the Coleoptera are a clade. Duane McKenna et al. 2015 used eight nuclear genes for 367 species from 172 of 183 Coleopteran families. They split the Adephaga into 2 clades, Hydradephaga and Geadephaga, broke up the Cucujoidea into 3 clades, and placed the Lymexyloidea within the Tenebrionoidea. The Polyphaga appear to date from the Triassic. Most extant beetle families appear to have arisen in the Cretaceous. The cladogram is based on McKenna 2015. The number of species in each group (mainly superfamilies) is shown in parentheses, and boldface if over 10,000. English names are given where possible. Dates of origin of major groups are shown in italics in millions of years ago (mya).

Digestive system

The digestive system of beetles is primarily adapted for a herbivorous diet. Digestion takes place mostly in the anterior midgut, although in predatory groups like the Carabidae, most digestion occurs in the crop by means of midgut enzymes. In the Elateridae, the predatory larvae defecate enzymes on their prey, with digestion being extraorally. The alimentary canal basically consists of a short, narrow pharynx, a widened expansion, the crop, and a poorly developed gizzard. After is the midgut, that varies in dimensions between species, with a large amount of cecum, and the hindgut, with varying lengths. There are typically four to six Malpighian tubules.

Nervous system

The nervous system in beetles contains all the types found in insects, varying between different species, from three thoracic and seven or eight abdominal ganglia which can be distinguished to that in which all the thoracic and abdominal ganglia are fused to form a composite structure.

Respiratory system

Like most insects, beetles inhale air, for the oxygen it contains, and exhale carbon dioxide, via a tracheal system. Air enters the body through spiracles, and circulates within the haemocoel in a system of tracheae and tracheoles, through whose walls the gases can diffuse.

Diving beetles, such as the Dytiscidae, carry a bubble of air with them when they dive. Such a bubble may be contained under the elytra or against the body by specialized hydrophobic hairs. The bubble covers at least some of the spiracles, permitting air to enter the tracheae. The function of the bubble is not so much to contain a store of air, but to act as a physical gill. The air that it traps is in contact with oxygenated water, so as the animal's consumption depletes the oxygen in the bubble, more oxygen can diffuse in to replenish it. Carbon dioxide is more soluble in water than either oxygen or nitrogen, so it readily diffuses out faster than in. Nitrogen is the most plentiful gas in the bubble, and the least soluble, so it constitutes a relatively static component of the bubble and acts as a stable medium for respiratory gases to accumulate in and pass through. Occasional visits to the surface are sufficient for the beetle to re-establish the constitution of the bubble.

Circulatory system

Like other insects, beetles have open circulatory systems, based on hemolymph rather than blood. As in other insects, a segmented tube-like heart is attached to the dorsal wall of the hemocoel. It has paired inlets or ostia at intervals down its length, and circulates the hemolymph from the main cavity of the haemocoel and out through the anterior cavity in the head.

Specialized organs

Different glands are specialized for different pheromones to attract mates. Pheromones from species of Rutelinea are produced from epithelial cells lining the inner surface of the apical abdominal segments; amino acid-based pheromones of Melolonthinae are produced from eversible glands on the abdominal apex. Other species produce different types of pheromones. Dermestids produce esters, and species of Elateridae produce fatty acid-derived aldehydes and acetates. To attract a mate, fireflies (Lampyridae) use modified fat body cells with transparent surfaces backed with reflective uric acid crystals to produce light by bioluminescence. Light production is highly efficient, by oxidation of luciferin catalyzed by enzymes (luciferases) in the presence of adenosine triphosphate (ATP) and oxygen, producing oxyluciferin, carbon dioxide, and light.

Tympanal organs or hearing organs consist of a membrane (tympanum) stretched across a frame backed by an air sac and associated sensory neurons, are found in two families. Several species of the genus Cicindela (Cicindelidae) have hearing organs on the dorsal surfaces of their first abdominal segments beneath the wings; two tribes in the Dynastinae (within the Scarabaeidae) have hearing organs just beneath their pronotal shields or neck membranes. Both families are sensitive to ultrasonic frequencies, with strong evidence indicating they function to detect the presence of bats by their ultrasonic echolocation.

Reproduction and development

Beetles are members of the superorder Endopterygota, and accordingly most of them undergo complete metamorphosis. The typical form of metamorphosis in beetles passes through four main stages: the egg, the larva, the pupa, and the imago or adult. The larvae are commonly called grubs and the pupa sometimes is called the chrysalis. In some species, the pupa may be enclosed in a cocoon constructed by the larva towards the end of its final instar. Going beyond "complete metamorphosis", however, some beetles, such as typical members of the families Meloidae and Rhipiphoridae, undergo hypermetamorphosis in which the first instar takes the form of a triungulin.

Mating

Beetles may display extremely intricate behavior when mating. Pheromone communication is likely to be important in the location of a mate.

Different species use different chemicals for their pheromones. Some scarab beetles (for example, Rutelinae) utilize pheromones derived from fatty acid synthesis, while other scarab beetles use amino acids and terpenoid compounds (for example, Melolonthinae). Another way beetles find mates is by producing light, bioluminescence. This special kind of mating call is confined to fireflies (Lampyridae) which have abdominal light-producing organs. The males and females engage in complex dialogue before mating; different species are separated by differences in flight patterns, duration, composition, and intensity of the light produced.

Before mating, males and females may engage in various forms of behavior. They may stridulate, or vibrate the objects they are on. In some species (for example, Meloidae), the male climbs onto the dorsum of the female and strokes his antennae on her head, palps, and antennae. In the genus Eupompha, the male draws his antennae along his longitudinal vertex. They may not mate at all if they do not perform the precopulatory ritual. This mating behaviour may be different amongst dispersed populations of the same species. For example, the mating of a Russian population of tansy beetle (Chysolina graminis) is preceded by an elaborate ritual involving the male tapping the female's eyes, pronotum and antennae with its antennae, which is not evident in the population of this species in the United Kingdom.

Conflict can play a part in the mating rituals of species such as burying beetles (Nicrophorus), where conflicts between males and females rage until only one of each is left, thus ensuring reproduction by the strongest and fittest. Many male beetles are territorial and fiercely defend their small patches of territory from intruding males. In such species, the male often has horns on the head or thorax, making its body length greater than that of a female. Pairing is generally quick, but in some cases lasts for several hours. During pairing, sperm cells are transferred to the female to fertilize the egg.

Egg

Essentially all beetles lay eggs, though some myrmecophilous Aleocharinae and some Chrysomelinae which live in mountains or the subarctic are ovoviviparous, laying eggs which hatch almost immediately. Beetle eggs generally have smooth surfaces and are soft, though the Cupedidae have hard eggs. Eggs vary widely between species: the eggs tend to be small in species with many instars (larval stages), and in those that lay large numbers of eggs. A female may lay from several dozen to several thousand eggs during her lifetime, depending on the extent of parental care. This ranges from the simple laying of eggs under a leaf, to the parental care provided by scarab beetles, which house, feed and protect their young. The Attelabidae roll leaves and lay their eggs inside the roll for protection.

Larva

The larva is usually the principal feeding stage of the beetle lifecycle. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles, while others feed within their food sources. Examples of internal feeders are most Buprestidae and longhorn beetles. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). The larval period varies between species, but can be as long as several years. The larvae of skin beetles undergo a degree of "reversed development" when starved, and later grow back to the previously attained level of maturity. The cycle can be repeated many times (see Biological immortality). Larval morphology is highly varied amongst species, with well-developed and sclerotized heads, distinguishable thoracic and abdominal segments (usually the tenth, though sometimes the eighth or ninth).

Beetle larvae can be differentiated from other insect larvae by their hardened, often darkened heads, the presence of chewing mouthparts, and spiracles along the sides of their bodies. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. Beetles with somewhat flattened, highly mobile larvae include the ground beetles and rove beetles; their larvae are described as 'campodeiform'. Some beetle larvae resemble hardened WoRMS with dark head capsules and minute legs. These are 'elateriform' larvae, and are found in the click beetle (Elateridae) and darkling beetle (Tenebrionidae) families. Some elateriform larvae of click beetles are known as wireworms. Beetles in the Scarabaeoidea have short, thick larvae described as 'Scarabaeiform', more commonly known as grubs.

All beetle larvae go through several instar stages, which are the developmental stages between each moult. In many species, the larvae simply increase in size with each successive instar as more food is consumed. 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 to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as hypermetamorphosis; it occurs in the Meloidae, Micromalthidae, and Ripiphoridae. The blister beetle Epicauta vittata (Meloidae), for example, has three distinct larval stages. Its first stage, the 'triungulin', has longer legs to go in search of the eggs of grasshoppers. After feeding for a week it moults to the second stage, called the 'caraboid stage', which resembles the larva of a carabid beetle. In another week it moults and assumes the appearance of a scarabaeid larva – the 'scarabaeidoid stage'. Its penultimate larval stage is the pseudo-pupa or the 'coarcate larva', which will overwinter and pupate fully the next Spring.

Pupa

As with all endopterygotes, beetle larvae pupate, and from these pupae emerge fully formed, sexually mature adult beetles, or imagos. Adults have extremely variable lifespans, from weeks to years, depending on the species. In some species, the pupa may go through all four forms during its development, called hypermetamorphosis (for example, Meloidae). Pupae always have no mandibles (are adecticous). In most, the appendages are not attached to the pupae; ones that do have appendages are mostly obtect, and the rest are exarate.

Locomotion

The elytra allow beetles to both fly and move through confined spaces, doing so by folding the delicate wings under the elytra while not flying, and folding their wings out just before take off. The unfolding and folding of the wings is operated by muscles attached to the wing base; as long as the tension on the radial and cubital veins remains, the wings remain straight. In some day-flying species (for example, Buprestidae, Scarabaeidae), flight does not include large amounts of lifting of the elytra, having the metathorac wings extended under the lateral elytra margins. The altitude reached by beetles in flight varies. One study investigating the flight altitude of the ladybird species Coccinella septempunctata and Harmonia axyridis using radar showed that, whilst the majority in flight over a single location were at 150–195 m above ground level, some reached altitudes of over 1100 m.

Aquatic beetles use several techniques for retaining air beneath the water's surface. Diving beetles (Dytiscidae) hold air between the abdomen and the elytra when diving. Hydrophilidae have hairs on their under surface that retain a layer of air against their bodies. Adult crawling water beetles use both their elytra and their hind coxae (the basal segment of the back legs) in air retention, while whirligig beetles simply carry an air bubble down with them whenever they dive.

Communication

Beetles have a variety of ways to communicate, including the use of pheromones. The mountain pine beetle emits a pheromone to attract other beetles to a tree. The mass of beetles are able to overcome the chemical defenses of the tree. After the tree's defenses have been exhausted, the beetles emits an anti-aggregation pheromone. The species can stridulate to communicate.

Parental care

Parental care is found in a few species of beetle, perhaps for protection against adverse conditions and predators. The rove beetle Bledius spectabilis lives in salt marshes, so the eggs and larvae are endangered by the rising tide. The maternal beetle patrols the eggs and larvae, burrowing to keep them from flooding and asphyxiating, and protects them from the predatory carabid beetle Dicheirotrichus gustavi and from the parasitoidal wasp Barycnemis blediator, which kills some 15% of the larvae.

Some dung beetles provide parental care, collecting animal feces, or "dung", and laying eggs within the food supply, an instance of mass provisioning. Some species do not leave after this stage, but remain to safeguard their offspring.

Feeding

Beetles are able to exploit the wide diversity of food sources available in their many habitats. Some are omnivores, eating both plants and animals. Other beetles are highly specialized in their diet. Many species of leaf beetles, longhorn beetles, and weevils are very host-specific, feeding on only a single species of plant. Ground beetles and rove beetles (Staphylinidae), among others, are primarily carnivorous and catch and consume many other arthropods and small prey, such as earthworms and snails. While most predatory beetles are generalists, a few species have more specific prey requirements or preferences.

Decaying organic matter is a primary diet for many species. This can range from dung, which is consumed by coprophagous species (such as certain scarab beetles in the Scarabaeidae), to dead animals, which are eaten by necrophagous species (such as the carrion beetles, Silphidae). Some beetles found in dung and carrion are in fact predatory. These include members of the Histeridae and Silphidae, preying on the larvae of coprophagous and necrophagous insects.

Anti-predator adaptations

Beetles, both adults and larvae, use a variety of anti-predator adaptations. These include camouflage, mimicry, toxicity, and defensive behavior. Camouflage is common and widespread among beetle families, especially those that feed on wood or vegetation, such as leaf beetles (Chrysomelidae) and weevils. In some species, sculpturing or various colored scales or hairs cause the beetle to resemble bird dung or other inedible objects. Many of those that live in sandy environments blend in with the coloration of the substrate. The giant African longhorn beetle (Petrognatha gigas) resembles the moss and bark of the tree it feeds on. Some longhorn beetles (Cerambycidae) are effective Batesian mimics of wasps. Beetles may combine their color mimicry with behavioral mimicry, acting like the wasps they already closely resemble. Many other beetles, including ladybirds, blister beetles, and lycid beetles secrete distasteful or toxic substances to make them unpalatable or poisonous, and are often aposematic, where bright or contrasting color patterns warn off predators; many beetles and other insects mimic these chemically protected species.

Chemical defense is important in some species, usually being advertised by bright colors. Some Tenebrionidae use the posture for releasing noxious chemicals to warn off predators. Chemical defense may serve purposes other than just protection from vertebrates, such as protection from a wide range of microbes. Some species sequester chemicals from the plants they feed on, incorporating them into their own defenses.

Other species have special glands to produce deterrent chemicals. The defensive glands of carabid ground beetles produce a variety of hydrocarbons, aldehydes, phenols, quinones, esters, and acids released from an opening at the end of the abdomen. African carabid beetles (for example, Anthia and Thermophilum – Thermophilum generally included within Anthia) employ the same chemicals as ants: formic acid. Bombardier beetles have well-developed pygidial glands that empty from the lateral edges of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers, one for hydroquinones and hydrogen peroxide, the other holding hydrogen peroxide and catalase enzymes. These chemicals mix and result in an explosive ejection, reaching a temperature of around 100 °C (212 °F), with the breakdown of hydroquinone to hydrogen, oxygen, and quinone. The oxygen propels the noxious chemical spray as a jet that can be aimed accurately at predators.

Large ground-dwelling beetles such as Carabidae, the rhinoceros beetle and the longhorn beetles defend themselves using strong mandibles, or heavily sclerotised (armored) spines or horns to deter or fight off predators. Many species of weevil that feed out in the open on leaves of plants react to attack by employing a "drop-off reflex". Some combine it with thanatosis, in which they close up their appendages and "play dead".

Parasitism

Over 1000 species of beetles are parasitic, predatory, or commensals in the nests of ants.

A few species of beetles are ectoparasitic on mammals. One such species, Platypsyllus castoris, parasitises beavers (Castor spp.). This beetle lives as a parasite both as a larva and as an adult, feeding on epidermal tissue and possibly on skin secretions and wound exudates. They are strikingly flattened dorsoventrally, no doubt as an adaptation for slipping between the beavers' hairs. They are wingless and eyeless, as are many other ectoparasites. Others are kleptoparasites of other invertebrates, such as the small hive beetle (Aethina tumida) that infests honey bee hives.

Pollination

Beetle-pollinated flowers are usually large, greenish or off-white in color, and heavily scented. Scents may be spicy, fruity, or similar to decaying organic material. Beetles were most likely the first insects to pollinate flowers. Most beetle-pollinated flowers are flattened or dish-shaped, with pollen easily accessible, although they may include traps to keep the beetle longer. The plants' ovaries are usually well protected from the biting mouthparts of their pollinators. The beetle families that habitually pollinate flowers are the Buprestidae, Cantharidae, Carambycidae, Cleridae, Dermestidae, Lycidae, Melyridae, Mordellidae, Nitidulidae and Scarabeidae. Beetles may be particularly important in some parts of the world such as semiarid areas of southern Africa and southern California and the montane grasslands of KwaZulu-Natal in South Africa.

Mutualism

Mutualism occurs in a few beetles, such as the ambrosia beetle, which partners with fungi to digest the wood of dead trees. The beetles excavate tunnels in dead trees in which they cultivate fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases spores of its fungal symbiont. The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery, so the weevils and the fungus both benefit. The beetles cannot eat the wood due to toxins, and uses its relationship with fungi to help overcome its host tree defenses and to provide nutrition for their larvae. Chemically mediated by a bacterially produced polyunsaturated peroxide, this mutualistic relationship between the beetle and the fungus is coevolved.

Eusociality

Eusociality involves cooperative brood care (including brood care of offspring from other individuals), overlapping generations within a colony of adults, and a division of labour into reproductive and non-reproductive groups. Few organisms outside Hymenoptera exhibit this behavior; the only beetle to do so is the weevil Austroplatypus incompertus. This Australian species lives in horizontal networks of tunnels, in the heartwood of Eucalyptus trees. It is one of more than 300 species of wood-boring Ambrosia beetles which distribute the spores of ambrosia fungi. The fungi grow in the beetles' tunnels, providing food for the beetles and their larvae; female offspring remain in the tunnels and maintain the fungal growth, probably never reproducing.

Tolerance of extreme temperatures

All insects are poikilothermic, so the ability of a few beetles to live in extreme environments depends on their resilience to unusually high or low temperatures. The bark beetle Pityogenes chalcographus can survive −39°C whilst overwintering beneath tree bark; the Alaskan beetle Cucujus clavipes puniceus is able to withstand −58°C; its larvae may survive −100°C. Conversely, desert dwelling beetles are adapted to tolerate high temperatures. For example, the Tenebrionid beetle Onymacris rugatipennis can withstand 50°C, with the help of an antifreeze protein. The Alaskan beetle Upis ceramboides can survive −60 °C: its cryoprotectants are xylomannan, a molecule consisting of a sugar bound to a fatty acid, and the sugar-alcohol, threitol.

In ancient cultures

Several species of dung beetle, especially the "sacred scarab", Scarabaeus sacer, were revered in Ancient Egypt. The hieroglyphic image of the beetle may have had existential, fictional, or ontologic significance. Images of the scarab in bone, ivory, stone, Egyptian faience, and precious metals are known from the Sixth Dynasty and up to the period of Roman rule. The scarab was of prime significance in the funerary cult of ancient Egypt. The scarab was linked to Khepri, the god of the rising sun, from the supposed resemblance of the rolling of the dung ball by the beetle to the rolling of the sun by the god. Some of ancient Egypt's neighbors adopted the scarab motif for seals of varying types. The best-known of these are the Judean LMLK seals, where eight of 21 designs contained scarab beetles, which were used exclusively to stamp impressions on storage jars during the reign of Hezekiah. Beetles are mentioned as a symbol of the sun, as in ancient Egypt, in Plutarch's 1st century Moralia. The Greek Magical Papyri of the 2nd century BC to the 5th century AD describe scarabs as an ingredient in a spell.

Pliny the Elder discusses beetles in his Natural History, describing the stag beetle: "Some insects, for the preservation of their wings, are covered with a erust (elytra) – the beetle, for instance, the wing of which is peculiarly fine and frail. To these insects a sting has been denied by Nature; but in one large kind we find horns of a remarkable length, two-pronged at the extremities, and forming pincers, which the animal closes when it is its intention to bite." The stag beetle is recorded in a Greek myth by Nicander and recalled by Antoninus Liberalis in which Cerambus is turned into a beetle: "He can be seen on trunks and has hook-teeth, ever moving his jaws together. He is black, long and has hard wings like a great dung beetle". The story concludes with the comment that the beetles were used as toys by young boys, and that the head was removed and worn as a pendant.

As pests

About 75% of beetle species are phytophagous in both the larval and adult stages. Many feed on economically important plants and stored plant products, including trees, cereals, tobacco, and dried fruits. Some, such as the boll weevil, which feeds on cotton buds and flowers, can cause extremely serious damage to agriculture. The boll weevil crossed the Rio Grande near Brownsville, Texas, to enter the United States from Mexico around 1892, and had reached southeastern Alabama by 1915. By the mid-1920s, it had entered all cotton-growing regions in the US, traveling 40 to 160 miles (60–260 km) per year. It remains the most destructive cotton pest in North America. Mississippi State University has estimated, since the boll weevil entered the United States, it has cost cotton producers about $13 billion, and in recent times about $300 million per year.

The bark beetle, elm leaf beetle and the Asian longhorned beetle are among the species that attack elm trees. Bark beetles carry Dutch elm disease as they move from infected breeding sites to healthy trees. The disease has devastated elm trees across Europe and North America.

Some species have become immune to insecticides. For example, the Colorado potato beetle, Leptinotarsa decemlineata, is a destructive pest of potato plants. Its hosts include other members of the Solanaceae, such as nightshade, tomato, eggplant and capsicum, as well as the potato. Different populations have between them developed resistance to all major classes of insecticide. The Colorado potato beetle was evaluated as a tool of entomological warfare during World War II, the idea being to use the beetle and its larvae to damage the crops of enemy nations. Germany tested its Colorado potato beetle weaponisation program south of Frankfurt, releasing 54,000 beetles.

Pests such as the death watch beetle, Xestobium rufovillosum (Anobiidae), is a serious pest of older wooden buildings in Europe. It attacks hardwoods such as oak and chestnut, always where some fungal decay has taken or is taking place. The actual introduction of the pest into buildings is thought to take place at the time of construction.

Other pest include the coconut hispine beetle, Brontispa longissima, which feeds on young leaves, seedlings and mature coconut trees, causing serious economic damage in the Philippines. The mountain pine beetle is a destructive pest of mature or weakened lodgepole pine, sometimes affecting large areas of Canada.

As beneficial resources

Beetles can be beneficial, usually by controlling the populations of pests. The larvae and adults of some species of lady beetles (Coccinellidae) feed on aphids that are pests. Other lady beetles feed on scale insects, whitefly and mealybugs. If normal food sources are scarce, they may feed on small caterpillars, young plant bugs, or honeydew and nectar. Ground beetles (Carabidae) are common predators of many insect pests, including fly eggs, caterpillars, and wireworms.

Dung beetles (Scarabidae) have been successfully used to reduce the populations of pestilent flies, such as Musca vetustissima and Haematobia exigua which are serious pests of cattle in Australia. The beetles make the dung unavailable to breeding pests by quickly rolling and burying it in the soil, with the added effect of improving soil fertility, tilth, and nutrient cycling. The Australian Dung Beetle Project (1965–1985), introduced species of dung beetle to Australia from South Africa and Europe to reduce populations of Musca vetustissima, following successful trials of this technique in Hawaii. The American Institute of Biological Sciences reports that dung beetles save the United States cattle industry an estimated US$380 million annually through burying above-ground livestock feces.

The Dermestidae are often used in taxidermy and in the preparation of scientific specimens, to clean soft tissue from bones. Multiple larvae may be used to assist in removing cartilage along with other soft tissue.

As food

Beetles are the most widely eaten insects, with about 344 species used as food, usually at the larval stage. The mealworm (the larva of the darkling beetle) and the rhinoceros beetle are among the species commonly eaten.

In art and adornment

Many beetles have beautiful and durable elytra that have been used as material in arts, with beetlewing the best example. Sometimes, they are incorporated into ritual objects for their religious significance. Whole beetles, either as-is or encased in clear plastic, are made into objects from cheap souvenirs such as key chains to expensive fine-art jewelry. In parts of Mexico, beetles of the genus Zopherus are made into living brooches by attaching costume jewelry and golden chains, which is made possible by the incredibly hard elytra and sedentary habits of the genus.

In entertainment

Fighting beetles are used for entertainment and gambling. This sport exploits the territorial behavior and mating Competition of certain species of large beetles. In the Chiang Mai district of northern Thailand, male Xylotrupes rhinoceros beetles are caught in the wild and trained for fighting. Females are held inside a log to stimulate the fighting males with their pheromones. These fights may be competitive and involve gambling both money and property. In South Korea the Dytiscidae species Cybister tripunctatus is used in a roulette-like game.

Beetles are sometimes used as instruments: the Onabasulu of Papua New Guinea historically used the weevil Rhynchophorus ferrugineus as a musical instrument by letting the human mouth serve as a variable resonance chamber for the wing vibrations of the live adult beetle.

As pets

Some species of beetle are kept as pets, for example diving beetles (Dytiscidae) may be kept in a domestic fresh water tank.

In Japan the practice of keeping horned rhinoceros beetles (Dynastinae) and stag beetles (Lucanidae) is particularly popular amongst young boys. Such is the popularity in Japan that vending machines dispensing live beetles were developed in 1999, each holding up to 100 stag beetles.

As things to collect

Beetle collecting became extremely popular in the Victorian era. The naturalist Alfred Russel Wallace collected (by his own count) a total of 83,200 beetles during the eight years described in his 1869 book The Malay Archipelago, including 2,000 species new to science.