Beetle
Beetles are insects that form the order Coleoptera, in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 described species, is the largest of all orders, constituting almost 40% of described arthropods and 25% of all known animal species; new species are discovered frequently, with estimates suggesting that there are between 0.9 and 2.1 million total species.
Found in almost every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are serious agricultural pests, such as the Colorado potato beetle, while others such as Coccinellidae eat aphids, scale insects, thrips, and other plant-sucking insects that damage crops. Some others also have unusual characteristics, such as fireflies, which use a light-emitting organ for mating and communication purposes.
Beetles typically have a particularly hard exoskeleton including the elytra, though some such as the rove beetles have very short elytra while blister beetles have softer elytra. The general anatomy of a beetle is quite uniform and typical of insects, although there are several examples of novelty, such as adaptations in water beetles which trap air bubbles under the elytra for use while diving. Beetles are holometabolans, which means that they undergo complete metamorphosis, with a series of conspicuous and relatively abrupt changes in body structure between hatching and becoming adult after a relatively immobile pupal stage. Some, such as stag beetles, have a marked sexual dimorphism, the males possessing enormously enlarged mandibles which they use to fight other males. Many beetles are aposematic, with bright colors and patterns warning of their toxicity, while others are harmless Batesian mimics of such insects. Many beetles, including those that live in sandy places, have effective camouflage.
Beetles are prominent in human culture, from the sacred scarabs of ancient Egypt to beetlewing art and use as pets or fighting insects for entertainment and gambling. Many beetle groups are brightly and attractively colored making them objects of collection and decorative displays. Over 300 species are used as food, mostly as larvae; species widely consumed include mealworms and rhinoceros beetle larvae. However, the major impact of beetles on human life is as agricultural, forestry, and horticultural pests. Serious pest species include the boll weevil of cotton, the Colorado potato beetle, the coconut hispine beetle, the mountain pine beetle, and many others. Most beetles, however, do not cause economic damage and some, such as numerous species of lady beetles, are beneficial by helping to control insect pests. The scientific study of beetles is known as coleopterology.
Etymology
The name of the taxonomic order, 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, leading to Middle English betylle. Another Old English name for beetle is ċeafor, chafer, used in names such as cockchafer, from the Proto-Germanic *kebrô.Distribution and diversity
Beetles are by far the largest order of insects: the roughly 400,000 species make up about 40% of all arthropod species so far described, and about 25% of all animal species. A 2015 study 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 from a minimum of 0.9 to a maximum of 2.1 million beetle species. The four estimates made use of host-specificity relationships, ratios with other taxa, plant:beetle ratios, and extrapolations based on body size by year of description.This immense diversity led the evolutionary biologist J. B. S. Haldane to quip, when some theologians asked him what could be inferred about the mind of the Christian God from the works of His Creation, "An inordinate fondness for beetles".
However, the ranking of beetles as most diverse has been challenged. Multiple studies posit that Diptera and/or Hymenoptera may have more species.
Beetles are found in nearly all habitats, including freshwater and coastal habitats, wherever 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. Tropical forest canopies have a large and diverse fauna of beetles, including Carabidae, Chrysomelidae, and Scarabaeidae.
Among the heaviest beetles, whose larval stage can attain a mass of at least and a length of, is the goliath beetle, Goliathus goliatus; in its adult stage it can weigh and reach up to in length. Adult elephant beetles, Megasoma elephas and Megasoma actaeon often reach and.
The longest beetle is the Hercules beetle Dynastes hercules, with a maximum overall length of at least 16.7 cm including the very long pronotal horn. The smallest recorded beetle and the smallest free-living insect, is the featherwing beetle Scydosella musawasensis which may measure as little as 325 μm in length.
Evolution
Late Paleozoic and Triassic
The oldest known beetle is Coleopsis, from the earliest Permian of Germany, around 295 million years ago. Early beetles from the Permian, which are collectively grouped into the "Protocoleoptera" are thought to have been xylophagous and wood boring. Fossils from this time have been found in Siberia 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. The earliest members of modern beetle lineages appeared during the Late Permian. In the Permian–Triassic extinction event at the end of the Permian, most "protocoleopteran" lineages became extinct. Beetle diversity did not recover to pre-extinction levels until the Middle Triassic.Jurassic
During the Jurassic, 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 and click beetles appeared. The first jewel beetles are present, but they remained rare until the Cretaceous.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 and rove beetles began to distribute into different patterns; the Carabidae predominantly occurred in the warm regions, while the Staphylinidae and click beetles preferred temperate climates. Likewise, predatory species of Cleroidea and Cucujoidea hunted their prey under the bark of trees together with the jewel beetles. The diversity of jewel beetles increased rapidly, as they were the primary consumers of wood, while longhorn beetles were rather rare: their diversity increased only towards the end of the Upper Cretaceous. The first coprophagous beetles are from the Upper Cretaceous and 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 were moderately diverse, although other early beetles were less, with the most widespread being the species of Coptoclavidae, which preyed on aquatic fly larvae.A 2020 review of the palaeoecological interpretations of fossil beetles from Cretaceous ambers has suggested that saproxylicity was the most common feeding strategy, with fungivorous species in particular appearing to dominate.
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.