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The car became widely known in its home country as the Kfer (German for "beetle", cognate with English chafer) and was later marketed under that name in Germany,[18] and as the Volkswagen in other countries. For example, in France it was known as the Coccinelle (French for ladybug).[19][20]

Beetles are insects that form the order Coleoptera (/koliptr/), 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 insects and 25% of all known animal species;[2] 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 (ladybirds or ladybugs) eat aphids, scale insects, thrips, and other plant-sucking insects that damage crops.

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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 pests include the boll weevil of cotton, the Colorado potato beetle, the coconut hispine beetle, and the mountain pine beetle. Most beetles, however, do not cause economic damage and many, such as the lady beetles and dung beetles are beneficial by helping to control insect pests.

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 animal species.[1][7][8][9][10][11] 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"[12] 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 (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 extrapolations based on body size by year of description (1.7 to 2.1 million).[12][13]

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".[14]

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.[18] Tropical forest canopies have a large and diverse fauna of beetles,[19] including Carabidae,[20] Chrysomelidae,[21] and Scarabaeidae.[22]

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[update]), is the featherwing beetle Scydosella musawasensis which may measure as little as 325 m in length.[25]

During the Jurassic (210 to 145 mya), there was a dramatic increase in the diversity of beetle families,[29] 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.[32] 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.[33] 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.[34][35][36] 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.

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.[32] 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,[41] while longhorn beetles (Cerambycidae) were rather rare: their diversity increased only towards the end of the Upper Cretaceous.[29] The first coprophagous beetles are from the Upper Cretaceous[42] and may have lived on the excrement of herbivorous dinosaurs.[43] 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.[29]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.[44]

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.[39] Lower Cretaceous sites include the Crato fossil beds in the Araripe basin in the Cear, 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.[29]

Beetle fossils are abundant in the Cenozoic; by the Quaternary (up to 1.6 mya), fossil species are identical to living ones, while from the Late Miocene (5.7 mya) the fossils are still so close to modern forms that they are most likely the ancestors of living species. The large oscillations in climate during the Quaternary caused beetles to change their geographic distributions so much that current location gives little clue to the biogeographical history of a species. It is evident that geographic isolation of populations must often have been broken as insects moved under the influence of changing climate, causing mixing of gene pools, rapid evolution, and extinctions, especially in middle latitudes.[46]

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. 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).[10][47] These polyphagan beetle groups 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.[48]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).[49]Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and the telephone-pole beetle.[50]The Archostemata have an exposed plate called the metatrochantin in front of the basal segment or coxa of the hind leg.[51] Myxophaga contains about 65 described species in four families, mostly very small, including Hydroscaphidae and the genus Sphaerius.[52] The myxophagan beetles are small and mostly alga-feeders. Their mouthparts are characteristic in lacking galeae and having a mobile tooth on their left mandible.[53] 006ab0faaa