The Cisuralian is the first series / epoch of the Permian . The Cisuralian was preceded by the Pennsylvanian and followed by the Guadalupian . The Cisuralian Epoch is named after the western slopes of the Ural Mountains in Russia and Kazakhstan and dates between 298.9 ± 0.15 – 272.3 ± 0.5 Ma .
80-570: In the regional stratigraphy of southwestern North America, the Cisuralian encompasses two series: the Wolfcampian (Asselian to mid-Artinskian) and Leonardian (mid-Artinskian to Kungurian). The series saw the appearance of beetles and flies and was a relatively stable warming period of about 21 million years. The Cisuralian is the first series or epoch of the Permian. The Cisuralian
160-421: 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 (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). This immense diversity led
240-448: A beetle is quite uniform, although specific organs and appendages vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen. Because there are so many species, identification is quite difficult, and relies on attributes including the shape of the antennae, the tarsal formulae and shapes of these small segments on
320-481: A beetle's environment. Beetle families may use antennae in different ways. For example, when moving quickly, tiger beetles may not be able to see very well and instead hold their antennae rigidly in front of them in order to avoid obstacles. Certain Cerambycidae use antennae to balance, and blister beetles may use them for grasping. Some aquatic beetle species may use antennae for gathering air and passing it under
400-709: 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
480-456: A length of up to 7 cm (2.8 in), is generally considered to be the largest fly in the world, while the smallest is Euryplatea nanaknihali , which at 0.4 mm (0.016 in) is smaller than a grain of salt. Brachycera are ecologically very diverse, with many being predatory at the larval stage and some being parasitic. Animals parasitised include molluscs , woodlice , millipedes , insects, mammals , and amphibians . Flies are
560-441: 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
640-406: 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
720-405: A meal. For visual course control, flies' optic flow field is analyzed by a set of motion-sensitive neurons. A subset of these neurons is thought to be involved in using the optic flow to estimate the parameters of self-motion, such as yaw, roll, and sideward translation. Other neurons are thought to be involved in analyzing the content of the visual scene itself, such as separating figures from
800-401: A mobile head, with a pair of large compound eyes , and mouthparts designed for piercing and sucking (mosquitoes, black flies and robber flies), or for lapping and sucking in the other groups. Their wing arrangement gives them great manoeuvrability in flight, and claws and pads on their feet enable them to cling to smooth surfaces. Flies undergo complete metamorphosis ; the eggs are often laid on
880-461: A mobile tooth on their left mandible. 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 thought to be
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#1732772915618960-490: A particularly hard exoskeleton and hard forewings ( elytra ) not usable for flying. Almost all beetles have mandibles that move in a horizontal plane. The mouthparts are rarely suctorial, though they are sometimes reduced; the maxillae always bear palps. The antennae usually have 11 or fewer segments, except in some groups like the Cerambycidae (longhorn beetles) and the Rhipiceridae (cicada parasite beetles). The coxae of
1040-708: A sister group to the beetles, having split from them in the Early Permian . Molecular phylogenetic analysis confirms that the Coleoptera are monophyletic. 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
1120-408: A string of beads , comb-like (either on one side or both, bipectinate), or toothed . The physical variation of antennae is important for the identification of many beetle groups. The Curculionidae have elbowed or geniculate antennae. Feather like flabellate antennae are a restricted form found in the Rhipiceridae and a few other families. The Silphidae have a capitate antennae with a spherical head at
1200-438: A variety of hosts. The larvae of the shore flies (Ephydridae) and some Chironomidae survive in extreme environments including glaciers ( Diamesa sp., Chironomidae ), hot springs, geysers, saline pools, sulphur pools, septic tanks and even crude oil ( Helaeomyia petrolei ). Adult hoverflies (Syrphidae) are well known for their mimicry and the larvae adopt diverse lifestyles including being inquiline scavengers inside
1280-485: Is Spilomyia longicornis , which is a fly but mimics a vespid wasp. Flies have a mobile head with a pair of large compound eyes on the sides of the head, and in most species, three small ocelli on the top. The compound eyes may be close together or widely separated, and in some instances are divided into a dorsal region and a ventral region, perhaps to assist in swarming behaviour. The antennae are well-developed but variable, being thread-like, feathery or comb-like in
1360-489: 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 name of the taxonomic order, Coleoptera, comes from
1440-438: 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 . The heaviest beetle, indeed the heaviest insect stage, is the larva of the goliath beetle , Goliathus goliatus , which can attain
1520-547: Is one of the major insect orders and of considerable ecological and human importance. Flies are major pollinators, second only to the bees and their Hymenopteran relatives. Flies may have been among the evolutionarily earliest pollinators responsible for early plant pollination . Fruit flies are used as model organisms in research, but less benignly, mosquitoes are vectors for malaria , dengue , West Nile fever , yellow fever , encephalitis , and other infectious diseases ; and houseflies , commensal with humans all over
1600-548: 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 325 μm in length. The oldest known beetle is Coleopsis , from the earliest Permian ( Asselian ) of Germany, around 295 million years ago. Early beetles from
1680-404: Is the largest of all orders, constituting almost 40% of described insects 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. However, the number of beetle species is challenged by the number of species in dipterans (flies) and hymenopterans (wasps). Found in almost every habitat except
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#17327729156181760-437: 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 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
1840-524: Is widely thought to be a member of Mecopterida , along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly Strepsiptera (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies. Diptera were traditionally broken down into two suborders, Nematocera and Brachycera , distinguished by
1920-1210: The Dytiscidae (diving beetles) , Haliplidae , and many species of Hydrophilidae , the legs, often the last pair, are modified for swimming, typically with rows of long hairs. Male diving beetles have suctorial cups on their forelegs that they use to grasp females. Other beetles have fossorial legs widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles ( Histeridae ). The hind legs of some beetles, such as flea beetles (within Chrysomelidae) and flea weevils (within Curculionidae), have enlarged femurs that help them leap. Fly Housefly (Muscidae) (top left) Haematopota pluvialis (Tabanidae) (top right) Ctenophora pectinicornis (Tipulidae) (mid left) Ochlerotatus notoscriptus (Culicidae) (mid right) Milesia crabroniformis (Syrphidae) (bottom left) Holcocephala fusca (Asilidae) (bottom right) Nematocera ( paraphyletic ) (inc Eudiptera ) Brachycera Flies are insects of
2000-531: The Greek koleopteros (κολεόπτερος), 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 ċeafor , chafer, used in names such as cockchafer , from
2080-631: The Jurassic , some 180 million years ago. A third radiation took place among the Schizophora at the start of the Paleogene , 66 million years ago. The phylogenetic position of Diptera has been controversial. The monophyly of holometabolous insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera
2160-704: The Late Carboniferous , and reached their apex in the Cisuralian remaining the dominant land animals for some 40 million years. A few continued into the Capitanian . They were succeeded by the therapsids . Beetle See subgroups of the order Coleoptera Beetles are insects that form the order Coleoptera ( / k oʊ l iː ˈ ɒ p t ər ə / ), 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,
2240-1071: The Mecopterida , alongside the Mecoptera , Siphonaptera , Lepidoptera and Trichoptera . The possession of a single pair of wings distinguishes most true flies from other insects with "fly" in their names. However, some true flies such as Hippoboscidae (louse flies) have become secondarily wingless. The cladogram represents the current consensus view. Hymenoptera (sawflies, wasps, ants, bees) [REDACTED] Raphidioptera (snakeflies) [REDACTED] Megaloptera (alderflies and allies) [REDACTED] Neuroptera (Lacewings and allies) [REDACTED] Coleoptera (beetles) [REDACTED] Strepsiptera (twisted-wing parasites) [REDACTED] Trichoptera (caddisflies) [REDACTED] Lepidoptera (butterflies, moths) [REDACTED] Diptera [REDACTED] Mecoptera (scorpionflies) [REDACTED] Siphonaptera (fleas) [REDACTED] The first true dipterans known are from
2320-666: The Middle Triassic (around 240 million years ago), and they became widespread during the Middle and Late Triassic . Modern flowering plants did not appear until the Cretaceous (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than nectar . Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on honeydew produced by sap-sucking bugs which were abundant at
2400-545: The Middle Triassic . During the Jurassic ( 210 to 145 mya ), 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
2480-519: The order Diptera , the name being derived from the Greek δι- di- "two", and πτερόν pteron "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced mechanosensory organs known as halteres , which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing more than 150,000 species including horse-flies , crane flies , hoverflies , mosquitoes and others. Flies have
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2560-431: 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
2640-415: The telephone-pole beetle . The Archostemata have an exposed plate called the metatrochantin in front of the basal segment or coxa of the hind leg. Myxophaga contains about 65 described species in four families, mostly very small, including Hydroscaphidae and the genus Sphaerius . The myxophagan beetles are small and mostly alga-feeders. Their mouthparts are characteristic in lacking galeae and having
2720-548: The thorax , bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures, and the third segment bears the halteres , which help to balance the insect during flight. The third tagma is the abdomen consisting of 11 segments, some of which may be fused, and with the 3 hindmost segments modified for reproduction. Some Dipterans are mimics and can only be distinguished from their models by very careful inspection. An example of this
2800-827: The Brachycera includes broader, more robust flies with short antennae. Many nematoceran larvae are aquatic. There are estimated to be a total of about 19,000 species of Diptera in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region and 19,000 in the Australasian region. While most species have restricted distributions, a few like the housefly ( Musca domestica ) are cosmopolitan. Gauromydas heros ( Asiloidea ), with
2880-707: 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 ,
2960-1403: The Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project. Ptychopteromorpha (phantom and primitive crane-flies) [REDACTED] Culicomorpha (mosquitoes, blackflies and midges) [REDACTED] Blephariceromorpha (net-winged midges, etc) [REDACTED] Bibionomorpha (gnats) [REDACTED] Psychodomorpha (drain flies, sand flies, etc) [REDACTED] Tipulomorpha (crane flies) [REDACTED] Stratiomyomorpha (soldier flies, etc) [REDACTED] Xylophagomorpha (stink flies, etc) [REDACTED] Tabanomorpha (horse flies, snipe flies, etc) [REDACTED] Nemestrinoidea [REDACTED] Asiloidea (robber flies, bee flies, etc) [REDACTED] Empidoidea (dance flies, etc) [REDACTED] Aschiza (in part) Phoroidea (flat-footed flies, etc) [REDACTED] Syrphoidea (hoverflies) [REDACTED] Hippoboscoidea (louse flies, etc) [REDACTED] Muscoidea (house flies, dung flies, etc) [REDACTED] Oestroidea (blow flies, flesh flies, etc) [REDACTED] Acalyptratae (marsh flies, etc) [REDACTED] Flies are often abundant and are found in almost all terrestrial habitats in
3040-572: The Permian, which are collectively grouped into the " Protocoleoptera " are thought to have been xylophagous (wood eating) 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
3120-483: The Proto-Germanic * kebrô ("beetle"; compare German Käfer , Dutch kever , Afrikaans kewer ). 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. A 2015 study provided four independent estimates of the total number of beetle species, giving a mean estimate of some 1.5 million with
3200-662: 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. 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
3280-535: The Soviet Union during WW2 when the Germans controlled the oil fields to the west. The International Chronostratigraphic Chart (v2018/07) provides a numerical age of 298.9 ± 0.15 – 272.3 ± 0.5 Ma. The base of the Cisuralian series and the Permian system is defined as the place in the stratigraphic record where fossils of the conodont Streptognathodus isolatus first appear. The global reference profile for
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3360-2009: 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 common names are given where possible. Dates of origin of major groups are shown in italics in millions of years ago (mya). Archostemata 160 mya (40) [REDACTED] Myxophaga 220 mya (94) [REDACTED] Hydradephaga (5,560) e.g. Dytiscidae (diving beetles) [REDACTED] Geadephaga ( 35,000 ) e.g. Carabidae (ground beetles) [REDACTED] Scirtoidea (800) + Derodontoidea (29) 200 mya [REDACTED] Staphylinidae 195 mya ( 48,000 , rove beetles) [REDACTED] Scarabaeoidea 145 mya ( 35,000 , scarabs, stag beetles, etc.) [REDACTED] Hydrophiloidea (2,800, water scavenger beetles) [REDACTED] Histeroidea (3,800, clown beetles) [REDACTED] Nosodendridae (70) Dascilloidea (180) [REDACTED] Buprestoidea ( 14,000 , jewel beetles) [REDACTED] Byrrhoidea (400, pill and turtle beetles, etc.) [REDACTED] Elateroidea ( 23,000 , click and soldier beetles, fireflies) [REDACTED] Bostrichoidea (3150, deathwatch, powderpost and skin beetles) [REDACTED] Coccinelloidea (6,000, ladybirds or lady beetles) [REDACTED] Tenebrionoidea 180 mya ( 35,000 , leaf/flower beetles, etc.) and Lymexyloidea [REDACTED] Cleroidea (9,900, checkered beetles and allies) [REDACTED] Cucujoidea (8,000) [REDACTED] Chrysomelidae ( 35,000 , leaf beetles) [REDACTED] Cerambycidae ( 25,000 , longhorn beetles) [REDACTED] Curculionoidea ( 97,000 , weevils) [REDACTED] Beetles are generally characterized by
3440-763: The Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia. 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
3520-797: 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. The Cretaceous saw the fragmenting of the southern landmass, with the opening of
3600-519: The antennae and the maxillary palps bear the main olfactory receptors, while the gustatory receptors are in the labium, pharynx, feet, wing margins and female genitalia, enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing. Flies that feed on blood have special sensory structures that can detect infrared emissions, and use them to home in on their hosts, Many blood-sucking flies can detect
3680-701: The appearance of beetles and flies. The coal swamps from the Carboniferous declined but the herbivores, Diadectes and Edaphosaurus persisted until the end of this series, approximately. The dry interior had small insectivores. Caseids and prototherapsid Tetraceratops made their appearance. The marine life was probably more diverse than modern times as the climate warmed. Unusual sharks such as Helicoprion continued in this series. Early Permian terrestrial faunas were dominated by pelycosaurs (a paraphyletic group of early synapsids ), diadectids , and temnospondyls , The pelycosaurs appeared during
3760-456: The aquatic whirligig beetles ( Gyrinidae ), where they are split to allow a view both above and below the waterline. A few Longhorn beetles ( Cerambycidae ) and weevils as well as some fireflies ( Rhagophthalmidae ) have divided eyes, while many have eyes that are notched, and a few have ocelli , small, simple eyes usually farther back on the head (on the vertex ); these are more common in larvae than in adults. The anatomical organization of
3840-447: The back part of the thorax is concealed by the beetle's wings . This further segmentation is usually best seen on the abdomen. The multisegmented legs end in two to five small segments called tarsi. Like many other insect orders, beetles have claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs have been variously adapted for other uses. Aquatic beetles including
3920-631: 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 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 (Gyrinidae) were moderately diverse, although other early beetles (e.g. Dytiscidae) were less, with
4000-717: The base (the GSSP or golden spike ) is located in the valley of the Aidaralash River, near Aqtöbe in the Ural Mountains of Kazakhstan. Gondwana collided with Laurussia and created the Alleghenian orogeny in present-day North America. In northwestern Europe, the Hercynian orogeny continued. This created the large supercontinent, Pangea , by the middle of the early Permian, which was to have an impact on
4080-433: The body whilst submerged. Equally, some families use antennae during mating, and a few species use them for defense. In the cerambycid Onychocerus albitarsis , the antennae have venom injecting structures used in defense, which is unique among arthropods . Antennae vary greatly in form, sometimes between the sexes, but are often similar within any given family. Antennae may be clubbed , threadlike , angled , shaped like
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#17327729156184160-703: The climate. At the start of the Permian, the Late Palaeozoic Ice Age , which began in the Carboniferous , was at its peak. Glaciers receded over the course of the late Cisuralian as the Earth's climate gradually warmed, particularly during the Artinskian Warming Event, drying the continent's interiors. The pan-tropical belt of Pangaea experienced particularly significant aridification during this epoch. The swampy fringes were mostly ferns, seed ferns, and lycophytes . The series saw
4240-555: The compound eyes may be modified and depends on whether a species is primarily crepuscular, or diurnally or nocturnally active. Ocelli are found in the adult carpet beetle (as a single central ocellus in Dermestidae ), some rove beetles ( Omaliinae ), and the Derodontidae . Beetle antennae are primarily organs of sensory perception and can detect motion, odor and chemical substances, but may also be used to physically feel
4320-404: The differences in antennae. The Nematocera are identified by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae. Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in
4400-431: The different families. The mouthparts are adapted for piercing and sucking, as in the black flies, mosquitoes and robber flies, and for lapping and sucking as in many other groups. Female horse-flies use knife-like mandibles and maxillae to make a cross-shaped incision in the host's skin and then lap up the blood that flows. The gut includes large diverticulae , allowing the insect to store small quantities of liquid after
4480-651: 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
4560-715: 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
4640-590: 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 (flies) and/or Hymenoptera (sawflies, wasps, ants and bees) may have more species. Beetles are found in nearly all habitats, including freshwater and coastal habitats, wherever vegetative foliage
4720-440: The first part is called the scape and the second part is the pedicel. The other segments are jointly called the flagellum. Beetles have mouthparts like those of grasshoppers . The mandibles 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 defence , below). Two pairs of finger-like appendages,
4800-749: The flies, principally in the family Cecidomyiidae (gall midges). Many flies (most importantly in the family Agromyzidae) lay their eggs in the mesophyll tissue of leaves with larvae feeding between the surfaces forming blisters and mines. Some families are mycophagous or fungus feeding. These include the cave dwelling Mycetophilidae (fungus gnats) whose larvae are the only diptera with bioluminescence. The Sciaridae are also fungus feeders. Some plants are pollinated by fungus feeding flies that visit fungus infected male flowers. The larvae of Megaselia scalaris (Phoridae) are almost omnivorous and consume such substances as paint and shoe polish. The Exorista mella (Walker) fly are considered generalists and parasitoids of
4880-527: The ground using motion parallax. The H1 neuron is responsible for detecting horizontal motion across the entire visual field of the fly, allowing the fly to generate and guide stabilizing motor corrections midflight with respect to yaw. The ocelli are concerned in the detection of changes in light intensity, enabling the fly to react swiftly to the approach of an object. Like other insects, flies have chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of
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#17327729156184960-431: The larval food-source and the larvae, which lack true limbs, develop in a protected environment, often inside their food source. Other species are ovoviviparous , opportunistically depositing hatched or hatching larvae instead of eggs on carrion , dung, decaying material, or open wounds of mammals. The pupa is a tough capsule from which the adult emerges when ready to do so; flies mostly have short lives as adults. Diptera
5040-431: The legs are usually located recessed within a coxal cavity. The genitalic structures are telescoped into the last abdominal segment in all extant beetles. Beetle larvae can often be confused with those of other holometabolan groups. The beetle's exoskeleton is made up of numerous plates, called sclerites , separated by thin sutures. This design provides armored defenses while maintaining flexibility. The general anatomy of
5120-407: The legs, the mouthparts, and the ventral plates (sterna, pleura, coxae). In many species accurate identification can only be made by examination of the unique male genitalic structures. The head, having mouthparts projecting forward or sometimes downturned, is usually heavily sclerotized and is sometimes very large. The eyes are compound and may display remarkable adaptability, as in the case of
5200-489: The maxillary and labial palpi, are found around the mouth in most beetles, serving to move food into the mouth. In many species, the mandibles are sexually dimorphic, with those of the males enlarged enormously compared with those of females of the same species. The thorax is segmented into the two discernible parts, the pro- and pterothorax. The pterothorax is the fused meso- and metathorax, which are commonly separated in other insect species, although flexibly articulate from
5280-436: 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
5360-419: The nests of social insects. Some brachycerans are agricultural pests, some bite animals and humans and suck their blood, and some transmit diseases. Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, bears the eyes, the antennae , and the mouthparts (the labrum, labium, mandible, and maxilla make up the mouthparts). The second tagma,
5440-406: The neural ganglia , and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder. Some flies such as the ectoparasitic Nycteribiidae and Streblidae are exceptional in having lost their wings and become flightless. The only other order of insects bearing a single pair of true, functional wings, in addition to any form of halteres, are
5520-490: 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
5600-410: The prothorax. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. The abdomen is everything posterior to the thorax. When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle section is a hard plate called the pronotum , which is only the front part of the thorax;
5680-405: The raised concentration of carbon dioxide that occurs near large animals. Some tachinid flies (Ormiinae) which are parasitoids of bush crickets , have sound receptors to help them locate their singing hosts. Diptera have one pair of fore wings on the mesothorax and a pair of halteres , or reduced hind wings, on the metathorax . A further adaptation for flight is the reduction in number of
5760-510: 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. Some others also have unusual characteristics, such as fireflies , which use
5840-568: The second largest group of pollinators after the Hymenoptera (bees, wasps and relatives). In wet and colder environments flies are significantly more important as pollinators. Compared to bees, they need less food as they do not need to provision their young. Many flowers that bear low nectar and those that have evolved trap pollination depend on flies. It is thought that some of the earliest pollinators of plants may have been flies. The greatest diversity of gall forming insects are found among
5920-664: 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
6000-702: 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
6080-606: The time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues. The basal clades in the Diptera include the Deuterophlebiidae and the enigmatic Nymphomyiidae . Three episodes of evolutionary radiation are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the Triassic , about 220 million years ago. Many lower Brachycera appeared in
6160-485: The tip. The Scarabaeidae typically have lamellate antennae with the terminal segments extended into long flat structures stacked together. The Carabidae typically have thread-like antennae. The antennae arises between the eye and the mandibles and in the Tenebrionidae, the antennae rise in front of a notch that breaks the usually circular outline of the compound eye. They are segmented and usually consist of 11 parts,
6240-473: The world apart from Antarctica. They include many familiar insects such as house flies, blow flies, mosquitoes, gnats, black flies, midges and fruit flies. More than 150,000 have been formally described and the actual species diversity is much greater, with the flies from many parts of the world yet to be studied intensively. The suborder Nematocera include generally small, slender insects with long antennae such as mosquitoes, gnats, midges and crane-flies, while
6320-607: The world, spread foodborne illnesses . Flies can be annoyances especially in some parts of the world where they can occur in large numbers, buzzing and settling on the skin or eyes to bite or seek fluids. Larger flies such as tsetse flies and screwworms cause significant economic harm to cattle. Blowfly larvae, known as gentles , and other dipteran larvae, known more generally as maggots , are used as fishing bait , as food for carnivorous animals, and in medicine in debridement , to clean wounds . Dipterans are holometabolans , insects that undergo radical metamorphosis. They belong to
6400-671: Was preceded by the last Pennsylvanian epoch ( Gzhelian ) and is followed by the Permian Guadalupian Epoch. The name "Cisuralian" was proposed in 1982, and approved by the International Subcommission on Permian Stratigraphy in 1996. The Cisuralian Epoch is named after the western slopes of the Ural Mountains in Russia and Kazakhstan. Limestones on the edge of Russian Platform and make up the Ishimbay oil fields. These oil fields were vital to
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