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Cicada
Temporal range: Late Triassic – recent
Linne's cicada, Neotibicen linnei
Calling song of Magicicada cassini
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Suborder: Auchenorrhyncha
Infraorder: Cicadomorpha
Superfamily: Cicadoidea
Latreille, 1802
Families

The cicadas (/sɪˈkɑːdəz, -ˈk-/) are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha,[a] along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two families, the Tettigarctidae, with two species in Australia, and the Cicadidae, with more than 3,000 species described from around the world; many species remain undescribed. Nearly all cicada species are annual cicadas with the exception of the few North American periodical cicada species, genus Magicicada, which in a given region emerge en masse every 13 or 17 years.

Cicadas have prominent eyes set wide apart, short antennae, and membranous front wings. They have an exceptionally loud song, produced in most species by the rapid buckling and unbuckling of drum-like tymbals. The earliest known fossil Cicadomorpha appeared in the Upper Permian period; extant species occur all around the world in temperate to tropical climates. They typically live in trees, feeding on watery sap from xylem tissue, and laying their eggs in a slit in the bark. Most cicadas are cryptic. The vast majority of species are active during the day as adults, with some calling at dawn or dusk. Only a rare few species are known to be nocturnal.

One exclusively North American genus, Magicicada (the periodical cicadas), which spend most of their lives as underground nymphs, emerge in predictable intervals of 13 or 17 years, depending on the species and the location. The unusual duration and synchronization of their emergence may reduce the number of cicadas lost to predation, both by making them a less reliably available prey (so that any predator that evolved to depend on cicadas for sustenance might starve waiting for their emergence), and by emerging in such huge numbers that they will satiate any remaining predators before losing enough of their number to threaten their survival as a species.[1]

The annual cicadas are species that emerge every year. Though these cicadas' life cycles can vary from 1 to 9 or more years as underground nymphs, their emergence above ground as adults is not synchronized, so some members of each species appear every year.[2]

Cicadas have been featured in literature since the time of Homer's Iliad and as motifs in art from the Chinese Shang dynasty.[3] They have also been used in myth and folklore as symbols of carefree living and immortality. The cicada is also mentioned in Hesiod's Shield (ll.393–394), in which it is said to sing when millet first ripens. Cicadas are eaten by humans in various parts of the world, including China, Myanmar, Malaysia, central Africa and parts of Mexico.[4]

Etymology

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The name is directly from the onomatopoeic Latin cicada.[5][6][b]

Taxonomy and diversity

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A 17-year cicada, Magicicada, Robert Evans Snodgrass, 1930[7]

The superfamily Cicadoidea is a sister of the Cercopoidea (the froghoppers). Cicadas are arranged into two families: the Tettigarctidae and Cicadidae. The two extant species of the Tettigarctidae include one in southern Australia and the other in Tasmania. The family Cicadidae is subdivided into the subfamilies Cicadettinae, Cicadinae, Derotettiginae, Tibicininae (or Tettigadinae), and Tettigomyiinae,[8] and they are found on all continents except Antarctica. Some previous works also included a family-level taxon called the Tibiceninae. The largest species is the Malaysian emperor cicada Megapomponia imperatoria; its wingspan is up to about 20 cm (8 in).[9] Cicadas are also notable for the great length of time some species take to mature.[10]

At least 3,000 cicada species are distributed worldwide, in essentially any habitat that has deciduous trees, with the majority being in the tropics. Most genera are restricted to a single biogeographical region, and many species have a very limited range. This high degree of endemism has been used to study the biogeography of complex island groups such as in Indonesia and Asia.[11] There are several hundred described species in Australia and New Zealand,[c] around 150 in South Africa, over 170 in America north of Mexico,[12] at least 800 in Latin America,[13] and over 200 in Southeast Asia and the Western Pacific.[14]

About 100 species occur in the Palaearctic. A few species are found in southern Europe,[10] and a single species was known from England, the New Forest cicada, Cicadetta montana, which also occurs in continental Europe.[15] Many species await formal description and many well-known species are yet to be studied carefully using modern acoustic analysis tools that allow their songs to be characterized.

A phylogenetic treatment suggested by a 2018 study.[8]

Many of the North American species are the annual or jarfly or dog-day cicadas, members of the Neotibicen, Megatibicen, or Hadoa genera, so named because they emerge in late July and August.[16] The best-known North American genus, however, may be Magicicada. These periodical cicadas have an extremely long life cycle of 13 or 17 years, with adults suddenly and briefly emerging in large numbers.[16][17]

Australian cicadas are found on tropical islands and cold coastal beaches around Tasmania, in tropical wetlands, high and low deserts, alpine areas of New South Wales and Victoria, large cities including Sydney, Melbourne, and Brisbane, and Tasmanian highlands and snowfields. Many of them have common names such as cherry nose, brown baker, red eye, greengrocer, yellow Monday, whisky drinker, double drummer, and black prince. The Australian greengrocer, Cyclochila australasiae, is among the loudest insects in the world.[18]

A chorus cicada, a species endemic to New Zealand
Cicadas in Japan

More than 40 species from five genera populate New Zealand, ranging from sea level to mountain tops, and all are endemic to New Zealand and its surrounding islands (Kermadec Islands, Chatham Islands). One species is found on Norfolk Island, which technically is part of Australia.[19] The closest relatives of the NZ cicadas live in New Caledonia and Australia.

Mesozoic fossil fore wing of Mesogereon superbum, Australia

Palaeontology

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Fossil Cicadomorpha first appeared in the Late Triassic. The superfamily Palaeontinoidea contains three families. The Upper Permian Dunstaniidae are found in Australia and South Africa, and also in younger rocks from China. The Upper Triassic Mesogereonidae are found in Australia and South Africa.[20] This group, though, is currently thought to be more distantly related to Cicadomorpha than previously thought.[21]

The giant cicada Prolystra lithographica from Germany Jurassic, about 145–150 million years ago

The Palaeontinidae or "giant cicadas" (though only distantly related to true cicadas) come from the Jurassic and Lower Cretaceous of Eurasia and South America.[20] The first of these was a fore wing discovered in the Taynton Limestone Formation of Oxfordshire, England; it was initially described as a butterfly in 1873, before being recognised as a cicada-like form and renamed Palaeontina oolitica.[22]

Tettigarctidae and Cicadidae had diverged from each other prior to or during the Jurassic, as evidenced by fossils related to both lineages present by the Middle Jurassic (~165 million years ago). The morphology of well preserved fossils of early relatives of Cicadidae from the mid Cretaceous Burmese amber of Myanmar suggests that unlike many modern cicadids, they were either silent or only made quiet sounds.[23] Most fossil Cicadidae are known from the Cenozoic,[24] and the oldest unambiguously identified modern cicadid is Davispia bearcreekensis (subfamily Tibicininae) from the Paleocene, around 56–59 million years ago.[25]

Biology

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Description

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A Japanese Min-min-zemi, called with onomatopoeia (Hyalessa maculaticollis, annual cicada)
A Japanese Nii-nii-zemi (ニイニイゼミ), called with onomatopoeia (Platypleura kaempferi, annual cicada

Cicadas are large insects made conspicuous by the courtship calls of the males. They are characterized by having three joints in their tarsi, and having small antennae with conical bases and three to six segments, including a seta at the tip.[26] The Auchenorrhyncha differ from other hemipterans by having a rostrum that arises from the posteroventral part of the head, complex sound-producing membranes, and a mechanism for linking the wings that involves a down-rolled edging on the rear of the fore wing and an upwardly protruding flap on the hind wing.[11]

Cicadas are feeble jumpers, and nymphs lack the ability to jump altogether. Another defining characteristic is the adaptations of the fore limbs of nymphs for underground life. The relict family Tettigarctidae differs from the Cicadidae in having the prothorax extending as far as the scutellum, and by lacking the tympanal apparatus.[11]

A black cicada just after molting in the garden of a private house (Midwest Saitama Prefecture, Japan)

The adult insect, known as an imago, is 2 to 5 cm (1 to 2 in) in total length in most species. The largest, the empress cicada (Megapomponia imperatoria), has a head-body length around 7 cm (2.8 in), and its wingspan is 18–20 cm (7–8 in).[10][27] Cicadas have prominent compound eyes set wide apart on the sides of the head. The short antennae protrude between the eyes or in front of them. They also have three small ocelli located on the top of the head in a triangle between the two large eyes; this distinguishes cicadas from other members of the Hemiptera. The mouthparts form a long, sharp rostrum that they insert into the plant to feed.[28] The postclypeus is a large, nose-like structure that lies between the eyes and makes up most of the front of the head; it contains the pumping musculature.[29]

The thorax has three segments and houses the powerful wing muscles. They have two pairs of membranous wings that may be hyaline, cloudy, or pigmented. The wing venation varies between species and may help in identification. The middle thoracic segment has an operculum on the underside, which may extend posteriorly and obscure parts of the abdomen. The abdomen is segmented, with the hindermost segments housing the reproductive organs, and terminates in females with a large, saw-edged ovipositor. In males, the abdomen is largely hollow and used as a resonating chamber.[28]

The surface of the fore wing is superhydrophobic; it is covered with minute, waxy cones, blunt spikes that create a water-repellent film. Rain rolls across the surface, removing dirt in the process. In the absence of rain, dew condenses on the wings. When the droplets coalesce, the cicada leaps several millimetres into the air, which also serves to clean the wings.[30] Bacteria landing on the wing surface are not repelled; rather, their membranes are torn apart by the nanoscale-sized spikes, making the wing surface the first-known biomaterial that can kill bacteria.[31]

Temperature regulation

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Desert cicadas such as Diceroprocta apache are unusual among insects in controlling their temperature by evaporative cooling, analogous to sweating in mammals. When their temperature rises above about 39 °C (102 °F), they suck excess sap from the food plants and extrude the excess water through pores in the tergum at a modest cost in energy. Such a rapid loss of water can be sustained only by feeding on water-rich xylem sap. At lower temperatures, feeding cicadas would normally need to excrete the excess water. By evaporative cooling, desert cicadas can reduce their bodily temperature by some 5 °C.[32][33] Some non-desert cicada species such as Magicicada tredecem also cool themselves evaporatively, but less dramatically.[34] Conversely, many other cicadas can voluntarily raise their body temperatures as much as 22 °C (40 °F) above ambient temperature.[35]

Song

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Cicada sound-producing organs and musculature:
a, Body of male from below, showing cover-plates;
b, From above, showing drumlike tymbals;
c, Section, muscles that vibrate tymbals;
d, A tymbal at rest;
e, Thrown into vibration, as when singing

The "singing" of male cicadas is produced principally and in the majority of species using a special structure called a tymbal, a pair of which lies below each side of the anterior abdominal region. The structure is buckled by muscular action and, being made of resilin, unbuckles rapidly on muscle relaxation, producing their characteristic sounds. Some cicadas, however, have mechanisms for stridulation, sometimes in addition to the tymbals. Here, the wings are rubbed over a series of midthoracic ridges. In the Chinese species Subpsaltria yangi, both males and females can stridulate.[36] The sounds may further be modulated by membranous coverings and by resonant cavities.[26]

The male abdomen in some species is largely hollow, and acts as a sound box. By rapidly vibrating these membranes, a cicada combines the clicks into apparently continuous notes, and enlarged chambers derived from the tracheae serve as resonating chambers with which it amplifies the sound. The cicada also modulates the song by positioning its abdomen toward or away from the substrate (their perch). Partly by the pattern in which it combines the clicks, each species produces its own distinctive mating songs and acoustic signals, ensuring that the song attracts only appropriate mates.[16] The tettigarctid (or hairy) cicadas have rudimentary tymbals in both sexes and do not produce airborne sounds, rather, both males and females produce vibrations that are transmitted through the tree they perch upon. They are considered as representing the original state from which other cicada communication has evolved.[37]

Average temperature of the natural habitat for the South American species Fidicina rana is about 29 °C (84 °F). During sound production, the temperature of the tymbal muscles was found to be significantly higher.[38] Many cicadas sing most actively during the hottest hours of a summer day; roughly a 24-hour cycle.[39] Most cicadas are diurnal in their calling and depend on external heat to warm them up, while a few are capable of raising their temperatures using muscle action and some species are known to call at dusk.[35] Kanakia gigas and Froggattoides typicus are among the few that are known to be truly nocturnal and there may be other nocturnal species living in tropical forests.[40][41]

Cicadas call from varying heights on trees. Where multiple species occur, the species may use different heights and timing of calling.[42][43] While the vast majority of cicadas call from above the ground, two Californian species, Okanagana pallidula and O. vanduzeei are known to call from hollows made at the base of the tree below the ground level. The adaptive significance is unclear, as the calls are not amplified or modified by the burrow structure, but this may avoid predation.[44]

Although only males produce the cicadas' distinctive sounds, both sexes have membranous structures called tympana (singular – tympanum) by which they detect sounds, the equivalent of having ears. Males disable their own tympana while calling, thereby preventing damage to their hearing; a necessity partly because some cicadas produce sounds up to 120 dB (SPL)[45] which is among the loudest of all insect-produced sounds.[46] The song is loud enough to cause permanent hearing loss in humans should the cicada be at "close range". In contrast, some small species have songs so high in pitch that they are inaudible to humans.[47]

For the human ear, telling precisely where a cicada song originates is often difficult. The pitch is nearly constant, the sound is continuous to the human ear, and cicadas sing in scattered groups. In addition to the mating song, many species have a distinct distress call, usually a broken and erratic sound emitted by the insect when seized or panicked. Some species also have courtship songs, generally quieter, and produced after a female has been drawn to the calling song. Males also produce encounter calls, whether in courtship or to maintain personal space within choruses.[48]

The songs of cicadas are considered by entomologists to be unique to a given species, and a number of resources exist to collect and analyse cicada sounds.[49]

Life cycle

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In some species of cicadas, the males remain in one location and call to attract females. Sometimes, several males aggregate and call in chorus. In other species, the males move from place to place, usually with quieter calls, while searching for females. The Tettigarctidae differ from other cicadas in producing vibrations in the substrate rather than audible sounds. After mating, the female cuts slits into the bark of a twig where she deposits her eggs.[11] Both male and female cicadas die within a few weeks after emerging from the soil. Although they have mouthparts and are able to consume some plant liquids for nutrition, the amount eaten is very small and the insects have a natural adult lifespan of less than two months.

When the eggs hatch, the newly hatched nymphs drop to the ground and burrow. Cicadas live underground as nymphs for most of their lives at depths of about 6–24 in (15–61 cm). Nymphs have strong front legs for digging and excavating chambers near to roots, where they feed on xylem sap. In the process, their bodies and interior of the burrow become coated in anal fluids. In wet habitats, larger species construct mud towers above ground to aerate their burrows. In the final nymphal instar, they construct an exit tunnel to the surface and emerge.[11] They then molt (shed their skins) on a nearby plant for the last time, and emerge as adults. The exuviae or abandoned exoskeletons remain, still clinging to the bark of the tree.[50]

Most cicadas go through a life cycle that lasts 2–5 years. Some species have much longer life cycles, such as the North American genus, Magicicada, which has a number of distinct "broods" that go through either a 17-year (Brood XIII), or in some parts of the region, a 13-year (Brood XIX) life cycle[51] The long life cycles may have developed as a response to predators, such as the cicada killer wasp and praying mantis.[52][53][54] A specialist predator with a shorter life cycle of at least two years could not reliably prey upon the cicadas;[55] for example, a 17-year cicada with a predator with a five-year life cycle will only be threatened by a peak predator population every 85 (5 × 17) years, while a non-prime cycle such as 15 would be endangered at every year of emergence.[56] An alternate hypothesis is that these long life cycles evolved during the ice ages so as to overcome cold spells, and that as species co-emerged and hybridized, they left distinct species that did not hybridize having periods matching prime numbers.[57] The 13- and 17-year cicadas only emerge in the midwestern and eastern US in the same year every 221 years (13 × 17), with 2024 being the first such year since 1803.[51]

Diet

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Cicada nymphs drink sap from the xylem of various species of trees, including oak, cypress, willow, ash, and maple. While common folklore indicates that adults do not eat, they actually do drink plant sap using their sucking mouthparts.[58][59]

Cicadas excrete fluid in streams of droplets due to their high volume consumption of xylem sap.[60] The jets of urine that cicadas produce have a velocity of up to 3 meters per second, making them the fastest among all assessed animals, including mammals like elephants and horses.[61][62]

Locomotion

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Cicadas, unlike other Auchenorrhyncha, are not adapted for jumping (saltation).[63] They have the usual insect modes of locomotion, walking and flight, but they do not walk or run well, and take to the wing to travel distances greater than a few centimetres.[11]

Predators, parasites, and pathogens

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Eastern cicada killer wasp (Sphecius speciosus) with cicada prey, United States

Cicadas are commonly eaten by birds and mammals,[64] as well as bats, wasps, mantises, spiders, and robber flies. In times of mass emergence of cicadas, various amphibians, fish, reptiles, mammals, and birds change their foraging habits so as to benefit from the glut. Newly hatched nymphs may be eaten by ants, and nymphs living underground are preyed on by burrowing mammals such as moles.[28] In northern Japan, brown bears prey on final instar nymphs of cicadas during summer by digging up the ground.[65] In Australia, cicadas are preyed on by the Australian cicada killer wasp (Exeirus lateritius), which stings and stuns cicadas high in the trees, making them drop to the ground, where the cicada hunter mounts and carries them, pushing with its hind legs, sometimes over a distance of 100 m, until they can be shoved down into its burrow, where the numb cicadas are placed onto one of many shelves in a "catacomb", to form the food stock for the wasp grub that grows out of the egg deposited there.[66] A katydid predator from Australia is capable of attracting singing male cicadas of a variety of species by imitating the timed click replies of sexually receptive female cicadas, which respond in pair formation by flicking their wings.[67] Their prime-number life cycle prevents predators with a life cycle of two or more years from synchronising with their emergence.[56]

Several fungal diseases infect and kill adult cicadas, while other fungi in the genera Ophiocordyceps and Isaria attack nymphs.[28] Massospora cicadina specifically attacks the adults of periodical cicadas, the spores remaining dormant in the soil between outbreaks.[68] This fungus is also capable of dosing cicadas with psilocybin, the psychedelic drug found in magic mushrooms, as well as cathinone, an alkaloid similar to various amphetamines. These chemicals alter the behaviour of the cicadas, driving males to copulate, including attempts with males, and is thought to be beneficial to the fungus, as the fungal spores are dispersed by a larger number of infected carriers.[69]

Plants can also defend themselves against cicadas. Although cicadas can feed on the roots of gymnosperms, it has been found that resinous conifers such as pine do not allow the eggs of Magicicada to hatch, the resin sealing up the egg cavities.[70][71]

Antipredator adaptations

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Cicada disruptively camouflaged on an olive tree

Cicadas use a variety of strategies to evade predators. Large cicadas can fly rapidly to escape if disturbed.[72] Many are extremely well camouflaged[72][73] to evade predators such as birds that hunt by sight. Being coloured like tree bark and disruptively patterned to break up their outlines, they are difficult to discern;[74] their partly transparent wings are held over the body and pressed close to the substrate. Some cicada species play dead when threatened.[75][76]

The day-flying cicada Huechys sanguinea warns off predators with its aposematic red and black coloration. (Southeast Asia)

Some cicadas such as Hemisciera maculipennis display bright deimatic flash coloration on their hind wings when threatened; the sudden contrast helps to startle predators, giving the cicadas time to escape.[77] Most cicadas are diurnal and rely on camouflage when at rest, but some species use aposematism-related Batesian mimicry, wearing the bright colors that warn of toxicity in other animals; the Malaysian Huechys sanguinea has conspicuous red and black warning coloration, is diurnal, and boldly flies about in full view of possible predators.[78]

Predators such as the sarcophagid fly Emblemasoma hunt cicadas by sound, being attracted to their songs.[79] Singing males soften their song so that the attention of the listener gets distracted to neighbouring louder singers, or cease singing altogether as a predator approaches. A loud cicada song, especially in chorus, has been asserted to repel predators, but observations of predator responses refute the claim.[80]

In human culture

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In art and literature

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Silver casket with writing utensils, made by the Nuremberg goldsmith Wenzel Jamnitzer (1507/08–1585): a silver cicada is at lower left.
Japanese snuff bottle in the form of a cicada, c. 1900

Cicadas have been featured in literature since the time of Homer's Iliad, and as motifs in decorative art from the Chinese Shang dynasty (1766–1122 BCE).[d] They are described by Aristotle in his History of Animals and by Pliny the Elder in his Natural History; their mechanism of sound production is mentioned by Hesiod in his poem "Works and Days": "when the Skolymus flowers, and the tuneful Tettix sitting on his tree in the weary summer season pours forth from under his wings his shrill song".[82] In the classic 14th-century Chinese novel Romance of the Three Kingdoms, Diaochan took her name from the sable (diāo) tails and jade decorations in the shape of cicadas (chán), which adorned the hats of high-level officials.

In the Japanese novel The Tale of Genji, the title character poetically likens one of his many love interests to a cicada for the way she delicately sheds her robe the way a cicada sheds its shell when molting. Cicada exuviae play a role in the manga Winter Cicada. Cicadas are a frequent subject of haiku, where, depending on type, they can indicate spring, summer, or autumn.[83] Shaun Tan's illustrated book Cicada tells the story of a hardworking but underappreciated cicada working in an office.[84] Branden Jacobs-Jenkins's play Appropriate takes place on an Arkansas farm in summer, and calls for the sounds of mating cicadas to underscore the entire show.[85]

In fashion

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Being lightweight, and with hooklike legs, the exuviae of cicadas can be used as hair or clothing accessories.[86]

As food and folk medicine

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Deep-fried Cryptotympana atrata in Shandong cuisine

Cicadas were eaten in Ancient Greece, and are consumed in selected regions in modern China, both as adults and (more often) as nymphs.[87] Cicadas are also eaten in Malaysia, Burma, North America, and central Africa, as well as the Balochistan region of Pakistan, especially in Ziarat.[88] Female cicadas are prized for being meatier.[47] Shells of cicadas are employed in traditional Chinese medicines,[89] claiming that they possess anti-convulsive, sedative, and hypothermic effects.[90] The 17-year "Onondaga Brood"[91] Magicicada is culturally important and a particular delicacy to the Onondaga people,[92] and are considered a novelty food item by modern consumers in several states.[93]

In music

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Cicadas are featured in the protest song "Como La Cigarra" ("Like the Cicada") written by Argentinian poet and composer María Elena Walsh. In the song, the cicada is a symbol of survival and defiance against death. The song was recorded by Mercedes Sosa, among other Latin American musicians.

In North America and Mexico, there is a well-known song, "La Cigarra" ("The Cicada"), written by Raymundo Perez Soto, which is a song in the Mariachi tradition, that romanticises the insect as a creature that sings until it dies.[94]

Brazilian artist Lenine with his track "Malvadeza" from the album Chão, creates a song built upon the sound of the cicada that can be heard along the track.[95]

Cicada sounds heavily feature on the 2021 album Solar Power by New Zealand artist Lorde. She described cicada song as being emblematic of the New Zealand summer.[96]

In mythology and folklore

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Jade cicada amulets. Western Han Dynasty 206 BCE – CE 8

Cicadas have been used as money, in folk medicine, to forecast the weather, to provide song (in China), and in folklore and myths around the world.[97] In France, the cicada represents the folklore of Provence and the Mediterranean cities.[98]

The cicada has represented insouciance since classical antiquity. Jean de La Fontaine began his collection of fables Les fables de La Fontaine with the story "La Cigale et la Fourmi" ("The Cicada and the Ant") based on one of Aesop's fables; in it, the cicada spends the summer singing, while the ant stores away food, and the cicada finds herself without food when the weather turns bitter.[99]

In Chinese tradition, the cicada (, chán) symbolises rebirth and immortality.[100] In the Chinese essay "Thirty-Six Stratagems", the phrase "to shed the golden cicada skin" (simplified Chinese: 金蝉脱壳; traditional Chinese: 金蟬脫殼; pinyin: jīnchán tuōqiào) is the poetic name for using a decoy (leaving the exuviae) to fool enemies.[101] In the Chinese classic novel Journey to the West (16th century), the protagonist Priest of Tang was named the Golden Cicada.[102]

In Japan, the cicada is associated with the summer season.[103] For many Japanese people, summer hasn't officially begun until the first songs of the cicada are heard.[104] According to Lafcadio Hearn, the song of Meimuna opalifera, called tsuku-tsuku boshi, is said to indicate the end of summer, and it is called so because of its particular call.[105]

In the Homeric Hymn to Aphrodite, the goddess Aphrodite retells the legend of how Eos, the goddess of the dawn, requested Zeus to let her lover Tithonus live forever as an immortal.[106] Zeus granted her request, but because Eos forgot to ask him to also make Tithonus ageless, Tithonus never died, but he did grow old.[106] Eventually, he became so tiny and shriveled that he turned into the first cicada.[106] The Greeks also used a cicada sitting on a harp as an emblem of music.[107]

In Kapampangan mythology in the Philippines, the goddess of dusk, Sisilim, is said to be greeted by the sounds and appearances of cicadas whenever she appears.[108]

As pests

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Cicadas feed on sap; they do not bite or sting in a true sense, but may occasionally mistake a person's arm for a plant limb and attempt to feed.[109] Male cicadas produce very loud calls that can damage human hearing.[110]

Cicadas are not major agricultural pests, but in some outbreak years, trees may be overwhelmed by the sheer numbers of females laying their eggs in the shoots. Small trees may wilt and larger trees may lose small branches.[28] Although in general, the feeding activities of the nymphs do little damage, during the year before an outbreak of periodic cicadas, the large nymphs feed heavily and plant growth may suffer.[111] Some species have turned from wild grasses to sugarcane, which affects the crop adversely, and in a few isolated cases, females have oviposited on cash crops such as date palms, grape vines, citrus trees, asparagus, and cotton.[28]

Cicadas sometimes cause damage to ornamental shrubs and trees, mainly in the form of scarring left on tree branches where the females have laid their eggs. Branches of young trees may die as a result.[112][113][failed verification]

See also

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Notes

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References

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Further reading

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Cicadas are belonging to the superfamily Cicadoidea, primarily the family within the order , characterized by their large size—typically 1 to 2 inches (2.5 to 5 cm) long—bullet-shaped bodies, transparent wings, and prominent eyes positioned on the sides of the head. They are renowned for the males' loud, species-specific calls produced by vibrating specialized membranes called tymbals, which are amplified by resonant cavities in the , serving primarily to attract mates. Distributed worldwide in tropical and temperate regions, cicadas include more than 3,000 described species, with estimates suggesting over 5,000 including undescribed taxa; nymphs spend most of their lives underground feeding on from before emerging as short-lived adults that feed on twig and reproduce. Cicadas are broadly classified into annual (or "dog-day") species, which emerge every summer in overlapping generations after 2 to 5 years underground, and periodical species in the genus Magicicada, which synchronize massive emergences every 13 or 17 years across eastern . Annual cicadas, such as those in the genus , are often green and black with dark eyes, active from July to September, and produce a continuous buzzing resembling a buzz-saw. In contrast, periodical cicadas feature black bodies, reddish eyes, and orange-tinged wings, emerging en masse—sometimes reaching 1.5 million individuals per acre—when soil temperatures hit about 64°F (18°C) in late spring, overwhelming predators through sheer numbers in a phenomenon known as . Ecologically, cicadas play a key role as herbivores, with nymphs potentially damaging young over years of feeding, while adult oviposition—where females use saw-like ovipositors to slit twigs and lay up to 600 eggs—can weaken branches, though populations rarely cause widespread harm. Harmless to humans, lacking the ability to bite or sting, cicadas have a fossil record with the superfamily dating back to the period over 200 million years ago, with their periodic emergences inspiring cultural references and scientific study into synchronized behaviors.

Etymology and nomenclature

Etymology

The English word "cicada" is derived from the Latin cicada, an onomatopoeic term mimicking the insect's characteristic chirping or buzzing sound produced by vibrating specialized membranes called tymbals. This Latin name likely originated as a Mediterranean , possibly influenced by earlier onomatopoeic forms in regional languages that echoed the rhythmic noises of these . In , the equivalent term was tétrix (τέττιξ), also onomatopoeic, capturing the shrill, repetitive calls of the creatures and used in classical texts to denote similar singing . Ancient writers provided early linguistic and descriptive references to cicadas, embedding their names in natural history. Aristotle, in his History of Animals (circa 350 BCE), described the lifecycle of the tétrix, including its transformation from grub to nymph (tettigometra) and adult stages, noting that males produce chirping sounds while females are silent, and distinguishing it from other insects based on its generation and vocalization. Similarly, Pliny the Elder, in Natural History (circa 77 CE), described the cicada in Latin, noting its lifecycle divisions into singing and silent varieties and its habitat preferences, while attributing its name to the audible emissions that define its presence. Related terms like "" in English have often been conflated with cicada due to superficial resemblances in swarming behavior and noise, though their etymologies differ; "locust" stems from Latin , originally denoting a type of or even a lobster-like , without direct phonetic ties to sounds. This confusion arose historically in English-speaking regions, where early observers applied "locust" broadly to noisy, emergent insects like cicadas, despite the latter's distinct onomatopoeic naming rooted in their specific vocalizations. The scientific nomenclature of cicadas evolved significantly with Carl Linnaeus's binomial system in (1758), where he established the genus for these , formalizing their based on morphological and auditory traits observed in . This Linnaean framework laid the foundation for the family Cicadidae, formally established by Batsch in 1789, shifting from folk etymologies to a standardized, sound-inspired that persists in modern .

Common names and classification

Cicadas are known by various regional common names that often reflect their distinctive buzzing sounds or local cultural associations. In the , they are frequently called "jar flies," a term derived from the verb "jar" meaning to buzz or vibrate harshly, mimicking the harsh, rattling noise produced by species like those in the genus Neotibicen during late summer. In , particularly in , cicadas are referred to as "cigale," a name rooted in Latin cicada and symbolizing good fortune and the vibrant summer landscape, where their songs are an iconic feature of the region's identity. In , the term "semi" is used, which phonetically evokes the cicadas' shrill calls and holds cultural significance as a symbol of summer and seasonal transience in and . In Spanish-speaking regions, cicadas are known as "cigarra" or "chicharra", terms that similarly reflect their characteristic loud, continuous buzzing sounds. These common names sometimes lead to confusion with other insects, particularly grasshoppers and true locusts, which are species of grasshoppers (: ) that form swarming plagues. Cicadas, belonging to the order , are often mistakenly called "locusts" in due to their mass emergences, fostering public perceptions of them as destructive pests akin to biblical plagues, despite being harmless herbivores that do not swarm or damage crops significantly. This persists regionally, affecting awareness and leading to unnecessary fear during periodical outbreaks. Scientifically, cicadas are classified using , a system established by in which each is denoted by a two-part Latin or Latinized name: the (capitalized) followed by the (lowercase), both italicized, and often accompanied by the (the describer's name and publication year). For example, the periodical cicada (Linnaeus, 1758) refers to the 17-year , with Linnaeus as the based on his original description. The for the Cicada, which anchors the family , is Cicada orni (Linnaeus, 1758), selected to represent the group's defining characteristics. The naming of cicadas adheres to the (ICZN), which ensures stability and universality in . Key principles include priority (the earliest valid name prevails), uniqueness (no two taxa share the same name), and the requirement for names to be in Latin form, with type specimens (holotypes) designated to fix the application of names; for cicadids, this applies to over 3,000 species across 400+ genera, preventing synonymy and resolving disputes through the .

Taxonomy and evolution

Classification and phylogeny

Cicadas belong to the order within the class Insecta, specifically in the suborder , infraorder , superfamily Cicadoidea, and family . This placement reflects their shared characteristics with other true bugs, such as piercing-sucking mouthparts, while distinguishing them through specialized morphological and behavioral traits like sound production. The superfamily Cicadoidea is divided into two families: the relict with only two extant species in , and the more diverse , which encompasses the vast majority of cicada species worldwide. The family Cicadidae includes approximately 3,000 described distributed across more than 250 . These are organized into five recognized subfamilies: Cicadinae, Cicadettinae (formerly referred to as Tettigettinae in some classifications), Tibicininae, Tettigomyiinae, and Derotettiginae. Prominent genera exemplify this diversity; for instance, the genus Cicada (type genus of Cicadinae) includes widespread , Magicicada (in Cicadettinae) comprises the of eastern known for synchronized emergences, and Quesada (in Tibicininae) features large, annual from the tropics. Tribal divisions within these subfamilies further refine the hierarchy, with over 50 tribes identified based on morphological and molecular evidence. Phylogenetic analyses of Cicadidae have relied on molecular markers, including 18S rRNA for ribosomal structure and subunit I (COI) for mitochondrial variation, to reconstruct evolutionary relationships. These studies confirm that cicadas diverged from other hemipteran lineages, particularly , around 300 million years ago during the late to early Permian, marking the early radiation of . Within Cicadoidea, the split between and occurred later, likely in the , with diversifying into its modern subfamilies through a combination of and ecological adaptations. In the 2020s, taxonomic revisions have refined classifications, particularly for species in Tibicininae, integrating genital morphology—such as pygofer and structures—with genetic data from multi-locus phylogenies. A key 2023 study redefined genera like Tibicinoides and Okanagana, transferring multiple species based on molecular evidence showing and using morphological traits to resolve rapid radiations in . These updates highlight ongoing efforts to align with evolutionary history, reducing paraphyletic groupings in regional faunas.

Diversity and distribution

Cicadas comprise over 3,000 described species worldwide, belonging to the superfamily Cicadoidea, with the vast majority occurring in tropical and subtropical habitats that provide suitable conditions for their life cycles. Species diversity peaks in tropical regions, where warm climates and abundant vegetation support high richness; for instance, alone harbors more than 200 species across diverse ecosystems, while features over 150 species, many endemic to forested islands and mainland tropics. Biogeographic hotspots of include the archipelagoes and the , where unique evolutionary radiations have produced specialized assemblages, such as the seven species of the Magicicada—all —confined to the . Globally, distribution patterns reflect climatic tolerances: periodical species like Magicicada are restricted to temperate zones in , enabling synchronized mass emergences, whereas most non-periodical species form continuous broods in tropical forests; cicadas are notably absent from polar regions, where cold temperatures and lack of host trees preclude survival. Recent taxonomic advances, driven by acoustic recordings and , have revealed new species in understudied tropical areas, including 12 from described in 2021 and 13 from in 2024, underscoring the untapped diversity in regions like hotspots and continental rainforests.

Fossil record

The fossil record of cicadas, encompassing the superfamily Cicadoidea within the broader , extends back to the Permian period, approximately 290 million years ago, marking the earliest known appearances of stem-group cicadomorphans. These primitive forms, such as members of the family Dysmorphoptilidae from deposits in and , exhibited early hemipteran traits including piercing-sucking mouthparts adapted for feeding and wing venation patterns indicative of jumping locomotion similar to modern leafhoppers. By the middle Permian, cicadomorphans had become dominant herbivores in certain ecosystems, with s from the Kungurian stage (around 283–272 Ma) in the Tunguska Basin of revealing inflated frontoclypeal structures and sessile morphologies that foreshadowed later cicada adaptations. During the era, cicadomorphans underwent significant diversification, with true cicadoids emerging by the . Exceptional preservation in the Daohugou Beds of , (approximately 165 Ma), has yielded genera such as Macrotettigarcta and Shuraboprosbole, showcasing advanced wing structures and body plans closely resembling extant cicadas. These fossils indicate the divergence of key lineages, including the relict family and the more derived , with evidence of early acoustic communication emerging in Permian dysmorphoptilids through specialized wing modifications for sound production—predating modern organs but hinting at ancestral stridulatory mechanisms in hemipterans. Cenozoic records provide insights into modern cicada morphologies, with amber-preserved specimens revealing detailed soft-tissue preservation. Eocene amber from the Baltic region contains nymphs of cicadids, approximately 44 million years old, displaying root-feeding behaviors and burrowing adaptations akin to contemporary species, while Dominican amber inclusions from the Oligocene-Miocene boundary (around 23–20 Ma) include adults like Minyscapheus dominicanus, with intact wing venation and ovipositor structures. A notable 2025 discovery from the Eocene Messel Pit in Germany revealed a well-preserved adult cicada fossil, approximately 47 million years old, providing evidence of early acoustic structures in European cicadids. These fossils highlight a post-Cretaceous-Paleogene recovery, where cicada diversity appears reduced immediately after the event—likely due to habitat disruptions from the extinction of non-avian dinosaurs and associated vegetation changes—but rebounded with the radiation of angiosperms in the Paleogene, facilitating host shifts to new xylem sources among root-feeding nymphs.

Physical characteristics

Morphology

Cicadas exhibit a typical body plan, divided into three distinct tagmata: the head, , and . The head is short and wide, featuring prominent compound eyes positioned wide apart on short eyestalks, which provide a broad field of vision, and three simple eyes (ocelli) arranged in a triangular formation on the vertex. Short antennae, consisting of 7-9 segments, arise between the eyes, and the mouthparts are modified into an elongated rostrum formed by the labium, which houses a bundle of piercing stylets for feeding. The is robust and segmented into pro-, meso-, and metathorax, bearing three pairs of legs adapted for walking and grasping—the forelegs are particularly prehensile—and two pairs of membranous wings. The forewings, attached to the mesothorax, are larger and typically transparent with visible venation, though some display patterned coloration; at rest, they are held roof-like over the . The hindwings, smaller and attached to the metathorax, are folded fan-like beneath the forewings when not in use. The abdomen comprises 10 segments and houses key external structures, including paired tympanal organs—thin, oval membranes located ventrally on the second segment—for auditory function. Males possess enlarged tymbals, ribbed membranes on the dorsolateral sides of the first abdominal segment, while females feature a robust, saw-like at the posterior end for inserting eggs into plant tissues. These sexual differences in abdominal structures facilitate , with variations in overall size and coloration addressed in subsequent discussions of dimorphism.

Size, coloration, and sexual dimorphism

Cicadas vary considerably in body size, with most measuring 1 to 5 cm in length. Larger , such as the Southeast Asian Tacua speciosa, attain a head-body length of 4.7–5.7 cm and a of 15–18 cm, making it one of the largest known cicadas. These size differences reflect adaptations to diverse habitats and ecological roles across the family's approximately 3,000 . Coloration in cicadas is typically dominated by shades of black, green, or brown, providing effective against tree bark and foliage. Distinctive patterns occur in certain taxa, including red compound eyes and orange-tinged wing veins in of the genus Magicicada. Intraspecific color variations are common, such as differences in (e.g., , , or ) and wing vein pigmentation, which may arise from genetic or environmental factors without altering species classification. Sexual dimorphism in cicadas primarily manifests in morphology and size rather than coloration. Females are often heavier and equipped with a strong, pointed for inserting eggs into tissue, while males possess a blunter and specialized structures for acoustic signaling. The degree of size dimorphism varies by and ; for instance, in some North American taxa, females exceed males in body mass and , enhancing their oviposition capabilities, whereas males may exhibit relatively greater maneuverability. Coloration differences between sexes are subtle and not widespread, though habitat-linked intraspecific morphs—such as greener individuals in forested environments versus browner forms in arid regions—can influence overall appearance across populations.

Physiology

Thermoregulation

Cicadas are ectothermic insects that primarily rely on behavioral thermoregulation to maintain body temperatures suitable for activity, typically optimizing around 30–35°C through basking in sunlight to absorb solar radiation or seeking shade to avoid overheating. Species adjust their body orientation, such as exposing lateral surfaces to the sun or using wings as a parasol, to fine-tune heat gain, with activity often suspended when ambient temperatures exceed thresholds like 40–42°C. This behavioral strategy allows cicadas to achieve thoracic temperatures up to 13°C above ambient during warm-up, enabling flight and other functions even in suboptimal conditions. Physiologically, cicadas supplement behavior with endothermic production via rapid contractions of thoracic flight muscles, raising body temperature by 10–12°C above ambient to support activities like singing, where metabolic rates increase significantly during choruses. For cooling, they employ cuticular by expelling water from fluids through spiracles, which can lower body temperature up to 6.7°C below ambient, aided by circulation to distribute or moisture. Maximum voluntary tolerance temperatures range from 36–38°C across , beyond which stress impairs function, including reduced calling above approximately 40°C due to onset near 46–47°C. In like Magicicada species, nymphs underground benefit from 's , which buffers extreme surface fluctuations and maintains stable conditions during their multi-year development, with triggered only when soil at 20 cm depth exceeds 18°C. As adults, they adapt by wing-spreading to capture solar heat at lower temperatures or fanning wings to enhance evaporative cooling in heat, helping sustain choruses in variable post- environments. These mechanisms underscore cicadas' resilience to thermal variability, though upper limits constrain activity in increasingly warm climates.

Sound production and communication

Cicadas produce sound primarily through a mechanism known as the tymbal mechanism, unique to males of the superfamily Cicadoidea. The tymbals consist of ribbed, stiff membranes located dorsolaterally on the anterior , specifically the first tergite, which are rapidly deformed by contractions of specialized tymbal muscles. These muscles vibrate the membranes at frequencies typically between 200 and 500 Hz, with each rapid buckling producing a brief click that combines into continuous song when repeated. The acts as an acoustic , amplifying the low-frequency components of the sound for efficient transmission over distances. Unlike crickets and many other insects that generate sounds via stridulation (rubbing body parts together, typically wings), producing rhythmic, intermittent chirps, cicadas produce a more continuous, high-pitched buzzing or droning noise using tymbals. Cicada songs are significantly louder than typical cricket calls, with individual males of the loudest species, such as the African cicada Brevisana brevis, reaching a mean sound pressure level of 106.7 dB at a distance of 50 cm. Male cicadas employ distinct types for communication, primarily to attract s and defend territories. Calling are long, repetitive phrases broadcast from perches to draw conspecifics into choruses, reaching peak intensities of up to 100 dB in dense aggregations of periodical such as Magicicada. Once a approaches, males switch to —shorter, more variable sequences—and elicit responses through duets, where s produce sharp wing-clicks by flicking their wings against the , signaling receptivity and aiding precise mate location. The acoustic repertoire of cicadas exhibits high diversity, with species-specific patterns in phrase duration, syllable rate, and dominant frequency facilitating mate recognition amid sympatric assemblages. For instance, different Magicicada species produce calls with unique temporal structures, such as the interrupted buzz of M. cassini versus the continuous whine of M. septendecim. Chorusing , where males synchronize calls within groups, enhances overall signal salience for females while reducing predation risk through temporal overlap that confounds predator localization. Recent bioacoustics research in the has uncovered geographic dialects in calling across populations, such as variations in phrase complexity linked to and isolation in genera like Tettigettalna, informing models of acoustic evolution and . Studies during 2024 mass emergences have also documented chorus intensities up to 86 dB via crowdsourced recordings, raising awareness of auditory discomfort for humans with differences, including those on the autism spectrum, though no permanent occurs at these levels.

Locomotion and sensory systems

Cicadas exhibit specialized locomotion adapted to their arboreal and subterranean lifestyles. Adult cicadas primarily rely on flight for dispersal and , powered by robust asynchronous flight muscles in the that enable rapid wing oscillations. These muscles drive the forewings and hindwings in a coupled motion, generating during the upstroke and managing drag in the downstroke to support forward flight and limited hovering. While capable of sustained flight over moderate distances, adults are notably clumsy on the ground, using their six spiny legs—equipped with tibial spines for grip—to walk or climb awkwardly, often preferring to launch into flight rather than navigate terrestrially. Nymphs, in contrast, are subterranean dwellers specialized for burrowing. Their powerful forelegs, modified with hook-like structures and robust segments, facilitate excavation through , allowing them to construct tunnels and chambers while feeding on root . These forelegs enable efficient forward digging, where is rolled into balls and pushed behind the body. Upon maturity, nymphs emerge from the at night, using their legs to crawl up vertical surfaces such as trunks to reach safe molting sites, minimizing predation risk during the transition to adulthood. Cicadas possess a suite of sensory organs that integrate visual, chemical, and vibratory cues for environmental . The prominent compound eyes, each comprising approximately 7,500 ommatidia in species like the Australian redeye cicada (Psaltoda moerens), provide a wide and excel at detecting motion, aiding in obstacle avoidance and predator detection during flight. Three dorsal ocelli serve as simple photoreceptors, primarily sensing light intensity and changes in illumination to orient the body and regulate daily activities, such as timing. Antennae, though short and bristle-like, house olfactory receptors that detect volatile plant compounds and potentially other chemical signals, contributing to host selection and habitat navigation. The tympanal organs on the function as auditory sensors for airborne sounds, with membranes that vibrate in response to acoustic signals from conspecifics. These sensory systems underpin key behaviors in locomotion. Optomotor responses, mediated by the compound eyes, stabilize flight paths by reflexively adjusting wing beats to counteract visual flow, ensuring straight-line trajectories during dispersal. Substrate vibration sensing via leg chordotonal organs allows cicadas to perceive and evade ground-based predators, triggering rapid flight initiation or postural changes with latencies as low as 6 ms. This integration enhances survival by linking sensory input directly to motor outputs, from burrowing adjustments in nymphs to evasive maneuvers in adults.

Life history and ecology

Life cycle and development

Cicadas undergo incomplete , also known as hemimetabolous development, consisting of three primary stages: , , and . This process lacks a distinct pupal stage, with resembling smaller, wingless versions of and undergoing gradual changes through molting. The cycle begins with deposition. Females use their specialized to cut slits into the bark of small twigs or branches, typically 1–2 years old, and lay batches of 20–30 eggs per slit, potentially creating dozens of such slits per female. Eggs incubate for 2–10 weeks, depending on species and environmental conditions, before hatching into first-instar . Upon hatching, the tiny drop to the ground and immediately into the using their forelegs, which are robust and adapted for digging with enlarged, spade-like tibiae and strong claws. Nymphs spend the majority of their lives underground, progressing through five instars over periods ranging from 2–5 years in annual species to 13 or 17 years in periodical species. During this subterranean phase, they feed on sap from plant roots using piercing-sucking mouthparts, excavating burrows typically 15 to 61 cm (6 to 24 in) deep; in wet soils, nymphs sometimes construct shallow mud chimneys (2 to 8 cm high) above the entrance for protection against flooding. Development is marked by periodic molts, where nymphs shed their to grow, with synchronization across populations often regulated by environmental cues such as soil temperature and moisture rather than strict . In the final , mature nymphs construct emergence tunnels to the surface, typically when soil temperatures reach about 18°C (64°F). Emergence culminates in the transformation to . Nymphs climb vertical surfaces like tree trunks or stems, anchor themselves, and undergo a final molt, splitting their along the back to reveal the pale, soft-bodied teneral . This molting , part of mass synchronous events in periodical species, can involve billions of individuals emerging over a few nights, overwhelming predators through sheer numbers. The new hardens within hours to days, during which adults remain vulnerable and do not fly or sing; morphological changes include the expansion and pigmentation of wings and eyes. Adult cicadas have a brief lifespan of 2–6 weeks above ground, dedicated almost entirely to . Males produce species-specific calls to attract females, occurs soon after hardening, and females then oviposit before dying; unemerged nymphs from previous cycles may persist underground. Variations in cycle length highlight adaptive strategies: annual species complete development in 2–5 years without strict , while periodical species like those in X (17-year) and XIX (13-year) align emergences precisely, as seen in the rare 2024 dual-brood event across the eastern U.S., where billions emerged in regions home to millions of people.

Periodical versus annual species

Periodical cicadas, belonging to the genus Magicicada, are characterized by highly synchronized emergences after fixed intervals of either 13 or 17 years, with seven divided into four 13-year and three 17-year primarily found in eastern . These prime-numbered cycles facilitate brood synchrony, where vast populations emerge simultaneously, a strategy hypothesized to minimize predation risk by disrupting potential predator population cycles. In contrast, annual cicadas exhibit overlapping life cycles of 2 to 5 years, resulting in consistent yearly emergences without strict synchrony, and they represent a far more diverse group globally, with thousands of species across various genera. For example, species in the genus (formerly Tibicen) are common in temperate regions of , where their staggered generations ensure a steady presence during summer months. The evolutionary advantages of these strategies differ markedly: achieve through mass emergences involving billions of individuals, overwhelming predators and allowing sufficient survivors to reproduce, whereas annual cicadas face ongoing predation pressure, favoring adaptations like and over numerical overwhelming. Recent genetic research from 2024 has modeled potential hybridization between 13- and 17-year during rare co-emergences, such as the 2024 event, suggesting genetic mixing could lead to hybrid broods with altered cycles. Additionally, studies indicate that may desynchronize periodical cycles by altering soil temperatures and emergence cues, potentially causing earlier or off-schedule emergences and reducing brood cohesion over time.

Diet and habitat preferences

Cicadas are feeders throughout their lives, relying on this fluid as their primary source of . Nymphs use specialized piercing-sucking mouthparts to extract from of trees and grasses, employing pump-like mechanisms to draw up the water-rich, nutrient-poor mixture containing and minerals. Adults continue this diet by feeding on from twigs and stems of host plants, a facilitated by the same mouthpart structures. Host plant specificity varies among cicada species and influences their distribution. Periodical cicadas, such as those in the genus Magicicada, preferentially feed on and oviposit in deciduous trees like oaks (Quercus spp.), maples (Acer spp.), and hickories (Carya spp.), which provide suitable root systems for nymphal development. In contrast, many annual cicada species in arid regions associate with grasses or conifers such as pines (Pinus spp.), adapting to sparser vegetation. Nymphs inhabit underground environments in well-drained soils, where they burrow to access root , with in urban areas posing a barrier to their movement and survival. Adults emerge into forested woodlands, edges, or urban fringes, particularly in temperate eastern North American deciduous forests for periodical species, while tropical and arid favor open grasslands or pine-dominated landscapes. The sap diet, composed of approximately 95% water and low in nutrients, necessitates specialized physiological adaptations for survival. Cicadas exhibit efficient to manage and from the hypotonic fluid, preventing or imbalances. Additionally, in their guts convert components into essential , supplementing the deficient diet and enabling long-term subterranean life in some species.

Interactions with other organisms

Predators, parasites, and pathogens

Cicadas face significant biotic threats from a diverse array of predators that target various life stages, particularly adults during mass emergences. Birds are the primary predators of adult periodical cicadas, with species such as starlings, , blue jays, and cuckoos consuming large numbers of emergents, often accounting for 15–40% of the adult population in the early phases of emergence. Mammals, including squirrels, raccoons, dogs, and moles (which prey on subterranean nymphs), also contribute substantially to mortality, while reptiles like feed on both nymphs and adults. predators, such as cicada killer wasps and spiders, further reduce populations, though overall predation rates do not scale proportionally with cicada density due to strategies. Parasitic organisms exploit cicadas across all developmental stages, often with host-specific adaptations. Parasitoid wasps, including trichogrammatids that target eggs and cecidomyiids that attack egg masses, impose limited but notable mortality on early life stages, though they do not significantly regulate overall populations. Nematodes, such as entomopathogenic species like those in the genus Steinernema, infect nymphs and adults in soil environments, causing internal damage and death, with records of natural infections in species like Diceroprocta apache. Mites, particularly the ectoparasitic Pyemotes herfsi, attach to adults and drain nutrients, leading to weakened hosts and secondary infections; during the 2024 Brood XIII emergence, mite infection rates reached up to 14% in some populations. In tropical cicada species, trematode flatworms occasionally parasitize nymphs, altering development and increasing vulnerability to other threats, though such infections are less documented in temperate broods. Bacterial endosymbionts like , detected in 41–57% of periodical cicadas during the 2024 Brood XIII emergence, are known to manipulate reproduction in some insect hosts. Pathogenic microbes, especially fungi, pose severe risks to cicada choruses, often causing episodic die-offs. The obligate fungal Massospora cicadina infects late-stage nymphs underground via resting s and manifests in adults as a "zombie" infection, replacing the posterior abdomen with a mass that manipulates behavior—inducing in males, who mimic female signals to attract mates and disperse conidiospores during prolonged flights. Infected individuals experience no apparent pain or reduced mobility, continuing to feed and mate despite organ destruction, which ultimately leads to sterility and death; infection rates during emergences typically range from 5–25%, with symptomatic cases causing up to 50% mortality in dense choruses. During the 2024 dual emergence of Broods XIII and XIX, fungal epidemics, primarily , reduced populations by 20–30% in affected U.S. Midwest and Southern areas, with DNA detection in 23% of asymptomatic cicadas indicating latent spread. Other pathogens include viruses that cause unspecified systemic infections in larvae and bacteria like those in the genus , which trigger septicemia in weakened adults, though these are less prevalent than fungal threats.

Defense mechanisms

Cicadas utilize a range of physical, behavioral, and physiological strategies to deter predators and resist parasites and pathogens. These adaptations are particularly crucial for species like , which emerge in massive numbers after long periods, making them vulnerable during short adult phases. Cryptic coloration serves as a primary visual defense for many cicada species, allowing them to blend seamlessly with tree bark, twigs, and foliage when at rest. This reduces detection by visual predators such as birds and , with adults and nymphs exhibiting mottled browns, grays, and greens that mimic their arboreal or subterranean environments. In some tropical species, such as those in the genus Hemisciera, adults employ deimatic displays by suddenly flashing brightly colored hindwings—often red or orange—creating a startling contrast that briefly disorients approaching threats, providing time for escape. Behavioral defenses are prominent in gregarious species, especially (Magicicada spp.), where mass chorusing and synchronous s overwhelm predators through a combination of acoustic saturation and numerical abundance. Males aggregate in choruses producing intense, synchronized calls that can reach deafening levels, confusing predators and diluting the risk to any individual by distributing attention across thousands or millions of individuals; densities have been recorded up to 1.5 million per acre, exemplifying . Disturbed adults often produce a defensive "alarm buzz" or struggle flight, further deterring close-range attacks. Chemical defenses in cicadas are less prominent than in related hemipterans but include unpalatable tissues in nymphs, which contain high levels of indigestible compounds and structural reinforcements that discourage burrowing predators like moles or from consuming them fully. Adults may release viscous fluids from oral or anal regions when handled, acting as a repellent to small predators, though this is not as potent as glandular secretions in true bugs. Immune responses in cicadas mirror broader insect mechanisms, with hemocytes in the hemocoel encapsulating invading parasites such as nematodes or fungal elements through multilayered cellular sheaths that isolate and melanize the intruder, preventing systemic spread. Recent research has highlighted humoral defenses, including antifungal peptides like cicadin isolated from juvenile cicada tissues, which exhibit potent activity against pathogenic fungi such as Botrytis cinerea at nanomolar concentrations, potentially aiding periodical species in resisting soil-borne infections during prolonged nymphal stages. These peptides contribute to innate immunity by disrupting fungal cell membranes, underscoring cicadas' adaptive resilience to microbial threats.

Ecological roles

Cicadas play a pivotal role in nutrient cycling within ecosystems, particularly through the mass mortality following periodical emergences. The of billions of adult carcasses introduces substantial pulses of , including and , into the . Studies estimate that these events can deposit 10 to 70 kg of per every 13 or 17 years, with similar contributions for representing up to 50% of annual leaf litter inputs in some . This nutrient enrichment enhances , stimulates microbial activity, and promotes productivity; for instance, experimental additions of cicada carcasses increased understory by 61% and foliar content by 20% compared to controls. Such inputs are especially vital in nutrient-poor soils, where they facilitate long-term and support post-emergence vegetation recovery. As a major prey resource, cicadas bolster dynamics and during their brief adult phase. The synchronized mass emergences provide a predictable, high-density pulse that sustains diverse predators, including over 80 species that shift behaviors to target cicadas, thereby reducing predation on alternative prey like caterpillars and allowing those populations to rebound. This trophic rewiring extends to mammals, reptiles, and , enhancing overall resilience and connectivity across communities. The scale of these events—often exceeding 1 million individuals per —positions cicadas as a keystone , preventing predator starvation in lean periods and promoting stable predator-prey balances over multi-year cycles. Cicadas interact with plants in ways that influence structure and function. Female oviposition scars branches by slicing slits for egg-laying, often causing localized dieback or "flagging," but long-term studies show negligible effects on radial growth or mortality across multiple species. In some cases, this injury prompts compensatory lateral branching, potentially leading to denser canopies. Meanwhile, both nymphal root-feeding and adult xylem sap consumption have minimal direct impacts on hydraulics or overall vigor, though chronic nymphal herbivory may subtly alter water and uptake without hindering net growth. These interactions, combined with soil bioturbation from nymph burrows, contribute to improved and water infiltration. Due to their long generation times and requirements, cicadas function as effective indicator species for . Periodical species demand large, unbroken patches of at least 52 hectares to sustain synchronized emergences, making them vulnerable to fragmentation from and . Populations are declining in regions affected by shifts, with altered temperature regimes disrupting developmental timing and survival; for example, some populations have dropped by up to 80% over the past decade due to warming and loss. Monitoring cicada abundances thus signals broader degradation, underscoring the need for conservation to preserve these roles.

Human interactions

Cultural and symbolic significance

In , cicadas have long symbolized rebirth and resurrection, attributed to their life cycle of emerging from the ground after years underground and shedding their exoskeleton. This association dates back to ancient times, with jade carvings of cicadas placed on the tongues of the deceased during the (206 BCE–220 CE) to ensure immortality and guide the soul in the . In ancient Greek mythology, cicadas were linked to summer and revered as sacred to Apollo, the god of and , with their shrill songs evoking and the heat of the season. , in (ca. 700 BCE), described the cicada's song as a marker of "the season of fatiguing summer," while Plato's Phaedrus (ca. 370 BCE) mythologized them as humans transformed into insects by the Muses for their devotion to song, granting them the ability to subsist on dew without food. Cicadas frequently appear in literature and art as emblems of transience and renewal. In Japanese , the 17th-century poet captured their evanescence, as in his verse "The cry of the cicada / Soaks into stone," where the insect's piercing call against unyielding rock symbolizes the brevity of life amid enduring nature. During the European Renaissance, insects like cicadas featured in still-life paintings as symbols, representing the fleeting nature of existence and the cycle of , often juxtaposed with fruits and flowers to underscore mortality. In , cicada motifs persist in tattoos, where the insect embodies personal transformation and rebirth, drawing on its metamorphic life stage to signify overcoming adversity and emerging renewed. In music, cicadas have inspired works that mimic or evoke their resonant calls. While Antonio Vivaldi's The Four Seasons (1725) programmatically depicts summer's oppressive heat in its "Summer" concerto—complete with string figures suggesting swarming insects and languid languor—no explicit cicada motif appears, though the piece's vivid portrayal of seasonal sounds has been interpreted as encompassing such natural choruses. In the 2020s, ambient and neoclassical compositions have directly incorporated cicada field recordings for atmospheric effect, as in Scott Buckley's "Cicadas" (2024), a delicate track blending piano, strings, and actual insect sounds to create a meditative soundscape of renewal and natural rhythm. Across traditions, cicadas often serve as harbingers of change or spiritual intermediaries. In Australian Aboriginal lore, natural indicators like emergences help predict seasonal shifts, including , though specific cicada stories emphasize their role in signaling the end of dry periods and the arrival of wetter weather. In some African and traditions, cicada songs are viewed as carriers of spiritual messages from ancestors or the divine, their persistent choruses interpreted as calls connecting the living to the departed.

As pests and economic impacts

Cicadas, particularly periodical species in , can cause significant damage through oviposition, where females use their ovipositors to make V-shaped in the bark of twigs and small branches to deposit eggs. These , typically in branches 3/16 to 7/16 inches in diameter, weaken the , leading to , dieback, and branch breakage in affected trees. Young orchards, such as those with apple, , or trees, are especially vulnerable during mass emergences, as the high density of egg-laying females can result in widespread flagging of foliage and reduced tree vigor. The wounds also create entry points for pathogens and secondary pests, exacerbating long-term horticultural losses in commercial fruit production. Economic impacts are primarily localized to and sectors, where oviposition damage reduces yields in orchards and nursery stock, necessitating replanting or . In urban and suburban settings, the intense chorusing of emerging adults—reaching up to 100 decibels—generates widespread complaints, disrupting daily life and potentially increasing municipal response costs, though direct effects on property values remain unquantified. During the dual-brood emergence in the eastern and midwestern U.S., these effects were amplified across 16 states. The 2025 emergence of Brood XIV across approximately 13 states, including Georgia, , , and others, similarly prompted preparations for potential tree damage, though reports indicated limited agricultural impacts overall. Management strategies focus on prevention rather than eradication, given the short adult lifespan and ecological benefits of cicadas. Cicada-proof netting with mesh smaller than 1/4 inch is recommended for young trees and shrubs up to 10 feet tall, installed before to exclude females from oviposition sites; this method is cost-effective for high-value orchards but labor-intensive for large areas. Insecticide sprays, such as or pyrethroids, can be applied during peak adult activity (first through third cover sprays) to reduce egg-laying, though experts advise against broad use due to harm to pollinators and predators. Biological controls leverage natural enemies, including birds, mammals, and insects like cicada killer wasps, which consume large numbers during emergences, minimizing the need for chemical interventions. In non-pest contexts, cicadas rarely reach damaging densities in tropical regions due to continuous predation by birds, reptiles, and parasitoids, which prevent population booms typical of temperate periodical . However, climate-driven range shifts and altered timing may expand cicada impacts northward or into new habitats, potentially increasing pest pressure in as warmer conditions favor survival and synchronization.

Uses in food, medicine, and other practices

Cicadas have been consumed as in various cultures, particularly in parts of and , where they are valued for their nutritional content. In , species such as Cryptotympana atrata are harvested and prepared in traditional dishes, including or stir-frying, providing a high-protein source with essential and fatty acids that support effects. These are eaten at different life stages, from protein-rich nymphs to adults, and recent studies highlight their sustainability as an alternative protein amid global challenges, noting low environmental impact compared to conventional . In , cicada , known as chan tui (the dried outer shell of Cryptotympana pustulata nymphs), have been used for centuries to treat respiratory conditions like by dispersing wind-heat and relieving spasms. These are often prepared as decoctions or teas to soothe and improve breathing. Pharmacological research in 2023 has confirmed properties in cicada extracts, attributing them to bioactive compounds that modulate immune responses and reduce airway in models. Beyond consumption, cicada shells serve practical purposes in crafts and industry. , live cicadas are commonly used as , hooked through the to attract species like bass, , and during emergences, leveraging their natural movement on the surface. Emerging biotechnological research explores extraction from cicada sloughs for applications in sustainable materials, such as wound dressings and nanocomposites, due to the polymer's and thermal stability. While cicadas are generally safe for human consumption when properly prepared, individuals with shellfish allergies should avoid them owing to shared chitin proteins that can trigger reactions. Wild-harvested specimens pose risks from environmental contaminants like pesticides or pathogens, necessitating thorough cleaning and cooking to mitigate potential health hazards.

References

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