| Cicada | |
|---|---|
 | |
| Linne's cicada,Neotibicen linnei | |
| Calling song ofMagicicada cassini | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Hemiptera |
| Suborder: | Auchenorrhyncha |
| Infraorder: | Cicadomorpha |
| Superfamily: | Cicadoidea Latreille, 1802 |
| Families | |
Thecicadas (/sɪˈkɑːdəz,-ˈkeɪ-/) are a superfamily, theCicadoidea, ofinsects in the orderHemiptera (true bugs). They are in the suborderAuchenorrhyncha,[a] along with smaller jumping bugs such asleafhoppers andfroghoppers. The superfamily is divided into two families, theTettigarctidae, with two species in Australia, and theCicadidae, with more than 3,000 speciesdescribed from around the world; many species remainundescribed. Nearly all cicada species areannual cicadas with the exception of the few North Americanperiodical 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-liketymbals. The earliest known fossil Cicadomorpha appeared in theUpper Permian period; extant species occur all around the world in temperate to tropical climates. They typically live in trees, feeding on watery sap fromxylem tissue, and laying their eggs in a slit in the bark. Most cicadas arecryptic. 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 (theperiodical 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 timing and synchronization of their emergence may reduce cicada losses topredation by making them less reliable prey and by overwhelming predators with sheer numbers before significant losses occur.[1]
Theannual cicadas are species that emerge every year. Though these cicadas' life cycles can vary from one to nine 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 ofHomer'sIliad and as motifs in art from the ChineseShang dynasty.[3] They have also been used in myth and folklore as symbols of carefree living and immortality. The cicada is also mentioned inHesiod'sShield (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, includingChina,Myanmar,Malaysia, centralAfrica and parts ofMexico.[4]
The name is directly from theonomatopoeicLatincicada.[5][6][b]
The superfamily Cicadoidea is a sister of theCercopoidea (the froghoppers). Cicadas are arranged into two families: the Tettigarctidae and Cicadidae. The twoextant species of the Tettigarctidae include one in southern Australia and the other inTasmania. The family Cicadidae is subdivided into the subfamiliesCicadettinae,Cicadinae,Derotettiginae,Tibicininae (or Tettigadinae), andTettigomyiinae,[8] and they are found on all continents except Antarctica. Some previous works also included a family-level taxon called theTibiceninae. The largest species is the Malaysian emperor cicadaMegapomponia 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 ofendemism 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 andNew 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 thePalaearctic. A few species are found in southern Europe,[10] and a single species was known from England, theNew 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 ordog-day cicadas, members of theNeotibicen,Megatibicen, orHadoa genera, so named because they emerge in late July and August.[16] The best-known North American genus, however, may beMagicicada. These periodical cicadas have an extremely long life cycle of 13 or 17 years, with adults suddenly and brieflyemerging 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 ofNew South Wales andVictoria, large cities includingSydney,Melbourne, andBrisbane, 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]
More than 40 species from five genera populate New Zealand, ranging from sea level to mountain tops, and all areendemic 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.
Fossil Cicadomorpha first appeared in theLate Triassic. The superfamilyPalaeontinoidea contains three families. The Upper Permian Dunstaniidae are found in Australia and South Africa, and also in younger rocks from China. TheUpper 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]
ThePalaeontinidae or "giant cicadas" (though only distantly related to true cicadas) come from theJurassic andLower Cretaceous of Eurasia and South America.[20] The first of these was a fore wing discovered in theTaynton Limestone Formation of Oxfordshire, England; it was initially described as a butterfly in 1873, before being recognised as a cicada-like form and renamedPalaeontina oolitica.[22]
Tettigarctidae and Cicadidae had diverged from each other prior to or during theJurassic, 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 CretaceousBurmese 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 isDavispia bearcreekensis (subfamily Tibicininae) from thePaleocene, around 56–59 million years ago.[25]
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Cicadas are large insects made conspicuous by the courtship calls of the males. They are characterized by having three joints in theirtarsi, and having smallantennae with conical bases and three to six segments, including aseta at the tip.[26] TheAuchenorrhyncha differ from other hemipterans by having arostrum 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 theprothorax extending as far as thescutellum, and by lacking the tympanal apparatus.[11]
The adult insect, known as animago, is 2 to 5 cm (1 to 2 in) in total length in most species. The largest, theempress 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 smallocelli 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, sharprostrum that they insert into the plant to feed.[28] Thepostclypeus 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 behyaline, cloudy, or pigmented. The wing venation varies between species and may help in identification. The middle thoracic segment has anoperculum 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-edgedovipositor. In males, the abdomen is largely hollow and used as aresonating chamber.[28]
The surface of the fore wing issuperhydrophobic; 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-knownbiomaterial that can kill bacteria.[31]
Desert cicadas such asDiceroprocta apache are unusual among insects in controlling their temperature byevaporative cooling, analogous tosweating 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 thetergum at a modest cost in energy. Such a rapid loss of water can be sustained only by feeding on water-richxylem 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 asMagicicada 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]
The "singing" of male cicadas is produced principally and in the majority of species using a special structure called atymbal, a pair of which lies below each side of theanteriorabdominal region. The structure is buckled by muscular action and, being made ofresilin, unbuckles rapidly on muscle relaxation, producing their characteristic sounds. Some cicadas, however, have mechanisms forstridulation, sometimes in addition to the tymbals. Here, the wings are rubbed over a series of midthoracic ridges. In the Chinese speciesSubpsaltria 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 asound box. By rapidly vibrating these membranes, a cicada combines the clicks into apparently continuous notes, and enlarged chambers derived from thetracheae serve asresonating chambers with which it amplifies the sound. The cicada also modulates the song by positioning its abdomen toward or away from thesubstrate (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] Thetettigarctid (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 theoriginal state from which other cicada communication has evolved.[37]
Average temperature of the natural habitat for the South American speciesFidicina 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 a24-hour cycle.[39] Most cicadas are diurnal in their calling anddepend 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 andFroggattoides 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 andO. 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 theburrow structure, but this may avoidpredation.[44]
Although only males produce the cicadas' distinctive sounds, both sexes have membranous structures calledtympana (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 permanenthearing 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]
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 thesubstrate 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 hatchednymphs 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 onxylem 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 nymphalinstar, they construct an exit tunnel to the surface and emerge.[11] They thenmolt (shed their skins) on a nearby plant for the last time, and emerge as adults. Theexuviae 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 topredators, such as thecicada killer wasp andpraying 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 matchingprime 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]
Cicada nymphs drink sap from the xylem of various species of trees, includingoak,cypress,willow,ash, andmaple. 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]
Cicadas, unlike other Auchenorrhyncha, are not adapted for jumping (saltation).[63] They have the usual insect modes oflocomotion, 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]
_with_Cicada.jpg)
Cicadas are commonly eaten by birds and mammals,[64] as well asbats,wasps,mantises,spiders, androbber 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] Akatydid 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 otherfungi in the generaOphiocordyceps andIsaria 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 withpsilocybin, thepsychedelic drug found inmagic mushrooms, as well ascathinone, analkaloid similar to variousamphetamines. 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 ofMagicicada to hatch, the resin sealing up the egg cavities.[70][71]
Cicadas use a variety of strategies to evade predators. Large cicadas can fly rapidly to escape if disturbed.[72] Many are extremely wellcamouflaged[72][73] to evade predators such as birds that hunt by sight. Being coloured like tree bark anddisruptively 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 speciesplay dead when threatened.[75][76]
Some cicadas such asHemisciera maculipennis display brightdeimatic flash coloration on their hind wings when threatened; the sudden contrast helps to startle predators, giving the cicadas time to escape.[77] Most cicadas arediurnal and rely on camouflage when at rest, but some species useaposematism-relatedBatesian mimicry, wearing the bright colors that warn of toxicity in other animals; the MalaysianHuechys sanguinea has conspicuous red and black warning coloration, is diurnal, and boldly flies about in full view of possible predators.[78]
Predators such as thesarcophagid flyEmblemasoma 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]
.jpg)
Cicadas have been featured in literature since the time of Homer'sIliad, and as motifs in decorative art from the Chinese Shang dynasty (1766–1122 BCE).[d] They are described byAristotle in hisHistory of Animals and byPliny the Elder in hisNatural History; their mechanism of sound production is mentioned byHesiod in his poem "Works and Days": "when theSkolymus flowers, and the tunefulTettix 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 novelRomance 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 novelThe 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 themangaWinter Cicada. Cicadas are a frequent subject ofhaiku, where, depending on type, they can indicate spring, summer, or autumn.[83]Shaun Tan's illustrated bookCicada tells the story of a hardworking but underappreciated cicada working in an office.[84]Branden Jacobs-Jenkins's playAppropriate takes place on an Arkansas farm in summer, and calls for the sounds of mating cicadas to underscore the entire show.[85]
Being lightweight, and with hooklike legs, theexuviae of cicadas can be used as hair or clothing accessories.[86]
Cicadas were eaten inAncient Greece, and are consumed in selected regions in modernChina, both as adults and (more often) as nymphs.[87] Cicadas are also eaten inMalaysia,Burma, North America, and central Africa, as well as theBalochistan region ofPakistan, especially inZiarat.[88] Female cicadas are prized for being meatier.[47] Shells of cicadas are employed intraditional 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 theOnondaga people,[92] and are considered a novelty food item by modern consumers in several states.[93]
Cicadas are featured in the protest song "Como La Cigarra" ("Like the Cicada") written by Argentinian poet and composerMaría Elena Walsh. In the song, the cicada is a symbol of survival and defiance against death. The song was recorded byMercedes Sosa, among other Latin American musicians.
In North America and Mexico, there is a well-known song, "La Cigarra" ("The Cicada"), written byRaymundo Perez Soto, which is a song in theMariachi tradition, that romanticises the insect as a creature that sings until it dies.[94]
Brazilian artistLenine with his track "Malvadeza" from the albumChão, 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 albumSolar Power by New Zealand artistLorde. She described cicada song as being emblematic of the New Zealand summer.[96]
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 thefolklore ofProvence and the Mediterranean cities.[98]
The cicada has representedinsouciance sinceclassical antiquity.Jean de La Fontaine began his collection of fablesLes fables de La Fontaine with the story "La Cigale et la Fourmi" ("The Cicada and the Ant") based on one ofAesop'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 (蟬,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 novelJourney 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 toLafcadio Hearn, the song ofMeimuna opalifera, calledtsuku-tsuku boshi, is said to indicate the end of summer, and it is called so because of its particular call.[105]
In theHomericHymn to Aphrodite, the goddessAphrodite retells the legend of howEos, the goddess of the dawn, requestedZeus to let her loverTithonus live forever as animmortal.[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 thePhilippines, the goddess of dusk, Sisilim, is said to be greeted by the sounds and appearances of cicadas whenever she appears.[108]
Cicadas feed onsap; they do notbite orsting 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 tosugarcane, which affects the crop adversely, and in a few isolated cases, females have oviposited oncash crops such asdate palms,grape vines,citrus trees,asparagus, andcotton.[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]
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