Cicada
Cicada Temporal range:
| |
---|---|
Annual cicada, Neotibicen linnei | |
Calling song of Magicicada cassini | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hemiptera |
Suborder: | Auchenorrhyncha |
Infraorder: | Cicadomorpha |
Superfamily: | Cicadoidea Latreille, 1802 |
Families | |
The cicadas (/sɪˈkɑːdəz, -ˈkeɪ-/) 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.
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
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, and central Africa.[citation needed]
Etymology
The name is directly from the onomatopoeic Latin cicada.[4][5][b]
Taxonomy and diversity
The superfamily Cicadoidea is a sister of the
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.[10] There are several hundred described species in Australia and New Zealand,[c] around 150 in South Africa, over 170 in America north of Mexico,[11] at least 800 in Latin America,[12] and over 200 in Southeast Asia and the Western Pacific.[13]
About 100 species occur in the Palaearctic. A few species are found in southern Europe,[9] and a single species was known from England, the New Forest cicada, Cicadetta montana, which also occurs in continental Europe.[14] 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.[7] |
Many of the North American species are the annual or jarfly or
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.[18] The closest relatives of the NZ cicadas live in New Caledonia and Australia.
Palaeontology
Fossil Cicadomorpha first appeared in the
The
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. 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.[22] Most fossil Cicadidae are known from the Cenozoic,[23] and the oldest unambiguously identified modern cicadid is Davispia bearcreekensis (subfamily Tibicininae) from the Paleocene, around 59–56 million years ago (Mya).[24]
Biology
Description
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.[25] 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.[10]
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.[10]
The adult insect, known as an
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.[27]
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, they leap several millimetres into the air, which also serves to clean the wings.[29] 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.[30]
Temperature regulation
Desert cicadas such as
Song
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.[35] The sounds may further be modulated by membranous coverings and by resonant cavities.[25]
The male abdomen in some species is largely hollow, and acts as a
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.[37] Many cicadas sing most actively during the hottest hours of a summer day; roughly a 24-hour cycle.[38] 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.[34] 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.[39][40]
Cicadas call from varying heights on trees. Where multiple species occur, the species may use different heights and timing of calling.[41][42] 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.[43]
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;[44] a necessity partly because some cicadas produce sounds up to 120 dB (SPL)[44] which is among the loudest of all insect-produced sounds.[45] 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.[46]
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.[47]
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.[48]
Life cycle
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.[10] After mating, the female cuts slits into the bark of a twig where she deposits her eggs.[10] 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 down to about 2.5 m (8 ft). 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.[10] They then moult (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.[49]
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,
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A teneral cicada that has just emerged and is waiting to dry before flying away
-
Newly emerged adult cicada held on human fingers
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Cicada exuviae after the adult cicada has left
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Cicada clinging to the bark of aneastern red cedar tree in Oklahoma
Diet
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.[57][58] Cicadas excrete fluid in streams of droplets due to their high volume consumption of xylem sap.[59]
Locomotion
Cicadas, unlike other Auchenorrhyncha, are not adapted for jumping (saltation).[60] 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.[10]
Predators, parasites, and pathogens
Cicadas are commonly eaten by birds and mammals,
Several fungal diseases infect and kill adult cicadas, while other
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.[67][68]
Antipredator adaptations
Cicadas use a variety of strategies to evade predators. Large cicadas can fly rapidly to escape if disturbed.
Some cicadas such as
Predators such as the
In human culture
In art and literature
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".[79] 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.[80] Shaun Tan's illustrated book Cicada tells the story of a hardworking but underappreciated cicada working in an office.[81] Branden Jacobs-Jenkins' play Appropriate takes place on an Arkansas farm in summer, and calls for the sounds of mating cicadas to underscore the entire show.[82]
In fashion
Being lightweight, and with hooklike legs, the exuviae of cicadas can be used as hair or clothing accessories.[83]
As food and folk medicine
Cicadas were eaten in
In music
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.[90]
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.[91]
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.[92]
In mythology and folklore
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.[93] In France, the cicada represents the folklore of Provence and the Mediterranean cities.[94]
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.[95]
In Chinese tradition, the cicada (蟬, chán) symbolises rebirth and immortality.[96] 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.[97] In the Chinese classic novel Journey to the West (16th century), the protagonist Priest of Tang was named the Golden Cicada.[98]
In Japan, the cicada is associated with the summer season.[99] For many Japanese people, summer hasn't officially begun until the first songs of the cicada are heard.[100] 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.[101]
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.[102] Zeus granted her request, but because Eos forgot to ask him to also make Tithonus ageless, Tithonus never died, but he did grow old.[102] Eventually, he became so tiny and shriveled that he turned into the first cicada.[102] The Greeks also used a cicada sitting on a harp as an emblem of music.[103]
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.[104]
As pests
Cicadas feed on
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.[27] 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.[107] 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.[27]
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.[108][109][failed verification]
See also
Notes
References
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Further reading
- Clausen, Lucy W. (1954). Insect Fact and Folklore. Macmillan. [ISBN missing]
- Egan, Rory B. (1994). "Cicada in Ancient Greece". Archived from the original on 10 November 2006. Retrieved 28 December 2006.
- Hoppensteadt, Frank C; Keller, Joseph B (1976). "Synchronization of periodical cicada emergences" (PDF). PMID 987617. Archived from the original(PDF) on 9 November 2016. Retrieved 3 August 2011.
- Main, Douglas (3 May 2021). "Trillions of cicadas are coming to the U.S. Here's why that's a good thing". National Geographic. Archived from the original on 3 May 2021.
- Myers, JG (1929). Insect Singers: A Natural History of the Cicadas. Routledge.[ISBN missing]
- Ramel, Gordon (2005). "The Singing Cicadas". Earth Life. Retrieved 31 January 2007.
- Riegel, Garland (1994). Cicada in Chinese Folklore. Melsheimer Entomological Series. Bug bios. Archived from the original on 10 November 2006. Retrieved 28 December 2006.
- Walker, Annette (2000). The Reed Handbook of Common New Zealand Insects. Reed. ISBN 978-0-7900-0718-2.
External links
- Cicada Mania – Website dedicated to cicadas, the most amazing insects in the world
- Massachusetts Cicadas describes behavior, sightings, photos, how to find guide, videos, periodical and annual cicada species information and distribution maps
- Cicadas.uconn.edu/ Brood mapping project – solicits records and observations from the general public
- Song recordings and information of cicadas of the United States and Canada
- Cicadas of Florida, Neocicada hieroglyphica, Tibicen, Diceroprocta and Melampsalta spp. at University of Florida/IFAS Featured Creatures
- Greater Cincinnati Cicada Information & Teaching Resources Archived 16 April 2013 at the Wayback Machine – College of Mt Saint Joseph Cicada Information Site
- DrMetcalf: a resource on cicadas, leafhoppers, planthoppers, spittlebugs, and treehoppers