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The cicadas () are a superfamily, the Cicadoidea, of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s in the order
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to arou ...
(true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as
leafhopper A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and ...
s and
froghopper The froghoppers, or the superfamily Cercopoidea, are a group of hemipteran insects in the suborder Auchenorrhyncha. Adults are capable of jumping many times their height and length, giving the group their common name, but they are best known ...
s. 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 drumlike tymbals. The earliest known fossil Cicadomorpha appeared in the
Upper Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
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 Cryptic may refer to: In science: * Cryptic species complex, a group of species that are very difficult to distinguish from one another * Crypsis, the ability of animals to blend in to avoid observation * Cryptic era, earliest period of the Earth ...
. 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 Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
, 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. The annual cicadas are species that emerge every year. Though these cicadas' life cycles can vary from 1–9 or more years as underground nymphs, their emergence above ground as adults is not synchronized, so some members of each species appear every year. Cicadas have been featured in literature since the time of
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
's ''
Iliad The ''Iliad'' (; grc, Ἰλιάς, Iliás, ; "a poem about Ilium") is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Ody ...
'' and as motifs in art from the Chinese
Shang dynasty The Shang dynasty (), also known as the Yin dynasty (), was a Chinese royal dynasty founded by Tang of Shang (Cheng Tang) that ruled in the Yellow River valley in the second millennium BC, traditionally succeeding the Xia dynasty a ...
. They have also been used in myth and folklore as symbols of carefree living and immortality. The cicada is also mentioned in
Hesiod Hesiod (; grc-gre, Ἡσίοδος ''Hēsíodos'') was an ancient Greek poet generally thought to have been active between 750 and 650 BC, around the same time as Homer. He is generally regarded by western authors as 'the first written poet i ...
's ''
Shield A shield is a piece of personal armour held in the hand, which may or may not be strapped to the wrist or forearm. Shields are used to intercept specific attacks, whether from close-ranged weaponry or projectiles such as arrows, by means of ...
'' (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 China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
,
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John Wells explai ...
,
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federal constitutional monarchy consists of thirteen states and three federal territories, separated by the South China Sea into two regions: Peninsular Malaysia and Borneo's East Mal ...
, central
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, and
Pakistani Balochistan Balochistan (; bal, بلۏچستان; ) is one of the four provinces of Pakistan. Located in the southwestern region of the country, Balochistan is the largest province of Pakistan by land area but is the least populated one. It shares land ...
.


Etymology

The name is directly from the
onomatopoeic Onomatopoeia is the process of creating a word that phonetically imitates, resembles, or suggests the sound that it describes. Such a word itself is also called an onomatopoeia. Common onomatopoeias include animal noises such as ''oink'', ''m ...
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''cicada''.


Taxonomy and diversity

The superfamily Cicadoidea is a sister of the Cercopoidea (the froghoppers). Cicadas are arranged into two families: the Tettigarctidae and Cicadidae. The two
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
species of the Tettigarctidae include one in southern Australia and the other in
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
. The family Cicadidae is subdivided into the subfamilies
Cicadettinae Cicadettinae is a subfamily of cicadas in the family Cicadidae. About 230 genera and 1,200 described species are placed in the Cicadettinae. Tribes These 14 tribes belong to the subfamily Cicadettinae: * Aragualnini Sanborn, 2018 * Carinetin ...
, Cicadinae, Derotettiginae,
Tibicininae Tibicininae is a subfamily of cicadas in the family Cicadidae. There are 5 tribes and at least 140 described species in Tibicininae. They are found in the Neotropics, the Nearctic, and the Palearctic. Tibicininae underwent a major revision in ...
(or Tettigadinae), and
Tettigomyiinae Tettigomyiinae is a subfamily of cicadas in the family Cicadidae, found in tropical Africa. There are about 16 genera and at least 100 described species in Tettigomyiinae. The tribes Tettigomyiini and Ydiellini were recently transferred to this ...
they are found on all continents except Antarctica. Some previous works also included a family-level taxon called the
Tibiceninae ''Tibiceninae'' is a historical subfamily name from the insect family Cicadidae (the true cicadas). It was first used by Distant (1889). As of 2021, the name became unavailable to zoological nomenclature due to the suppression of its type genus '' ...
. The largest species is the Malaysian emperor cicada ''
Megapomponia imperatoria The empress cicada (''Megapomponia imperatoria'') is a species of cicada from Southeast Asia. It is the largest species of cicada with a head-body length of about and a wingspan of .Flindt, R. (2006). ''Amazing Numbers in Biology'', p. 10. Ref ...
''; its wingspan is up to about . Cicadas are also notable for the great length of time some species take to mature. At least 3000 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 Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
has been used to study the biogeography of complex island groups such as in Indonesia and Asia. There are several hundred described species in Australia and
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
, around 150 in South Africa, over 170 in America north of Mexico, at least 800 in Latin America, and over 200 in Southeast Asia and the Western Pacific. About 100 species occur in the Palaearctic. A few species are found in southern Europe, and a single species was known from England, the New Forest cicada, ''Cicadetta montana'', which also occurs in continental Europe. 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. Many of the North American species are the annual or jarfly or dog-day cicadas, members of the ''
Neotibicen Cicadas of the genus ''Neotibicen'' are large-bodied insects of the family Cicadidae that appear in summer or early fall in eastern North America. Common names include cicada, harvestfly, jar fly, and the misnomer locust. In 2015, these species ...
'', ''
Megatibicen ''Megatibicen'' is a genus of cicadas in the family Cicadidae, with about 10 described species. Until 2016, these species were included in the genus ''Tibicen'' (now genus ''Lyristes'' Horvath, 1926) and then briefly in ''Neotibicen''. The spec ...
'', or ''
Hadoa The genus ''Hadoa'' comprises large-bodied Cicadidae occurring in Western North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Ar ...
'' genera, so named because they emerge in late July and August. The best-known North American genus, however, may be ''
Magicicada The term periodical cicada is commonly used to refer to any of the seven species of the genus ''Magicicada'' of eastern North America, the 13- and 17-year cicadas. They are called periodical because nearly all individuals in a local population a ...
''. These periodical cicadas have an extremely long life cycle of 13 or 17 years, with adults suddenly and briefly emerging in large numbers. 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 ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
and Victoria, large cities including
Sydney Sydney ( ) is the capital city of the state of New South Wales, and the most populous city in both Australia and Oceania. Located on Australia's east coast, the metropolis surrounds Sydney Harbour and extends about towards the Blue Mounta ...
,
Melbourne Melbourne ( ; Boonwurrung/ Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a metro ...
, and
Brisbane Brisbane ( ) is the capital and most populous city of the Australian state of Queensland, and the third-most populous city in Australia and Oceania, with a population of approximately 2.6 million. Brisbane lies at the centre of the South ...
, 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. More than 40 species from five genera populate New Zealand, ranging from sea level to mountain tops, and all are
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to New Zealand and its surrounding islands ( Kermadec Islands,
Chatham Islands The Chatham Islands ( ) (Moriori: ''Rēkohu'', 'Misty Sun'; mi, Wharekauri) are an archipelago in the Pacific Ocean about east of New Zealand's South Island. They are administered as part of New Zealand. The archipelago consists of about t ...
). One species is found on Norfolk Island, which technically is part of Australia. The closest relatives of the NZ cicadas live in New Caledonia and Australia.


Palaeontology

Fossil Cicadomorpha first appeared in the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
. 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. This group, though, is currently thought to be more distantly related to Cicadomorpha than previously thought. The Palaeontinidae or "giant cicadas" come from the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
and
Lower Cretaceous Lower may refer to: * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also *Nizhny Nizhny (russian: Ни́жний; masculine), Nizhnyaya (; feminine), or Nizhneye (russian: Н� ...
of Eurasia and South America. 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''. Most fossil Cicadidae are known from the Cenozoic, and the oldest unambiguously identified specimen is ''Davispia bearcreekensis'' (subfamily Tibicininae) from 59–56 million years ago (Mya). One fossil genus and species (''Burmacicada protera'') based on a first-instar nymph has recently been reported from 98–99 Mya in the Late Cretaceous, although questions remain about its assignment to the Cicadidae.


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 In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
at the tip. 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. 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. The adult insect, known as an
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the ...
, is in total length in most species. The largest, the empress cicada (''Megapomponia imperatoria''), has a head-body length around , and its wingspan is . 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. 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. The thorax has three segments and houses the powerful wing muscles. They have two pairs of membranous wings that may be
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
, 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 The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. In males, the abdomen is largely hollow and used as a resonating chamber. 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.
Bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
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 A biomaterial is a substance that has been engineered to interact with biological systems for a medical purpose, either a therapeutic (treat, augment, repair, or replace a tissue function of the body) or a diagnostic one. As a science, biomateria ...
that can kill bacteria.


Temperature regulation

Desert cicadas such as '' Diceroprocta apache'' are unusual among insects in controlling their temperature by
evaporative cooling An evaporative cooler (also known as evaporative air conditioner, swamp cooler, swamp box, desert cooler and wet air cooler) is a device that cools air through the evaporation of water. Evaporative cooling differs from other air conditioning ...
, analogous to
sweating Perspiration, also known as sweating, is the production of fluids secreted by the sweat glands in the skin of mammals. Two types of sweat glands can be found in humans: eccrine glands and apocrine glands. The eccrine sweat glands are distri ...
in mammals. When their temperature rises above about , they suck excess sap from the food plants and extrude the excess water through pores in the
tergum A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or ' ...
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. Some non-desert cicada species such as ''Magicicada tredecem'' also cool themselves evaporatively, but less dramatically. Conversely, many other cicadas can voluntarily raise their body temperatures as much as above ambient temperature.


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 Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
abdominal The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
region. The structure is buckled by muscular action and, being made of
resilin Resilin is an elastomeric protein found in many insects and other arthropods. It provides soft rubber-elasticity to mechanically active organs and tissue; for example, it enables insects of many species to jump or pivot their wings efficiently. ...
, 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. The sounds may further be modulated by membranous coverings and by resonant cavities. 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 The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
serve as resonance chambers with which it amplifies the sound. The cicada also modulates the song by positioning its abdomen toward or away from the substrate. 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. The tettigarctid (or hairy) cicadas ''
Tettigarcta crinita ''Tettigarcta'' is a genus of cicadas in the family Tettigarctidae. There are two described species in ''Tettigarcta'', one found in mainland Australia and one on the island of Tasmania. These two species are the only living species of the family ...
'' of Australia and ''T. tomentosa'' have rudimentary tymbals in both sexes and do not produce airborne sounds. Both males and females produce vibrations that are transmitted through the tree substrate. They are considered as representing the original state from which other cicada communication has evolved. Average temperature of the natural habitat for the South American species ''Fidicina rana'' is about . During sound production, the temperature of the tymbal muscles was found to be significantly higher. Many cicadas sing most actively during the hottest hours of a summer day; roughly a 24-hour cycle. 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. ''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. Cicadas call from varying heights on trees. Where multiple species occur, the species may use different heights and timing of calling. 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 Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
. 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) Sound pressure or acoustic pressure is the local pressure deviation from the ambient (average or equilibrium) atmospheric pressure, caused by a sound wave. In air, sound pressure can be measured using a microphone, and in water with a hydrophone ...
which is among the loudest of all insect-produced sounds. The song is loud enough to cause permanent
hearing loss Hearing loss is a partial or total inability to hear. Hearing loss may be present at birth or acquired at any time afterwards. Hearing loss may occur in one or both ears. In children, hearing problems can affect the ability to acquire spoken l ...
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. 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. 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.


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. After mating, the female cuts slits into the bark of a twig where she deposits her eggs. 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 . Nymphs have strong front legs for digging and excavating chambers in close proximity 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 An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
, they construct an exit tunnel to the surface and emerge. 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. 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, or in some parts of the region, a 13-year life cycle. The long life cycles may have developed as a response to predators, such as the cicada killer wasp and praying mantis. A specialist predator with a shorter life cycle of at least two years could not reliably prey upon the cicadas. 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. File:Emerging cicada.jpg, A teneral cicada that has just emerged and is waiting to dry before flying away File:Cicada Close-Up.jpg, Newly emerged adult cicada held on human fingers File:Cicada skin.jpg, Cicada exuviae after the adult cicada has left File:Cicada on an eastern red cedar tree in Oklahoma.jpg, Cicada clinging to the bark of an eastern red cedar tree in
Oklahoma Oklahoma (; Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a state in the South Central region of the United States, bordered by Texas on the south and west, Kansas on the north, Missouri on the northeast, Arkansas on the east, New ...


Diet

Cicada nymphs drink sap from the xylem of various species of trees, including oak,
cypress Cypress is a common name for various coniferous trees or shrubs of northern temperate regions that belong to the family Cupressaceae. The word ''cypress'' is derived from Old French ''cipres'', which was imported from Latin ''cypressus'', the l ...
,
willow Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist so ...
,
ash Ash or ashes are the solid remnants of fires. Specifically, ''ash'' refers to all non-aqueous, non-gaseous residues that remain after something burns. In analytical chemistry, to analyse the mineral and metal content of chemical samples, ash ...
, and
maple ''Acer'' () is a genus of trees and shrubs commonly known as maples. The genus is placed in the family Sapindaceae.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 nd more or less continuously updated since h ...
. While common folklore indicates that adults do not eat, they actually do drink plant sap using their sucking mouthparts.


Locomotion

Cicadas, unlike other Auchenorrhyncha, are not adapted for jumping (saltation). 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.


Predators, parasites, and pathogens

Cicadas are commonly eaten by birds and mammals, as well as bats,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s, mantises,
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, and
robber flies The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking hypopharynx. The name "robber flies" reflects their notoriously aggressive pr ...
. 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. In northern Japan, brown bears prey on final instar nymphs of cicadas during summer by digging up the ground. 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. A
katydid Insects in the family Tettigoniidae are commonly called katydids (especially in North America), or bush crickets. They have previously been known as "long-horned grasshoppers". More than 8,000 species are known. Part of the suborder Ensifera, ...
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. Several fungal diseases infect and kill adult cicadas, while other
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
in the genera ''
Ophiocordyceps ''Ophiocordyceps'' is a genus of fungi within the family Ophiocordycipitaceae. The widespread genus, first described scientifically by British mycologist Tom Petch in 1931, contains about 140 species that grow on insects. Anamorphic genera that c ...
'' and '' Isaria'' attack nymphs. '' Massospora cicadina'' specifically attacks the adults of periodical cicadas, the spores remaining dormant in the soil between outbreaks. This fungus is also capable of dosing cicadas with psilocybin, the
psychedelic drug Psychedelics are a subclass of hallucinogenic drugs whose primary effect is to trigger non-ordinary states of consciousness (known as psychedelic experiences or "trips").Pollan, Michael (2018). ''How to Change Your Mind: What the New Science o ...
found in magic mushrooms, as well as
cathinone Cathinone (also known as benzoylethanamine, or β-keto-amphetamine) is a monoamine alkaloid found in the shrub ''Catha edulis'' (khat) and is chemically similar to ephedrine, cathine, methcathinone and other amphetamines. It is probably the mai ...
, an
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
similar to various
amphetamines Substituted amphetamines are a class of compounds based upon the amphetamine structure; it includes all derivative compounds which are formed by replacing, or substituting, one or more hydrogen atoms in the amphetamine core structure with sub ...
. 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. 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.


Antipredator adaptations

Cicadas use a variety of strategies to evade predators. Large cicadas can fly rapidly to escape if disturbed. Many are extremely well
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d 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; their partly transparent wings are held over the body and pressed close to the substrate. Some cicada species play dead when threatened. Some cicadas such as '' Hemisciera maculipennis'' display bright
deimatic Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
flash coloration on their hind wings when threatened; the sudden contrast helps to startle predators, giving the cicadas time to escape. Most cicadas are diurnal and rely on camouflage when at rest, but some species use
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
-related
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on ...
, wearing the bright colors that warn of toxicity in other animals; the Malaysian ''
Huechys sanguinea ''Huechys sanguinea'', commonly known as the black and scarlet cicada, is a species of cicada belonging to the family Cicadidae Cicadidae, the true cicadas, is the largest family of cicadas, with more than 3,200 species worldwide. The oldest k ...
'' has conspicuous red and black warning coloration, is diurnal, and boldly flies about in full view of possible predators. Predators such as the
sarcophagid Sarcophagidae () are a family of flies commonly known as flesh flies. They differ from most flies in that they are ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on carrion, dung, decaying material, or ...
fly '' Emblemasoma'' hunt cicadas by sound, being attracted to their songs. 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.


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). They are described by
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
in his ''
History of Animals ''History of Animals'' ( grc-gre, Τῶν περὶ τὰ ζῷα ἱστοριῶν, ''Ton peri ta zoia historion'', "Inquiries on Animals"; la, Historia Animalium, "History of Animals") is one of the major Aristotle's biology, texts on biolo ...
'' and by
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
in his '' Natural History''; their mechanism of sound production is mentioned by
Hesiod Hesiod (; grc-gre, Ἡσίοδος ''Hēsíodos'') was an ancient Greek poet generally thought to have been active between 750 and 650 BC, around the same time as Homer. He is generally regarded by western authors as 'the first written poet i ...
in his poem "
Works and Days ''Works and Days'' ( grc, Ἔργα καὶ Ἡμέραι, Érga kaì Hēmérai)The ''Works and Days'' is sometimes called by the Latin translation of the title, ''Opera et Dies''. Common abbreviations are ''WD'' and ''Op''. for ''Opera''. is a ...
": "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". In the classic 14th-century Chinese novel ''
Romance of the Three Kingdoms ''Romance of the Three Kingdoms'' () is a 14th-century historical novel attributed to Luo Guanzhong. It is set in the turbulent years towards the end of the Han dynasty and the Three Kingdoms period in Chinese history, starting in 184 AD ...
'',
Diaochan Diaochan was one of the Four Beauties of ancient China. Although based on a minor historical personage, she is mostly a fictional character. She is best known for her role in the 14th-century historical novel ''Romance of the Three Kingdoms'', ...
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 Manga ( Japanese: 漫画 ) are comics or graphic novels originating from Japan. Most manga conform to a style developed in Japan in the late 19th century, and the form has a long prehistory in earlier Japanese art. The term ''manga'' is ...
''
Winter Cicada is a Japanese anime OVA loosely based on Youka Nitta's manga series . The U.S. broadcast rights to the OVA have been licensed to Logo, MTV Networks' LGBT cable channel, and premiered on October 31, 2008. Synopsis This three part ''yaoi' ...
''. Cicadas are a frequent subject of ''
haiku is a type of short form poetry originally from Japan. Traditional Japanese haiku consist of three phrases that contain a '' kireji'', or "cutting word", 17 '' on'' (phonetic units similar to syllables) in a 5, 7, 5 pattern, and a '' kigo'', or ...
'', where, depending on type, they can indicate spring, summer, or autumn..
Shaun Tan Shaun Tan (born 1973) is an Australian artist, writer and film maker. He won an Academy Award for '' The Lost Thing'', a 2011 animated film adaptation of a 2000 picture book he wrote and illustrated. Other books he has written and illustrated inc ...
's illustrated book ''Cicada'' tells the story of a hardworking but underappreciated cicada working in an office. 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.


In fashion

Being lightweight and (on the legs) hooklike, the exuviae of cicadas can be used as hair or clothing accessories.


As food and folk medicine

Deep-fried ''Cryptotympana atrata'' in Shandong cuisine Cicadas were eaten in
Ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cu ...
, and are consumed today in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, both as adults and (more often) as nymphs. Cicadas are also eaten in
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federal constitutional monarchy consists of thirteen states and three federal territories, separated by the South China Sea into two regions: Peninsular Malaysia and Borneo's East Mal ...
,
Burma Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, North America, and central Africa, as well as the
Balochistan Balochistan ( ; bal, بلۏچستان; also romanised as Baluchistan and Baluchestan) is a historical region in Western and South Asia, located in the Iranian plateau's far southeast and bordering the Indian Plate and the Arabian Sea coastline. ...
region of
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
, especially in Ziarat. Female cicadas are prized for being meatier. Shells of cicadas are employed in
traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...
s. The 17-year "Onondaga Brood" ''Magicicada'' is culturally important and a particular delicacy to the
Onondaga people The Onondaga people ( Onondaga: , ''Hill Place people'') are one of the original five constituent nations of the Iroquois (''Haudenosaunee'') Confederacy in northeast North America. Their traditional homeland is in and around present-day Onondag ...
, and are considered a novelty food item by modern consumers in several states.


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 Haydée Mercedes Sosa (; 9 July 1935
at BrainyHistory.com
– 4 October 2009), sometimes known as ' ...
, among other Latin American musicians. In North America and Mexico, there is a well-known song, "" ("The Cicada"), written by Raymundo Perez Soto, which is a song in the
Mariachi Mariachi (, , ) is a genre of regional Mexican music that dates back to at least the 18th century, evolving over time in the countryside of various regions of western Mexico. The usual mariachi group today consists of as many as eight violins, t ...
tradition, that romanticises the insect as a creature that sings until it dies. 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. Cicada sounds heavily feature on the 2021 album ''
Solar Power Solar power is the conversion of energy from sunlight into electricity, either directly using photovoltaics (PV) or indirectly using concentrated solar power. Photovoltaic cells convert light into an electric current using the photovolta ...
'' by New Zealand artist
Lorde Ella Marija Lani Yelich-O'Connor (born 7 November 1996), known professionally as Lorde ( ), is a New Zealand singer-songwriter. Taking inspiration from aristocracy for her stage name, she is known for her unconventional musical styles and i ...
. She described cicada song as being emblematic of the New Zealand summer.


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. In France, the cicada represents the
folklore Folklore is shared by a particular group of people; it encompasses the traditions common to that culture, subculture or group. This includes oral traditions such as tales, legends, proverbs and jokes. They include material culture, rangin ...
of
Provence Provence (, , , , ; oc, Provença or ''Prouvènço'' , ) is a geographical region and historical province of southeastern France, which extends from the left bank of the lower Rhône to the west to the Italian border to the east; it is bo ...
and the Mediterranean cities. The cicada has represented since
classical antiquity Classical antiquity (also the classical era, classical period or classical age) is the period of cultural history between the 8th century BC and the 5th century AD centred on the Mediterranean Sea, comprising the interlocking civilizations of ...
.
Jean de La Fontaine Jean de La Fontaine (, , ; 8 July 162113 April 1695) was a French fabulist and one of the most widely read French poets of the 17th century. He is known above all for his ''Fables'', which provided a model for subsequent fabulists across Euro ...
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 finds herself without food when the weather turns bitter. In Chinese tradition, the cicada (, ''chán'') symbolises rebirth and immortality. In the Chinese essay " Thirty-Six Stratagems", the phrase "to shed the golden cicada skin" () is the poetic name for using a decoy (leaving the exuviae) to fool enemies. In the Chinese classic novel '' Journey to the West'' (16th century), the protagonist Priest of Tang was named the Golden Cicada. In Japan, the cicada is associated with the summer season. For many Japanese people, summer hasn't officially begun until the first songs of the cicada are heard. According to
Lafcadio Hearn , born Patrick Lafcadio Hearn (; el, Πατρίκιος Λευκάδιος Χέρν, Patríkios Lefkádios Chérn, Irish: Pádraig Lafcadio O'hEarain), was an Irish- Greek- Japanese writer, translator, and teacher who introduced the culture a ...
, 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. In the Homeric ''Hymn to Aphrodite'', the goddess
Aphrodite Aphrodite ( ; grc-gre, Ἀφροδίτη, Aphrodítē; , , ) is an ancient Greek goddess associated with love, lust, beauty, pleasure, passion, and procreation. She was syncretized with the Roman goddess . Aphrodite's major symbols incl ...
retells the legend of how Eos, the goddess of the dawn, requested
Zeus Zeus or , , ; grc, Δῐός, ''Diós'', label= genitive Boeotian Aeolic and Laconian grc-dor, Δεύς, Deús ; grc, Δέος, ''Déos'', label= genitive el, Δίας, ''Días'' () is the sky and thunder god in ancient Greek relig ...
to let her lover
Tithonus In Greek mythology, Tithonus ( or ; grc, Τιθωνός, Tithonos) was the lover of Eos, Goddess of the Dawn. He was a prince of Troy, the son of King Laomedon by the Naiad Strymo (Στρυμώ). The mythology reflected by the fifth-century vas ...
live forever as an immortal. Zeus granted her request, but because Eos forgot to ask him to also make Tithonus ageless, Tithonus never died, but he did grow old. Eventually, he became so tiny and shriveled that he turned into the first cicada. The Greeks also used a cicada sitting on a harp as an emblem of music. In Kapampangan mythology in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
, the goddess of dusk, Sisilim, is said to be greeted by the sounds and appearances of cicadas whenever she appears.


As pests

Cicadas feed on sap; they do not bite or
sting Sting may refer to: * Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger * Irritating hairs or prickles of a stinging plant, or the plant itself Fictional characters and entities * Sting (Middle-earth ...
in a true sense, but may occasionally mistake a person's arm for a plant limb and attempt to feed. Male cicadas produce very loud calls that can damage human hearing. 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. 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. Some species have turned from wild grasses to
sugarcane Sugarcane or sugar cane is a species of (often hybrid) tall, perennial grass (in the genus '' Saccharum'', tribe Andropogoneae) that is used for sugar production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fibrous stalk ...
, which affects the crop adversely, and in a few isolated cases, females have oviposited on
cash crop A cash crop or profit crop is an agricultural crop which is grown to sell for profit. It is typically purchased by parties separate from a farm. The term is used to differentiate marketed crops from staple crop (or "subsistence crop") in subsist ...
s such as
date palm ''Phoenix dactylifera'', commonly known as date or date palm, is a flowering plant species in the palm family, Arecaceae, cultivated for its edible sweet fruit called dates. The species is widely cultivated across northern Africa, the Middle Eas ...
s,
grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus '' Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began perhaps 8,000 years a ...
vines,
citrus ''Citrus'' is a genus of flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, lemons, grapefruits, pomelos, and limes. The genus ''Citrus'' is native to ...
trees, asparagus, and
cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor pe ...
. 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..


See also

* Cicada 3301 * List of Cicadidae genera


Notes


References


Further reading

* * * * * * * *


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

* ttps://entnemdept.ufl.edu/creatures/misc/bugs/cicadas.htm Cicadas of Florida, ''Neocicada hieroglyphica'', ''Tibicen'', ''Diceroprocta'' and ''Melampsalta'' spp.at
University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...
/ IFAS ''Featured Creatures''
Greater Cincinnati Cicada Information & Teaching Resources
– College of Mt Saint Joseph Cicada Information Site

{{Taxonbar, from=Q1947892 Cicadomorpha Edible insects Insects in culture Extant Late Triassic first appearances