Magicicada
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The term periodical cicada is commonly used to refer to any of the seven species of the genus ''Magicicada'' of eastern
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 Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, the 13- and 17-year cicadas. They are called periodical because nearly all individuals in a local population are developmentally synchronized and emerge in the same year. Although they are sometimes called "
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
s", this is a misnomer, as
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s belong to the taxonomic order
Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
(true bugs), suborder
Auchenorrhyncha The Auchenorrhyncha suborder of the Hemiptera contains most of the familiar members of what was called the "Homoptera" – groups such as cicadas, leafhoppers, treehoppers, planthoppers, and spittlebugs. The aphids and scale insects are the ot ...
, while locusts are grasshoppers belonging to the order
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassho ...
. ''Magicicada'' belongs to the cicada tribe
Lamotialnini Lamotialnini is a tribe of cicadas in the family Cicadidae. There are about 19 genera and at least 90 described species in Lamotialnini, occurring worldwide except South America. Genera These 19 genera belong to the tribe Lamotialnini: * '' Ab ...
, a group of
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
with representatives in Australia, Africa, and Asia, as well as the Americas. ''Magicicada'' species spend around 99.5% of their long lives underground in an immature state called a
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
. While underground the nymphs feed on
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
fluids from the roots of
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
forest trees in the eastern United States. In the spring of their 13th or 17th year mature cicada nymphs emerge between late April and early June at a given locality, synchronously and in tremendous numbers. The adults are active for only about four to six weeks after the unusually prolonged developmental phase. The males aggregate in chorus centers and call there to attract mates. Mated females lay eggs in the stems of woody plants. Within two months of the original emergence, the life cycle is complete and the adult cicadas die. Later in that same summer the eggs hatch and the new nymphs burrow underground to develop for the next 13 or 17 years. Periodical emergences are also reported for the "World Cup cicada" ''Chremistica ribhoi'' (every four years) in northeast India and for a cicada species from Fiji, ''Raiateana knowlesi'' (every eight years).


Description

The winged
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 f ...
(adult) periodical cicada has red eyes and a black dorsal thorax. The wings are translucent with orange veins. The underside of the abdomen may be black, orange, or striped with orange and black, depending on the species. Adults are typically , depending on species, generally about 75% the size of most of the annual cicada species found in the same region. Mature females are slightly larger than males. ''Magicicada'' males typically form large aggregations that sing in chorus to attract receptive females. Different species have different characteristic calling songs. The call of
decim periodical cicadas Decim periodical cicadas is a term used to group three closely related species of periodical cicadas: ''Magicicada septendecim'', '' Magicicada tredecim'', and '' Magicicada neotredecim''. ''M. septendecim'', first described by Carl Linnaeus, has ...
is said to resemble someone calling "weeeee-whoa" or "Pharaoh". The cassini and decula periodical cicadas (including ''M. tredecula'') have songs that intersperse buzzing and ticking sounds. Cicadas cannot sting and do not normally bite. Like other
Auchenorrhyncha The Auchenorrhyncha suborder of the Hemiptera contains most of the familiar members of what was called the "Homoptera" – groups such as cicadas, leafhoppers, treehoppers, planthoppers, and spittlebugs. The aphids and scale insects are the ot ...
(true) bugs, they have mouthparts used to pierce plants and suck their sap. These mouthparts are used during the nymph stage to tap underground roots for water, minerals and carbohydrates and in the adult stage to acquire nutrients and water from plant stems. An adult cicada's proboscis can pierce human skin when it is handled, which is painful, but in no other way harmful. Cicadas are neither
venomous Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
nor
poisonous Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
and there is no evidence that they or their bites can transmit diseases. Cicadas pose little threat to mature vegetation, although planting new trees or shrubs is best postponed until after an expected emergence of the periodical cicadas. Mature plants rarely suffer lasting damage, although twig die-off or flagging may result from egg-laying. Young trees or shrubs can be covered with
cheesecloth Cheesecloth is a loose-woven gauze-like carded cotton cloth used primarily in cheesemaking and cooking. Grades Cheesecloth is available in at least seven different grades, from open to extra-fine weave. Grades are distinguished by the numbe ...
or other mesh netting with holes that are in diameter or smaller to prevent damage during the oviposition period, which begins about a week after the first adults emerge and lasts until all females have died. Massospora cicadina is a fungal pathogen that infects only 13 and 17 year periodical cicadas. Infection results in a "plug" of spores that replaces the end of the cicada's abdomen while it is still alive, leading to infertility, disease transmission, and eventual death of the cicada.


Life cycle

Nearly all cicadas spend years underground as juveniles, before emerging above ground for a short adult stage of several weeks to a few months. The seven periodical cicada species are so named because, in any one location, all members of the population are developmentally synchronized—they emerge as adults all at once in the same year. This periodicity is especially remarkable because their life cycles are so long—13 or 17 years. No other species of cicada in the world (among perhaps 3,000 species) synchronize their development in this way. In contrast, for nonperiodical species, some adults mature each summer and emerge while the rest of the population continues to develop underground. Many people refer to these nonperiodical species as
annual cicada Annual cicadas are North American Cicadidae species that appear every summer. The life cycle of a so-called annual cicada typically spans 2 to 5 years; they are "annual" only in the sense that members of the species reappear annually. The name is u ...
s because some are seen every summer. This may lead some to conclude that the non-periodic cicadas have life cycles of one year. This is incorrect. The few known life cycles of "annual" species range from two to 10 years, although some could be longer. The nymphs of the periodical cicadas live underground, usually within of the surface, feeding on the juices of plant roots. The nymphs of the periodical cicada undergo five
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 ass ...
stages in their development underground. The difference in the 13- and 17-year life cycle is said to be the time needed for the second instar to mature. When underground the nymphs move deeper below ground, detecting and then feeding on larger roots as they mature. In late April to early June of the emergence year, mature fifth-instar nymphs construct tunnels to the surface and wait for the soil temperature to reach a critical value. In some situations, nymphs extend mud turrets up to several inches above the soil surface. The function of these turrets is not known, but the phenomenon has been observed in some nonperiodical cicadas, as well as other tunneling insects. The nymphs first emerge on a spring evening when the soil temperature at around of depth is above . The
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
emergence is thought to be related to the fact that maximum soil temperatures lag behind maximum
insolation Solar irradiance is the power per unit area (surface power density) received from the Sun in the form of electromagnetic radiation in the wavelength range of the measuring instrument. Solar irradiance is measured in watts per square metre (W/m ...
by several hours, conveniently providing some protection for the flightless nymphs against diurnal sight predators such as birds. For the rest of their lives the mature periodical cicadas will be strongly diurnal, with song often nearly ceasing at night. During most years in the United States this emergence cue translates to late April or early May in the far south, and late May to early June in the far north. Emerging nymphs may
molt In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
in the grass or climb from a few centimeters to more than 100 feet (30 m) to find a suitable vertical surface to complete their transformation into adults. After securing themselves to tree trunks, the walls of buildings, telephone poles, fenceposts, hanging foliage, and even stationary automobile tires, the nymphs undergo a final molt and then spend about six days in the trees to await the complete hardening of their wings and
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
s. Just after emerging from this final molt the
teneral Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
adults are off-white, but darken within an hour. Adult periodical cicadas live for only a few weeks; by mid-July, all have died. Their ephemeral adult forms are adapted for one purpose: reproduction. Like other cicadas the males produce a very loud species-specific mating song using their tymbals. Singing males of the same ''Magicicada'' species tend to form aggregations called ''choruses'' whose collective songs are attractive to females. Males in these choruses alternate bouts of singing with short flights from tree to tree in search of receptive females. Most matings occur in so-called chorus trees. Receptive females respond to the calls of conspecific males with timed wing-flicks (visual signaling is apparently a necessity in the midst of the males' song) which attract the males for mating. The sound of a chorus can be literally deafening and depending on the number of males composing it, may reach 100  dB in the immediate vicinity. In addition to their "calling" or "congregating" songs, males produce a distinctive courtship song when approaching an individual female. Both males and females can mate multiple times, although most females seem to mate only once. After mating, the female cuts V-shaped slits in the bark of young twigs and lays about 20 eggs in each, for a total clutch of 600 or more. After about 6–10 weeks, the eggs hatch and the nymphs drop to the ground, where they burrow and begin another 13- or 17-year cycle. File:Magicicada emergence holes.jpg, ''Magicicada'' nymph emergence holes File:Brood X emergence turrets.jpg, Mud turrets that emergent
Brood X Brood X (Brood 10), the Great Eastern Brood, is one of 15 broods of periodical cicadas that appear regularly throughout the eastern United States. The brood's first major emergence after 2021 is predicted to occur during 2038. Characteristics ...
''Magicicada'' nymphs created in
Potomac, Maryland Potomac () is a census-designated place (CDP) in Montgomery County, Maryland, United States, named after the nearby Potomac River. Potomac is the seventh most educated small town in America, based on percentage of residents with postsecondary deg ...
near Washington, D.C. (June 30, 2021) File:Premolt Brood XIII.JPG, Brood XIII ''Magicicada'' nymph prior to final molt in suburban Chicago (May 24, 2007) File:MoltingMagicicadaCrop.jpg, ''Magicicada'' molting File:White cicada.jpg,
Teneral Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
adult An adult is a human or other animal that has reached full growth. In human context, the term ''adult'' has meanings associated with social and legal concepts. In contrast to a " minor", a legal adult is a person who has attained the age of major ...
Brood XIII ''Magicicada'' and
exuviae In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have moulted. The exuviae of an animal can be important to biologists as they can often b ...
after
molting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
in
Highland Park, Illinois Highland Park is a suburban city located in the southeastern part of Lake County, Illinois, United States, about north of downtown Chicago. Per the 2020 census, the population was 30,176. Highland Park is one of several municipalities located o ...
near Chicago. (May 2007) File:Newly molted Brood XIII.JPG, Teneral adult Brood XIII ''Magicicada'' in suburban Chicago (May 24, 2007) File:Magicicada young.jpg, Mass of ''Magicicada'' Teneral adults and exuviae on vegetation File:SD-1071.jpg, An adult Brood X ''Magicicada septendecim'' in
Princeton, New Jersey Princeton is a municipality with a borough form of government in Mercer County, in the U.S. state of New Jersey. It was established on January 1, 2013, through the consolidation of the Borough of Princeton and Princeton Township, both of whi ...
(June 6, 2004) File:Cicada Sex Brood X 2021-05-31 092614 1 crop.jpg, Two Brood X ''Magicicada''s mating in
Bethesda, Maryland Bethesda () is an unincorporated, census-designated place in southern Montgomery County, Maryland. It is located just northwest of Washington, D.C. It takes its name from a local church, the Bethesda Meeting House (1820, rebuilt 1849), which in ...
near Washington, D.C. (May 31, 2021) File:Magicicada ovipositing.jpg, A Brood X ''Magicicada'' ovipositing eggs in a tree branch near
Baltimore Baltimore ( , locally: or ) is the List of municipalities in Maryland, most populous city in the U.S. state of Maryland, fourth most populous city in the Mid-Atlantic (United States), Mid-Atlantic, and List of United States cities by popula ...
, Maryland (May 26, 2021) File:17 Year Cicada - Brood X laying eggs in a tree branch 2021-06-01 13 59 48.webm, A Brood X ''Magicicada'' laying eggs in a tree branch (video) (June 1, 2021) File:cicada egg slits 20040606 200213 1.jpg, ''Magicicada'' egg slits (circled in red)


Predator satiation survival strategy

The nymphs emerge in very large numbers at nearly the same time, sometimes more than 1.5 million individuals per
acre The acre is a unit of land area used in the imperial Imperial is that which relates to an empire, emperor, or imperialism. Imperial or The Imperial may also refer to: Places United States * Imperial, California * Imperial, Missouri * Imp ...
(>370/m2). Their mass emergence is, among other things, a survival trait called
predator satiation Predator satiation (less commonly called predator saturation) is an anti-predator adaptation in which prey briefly occur at high population densities, reducing the probability of an individual organism being eaten. When predators are flooded with ...
. The details of this strategy are simple: for the first week after emergence the periodical cicadas are easy prey for
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s,
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s,
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s,
dogs The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
and other small and large
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. In their present range the periodical cicadas have no effective predators, and all other animals feeding on them after emergence quickly fade into irrelevance with respect to their impact on total cicada populations. Early entomologists maintained that the cicadas' overall survival mechanism was simply to overwhelm predators by their sheer numbers, ensuring the survival of most of the individuals. Later, the fact that the developmental periods were both a
prime number A prime number (or a prime) is a natural number greater than 1 that is not a product of two smaller natural numbers. A natural number greater than 1 that is not prime is called a composite number. For example, 5 is prime because the only ways ...
of years (13 and 17) was hypothesized to be a predator avoidance strategy, one adopted to eliminate the possibility of potential predators receiving periodic population boosts by synchronizing their own generations to
divisor In mathematics, a divisor of an integer n, also called a factor of n, is an integer m that may be multiplied by some integer to produce n. In this case, one also says that n is a multiple of m. An integer n is divisible or evenly divisible by ...
s of the cicada emergence period. On this prime number hypothesis, a predator with a three-year reproductive cycle, which happened to coincide with a brood emergence in a given year, will have gone through either four cycles plus one year (12 + 1) or five cycles plus two years (15 + 2) by the next time that brood emerges. In this way prime-numbered broods exhibit a strategy to ensure that they nearly always emerge when some portion of the predators they will confront are sexually immature and therefore incapable of taking maximum advantage of the momentarily limitless food supply. Another viewpoint turns this hypothesis back onto the cicada broods themselves. It posits that the prime-numbered developmental times represent an adaptation to prevent hybridization ''between'' broods. It is hypothesized that this unusual method of sequestering different populations in time arose when conditions were extremely harsh. Under those conditions the mutation producing extremely long development times became so valuable that cicadas which possessed it found it beneficial to protect themselves from mating with cicadas that lacked the long-development trait. In this way, the long-developing cicadas retained a trait allowing them to survive the period of heavy selection pressure (i.e., harsh conditions) brought on by isolated and lowered populations during the period immediately following the retreat of glaciers (in the case of periodic cicadas, the North American Pleistocene glacial stadia). When seen in this light, their mass emergence and the predator satiation strategy that follows from this serves only to maintain the much ''longer-term'' survival strategy of protecting their long-development trait from hybridizations that might dilute it. This hybridization hypothesis was subsequently supported through a series of mathematical models and remains the most widely-accepted explanation for the unusually lengthy and mathematically sophisticated survival strategy of these insects. The length of the cycle was hypothesized to be controlled by a single gene locus, with the 13-year cycle dominant to the 17-year one, but this interpretation remains controversial and unsubstantiated at the level of DNA.


Impact on other populations

Cycles in cicada populations are significant enough to affect other animal and plant populations. For example, tree growth has been observed to decline the year before the emergence of a brood because of the increased feeding on roots by the growing nymphs.
Moles Moles can refer to: * Moles de Xert, a mountain range in the Baix Maestrat comarca, Valencian Community, Spain * The Moles (Australian band) *The Moles, alter ego of Scottish band Simon Dupree and the Big Sound People *Abraham Moles, French engin ...
, which feed on nymphs, have been observed to do well during the year before an emergence, but suffer population declines the following year because of the reduced food source.
Wild turkey The wild turkey (''Meleagris gallopavo'') is an Upland game bird, upland ground bird native to North America, one of two extant species of Turkey (bird), turkey and the heaviest member of the order Galliformes. It is the ancestor to the domestic ...
populations respond favorably to increased nutrition in their food supply from gorging on cicada adults on the ground at the end of their life cycles. Uneaten carcasses of periodic cicadas decompose on the ground, providing a resource pulse of nutrients to the forest community. Cicada broods may also have a negative impact.
Eastern gray squirrel The eastern gray squirrel (''Sciurus carolinensis''), also known, particularly outside of North America, as simply the grey squirrel, is a tree squirrel in the genus ''Sciurus''. It is native to eastern North America, where it is the most prodi ...
populations have been negatively affected, because the egg-laying activity of female cicadas damaged upcoming mast crops.


Broods

Periodical cicadas are grouped into geographic
broods Broods is a New Zealand musical duo from Nelson, composed of Georgia Josiena Nott on lead vocals, with older brother and multi-instrumentalist Caleb Allan Joseph Nott on production and backing vocals. They released the single "Bridges", which w ...
based on the calendar year when they emerge. For example, in 2014, the 13-year brood XXII emerged in Louisiana and the 17-year brood III emerged in western Illinois and eastern Iowa. In 1907,
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
Charles Lester Marlatt assigned Roman numerals to 30 different broods of periodical cicadas: 17 distinct broods with a 17-year life cycle, to which he assigned brood numbers I through XVII (with emerging years 1893 through 1909); plus 13 broods with a 13-year cycle, to which he assigned brood numbers XVIII through XXX (1893 through 1905). Marlatt noted that the 17-year broods are generally more northerly than are the 13-year broods. Many of these hypothetical 30 broods have not been observed. Marlatt noted that some cicada populations (especially Brood XI in the valley of the
Connecticut River The Connecticut River is the longest river in the New England region of the United States, flowing roughly southward for through four states. It rises 300 yards (270 m) south of the U.S. border with Quebec, Canada, and discharges at Long Island ...
in
Massachusetts Massachusetts (Massachusett language, Massachusett: ''Muhsachuweesut assachusett writing systems, məhswatʃəwiːsət'' English: , ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England ...
and
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York to the west, and Long Island Sound to the south. Its cap ...
) were disappearing, a fact that he attributed to the reduction in forests and the introduction and proliferation of insect-eating "English sparrows" (House sparrows, ''Passer domesticus'') that had followed the European settlement of North America. Two of the broods that Marlatt named (Broods XI and XXI) are now extinct. His numbering scheme has been retained for convenience (and because it clearly separates 13- and 17-year life cycles), although only 15 broods are known to survive today.


Map of brood locations


Taxonomy


Phylogeny

''Magicicada'' is a member of the cicada
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
Lamotialnini Lamotialnini is a tribe of cicadas in the family Cicadidae. There are about 19 genera and at least 90 described species in Lamotialnini, occurring worldwide except South America. Genera These 19 genera belong to the tribe Lamotialnini: * '' Ab ...
, which is distributed globally aside from
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. Despite ''Magicicada'' being only found in eastern
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 Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, its closest relatives are thought to be the genera ''
Tryella ''Tryella'' is a genus of 14 species of cicada found in Australia and New Guinea. For many years, the members were classified in the genus '' Abricta''. However, recent review of the genus has shown it to be a disparate group of species, and the ...
'' and '' Aleeta'' from
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, with ''Magicicada'' being
sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...
to the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
containing ''Tryella'' and ''Aleeta''. Within the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
, its closest relative is thought to be the genus '' Chrysolasia'' from
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by H ...
.


Species

Seven recognized species are placed within ''Magicicada''—three 17-year species and four 13-year species. These seven species are also sometimes grouped differently into three subgroups, the so-called Decim species group, Cassini species group, and Decula species group, reflecting strong similarities of each 17-year species with one or more species with a 13-year cycle.


Evolution and speciation

Not only are the periodical cicada life cycles curious for their use of the prime numbers 13 or 17, but their evolution is also intricately tied to one- and four-year changes in their life cycles. One-year changes are less common than four-year changes and are probably tied to variation in local climatic conditions. Four-year early and late emergences are common and involve a much larger proportion of the population than one-year changes. The different species are well-understood to have originated from a process of
allochronic speciation Allochronic speciation (also known as allochronic isolation, or temporal isolation) is a form of speciation (specifically ecological speciation) arising from reproductive isolation that occurs due to a change in breeding time that reduces or eli ...
, in which species subpopulations that are isolated from one another in time eventually become
reproductively isolated The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offsprin ...
as well. Recent research suggests that in extant periodical cicadas, the 13- and 17-year life cycles evolved at least eight different times in the last 4 million years and that different species with identical life cycles developed their overlapping geographic distribution by synchronizing their life cycles to the existing dominant populations. The same study estimates that the Decim species group split from the common ancestor of the Decula plus Cassini species groups around 4 million years ago (Mya). At around 2.5 Mya, the Cassini and Decula groups split from each other. The Sota ''et al.'' (2013) paper also calculates that the first separation of extant 13-year cicadas from 17-year cicadas took place in the Decim group about 530,000 years ago when the southern ''M. tredecim'' split from the northern ''M. septendecim''. The second noteworthy event took place about 320,000 years ago with the split of the western Cassini group from its conspecifics to the east. The Decim and the Decula clades experienced similar western splits, but these are estimated to have taken place 270,000 and 230,000 years ago, respectively. The 13- and 17-year splits in Cassini and Decula took place after these events. The 17-year cicadas largely occupy formerly glaciated territory, and as a result their phylogeographic relationships reflect the effects of repeated contraction into glacial refugia (small islands of suitable habitat) and subsequent re-expansion during multiple interglacial periods. In each species group, Decim, Cassini, and Decula, the signature of the glacial periods is manifested today in three phylogeographic genetic subdivisions: one subgroup east of the Appalachians, one midwestern, and one on the far western edge of their range. The Sota ''et al.'' data suggest that the founders of the southern 13-year cicada populations seen today originated from the Decim group. These were later joined by Cassini originating from the western Cassini clade and Decula originating from eastern, middle, and western Decula clades. As Cassini and Decula invaded the south, they became synchronized with the resident ''M. tredecim''. Today, these Cassini and Decula are known as ''M. tredecassini'' and ''M. tredecula''. More data is needed to lend support to this hypothesis and others hypotheses related to more recent 13- and 17-year splits involving ''M. neotredecim'' and ''M. tredecim''.


Distribution

The 17-year periodical cicadas are distributed across the Eastern, upper Midwestern, and Great Plains states within the U.S., while the 13-year cicadas occur in the Southern and Mississippi Valley states, but some may overlap slightly. For example, broods IV (17-year cycle) and XIX (13-year cycle) overlap in western Missouri and eastern Oklahoma.See Figure 1, p. 107 ''in'' Their emergences should again coincide in 2219, 2440, 2661, etc., as they did in 1998 (although distributions change slightly from generation to generation and older distribution maps can be unreliable.). An effort that the
National Geographic Society The National Geographic Society (NGS), headquartered in Washington, D.C., United States, is one of the largest non-profit scientific and educational organizations in the world. Founded in 1888, its interests include geography, archaeology, and ...
is sponsoring is currently underway (as of April 2021) at the
University of Connecticut The University of Connecticut (UConn) is a public land-grant research university in Storrs, Connecticut, a village in the town of Mansfield. The primary 4,400-acre (17.8 km2) campus is in Storrs, approximately a half hour's drive from Hart ...
to generate new distribution maps of all periodical cicada broods. The effort uses
crowdsourced Crowdsourcing involves a large group of dispersed participants contributing or producing goods or services—including ideas, votes, micro-tasks, and finances—for payment or as volunteers. Contemporary crowdsourcing often involves digita ...
data and records that
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
s and volunteers collect.


Symbiosis

''Magicicada'' are unable to obtain all of the essential amino acids from the dilute xylem fluid that they feed upon, and instead rely upon
endosymbiotic An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
bacteria that provide essential vitamins and nutrients for growth. Bacteria in the genus ''Hodgkinia'' live inside periodical cicadas, and grow and divide for years before punctuated cicada reproduction events impose
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charle ...
on these bacteria to maintain a mutually beneficial relationship. As a result, the genome of ''Hodgkinia'' has fractionated into three independent bacterial species each containing only a subset of genes essential for this symbiosis. The host now requires all three subgroups of symbionts, as only the complete complement of all three subgroups provides the host with all its essential nutrients. The ''Hodgkinia''–''Magicicada'' symbiosis is a powerful example of how bacterial endosymbionts drive the evolution of their hosts.


History

The first known account of a large emergence of cicadas appeared in a 1633 report by William Bradford, the governor of the
Plymouth Colony Plymouth Colony (sometimes Plimouth) was, from 1620 to 1691, the British America, first permanent English colony in New England and the second permanent English colony in North America, after the Jamestown Colony. It was first settled by the pa ...
, which had been established in 1620 within the future state of Massachusetts. After describing a "pestilent fever" that had swept through the colony and neighboring Indians, the report stated:
It is to be observed that, the spring before this sickness, there was a numerous company of ''Flies'' which were like for bigness unto wasps or ''Bumble-Bees''; they came out of little holes in the ground, and did eat up the green things, and made such a constant yelling noise as made the woods ring of them, and ready to deafen the hearers; they were not any seen or heard by the ''English'' in this country before this time; but the ''Indians'' told them that sickness would follow, and so it did, very hot, in the months of ''June'', ''July'', and ''August'' of that summer.
(Elaborating on an observation that Marlatt reported in 1907, Gene Kritsky has suggested that Bradford's report is misdated, as Broods XI and XIV would have emerged in Plymouth in 1631 and 1634, respectively, while no presently known brood would have emerged there in 1633.) Historical accounts cite reports of 15- to 17-year recurrences of enormous numbers of noisy emergent cicadas ("locusts") written as early as 1733.
John Bartram John Bartram (March 23, 1699 – September 22, 1777) was an American botanist, horticulturist, and explorer, based in Philadelphia, Pennsylvania, for most of his career. Swedish botanist and taxonomist Carl Linnaeus said he was the "greatest na ...
, a noted Philadelphia
botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
and
horticulturist Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
, was among the early writers that described the insect's life cycle, appearance and characteristics. On May 9, 1715, the Rev. Andreas Sandel, the pastor of
Philadelphia Philadelphia, often called Philly, is the largest city in the Commonwealth of Pennsylvania, the sixth-largest city in the U.S., the second-largest city in both the Northeast megalopolis and Mid-Atlantic regions after New York City. Sinc ...
's "Gloria Dei" Swedish
Lutheran Lutheranism is one of the largest branches of Protestantism, identifying primarily with the theology of Martin Luther, the 16th-century German monk and reformer whose efforts to reform the theology and practice of the Catholic Church launched th ...
Church, described in his journal an emergence of Brood X.
Pehr Kalm Pehr Kalm (6 March 1716 – 16 November 1779), also known as Peter Kalm, was a Swedish explorer, botanist, naturalist, and agricultural economist. He was one of the most important apostles of Carl Linnaeus. In 1747, he was commissioned by the ...
, a
Finnish Finnish may refer to: * Something or someone from, or related to Finland * Culture of Finland * Finnish people or Finns, the primary ethnic group in Finland * Finnish language, the national language of the Finnish people * Finnish cuisine See also ...
naturalist visiting Pennsylvania and New Jersey in 1749 on behalf of the
Royal Swedish Academy of Sciences The Royal Swedish Academy of Sciences ( sv, Kungliga Vetenskapsakademien) is one of the Swedish Royal Academies, royal academies of Sweden. Founded on 2 June 1739, it is an independent, non-governmental scientific organization that takes special ...
, observed in late May another emergence of that brood. Republished by When reporting the event in a paper that a Swedish academic journal published in 1756, Kalm wrote: Kalm then described Rev. Sandel's report and one that he had obtained from
Benjamin Franklin Benjamin Franklin ( April 17, 1790) was an American polymath who was active as a writer, scientist, inventor, statesman, diplomat, printer, publisher, and political philosopher. Encyclopædia Britannica, Wood, 2021 Among the leading inte ...
that had recorded in Philadelphia the emergence from the ground of large numbers of cicadas during early May 1732. He noted that the people who had prepared these documents had made no such reports in other years. Kalm further noted that others had informed him that they had seen cicadas only occasionally before the insects emerged from the ground in Pennsylvania in large swarms on May 22, 1749. He additionally stated that he had not heard any cicadas in Pennsylvania and New Jersey in 1750 in the same months and areas in which he had heard many in 1749. The 1715 and 1732 reports, when coupled with his own 1749 and 1750 observations, supported the previous "general opinion" that he had cited. Kalm summarized his findings in a book translated into English and published in London in 1771, stating: Based on Kalm's account and a specimen that Kalm had provided, in 1758
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
named the insect ''Cicada septendecim'' in the tenth edition of his ''Systema Naturae''. Moses Bartram, a son of John Bartram, described the next appearance of the brood (Brood X) that Kalm had observed in 1749 in an article entitled ''Observations on the cicada, or locust of America, which appears periodically once in 16 or 17 years'' that he wrote in 1766. Bartram's article, which a London journal published in 1768, noted that upon hatching from eggs deposited in the twigs of trees, the young insects ran down to the earth and "entered the first opening that they could find". He reported that he had been able to discover them below the surface, but that others had reportedly found them deep. In 1775,
Thomas Jefferson Thomas Jefferson (April 13, 1743 – July 4, 1826) was an American statesman, diplomat, lawyer, architect, philosopher, and Founding Fathers of the United States, Founding Father who served as the third president of the United States from 18 ...
recorded in his "Garden Book" Brood II's 17-year periodicity, writing that an acquaintance remembered "great locust years" in 1724 and 1741, that he and others recalled another such year in 1758 and that the insects had again emerged from the ground at
Monticello Monticello ( ) was the primary plantation of Founding Father Thomas Jefferson, the third president of the United States, who began designing Monticello after inheriting land from his father at age 26. Located just outside Charlottesville, V ...
in 1775. He noted that the females lay their eggs in the small twigs of trees while above ground. In April 1800,
Benjamin Banneker Benjamin Banneker (November 9, 1731October 19, 1806) was an African-American naturalist, mathematician, astronomer and almanac author. He was a landowner who also worked as a surveyor and farmer. Born in Baltimore County, Maryland, to a fr ...
, who lived near Ellicott's Mills, Maryland, wrote in his record book that he recalled a "great locust year" in 1749, a second in 1766 during which the insects appeared to be "full as numerous as the first", and a third in 1783. He predicted that the insects (Brood X) "may be expected again in they year 1800 which is Seventeen Since their third appearance to me". Describing an effect that the
pathogenic fungus Pathogenic fungi are fungi that cause disease in humans or other organisms. Approximately 300 fungi are known to be pathogenic to humans. Markedly more fungi are known to be pathogenic to plant life than those of the animal kingdom. The study of fu ...
, '' Massospora cicadina'', has on its
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
, Banneker's record book stated that the insects:
.... begin to Sing or make a noise from first they come out of the Earth till they die. The hindermost part rots off, and it does not appear to be any pain to them, for they still continue on Singing till they die.
In 1845, Dr. D.L. Pharas of
Woodville, Mississippi Woodville is a town in and the county seat of Wilkinson County, Mississippi, United States. Its population as of 2020 was 928. History This historic town, one of the oldest in Mississippi, is set among the rolling hills and pastures of Wilkins ...
, announced the 13-year periodicity of the southern cicada broods in a local newspaper, the '' Woodville Republican''. In 1858, Pharas placed the title ''Cicada tredecim'' in a subsequent article that the newspaper published on the subject. Ten years later, the ''American Entomologist'' published in December 1868 a paper that
Benjamin Dann Walsh Benjamin Dann Walsh (September 21, 1808 – November 18, 1869) was an English-born American entomologist who served as the first official state entomologist in Illinois.Mallis (1971) He was a leading influence during a time of significant tra ...
and
Charles Valentine Riley Charles Valentine Riley (18 September 1843 – 14 September 1895) was a British-born American entomologist and artist. He was one of the first individuals to use biological pest control and authored over 2,400 publications. He convinced Congress ...
had written that also reported the 13-year periodicity of the southern cicada broods. Walsh's and Riley's paper, which ''
Scientific American ''Scientific American'', informally abbreviated ''SciAm'' or sometimes ''SA'', is an American popular science magazine. Many famous scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it. In print since 1845, it i ...
'' reprinted in January 1869, illustrated the interior and exterior characteristics of the nymphs' emergence holes and raised turrets. Their article, which did not cite Pharas' reports, was the first to describe the southern cicadas' 13-year periodicity that received widespread attention. Riley later acknowledged Pharas' work in an 1885 publication on periodical cicadas that he authored. In 1998, an emergence contained a brood of 17-year cicadas (Brood IV) in western Missouri and a brood of 13-year cicadas (Brood XIX) over much of the rest of the state. Each of the broods are the state's largest of their types. As the territories of the two broods overlap (converge) in some areas, the convergence was the state's first since 1777.


Use as human food

''Magicicada'' species are edible when cooked for people who lack allergies to similar foods. A number of recipes are available for this purpose. Some recommend collecting the insects shortly after molting while still soft. Others exhibit preferences for emergent nymphs or hardened adults. The insects have historically been eaten by Native Americans, who roasted them in hot ovens, stirring them until they were well browned. Marlatt wrote in 1907:


Notes


References

*


Further reading

*


External links

*
''Cicada Mania''
*
GIGAmacro
has a zoomable, very high-resolution image of th
male, female & nymph cicada''InsectSingers.com''
Recordings of species-specific songs of many North American cicada species. *Liebhold, A.M.; Bohne, M.J.; Lilja, R.L.
Active Periodical Cicada Broods of the United States
(map). ''USDA Forest Service Northern Research Station, Northeastern Area State and Private Forestry''. 2013.
Magicicada.org Brood mapping project
General periodical cicada information. Also solicits records and observations from the general public *
''Massachusetts Cicadas''
describes behavior, sightings, photos, "how to find" guide, videos and distribution maps of New England and U.S. periodical and annual cicada species including Brood X, Brood XIII, Brood XIV and Brood XIX {{Authority control Cicadas Edible insects Lamotialnini Native American cuisine Periodic phenomena Antipredator adaptations