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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, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, 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 Latin ''locusta'', locust or lobster) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstances they b ...
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 of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
(true bugs), suborder Auchenorrhyncha, 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 – gras ...
. ''Magicicada'' belongs to the cicada tribe Lamotialnini, a group of
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
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 (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...
. While underground, the nymphs feed on
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
fluids from the roots of broadleaf 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 (depending on latitude), 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 4'' ''years) in northeast India and for a cicada species from Fiji, '' Raiateana knowlesi'' (every 8'' ''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 ("imaginal" being "imago" in adjective form), the stage in wh ...
(adult) periodical cicada has two red compound eyes, three small ocelli, 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'', Species description, first described by C ...
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 (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 A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
can pierce human skin when it is handled, which is painful but in no other way harmful. Cicadas are neither
venom 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 ...
ous nor poisonous and there is no evidence that they or their bites can transmit diseases. Oviposition by female periodical cicadas damages pencil-sized twigs of woody vegetation. Mature trees rarely suffer lasting damage, although peripheral twig die-off or "flagging" may result. Planting young trees or shrubs is best postponed until after an expected emergence of the periodical cicadas. Existing young trees or shrubs can be covered with cheesecloth 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.


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. 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 cicadas because some are seen every summer. This may lead some to conclude that the non-periodic cicadas have life cycles of 1'' ''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, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
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. The nymphs seem to track the number of years by detecting the changes in the
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
caused by
abscission Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is the intentional shedding of a body part, such as the shedding of a claw, husk, or the autotomy of a ...
of the tree. This was supported experimentally by inducing a grove of trees to go through two cycles of losing and re-growing leaves in one calendar year. Cicadas feeding on those trees emerged after 16 years instead of 17. 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 (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
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 ...
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 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 () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
s. Just after emerging from this final molt the teneral 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 ''Magicicada'' nymphs created in
Potomac, Maryland Potomac () is an Unincorporated area#United States, unincorporated community and census-designated place in Montgomery County, Maryland, United States. As of the 2020 United States census, 2020 census, it had a population of 47,018. It is named a ...
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
adult An adult is an animal that has reached full growth. The biological definition of the word means an animal reaching sexual maturity and thus capable of reproduction. In the human context, the term ''adult'' has meanings associated with social an ...
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 molted. The exuviae of an animal can be important to biologists as they can often be ...
after molting in
Highland Park, Illinois Highland Park is a suburban city located in southeastern Lake County, Illinois, United States, about north of downtown Chicago. Per the 2020 United States census, 2020 census, the population was 30,176. Highland Park is one of several municipali ...
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 The Municipality of Princeton is a Borough (New Jersey), borough in Mercer County, New Jersey, United States. It was established on January 1, 2013, through the consolidation of the Borough of Princeton, New Jersey, Borough of Princeton and Pri ...
(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, United States. Located just northwest of Washington, D.C., it is a major business and government center of the Washington metropolitan region ...
near Washington, D.C. (May 31, 2021) File:Magicicada ovipositing.jpg, A Brood X ''Magicicada'' ovipositing eggs in a tree branch near
Baltimore Baltimore is the most populous city in the U.S. state of Maryland. With a population of 585,708 at the 2020 census and estimated at 568,271 in 2024, it is the 30th-most populous U.S. city. The Baltimore metropolitan area is the 20th-large ...
, 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 measurement, unit of land area used in the Imperial units, British imperial and the United States customary units#Area, United States customary systems. It is traditionally defined as the area of one Chain (unit), ch ...
(>370/m2). Their mass emergence is, among other things, a survival trait called predator satiation. The details of this strategy are simple: for the first week after emergence the periodical cicadas are easy prey for
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s,
squirrel Squirrels are members of the family Sciuridae (), a family that includes small or medium-sized rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrel ...
s,
cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
s,
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers. ...
s and other small and large
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s. In their present range the periodical cicadas have no effective predators, and all other animals feeding on them after emergence quickly become irrelevant 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 each a
prime number A prime number (or a prime) is a natural number greater than 1 that is not a Product (mathematics), 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 ...
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 divisibl ...
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 periodical 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 Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
.


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, 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 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 turkey (''M. g. dom ...
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 periodical 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 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 are a musical duo from Nelson, New Zealand, 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 (Broo ...
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 (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
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 into Long Isl ...
in
Massachusetts Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...
and
Connecticut Connecticut ( ) is a U.S. state, state in the New England region of the Northeastern United States. It borders Rhode Island to the east, Massachusetts to the north, New York (state), New York to the west, and Long Island Sound to the south. ...
) 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) have become 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. Periodical cicadas that emerge outside the expected time frame are called stragglers. Although they can emerge at any time, they usually do so one or four years before or after most other members of their broods emerge. Stragglers with a 17-year life cycle typically emerge four years early. Those with a 13-year cycle typically emerge four years late. The emergence of stragglers may in theory be indicative of a brood shifting from a 17-year cycle to a 13-year one. Brood XIII of the 17-year cicada, which reputably has the largest emergence of cicadas by size known anywhere, and Brood XIX of the 13-year cicada, arguably the largest (by geographic extent) of all periodical cicada broods, were expected to emerge together in 2024 for the first time since 1803. However, the two broods were not expected to overlap except potentially in a thin area in central and eastern Illinois ( Macon, Sangamon, Livingston, and Logan counties). The next such dual emergence of these two particular broods will occur in 2245, 221 years after 2024. Many other 13-year and 17-year broods emerge during the same years, but the broods are not geographically close.


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 use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
Lamotialnini, 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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. Despite ''Magicicada'' being only found in eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, 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 ''Aleeta'' is a genus of cicadas with the Monotypic taxon, only species ''Aleeta curvicosta'' (commonly known as the floury baker or floury miller, known until 2003 as ''Abricta curvicosta''), one of Australia's most familiar insects. Native to ...
'' from
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, with ''Magicicada'' being
sister A sister is a woman or a girl who shares parents or a parent 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 ref ...
to the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
containing ''Tryella'' and ''Aleeta''. Within the
Americas The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.''Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sing ...
, its closest relative is thought to be the genus '' Chrysolasia'' from
Guatemala Guatemala, officially the Republic of Guatemala, is a country in Central America. It is bordered to the north and west by Mexico, to the northeast by Belize, to the east by Honduras, and to the southeast by El Salvador. It is hydrologically b ...
.


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 elimi ...
, in which species subpopulations that are isolated from one another in time eventually become reproductively isolated as well. 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 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 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''. 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 from the Eastern states, across the
Ohio Valley The Ohio River () is a river in the United States. It is located at the boundary of the Midwestern and Southern United States, flowing in a southwesterly direction from Pittsburgh, Pennsylvania, to its mouth on the Mississippi River in Cairo, ...
, to the Great Plains states and north to the edges of the
Upper Midwest The Upper Midwest is a northern subregion of the U.S. Census Bureau's Midwestern United States. Although the exact boundaries are not uniformly agreed upon, the region is usually defined to include the states of Iowa, Michigan, Minnesota and Wi ...
, while the 13-year cicadas occur in the Southern and
Mississippi Valley The Mississippi River is the main stem, primary river of the largest drainage basin in the United States. It is the second-longest river in the United States, behind only the Missouri River, Missouri. From its traditional source of Lake Ita ...
states, with some slight overlap of the two groups. 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 sponsored by the
National Geographic Society The National Geographic Society, headquartered in Washington, D.C., United States, is one of the largest nonprofit scientific and educational organizations in the world. Founded in 1888, its interests include geography, archaeology, natural sc ...
is underway as of April 2021 at the
University of Connecticut The University of Connecticut (UConn) is a public land-grant research university system with its main campus in Storrs, Connecticut, United States. It was founded in 1881 as the Storrs Agricultural School, named after two benefactors. In 1893, ...
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 digit ...
data and records that
entomologist Entomology (from Ancient Greek ἔντομον (''éntomon''), meaning "insect", and -logy from λόγος (''lógos''), meaning "study") is the branch of zoology that focuses on insects. Those who study entomology are known as entomologists. In ...
s and volunteers collect.


Parasites, pests and pathogens

Although it usually feeds on oak leaf gall midge ('' Polystepha pilulae'') larvae and other insects, the oak leaf gall mite ("itch mite") ('' Pyemotes herfsi'') becomes an
ectoparasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
of periodical cicada eggs when these are available. After cicadas deposit their eggs in the branches of trees, feeding mites reproduce and their numbers increase.Multiple sources: * * * * * First report of ''Pyemotes herfsi'' parasitism on periodical cicada eggs. * * * * * Contains 1:38 minute video showing images and descriptions of ''Pyemotes herfsi'' bites. * * After cicada emergences have ended, many people have therefore developed rashes, pustules, intense itching and other mite bite sequelae on their upper
torso The torso or trunk is an anatomical terminology, anatomical term for the central part, or the core (anatomy), core, of the body (biology), body of many animals (including human beings), from which the head, neck, limb (anatomy), limbs, tail an ...
, head, neck and arms. Rashes and itching peaked after several days, but lasted as long as two weeks. Anti-itch treatments, including calamine lotion and topical steroid creams, did not relieve the itching. '' Massospora cicadina'' is a pathogenic fungus 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.


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 an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), which live in the root ...
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 Heredity, heritable traits characteristic of a population over generation ...
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 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 spelled Plimouth) was the first permanent English colony in New England from 1620 and the third permanent English colony in America, after Newfoundland and the Jamestown Colony. It was settled by the passengers on t ...
, which had been established in 1620 within the future state of
Massachusetts Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...
. 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 (June 3, 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 natu ...
, a noted
Philadelphia Philadelphia ( ), colloquially referred to as Philly, is the List of municipalities in Pennsylvania, most populous city in the U.S. state of Pennsylvania and the List of United States cities by population, sixth-most populous city in the Unit ...
botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
and
horticulturist Horticulture (from ) is the art and science of growing fruits, vegetables, flowers, trees, shrubs and ornamental plants. Horticulture is commonly associated with the more professional and technical aspects of plant cultivation on a smaller and mo ...
, was among the early writers that described the insect's life cycle, appearance and characteristics. On May 9, 1715, Andreas Sandel, the pastor of Philadelphia's "Gloria Dei" Swedish
Lutheran Lutheranism is a major branch of Protestantism that emerged under the work of Martin Luther, the 16th-century German friar and Protestant Reformers, reformer whose efforts to reform the theology and practices of the Catholic Church launched ...
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-Finnish List of explorers, explorer, botany, botanist, natural history, naturalist, and Agricultural economics, agricultural economist. He was one of the most ...
, a Finnish naturalist visiting Pennsylvania and New Jersey in 1749 on behalf of the
Royal Swedish Academy of Sciences The Royal Swedish Academy of Sciences () 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 responsibility for promoting nat ...
, 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 Sandel's report and one that he had obtained from
Benjamin Franklin Benjamin Franklin (April 17, 1790) was an American polymath: a writer, scientist, inventor, statesman, diplomat, printer, publisher and Political philosophy, political philosopher.#britannica, Encyclopædia Britannica, Wood, 2021 Among the m ...
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 ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
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 (, 1743July 4, 1826) was an American Founding Fathers of the United States, Founding Father and the third president of the United States from 1801 to 1809. He was the primary author of the United States Declaration of Indepe ...
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 residence and plantation of Thomas Jefferson, a Founding Father, author of the Declaration of Independence, and the third president of the United States. Jefferson began designing Monticello after inheriting l ...
in 1775. He noted that the females lay their eggs in the small twigs of trees while above ground. The 1780 emergence of the Brood VII cicadas (also known as the Onondaga brood) during the
American Revolutionary War The American Revolutionary War (April 19, 1775 – September 3, 1783), also known as the Revolutionary War or American War of Independence, was the armed conflict that comprised the final eight years of the broader American Revolution, in which Am ...
, coincided with the aftermath of the military operation known as the
Sullivan Expedition The 1779 Sullivan Expedition (also known as the Sullivan-Clinton Expedition, the Sullivan Campaign, and the Sullivan-Clinton Campaign) was a United States military campaign under the command of General John Sullivan (general), John Sullivan duri ...
which devastated the indigenous Onondagan communities and destroyed their crops. The sudden arrival of such a substantial quantity of the cicadas provided a source of sustenance for the Onondaga people who were experiencing severe food insecurity following the Sullivan campaigns and the subsequent brutal winter. The seemingly miraculous arrival of the cicadas is commemorated by the Onondaga as though it were an intervention by the Creator to ensure their survival after such a traumatizing, catastrophic event. In April 1800,
Benjamin Banneker Benjamin Banneker (November 9, 1731October 19, 1806) was an American Natural history, naturalist, mathematician, astronomer and almanac author. A Land tenure, landowner, he also worked as a surveying, surveyor and farmer. Born in Baltimore Co ...
, 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, '' 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 * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
, 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, D.L. Pharas of
Woodville, Mississippi Woodville is one of the oldest towns in Mississippi and is the county seat of Wilkinson County, Mississippi, United States. Its population as of 2020 was 928. Geography According to the United States Census Bureau, the town has a total area of ...
, 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 scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it, with more than 150 Nobel Pri ...
'' 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. In 2007 and 2008, Edmond Zaborski, a research scientist with the Illinois Natural History Survey, reported that the oak leaf gall mite ("itch mite") (''Pyemotes herfsi'') is an ectoparasite of periodical cicada eggs. While investigating with the help of others the mysterious itchy welts and rashes that people were developing in
Chicago Chicago is the List of municipalities in Illinois, most populous city in the U.S. state of Illinois and in the Midwestern United States. With a population of 2,746,388, as of the 2020 United States census, 2020 census, it is the List of Unite ...
's suburbs after the end of a 2007 Brood XIII emergence, he attributed the event to bites by mites whose populations had quickly increased while parasitizing those eggs. Similar events occurred in
Cincinnati Cincinnati ( ; colloquially nicknamed Cincy) is a city in Hamilton County, Ohio, United States, and its county seat. Settled in 1788, the city is located on the northern side of the confluence of the Licking River (Kentucky), Licking and Ohio Ri ...
after a Brood XIV emergence ended in 2008, in
Cleveland Cleveland is a city in the U.S. state of Ohio and the county seat of Cuyahoga County. Located along the southern shore of Lake Erie, it is situated across the Canada–U.S. maritime border and approximately west of the Ohio-Pennsylvania st ...
and elsewhere in northern and eastern
Ohio Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
after a Brood V emergence ended in 2016, in the
Washington, D.C. Washington, D.C., formally the District of Columbia and commonly known as Washington or D.C., is the capital city and federal district of the United States. The city is on the Potomac River, across from Virginia, and shares land borders with ...
, area after a Brood X emergence ended in 2021, and again in the Chicago area after the next Brood XIII emergence ended in 2024.


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 fried them or roasted them in hot ovens, stirring them until they were well browned. Republished by Marlatt wrote in 1907:


Notes


References

*


Further reading

* Wikipedia
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 ...
page. *


External links


''The Periodical Cicada Page''
Informational page about periodical cicadas that supersedes www.magicicada.org. Has maps and 3-D models. *
''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. *
''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