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Periodical cicadas
Periodical cicadas
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Periodical cicada
Specimen of Magicicada septendecim in the Bavarian State Collection of Zoology, Munich (2015)
A Magicicada chorus with M. septendecim, M. cassini, and M. septendecula
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Suborder: Auchenorrhyncha
Family: Cicadidae
Subfamily: Cicadettinae
Tribe: Lamotialnini
Genus: Magicicada
W. T. Davis, 1925
Type species
Magicicada septendecim[1]

The term periodical cicada is commonly used to refer to any of the seven species of the genus Magicicada of eastern North America, the 13- and 17-year cicadas. They are called periodical because nearly all individuals in a local population are developmentally synchronized and emerge in the same year. Although they are sometimes called "locusts", this is a misnomer, as cicadas belong to the taxonomic order Hemiptera (true bugs), suborder Auchenorrhyncha, while locusts are grasshoppers belonging to the order Orthoptera.[2] Magicicada belongs to the cicada tribe Lamotialnini, a group of genera with representatives in Australia, Africa, and Asia, as well as the Americas.[3]

Magicicada species spend around 99.5% of their long lives underground in an immature state called a nymph. While underground, the nymphs feed on xylem fluids from the roots of broadleaf forest trees in the eastern United States.[4] 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.[5][6] The adults are active for only about four to six weeks after the unusually prolonged developmental phase.[7]

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)[8] in northeast India and for a cicada species from Fiji, Raiateana knowlesi (every 8 years).[9]

Description

[edit]
Many Brood X periodical cicadas (Magicicada) (video with sound)

The winged imago (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.[10]

Adults are typically 2.4 to 3.3 cm (0.9 to 1.3 in), 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.[11]

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 is said to resemble someone calling "weeeee-whoa" or "Pharaoh".[12] The cassini and decula periodical cicadas (including M. tredecula) have songs that intersperse buzzing and ticking sounds.[11]

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 can pierce human skin when it is handled, which is painful but in no other way harmful. Cicadas are neither venomous nor poisonous and there is no evidence that they or their bites can transmit diseases.[13]

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.[14] 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 38 in (1.0 cm) in diameter or smaller to prevent damage during the oviposition period,[15] which begins about a week after the first adults emerge and lasts until all females have died.

Life cycle

[edit]
Transformation from mature nymph to adult
Time-lapse of final molt and darkening, over 4.5 hours
Emergence! Nearly all at once. Many do not survive, but with mass emergence, many will reach maturity to start the next generation.
Adult cicada female creating a slit in twig and inserting eggs. The sound is of thousands of cicadas.

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.[citation needed]

The nymphs of the periodical cicadas live underground, usually within 2 ft (61 cm) of the surface, feeding on the juices of plant roots.[16] The nymphs of the periodical cicada undergo five instar 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.[17]

The nymphs seem to track the number of years by detecting the changes in the xylem caused by abscission 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.[9]

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.[7][18] In some situations, nymphs extend mud turrets up to several inches above the soil surface.[19] The function of these turrets is not known, but the phenomenon has been observed in some nonperiodical cicadas, as well as other tunneling insects.[20]

The nymphs first emerge on a spring evening when the soil temperature at around 20 cm (8 in) of depth is above 17.9 °C (64 °F). The crepuscular emergence is thought to be related to the fact that maximum soil temperatures lag behind maximum insolation 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 exoskeletons. 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.[2] Most matings occur in so-called chorus trees.[10]

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.[21] 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.[10]

Both males and females can mate multiple times, although most females seem to mate only once [citation needed]. 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.

Predator satiation survival strategy

[edit]

The nymphs emerge in very large numbers at nearly the same time, sometimes more than 1.5 million individuals per acre (370 individuals per m2).[22] Their mass emergence is, among other things, an adaptation called predator satiation. Although periodical cicadas are easy prey for reptiles, birds, squirrels, cats, dogs and other small and large mammals[7][23], there are after synchronized emergence simply too many individuals for the predators to consume; many individuals thus remain behind to procreate.

It has been hypothesized that the prime-number development times (13 and 17 years) improve avoidance of predators with shorter reproductive cycles and for this reason have been selected for. A predator with, for example, a three-year reproductive cycle, which happened to benefit from 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 the same brood emerges. In this way cicada generations 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.[24]

A second hypothesis posits that the prime-numbered developmental times are an adaptation that prevents hybridization between broods. Under extremely harsh conditions, mutations producing extremely long development times are selected for. A mechanism, such as reproducing only after prime-numbered intervals, that reduces the frequency of cicadas mating with cicadas that may lack the long-development trait will also be selected for. The North American Pleistocene glacial stadia are instances of such extremely harsh conditions. On this hypothesis, predator satiation reinforces a longer-term survival strategy of protecting the long-development trait from hybridizations that might dilute it.[25] This hypothesis has been supported by mathematical modeling.[26]

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,[27] but this interpretation remains controversial and unsubstantiated at the level of DNA.

Impact on other populations

[edit]

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.[28] 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.[29] Wild turkey 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.[28]

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.[30]

Broods

[edit]

Periodical cicadas are grouped into geographic broods 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 a 1907 journal article, entomologist 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).[31] Marlatt noted that the 17-year broods are generally more northerly than are the 13-year broods.[32]

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 in Massachusetts and Connecticut) 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.[33] 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.[34]

Name Nickname Cycle (yrs) Last emergence Next emergence Extent
Brood I Blue Ridge brood 17 2012 2029 Western Virginia, West Virginia
Brood II East Coast brood 17 2013 2030 Connecticut, Maryland, North Carolina, New Jersey, New York, Pennsylvania, Delaware, Virginia, District of Columbia
Brood III Iowan brood 17 2014 2031 Iowa
Brood IV Kansan brood 17 2015 2032 Eastern Nebraska, southwestern Iowa, eastern Kansas, western Missouri, Oklahoma, north Texas[35]
Brood V 17 2016 2033 Eastern Ohio, Western Maryland, Southwestern Pennsylvania, Northwestern Virginia, West Virginia, New York (Suffolk County)[36]
Brood VI 17 2017 2034 Northern Georgia, western North Carolina, northwestern South Carolina
Brood VII Onondaga brood 17 2018 2035 Central New York (Onondaga, Cayuga, Seneca, Ontario, Yates counties)[Note 1]
Brood VIII 17 2019 2036 Eastern Ohio, western Pennsylvania, northern West Virginia
Brood IX 17 2020 2037 southwestern Virginia, southern West Virginia, western North Carolina
Brood X Great eastern brood 17 2021 2038 New York, New Jersey, Pennsylvania, Delaware, Maryland, District of Columbia, Virginia, West Virginia, North Carolina, Georgia, Tennessee, Kentucky, Ohio, Indiana, Illinois, Michigan[37][Note 2]
Brood XI 17 1954 Extinct Connecticut, Massachusetts, Rhode Island. Last seen in 1954 in Ashford, Connecticut along the Fenton River
Brood XIII Northern Illinois brood 17 2024 2041 Northern Illinois and in parts of Iowa, Wisconsin, and Indiana[Note 3]
Brood XIV 17 2025 2042 Southern Ohio, Kentucky, Tennessee, Massachusetts, Maryland, North Carolina, Pennsylvania, northern Georgia, Southwestern Virginia and West Virginia, and parts of New York and New Jersey
Brood XIX Great Southern Brood 13 2024 2037 Alabama, Arkansas, Georgia, Indiana, Illinois, Kentucky, Louisiana, Maryland, Missouri, Mississippi, North Carolina, Oklahoma, South Carolina, Tennessee, and Virginia[Note 4]
Brood XXI Floridian Brood 13 1870 Extinct Last recorded in 1870, historical range included the Florida panhandle[41]
Brood XXII Baton Rouge Brood[42] 13 2014 2027 Louisiana, Mississippi[Note 5]
Brood XXIII Mississippi Valley Brood[43] 13 2015 2028 Arkansas, Illinois, Indiana, Kentucky, Louisiana, Missouri, Mississippi, Tennessee
  1. ^ Consists only of M. septendecim
  2. ^ A premature emergence occurred in 2017.[38]
  3. ^ Reputedly has the largest emergence of cicadas by size known anywhere. A premature emergence occurred in 2020.[39]
  4. ^ Arguably the largest (by geographic extent) of all periodical cicada broods.[40]
  5. ^ This 13-year brood does not include M. neotredecim.

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.[44] Stragglers with a 17-year life cycle typically emerge four years early. Those with a 13-year cycle typically emerge four years late.[45] The emergence of stragglers may in theory be indicative of a brood shifting from a 17-year cycle to a 13-year one.[46]

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).[47] 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.[48]

Map of brood locations

[edit]
County-by-county map showing the locations of cicada broods, published May 2013
USDA Forest Service map of periodical cicada brood locations by county and timing of next emergence (as of 2024)

Taxonomy

[edit]

Phylogeny

[edit]

Magicicada is a member of the cicada tribe Lamotialnini, which is distributed globally aside from South America. Despite Magicicada being only found in eastern North America, its closest relatives are thought to be the genera Tryella and Aleeta from Australia, with Magicicada being sister to the clade containing Tryella and Aleeta.[49] Within the Americas, its closest relative is thought to be the genus Chrysolasia from Guatemala.[50]

Species

[edit]

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.[51]

17-year cycle Species
group
13-year cycle
Image Scientific name Common name Distribution Image Scientific name Common name Distribution
M. septendecim
(Linnaeus, 1758)
17-year locust,
Pharaoh cicada
Canada,
United States
Decim M. tredecim
(Walsh & Riley, 1868)
Southeastern
United States
M. neotredecim
Marshall & Cooley, 2000
United States
M. cassini [52]
(Fisher, 1852)
17-year cicada,
dwarf periodical cicada
United States Cassini M. tredecassini
Alexander & Moore, 1962
United States
M. septendecula
Alexander & Moore, 1962
United States Decula M. tredecula
Alexander & Moore, 1962
United States

Evolution and speciation

[edit]

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.[25][27] 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,[53][54] 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.[54] 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

[edit]

The 17-year periodical cicadas are distributed from the Eastern states, across the Ohio Valley, to the Great Plains states and north to the edges of the Upper Midwest, while the 13-year cicadas occur in the Southern and Mississippi Valley 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.[55][56] Their emergences should again coincide in 2219, 2440, 2661, etc., as they did in 1998[57] (although distributions change slightly from generation to generation and older distribution maps can be unreliable[56]).

An effort sponsored by the National Geographic Society is underway as of April 2021 at the University of Connecticut to generate new distribution maps of all periodical cicada broods.[58] The effort uses crowdsourced data and records that entomologists and volunteers collect.[59]

Parasites, pests and pathogens

[edit]

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 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.[60]

A Brood X Magicicada with abdominal Massospora cicadina infection in Bethesda, Maryland (May 31, 2021)

After cicada emergences have ended, many people have therefore developed rashes, pustules, intense itching and other mite bite sequelae on their upper torso, 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.[60]

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.[61]

Symbiosis

[edit]

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 bacteria that provide essential vitamins and nutrients for growth.[62] Bacteria in the genus Hodgkinia live inside periodical cicadas, and grow and divide for years before punctuated cicada reproduction events impose natural selection 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.[63] The HodgkiniaMagicicada symbiosis is a powerful example of how bacterial endosymbionts drive the evolution of their hosts.

History

[edit]

Marlatt wrote in his 1907 journal article that the earliest published account of the periodical cicada which had come under his observation appeared in a 1666 issue of the journal Philosophical Transactions of the Royal Society,[32] which at the time had the name Philosophical Transactions.[64] The account stated:

A great Observer, who hath lived long in New England, did upon occasion, relate to a Friend of his in London, where he lately was, That some few Years since there was such a swarm of a certain sort of Insects in that English Colony, that for the space of 200 Miles they poyson'd and destroyed all the Trees of that Country; there being found innumerable little holes in the ground, out of which those Insects broke forth in the form of Maggots, which turned into Flyes that had a kind of taile or sting, which they struck into the Tree, and thereby envenomed and killed it.[64]

Marlatt also wrote that the next report of the cicada appeared in a work entitled New-Englands Memoriall, which was printed in Cambridge, Massachusetts in 1669. After describing a "pestilent fever" that had swept through the Plymouth Colony and neighboring Indians in 1633, the New-Englands Memoriall's account 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.[32][65]

(Elaborating on an observation that Marlatt had reported in 1907,[32] Gene Kritsky suggested in 2001 that the account of the 1633 emergence 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. Kritsky also noted that William Bradford, the governor of the Plymouth Colony, had reportedly written in 1633 the same account of the cicada emergence that the New-Englands Memoriall published in 1669.[66] However, a reprint of Bradford's History of Plymouth Plantation: 1606-1646 contains a different account of that emergence.[67])

Historical accounts cite reports of 15- to 17-year recurrences of enormous numbers of noisy emergent cicadas ("locusts") written as early as 1733.[68][69] John Bartram, a noted Philadelphia botanist and horticulturist, was among the early writers that described the insect's life cycle, appearance and characteristics.[70]

On May 9, 1715, Andreas Sandel, the pastor of Philadelphia's "Gloria Dei" Swedish Lutheran Church, described in his journal an emergence of Brood X.[71] Pehr Kalm, a Finnish naturalist visiting Pennsylvania and New Jersey in 1749 on behalf of the Royal Swedish Academy of Sciences, observed in late May another emergence of that brood.[72][73] When reporting the event in a paper that a Swedish academic journal published in 1756, Kalm wrote:

The general opinion is that these insects appear in these fantastic numbers in every seventeenth year. Meanwhile, except for an occasional one which may appear in the summer, they remain underground.
There is considerable evidence that these insects appear every seventeenth year in Pennsylvania.[73]

Kalm then described Sandel's report and one that he had obtained from Benjamin Franklin 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.[73]

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.[73] 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.[73] 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,[74] stating:

There are a kind of Locusts which about every seventeen years come hither in incredible numbers ... In the interval between the years when they are so numerous, they are only seen or heard single in the woods.[68][75]

Based on Kalm's account and a specimen that Kalm had provided, in 1758 Carl Linnaeus named the insect Cicada septendecim in the tenth edition of his Systema Naturae.[10][76]

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 10 feet (3 m) below the surface, but that others had reportedly found them 30 feet (9 m) deep.[77]

In 1775, Thomas Jefferson 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 in 1775. He noted that the females lay their eggs in the small twigs of trees while above ground.[78]

The 1780 emergence of the Brood VII cicadas (also known as the Onondaga brood) during the American Revolutionary War, coincided with the aftermath of the military operation known as the Sullivan Expedition 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.[79] 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.[80]

A Brood X Magicicada with abdominal Massospora cicadina infection in Takoma Park, Maryland (May 31, 2021)

In April 1800, Benjamin Banneker, 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".[81] Describing an effect that the pathogenic fungus, Massospora cicadina, has on its host,[82] 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.[83]

In 1845, Dr. D.L. Pharas of Woodville, Mississippi, announced the 13-year periodicity of the southern cicada broods in a local newspaper, the Woodville Republican.[32] In 1858, Pharas placed the title Cicada tredecim in a subsequent article that the newspaper published on the subject 13 years later.[32]

Ten years afterwards, Benjamin Dann Walsh and Charles Valentine Riley authored a paper that the American Entomologist published in December 1868 which also reported the 13-year periodicity of the southern cicada broods. Walsh and Riley "for convenience sake", named the 13-year brood Cicada tredecim, in contradistinction to Cicada septemdecim, the 17-year brood.[32][84]

Walsh's and Riley's paper, which Scientific American reprinted with some revisions in January 1869, illustrated the interior and exterior characteristics of the nymphs' emergence holes and raised turrets.[85] Their articles, which did not cite Pharas' reports, were the first to describe the southern cicadas' 13-year periodicity that received widespread attention.[32] Riley later acknowledged Pharas' work in an 1885 publication on periodical cicadas that he authored.[32][86]

Itch mite bites

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.[87]

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'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.[88] Similar events occurred in Cincinnati after a Brood XIV emergence ended in 2008,[89] in Cleveland and elsewhere in northern and eastern Ohio after a Brood V emergence ended in 2016,[90] in the Washington, D.C., area after a Brood X emergence ended in 2021,[91] and again in the Chicago area after the next Brood XIII emergence ended in 2024.[92]

Use as human food

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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.[93]

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.[94][95] Marlatt wrote in 1907:

The use of the newly emerged and succulent cicadas as an article of human diet has merely a theoretical interest, because, if for no other reason, they occur too rarely to have any real value. There is also the much stronger objection in the instinctive repugnance which all insects seem to inspire as an article of food to most civilized nations. Theoretically, the Cicada, collected at the proper time and suitably dressed and served, should be a rather attractive food. The larvae have lived solely on vegetable matter of the cleanest and most whole-some sort, and supposedly, therefore, would be much more palatable and suitable for food than the oyster, with its scavenger habit of living in the muddy ooze of river bottoms, or many other animals which are highly prized and which have not half so clean a record as the periodical Cicada.[95]

Notes

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References

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Further reading

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Periodical cicadas, belonging to the genus Magicicada, are of in the family , native exclusively to eastern , and are distinguished by their extraordinary life cycles of either 13 or 17 years, during which nymphs remain underground feeding on root xylem before emerging en masse as adults in synchronized . These emergences, occurring periodically across specific geographic regions, involve densities reaching up to 1.5 million individuals per acre, serving as a predator-satiation strategy to overwhelm natural enemies. Adults feature striking black bodies, red eyes, and orange-tinged wing veins, with males producing species-specific choruses through organs to attract mates. The life cycle of periodical cicadas is among the longest of any , with nymphs hatching from eggs laid in tree twigs and burrowing into shortly after, where they develop over 13 or 17 years by sucking fluids from . is triggered when temperatures at about 8 inches depth reach approximately 64°F (18°C), typically in late spring from to , depending on and . Upon surfacing, nymphs molt into winged adults that live only 3–6 weeks, during which time females oviposit 400–600 eggs into slits in pencil-thin branches, potentially causing minor damage known as "flagging." Nymphs of the next generation hatch in 6–10 weeks, drop to the ground, and begin the cycle anew. The seven Magicicada species are divided into three species groups—decula, cassini, and decim—with three species exhibiting 17-year cycles (generally in northern ranges) and four showing 13-year cycles (more common in southern and midwestern areas), though distributions overlap in parts of the central U.S. These form 15 distinct , labeled I–XVII for 17-year cycles and XVIII–XXX for 13-year cycles, each brood emerging on a unique 13- or 17-year schedule in predefined regions, ensuring no two broods overlap temporally. Genetic studies indicate that broods arose through periodic shifts in cycle length, with the originating around 3.9 million years ago. Ecologically, periodical cicadas play a key role in forest ecosystems by aerating through their burrows and providing a massive, periodic food source for birds, mammals, and other predators, which boosts predator populations during years. While harmless to humans—they neither bite nor sting—their mass appearances have cultural significance, appearing in indigenous oral traditions, and occasional "stragglers" (off-cycle emergences) can occur but in low numbers insufficient for successful . No effective chemical controls are typically needed due to their brief adult phase and natural decline post-oviposition, though netting may protect young trees in affected areas.

Physical Characteristics

Morphology and Identification

Periodical cicadas exhibit a distinctive morphology adapted to their subterranean and arboreal lifestyles. Adult individuals measure 2.5 to 3.5 cm in length, possessing a robust build with broad heads, large compound eyes that are typically but can vary to orange or , three simple ocelli positioned on the vertex, and short bristle-like antennae. Their wings are and span approximately 3 to 8 cm when fully extended, often held roof-like over the at rest, featuring prominent orange or reddish veins and a characteristic "W"-shaped marking near the tips of the forewings. Sexual dimorphism is evident in the abdominal structures related to and communication. Males possess tymbals—paired ribbed membranes on the sides of the first abdominal segment—that enable sound production through rapid vibration, while females lack these organs but have a robust, pointed terminating in a sword-like used to slit bark for egg deposition. The overall body is predominantly black, providing against trunks, with the eyes and wing veins offering key visual cues. Nymphs, the immature stage, are wingless and adapted for underground life, measuring 2.5 to 5 cm when mature, with a pale, - or crayfish-like appearance. They feature strong, rake-like forelegs equipped with spines and notches for efficient burrowing through , while the hind legs are suited for propulsion in their subterranean tunnels. Identification of periodical cicadas relies on a of morphological traits that distinguish them from cicadas and among . Key features include the robust body size, striking eye coloration, and orange-tinged wing veins, with variations such as larger body size in Magicicada septendecim (up to 3.5 cm) and more pronounced orange venation compared to smaller like M. cassini. The presence of three ocelli and the specific wing vein patterns further aid in confirming affiliation, though eye color polymorphisms (e.g., rare black-eyed individuals) can occur naturally across populations.

Coloration and Sexual Dimorphism

Periodical cicadas of the Magicicada exhibit a striking coloration dominated by a glossy black , complemented by vivid to orange compound eyes, orange tarsi on the legs, and orange venation in the translucent wings. The often features orange markings that vary significantly among , serving as key visual traits alongside their overall robust, cylindrical body form. These color elements contribute to against forest floors and during emergences, while also aiding in recognition. Species-specific variations in coloration are most evident in the extent and pattern of orange pigmentation on the abdominal venter. The Decim group (M. septendecim, M. neotredecim, M. tredecim) displays broad orange stripes across most abdominal segments, often with an additional orange patch posterior to the eyes. In contrast, the Decula group (M. septendecula, M. tredecula) has narrower, more defined orange stripes confined to fewer segments, lacking the postocular orange spot. The Cassini group (M. cassini, M. tredecassini) is notably more melanistic, with a fully abdomen devoid of orange stripes, though rare individuals in western populations may show faint yellowish ventral marks. is uniformly red in typical individuals, but natural genetic variations produce rare white- or blue-eyed forms, estimated at about 1 in 1,000 emergences, without altering other pigmentation. Sexual dimorphism in periodical cicadas primarily manifests in body size, with females consistently larger than males across all , a pattern linked to the demands of egg production and oviposition. For instance, in M. septendecim, females average longer forewing lengths than males, enhancing their durability for laying up to 600 . Coloration remains largely uniform between sexes within species, though subtle differences in orange intensity may occur due to individual variation rather than consistent dimorphism. These coloration patterns play a crucial role in field identification, allowing rapid distinction among sympatric species during mass emergences. For example, the broad orange abdominal stripes of M. septendecim—known as the "Pharaoh" type—contrast sharply with the stripe-less black abdomen of M. cassini, enabling observers to differentiate them without relying solely on song or size. Such visual markers are particularly useful in mixed-brood areas, where 13-year and 17-year species may co-occur.

Taxonomy and Evolution

Species Classification

Periodical cicadas belong to the genus Magicicada within the family Cicadidae, encompassing seven recognized species endemic to eastern North America. These species are classified into two main groups based on their life cycle durations: three species with a 17-year cycle and four with a 13-year cycle. This taxonomic division reflects parallel evolutionary lineages, with each group containing species that share similar morphological and acoustic traits but differ in periodicity. The classification was initially established through comparative studies of morphology, songs, and life histories, with subsequent refinements identifying additional species. The 17-year species include (Linnaeus, 1758), (Fisher, 1852), and Magicicada septendecula (Alexander and Moore, 1962). The 13-year species comprise Magicicada tredecim (Walsh and Riley, 1868), Magicicada neotredecim (Marshall and Cooley, 2000), Magicicada tredecassini (Alexander and Moore, 1962), and Magicicada tredecula (Alexander and Moore, 1962). These species are further subgrouped into -decim, -cassini, and -decula categories based on shared characteristics such as body size and coloration patterns.
SpeciesGroupCycle LengthKey Distinctions
M. septendecim-decim17 yearsLargest size; broad orange abdominal stripes; characteristic "wee-oh" or "" calling song with a low-pitched followed by a higher one.
M. cassini-cassini17 yearsLacks abdominal stripes; continuous buzzing call with trailing clicks.
M. septendecula-decula17 yearsSmaller size; narrow orange abdominal stripes; similar song to M. cassini but with shorter phrases.
M. tredecim-decim13 yearsSimilar to M. septendecim but smaller; broad stripes; lower-pitched song in overlap zones.
M. neotredecim-decim13 yearsResembles M. tredecim but with darker stripes and higher-pitched song for species recognition.
M. tredecassini-cassini13 yearsNo abdominal stripes; buzzing call akin to M. cassini.
M. tredecula-decula13 yearsNarrow stripes; song similar to M. cassini and M. septendecula.
Key distinctions among species include cycle length, which synchronizes emergences within ; acoustic signals, where males produce species-specific calling to attract mates; and subtle morphological traits such as wing length, body size, and the extent of orange pigmentation on the . For instance, -decim species tend to be larger with more prominent orange markings, while -cassini species exhibit black abdomens and synchronized chorusing behaviors. These features aid identification during mass emergences, though overlap in sympatric areas requires combined analysis of and morphology. Interspecies mating is rare in natural settings due to strong premating isolation via divergent songs and female preferences, maintaining distinct genetic lineages despite occasional experimental hybrids. Magicicada species are generally considered secure at the global level due to their wide distribution and large emergence populations, though some broods have gone extinct and others face threats from habitat loss and fragmentation. At least three broods (I, XI, and parts of others) are considered extinct due to habitat loss.

Phylogenetic Relationships

The genus Magicicada forms a monophyletic within the family , as evidenced by analyses of both mitochondrial and nuclear genetic markers. The periodical cicadas are organized into three distinct groups—Decim, Cassini, and Decula—each typically comprising one 17-year and one or two 13-year . Phylogenetic reconstructions indicate that the Decim group occupies a basal position, serving as the lineage to a formed by the Cassini and Decula groups. The divergence between Decim and this is estimated at approximately 3.9 million years ago, while the split between Cassini and Decula occurred around 2.5 million years ago. Within each species group, the 17-year lineages are basal, with the 13-year species derived from ancestors resembling modern 17-year forms, such as M. septendecim in the Decim group. For instance, in Decim, M. neotredecim (13-year) branches as the sister taxon to M. septendecim (17-year), while M. tredecim (13-year) represents a more basal divergence within the group. Similar derived positions for 13-year species are observed in Cassini (M. tredecassini from M. cassini) and Decula (M. tredecula from M. septendecula). These relationships highlight parallel evolutionary patterns across groups, where 13-year cycles emerged independently from 17-year ancestors. Genetic evidence from (including COI, COII, and tRNA-Leu) and nuclear loci (such as 18S rRNA, wingless, EF1-α, and ) underscores the close affinities between 13-year and 17-year taxa within groups, with shared haplotypes and low sequence indicating recent origins and occasional . (AFLP) markers further confirm differentiation primarily by life cycle and geographic region rather than strict boundaries. analyses, employing substitution rates of approximately 0.024 substitutions per site per million years and calibrated against the most recent common ancestor (MRCA) of the groups, date the initial 13-year s to the Pleistocene epoch. The earliest such split, within Decim, is timed to about 0.5 million years ago, with subsequent events in Cassini and Decula occurring 0.1–0.2 million years ago, aligning with glacial-interglacial fluctuations that likely facilitated sympatric through allochronic isolation.

Evolutionary Origins and Speciation

Periodical cicadas of the genus Magicicada evolved in eastern from ancestors with more variable, annual-like life cycles, with estimates placing the around 3.6 million years ago. This is supported by analyses showing sequence consistent with a mid-Pliocene origin. The transition to strict periodicity likely occurred through the rigidification of developmental cycles, where variable emergence times in progenitor populations became synchronized and extended, possibly in response to climatic cooling during Pleistocene glacial periods that reduced nymphal growth rates and prolonged . Simulations suggest this fixation of long cycles could have taken 10,000 to 30,000 years under selective pressures favoring synchronization. The adoption of prime-numbered cycles—13 years in southern populations and 17 years in northern ones—represents an adaptive refinement following the establishment of periodicity. These odd, prime lengths minimize periodic overlap with predators possessing shorter cycles (e.g., 2–4 years), reducing the frequency of synchronized attacks and enhancing survival through intermittent during mass emergences. Modeling indicates that such cycle lengths provide an evolutionary advantage by avoiding regular coincidences with potential predators' life histories, contributing to the persistence of these traits over millions of years. Speciation within Magicicada is driven primarily by allochronic isolation, where shifts in cycle length (e.g., from 17 to 13 years via 4-year accelerations) create temporal barriers to mating, leading to reproductive in songs and morphologies. Hybridization remains rare due to the asynchrony of broods, which prevents between differing cycles despite occasional spatial overlap. Recent genetic studies reveal a rapid post-glacial radiation, with phylogeographic patterns indicating population expansions from southern refugia around 10,000 years ago, coinciding with warming and northern range recolonization. Fossil evidence for Magicicada is scarce, but related cicada lineages have records from the (approximately 8–5 million years ago), underscoring the ancient origins of the group's developmental strategies.

Life Cycle

Nymphal Development

Periodical cicada nymphs spend the majority of their life cycle—either 13 or 17 years, depending on the species—underground, progressing through five distinct stages of development. This prolonged nymphal phase allows for gradual growth in the subterranean environment, with molting events synchronized across individuals within a brood based on physiological age, which is determined by the accumulation of thermal units or degree-days above a developmental threshold. The exact mechanism for tracking this cumulative temperature remains under study, but it ensures that nymphs reach maturity in unison, minimizing predation risks during the vulnerable transition to adulthood. During this underground period, nymphs sustain themselves by feeding on xylem sap drawn from the roots of deciduous trees and shrubs via a specialized proboscis that pierces vascular tissues. fluid is nutrient-poor, consisting primarily of water, minerals, and trace , which necessitates large volumes of intake and contributes to the exceptionally slow growth rate observed over their multi-year development. This feeding strategy ties nymphal survival directly to the health and availability of host root systems in forested or wooded habitats. Nymphs are adept burrowers, using robust forelegs to excavate tunnels typically at depths of 5 to 60 cm (2 to 24 inches), where they remain close to root networks for feeding access. In areas with high , particularly during rainy periods, they construct mud or turrets—small soil mounds up to 15 cm tall—above their exit holes to prevent flooding and maintain within the . Burrowing depth and the frequency of chimney building are influenced by local levels and the density of , with denser root zones allowing shallower tunnels and more efficient . Toward the end of their development, environmental cues such as rising temperatures play a critical role in signaling preparation for the final molt. In the spring of their year, fifth-instar nymphs initiate the or enlargement of vertical exit tunnels a few weeks before , positioning themselves just below the surface. occurs when temperatures at a depth of about 20 cm (8 inches) reach approximately 18°C (64°F). This threshold ensures synchronized activity in response to seasonal warming, optimizing conditions for the transition to the adult stage.

Emergence and Adult Behavior

Periodical cicadas emerge en masse from the when temperatures at a depth of 7-8 inches reach approximately 64°F (18°C), typically after a warming period in . Nymphs, having completed their underground development, exit burrows primarily after sunset, climb nearby such as trunks or shrubs, and undergo their final molt to become adults. This transformation produces pale, soft-bodied teneral adults that require 4-6 days to fully harden and darken before engaging in reproductive activities. As adults, periodical cicadas have a brief lifespan of 4-6 weeks, during which their primary focus is reproduction. Males aggregate in choruses, producing species-specific songs through tymbal vibrations to attract females; these choruses can reach sound levels exceeding 90 decibels. Once mated, females use their ovipositors to carve slits into living twigs of deciduous trees, creating Y-shaped nests where they deposit 20-30 eggs each, potentially laying up to 600 eggs across dozens of such slits. The eggs hatch after 6-10 weeks, and the resulting nymphs drop to the ground, burrowing into the soil to begin their long subterranean phase. Following oviposition, adults undergo rapid , with both males and females dying within weeks of , often leaving behind empty exoskeletons () on vegetation as remnants of the event.

Broods and Distribution

Brood Cycles and Synchronization

Periodical cicadas are organized into distinct broods, each representing a synchronized population that emerges on a precise 13- or 17-year cycle. The brood numbering system, established by entomologist C. L. Marlatt in 1902, designates 30 possible broods using I through XXX, with I–XVII reserved for 17-year cycles and XVIII–XXX for 13-year cycles. Currently, 12 broods are active on the 17-year cycle, including notable examples like , which emerged across 15 states in 2021, and Brood XIV, which emerged in 2025 in parts of the eastern U.S. In contrast, only three broods remain active on the 13-year cycle: Broods XIX, XXII, and XXIII, with —the largest—emerging in 2024 across 13 southern and midwestern states. Several broods are considered extinct or empty, such as the 17-year Brood XI (last recorded in 1954) and various 13-year broods like XXI (last in 1870), due to habitat loss and other factors. Within each brood, all individuals emerge synchronously after exactly 13 or 17 years underground, a phenomenon driven by precise developmental timing that ensures mass appearances typically spanning late to early , depending on and soil temperature. This synchronization is remarkably tight, with populations in a given region emerging en masse to overwhelm predators through sheer numbers, though off-cycle emergences known as stragglers—individuals appearing one to four years early or late—do occur rarely, often in low densities and with reduced , rendering them non-viable for sustaining the brood. Broods lack sub-broods or internal divisions; instead, their separation is maintained by geographic isolation, where populations in adjacent areas are offset by years in their cycles—for instance, Brood II (next in 2030) and Brood III (next in 2031) occupy overlapping but distinct ranges in the Northeast, preventing overlap and interbreeding. Rare dual emergences highlight the independence of these cycles, as 13- and 17-year broods occasionally coincide due to their prime-number periodicity. In 2024, (17-year) in the Midwest and (13-year) in the South emerged simultaneously across overlapping regions like and , creating unprecedented densities estimated in the trillions and marking the first such event since 1803. These overlaps amplify ecological impacts but do not disrupt long-term brood integrity, as geographic barriers limit widespread hybridization.

Geographical Range

Periodical cicadas are endemic to , with their primary range spanning the eastern and , from northern Georgia northward to Iowa and generally confined to areas east of the . This distribution encompasses states including , , , , , , , New York, , , , , , and , among others. The range is discontinuous and patchy, shaped by historical events such as the Pleistocene glaciation, which restricted populations to southern refugia during ice ages, and later recolonization that left gaps in formerly glaciated northern territories. Additionally, widespread during European settlement in the 18th and 19th centuries fragmented suitable habitats, leading to local extirpations and further discontinuities in brood distributions. These insects exhibit strong habitat preferences for deciduous woodlands, where underground nymphs feed on the xylem sap of roots from hardwood trees such as oaks (Quercus spp.), maples (Acer spp.), hickories (Carya spp.), and willows (Salix spp.). They thrive in areas with well-drained, loamy soils that support these host plants and allow for nymphal burrowing up to 2 meters deep. Periodical cicadas are notably absent from coniferous forests, arid or semi-arid regions like the southwestern deserts, and intensively agricultural or urbanized landscapes lacking mature deciduous vegetation. Over time, the overall range has contracted due to habitat loss from and since European colonization. may influence future distributions, with warmer temperatures potentially enabling earlier emergences and slight expansions into marginal habitats, though ongoing poses a countervailing risk of further decline. In regions of , such as parts of and the Midwest, 13-year and 17-year coexist spatially but emerge asynchronously, preventing direct temporal overlap in mass events.

Mapping Emergence Locations

Mapping emergence locations of periodical cicadas relies on specialized databases and (GIS) tools that compile historical records, verified observations, and projected cycles to visualize brood distributions across the . The of Connecticut's Periodical Cicada Information Pages maintain an interactive database featuring point-based maps derived from field-verified presence and absence data, avoiding generalized boundaries to reflect the patchy nature of emergences. These maps use symbols to denote confirmed cicada occurrences (e.g., cicada icons), absences (red symbols), and historical records from sources like Simon (1988) and Marlatt (1923), allowing users to assess density gradients and potential overlap zones between broods. The USDA Forest Service provides complementary GIS layers through its Enterprise , offering county-level polygons for active that integrate historical data with expected future cycles, such as 13- and 17-year schedules. These static and dynamic maps, updated as of with references to seminal works like Marlatt (1907) and Koenig et al. (2011), facilitate by highlighting the geographic extent of and their relationships to habitat loss. For instance, Brood X's 2021 was mapped across a vast area including dense populations in and , where interactive UConn visualizations showed high-density choruses in urban-adjacent woodlands and lower densities near state borders. The concurrent 2024 emergences of Broods XIII and XIX exemplified dual-brood mapping challenges, with GIS layers depicting Brood XIII's core in —verified through Stannard (1975) delineations and recent citizen reports—and Brood XIX's expansive footprint extending into Missouri's Ozark regions, where density gradients tapered from full emergences in river valleys to stragglers in peripheral counties. The 2025 emergence of Brood XIV occurred in four distinct patches as projected via UConn's point maps, including a large central area from to southern and smaller disjunct populations in central , with visualizations emphasizing overlap risks near boundaries shared with . Historical mapping reveals shifts due to , as seen with extinct Brood XXI, last recorded in 1870 along Florida's Valley and now mapped as absent using blue symbols for low-certainty historical sites, linking its decline to and . apps and protocols, such as those outlined by Cooley et al. (2013), have enhanced accuracy by georeferenced photos, audio recordings, and density assessments (categorized from stragglers to full choruses), refining older maps that overestimated extents by including off-cycle individuals. These tools collectively support predictive modeling for future emergences, with interactive features on platforms like UConn's site enabling users to zoom into density hotspots and export data for local risk assessments.

Ecology and Interactions

Predator Satiation Mechanism

Periodical cicadas employ a strategy through synchronized mass emergences, where billions of adults surface simultaneously across large areas, overwhelming the consumption capacity of predators such as birds and small mammals. This tactic ensures that, despite heavy initial predation, a sufficient number of individuals survive to reproduce, as predators become temporarily satiated and unable to consume the entire population. Empirical evidence supports this mechanism, with studies documenting temporary booms in predator populations following emergences due to the abundant food supply. For instance, during the 2021 Brood X emergence, over 80 bird species shifted their diets to cicadas, leading to higher nestling survival rates and subsequent increases in populations of species like blue jays and grackles one to three years later. The mathematical foundation of satiation relies on density-dependent predation models, such as the Holling Type II functional response, where predator intake rate increases with prey density but plateaus at a satiation threshold beyond which additional prey have minimal impact on consumption. Survival rates become positively density-dependent above critical emergence densities, typically exceeding 1 million individuals per acre, allowing a substantial fraction of the brood to evade predation. For example, at high densities during the 1985 Brood IX emergence, avian predators consumed only 15-40% of available cicadas after initial satiation, compared to near-total predation at lower densities. This strategy is most effective against generalist predators that opportunistically feed on cicadas but is less reliable against specialized predators adapted to exploit them. Additionally, off-schedule "straggler" cicadas emerging outside the main brood are highly vulnerable, facing predation rates up to 20 times higher than synchronous individuals due to insufficient numbers for satiation.

Ecosystem Impacts

Periodical cicada emergences deliver a massive pulse of nutrients to forest ecosystems through the decomposition of adult carcasses, which can number in the millions per (up to approximately 3.7 million) and contain approximately 10% by dry weight, exceeding typical litter. This influx enriches with and , elevating plant-available nutrient levels and stimulating microbial activity that enhances nutrient cycling. For instance, studies have documented increased and availability in soils following emergences, leading to higher foliar concentrations in plants, with some species showing up to a 20% increase in content. While female cicadas cause localized tree damage by slashing slits into twigs and branches during oviposition, resulting in branch dieback known as "flagging" where affected tips wilt and die, this injury is typically superficial in mature forests and does not compromise overall health. The nutrient enrichment from decomposing adults often outweighs these effects, promoting aboveground plant biomass growth by up to 50% in some cases and improving long-term forest productivity through enhanced . Trophically, the enormous input—equivalent to several tons per —triggers surges in predator populations, with birds and small mammals like exhibiting increased and abundance during years due to the superabundant source. Over 80 shift behaviors to exploit cicadas, temporarily reducing predation on other and causing herbivore populations, such as caterpillars, to double in density, which in turn boosts leaf herbivory on trees like oaks. This dominance temporarily disrupts herbivore dynamics, as the cicada pulse overshadows alternative resources for generalist consumers. Long-term, these events enhance biodiversity by rewiring food webs and providing sporadic boosts to multiple trophic levels, with 2023 research demonstrating that the biomass influx acts as a "trophic ," altering community interactions for years afterward, such as reduced production in oaks due to heightened herbivory. years thus foster greater ecological connectivity, supporting diverse wildlife while cycling nutrients that sustain resilience over decadal cycles. Post-2024 studies on the dual Broods XIII and XIX continue to investigate similar trophic and nutrient effects, with results expected to align with patterns observed in prior events like .

Parasites, Pathogens, and Pests

Periodical cicadas are primarily affected by the fungal pathogen Massospora cicadina, which infects adults during emergence and induces altered behaviors often described as "zombie-like." This fungus replaces the cicada's abdomen and genitals with a mass of spores, rendering the host sterile while manipulating its actions to facilitate spore dispersal through continued mating attempts, including hypersexual behavior in males that mimic female signals to attract others. In dense broods, infection rates can reach up to 23%, with higher prevalence in areas of high cicada density, though asymptomatic carriers also contribute to spread. Other parasites, including protozoans and nematodes, occasionally infect periodical cicadas but exert limited overall impact due to the adults' brief lifespan of 2–5 weeks. Nematodes such as those in the genus Mermis can parasitize nymphs or adults, reducing host longevity by feeding internally, while like gregarine species may cause sublethal effects on gut function; however, these are rare and do not significantly regulate population sizes given the synchronized mass emergence. Mites, particularly Pyemotes herfsi, attach to adults and feed on , shortening lifespan but affecting only a small fraction of individuals in most broods. As pests, periodical cicadas pose minor threats mainly through egg-laying damage to trees, where females use their ovipositors to slit twigs of young saplings, orchards, and shrubs, causing wilting and dieback known as "flagging." This is most concerning in commercial orchards and nurseries, potentially killing branches on trees under 10 feet tall, though mature forests recover quickly without intervention. There are no significant human health risks from periodical cicadas, as they lack , do not bite, and their cast skins or remains pose no toxicological concerns. Recent 2024 studies on cicadas revealed that bacterial diversity varies by species and location across forest preserves, with higher diversity linked to lower Massospora cicadina infection rates, suggesting a potential role in resistance through microbial or immune modulation. These findings highlight how environmental factors influence microbial communities, which may buffer against parasitic pressures during vulnerable emergence periods.

Symbiotic Associations

Periodical cicadas maintain mutualistic relationships with microbial symbionts that are essential for their survival on a nutrient-poor diet of sap during the prolonged nymphal stage. The primary bacterial endosymbionts, Sulcia muelleri and Hodgkinia cicadicola, reside in specialized bacteriomes within the cicada's gut and synthesize essential and vitamins absent or scarce in xylem fluid. These ancient co-obligates enable the cicadas to extract sufficient from over 13 or 17 years underground, a that has co-evolved over millions of years to support their unique life cycles. Without these symbionts, the cicadas could not complete their development, highlighting the intimate integration of host and microbe genomes for metabolic provisioning. Recent analyses of the gut s in periodical cicadas reveal species-specific variations that influence nutritional efficiency and host fitness. During the 2024 emergence of , sampling across ecologically distinct sites in showed that Sulcia and Hodgkinia dominated profiles but exhibited differences in relative abundance and associated taxa among Magicicada species, such as M. septendecim and M. cassini. These variations, potentially shaped by local environmental factors and host genetics, underscore how microbiome composition adapts to support digestion in diverse populations. Beyond the core endosymbionts, transient gut contribute to secondary metabolic functions, though the obligates remain pivotal for long-term nutrient supplementation. Fungal symbionts also play a role in cicada , particularly in lineages where bacterial partners are reduced or absent, providing an alternative pathway for essential acquisition. Yeast-like fungal symbionts (YLS), harbored in fat bodies or bacteriomes, supply and , compensating for gaps in bacterial provisioning and enhancing overall host resilience during root-feeding. In periodical cicadas, such fungal associations may indirectly bolster host plant through endophytic fungi in , which improve uptake and stress tolerance in hosts like oaks, thereby sustaining the resources critical for nymphal development. These fungal partnerships, while less dominant than bacterial ones in Magicicada, illustrate the flexibility of symbiotic networks in enabling extended subterranean life. A minor reciprocal interaction occurs between emerging adult periodical cicadas and , where cicadas excrete honeydew-like fluids from feeding that serve as a source for . In response, may deter certain herbivores from host trees, indirectly benefiting cicada oviposition sites, though this mutualism is opportunistic and not as structured as in aphid-ant systems. Such exchanges contribute to broader trophic dynamics but remain peripheral to the cicadas' primary microbial dependencies. The evolutionary history of these symbioses reveals deep co-speciation, with bacterial and fungal partners undergoing genome streamlining to match the cicadas' extreme life cycles. Ancient acquisitions of Sulcia and Hodgkinia, dating back over 100 million years in Auchenorrhyncha lineages, have fragmented in long-cycle species like periodical cicadas, leading to population-level symbiont diversity that supports prolonged nymphal dormancy. No obligate symbionts beyond the core bacteria have been identified as universally required, but the system's modularity—evident in recurrent fungal recruitments—has facilitated the evolution of 13- and 17-year periodicities by optimizing nutrient extraction from impoverished diets. This co-evolutionary framework underscores how symbioses underpin the ecological success of periodical cicadas.

Human Aspects

Historical Observations

Native American oral histories document periodical cicadas as recurring natural phenomena, with accounts of their massive emergences predating European contact in the 1600s; these recognized the insects' synchronized cycles, distinguishing them from annual species and integrating observations into cultural narratives across eastern North American tribes. The earliest written European observations date to 1634, when governor William Bradford described swarms of "flies" emerging en masse from the soil in , likening them to biblical plagues but noting their 17-year recurrence based on local reports. By the late , naturalist documented a Brood I emergence in 1800 through detailed engravings and museum displays in , capturing the insects' life stages and contributing to early visual records of their . In the mid-18th century, Swedish naturalist Pehr Kalm's fieldwork in 1748–1749 provided the first systematic scientific study, confirming the 17-year cycle through observations in New Jersey and predicting future emergences that were verified in 1766. Also in the 18th century, African American astronomer and naturalist Benjamin Banneker documented periodical cicada emergences in Maryland and predicted future cycles based on local observations, contributing to early understanding of their periodicity. During the 19th century, entomologists Benjamin D. Walsh and Charles V. Riley advanced knowledge through their 1868 study of Brood X, producing the first comprehensive brood maps based on field reports from across the Midwest and East, which delineated geographical distributions and cycles. Misconceptions persisted, with periodical cicadas often called "locusts" due to their swarming behavior, until C. L. Marlatt's 1907 U.S. Department of Agriculture bulletin clarified their true identity as harmless , not destructive , while expanding on distribution and with updated maps. Key events included accurate predictions of emergences since Kalm's work, demonstrating the reliability of the 13- and 17-year cycles; however, habitat loss began impacting broods, with Brood XI showing signs of decline by the early and failing to emerge after 1954, marking it as extinct due to in .

Culinary and Cultural Uses

Periodical cicadas are and have been consumed by humans in various cultures, particularly when harvested as tenerals shortly after molting, when their exoskeletons are soft and easier to digest. Preparation methods include blanching to remove potential from , followed by , , or , which yields a nutty flavor. Regional recipes, such as cicada tacos from Appalachian and Midwestern traditions, involve marinating the in lime and chili before and serving in corn tortillas with toppings like and cilantro. These pose a low risk for most , though individuals with shellfish allergies should exercise caution due to structural similarities in ; no major toxicity has been reported when properly prepared. Nutritionally, periodical cicadas offer high protein content, approximately 50% by dry weight, along with essential minerals, vitamins, and low levels of fat and , making them comparable to lean meats like . Their abundance during supports sustainable harvesting potential as an alternative protein source, requiring fewer resources than traditional and contributing to reduced environmental impact. For instance, a single can produce billions of individuals, allowing for opportunistic collection without threatening populations. In cultural contexts, periodical cicadas have often been conflated with biblical locusts, symbolizing plagues or , as seen in interpretations of plagues where early European settlers mistook emergences for the described swarms. This association persists in , with some Native American groups viewing cicadas as harbingers of renewal or immortality due to their underground life and dramatic rebirth. Modern media has amplified their symbolic role, particularly during the 2024 dual-brood emergence dubbed the "cicadapocalypse," which generated widespread hype in outlets portraying as an overwhelming natural spectacle. Emergence years often inspire community festivals, such as the 2021 Brood X Cicadafest events in and , featuring educational programs, art exhibits, and culinary demonstrations to celebrate the phenomenon.

Recent Research and Conservation

The 2024 emergence marked a rare dual event involving (17-year cycle) and (13-year cycle), the first major overlap since 1803, spanning 221 years and covering parts of the Midwest and Southeast . This synchrony provided an unprecedented opportunity to study inter-brood interactions, with researchers documenting amplified trophic boosts as the combined —estimated in the trillions—disrupted webs, prompting shifts in avian foraging behaviors across over 80 bird species and enhancing predator populations for years post-emergence. analyses from samples revealed significant variation by species and location, with distinct bacterial communities in ecologically diverse preserves, potentially influencing host fitness and adaptation during mass emergences; similar investigations into highlighted symbiotic microbial roles in nutrient cycling amid the heightened ecological pulse. Climate change poses emerging threats to periodical cicada life cycles, with rising temperatures likely advancing timing by altering warmth thresholds and potentially desynchronizing through accelerated development. Evolutionary models suggest that prolonged warming could shift 17-year toward 13-year cycles or cause straggling, disrupting the precise synchronization that underpins . Habitat suitability projections indicate northward range expansions for some , with single-generation shifts already documented at rates of several kilometers per decade, potentially leading to 10-20% distributional changes by 2100 under moderate warming scenarios. Conservation efforts emphasize preservation, as periodical cicadas depend on contiguous eastern forests; initiatives in fragmented landscapes have been linked to recovery by restoring woodland connectivity essential for brood persistence. Monitoring programs target locally extinct or contracting broods, such as Brood XI (last recorded in 1954) and Brood VII (historically diminished in ), using historical records and field surveys to assess revival potential amid habitat loss. platforms like have proven vital, aggregating thousands of observations during the 2024 emergences to map distributions, detect stragglers, and inform predictive models for future broods. Genomic research has advanced understanding of cycle evolution, with a 2024 chromosome-level assembly of Magicicada septendecula revealing genetic mechanisms underlying periodical timing and , including reduced genomes in bacterial partners that stabilize long underground development. Papers from 2025 explore multigenerational trends, analyzing data across 1987, 2004, and 2021 emergences to quantify density declines in urban habitats and phenological advances linked to warming. Concurrent studies on pathogen dynamics highlight the fungal pathogen , synchronized to cycles, whose spore transmission surges during mass events; profiling from 2024 samples further elucidates how microbial communities modulate infection resistance over generations.

References

  1. https://extension.[illinois](/page/Illinois).edu/blogs/commercial-fruit-and-vegetable-growers/2024-02-20-considering-eating-periodical-cicada
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