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Periodical cicadas

Periodical cicadas, belonging to the genus Magicicada, are seven species of insects in the family Cicadidae, native exclusively to eastern North America, 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 broods.^[1]^[2] 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.^[1] Adults feature striking black bodies, red eyes, and orange-tinged wing veins, with males producing species-specific choruses through tymbal organs to attract mates.^[1]^[3] The life cycle of periodical cicadas is among the longest of any insect, with nymphs hatching from eggs laid in tree twigs and burrowing into soil shortly after, where they develop over 13 or 17 years by sucking fluids from deciduous tree roots.^[2]^[3] Emergence is triggered when soil temperatures at about 8 inches depth reach approximately 64°F (18°C), typically in late spring from April to June, depending on latitude and weather.^[1]^[2] 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 tree damage known as "flagging."^[3]^[2] Nymphs of the next generation hatch in 6–10 weeks, drop to the ground, and begin the cycle anew.^[3] 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.^[1]^[2] These species form 15 distinct broods, labeled Roman numerals 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.^[3]^[2] Genetic studies indicate that broods arose through periodic shifts in cycle length, with the genus originating around 3.9 million years ago.^[1] Ecologically, periodical cicadas play a key role in forest ecosystems by aerating soil through their burrows and providing a massive, periodic food source for birds, mammals, and other predators, which boosts predator populations during emergence years.^[1] 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 reproduction.^[1]^[2] 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.^[3]

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 red but can vary to orange or black, three simple ocelli positioned on the vertex, and short bristle-like antennae.^[4]^[5] Their wings are hyaline and span approximately 3 to 8 cm when fully extended, often held roof-like over the abdomen at rest, featuring prominent orange or reddish veins and a characteristic black "W"-shaped marking near the tips of the forewings.^[6]^[4] Sexual dimorphism is evident in the abdominal structures related to reproduction 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 abdomen terminating in a sword-like ovipositor used to slit tree bark for egg deposition.^[4]^[6] The overall body is predominantly black, providing camouflage against tree trunks, with the eyes and wing veins offering key visual cues.^[7] Nymphs, the immature stage, are wingless and adapted for underground life, measuring 2.5 to 5 cm when mature, with a pale, ant- or crayfish-like appearance.^[8] They feature strong, rake-like forelegs equipped with spines and notches for efficient burrowing through soil, while the hind legs are suited for propulsion in their subterranean tunnels.^[8]^[9] Identification of periodical cicadas relies on a combination of morphological traits that distinguish them from annual cicadas and among species. Key features include the robust body size, striking eye coloration, and orange-tinged wing veins, with species variations such as larger body size in Magicicada septendecim (up to 3.5 cm) and more pronounced orange venation compared to smaller species like M. cassini.^[6]^[4] The presence of three ocelli and the specific wing vein patterns further aid in confirming genus affiliation, though eye color polymorphisms (e.g., rare black-eyed individuals) can occur naturally across populations.^[5]

Coloration and Sexual Dimorphism

Periodical cicadas of the genus Magicicada exhibit a striking coloration dominated by a glossy black exoskeleton, complemented by vivid red to orange compound eyes, orange tarsi on the legs, and orange venation in the translucent wings.^[2] The abdomen often features orange markings that vary significantly among species, serving as key visual traits alongside their overall robust, cylindrical body form.^[1] These color elements contribute to camouflage against forest floors and understory vegetation during emergences, while also aiding in species recognition.^[10] 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.^[10] In contrast, the Decula group (M. septendecula, M. tredecula) has narrower, more defined orange stripes confined to fewer segments, lacking the postocular orange spot.^[11] The Cassini group (M. cassini, M. tredecassini) is notably more melanistic, with a fully black abdomen devoid of orange stripes, though rare individuals in western populations may show faint yellowish ventral marks.^[12] Eye color 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.^[5] Sexual dimorphism in periodical cicadas primarily manifests in body size, with females consistently larger than males across all species, a pattern linked to the demands of egg production and oviposition.^[13] For instance, in M. septendecim, females average longer forewing lengths than males, enhancing their durability for laying up to 600 eggs.^[14] Coloration remains largely uniform between sexes within species, though subtle differences in orange intensity may occur due to individual variation rather than consistent dimorphism.^[10] 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.^[15] Such visual markers are particularly useful in mixed-brood areas, where 13-year and 17-year species may co-occur.^[10]

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.^[16] The 17-year species include Magicicada septendecim (Linnaeus, 1758), Magicicada cassini (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.^[10]

Species	Group	Cycle Length	Key Distinctions
M. septendecim	-decim	17 years	Largest size; broad orange abdominal stripes; characteristic "wee-oh" or "pharaoh" calling song with a low-pitched phrase followed by a higher one.
M. cassini	-cassini	17 years	Lacks abdominal stripes; continuous buzzing call with trailing clicks.
M. septendecula	-decula	17 years	Smaller size; narrow orange abdominal stripes; similar song to M. cassini but with shorter phrases.
M. tredecim	-decim	13 years	Similar to M. septendecim but smaller; broad stripes; lower-pitched song in overlap zones.
M. neotredecim	-decim	13 years	Resembles M. tredecim but with darker stripes and higher-pitched song for species recognition.
M. tredecassini	-cassini	13 years	No abdominal stripes; buzzing call akin to M. cassini.
M. tredecula	-decula	13 years	Narrow stripes; song similar to M. cassini and M. septendecula.

Key distinctions among species include cycle length, which synchronizes emergences within broods; acoustic signals, where males produce species-specific calling songs to attract mates; and subtle morphological traits such as wing length, body size, and the extent of orange pigmentation on the abdomen. 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 songs and morphology.^[10]^[17]^[15] 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.^[18]^[19]^[20]^[21]

Phylogenetic Relationships

The genus Magicicada forms a monophyletic clade within the family Cicadidae, as evidenced by analyses of both mitochondrial and nuclear genetic markers.^[22] The periodical cicadas are organized into three distinct species groups—Decim, Cassini, and Decula—each typically comprising one 17-year species and one or two 13-year species. Phylogenetic reconstructions indicate that the Decim group occupies a basal position, serving as the sister lineage to a clade formed by the Cassini and Decula groups. The divergence between Decim and this sister clade is estimated at approximately 3.9 million years ago, while the split between Cassini and Decula occurred around 2.5 million years ago.^[22]^[23] 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.^[24]^[22] Genetic evidence from mitochondrial DNA (including COI, COII, and tRNA-Leu) and nuclear loci (such as 18S rRNA, wingless, EF1-α, and calmodulin) underscores the close affinities between 13-year and 17-year taxa within groups, with shared haplotypes and low sequence divergence indicating recent origins and occasional gene flow. Amplified fragment length polymorphism (AFLP) markers further confirm differentiation primarily by life cycle and geographic region rather than strict species boundaries.^[22] Molecular clock 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 divergences 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 climate fluctuations that likely facilitated sympatric divergence through allochronic mating isolation.^[23]^[22]

Evolutionary Origins and Speciation

Periodical cicadas of the genus Magicicada evolved in eastern North America from ancestors with more variable, annual-like life cycles, with molecular clock estimates placing the most recent common ancestor around 3.6 million years ago. This divergence is supported by mitochondrial DNA analyses showing sequence divergence 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 underground development. 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 predator satiation 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 character displacement in songs and morphologies. Hybridization remains rare due to the asynchrony of broods, which prevents gene flow 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 Holocene warming and northern range recolonization. Fossil evidence for Magicicada is scarce, but related cicada lineages have records from the Miocene (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 instar stages of development.^[25] 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.^[26] 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.^[27] 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.^[28] Xylem fluid is nutrient-poor, consisting primarily of water, minerals, and trace amino acids, 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.^[28] 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.^[29] In areas with high soil moisture, particularly during rainy periods, they construct mud chimneys or turrets—small soil mounds up to 15 cm tall—above their exit holes to prevent flooding and maintain airflow within the burrow.^[30] Burrowing depth and the frequency of chimney building are influenced by local soil moisture levels and the density of tree roots, with denser root zones allowing shallower tunnels and more efficient foraging. Toward the end of their development, environmental cues such as rising soil temperatures play a critical role in signaling preparation for the final molt. In the spring of their emergence year, fifth-instar nymphs initiate the construction or enlargement of vertical exit tunnels a few weeks before emergence, positioning themselves just below the surface.^[31]^[4] Emergence occurs when soil temperatures at a depth of about 20 cm (8 inches) reach approximately 18°C (64°F).^[2] This temperature threshold ensures synchronized activity in response to seasonal warming, optimizing conditions for the transition to the adult stage.^[26]

Emergence and Adult Behavior

Periodical cicadas emerge en masse from the soil when temperatures at a depth of 7-8 inches reach approximately 64°F (18°C), typically after a warming period in late spring.^[1] Nymphs, having completed their underground development, exit burrows primarily after sunset, climb nearby vegetation such as tree trunks or shrubs, and undergo their final molt to become adults.^[1] This transformation produces pale, soft-bodied teneral adults that require 4-6 days to fully harden and darken before engaging in reproductive activities.^[1] As adults, periodical cicadas have a brief lifespan of 4-6 weeks, during which their primary focus is reproduction.^[32] Males aggregate in choruses, producing species-specific songs through tymbal vibrations to attract females; these choruses can reach sound levels exceeding 90 decibels.^[33] 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.^[1] 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.^[1] Following oviposition, adults undergo rapid senescence, with both males and females dying within weeks of emergence, often leaving behind empty exoskeletons (exuviae) on vegetation as remnants of the event.^[1]

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 Roman numerals 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 Brood X, 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 Brood XIX—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.^[34]^[35] 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 April to early June, depending on latitude 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 reproductive success, 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.^[34]^[36] 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, Brood XIII (17-year) in the Midwest and Brood XIX (13-year) in the South emerged simultaneously across overlapping regions like Illinois and Indiana, 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.^[37]^[38]

Geographical Range

Periodical cicadas are endemic to North America, with their primary range spanning the eastern and midwestern United States, from northern Georgia northward to Iowa and generally confined to areas east of the Great Plains. This distribution encompasses states including Alabama, Arkansas, Illinois, Indiana, Kentucky, Maryland, Missouri, New York, North Carolina, Ohio, Pennsylvania, Tennessee, Virginia, and West Virginia, 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 deforestation during European settlement in the 18th and 19th centuries fragmented suitable habitats, leading to local extirpations and further discontinuities in brood distributions.^[39]^[26] 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.^[39]^[6]^[40] Over time, the overall range has contracted due to habitat loss from agricultural expansion and urbanization since European colonization. Climate change may influence future distributions, with warmer temperatures potentially enabling earlier emergences and slight expansions into marginal habitats, though ongoing habitat fragmentation poses a countervailing risk of further decline. In regions of sympatry, such as parts of Oklahoma and the Midwest, 13-year and 17-year broods coexist spatially but emerge asynchronously, preventing direct temporal overlap in mass events.^[39]^[1]^[41]

Mapping Emergence Locations

Mapping emergence locations of periodical cicadas relies on specialized databases and geographic information system (GIS) tools that compile historical records, verified observations, and projected cycles to visualize brood distributions across the eastern United States. The University 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.^[42] These maps use symbols to denote confirmed cicada occurrences (e.g., black 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.^[34] The USDA Forest Service provides complementary GIS layers through its Enterprise Data Warehouse, offering county-level polygons for active broods that integrate historical emergence data with expected future cycles, such as 13- and 17-year schedules.^[35] These static and dynamic maps, updated as of 2013 with references to seminal works like Marlatt (1907) and Koenig et al. (2011), facilitate spatial planning by highlighting the geographic extent of broods and their relationships to habitat loss.^[43] For instance, Brood X's 2021 emergence was mapped across a vast area including dense populations in Ohio and Pennsylvania, where interactive UConn visualizations showed high-density choruses in urban-adjacent woodlands and lower densities near state borders.^[44] The concurrent 2024 emergences of Broods XIII and XIX exemplified dual-brood mapping challenges, with GIS layers depicting Brood XIII's core in northern Illinois—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.^[45]^[46] The 2025 emergence of Brood XIV occurred in four distinct patches as projected via UConn's point maps, including a large central area from northeast Georgia to southern Ohio and smaller disjunct populations in central Pennsylvania, with visualizations emphasizing overlap risks near boundaries shared with Brood XIX.^[47] Historical mapping reveals shifts due to habitat fragmentation, as seen with extinct Brood XXI, last recorded in 1870 along Florida's Apalachicola River Valley and now mapped as absent using blue symbols for low-certainty historical sites, linking its decline to agricultural expansion and urbanization.^[48] Citizen science apps and protocols, such as those outlined by Cooley et al. (2013), have enhanced accuracy by crowdsourcing georeferenced photos, audio recordings, and density assessments (categorized from stragglers to full choruses), refining older maps that overestimated extents by including off-cycle individuals.^[42] 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.^[31]

Ecology and Interactions

Predator Satiation Mechanism

Periodical cicadas employ a predator satiation 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.^[49]^[50] 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.^[49]^[50] 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.^[51]^[52]^[1] 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.^[36]^[53]

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 hectare (up to approximately 3.7 million) and contain approximately 10% nitrogen by dry weight, exceeding typical leaf litter. This influx enriches soil with nitrogen and phosphorus, elevating plant-available nutrient levels and stimulating microbial activity that enhances nutrient cycling. For instance, studies have documented increased ammonium and nitrate availability in soils following emergences, leading to higher foliar nitrogen concentrations in understory plants, with some species showing up to a 20% increase in leaf nitrogen content.^[54]^[55]^[49]^[1] 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 tree 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 soil fertility.^[7]^[56]^[49] Trophically, the enormous biomass input—equivalent to several tons per hectare—triggers surges in predator populations, with birds and small mammals like shrews exhibiting increased reproduction and abundance during emergence years due to the superabundant food source. Over 80 bird species shift foraging behaviors to exploit cicadas, temporarily reducing predation on other insects 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 food resources for generalist consumers.^[49]^[54]^[57] Long-term, these events enhance forest 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 veto," altering community interactions for years afterward, such as reduced acorn production in oaks due to heightened herbivory. Emergence years thus foster greater ecological connectivity, supporting diverse wildlife while cycling nutrients that sustain forest resilience over decadal cycles. Post-2024 studies on the dual Broods XIII and XIX emergence continue to investigate similar trophic and nutrient effects, with results expected to align with patterns observed in prior events like Brood X.^[49]^[58]^[59]

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.^[60]^[61] 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 protozoan infections 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 hemolymph, shortening lifespan but affecting only a small fraction of individuals in most broods.^[62]^[63] 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 injury 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 venom, do not bite, and their cast skins or remains pose no toxicological concerns.^[64]^[65]^[7] Recent 2024 studies on Brood XIII cicadas revealed that bacterial microbiome diversity varies by species and location across northern Illinois forest preserves, with higher diversity linked to lower Massospora cicadina infection rates, suggesting a potential role in pathogen resistance through microbial competition or immune modulation. These findings highlight how environmental factors influence microbial communities, which may buffer against parasitic pressures during vulnerable emergence periods.^[61]

Symbiotic Associations

Periodical cicadas maintain mutualistic relationships with microbial symbionts that are essential for their survival on a nutrient-poor diet of xylem 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 amino acids and vitamins absent or scarce in xylem fluid.^[66] These ancient co-obligates enable the cicadas to extract sufficient nutrition from tree roots over 13 or 17 years underground, a symbiosis that has co-evolved over millions of years to support their unique life cycles.^[67] Without these symbionts, the cicadas could not complete their development, highlighting the intimate integration of host and microbe genomes for metabolic provisioning.^[68] Recent analyses of the gut microbiomes in periodical cicadas reveal species-specific variations that influence nutritional efficiency and host fitness. During the 2024 emergence of Brood XIII, sampling across ecologically distinct sites in northern Illinois 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.^[61] These variations, potentially shaped by local environmental factors and host genetics, underscore how microbiome composition adapts to support xylem digestion in diverse populations.^[69] Beyond the core endosymbionts, transient gut bacteria contribute to secondary metabolic functions, though the obligates remain pivotal for long-term nutrient supplementation.^[66] Fungal symbionts also play a role in cicada nutrition, particularly in lineages where bacterial partners are reduced or absent, providing an alternative pathway for essential nutrient acquisition. Yeast-like fungal symbionts (YLS), harbored in fat bodies or bacteriomes, supply amino acids and carotenoids, compensating for gaps in bacterial provisioning and enhancing overall host resilience during root-feeding.^[70] In periodical cicadas, such fungal associations may indirectly bolster host plant health through endophytic fungi in tree roots, which improve nutrient uptake and stress tolerance in hosts like oaks, thereby sustaining the xylem resources critical for nymphal development.^[71] These fungal partnerships, while less dominant than bacterial ones in Magicicada, illustrate the flexibility of symbiotic networks in enabling extended subterranean life.^[72] A minor reciprocal interaction occurs between emerging adult periodical cicadas and ants, where cicadas excrete honeydew-like fluids from xylem feeding that serve as a food source for ants. In response, ants 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.^[73] Such exchanges contribute to broader trophic dynamics but remain peripheral to the cicadas' primary microbial dependencies.^[74] 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.^[68] 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.^[70] This co-evolutionary framework underscores how symbioses underpin the ecological success of periodical cicadas.^[75]

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 indigenous knowledge systems recognized the insects' synchronized cycles, distinguishing them from annual species and integrating observations into cultural narratives across eastern North American tribes.^[76] The earliest written European observations date to 1634, when Plymouth Colony governor William Bradford described swarms of "flies" emerging en masse from the soil in New England, likening them to biblical plagues but noting their 17-year recurrence based on local reports. By the late 18th century, naturalist Charles Willson Peale documented a Brood I emergence in 1800 through detailed engravings and museum displays in Philadelphia, capturing the insects' life stages and contributing to early visual records of their biology. 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.^[77] 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.^[77] 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.^[78] 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 Hemiptera, not destructive Acrididae, while expanding on distribution and ecology with updated maps.^[79] 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 20th century and failing to emerge after 1954, marking it as extinct due to deforestation in Connecticut.^[80]

Culinary and Cultural Uses

Periodical cicadas are edible 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.^[81] Preparation methods include blanching to remove potential bacteria from soil, followed by roasting, frying, or sautéing, which yields a nutty flavor.^[81] Regional recipes, such as cicada tacos from Appalachian and Midwestern traditions, involve marinating the insects in lime and chili before frying and serving in corn tortillas with toppings like avocado and cilantro.^[82] These insects pose a low allergen risk for most people, though individuals with shellfish allergies should exercise caution due to structural similarities in chitin; no major toxicity has been reported when properly prepared.^[81] Nutritionally, periodical cicadas offer high protein content, approximately 50% by dry weight, along with essential minerals, vitamins, and low levels of fat and cholesterol, making them comparable to lean meats like beef.^[83] Their abundance during emergences supports sustainable harvesting potential as an alternative protein source, requiring fewer resources than traditional livestock and contributing to reduced environmental impact.^[83] For instance, a single emergence 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 divine judgment, as seen in interpretations of the Exodus plagues where early European settlers mistook emergences for the described swarms.^[85] This association persists in folklore, with some Native American groups viewing cicadas as harbingers of renewal or immortality due to their underground life and dramatic rebirth.^[76] Modern media has amplified their symbolic role, particularly during the 2024 dual-brood emergence dubbed the "cicadapocalypse," which generated widespread hype in outlets portraying the event as an overwhelming natural spectacle.^[86] Emergence years often inspire community festivals, such as the 2021 Brood X Cicadafest events in Maryland and Ohio, featuring educational programs, art exhibits, and culinary demonstrations to celebrate the phenomenon.^[87]

Recent Research and Conservation

The 2024 emergence marked a rare dual event involving Brood XIII (17-year cycle) and Brood XIX (13-year cycle), the first major overlap since 1803, spanning 221 years and covering parts of the Midwest and Southeast United States.^[38]^[88] This synchrony provided an unprecedented opportunity to study inter-brood interactions, with researchers documenting amplified trophic boosts as the combined biomass—estimated in the trillions—disrupted forest food webs, prompting shifts in avian foraging behaviors across over 80 bird species and enhancing predator populations for years post-emergence.^[89]^[49] Microbiome analyses from Brood XIII samples revealed significant variation by species and location, with distinct bacterial communities in ecologically diverse forest preserves, potentially influencing host fitness and adaptation during mass emergences; similar investigations into Brood XIX highlighted symbiotic microbial roles in nutrient cycling amid the heightened ecological pulse.^[61]^[90] Climate change poses emerging threats to periodical cicada life cycles, with rising temperatures likely advancing emergence timing by altering soil warmth thresholds and potentially desynchronizing broods through accelerated development.^[91]^[39] Evolutionary models suggest that prolonged warming could shift 17-year broods toward 13-year cycles or cause straggling, disrupting the precise synchronization that underpins predator satiation.^[92] Habitat suitability projections indicate northward range expansions for some broods, 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.^[93]^[94] Conservation efforts emphasize habitat preservation, as periodical cicadas depend on contiguous eastern deciduous forests; reforestation initiatives in fragmented landscapes have been linked to population recovery by restoring woodland connectivity essential for brood persistence.^[39]^[95] Monitoring programs target locally extinct or contracting broods, such as Brood XI (last recorded in 1954) and Brood VII (historically diminished in upstate New York), using historical records and field surveys to assess revival potential amid habitat loss.^[80]^[96] Citizen science platforms like iNaturalist have proven vital, aggregating thousands of observations during the 2024 emergences to map distributions, detect stragglers, and inform predictive models for future broods.^[97] Genomic research has advanced understanding of cycle evolution, with a 2024 chromosome-level assembly of Magicicada septendecula revealing genetic mechanisms underlying periodical timing and symbiosis, including reduced genomes in bacterial partners that stabilize long underground development.^[98] Papers from 2025 explore multigenerational trends, analyzing Brood X data across 1987, 2004, and 2021 emergences to quantify density declines in urban habitats and phenological advances linked to warming.^[99] Concurrent studies on pathogen dynamics highlight the fungal pathogen Massospora cicadina, synchronized to cicada cycles, whose spore transmission surges during mass events; microbiome profiling from 2024 Brood XIII samples further elucidates how microbial communities modulate infection resistance over generations.^[100]^[61]

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Billions of cicadas will emerge in rare 2024 double-brood event
Jan 20, 2024 · Billions of the winged insects will make an appearance across the Midwest and the Southeast, beginning in some places in late April, for a raucous mating ...Missing: hype | Show results with:hype
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Cicada dual emergence brings chaos to the food chain - BBC
May 6, 2024 · ... broods – Brood XIII and Brood XIX – are emerging at the same time in 2024: the first time they have done so simultaneously since 1803. Such ...
[89]
(PDF) Microbiomes of 2024's Periodical Cicada Brood XIII Vary By ...
Oct 21, 2025 · During the 2024 emergence of Brood XIII in northern Illinois, 17-year cicadas were sampled from four ecologically distinct forest preserves.
[90]
Cicadas' cycles may be affected by climate change, scientists suggest
Apr 29, 2024 · The synchronized emergence or periodic cicadas is an evolutionary strategy. But some scientists think global warming could be affecting the insects' cycles.
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DOES CLIMATE CHANGE AFFECT CICADAS?
Apr 26, 2024 · Warming temperatures could cause periodical broods of cicadas to come out earlier in the spring. In fact, it wouldn't be a first-time occurrence ...
[92]
[PDF] Documenting Single-Generation Range Shifts of Periodical Cicada ...
Apr 13, 2021 · Quantifying range shifts, however, is difficult because of the challenge of using point data to infer species ranges (Yalcin and Leroux 2017) ...
[93]
[PDF] Cicadian Rhythms: The 2024 Cicada Emergence
Sep 9, 2025 · In Cicadian Rhythms, Farah Davis recounts the rare 2024 dual emergence of cicada broods XIII and XIX, which appeared weeks early due to climate ...<|separator|>
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Periodical cicada emergence resource pulse tracks forest expansion ...
Jul 12, 2019 · We estimated 35.2 million cicadas emerged across 159 ha in 2015, almost 2× more than the 19.6 million across 98 ha in 1998. The 2015 emergence ...
[95]
(PDF) The historical contraction of periodical cicada Brood VII ...
Aug 6, 2025 · We discuss periodical cicada (Magicicada) Brood VII, which is becoming extinct from much of its historically-reported range in upstate New ...<|separator|>
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2024 Magicicada: Broods XIII and XIX - iNaturalist
These two broods include all seven species of periodical cicadas, providing a rare opportunity to study the species' distribution and interactions.
[97]
Chromosome-Level Genome Assembly and Annotation of a ...
Jan 8, 2024 · We present a high-quality assembly and annotation of the periodical cicada species, Magicicada septendecula (Hemiptera: Auchenorrhyncha: Cicadidae).
[98]
Multigenerational emergence trends of Brood X Magicicada spp ...
Sep 24, 2025 · We investigated trends in population density and emergence phenology of Brood X cicadas across three reproductive events (1987, 2004, and 2021) ...Missing: pathogen | Show results with:pathogen
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Mind Control, Mutilation and Death. The Fungal Fate That Lurks in ...
Jun 13, 2024 · M. cicadina —A Fungal Zombie Maker. M. cicadina is the only known pathogen uniquely synchronized to periodical cicadas' life cycle. Cicada ...<|separator|>