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Cicada

Cicadas are insects belonging to the superfamily Cicadoidea, primarily the family Cicadidae within the order Hemiptera, characterized by their large size—typically 1 to 2 inches (2.5 to 5 cm) long—bullet-shaped bodies, transparent wings, and prominent eyes positioned on the sides of the head.^[1]^[2] They are renowned for the males' loud, species-specific calls produced by vibrating specialized membranes called tymbals, which are amplified by resonant cavities in the abdomen, serving primarily to attract mates.^[1]^[3] Distributed worldwide in tropical and temperate regions, cicadas include more than 3,000 described species, with estimates suggesting over 5,000 including undescribed taxa; nymphs spend most of their lives underground feeding on xylem sap from plant roots before emerging as short-lived adults that feed on twig sap and reproduce.^[1]^[2]^[3] Cicadas are broadly classified into annual (or "dog-day") species, which emerge every summer in overlapping generations after 2 to 5 years underground, and periodical species in the genus Magicicada, which synchronize massive emergences every 13 or 17 years across eastern North America.^[3]^[2] Annual cicadas, such as those in the genus Neotibicen, are often green and black with dark eyes, active from July to September, and produce a continuous buzzing song resembling a buzz-saw.^[2] In contrast, periodical cicadas feature black bodies, reddish eyes, and orange-tinged wings, emerging en masse—sometimes reaching 1.5 million individuals per acre—when soil temperatures hit about 64°F (18°C) in late spring, overwhelming predators through sheer numbers in a phenomenon known as predator satiation.^[3]^[2] Ecologically, cicadas play a key role as herbivores, with nymphs potentially damaging young tree roots over years of feeding, while adult oviposition—where females use saw-like ovipositors to slit twigs and lay up to 600 eggs—can weaken branches, though populations rarely cause widespread harm.^[1]^[3] Harmless to humans, lacking the ability to bite or sting, cicadas have a fossil record with the superfamily dating back to the Triassic period over 200 million years ago, with their periodic emergences inspiring cultural references and scientific study into synchronized behaviors.^[3]^[1]

Etymology and nomenclature

Etymology

The English word "cicada" is derived from the Latin cicada, an onomatopoeic term mimicking the insect's characteristic chirping or buzzing sound produced by vibrating specialized membranes called tymbals.^[4]^[5] This Latin name likely originated as a Mediterranean loanword, possibly influenced by earlier onomatopoeic forms in regional languages that echoed the rhythmic noises of these insects.^[4] In Ancient Greek, the equivalent term was tétrix (τέττιξ), also onomatopoeic, capturing the shrill, repetitive calls of the creatures and used in classical texts to denote similar singing insects. Ancient writers provided early linguistic and descriptive references to cicadas, embedding their names in natural history. Aristotle, in his History of Animals (circa 350 BCE), described the lifecycle of the tétrix, including its transformation from grub to nymph (tettigometra) and adult stages, noting that males produce chirping sounds while females are silent, and distinguishing it from other insects based on its generation and vocalization. Similarly, Pliny the Elder, in Natural History (circa 77 CE), described the cicada in Latin, noting its lifecycle divisions into singing and silent varieties and its habitat preferences, while attributing its name to the audible emissions that define its presence.^[6] Related terms like "locust" in English have often been conflated with cicada due to superficial resemblances in swarming behavior and noise, though their etymologies differ; "locust" stems from Latin locusta, originally denoting a type of grasshopper or even a lobster-like crustacean, without direct phonetic ties to stridulation sounds. This confusion arose historically in English-speaking regions, where early observers applied "locust" broadly to noisy, emergent insects like cicadas, despite the latter's distinct onomatopoeic naming rooted in their specific vocalizations.^[7] The scientific nomenclature of cicadas evolved significantly with Carl Linnaeus's binomial system in Systema Naturae (1758), where he established the genus Cicada for these insects, formalizing their classification based on morphological and auditory traits observed in European species.^[8] This Linnaean framework laid the foundation for the family Cicadidae, formally established by Batsch in 1789, shifting from folk etymologies to a standardized, sound-inspired taxonomy that persists in modern entomology.^[8]

Common names and classification

Cicadas are known by various regional common names that often reflect their distinctive buzzing sounds or local cultural associations. In the southern United States, they are frequently called "jar flies," a term derived from the old English verb "jar" meaning to buzz or vibrate harshly, mimicking the harsh, rattling noise produced by species like those in the genus Neotibicen during late summer.^[9] In France, particularly in Provence, cicadas are referred to as "cigale," a name rooted in Latin cicada and symbolizing good fortune and the vibrant summer landscape, where their songs are an iconic feature of the region's identity.^[10] In Japan, the term "semi" is used, which phonetically evokes the cicadas' shrill calls and holds cultural significance as a symbol of summer and seasonal transience in literature and folklore.^[11] These common names sometimes lead to confusion with other insects, particularly grasshoppers and true locusts, which are species of grasshoppers (Orthoptera: Acrididae) that form swarming plagues. Cicadas, belonging to the order Hemiptera, are often mistakenly called "locusts" in North America due to their mass emergences, fostering public perceptions of them as destructive pests akin to biblical plagues, despite being harmless herbivores that do not swarm or damage crops significantly.^[12] This misnomer persists regionally, affecting awareness and leading to unnecessary fear during periodical outbreaks.^[13] Scientifically, cicadas are classified using binomial nomenclature, a system established by Carl Linnaeus in which each species is denoted by a two-part Latin or Latinized name: the genus (capitalized) followed by the species epithet (lowercase), both italicized, and often accompanied by the authority (the describer's name and publication year). For example, the periodical cicada Magicicada septendecim (Linnaeus, 1758) refers to the 17-year species, with Linnaeus as the authority based on his original description.^[14] The type species for the genus Cicada, which anchors the family Cicadidae, is Cicada orni (Linnaeus, 1758), selected to represent the group's defining characteristics.^[15] The naming of cicadas adheres to the International Code of Zoological Nomenclature (ICZN), which ensures stability and universality in zoological taxonomy. Key principles include priority (the earliest valid name prevails), uniqueness (no two taxa share the same name), and the requirement for names to be in Latin form, with type specimens (holotypes) designated to fix the application of names; for cicadids, this applies to over 3,000 species across 400+ genera, preventing synonymy and resolving disputes through the International Commission on Zoological Nomenclature.

Taxonomy and evolution

Classification and phylogeny

Cicadas belong to the order Hemiptera within the class Insecta, specifically in the suborder Auchenorrhyncha, infraorder Cicadomorpha, superfamily Cicadoidea, and family Cicadidae.^[15] This placement reflects their shared characteristics with other true bugs, such as piercing-sucking mouthparts, while distinguishing them through specialized morphological and behavioral traits like tymbal sound production.^[16] The superfamily Cicadoidea is divided into two families: the relict Tettigarctidae with only two extant species in Australia, and the more diverse Cicadidae, which encompasses the vast majority of cicada species worldwide.^[17] The family Cicadidae includes approximately 3,000 described species distributed across more than 250 genera.^[16] These are organized into five recognized subfamilies: Cicadinae, Cicadettinae (formerly referred to as Tettigettinae in some classifications), Tibicininae, Tettigomyiinae, and Derotettiginae.^[18] Prominent genera exemplify this diversity; for instance, the genus Cicada (type genus of Cicadinae) includes widespread Old World species, Magicicada (in Cicadettinae) comprises the periodical cicadas of eastern North America known for synchronized emergences, and Quesada (in Tibicininae) features large, annual species from the New World tropics.^[17] Tribal divisions within these subfamilies further refine the hierarchy, with over 50 tribes identified based on morphological and molecular evidence.^[17] Phylogenetic analyses of Cicadidae have relied on molecular markers, including 18S rRNA for ribosomal structure and cytochrome c oxidase subunit I (COI) for mitochondrial variation, to reconstruct evolutionary relationships.^[17] These studies confirm that cicadas diverged from other hemipteran lineages, particularly Sternorrhyncha, around 300 million years ago during the late Carboniferous to early Permian, marking the early radiation of Auchenorrhyncha.^[19] Within Cicadoidea, the split between Tettigarctidae and Cicadidae occurred later, likely in the Mesozoic, with Cicadidae diversifying into its modern subfamilies through a combination of continental drift and ecological adaptations.^[19] In the 2020s, taxonomic revisions have refined classifications, particularly for New World species in Tibicininae, integrating genital morphology—such as pygofer and aedeagus structures—with genetic data from multi-locus phylogenies.^[20] A key 2023 study redefined genera like Tibicinoides and Okanagana, transferring multiple species based on molecular evidence showing polyphyly and using morphological traits to resolve rapid radiations in North America.^[20] These updates highlight ongoing efforts to align nomenclature with evolutionary history, reducing paraphyletic groupings in regional faunas.^[20]

Diversity and distribution

Cicadas comprise over 3,000 described species worldwide, belonging to the superfamily Cicadoidea, with the vast majority occurring in tropical and subtropical habitats that provide suitable conditions for their life cycles. Species diversity peaks in tropical regions, where warm climates and abundant vegetation support high richness; for instance, Australia alone harbors more than 200 species across diverse ecosystems, while Southeast Asia features over 150 species, many endemic to forested islands and mainland tropics.^[21] Biogeographic hotspots of endemism include the Indo-Pacific archipelagoes and the Americas, where unique evolutionary radiations have produced specialized assemblages, such as the seven species of the genus Magicicada—all periodical cicadas—confined to the eastern United States.^[14] Globally, distribution patterns reflect climatic tolerances: periodical species like Magicicada are restricted to temperate zones in North America, enabling synchronized mass emergences, whereas most non-periodical species form continuous broods in tropical forests; cicadas are notably absent from polar regions, where cold temperatures and lack of host trees preclude survival.^[22]^[23] Recent taxonomic advances, driven by acoustic recordings and DNA barcoding, have revealed new species in understudied tropical areas, including 12 from Madagascar described in 2021 and 13 from Thailand in 2024, underscoring the untapped diversity in regions like island hotspots and continental rainforests.

Fossil record

The fossil record of cicadas, encompassing the superfamily Cicadoidea within the broader Cicadomorpha, extends back to the Permian period, approximately 290 million years ago, marking the earliest known appearances of stem-group cicadomorphans. These primitive forms, such as members of the family Dysmorphoptilidae from deposits in European Russia and Siberia, exhibited early hemipteran traits including piercing-sucking mouthparts adapted for xylem feeding and wing venation patterns indicative of jumping locomotion similar to modern leafhoppers. By the middle Permian, cicadomorphans had become dominant herbivores in certain ecosystems, with fossils from the Kungurian stage (around 283–272 Ma) in the Tunguska Basin of Russia revealing inflated frontoclypeal structures and sessile nymph morphologies that foreshadowed later cicada adaptations.^[24]^[25] During the Mesozoic era, cicadomorphans underwent significant diversification, with true cicadoids emerging by the Middle Jurassic. Exceptional preservation in the Daohugou Beds of Inner Mongolia, China (approximately 165 Ma), has yielded genera such as Macrotettigarcta and Shuraboprosbole, showcasing advanced wing structures and body plans closely resembling extant cicadas. These fossils indicate the divergence of key lineages, including the relict family Tettigarctidae and the more derived Cicadidae, with evidence of early acoustic communication emerging in Permian dysmorphoptilids through specialized wing modifications for sound production—predating modern tymbal organs but hinting at ancestral stridulatory mechanisms in hemipterans.^[26]^[25] Cenozoic records provide insights into modern cicada morphologies, with amber-preserved specimens revealing detailed soft-tissue preservation. Eocene amber from the Baltic region contains nymphs of cicadids, approximately 44 million years old, displaying root-feeding behaviors and burrowing adaptations akin to contemporary species, while Dominican amber inclusions from the Oligocene-Miocene boundary (around 23–20 Ma) include adults like Minyscapheus dominicanus, with intact wing venation and ovipositor structures. A notable 2025 discovery from the Eocene Messel Pit in Germany revealed a well-preserved adult cicada fossil, approximately 47 million years old, providing evidence of early acoustic structures in European cicadids.^[27] These fossils highlight a post-Cretaceous-Paleogene recovery, where cicada diversity appears reduced immediately after the event—likely due to habitat disruptions from the extinction of non-avian dinosaurs and associated vegetation changes—but rebounded with the radiation of angiosperms in the Paleogene, facilitating host shifts to new xylem sources among root-feeding nymphs.^[26]

Physical characteristics

Morphology

Cicadas exhibit a typical insect body plan, divided into three distinct tagmata: the head, thorax, and abdomen. The head is short and wide, featuring prominent compound eyes positioned wide apart on short eyestalks, which provide a broad field of vision, and three simple eyes (ocelli) arranged in a triangular formation on the vertex. Short antennae, consisting of 7-9 segments, arise between the eyes, and the mouthparts are modified into an elongated rostrum formed by the labium, which houses a bundle of piercing stylets for feeding.^[28]^[29] The thorax is robust and segmented into pro-, meso-, and metathorax, bearing three pairs of legs adapted for walking and grasping—the forelegs are particularly prehensile—and two pairs of membranous wings. The forewings, attached to the mesothorax, are larger and typically transparent with visible venation, though some species display patterned coloration; at rest, they are held roof-like over the abdomen. The hindwings, smaller and attached to the metathorax, are folded fan-like beneath the forewings when not in use.^[28]^[30] The abdomen comprises 10 segments and houses key external structures, including paired tympanal organs—thin, oval membranes located ventrally on the second segment—for auditory function. Males possess enlarged tymbals, ribbed membranes on the dorsolateral sides of the first abdominal segment, while females feature a robust, saw-like ovipositor at the posterior end for inserting eggs into plant tissues. These sexual differences in abdominal structures facilitate reproduction, with variations in overall size and coloration addressed in subsequent discussions of dimorphism.^[28]^[31]

Size, coloration, and sexual dimorphism

Cicadas vary considerably in body size, with most species measuring 1 to 5 cm in length.^[32] Larger species, such as the Southeast Asian Tacua speciosa, attain a head-body length of 4.7–5.7 cm and a wingspan of 15–18 cm, making it one of the largest known cicadas.^[33] These size differences reflect adaptations to diverse habitats and ecological roles across the family's approximately 3,000 species. Coloration in cicadas is typically dominated by shades of black, green, or brown, providing effective camouflage against tree bark and foliage.^[34] Distinctive patterns occur in certain taxa, including red compound eyes and orange-tinged wing veins in periodical cicadas of the genus Magicicada.^[3] Intraspecific color variations are common, such as differences in eye color (e.g., white, blue, or red) and wing vein pigmentation, which may arise from genetic or environmental factors without altering species classification.^[35] Sexual dimorphism in cicadas primarily manifests in morphology and size rather than coloration. Females are often heavier and equipped with a strong, pointed ovipositor for inserting eggs into plant tissue, while males possess a blunter abdomen and specialized tymbal structures for acoustic signaling.^[3] The degree of size dimorphism varies by species and population; for instance, in some North American taxa, females exceed males in body mass and wing loading, enhancing their oviposition capabilities, whereas males may exhibit relatively greater maneuverability.^[36]^[37] Coloration differences between sexes are subtle and not widespread, though habitat-linked intraspecific morphs—such as greener individuals in forested environments versus browner forms in arid regions—can influence overall appearance across populations.^[38]

Physiology

Thermoregulation

Cicadas are ectothermic insects that primarily rely on behavioral thermoregulation to maintain body temperatures suitable for activity, typically optimizing around 30–35°C through basking in sunlight to absorb solar radiation or seeking shade to avoid overheating.^[39] Species adjust their body orientation, such as exposing lateral surfaces to the sun or using wings as a parasol, to fine-tune heat gain, with activity often suspended when ambient temperatures exceed thresholds like 40–42°C.^[39] This behavioral strategy allows cicadas to achieve thoracic temperatures up to 13°C above ambient during warm-up, enabling flight and other functions even in suboptimal conditions.^[40] Physiologically, cicadas supplement behavior with endothermic heat production via rapid contractions of thoracic flight muscles, raising body temperature by 10–12°C above ambient to support activities like singing, where metabolic rates increase significantly during choruses.^[40] For cooling, they employ cuticular evaporation by expelling water from xylem fluids through spiracles, which can lower body temperature up to 6.7°C below ambient, aided by hemolymph circulation to distribute heat or moisture.^[39] Maximum voluntary tolerance temperatures range from 36–38°C across species, beyond which heat stress impairs function, including reduced calling above approximately 40°C due to torpor onset near 46–47°C.^[41]^[42] In periodical cicadas like Magicicada species, nymphs underground benefit from soil's thermal insulation, which buffers extreme surface fluctuations and maintains stable conditions during their multi-year development, with emergence triggered only when soil at 20 cm depth exceeds 18°C.^[39] As adults, they adapt by wing-spreading to capture solar heat at lower temperatures or fanning wings to enhance evaporative cooling in heat, helping sustain choruses in variable post-emergence environments.^[39] These mechanisms underscore cicadas' resilience to thermal variability, though upper limits constrain activity in increasingly warm climates.^[39]

Sound production and communication

Cicadas produce sound primarily through a mechanism known as tymbal stridulation, unique to males of the order. The tymbals consist of ribbed, stiff membranes located dorsolaterally on the anterior abdomen, specifically the first tergite, which are rapidly deformed by contractions of specialized tymbal muscles. These muscles vibrate the membranes at frequencies typically between 200 and 500 Hz, with each rapid buckling producing a brief click that combines into continuous song when repeated. The abdomen acts as an acoustic resonator, amplifying the low-frequency components of the sound for efficient transmission over distances.^[43]^[44]^[45] Male cicadas employ distinct song types for communication, primarily to attract females and defend territories. Calling songs are long, repetitive phrases broadcast from perches to draw conspecifics into choruses, reaching peak intensities of up to 100 dB in dense aggregations of periodical species such as Magicicada. Once a female approaches, males switch to courtship songs—shorter, more variable sequences—and elicit responses through duets, where females produce sharp wing-clicks by flicking their wings against the thorax, signaling receptivity and aiding precise mate location.^[46]^[47]^[48] The acoustic repertoire of cicadas exhibits high diversity, with species-specific patterns in phrase duration, syllable rate, and dominant frequency facilitating mate recognition amid sympatric assemblages. For instance, different Magicicada species produce calls with unique temporal structures, such as the interrupted buzz of M. cassini versus the continuous whine of M. septendecim. Chorusing behavior, where males synchronize calls within groups, enhances overall signal salience for females while reducing predation risk through temporal overlap that confounds predator localization.^[48]^[46]^[49] Recent bioacoustics research in the 2020s has uncovered geographic dialects in calling song structure across populations, such as variations in phrase complexity linked to habitat and isolation in genera like Tettigettalna, informing models of acoustic evolution and speciation. Studies during 2024 mass emergences have also documented chorus intensities up to 86 dB via crowdsourced recordings, raising awareness of auditory discomfort for humans with sensory processing differences, including those on the autism spectrum, though no permanent hearing loss occurs at these levels.^[50]^[51]^[52]

Locomotion and sensory systems

Cicadas exhibit specialized locomotion adapted to their arboreal and subterranean lifestyles. Adult cicadas primarily rely on flight for dispersal and mating, powered by robust asynchronous flight muscles in the thorax that enable rapid wing oscillations.^[53] These muscles drive the forewings and hindwings in a coupled motion, generating thrust during the upstroke and managing drag in the downstroke to support forward flight and limited hovering.^[30] While capable of sustained flight over moderate distances, adults are notably clumsy on the ground, using their six spiny legs—equipped with tibial spines for grip—to walk or climb awkwardly, often preferring to launch into flight rather than navigate terrestrially.^[54] Nymphs, in contrast, are subterranean dwellers specialized for burrowing. Their powerful forelegs, modified with hook-like structures and robust segments, facilitate excavation through soil, allowing them to construct tunnels and chambers while feeding on root xylem.^[55] These forelegs enable efficient forward digging, where sediment is rolled into balls and pushed behind the body.^[56] Upon maturity, nymphs emerge from the soil at night, using their legs to crawl up vertical surfaces such as tree trunks to reach safe molting sites, minimizing predation risk during the transition to adulthood.^[57] Cicadas possess a suite of sensory organs that integrate visual, chemical, and vibratory cues for environmental perception. The prominent compound eyes, each comprising approximately 7,500 ommatidia in species like the Australian redeye cicada (Psaltoda moerens), provide a wide field of view and excel at detecting motion, aiding in obstacle avoidance and predator detection during flight.^[58] Three dorsal ocelli serve as simple photoreceptors, primarily sensing light intensity and changes in illumination to orient the body and regulate daily activities, such as emergence timing.^[59] Antennae, though short and bristle-like, house olfactory receptors that detect volatile plant compounds and potentially other chemical signals, contributing to host selection and habitat navigation.^[60] The tympanal organs on the abdomen function as auditory sensors for airborne sounds, with membranes that vibrate in response to acoustic signals from conspecifics.^[61] These sensory systems underpin key behaviors in locomotion. Optomotor responses, mediated by the compound eyes, stabilize flight paths by reflexively adjusting wing beats to counteract visual flow, ensuring straight-line trajectories during dispersal.^[62] Substrate vibration sensing via leg chordotonal organs allows cicadas to perceive and evade ground-based predators, triggering rapid flight initiation or postural changes with latencies as low as 6 ms.^[63] This integration enhances survival by linking sensory input directly to motor outputs, from burrowing adjustments in nymphs to evasive maneuvers in adults.^[64]

Life history and ecology

Life cycle and development

Cicadas undergo incomplete metamorphosis, also known as hemimetabolous development, consisting of three primary stages: egg, nymph, and adult.^[3] This process lacks a distinct pupal stage, with nymphs resembling smaller, wingless versions of adults and undergoing gradual changes through molting.^[65] The cycle begins with egg deposition. Females use their specialized ovipositor to cut slits into the bark of small tree twigs or branches, typically 1–2 years old, and lay batches of 20–30 eggs per slit, potentially creating dozens of such slits per female.^[66] Eggs incubate for 2–10 weeks, depending on species and environmental conditions, before hatching into first-instar nymphs.^[67] Upon hatching, the tiny nymphs drop to the ground and immediately burrow into the soil using their fossorial forelegs, which are robust and adapted for digging with enlarged, spade-like tibiae and strong claws.^[38] Nymphs spend the majority of their lives underground, progressing through five instars over periods ranging from 2–5 years in annual species to 13 or 17 years in periodical species.^[66] During this subterranean phase, they feed on xylem sap from plant roots using piercing-sucking mouthparts, excavating burrows typically 15 to 61 cm (6 to 24 in) deep; in wet soils, nymphs sometimes construct shallow mud chimneys (2 to 8 cm high) above the entrance for protection against flooding.^[65]^[68] Development is marked by periodic molts, where nymphs shed their exoskeleton to grow, with synchronization across populations often regulated by environmental cues such as soil temperature and moisture rather than strict diapause.^[3] In the final instar, mature nymphs construct emergence tunnels to the surface, typically when soil temperatures reach about 18°C (64°F).^[66] Emergence culminates in the transformation to adulthood. Nymphs climb vertical surfaces like tree trunks or stems, anchor themselves, and undergo a final molt, splitting their exoskeleton along the back to reveal the pale, soft-bodied teneral adult.^[65] This molting process, part of mass synchronous events in periodical species, can involve billions of individuals emerging over a few nights, overwhelming predators through sheer numbers.^[69] The new exoskeleton hardens within hours to days, during which adults remain vulnerable and do not fly or sing; morphological changes include the expansion and pigmentation of wings and eyes.^[3] Adult cicadas have a brief lifespan of 2–6 weeks above ground, dedicated almost entirely to reproduction.^[70] Males produce species-specific calls to attract females, mating occurs soon after hardening, and females then oviposit before dying; unemerged nymphs from previous cycles may persist underground.^[65] Variations in cycle length highlight adaptive strategies: annual species complete development in 2–5 years without strict synchronization, while periodical species like those in Broods X (17-year) and XIX (13-year) align emergences precisely, as seen in the rare 2024 dual-brood event across the eastern U.S., where billions emerged in regions home to millions of people.^[69]

Periodical versus annual species

Periodical cicadas, belonging to the genus Magicicada, are characterized by highly synchronized emergences after fixed intervals of either 13 or 17 years, with seven species divided into four 13-year species and three 17-year species primarily found in eastern North America.^[14]^[71] These prime-numbered cycles facilitate brood synchrony, where vast populations emerge simultaneously, a strategy hypothesized to minimize predation risk by disrupting potential predator population cycles.^[72]^[73] In contrast, annual cicadas exhibit overlapping life cycles of 2 to 5 years, resulting in consistent yearly emergences without strict synchrony, and they represent a far more diverse group globally, with thousands of species across various genera.^[57]^[74] For example, species in the genus Neotibicen (formerly Tibicen) are common in temperate regions of North America, where their staggered generations ensure a steady presence during summer months.^[75]^[76] The evolutionary advantages of these strategies differ markedly: periodical cicadas achieve predator satiation through mass emergences involving billions of individuals, overwhelming predators and allowing sufficient survivors to reproduce, whereas annual cicadas face ongoing predation pressure, favoring adaptations like camouflage and crypsis over numerical overwhelming.^[77]^[78]^[79] Recent genetic research from 2024 has modeled potential hybridization between 13- and 17-year broods during rare co-emergences, such as the 2024 event, suggesting genetic mixing could lead to hybrid broods with altered cycles.^[80] Additionally, studies indicate that climate change may desynchronize periodical cycles by altering soil temperatures and emergence cues, potentially causing earlier or off-schedule emergences and reducing brood cohesion over time.^[81]^[82]

Diet and habitat preferences

Cicadas are xylem sap feeders throughout their lives, relying on this fluid as their primary source of nutrition. Nymphs use specialized piercing-sucking mouthparts to extract xylem sap from the roots of trees and grasses, employing pump-like mechanisms to draw up the water-rich, nutrient-poor mixture containing amino acids and minerals.^[83]^[84] Adults continue this diet by feeding on xylem sap from twigs and stems of host plants, a behavior facilitated by the same mouthpart structures.^[85]^[83] Host plant specificity varies among cicada species and influences their distribution. Periodical cicadas, such as those in the genus Magicicada, preferentially feed on and oviposit in deciduous trees like oaks (Quercus spp.), maples (Acer spp.), and hickories (Carya spp.), which provide suitable root systems for nymphal development.^[86]^[87] In contrast, many annual cicada species in arid regions associate with grasses or conifers such as pines (Pinus spp.), adapting to sparser vegetation.^[88]^[16] Nymphs inhabit underground environments in well-drained soils, where they burrow to access root xylem, with soil compaction in urban areas posing a barrier to their movement and survival.^[89]^[90] Adults emerge into forested woodlands, forest edges, or urban fringes, particularly in temperate eastern North American deciduous forests for periodical species, while tropical and arid annuals favor open grasslands or pine-dominated landscapes.^[81]^[91]^[88] The xylem sap diet, composed of approximately 95% water and low in nutrients, necessitates specialized physiological adaptations for survival. Cicadas exhibit efficient osmoregulation to manage ion and water balance from the hypotonic fluid, preventing dehydration or electrolyte imbalances.^[92]^[84] Additionally, symbiotic bacteria in their guts convert xylem components into essential amino acids, supplementing the deficient diet and enabling long-term subterranean life in some species.^[93]

Interactions with other organisms

Predators, parasites, and pathogens

Cicadas face significant biotic threats from a diverse array of predators that target various life stages, particularly adults during mass emergences. Birds are the primary predators of adult periodical cicadas, with species such as starlings, robins, blue jays, and cuckoos consuming large numbers of emergents, often accounting for 15–40% of the adult population in the early phases of emergence. Mammals, including squirrels, raccoons, dogs, and moles (which prey on subterranean nymphs), also contribute substantially to mortality, while reptiles like turtles feed on both nymphs and adults. Arthropod predators, such as cicada killer wasps and spiders, further reduce populations, though overall predation rates do not scale proportionally with cicada density due to predator satiation strategies.^[77]^[94]^[95] Parasitic organisms exploit cicadas across all developmental stages, often with host-specific adaptations. Parasitoid wasps, including trichogrammatids that target eggs and cecidomyiids that attack egg masses, impose limited but notable mortality on early life stages, though they do not significantly regulate overall populations. Nematodes, such as entomopathogenic species like those in the genus Steinernema, infect nymphs and adults in soil environments, causing internal damage and death, with records of natural infections in species like Diceroprocta apache. Mites, particularly the ectoparasitic Pyemotes herfsi, attach to adults and drain nutrients, leading to weakened hosts and secondary infections; during the 2024 Brood XIII emergence, mite infection rates reached up to 14% in some Illinois populations. In tropical cicada species, trematode flatworms occasionally parasitize nymphs, altering development and increasing vulnerability to other threats, though such infections are less documented in temperate broods. Bacterial endosymbionts like Wolbachia pipientis, detected in 41–57% of periodical cicadas during the 2024 Brood XIII emergence, are known to manipulate reproduction in some insect hosts.^[77]^[96]^[97]^[98] Pathogenic microbes, especially fungi, pose severe risks to cicada choruses, often causing episodic die-offs. The obligate fungal pathogen Massospora cicadina infects late-stage nymphs underground via resting spores and manifests in adults as a "zombie" infection, replacing the posterior abdomen with a spore mass that manipulates behavior—inducing hypersexuality in males, who mimic female signals to attract mates and disperse conidiospores during prolonged flights. Infected individuals experience no apparent pain or reduced mobility, continuing to feed and mate despite organ destruction, which ultimately leads to sterility and death; infection rates during emergences typically range from 5–25%, with symptomatic cases causing up to 50% mortality in dense choruses. During the 2024 dual emergence of Broods XIII and XIX, fungal epidemics, primarily Massospora, reduced populations by 20–30% in affected U.S. Midwest and Southern areas, with DNA detection in 23% of asymptomatic cicadas indicating latent spread. Other pathogens include viruses that cause unspecified systemic infections in larvae and bacteria like those in the genus Serratia, which trigger septicemia in weakened adults, though these are less prevalent than fungal threats.^[77]^[99]^[100]^[101]^[102]

Defense mechanisms

Cicadas utilize a range of physical, behavioral, and physiological strategies to deter predators and resist parasites and pathogens. These adaptations are particularly crucial for species like periodical cicadas, which emerge in massive numbers after long underground development periods, making them vulnerable during short adult phases. Cryptic coloration serves as a primary visual defense for many cicada species, allowing them to blend seamlessly with tree bark, twigs, and foliage when at rest. This camouflage reduces detection by visual predators such as birds and lizards, with adults and nymphs exhibiting mottled browns, grays, and greens that mimic their arboreal or subterranean environments. ^[103] In some tropical species, such as those in the genus Hemisciera, adults employ deimatic displays by suddenly flashing brightly colored hindwings—often red or orange—creating a startling contrast that briefly disorients approaching threats, providing time for escape. ^[32] Behavioral defenses are prominent in gregarious species, especially periodical cicadas (Magicicada spp.), where mass chorusing and synchronous emergences overwhelm predators through a combination of acoustic saturation and numerical abundance. Males aggregate in choruses producing intense, synchronized calls that can reach deafening levels, confusing predators and diluting the risk to any individual by distributing attention across thousands or millions of individuals; emergence densities have been recorded up to 1.5 million per acre, exemplifying predator satiation. ^[32] ^[104] Disturbed adults often produce a defensive "alarm buzz" or struggle flight, further deterring close-range attacks. ^[105] Chemical defenses in cicadas are less prominent than in related hemipterans but include unpalatable tissues in nymphs, which contain high levels of indigestible compounds and structural reinforcements that discourage burrowing predators like moles or ants from consuming them fully. ^[106] Adults may release viscous fluids from oral or anal regions when handled, acting as a repellent to small arthropod predators, though this is not as potent as glandular secretions in true bugs. ^[92] Immune responses in cicadas mirror broader insect mechanisms, with hemocytes in the hemocoel encapsulating invading parasites such as nematodes or fungal elements through multilayered cellular sheaths that isolate and melanize the intruder, preventing systemic spread. ^[107] Recent research has highlighted humoral defenses, including antifungal peptides like cicadin isolated from juvenile cicada tissues, which exhibit potent activity against pathogenic fungi such as Botrytis cinerea at nanomolar concentrations, potentially aiding periodical species in resisting soil-borne infections during prolonged nymphal stages. ^[108] These peptides contribute to innate immunity by disrupting fungal cell membranes, underscoring cicadas' adaptive resilience to microbial threats. ^[109]

Ecological roles

Cicadas play a pivotal role in nutrient cycling within forest ecosystems, particularly through the mass mortality following periodical emergences. The decomposition of billions of adult carcasses introduces substantial pulses of organic matter, including nitrogen and phosphorus, into the soil. Studies estimate that these events can deposit 10 to 70 kg of nitrogen per hectare every 13 or 17 years, with similar contributions for phosphorus representing up to 50% of annual leaf litter inputs in some woodland streams.^[110]^[111] This nutrient enrichment enhances soil fertility, stimulates microbial activity, and promotes plant productivity; for instance, experimental additions of cicada carcasses increased understory plant biomass by 61% and foliar nitrogen content by 20% compared to controls. Such inputs are especially vital in nutrient-poor forest soils, where they facilitate long-term carbon sequestration and support post-emergence vegetation recovery. As a major prey resource, cicadas bolster food web dynamics and biodiversity during their brief adult phase. The synchronized mass emergences provide a predictable, high-density food pulse that sustains diverse predators, including over 80 bird species that shift foraging behaviors to target cicadas, thereby reducing predation on alternative prey like caterpillars and allowing those populations to rebound.^[91] This trophic rewiring extends to mammals, reptiles, and invertebrates, enhancing overall ecosystem resilience and connectivity across forest communities. The scale of these events—often exceeding 1 million individuals per hectare—positions cicadas as a keystone subsidy, preventing predator starvation in lean periods and promoting stable predator-prey balances over multi-year cycles.^[112] Cicadas interact with plants in ways that influence forest structure and function. Female oviposition scars branches by slicing slits for egg-laying, often causing localized dieback or "flagging," but long-term studies show negligible effects on radial tree growth or mortality across multiple species.^[113] In some cases, this injury prompts compensatory lateral branching, potentially leading to denser canopies. Meanwhile, both nymphal root-feeding and adult xylem sap consumption have minimal direct impacts on tree hydraulics or overall vigor, though chronic nymphal herbivory may subtly alter water and nutrient uptake without hindering net growth.^[114] These interactions, combined with soil bioturbation from nymph burrows, contribute to improved soil structure and water infiltration.^[115] Due to their long generation times and habitat requirements, cicadas function as effective indicator species for environmental health. Periodical species demand large, unbroken forest patches of at least 52 hectares to sustain synchronized emergences, making them vulnerable to fragmentation from urbanization and agriculture.^[81] Populations are declining in regions affected by climate shifts, with altered temperature regimes disrupting developmental timing and survival; for example, some annual cicada populations have dropped by up to 80% over the past decade due to warming and habitat loss.^[116] Monitoring cicada abundances thus signals broader ecosystem degradation, underscoring the need for conservation to preserve these roles.

Human interactions

Cultural and symbolic significance

In Chinese mythology, cicadas have long symbolized rebirth and resurrection, attributed to their life cycle of emerging from the ground after years underground and shedding their exoskeleton. This association dates back to ancient times, with jade carvings of cicadas placed on the tongues of the deceased during the Han dynasty (206 BCE–220 CE) to ensure immortality and guide the soul in the afterlife.^[117]^[118] In ancient Greek mythology, cicadas were linked to summer and revered as sacred to Apollo, the god of music and poetry, with their shrill songs evoking divine inspiration and the heat of the season. Hesiod, in Works and Days (ca. 700 BCE), described the cicada's song as a marker of "the season of fatiguing summer," while Plato's Phaedrus (ca. 370 BCE) mythologized them as humans transformed into insects by the Muses for their devotion to song, granting them the ability to subsist on dew without food.^[119]^[120] Cicadas frequently appear in literature and art as emblems of transience and renewal. In Japanese haiku, the 17th-century poet Matsuo Bashō captured their evanescence, as in his verse "The cry of the cicada / Soaks into stone," where the insect's piercing call against unyielding rock symbolizes the brevity of life amid enduring nature.^[121] During the European Renaissance, insects like cicadas featured in still-life paintings as vanitas symbols, representing the fleeting nature of existence and the cycle of life and death, often juxtaposed with fruits and flowers to underscore mortality.^[122] In contemporary art, cicada motifs persist in tattoos, where the insect embodies personal transformation and rebirth, drawing on its metamorphic life stage to signify overcoming adversity and emerging renewed.^[123] In music, cicadas have inspired works that mimic or evoke their resonant calls. While Antonio Vivaldi's The Four Seasons (1725) programmatically depicts summer's oppressive heat in its "Summer" concerto—complete with string figures suggesting swarming insects and languid languor—no explicit cicada motif appears, though the piece's vivid portrayal of seasonal sounds has been interpreted as encompassing such natural choruses.^[124] In the 2020s, ambient and neoclassical compositions have directly incorporated cicada field recordings for atmospheric effect, as in Scott Buckley's "Cicadas" (2024), a delicate track blending piano, strings, and actual insect sounds to create a meditative soundscape of renewal and natural rhythm.^[125] Across folklore traditions, cicadas often serve as harbingers of change or spiritual intermediaries. In Australian Aboriginal lore, natural indicators like insect emergences help predict seasonal shifts, including rain, though specific cicada stories emphasize their role in signaling the end of dry periods and the arrival of wetter weather. In some African and Caribbean traditions, cicada songs are viewed as carriers of spiritual messages from ancestors or the divine, their persistent choruses interpreted as calls connecting the living to the departed.^[126]^[127]

As pests and economic impacts

Cicadas, particularly periodical species in North America, can cause significant damage through oviposition, where females use their ovipositors to make V-shaped slits in the bark of twigs and small branches to deposit eggs. These slits, typically in branches 3/16 to 7/16 inches in diameter, weaken the vascular tissue, leading to wilting, dieback, and branch breakage in affected trees. Young orchards, such as those with apple, peach, or pear trees, are especially vulnerable during mass emergences, as the high density of egg-laying females can result in widespread flagging of foliage and reduced tree vigor.^[128]^[129]^[130] The wounds also create entry points for pathogens and secondary pests, exacerbating long-term horticultural losses in commercial fruit production.^[129] Economic impacts are primarily localized to agriculture and forestry sectors, where oviposition damage reduces yields in orchards and nursery stock, necessitating replanting or pruning. In urban and suburban settings, the intense chorusing of emerging adults—reaching up to 100 decibels—generates widespread noise complaints, disrupting daily life and potentially increasing municipal response costs, though direct effects on property values remain unquantified. During the 2024 dual-brood emergence in the eastern and midwestern U.S., these effects were amplified across 16 states.^[131]^[132]^[133] The 2025 emergence of Brood XIV across approximately 13 states, including Georgia, Kentucky, Indiana, and others, similarly prompted preparations for potential tree damage, though reports indicated limited agricultural impacts overall.^[134] Management strategies focus on prevention rather than eradication, given the short adult lifespan and ecological benefits of cicadas. Cicada-proof netting with mesh smaller than 1/4 inch is recommended for young trees and shrubs up to 10 feet tall, installed before emergence to exclude females from oviposition sites; this method is cost-effective for high-value orchards but labor-intensive for large areas. Insecticide sprays, such as carbaryl or pyrethroids, can be applied during peak adult activity (first through third cover sprays) to reduce egg-laying, though experts advise against broad use due to harm to pollinators and predators. Biological controls leverage natural enemies, including birds, mammals, and insects like cicada killer wasps, which consume large numbers during emergences, minimizing the need for chemical interventions.^[135]^[136]^[132] In non-pest contexts, cicadas rarely reach damaging densities in tropical regions due to continuous predation by birds, reptiles, and parasitoids, which prevent population booms typical of temperate periodical species. However, climate-driven range shifts and altered emergence timing may expand cicada impacts northward or into new habitats, potentially increasing pest pressure in agriculture as warmer conditions favor survival and synchronization.^[137]^[138]

Uses in food, medicine, and other practices

Cicadas have been consumed as food in various cultures, particularly in parts of Asia and Africa, where they are valued for their nutritional content. In China, species such as Cryptotympana atrata are harvested and prepared in traditional dishes, including roasting or stir-frying, providing a high-protein source with essential amino acids and fatty acids that support anti-inflammatory effects.^[139]^[140] These insects are eaten at different life stages, from protein-rich nymphs to adults, and recent studies highlight their sustainability as an alternative protein amid global food security challenges, noting low environmental impact compared to conventional livestock.^[141] In traditional Chinese medicine, cicada exuviae, known as chan tui (the dried outer shell of Cryptotympana pustulata nymphs), have been used for centuries to treat respiratory conditions like asthma by dispersing wind-heat and relieving spasms. These exuviae are often prepared as decoctions or teas to soothe throat irritation and improve breathing. Pharmacological research in 2023 has confirmed anti-inflammatory properties in cicada extracts, attributing them to bioactive compounds that modulate immune responses and reduce airway inflammation in asthma models.^[142]^[143]^[139] Beyond consumption, cicada shells serve practical purposes in crafts and industry. In the United States, live cicadas are commonly used as fishing bait, hooked through the thorax to attract species like bass, trout, and catfish during emergences, leveraging their natural movement on the water surface. Emerging biotechnological research explores chitin extraction from cicada sloughs for applications in sustainable materials, such as wound dressings and nanocomposites, due to the polymer's biocompatibility and thermal stability.^[144]^[145]^[146] While cicadas are generally safe for human consumption when properly prepared, individuals with shellfish allergies should avoid them owing to shared chitin proteins that can trigger reactions. Wild-harvested specimens pose risks from environmental contaminants like pesticides or pathogens, necessitating thorough cleaning and cooking to mitigate potential health hazards.^[147]^[148]

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Feb 3, 2025 · Broods of periodical cicada adults have synchronized emergence events every 13 or 17 years. These events can be loud and very noticeable to ...
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The Tibicen auletes is the largest species of the Tibicen cicadas (largest in terms of physical size). Visit Insect Singers to hear its song. Neotibicen auletes ...
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Periodical cicadas disrupt trophic dynamics through ... - Science
Oct 19, 2023 · Cicada emergence reduces avian predation on forest caterpillars. As anticipated, heavy consumption of cicadas by a diverse assemblage of ...
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Jul 7, 2021 · Birds are the primary predator of periodical cicadas, but reptiles and mammals such as squirrels, raccoons, and dogs readily feed on cicadas ...
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INFECTION RATE OF TWO PARASITES BETWEEN BROOD XIII ...
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Acute resource pulses from periodical cicadas propagate to ...
Cicada addition did change the community composition of soil nematodes and increased the abundance of bacterial‐ and fungal‐feeding nematodes, while plant ...
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Deposition and decomposition of periodical cicadas (Homoptera
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Periodical cicada emergence resource pulse tracks forest expansion ...
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The effects of pulsed fertilization and chronic herbivory by periodical ...
Apr 22, 2019 · We found that cicada fertilization strongly increased tree growth in the year of emergence, creating differences in tree size that persisted at least 2 yr ...
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Plan ahead to protect young fruit trees from 17-year periodical ...
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May 10, 2021 · These terrestrial cicadas will offer excellent dry fly fishing opportunities on almost any body of water near their emergence.
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Warning: Cicadas can be a tasty treat, but they can be hazardous to ...
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Consumers relying on wild harvested edible insects are particularly at risk of pesticide food poisoning. In Thailand for example, pesticide contaminated insects ...<|control11|><|separator|>