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Cicadomorpha

Cicadomorpha is an infraorder of insects in the order Hemiptera, suborder Auchenorrhyncha, consisting of approximately 33,000 extant species of plant sap-feeding bugs distributed worldwide.^[1] These insects are characterized by morphological features such as an enlarged postclypeus, a small antennal pedicel lacking conspicuous sensilla, an aristate flagellum, absent tegulae, forewings with usually separate anal veins, and narrowly separated small middle coxae.^[2] All species are phytophagous, primarily feeding on xylem or phloem sap, and many produce audible sounds through specialized tymbal organs for communication.^[2] Taxonomically, Cicadomorpha is divided into three main superfamilies: Cicadoidea (cicadas), Cercopoidea (spittlebugs), and Membracoidea (leafhoppers and treehoppers).^[2] The superfamily Cicadoidea includes two families, Cicadidae (with over 3,000 species) and the relict Tettigarctidae (two species); Cercopoidea encompasses six families such as Aphrophoridae and Cercopidae; and Membracoidea contains five families, notably the diverse Cicadellidae (over 22,000 species) and Membracidae (treehoppers).^[1] This classification reflects well-established morphological and molecular phylogenetic evidence, with Cicadomorpha dating back to the Permian period in the fossil record.^[2] Ecologically, Cicadomorpha species occupy diverse habitats from tropical rainforests to temperate grasslands, often exhibiting high diversity in canopy layers where over 90% of tropical species remain undescribed.^[2] They play key roles in ecosystems as herbivores and prey for predators, but many are economically significant pests due to direct feeding damage or as vectors of plant pathogens like Xylella fastidiosa, causing diseases such as Pierce's disease in grapes and citrus variegated chlorosis, with annual losses exceeding $100 million in affected regions.

Taxonomy and Phylogeny

Definition and Higher Classification

Cicadomorpha is a monophyletic infraorder within the suborder Auchenorrhyncha of the order Hemiptera, encompassing a diverse array of plant-sap feeding insects including cicadas, spittlebugs, leafhoppers, and treehoppers.^[3] This group is positioned as the sister clade to Fulgoromorpha within Auchenorrhyncha, with an estimated diversity exceeding 35,000 described species worldwide.^[4] Unlike the suborder Sternorrhyncha, which includes aphids, scale insects, and whiteflies, Cicadomorpha are distinguished by key morphological adaptations such as a specialized filter chamber in the gut that facilitates rapid filtration and excretion of excess water from xylem sap, differing structurally from the filter chamber in Sternorrhyncha, as well as the presence of specialized sound-producing organs like tymbals in certain lineages such as cicadas.^[5]^[6] Diagnostic traits unique to Cicadomorpha include paired ocelli located on the vertex of the head, tarsi composed of two segments in adults, and a labium divided into three segments, which collectively aid in distinguishing them from other hemipteran groups.^[7] These features, combined with an enlarged postclypeus and fused basal veins in the forewing (Sc and R), underscore their monophyletic status and adaptation to phytophagous lifestyles.^[4] Historically, Cicadomorpha were included in the polyphyletic assemblage Homoptera, which grouped Auchenorrhyncha and Sternorrhyncha together based on superficial similarities like piercing-sucking mouthparts.^[8] Cladistic analyses from the 1990s onward, integrating morphological characters and early molecular data such as 18S rDNA sequences, revealed Homoptera to be paraphyletic and prompted the reclassification of Auchenorrhyncha (including Cicadomorpha) as a distinct suborder within the monophyletic Hemiptera.^[9]^[10] This shift emphasized the evolutionary divergence between Auchenorrhyncha and Sternorrhyncha, dating back to at least the Permian period.^[8] The infraorder Cicadomorpha is further divided into three primary superfamilies: Cicadoidea, Cercopoidea, and Membracoidea.^[9]

Superfamilies and Families

Cicadomorpha is classified into three main superfamilies: Cicadoidea, Cercopoidea, and Membracoidea, encompassing a total of approximately 35,000 described species distributed across 12 families.^[11] The superfamily Membracoidea accounts for roughly 80% of these species, reflecting its exceptional diversity, while the remaining superfamilies contribute smaller but ecologically significant portions.^[11] Taxonomic classifications vary slightly, with some authorities recognizing 8 to 10 families by subsuming certain smaller groups, but molecular and morphological analyses consistently support the tri-superfamily structure.^[12] The superfamily Cicadoidea includes two families: Tettigarctidae, with only two extant species restricted to Australia, and Cicadidae, comprising about 3,000 species worldwide.^[11]^[13] Cicadidae species are typically large-bodied insects, often exceeding 2 cm in length, and are notable for their loud acoustic signals produced by males during mating; certain North American species exhibit synchronized periodical emergences lasting 13 or 17 years underground as nymphs.^[13] Superfamily Cercopoidea consists of five families—Aphrophoridae, Cercopidae, Clastopteridae, Epipygidae, and Machaerotidae—totaling around 3,000 species globally.^[11]^[14] The superfamily is characterized by the nymphs' production of protective spittle masses, formed from plant sap and excretions, which shield them from predators and desiccation during development on herbaceous hosts.^[14] Cercopidae, the largest family within this group, includes the common spittlebugs, many of which are polyphagous on grasses and forbs. The superfamily Membracoidea is the most species-rich, with five families: Cicadellidae, Membracidae, Aetalionidae, Melizoderidae, and Myerslopiidae, representing over 26,000 species.^[11] Cicadellidae (leafhoppers) dominates with approximately 23,000 species, exhibiting wedge-shaped bodies adapted for agile movement on foliage and a wide range of host plants, often with high specificity that influences their ecological roles as vectors of plant pathogens.^[15] Membracidae (treehoppers), with about 3,200 species, features exaggerated pronotal structures mimicking thorns or other plant parts for camouflage, primarily on woody hosts in tropical regions.^[16] Aetalionidae includes around 42 species with broad, shelf-like expansions on the pronotum, while Melizoderidae and Myerslopiidae are smaller, with fewer than 10 species each, mostly Neotropical. This superfamily's diversity underscores its adaptive radiation, particularly in exploiting varied plant associations.^[16] Recent taxonomic revisions within Cicadomorpha, driven by molecular data such as transcriptome sequencing, have refined family boundaries and confirmed the monophyly of these superfamilies, with particular emphasis on resolving relationships in Membracoidea.^[17] For instance, anchored phylogenomic analyses have integrated former subfamilies like those previously under Hylicidae into the expanded Cicadellidae, stabilizing the classification amid ongoing discoveries of new species.^[18]

Evolutionary Relationships

The monophyly of Cicadomorpha is robustly supported by both morphological and molecular synapomorphies. Morphologically, the presence of tymbal organs—specialized buckling structures on the abdominal terga used for sound production—represents a key synapomorphy, observed across nearly all families and likely originating as a shared derived trait early in the lineage's evolution.^[19] Molecular evidence further corroborates this, including shared mitochondrial gene arrangements and sequence data from markers such as 18S rRNA and cytochrome c oxidase subunit I (COI), which consistently recover Cicadomorpha as a cohesive clade in analyses spanning 2017 to 2024.^[20]^[21] Within the suborder Auchenorrhyncha, Cicadomorpha forms the sister group to Fulgoromorpha, a relationship affirmed by comprehensive mitogenomic and phylogenomic datasets that resolve Auchenorrhyncha as monophyletic.^[21] Internally, the phylogeny positions Membracoidea (leafhoppers and treehoppers) as the basal superfamily, sister to a clade comprising Cercopoidea (spittlebugs) and Cicadoidea (cicadas), with the latter two more closely related and thus Cicadoidea appearing as the most derived.^[20]^[21] This branching pattern aligns with the three-superfamily structure long recognized in Cicadomorpha. Early phylogenetic studies revealed conflicts between molecular and morphological approaches, particularly regarding the placement of Cercopoidea; for instance, a 2004 molecular analysis based on limited markers suggested Cercopoidea might fall outside the core Cicadomorpha clade, diverging from morphological consensus that emphasized shared feeding and ovipositor traits.^[22] These discrepancies arose from single-gene analyses and sparse taxon sampling, which often produced unstable topologies. Subsequent expanded mitogenome datasets and phylogenomic approaches, including those from 2024 incorporating hundreds of nuclear loci, have resolved these issues, strongly confirming the monophyly of Cicadomorpha and the (Membracoidea + (Cercopoidea + Cicadoidea)) topology across multiple analytical methods.^[20]^[21] Divergence time estimates place the origin of Cicadomorpha in the Permian, approximately 300 million years ago, coinciding with the early diversification of vascular plants. Major radiations within the group occurred during the Cretaceous, paralleling the explosive evolution of angiosperms, which provided new host opportunities and drove adaptive shifts in feeding and habitat use among cicadomorphans.^[9]^[21]

Morphology

External Features

Adult Cicadomorpha exhibit a diverse array of external features, though shared traits reflect their monophyletic origin within the Hemiptera. The body is typically elongate and often wedge-shaped, particularly in the dominant Membracoidea superfamily, which includes leafhoppers and treehoppers, facilitating movement on plant surfaces. The head is prominent, bearing large compound eyes positioned laterally for wide visual fields and three ocelli arranged in a triangle on the vertex, a characteristic feature across the infraorder. Antennae are short and bristle-like, consisting of two basal segments and a flagellum that varies in length but remains inconspicuous.^[9]^[1] Wings comprise two pairs: the forewings, known as tegmina, are leathery or membranous and held roof-like over the body at rest, while the hindwings are folded fan-like beneath them. Venation patterns are diagnostically important for superfamily identification; for instance, Cicadoidea (cicadas) display reticulate venation with numerous crossveins forming a net-like structure, contrasting with the more parallel veins and anteapical cells in Cicadellidae of Membracoidea.^[4]^[9] Legs are adapted for diverse locomotion, with tarsi typically two-segmented, a key autapomorphy of Auchenorrhyncha. In Membracoidea, particularly leafhoppers, the hind femora are enlarged and muscular, enabling powerful jumps for escape; fore and mid legs are slender for walking. Coloration varies widely for camouflage or signaling, ranging from cryptic greens and browns in foliage-dwelling leafhoppers to vibrant patterns in some cicadas. Body size spans approximately 2–70 mm, with minute leafhoppers at the lower end and larger cicadas at the upper.^[23] Sexual dimorphism often manifests in wing length, with females typically shorter-winged than males in many species.^[4]^[9]^[24] Nymphs generally resemble adults in overall form but are apterous, lacking wings, and possess ambulatory legs suited for crawling in many groups, though those in Membracoidea have enlarged hind femora adapted for jumping. In Cercopoidea (spittlebugs), nymphs feature specialized structures for producing protective spittle masses, including modified anal and abdominal glands, while those in Cicadoidea have fossorial forelegs for burrowing.^[25]^[9]^[26]

Internal Anatomy and Adaptations

The digestive system of Cicadomorpha is highly specialized for processing dilute plant sap, featuring a prominent filter chamber in the midgut that enables efficient nutrient concentration. This structure, formed by the looping of the anterior and posterior midgut segments along with the base of the Malpighian tubules and hindgut, facilitates rapid filtration and reabsorption of water and ions, allowing the insects to extract essential amino acids and sugars from low-nutrient xylem or phloem fluids.^[27] In species like the leafhopper Homalodisca vitripennis, the filter chamber is lined with microvilli-rich epithelium supported by mitochondria, enhancing active transport processes.^[28] The midgut itself includes a conical section for initial digestion and a looped portion with perimicrovillar membranes that further aid in nutrient uptake, adapting these insects to their phloem- or xylem-feeding lifestyles across superfamilies such as Cicadoidea and Membracoidea.^[27] Complementing this, the Malpighian tubules in Cicadomorpha are adapted for osmoregulation, managing the excess water intake from sap diets through selective ion reabsorption and waste excretion. These four tubules, originating from the filter chamber junction, are divided into proximal and distal regions with secretory cells containing vesicles and sparse microvilli, enabling the production of uric acid and maintenance of hemolymph balance in hypotonic environments.^[29] Such adaptations are particularly vital in xylem-feeders like those in Cercopoidea, where nutrient scarcity demands precise homeostasis.^[30] The circulatory system in Cicadomorpha follows the typical insect open hemocoel pattern, where hemolymph bathes internal organs directly without enclosed vessels, pumped by a dorsal vessel extending from the abdomen to the head. This system efficiently distributes nutrients, hormones, and immune factors throughout the body cavity, supporting the high metabolic demands of sap-feeding and sound production.^[31] Respiratory functions rely on a tracheal network branching from thoracic and abdominal spiracles, delivering oxygen via diffusion to tissues; in nymphs of Cercopoidea, this system includes adaptations for gas exchange within the moist spittle mass, where the abdomen tip extends externally to access atmospheric oxygen through spiracles while the body remains protected in foam.^[32] Sound production in Cicadomorpha centers on tymbal organs, paired buckling membranes typically located on the first two abdominal terga, vibrated by specialized tensor muscles to generate vibroacoustic signals. These organs feature a convex dome with sclerotized ribs that produce click-like pulses during inward buckling, amplified by adjacent air sacs and resonant structures for species-specific calls.^[19] Variation occurs across superfamilies: in Cicadoidea (cicadas), the tymbals are complex with multiple ribs and extensive air sacs enabling loud airborne sounds exceeding 100 dB, while in Membracoidea and Cercopoidea, simpler ribbed membranes produce primarily substrate-borne vibrations for communication.^[33] This synapomorphic trait is nearly ubiquitous in the suborder, underscoring its evolutionary role in acoustic signaling.^[34] Reproductive anatomy includes paired gonads tailored to the group's oviparous strategies, with males possessing two testes each containing variable numbers of seminal follicles (typically 6–100+ per testis, depending on superfamily) that produce spermatocytes funneled through vasa deferentia to seminal vesicles and accessory glands.^[35] Females have paired ovaries with 3–80 ovarioles per ovary, supplying eggs via lateral oviducts to a common vagina, often equipped with a spermatheca for sperm storage.^[36] The female ovipositor, formed by valvulae from abdominal segments 8 and 9, is a saw-like structure adapted for piercing plant tissues to deposit eggs, as seen in cicadas where it cuts slits in twigs for embedding clutches of 10–20 eggs.^[37] Sensory adaptations in Cicadomorpha emphasize mechanoreception, with Johnston's organ in the antennal pedicel serving as a key vibration detector through arrays of scolopidia that transduce antennal movements into neural signals. This chordotonal organ, containing hundreds to thousands of sensory cells, enables detection of substrate vibrations from conspecifics or environmental cues, facilitating navigation and communication in plant-dwelling species.^[38] In Membracoidea, such as treehoppers, Johnston's organ integrates with other chordotonal arrays to perceive host plant-borne vibrations, allowing precise localization during mate searching and host assessment via signal propagation through stems and leaves.^[39]

Biology and Life History

Feeding Mechanisms

Cicadomorpha possess specialized piercing-sucking mouthparts adapted for extracting plant sap, consisting of a rostrum that encloses a bundle of four interlocking stylets: two mandibular and two maxillary, which together form a food canal and a salivary canal. These stylets enable penetration of plant tissues, allowing access to vascular elements such as phloem sieve tubes or xylem vessels, with the mandibular stylets providing structural support and the maxillary ones facilitating ingestion and salivation.^[40] During penetration, the insects inject watery saliva through the salivary canal to lubricate the path and form a protective sheath around the stylets, preventing plant defenses from clogging the bundle; this sheath is composed of gelling salivary proteins that solidify upon contact with plant substrates.^[41] Salivary enzymes, including pectinases and cellulases, are also secreted to degrade cell walls and facilitate tissue access, while other proteins counteract plant oxidative responses and aid in extracellular digestion of sap components.^[42] Nutrient acquisition strategies vary across Cicadomorpha superfamilies, reflecting dietary specialization on either xylem or phloem sap. Xylem feeders, such as cicadas (Cicadoidea) and spittlebugs (Cercopoidea), ingest large volumes of dilute, mineral-rich xylem fluid under low or negative pressure, requiring continuous pumping via cibarial and pharyngeal muscles to overcome tension and obtain sufficient hydration and ions, though amino acids are scarce.^[43] In contrast, phloem specialists like leafhoppers and treehoppers (Membracoidea) target nutrient-dense phloem sap, which flows under positive turgor pressure and is rich in sugars such as sucrose, allowing passive ingestion once sieve tubes are reached; some species exhibit mixed feeding, ingesting both types sequentially during a single probe.^[44] To compensate for nutritional imbalances in these diets—particularly the low essential amino acid content—Cicadomorpha harbor obligate symbiotic bacteria, such as Candidatus Sulcia muelleri and Hodgkinia cicadicola in cicadas, housed in specialized gut bacteriomes that synthesize missing nutrients through complementary metabolic pathways.^[45] Excess sugars from phloem feeding or water from xylem ingestion are managed through rapid excretion as honeydew, a sticky droplet produced via the anus after filtration in the gut; this waste product often attracts ants in mutualistic trophobiosis, where ants protect the insects in exchange for the carbohydrate-rich secretion.^[46] Host plant specificity ranges from polyphagous, as seen in many leafhoppers that feed across multiple plant families, to monophagous in certain treehopper species restricted to specific hosts, influencing feeding efficiency and population dynamics.^[47] Insects employ behavioral probing sequences, characterized by brief intracellular punctures (X-waves in electropenetration graphs) to sample mesophyll cells and locate suitable vascular bundles, with phloem ingestion typically following successful pathway phases that avoid resistant tissues.^[48] Nymphal feeding differs markedly from adults, with most Cicadomorpha nymphs residing in soil or concealed on stems and roots, using their stylets to tap into xylem or occasionally phloem for prolonged sap ingestion that supports extended development periods.^[49] For instance, periodical cicada nymphs exclusively consume root xylem sap, excavating galleries around host roots to maintain access over years.^[49] This subterranean habit contrasts with adult foliar or twig feeding, and the gut's filter chamber— a modified midgut structure—plays a key role in both stages by rapidly reabsorbing water and concentrating solutes from the ingested fluid.^[3]

Reproduction and Development

Mating in Cicadomorpha primarily relies on acoustic and vibrational signals produced by males to attract females, with species-specific patterns ensuring reproductive isolation. In cicadas (Cicadoidea), males generate loud airborne songs using specialized tymbal organs, which are ribbed membranes on the abdomen that buckle and produce clicks when vibrated by thoracic muscles; these songs vary in frequency, duration, and pattern across species to convey identity and fitness.^[50] In contrast, smaller groups like leafhoppers and treehoppers (Membracoidea) predominantly use substrate-borne vibrations transmitted through plant stems, created by stridulation or tymbal-like mechanisms, allowing females to detect signals via subgenual organs in their legs; these vibrations often form duets where females reply to initiate courtship.^[50] Froghoppers (Cercopoidea) employ similar vibrational communication, though less studied, emphasizing the role of plant substrates in mediating pair formation across the suborder.^[51] Following mating, females of Cicadomorpha use a serrated, saw-like ovipositor to insert eggs into plant tissues, minimizing desiccation and predation risks. This oviposition behavior involves slashing slits in stems, leaves, or bark, where eggs are deposited in linear batches; clutch sizes typically range from 10 to 100 eggs per site, varying by species and environmental conditions, with cicadas often laying larger clutches in woody plants compared to the smaller batches of leafhoppers in herbaceous tissues.^[6] Multiple oviposition sites may be used by a single female over her lifespan, enhancing offspring dispersal while exploiting host plant resources.^[52] Development in Cicadomorpha follows a hemimetabolous pattern of incomplete metamorphosis, lacking a pupal stage and progressing through egg, nymphal, and adult phases. Nymphs undergo 5 to 7 instars, molting to increase size and develop wing pads, which appear as small external buds in later stages and expand progressively; early instars are wingless and resemble miniature adults but with reduced genitalia and mobility adapted for plant-dwelling.^[53] Wing pad development in the final instars signals impending adulthood, with ecdysis revealing fully functional wings in emergent adults.^[54] Parental care is generally absent in most Cicadomorpha, but subsocial behaviors occur in some treehoppers (Membracidae), where females guard egg masses and early nymphs against predators using defensive postures, vibrations, or kicking; this maternal attendance can last until nymphs disperse, improving survival rates in aggregated broods.^[55] Such care is rare elsewhere in the suborder, with no biparental or male-only examples reported.^[56] Sex determination in Cicadomorpha typically follows an XO-XX system, where females are homogametic (XX) and males heterogametic (XO), with the absence of one X chromosome in males leading to sex-specific traits; this system predominates across superfamilies and supports genetic diversity in bisexual populations.^[57] Parthenogenesis is rare, occurring sporadically in a few leafhopper species via thelytoky (unfertilized eggs developing into females), often linked to polyploidy or bacterial infections, but it does not dominate reproductive strategies in the suborder.^[58]

Life Cycle Patterns

Cicadomorpha exhibit a range of life cycle strategies adapted to diverse environmental conditions, with durations varying from annual to multiyear patterns across its superfamilies. In Membracoidea, which includes leafhoppers and treehoppers, most species complete their development within one to two years, often displaying multivoltine life histories in tropical and subtropical regions where multiple generations can occur annually without strong seasonal synchronization.^[59] For example, the treehopper Guayaquila projecta requires approximately 75 days from egg to adult under laboratory conditions, allowing for several overlapping generations in warmer climates.^[60] In temperate zones, these species tend toward univoltine cycles, with overwintering often occurring as diapausing eggs or nymphs to survive cold periods.^[61] In contrast, Cicadoidea, encompassing cicadas, feature prolonged subterranean nymphal phases lasting 2 to 17 years, during which nymphs feed on root xylem fluids underground before emerging as adults for a brief 2- to 6-week period.^[62] Periodical cicadas of the genus Magicicada exemplify extreme synchrony with prime-numbered cycles of 13 or 17 years, an adaptation that minimizes overlap with predator cycles and hybridizations, enhancing population persistence.^[63] Emergence is triggered by soil temperatures reaching approximately 18°C (64°F) at an 8-inch depth, leading to mass synchronized events that facilitate predator satiation, where overwhelming numbers reduce per-individual predation risk despite high overall mortality from predators consuming 15-40% of adults post-reproduction.^[62]^[63] Cercopoidea, known as spittlebugs or froghoppers, typically follow 1- to 3-year cycles, with nymphs developing over 4-9 weeks across five instars while feeding on xylem sap.^[64] Nymphs produce protective spittle masses from plant fluids mixed with mucopolysaccharides and proteins, shielding them from desiccation, predators, parasitoids, and extreme temperatures during this vulnerable stage.^[64] Voltinism varies geographically, with up to six generations per year in humid tropical areas, while temperate populations are often univoltine or bivoltine, sometimes incorporating diapause in overwintering eggs or early nymphs.^[64] Across Cicadomorpha, voltinism shifts from univoltine in temperate latitudes—where diapause in nymphs or eggs aligns development with seasonal availability—to multivoltine in the tropics, reflecting adaptations to temperature and resource predictability.^[59] Emergence synchrony, particularly in periodical species, relies on environmental cues like soil temperature to coordinate mass events, promoting predator satiation as a key survival mechanism.^[63] These patterns underscore the group's evolutionary flexibility in timing life stages to optimize feeding, protection, and reproduction amid varying ecological pressures.

Ecology and Distribution

Habitats and Global Range

Cicadomorpha, comprising approximately 33,000 extant described species of plant sap-feeding insects, are cosmopolitan in distribution, occurring on all continents except Antarctica and ranging from equatorial regions to temperate zones, though they are absent from polar extremes.^[1]^[65] They inhabit diverse terrestrial environments associated with vascular vegetation, including forests, grasslands, deserts, wetlands, and even urban areas, wherever suitable host plants are present.^[9]^[15] Nymphs of many species, particularly in families like Cicadidae and Cercopidae, are often subterranean or root-dwelling, burrowing into soil to feed on plant roots, while adults are typically arboreal or herbaceous.^[66]^[67] The group's highest species diversity is concentrated in tropical regions, with significant radiations in Asia and South America, where over half of all Cicadomorpha species occur, driven by abundant vegetation and climatic stability.^[68]^[69] Biogeographic patterns include notable endemism in Australia, which hosts unique cicada assemblages with 750–1,000 species, many restricted to eucalypt-dominated habitats.^[70] Some species, such as the glassy-winged sharpshooter (Homalodisca vitripennis) in the Cicadellidae, have become invasive outside their native ranges, spreading to new regions like North America and Europe.^[71] Cicadomorpha exhibit a broad altitudinal range, from sea level to elevations exceeding 4,000 meters, as seen in Andean populations of cicadas and leafhoppers adapted to high-altitude páramos and puna ecosystems.^[72] Adaptations to arid environments, such as xylem-feeding that allows survival in water-stressed plants, enable persistence in deserts and semi-arid scrublands across Africa, Australia, and the Americas.^[9] Their global expansion correlates with the mid-Cretaceous radiation of angiosperms, which provided diverse host plants and facilitated diversification into new habitats following gymnosperm declines.^[21]

Ecological Interactions

Cicadomorpha species primarily interact with plants through phloem or xylem sap feeding, which extracts nutrients but often results in minor direct damage such as localized wilting or reduced plant vigor. This feeding punctures vascular tissues, prompting host plants to activate defense responses including the production of reactive oxygen species, jasmonic acid signaling, and volatile organic compounds that may attract natural enemies of the insects. For instance, leafhoppers in the family Cicadellidae induce systemic acquired resistance in crops like rice, enhancing the plant's expression of pathogenesis-related proteins. While such interactions are generally antagonistic, mutualistic associations occur in some groups; treehoppers (Membracidae) and certain leafhoppers excrete honeydew, a sugary byproduct that attracts ants, which in turn defend the insects from predators in exchange for the resource. These ant-tending behaviors are documented in species like Umbonia crassicornis, where ants actively remove competitors and parasitoids from nymph clusters.^[73]^[74]^[75] Predators and parasitoids exert significant pressure on Cicadomorpha populations across life stages. Vertebrate predators such as birds (e.g., warblers and sparrows) and small mammals target adult cicadas and leafhoppers, while invertebrates like spiders (e.g., orb-weavers) and predatory wasps (e.g., Cicada killer wasps, Sphecius speciosus) consume nymphs and adults. Parasitoids, predominantly hymenopterans from families like Dryinidae and Mymaridae, oviposit into eggs or nymphs of leafhoppers and treehoppers, with species such as Gonatopus flavifemur achieving high parasitism rates in grassland habitats. Some Cicadomorpha counter these threats with chemical defenses, including volatile emissions from disturbed tissues, and physical barriers like waxy coatings on nymphs of spittlebugs (Cercopidae), which reduce desiccation and deter small parasitoids. These interactions help regulate population densities, preventing outbreaks in natural ecosystems.^[76]^[15]^[77] In food webs, Cicadomorpha serve as a foundational prey base for multiple trophic levels, supporting biodiversity in terrestrial and riparian ecosystems. Nymphs and adults are consumed by a diverse array of predators, from insects to amphibians and birds, contributing to energy transfer; for example, periodical cicada emergences (Magicicada spp.) provide a pulsed resource boom that sustains bird populations and influences foraging behaviors across over 80 species. Their frass and decomposing carcasses enhance nutrient cycling by returning nitrogen and phosphorus to soils, stimulating microbial activity and plant growth—studies during 2021 emergences showed increased soil nematode abundances and altered bacterial-fungal ratios due to cicada inputs. This role is particularly pronounced in forests, where mass emergences temporarily "rewire" trophic interactions, reducing herbivory on foliage as predators shift focus.^[78]^[79]^[80] Cicadomorpha act as vectors for plant pathogens, including phytoplasmas and viruses, acquired during sap feeding and transmitted to new hosts. Leafhoppers like Scaphoideus titanus efficiently spread Candidatus Phytoplasma vitis, causing flavescence dorée in grapevines, while other species vector aster yellows phytoplasmas affecting broadleaf plants. Transmission is persistent or propagative, with insects harboring pathogens in salivary glands, though not all species are competent vectors. Intraspecific competition within Cicadomorpha often involves resource partitioning on host plants, where individuals exploit different plant parts, tissues, or phenological stages to minimize overlap; for example, ovipositing female cicadas select distinct branch sizes or heights, reducing egg-laying interference and larval crowding on limited xylem resources. Such partitioning is evident in grassland leafhoppers, where temporal shifts in adult activity allow coexistence on shared grasses without severe density-dependent mortality.^[81]^[82]^[83]

Economic Significance

Cicadomorpha, particularly species in the family Cicadellidae (leafhoppers), represent significant agricultural pests through direct feeding damage on major crops such as grapes and rice, where nymphs and adults extract plant sap, leading to stippling, chlorosis, and reduced photosynthesis.^[84]^[85] Treehoppers (Membracidae) similarly inflict damage on ornamental plants by sucking sap from stems and leaves, causing defoliation, honeydew excretion, and subsequent sooty mold growth that impairs aesthetics and plant health.^[86] Many Cicadomorpha species serve as vectors for devastating plant pathogens, exacerbating their economic toll. Leafhoppers transmit Xylella fastidiosa, the causative agent of Pierce's disease in vineyards, which blocks xylem vessels and leads to vine decline and death.^[71] They also vector aster yellows phytoplasma, affecting crops like carrots, lettuce, and cereals by inducing witches' broom symptoms and yield reductions.^[87] Collectively, these diseases result in significant economic losses, with potential annual economic losses exceeding €5.5 billion in Europe alone from X. fastidiosa in olive and grape production.^[88] Management of Cicadomorpha pests relies on integrated pest management (IPM) strategies, combining cultural practices, monitoring with sticky traps, targeted insecticides, and biological controls such as egg parasitoids in the genus Anagrus.^[89]^[90] While predominantly detrimental, Cicadomorpha offer minor benefits, including incidental pollination by some flower-visiting species and cultural significance through cicada emergences celebrated in festivals like CicadaFest in the United States.^[91] No major beneficial species within the group contribute substantially to agriculture or ecosystems in economic terms. Invasive Cicadomorpha species amplify these impacts, such as Erythroneura ziczac (three-banded leafhopper), which spread to California vineyards via international trade since the early 2010s, intensifying feeding damage on grapes.

Fossil Record and Evolution

Known Fossils

The fossil record of Cicadomorpha begins in the late Carboniferous (approximately 308–299 Ma), extending through the Permian period (299–252 Ma), with early representatives documented from deposits in European Russia and China, including members of the Dysmorphoptilidae family that display primitive wing venation featuring extensive crossveins and reticulate patterns indicative of basal hemipteran morphology.^[92]^[93] These specimens, often preserved as wing impressions, highlight the group's initial diversification among Paleozoic flora.^[9] Mesozoic deposits reveal a marked increase in Cicadomorpha diversity during the Jurassic and Cretaceous periods, with over 200 fossil species described across various superfamilies, including extinct lineages like the Palaeontinidae, which exhibit large body sizes and specialized venation suggesting adaptations to early gymnosperm hosts.^[94] Amber inclusions from the mid-Cretaceous (ca. 99 Ma) Burmese amber of Myanmar preserve early Cicadidae, such as Burmacicada protera, showcasing detailed foreleg structures akin to modern digging behaviors in nymphs.^[95] Jurassic compression fossils from sites like Daohugou in China further document primitive cercopoids with elongated wings.^[96] Cenozoic records demonstrate continuity and refinement of Cicadomorpha forms, with Eocene (ca. 50 Ma) compression fossils from the Green River Formation in North America yielding extant-like leafhoppers (Cicadellidae) that closely resemble modern species in wing shape and body proportions, underscoring long-term morphological stability. Oligocene (ca. 34–44 Ma) Baltic amber inclusions preserve cicadas with intact sound-producing organs, including tymbals and opercula, allowing reconstruction of their acoustic apparatus.^[97] These amber specimens also capture treehoppers (Membracidae) from Eocene deposits, evidencing rapid diversification of pronotal ornamentation during the early Paleogene. Fossils of Cicadomorpha occur primarily through compression and impression in fine-grained sediments, as well as exceptional three-dimensional preservation in amber, though challenges arise in superfamily-level identification due to the predominance of isolated wing fossils that obscure body and genitalic features critical for higher taxonomy.^[9] Notable basal taxa include Palaeontinidae from Triassic to Cretaceous strata, representing early divergent forms with robust venation, while Eocene Membracidae fossils indicate accelerated cladogenesis linked to angiosperm expansion.^[98]^[99]

Phylogenetic Origins and Diversification

The Cicadomorpha suborder traces its origins to Paleozoic ancestors within the Hemiptera order, with Bayesian analyses of the fossil record estimating the emergence of Cicadomorpha around 308 million years ago (Ma) in the late Carboniferous.^[100] This timeline aligns with the broader Hemiptera radiation during the Pennsylvanian period, approximately 321 Ma, where early lineages like Aviorrhynchidae represent basal forms.^[10] By the middle Permian, around 280 Ma, Cicadomorpha had developed key innovations such as piercing-sucking mouthparts and enlarged cibarial muscles adapted for xylem sap feeding, alongside early jumping capabilities in hind legs that facilitated escape from predators.^[9] These traits marked a shift from generalized hemipteran feeding to specialized filter-feeding mechanisms, with modern superfamilies (Cicadoidea, Cercopoidea, Membracoidea) diversifying by the Triassic, approximately 252–201 Ma, as evidenced by the first records of cicada-like, spittlebug-like, and leafhopper-like forms.^[9] Diversification accelerated dramatically during the Cretaceous period (145–66 Ma), coinciding with the explosive radiation of angiosperms and driving co-evolutionary dynamics with host plants that spurred splits within superfamilies.^[101] For instance, cicadas (Cicadoidea) underwent a host shift from gymnosperms to angiosperms around the mid-Cretaceous (~99 Ma), enabling exploitation of new phloem-rich resources and contributing to family-level expansions in Cercopoidea and Membracoidea.^[101] This period saw the origin of many modern genera, with global tectonic shifts and vegetation changes further promoting lineage proliferation, as reflected in fossil assemblages from amber deposits.^[9] The end-Permian mass extinction (~252 Ma) imposed minor losses on Cicadomorpha, with genus-level extinction rates approximately 6-fold above background levels for herbivorous lineages like Hemiptera, yet the group survived through adaptive shifts to resilient vegetation post-extinction.^[102] These transitions facilitated recovery and set the stage for Mesozoic dominance. In contemporary ecosystems, Cicadomorpha exhibit high speciation rates particularly in tropical regions, where over 33,000 species thrive, fueled by ongoing host shifts and microhabitat adaptations.^[103] Recent 2024 phylogenomic studies on Membracoidea highlight this persistence, revealing mid-Cretaceous origins (~140–90 Ma) and multiple independent shifts from arboreal to litter/soil hosts, underscoring active diversification via plant-insect interactions.^[104] Future refinements to Cicadomorpha evolutionary timelines will require expanded molecular clock calibrations incorporating fossil data, as current models reveal biases from underutilizing extinct diversity and highlight discrepancies between molecular estimates and the rich Permian-to-Cretaceous record.^[99]

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