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Neotibicen

Neotibicen is a of large-bodied annual cicadas in the family , subfamily Cicadinae, and tribe Cryptotympanini, native primarily to eastern and central , where adults emerge during the late summer "" and produce loud, species-specific calling songs to attract mates. The genus comprises 13 recognized , including N. tibicen, N. lyricen, and N. canicularis, all characterized by robust bodies, broad heads, prominent compound eyes, and clear wings with distinct venation patterns. Established in through molecular phylogenetic and morphological analyses, Neotibicen was separated from the polyphyletic genus Tibicen to reflect the distinct North American clade, with diversification estimated to have occurred after the mid-Miocene. These cicadas, commonly known as dog-day, harvest, or jar flies (though not true locusts), inhabit a range of environments from woodlands and forests to urban areas across the , southeastern , and parts of , with one species, N. bermudianus, historically occurring in but now considered extinct. Species exhibit annual life cycles, with nymphs spending 2–5 years underground feeding on root before emerging en masse to molt into adults that live for several weeks, during which males perch on vegetation and produce continuous trills or buzzes using specialized organs. Notable for their ecological role in soil aeration and as prey for , bats, and spiders, Neotibicen species show variation in coloration—often featuring green, black, or pruinose (frosted) markings—and some form hybrid zones where ranges overlap, such as between N. tibicen .

Taxonomy

Classification History

The genus Neotibicen was historically classified within the broader genus Tibicen Latreille, , which encompassed a diverse array of North American cicadas north of , along with some Eurasian species, based primarily on superficial morphological similarities such as body size and coloration. Early subdivisions within Tibicen were proposed through morphological analyses, such as those by () identifying eastern, central, and western groups, but lacked robust phylogenetic support until molecular data became available. In , a comprehensive study by , , Moulds, and utilized integrated morphological, molecular (mitochondrial and nuclear EF-1α sequences from 35 North American ), and acoustic analyses to revise the of Tibicen, establishing Neotibicen Hill & Moulds as a new for 21 eastern and central North American , including the Cicada canicularis Harris, 1841. This reclassification split Tibicen sensu lato into several distinct genera, including Neotibicen for the focal , Hadoa Moulds for 20 southwestern (e.g., Tibicen duryi , 1917), and Lyristes Stål for certain Eurasian taxa, with the changes justified by cladistic analysis of 27 morphological characters (e.g., head width, operculum shape, male genitalia) and phylogenetic trees showing strong support (bootstrap values >90% for Neotibicen). Species-specific calling songs provided additional diagnostic evidence, reinforcing boundaries where was ambiguous. Phylogenetically, Neotibicen is positioned within the subtribe Tacuina Distant, 1904, of the tribe Cryptotympanini in the subfamily Cicadinae, forming a monophyletic closely related to other North American genera such as Hadoa, Megatibicen, Cacama, and Cornuplura, based on multi-gene analyses indicating shared Asian ancestry and divergence during invasions. This placement highlights Neotibicen's affinities with southwestern and periodic lineages, with molecular divergence estimates suggesting the clade originated 15–20 million years ago. Post-2015 updates have refined Neotibicen's boundaries through targeted bioacoustic and morphological studies; for instance, in , Sanborn and Heath erected Megatibicen for larger-bodied species previously assigned to Neotibicen (e.g., Tibicen auletes Germar, 1830), based on body size metrics and operculum morphology exceeding those of typical Neotibicen. Additionally, a bioacoustic investigation by Marshall and Hill described Neotibicen similaris apalachicola as a new from the Apalachicola region of the , distinguished by a divergent male courtship song despite morphological , and noted hybrid zones with the nominate form. These revisions underscore the role of acoustic data in resolving cryptic diversity within the .

Species and Subspecies

The genus Neotibicen comprises 13 recognized , primarily distributed across , with one endemic to ; these annual cicadas were delineated as a distinct genus in a 2015 phylogenetic revision based on molecular, morphological, and acoustic data. The species are characterized by their summer emergence and distinctive calls, though specific traits such as song structure aid in differentiation. The following table lists the recognized species, including scientific names, authorities, and common names where established:
Scientific NameAuthorityCommon Name
Neotibicen auriferus(Say, 1825)Plains dog-day cicada
Neotibicen bermudianus(Verrill, 1902)
Neotibicen canicularis(Harris, 1841)
Neotibicen davisi(Smith & Grossbeck, 1907)
Neotibicen latifasciatus(Davis, 1915)Coastal scissor-grinder cicada
Neotibicen linnei(Smith & Grossbeck, 1907)
Neotibicen lyricen(De Geer, 1773)Lyric cicada
Neotibicen pruinosus(Say, 1825)Scissor-grinder cicada
Neotibicen robinsonianus(Davis, 1922)Robinson's cicada
Neotibicen similaris(Smith & Grossbeck, 1907)Similar dog-day cicada
Neotibicen superbus(Fitch, 1855)Superb dog-day cicada
Neotibicen tibicen(Linnaeus, 1758)Swamp cicada
Neotibicen winnemanna(Davis, 1912)Winnemanna's cicada
Several species bear etymologies tied to their songs or seasonal associations; for instance, N. canicularis derives from the Latin canicula (little dog), referencing the "dog days" of late summer when it sings most prominently. Similarly, N. tibicen stems from the Latin tibicen (flute-player), alluding to its piping call, while N. lyricen evokes the lyre-like quality of its . Subspecies are recognized within several based on regional variations in acoustics and ; notable examples include N. tibicen tibicen (the nominate subspecies) and N. tibicen australis (southern form), as well as N. lyricen lyricen, N. lyricen engelhardti, and N. pruinosus fulvus. A recent addition is N. similaris apalachicola (Marshall & Hill, ), described from the Apalachicola region of the southeastern U.S. using song analysis and evidence, forming hybrid zones with the nominate N. similaris despite distinct calls. As of 2025, no further taxonomic revisions have elevated additional debated variants to status, though ongoing genetic studies continue to refine boundaries within the genus.

Morphology and Description

Adult Characteristics

Adult Neotibicen cicadas are large, robust typically measuring 25-51 mm in body length, characterized by a sturdy build and broad heads that contribute to their distinctive appearance among cicadids. Their coloration is predominantly green, often accented with black, brown, or pruinose (frosty white) markings that provide against tree bark and foliage. These patterns vary subtly across individuals and populations, reflecting adaptations to diverse woodland environments. Species-specific coloration enhances identification; for instance, N. canicularis often displays prominent black dorsal stripes along the and , contrasting with its green base, while N. auriferus exhibits golden-tan hues blended with greenish tones, particularly in prairie populations. These variations arise from genetic and environmental factors. Key morphological features include prominent red or black compound eyes positioned laterally for wide visual fields, complemented by three ocelli on the for additional light detection. The legs are robust and spined, adapted for secure clinging to during perching and calling. Sexual dimorphism is evident, with males possessing tymbals—ribbed membranes for sound production—and opercula that partially cover these organs on the abdomen's underside, structures briefly referenced in behavioral contexts for acoustic signaling. The wings are transparent with conspicuous dark veins, spanning 50-80 mm, and are held in a characteristic roof-like position over the body at rest, aiding in flight and .

Nymphal Characteristics

Neotibicen nymphs, adapted for a subterranean , reach maturity at lengths of 15–30 mm and exhibit a robust, arched resembling that of a , which facilitates burrowing through . Their coloration is typically dull brown or gray, providing effective against soil backgrounds, complemented by a thick, protective that withstands underground pressures. Prominent adaptations include powerful forelegs equipped with rake-like spines and teeth on the femora and tibiae for excavating tunnels, piercing-sucking mouthparts designed to extract fluids from plant roots, and the development of rudimentary wing pads visible in later instars. These nymphs undergo five developmental instars, during which morphological changes occur, such as progressive increases in body segmentation, from opaque white to functional, and overall size enlargement across stages.

Distribution and Habitat

Geographic Range

The genus Neotibicen is distributed primarily across eastern and central , spanning from southern , including , southward through the to the Gulf Coast, and extending westward to the . This range encompasses diverse regions such as the Midwest, Southeast, and areas, where the cicadas are most abundant in summer months. Historically, Neotibicen included a now-extirpated population in , represented by the endemic species N. bermudianus, which disappeared due to and the near-eradication of its primary host plant, the Bermuda cedar (), following a in the 1940s. The core of the genus's current distribution lies in the deciduous forests of the U.S. Midwest and Southeast, where multiple species overlap. Species distributions vary within this overall range; for instance, N. tibicen is widespread across much of the eastern and , from the Atlantic coast to the valley. In contrast, N. auriferus is more restricted to central U.S. states, including the regions of , , and . Other species, such as N. canicularis, extend northward into southern and southward to northern , while southeastern endemics like N. robinsonianus are concentrated in highland areas of , , and . The geographic range of Neotibicen is influenced by suitability, particularly warm, humid summers that support and , as well as the availability of host trees—such as oaks (Quercus spp.) and maples (Acer spp.)—essential for egg-laying and nymphal feeding on root . Recent modeling suggests potential northward expansions for some species, like N. winnemanna, driven by warming temperatures associated with , though no widespread shifts have been documented as of 2025.

Habitat Preferences

Neotibicen species primarily inhabit woodlands, edges, and areas with mature trees, including urban and suburban settings, parks, orchards, and oak-hickory woodlands. These cicadas favor environments that support their , such as open , edges, and streamside habitats, where nymphs can burrow and feed undisturbed for several years. For instance, Neotibicen lyricen is commonly associated with upland forests and wooded residential areas, while Neotibicen pruinosus thrives in mixed woodlands and suburban yards with trees. Nymphs of Neotibicen require fertile soils that sustain complex root systems of host plants, typically loamy or well-drained types conducive to burrowing and long-term development. They feed on the xylem sap from roots of various deciduous trees, including oaks (Quercus spp.), maples (Acer spp.), hickories (Carya spp.), ashes (Fraxinus spp.), elms (Ulmus spp.), beeches (Fagus spp.), persimmons (Diospyros spp.), walnuts (Juglans spp.), and even peach trees (Prunus persica). Adults oviposit eggs into the twigs of similar deciduous hardwoods, ensuring proximity to suitable root hosts for the emerging nymphs, though some species occasionally use conifers. Certain species exhibit specific microhabitat preferences influenced by moisture and vegetation structure; for example, Neotibicen tibicen is often found in swampy or marshy lowlands near water bodies, such as river banks or edges with shrubs and tall weeds. In contrast, species like Neotibicen canicularis occupy a broader range, from thick forests to settings, as long as trees are present for perching and calling. These habitats generally occur in warm, humid climates during summer, with adults emerging when soil temperatures reach about 18°C (64°F) at 8-inch depths, supporting their preference for temperate to subtropical regions across eastern .

Life Cycle

Developmental Stages

Neotibicen cicadas, like other members of the family , undergo hemimetabolous (incomplete) , characterized by three primary developmental stages: egg, , and , without a distinct pupal phase as seen in holometabolous insects. This allows nymphs to gradually resemble adults through a series of molts, with wing development occurring progressively during the nymphal instars rather than in a separate transformative stage. The egg stage begins when adult females use their to create slits in the of twigs, typically on trees or shrubs, and deposit clusters of 10 to 20 eggs per slit, with a single female capable of laying up to several hundred eggs over multiple sites. These eggs are elongated and cigar-shaped, measuring approximately 1.5 to 3 mm in length, and are creamy white in color. occurs after 6 to 10 weeks, depending on environmental conditions such as and , at which point the first-instar nymphs emerge and drop to the ground to begin their subterranean life. Upon reaching the soil, the nymphs burrow downward and initiate the longest phase of their development, lasting 2 to 5 years for Neotibicen species, during which they feed exclusively on drawn from the roots of trees and shrubs using piercing-sucking mouthparts. Nymphs undergo five instars, molting four times to accommodate growth; early instars are small and pale, while later ones develop more robust forelegs adapted for digging and visible wing pads that foreshadow the adult form. Throughout this period, they remain underground, constructing tunnels and chambers, with feeding and molting synchronized to seasonal root availability; detailed morphological features of these nymphs, such as their forelegs, are described elsewhere. In the final nymphal instar, individuals construct an emergence tunnel to the soil surface and form a preparatory chamber, where they undergo physiological changes including the expansion and pigmentation of wing pads in preparation for the terminal molt. This pre-emergence phase is brief, lasting days to weeks, and culminates in the nymph crawling out at night, climbing a vertical surface, and shedding its to reveal the adult; the discarded nymphal skin, or , remains as evidence of this transformation. The adult stage is short-lived, typically enduring 4 to 6 weeks, during which the primary focus is , though adults actively feed on or from trees and flowers to sustain energy needs, countering earlier misconceptions of non-feeding. Freshly molted adults are pale and soft but harden and expand their wings within hours, achieving full coloration and functionality soon after; this stage enables dispersal, calling, and oviposition to perpetuate the cycle.

Cycle Duration and Emergence

Neotibicen species exhibit an life cycle characterized by a prolonged subterranean nymphal lasting 2 to 5 years, in to the 13- or 17-year cycles of in the genus Magicicada. This extended underground development involves nymphs feeding on root , with overlapping cohorts from successive years ensuring that adults emerge annually rather than synchronously in massive broods. The asynchrony among year classes results in consistent yearly appearances across their range, though local populations may show partial synchronization within a season. Emergence typically occurs from late summer to early fall, spanning through , depending on and local conditions. This timing is primarily triggered by soil temperatures reaching approximately 18–20°C (64–68°F) at depths of 15–20 cm, often following warm rains that facilitate nymphal exit from the . In local areas, emergences are relatively synchronous over a few weeks, creating noticeable concentrations of adults, with males generally appearing slightly ahead of females to establish calling sites.

Behavior

Communication and Sound Production

Males of the Neotibicen produce species-specific calls primarily for attraction and species recognition using specialized organs located on the sides of the . These organs consist of ribbed membranes that are rapidly vibrated by contractions of internal tymbal muscles, generating a series of clicks as the ribs buckle inward and snap back. The resulting sound is amplified by large within the abdominal cavity, which act as resonators to enhance volume and projection. Call characteristics vary distinctly among Neotibicen species, aiding in . For instance, N. canicularis produces a continuous, high-pitched whining resembling an electric saw buzz, lasting about 15 seconds and starting softly before peaking and tapering off, with a dominant frequency around 7 kHz and intensities reaching up to 106 dB at close range. In contrast, N. lyricen emits a buzzy, rattling without strong pulsations, often described as lyrical due to its smoother phrasing, extending 30 seconds to over a minute with gradual volume increases. These songs generally span frequencies of 2-10 kHz and can exceed 100 dB in choruses. Calling typically occurs in late afternoon or early evening during warm weather, when males perch in and synchronize into choruses to amplify collective signaling and deter predators. These diurnal patterns align with peak activity in woodlands, where group singing creates a pervasive auditory . Females respond to appropriate conspecific calls by flicking their wings in timed patterns, producing brief clicking sounds that signal receptivity and guide males to their location. Such acoustic divergence plays a key role in Neotibicen , as distinct traits prevent interbreeding even in sympatric zones, with studies revealing cryptic taxa differentiated primarily by call structure despite morphological similarities. For example, hybrid zones between show limited due to mismatched acoustic signals, underscoring sound's barrier to .

Mating Behaviors

Males of Neotibicen species typically initiate by perching on elevated , such as trunks or branches, and producing loud, species-specific calls to attract receptive females from a distance. Females respond to these calls by approaching the male and signaling interest through rapid wing flicks, which generate a subtle clicking sound detectable at close range. Upon arrival, the pair engages in physical contact to confirm mutual receptivity before copulation commences. Mating in Neotibicen involves the male transferring sperm via his to the female's . Females often multiple times with different males to maximize fertilization success and in their offspring. Following , females select living twigs of trees or shrubs—typically pencil-sized in diameter—and use their saw-like to make parallel slits in the bark, depositing eggs in clusters of 10 to 20 per slit. A single female lays 200 to 600 eggs across multiple sites over several weeks, a process that can weaken twigs and cause flagging, where affected branches wilt and die back, though significant tree damage is rare. Species isolation in Neotibicen is primarily maintained through acoustic and behavioral barriers, where distinct calling patterns and female response preferences prevent interspecific mating. These prezygotic mechanisms ensure reproductive fidelity, though rare hybridization occurs in contact zones between closely related taxa, such as N. tibicen and a newly described subspecies.

Ecology

Predators and Parasites

Neotibicen cicadas face predation from a variety of vertebrates and throughout their . Adult cicadas are commonly consumed by birds such as , blue jays, and cardinals, which target them during flight or while perched on . Mammals like squirrels and moles prey on both adults and emerging nymphs, often near bases where nymphs climb to molt. Among insects, the eastern cicada killer wasp (Sphecius speciosus) is a primary predator of adult Neotibicen, with females stinging and paralyzing s to provision nests for their larvae; this wasp specializes in annual cicadas and does not attack periodical species. Parasitic organisms also exert significant pressure on Neotibicen populations. Fungi, including species in the genus , can infect adults, leading to sterility by replacing genital structures with spore masses that spread the pathogen during mating attempts, though such infections are more commonly documented in related genera. Nematodes infest both nymphs and adults, burrowing into tissues and potentially reducing and . Mites frequently parasitize , attaching to the body surface or entering orifices to feed on , which can weaken hosts and increase susceptibility to other threats. Additionally, sarcophagid flies such as Emblemasoma erro larviposit eggs into adult , with larvae developing internally and emerging to pupate, often resulting in host death; while typically targeting males via acoustic , of females has been observed in . Neotibicen employ several behavioral and morphological defenses against these threats. Their bark-like coloration provides against visual predators like birds and squirrels, blending seamlessly with trunks. Sudden flashes of bright underwings during flight serve as startle displays to deter approaching predators. Annual emergences, though less synchronized than in , occur in sufficient numbers to overwhelm predators locally through satiation, ensuring some individuals survive to reproduce. Predation and contribute to high mortality rates among Neotibicen, with estimates indicating up to 90% of emerging may succumb to these factors during outbreaks, compounded by vulnerabilities in soft-bodied nymphs and short lifespans. Despite this, populations remain resilient due to the high of females, who can lay 200–600 eggs per individual, compensating for losses and maintaining stable numbers across their range.

Ecological Role and Human Interactions

Neotibicen species play a key role in nutrient cycling within their ecosystems, as nymphs feed on xylem sap from tree roots for years underground, extracting nutrients that are later returned to the soil upon adult emergence. When adults emerge in summer, their bodies, molted exuviae, and eventual decomposition deposit significant biomass—often in large numbers—enriching forest floors and grasslands with nitrogen and other elements, thereby enhancing soil fertility and supporting plant growth. This pulsed return of nutrients can stimulate microbial activity and benefit understory vegetation, particularly in deciduous woodlands where Neotibicen are prevalent. As a prey base for , Neotibicen serve as an important food source for various predators, including , mammals, spiders, and specialized like the Eastern Cicada Killer wasp (), which provisions nests with paralyzed adults. Their abundance during annual emergences provides seasonal energy boosts to these consumers, contributing to dynamics in eastern North American forests and prairies. Neotibicen cicadas function as indicator species for quality due to their specific requirements for mature trees and undisturbed soil, making them sensitive to . Populations decline in fragmented or polluted areas, allowing researchers to use their presence and density in monitoring programs to assess health and integrity in regions like the Midwest and Southeast. In interactions, Neotibicen hold cultural significance in North , often symbolizing summer's arrival and renewal, with their loud calls evoking themes of transformation akin to broader symbolism in and early settler traditions. They occasionally serve as informal indicators of seasons in rural areas, signaling the end of crop growth with their mid-summer choruses. As pests, females cause minor damage by slitting twigs for egg-laying, which can lead to dieback in young orchard trees like apples and peaches, though this rarely poses major economic threats due to targeted management like netting. Conservation efforts for Neotibicen remain stable overall, with most species classified as least concern, but certain subspecies face vulnerability from that reduces suitable habitats. is expected to drive northward range shifts in species like Neotibicen tibicen, as indicated by ecophysiological studies, with recent observations suggesting expansions for related species such as N. winnemanna.

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