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Neotibicen linnei

Neotibicen linnei, commonly known as Linne's cicada, is a species of large-bodied annual cicada in the genus Neotibicen within the family Cicadidae.^[1] It is characterized by a robust black body with green or dull yellow markings, a body length of 28–34 mm, prominent eyes, a strongly bowed costal margin on the forewings, elongated male opercula, and an uninterrupted black face mask that is variable in southern populations.^[2]^[1] The species produces a distinctive song described as a rapid "Zeger-Zeger-Zeger" or whirring rattle, which aids in its identification and distinguishes it from similar species like Neotibicen pruinosus.^[1]^[3] First described as Cicada linnei in 1907 by John B. Smith and John A. Grossbeck based on specimens from Massachusetts, the species was long confused with Cicada tibicen of Linnaeus but recognized as distinct due to differences in size, thoracic markings, and wing venation.^[2] In 2015, the genus Neotibicen was established through molecular phylogenetic analysis, transferring N. linnei from the former genus Tibicen to reflect its evolutionary relationships within the Cryptotympanini tribe; this revision highlighted Neotibicen as a clade of medium to large cicadas with bulbous male basisternum 3 and protruding opercula.^[4] The species belongs to the east-central North American Neotibicen clade, with divergence estimated around the late Miocene approximately 11 million years ago.^[4] N. linnei is widely distributed across the eastern United States, from Massachusetts and New York westward to Michigan, Illinois, and Nebraska, southward to Florida, Texas, and Tennessee, and northward into adjacent southern Canada.^[1]^[4] It occurs in diverse habitats, primarily deciduous forests and woodlands with hardwoods such as oaks and hickories, though some populations in central Tennessee associate with junipers; adults feed on tree sap from various hardwoods and conifers.^[1]^[4] Unlike periodical cicadas, N. linnei has an annual life cycle, with nymphs developing underground for 2–5 years before emerging as adults primarily from June to October in most regions, extending to September–November in central Florida.^[1] The species exhibits sexual dimorphism, with males featuring larger basal abdominal spots, a longer second abdominal segment, and light-colored opercula as long as they are broad; females lack these spots and have similar genitalia to related species like N. canicularis.^[2] Potential hybridization with congeners such as N. pruinosus and N. winnemanna has been noted in overlap zones, particularly in the upper mid-South, though further taxonomic study is needed to clarify boundaries.^[1] As part of the "dog-day cicadas," N. linnei contributes to the summer soundscape of eastern North American forests, playing a role in ecosystems as prey for birds and predators while aiding in nutrient cycling through leaf litter from cast nymphal skins.^[3]

Taxonomy and nomenclature

Classification

Neotibicen linnei is classified in the domain Eukarya, kingdom Animalia, phylum Arthropoda, class Insecta, order Hemiptera, superfamily Cicadoidea, family Cicadidae, subfamily Cicadinae, tribe Cryptotympanini, subtribe Cryptotympanina, genus Neotibicen, and species N. linnei.^[5] This hierarchical placement situates the species among the true cicadas, distinguished by their sap-feeding habits and acoustic signaling behaviors typical of the Hemiptera order.^[6] The species was originally described as Cicada linnei by entomologists John B. Smith and John A. Grossbeck in 1907, based on specimens from Massachusetts.^[5] In 2015, a comprehensive molecular phylogenetic study led to significant taxonomic revisions within the North American cicadas of the former Tibicen genus, resulting in the establishment of Neotibicen as a new genus to encompass a monophyletic clade of larger annual species, including linnei. This change was driven by analyses of mitochondrial and nuclear DNA sequences that revealed distinct evolutionary lineages not reflected in the prior broad Tibicen classification.^[4] Placement within the tribe Cryptotympanini relies on morphological diagnostics, including the specialized tymbal apparatus—a ribbed membrane on the abdominal base used for sound production—and distinctive forewing venation patterns, such as the alignment of the radial and ulnar veins, which are hallmarks of this tribe's annual cicadas in contrast to periodical or other subtribes. These features, combined with phylogenetic evidence, confirm N. linnei's position among the Cryptotympanina subtribe, which includes other North American genera like Diceroprocta and Megatibicen.^[5]

Etymology and synonyms

The genus name Neotibicen derives from the Greek prefix "neo-" meaning "new," combined with Tibicen, a Latin term referring to a flute-player or piper, alluding to the characteristic loud calls of cicadas in this group.^[7] The species epithet linnei is the genitive form of "Linnaeus" (the Swedish form being Linné), honoring the 18th-century taxonomist Carl Linnaeus, as early specimens were misidentified as representing his species Cicada tibicen from 1758.^[1] Neotibicen linnei was first described as Cicada linnei in 1907 by John B. Smith and John A. Grossbeck, based on male specimens collected in Massachusetts.^[1] The description addressed the prior confusion with Linnaeus's Cicada tibicen, which actually refers to a different species now known as Neotibicen tibicen. This misattribution was further clarified in 1918 by William T. Davis, who provided a detailed key and additional observations on southeastern populations in his work on Mississippi cicadas.^[8] The primary synonym is Tibicen linnei (Smith & Grossbeck, 1907), reflecting the pre-2015 classification before the genus was revised to Neotibicen by Marshall et al..^[6]^[4] There are no major junior synonyms, though early literature often conflated N. linnei with Neotibicen pruinosus due to overlapping coloration and morphology, particularly in females where wing venation and pruinosity can appear similar.^[9]

Physical description

Morphology

Neotibicen linnei is a large-bodied cicada with a robust, heavy build characteristic of annual cicadas in the genus Neotibicen. Adults typically measure 28–34 mm in body length, with a wingspan reaching up to 70 mm. The body features a broad head and a three-segmented rostrum used for feeding on plant sap. Robust legs are adapted for clinging to tree bark, while the forewings exhibit a prominent bend near the middle, aiding in flight and perching. Males possess lobate opercula covering the tymbal organs, and the abdomen includes a black longitudinal band along the ventral surface.^[3]^[1]^[2] Sexual dimorphism is evident in reproductive structures. Males have larger, more prominent lobate opercula and tymbals on the abdomen for sound production, with the second abdominal segment being longer than in females. Females lack tymbals but feature a robust ovipositor for inserting eggs into twigs.^[3]^[2] Nymphs of N. linnei are wingless and adapted for a burrowing lifestyle underground. They possess front legs modified into fossorial structures with strong claws for digging through soil to access root xylem. The body is pale with dark markings, facilitating camouflage in the subterranean environment.^[10]^[11]

Coloration and variation

Adult Neotibicen linnei exhibit a coloration pattern dominated by black on the body, accented by green or greenish-yellow markings on the thorax and head, with translucent wings featuring dark veins.^[12] Newly emerged adults display a bright blue-green hue and soft texture, which darkens to a matte black-green within 1-2 hours as the exoskeleton hardens and wings expand.^[13] Coloration varies with age, as the vibrant greens fade in older specimens, resulting in brownish and black tones upon preservation or senescence. Some specimens exhibit pruinose (frosty white) patches on the abdomen, particularly larger basal spots in males.^[12]^[2]^[1] Nymphs of N. linnei are pale yellowish or cream-colored, facilitating camouflage in the subterranean environment.^[14] In comparison to the similar Neotibicen pruinosus, N. linnei features less extensive green markings and a sharper bend in the forewing near the middle, rather than the more gradual curve seen in N. pruinosus.^[3]

Distribution and habitat

Geographic range

Neotibicen linnei is native to the eastern United States and adjacent southern Canada, primarily inhabiting deciduous forest and grassland ecosystems east of the 100th meridian. Its range spans from southern New England (including Massachusetts) and the Great Lakes region (including Michigan and Illinois) southward through the Mid-Atlantic and Southeast to Texas and Tennessee, with confirmed occurrences in states including New York, Maryland, West Virginia, Georgia, Mississippi, Alabama, Florida (northern and central portions), Arkansas, Louisiana, and others. The species is absent west of the Mississippi River in most areas, though isolated records exist in states like Kansas and Nebraska; it does not extend into the extreme southwestern or far northern regions.^[15] The species was first described in 1907 by John B. Smith and John A. Grossbeck based on specimens collected from several localities in New York and New Jersey. Subsequent documentation has expanded knowledge of its distribution through museum records and field observations, with citizen science platforms like iNaturalist revealing its presence in urban environments, such as Washington, D.C., where it occurs in wooded parks and green spaces. These efforts have confirmed its adaptability to fragmented landscapes within its core range.^[1]^[16] The range includes established populations in southern Ontario, Canada, such as the Niagara Peninsula and Long Point. Ongoing monitoring through platforms like iNaturalist continues to support its consistent presence in eastern North America.^[17]

Preferred habitats

Neotibicen linnei primarily inhabits deciduous forests dominated by hardwood trees such as oaks (Quercus spp.), maples (Acer spp.), and sweetgums (Liquidambar styraciflua), where it is commonly observed in the eastern and midwestern United States.^[13]^[18] This species also thrives in urban woodlots, parks, and suburban areas featuring mature deciduous trees, demonstrating a notable tolerance for human-modified landscapes compared to more strictly woodland-dependent cicadas.^[13]^[18] This preference for deciduous vegetation supports the species' proliferation in both natural forest edges and planted urban green spaces with adequate tree cover.^[13]

Life cycle and behavior

Development and emergence

Neotibicen linnei is an annual cicada with a life cycle typically spanning 2 to 5 years, during which nymphs develop underground in an unsynchronized manner, leading to yearly emergences rather than mass outbreaks seen in periodical cicadas.^[19]^[20] The species feeds on root xylem as nymphs, contributing minimally to tree damage over this extended subterranean phase.^[21] Females oviposit 10 to 25 eggs per slit in the bark of twigs using their specialized ovipositor, creating linear incisions that can cause minor twig dieback known as flagging.^[22] Eggs hatch after 6 to 10 weeks, with the tiny nymphs dropping to the soil surface and burrowing downward to feed on root fluids.^[23] Nymphs undergo five instars over 2 to 5 years, burrowing to depths of 15 to 60 cm while tapping into tree root xylem for sustenance. Late in the season, typically from late June to September and peaking in July to August, mature fifth-instar nymphs emerge from the soil at night, climb vertical surfaces such as tree trunks, and molt into adults, leaving behind characteristic exoskeletons or "shells."^[3]^[24] Adult emergence often occurs about one hour after sunset, particularly on evenings following periods of male singing activity.^[13] Newly ecdysed adults remain vulnerable as they dry and harden for 1 to 2 hours before taking flight.^[25] The asynchrony of developmental cohorts ensures staggered annual emergences, maintaining population stability without synchronized booms. Adults live for about 4-6 weeks, during which they mate and feed on plant sap.^[20]^[26]

Mating and song

Adult Neotibicen linnei exhibit distinct mating behaviors centered on acoustic signaling. Males perch in trees or shrubs and produce calls to attract receptive females, often forming choruses in groups during late afternoon to dusk to amplify their signals and increase mating success.^[27]^[28] Females respond to these calls by flicking their wings, producing a subtle snapping sound that signals interest and guides males to their location.^[29]^[30] Once paired, copulation occurs, after which the male detaches and the female proceeds independently to oviposit eggs in tree slits.^[31] The species' song is a key element of courtship, characterized by a high-pitched, pulsating call with a peak frequency around 7 kHz. It begins as a soft, 5-second buzz that gradually increases in intensity, transitioning into a 10-second crescendo-decrescendo rattle resembling the sound of a saltshaker, before ending abruptly; the full call lasts approximately 15 seconds, often interspersed with softer buzzes between phrases.^[13]^[32] This song is generated by the rapid buckling and unbuckling of tymbals, specialized ribbed membranes on the male's abdomen. Males sing throughout the day during warm weather above 25°C, with variations in rhythm and intensity among individuals and across locations that facilitate species recognition and mate selection.^[13] In courtship, males actively defend small singing territories from rivals, perching prominently to broadcast their calls. Females evaluate potential mates based on the quality, volume, and species-specific patterns of the song, approaching only those whose signals meet their preferences for vigor and suitability.^[33]^[34]

Ecology

Interactions with other species

Neotibicen linnei serves as primary prey for the Eastern Cicada Killer wasp (Sphecius speciosus), a solitary predator that stings and paralyzes adult cicadas before transporting them to underground nests to provision its larvae.^[35] Adults and nymphs are also consumed by various birds, such as songbirds, and mammals including squirrels, which exploit the cicadas during emergence periods. Nymphs face predation underground from ground-dwelling arthropods like ants, ground beetles, and mole crickets.^[36] Parasitism affects N. linnei at multiple life stages, with sarcophagid flies such as Emblemasoma erro targeting adults by ovipositing eggs on the host, leading to larval development that consumes the cicada's tissues.^[37] In regions of sympatry, N. linnei co-occurs with congeners like N. tibicen, but direct competition is minimized through temporal asynchrony in emergence and spatial partitioning in habitat micro-niches, such as woodland edges versus open areas.^[38] Species recognition is reinforced by distinct calling songs, with N. linnei's wavering, salt-shaker-like trill differing from the scissor-grinder rasp of N. pruinosus.^[13] Defensive strategies in N. linnei include a disturbance stridulation, producing a sharp protest buzz when seized, potentially startling predators and allowing escape; some species in the genus also flash hindwing patterns as a deimatic display.^[39]

Role in ecosystem

Neotibicen linnei plays a significant role in nutrient cycling within its habitats. The nymphs, which spend several years underground feeding on root xylem sap, create burrows that aerate the soil, improving water infiltration and oxygen availability to plant roots.^[40] Upon emergence, the adults die off after a short period, contributing a pulse of biomass that decomposes and enriches the soil with organic matter and nutrients, supporting microbial decomposers and overall soil health.^[40]^[41] In plant interactions, female N. linnei lay eggs by using their ovipositor to make slits in the bark of twigs on deciduous hardwoods, causing minor damage such as branch flagging, particularly on younger trees; however, established host trees typically recover, and the wounds can stimulate lateral branching and new growth over time.^[14]^[42] While adults feed on twig sap rather than floral nectar, this activity does not significantly harm mature trees and indirectly supports host plant vigor by not competing with other herbivores.^[14] As a biodiversity indicator, the abundance of N. linnei reflects the health of deciduous forests and urban green spaces, where its presence signals intact soil quality and minimal disturbance from factors like pesticide use or habitat fragmentation; population declines may indicate environmental degradation.^[43] This species serves as a key prey base for higher trophic levels, including birds, mammals, and predatory insects, thereby supporting food web dynamics without posing a major pest threat due to its limited plant damage.^[23]^[44] Additionally, the characteristic songs of emerging adults contribute to the acoustic biodiversity of summer woodlands, enhancing the sensory profile of healthy ecosystems.^[41]

References

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### Habitat and Host Tree Information for Neotibicen linnei
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