Lycorma
Lycorma Stål, 1863, is a genus of planthoppers in the family Fulgoridae (subfamily Aphaeninae, tribe Aphaenini), comprising four valid extant species native to Asia.[1][2] These insects are characterized by their distinctive appearance, including elongated bodies and, in some species, colorful wings with spotted patterns.[3] The type species is Lycorma imperialis (White, 1846), originally described as Aphana imperialis.[1] The four recognized species are Lycorma delicatula (White, 1845), Lycorma imperialis (White, 1846), Lycorma meliae Kato, 1929, and Lycorma punicea (Hope, 1843).[2] Members of the genus are primarily distributed in the Indomalayan region and temperate parts of China, with L. delicatula also adventive to South Korea, Japan, and the United States (where it has established populations in at least 18 states as of 2025).[2][4][5] As phloem-feeding herbivores, Lycorma species can impact host plants by extracting sap, producing honeydew that promotes sooty mold, and in the case of L. delicatula, causing significant economic damage to vineyards, orchards, and forestry through direct feeding and indirect effects like plant stress and secondary infections.[6][3] L. delicatula, commonly known as the spotted lanternfly, is the most studied and notorious member of the genus due to its invasive spread and polyphagous feeding on over 70 plant species, including preferred hosts like Ailanthus altissima (tree of heaven) and Vitis spp. (grapes). As of late 2025, its U.S. populations continue to expand, with detections in additional regions.[4][7][8] Management efforts for invasive populations focus on biological control agents, such as the egg parasitoid Anastatus orientalis, and chemical insecticides, alongside regulatory quarantines to limit further spread.[9][2]Taxonomy
Classification
The genus Lycorma belongs to the kingdom Animalia, phylum Arthropoda, class Insecta, order Hemiptera, suborder Auchenorrhyncha, superfamily Fulgoroidea, family Fulgoridae, subfamily Aphaeninae.[2][4] Within the family Fulgoridae, commonly known as lanternflies, Lycorma is classified among large planthoppers characterized by piercing-sucking mouthparts adapted for feeding on plant sap and enlarged hind legs modified for jumping.[10][11] The genus Lycorma Stål, 1863, has no major synonyms recorded in taxonomic databases.[1] Its type species is Lycorma imperialis (White, 1846), originally described as Aphana imperialis.[1][2] Lycorma delicatula, the spotted lanternfly, represents the most extensively studied species within the genus due to its invasive status.[4]Nomenclature and history
The genus Lycorma was established by the Swedish entomologist Carl Stål in 1863 in his publication on Fulgoridae in the Stettiner Entomologische Zeitung.[1] The name derives from the ancient Greek Lycormas River (modern Evinos River in Aetolia), which in mythology was associated with the river god Evenus.[1] The type species, Lycorma imperialis, was originally described as Aphaena imperialis by Adam White in 1846 based on specimens from China, and its status as type species was formalized by William Lucas Distant in 1906.[1] Prior to the genus's creation, the first species now placed in Lycorma was described by Frederick William Hope in 1843 as Lystra punicea from specimens collected in Sylhet (present-day Bangladesh). Between 1845 and 1863, additional species were described and reclassified by entomologists including White, John Obadiah Westwood, and Francis Walker, often under genera such as Aphaena or Lystra, before Stål consolidated them into Lycorma.[1] A notable early nomenclatural debate involved Lycorma delicatula, originally described as Aphaena delicatula by White in 1845 from Chinese specimens; Stål simultaneously proposed the junior synonym Lycorma jole in 1863, later treated as a subspecies variant (L. delicatula jole) by Louis Maurice Adolphe Lallemand in 1963.[12] This synonymy was confirmed in 1996 by Shigeyuki Nagai and Thierry Porion, who recognized Lycorma as comprising four valid species without subspecies.[2] Key revisions continued into the 21st century, with You-Sheng Lin and colleagues in 2023 providing an updated taxonomic treatment of Fulgoridae in Taiwan, synonymizing Lycorma olivacea (Kato, 1929) under L. meliae and reinforcing the genus's four-species composition based on morphological and distributional evidence.[13] The invasive status of L. delicatula has driven recent molecular and phylogenetic studies, enhancing understanding of Lycorma's taxonomy.[14]Description
Adult morphology
Adult Lycorma individuals, particularly L. delicatula (the most studied species), exhibit a robust body structure typical of fulgorid planthoppers, with adults measuring 20–27 mm in length from head to the folded wing tips (other species may vary; e.g., L. imperialis is 16–21 mm), females generally larger at 24–27 mm compared to males at 20–23 mm.[7][15][16] The body is elongated and somewhat moth-like in appearance, with forewings overlapping and covering the hindwings at rest, providing a compact profile. Hind legs are enlarged and black, adapted for jumping with strong femora, enabling rapid escape responses.[17][18] The head is black to dark gray, bearing large compound eyes for wide visual fields and three ocelli arranged in a triangle for additional light detection, as characteristic of Auchenorrhyncha.[19] Antennae are short and filiform, often reddish and positioned below the eyes. Mouthparts form a long, segmented proboscis suited for sap-feeding: a tubular four- or five-segmented labium sheathes interlocked stylets, with the labium reaching up to 13 mm in length in adults; the stylet bundle includes maxillary stylets for puncturing plant tissues and mandibular stylets with sensory structures for host location.[19][20] Wings display dimorphism adapted for camouflage and defense: forewings are mottled gray-brown with black spots and reticulated tips, blending with tree bark for crypsis; hindwings, concealed beneath, are brightly colored with a proximal crimson or red section marked by black spots, transitioning to a black distal third separated by a bluish-white band, serving as aposematic warning signals when displayed.[7][15] Wingspan extends to approximately 50 mm when spread.[17] The abdomen is segmented, with seven visible sternites in females and six in males, featuring yellow ventral surfaces striped with black or brown bands. Sexual dimorphism is evident in abdominal width and genitalia: females possess broader abdomens and red valvifers at the distal margin for oviposition, while males have narrower forms; antenna length may vary slightly, but overall wing span aligns with body size differences.[15] Coloration patterns across the genus emphasize cryptic forewings for daytime concealment and vivid hindwings for predator deterrence, though species like L. imperialis exhibit brighter red and yellow hues in hindwings compared to the more subdued tones in L. delicatula.[7]Immature stages
The immature stages of Lycorma species, particularly Lycorma delicatula, encompass the egg and four nymphal instars, each exhibiting distinct morphological traits adapted for survival and development. Eggs are laid in masses containing 30 to 50 individuals, arranged in vertical rows and covered by a grayish-brown waxy secretion that provides camouflage and protection against desiccation.[21] These egg masses, measuring approximately 2.5 cm in length, are typically attached to the bark of host trees or other substrates, with the wax coating cracking over time to reveal the brownish, seed-like eggs beneath.[22] Upon hatching, first-instar nymphs measure about 3 mm in length and are wingless, featuring a predominantly black body adorned with prominent white spots on the dorsal surface and legs.[23] These spots serve as a key identifying feature, intensifying in visibility across the early instars as the nymphs grow to roughly double in size with each molt, reaching up to 8 mm by the third instar.[24] The first three instars maintain this black-and-white coloration, with no wing development, and exhibit gregarious tendencies that facilitate collective feeding and defense.[25] In the fourth and final nymphal instar, individuals grow to 12-15 mm, developing external wing pads as precursors to adult wings, while shifting to a reddish body marked with black and white spots for enhanced camouflage among foliage.[23] This instar also shows increased mobility compared to earlier stages, with elongated legs supporting stronger jumps, though the overall body remains wingless until the subsequent molt to adulthood.[24]Species
Recognized species
The genus Lycorma Stål, 1863 currently includes four recognized species, as established by Nagai and Porion (1996) and updated by Lin et al. (2023).[2][13] All species are native to Asia, though L. delicatula has become widely invasive outside its range.[2]- Lycorma delicatula (White, 1845), commonly known as the spotted lanternfly, was originally described as Aphaena delicatula. Adults measure about 25 mm in length, with grayish forewings marked by black spots and hindwings featuring red basal areas bordered by black and white bands.[17][2]
- Lycorma imperialis (White, 1846), the type species of the genus (originally Aphana imperialis), is similar in size to L. delicatula (around 25 mm long) but distinguished by darker forewing coloration.[2][26]
- Lycorma punicea (Hope, 1843), originally described from specimens in India, reaches adult lengths of approximately 20–25 mm and exhibits reddish hues on the wings and body.[2]
- Lycorma meliae (Kato, 1929), endemic to Taiwan and including the junior synonym L. olivacea Kato, 1929, is the largest in the genus, with adults averaging 25.7 mm in males (22.2–28.5 mm) and 28.9 mm in females (26.5–32.2 mm) in body length; it displays variable color morphs, including brown to dark brown or blue forms with spotted patterns.[13][2]