Genus Lycorma Stal, 1863

[back to North American Fulgoridae]

Family Fulgoridae Latreille, 1807

Subfamily Aphaeninae Blanchard, 1847

Tribe Aphaenini Blanchard, 1847

Genus Lycorma Stål, 1863: 234.

Type species (in original combination): Aphana imperialis White, 1846: 330.

Synonyms: None.

Recognized species

[Genus Lycorma in Metcalf catalog – Metcalf 1947: 163], see also 3i TaxonPages].  The taxonomy below adopts the view of Nagai & Porion (1996), updated by Lin et al. (2023), of Lycorma consisting of four species and no subspecies.

Lycorma delicatula (White, 1845) (Spotted lanternfly, SLF) – Bangladesh, China (Anhui, Beijing Fujian, Gansu, Guangdong, Hebei, Henan, Hubei, Jiangsu, Shaanxi, Shandong, Sichuan, Tianjin, Yunnan, Zhejiang); Japan (Honshu, Kyushu); Nansei-Shoto (Ryukyu Islands), South Korea; Vietnam; USA (currently 17 states – DE, CT, IL, IN, KY, MA, MD, MI, NC, NJ, NY, OH, PA, RI, TN, VA, WV; individual records (not populations) in Vermont, New Hampshire, Wisconsin, Florida,  Utah and California); Also Ontario and Nova Scotia in Canada.
= Aphaena delicatula White, 1845: 37.
= Lycorma delicatula (White, 1845), comb. by Stål 1863: 234.
= Lycorma delicatulum (White, 1845) incorrect subsequent spelling of Lycorma delicatula (White, 1845) in Matsumura 1931: 1269.
= Lycorma jole Stål, 1863: 234 (subjective synonym of Aphaena delicatula White, 1845) in Nagai & Porion, 1996a: 21.
= Lycorma operosa (Walker, 1858: 46) (subjective synonym of Aphaena delicatula White, 1845) in Nagai & Porion, 1996a: 21.

Lycorma imperialis (White, 1846: 330) – Bangladesh, India (Assam, Darjiling, Sillim), China
= Aphana imperialis White, 1846: 330.
= Aphaena imperialis (White, 1846), comb. by Westwood 1848: 74.
= Lycorma imperialis (White, 1846), comb. by Stål 1863: 234.
= Aphaena placabilis Walker, 1858.
= Lycorma imperialis placabilis (Walker, 1858), comb. by Distant 1906: 206.
= Lycorma imperialis var. placabilis (Walker, 1858), comb. by Metcalf 1947: 166.
= Lycorma imperialis var. placabilis (Walker, 1858) synonym of Lycorma imperialis (White, 1846) by Nagai & Porion 1996: 21.

Lycorma meliae Kato, 1929 – Taiwan
= Lycorma olivacea Kato, 1929, synonym by Lin et al. (2023: 13).

Lycorma punicea (Hope, 1843) – Bangladesh (as “Silhet”), China, India (Assam)
= Lystra punicea Hope, 1843: 133.
= Aphana punicea (Hope, 1843), comb. by Schaum 1850: 66.
= Aphaena delectabilis Walker, 1858.
= Aphaena delectabilis Walker, 1858, a synonym of Lycorma punicea (Hope, 1843) by Stål 1862: 485.
= Lycorma punicea (Hope, 1843), comb. by Stål 1863: 234.
= Lycorma imperialis punicea (Hope, 1843), revised status (subspecies) by Lallemand 1963: 45. (error, punicea predates imperialus).

Distribution

Primarily Indomalayan (for genus), but also temperate China.  One species – Lycorma delicatula, the spotted lanternfly [SLF] – adventive to Korea, Japan and USA.

Spotted lanternfly in the US

StopSLF.org is now live, a hub for Spotted Lanternfly information.
SLF in the NYT.
Penn State Extension for SLF.
Homeowner control recommendations (Penn State University Extension) – fact sheet with the current (30 June 2020) recommendations for residents: https://extension.psu.edu/spotted-lanternfly-management-for-homeowners.
SLF Detection map: see Cornell. This map is ‘official’ so may omit reports that state agencies haven’t seen or confirmed.
An article to explain the many options: https://extension.psu.edu/what-to-do-about-spotted-lanternfly-on-ornamental-trees-and-plants-in-residential-and-public-landscapes

There are now many websites on the spotted lanternfly. now too many to list here.  See “stop SLF ” which is intended as a clearinghouse of information..

California – no populations, but there have been several accounts of dead adults (or not-so-dead) arriving on aircraft.  CA dept. of Ag is taking the threat seriously; Mark Hoddle of UCR Riverside is working on it.
Connecticut (populations in Fairfield & New Haven Counties); reports widespread.
Delaware  (all 3 counties).
Indiana. Switzerland County (report new findings in Indiana to the Indiana Department of Natural Resources. Call 866-NO EXOTIC (866-663-9684) or email DEPP@dnr.IN.gov).
Maryland –  Now reported in 11 counties, the entire northern border of the state (except westernmost counties of Allegany and Garret), and the adjacent counties. It soon will be easier to list the counties it is not found in.
Massachusetts. Now in Worchester County (see MASS DAR). (pdf in case it gets archived – 22 Feb 2019 – dead specimen, Boston) (see also PDF)
New Jersey – NJ Dept of Ag – Essentially statewide, Cape May County does not yet have a reported population.
New York  – Dept. of Ag and Markets, Dept. of Environmental Conservation (Southeast corner of state plus Tomkins and Broome).
North Carolina – They have shown up in the state (dead or mostly dead), but no confirmed populations (NCDA SpottedLanternflyPestWatch.pdf (ncagr.gov)).
Ohio (populations in Cuyahoga and Jefferson Counties), see Ohio Department of Agriculture.
Pennsylvania – All of southeastern Pennsylvania and west along PA turnpike to Ohio border)- updates here; (map updated here: https://nysipm.cornell.edu/environment/invasive-species-exotic-pests/spotted-lanternfly/).
Rhode Island –  Detected in Warwick, but not a population yet, maybe (If you have found one, report it here).
Virginia – 6 counties in the northern part of the state (the population was detected in Frederick County, Virginia, on Jan. 10, 2018).
West Virginia – 4 counties in the eastern part of the state

State-level information and reporting

General steps you may take:

Collect a specimen: Turn the adult specimen or egg mass into your state’s department Entomology Lab for verification. First, place the sample in alcohol or hand sanitizer in a leak-proof container.

Take a picture: Submit photographs to the local cooperative extension service.

Report a site: Call the local Bad Bug hotline with details of the siting and your contact information.

Economic Importance

Lycorma delicatula was found in Berks Co., Pennsylvania.  In September 2014, the Pennsylvania Department of Agriculture has confirmed the presence of Lycorma delicatula (White, 1845) near Reading, PA.  This species probably came from South Korea, and may have arrived 2 years before (hence 2013 or maybe 2012).

Lycorma delicatula arrived in South Korea in 2006 (and subsequently Japan) (see Han et al. 2008), where it has become broadly established.  The primary host of the species is the tree of heaven (Ailanthus altissima) but feeds/reproduces on a variety of plants (mostly trees).  Particularly troublesome is that grapes (Vitis sp) are a particularly preferred host, but a number of tree fruits (apple, cherry, peach) are on the host list as well. The species sequesters toxins (esp. from its preferred host) and generally, birds will not consume this species.

Unfortunately, this species has been found in large numbers in a Pennsylvania apple orchard – a worrisome observation whose implications are not yet clear. At least 2 vineyards have been lost due to spotted lanternfly in the invasion area.

Webpage from PA Department of Agriculture on Lycorma detection and control efforts.

There is some concern that bees may collect honeydew from Lycorma (Lycorma delicatula and honey bees – honeydew).

Parasites and Predators

Known to be parasitized by a eupelmid wasp Anastatus orientalis (which is an egg parasite, see Kim et al. 2011, Choi et al. 2014 cited below) and a dryinid wasp now under the name Dryinus sinicus Olmi, 1987.  Other Dryinus species are also reported as parasites/possible parasites of Lycorma or related taxa. There is some evident of recruitment of egg parasites – several taxa have been reared from SLF eggs, but mostly isolated individuals. Recently, the indigenous Anastatus species, A. reduvii (a parasite of brown marmorated stink bug eggs), has been reared from SLF eggs in numbers, but the parasitism rate remains low (>1%).

There are a variety of reports of insects and birds eating spotted lanternflies.  It does appear that wheel bugs (Arilus cristatus (L.), mantids, spiders and the predatory stink bug Apoecilus cynicus (Say) (Barringer and Smyers 2016) will take them.

Recently Clifton et al. (2019) reported a large die-off of spotted lanternfly caused by an coepizootic of two fungal pathogens – Batkoa major (Division Zoopagomycota, Order Entomophthorales) and Beauveria bassiana (Division Ascomycota, Order Hypocreales).

Ooencyrtus kuvanae (Howard) (Encyrtidae)has been reared from eggs of SLF in Pennsylvania (Liu & Mottern 2017).

Figure 1 from Clifton et al. 2019: Two fungal pathogens causing a coepizootic in an L. delicatula population. (A) L. delicatula during the epizootic: only one of these adults was alive and the remainder had been killed by B. major. A similar degree of mortality was observed on trees throughout the site. (B) Rhizoids from B. major (arrow) attaching a dead adult to a tree. Image courtesy of Kelly Murman (photographer). (C) Adult cadaver with wings and legs extended outward as B. major conidia are released from the abdomen. (D) Adult killed by B. bassiana. (E) Percent L. delicatula killed by either pathogen on the trees or surrounding ground.

Plant associations

Lycorma meliae Kato, 1929 – Melia azedarach L. (Sapindales, Meliaceae) according to Kato (1933)

The primary hosts for Lycorma delicatula appear to be Ailanthus and grape. The EPPO host list has been updated for SLF (PDF).

Working spotted lanternfly host list from Lawrence Barrenger of PDI (as of 1 Aug 2019)

SLF Host List 2019 (2).

Table from Dara et al. 2015

Host table from Park et al. (2009).

Recognition

There are no other species in our area similar to Lycorma.

Below, Lycorma delicatula, first instar, photographed in Berks Co., PA on May 12, 2015 by Greg Hoover, Dept. of Entomology, Penn State University

 

Lycorma delicatula, first instar, photographed in Berks Co., PA on May 12, 2015 by Greg Hoover, Dept. of Entomology, Penn State University

Online Resources

3i TaxonPages.
EPPO.
PDA.
Pest news.
CDFA.
EOL.
Massachusetts Massachusets introduced pest outreach project.
Entomology Today.
Bugwoodwiki.
Pesttracker.
Wikipedia.
University of Florida featured creature.
StopSLF.
iNaturalist.
Bugguide.
Video at Reading Eagle (newspaper). Article.
Adult photos http://hojae.net/520.
Nymph photos  4th instar. earlier instars.

Collecting

Easily found on the tree of heaven within the invasion area, also on grape.  When numbers are higher, they are on red and silver maple, among other species.

Molecular resources

There is now a great deal of molecular data for the spotted lanternfly.

A complete mitogenome (pub here) is now available for the spotted lanternfly (Genbank data).

Lycorma at GenBank. (as of 7 Jan. 2020, 634 hits)

Lycorma at Barcode of life. (COI barcode gene, as of 7 Jan 2020, 137 sequences)

Selected references

This reference list includes peer-reviewed literature, technical documents, theses, popular articles, and some other gray literature.

Adam, J. and B. Overton. 2023. Forgotten fungi that could be used to control the spread of the Spotted Lanternfly (Hemiptera: Fulgoridae). Fungi 15(5): 41-50.

Adams, J. 2022. State-Level Challenges to Invasive Insect Management: A Case Study of Spotted Lanternfly in the Northeast. Masters Theses. Oregon State University. Corvallis, Oregon.

Allen, M.C., A.L. Nielsen, D.L. Peterson, and J.L. Lockwood. 2021. Terrestrial eDNA survey outperforms conventional approach for detecting an invasive pest insect within an agricultural ecosystem. Environmental DNA (2021): (PrePrint, 11 pp.). https://doi.org/10.1002/edn3.231. [Lycorma delicatula]

American Nurseryman staff. A colorful new pest. American Nurseryman Jan 2015, pp. 6-7.

AN, Hyunjin, Sung Ju CHO, Saera OH, and Jae-Min JUNG. 2018. Economic impacts of invasive pests under climate change: A case of Lycorma delicatula. Journal of the Korea Academia-Industrial Cooperation Society 19(7): 415-422. https://doi.org/10.5762/KAIS.2018.19.7.415.

Anonymous. 2014. Pest Alert, Spotted Lanternfly (Lycorma delicatula). United States Department of Agriculture APHIS 81-35-024.  pdf.

Anonymous. 2022. Spotted lanternfly found in Michigan. Newsletter of the Michigan Entomological Society 66(3): 1.

Avanesyan, A. and W.O. Lamp. 2020. Use of molecular gut content analysis to decipher the range of food plants of the invasive spotted lanternfly, Lycorma delicatula. Insects 11: 215; 1-13. https://doi.org/10.3390/insects11040215 (ePub Artilce#215, 13 pp.)

Avanesyan, A., T. K. Maugel, and W.O. Lamp. 2019. External morphology and developmental changes of tarsal tips and mouthparts of the invasive spotted lanternfly, Lycorma delicatula. PLoS ONE 14(12) (article e0226995: 26 pp. https://doi.org/10.1371/journal.pone.0226995

Avanesyan, A., C. McPherson, and W. O. Lamp. 2022. Analysis of plant trait data of host plants of Lycorma delicatula in the US suggests evidence for ecological fitting. Forests 13(12): 2017; 1-16. https://doi.org/10.3390/f13122017.

Aviles-Rosa, E. O., M. Nita, E. Feuerbacher, and N. J. Hall. 2023. An evaluation of Spotted Lanternfly (Lycorma delicatula) detection dog training and performance. Applied Animal Behaviour Science 258: 105816; 1-11. https://doi.org/10.1016/j.applanim.2022.105816.

Aviles-Rosa, E.O., S.A. Kane, M. Nita, E. Feuerbacher, and N.J. Hall. 2023. Olfactory threshold of dogs (Canis lupus familiaris) to cold-killed spotted lantern fly eggs. Applied Animal Behaviour Science 261: 105880. https://doi.org/10.1016/j.applanim.2023.105880.

Baek, S.-H & Joung, O & Lee, H.-Y & Shin, J.-C & Choi, W.-S & Lee, T.H. & Hwang, J.-S & Nam, S.-H & Son, H.-U & Lee, S.-H. 2018. Anti-oxidative fraction of Lycorma delicatula alleviates inflammatory indicators. Natural Product Communications 13: 431-434. (researchgate)

Baker, T. C., E. C. Smyers, J. M. Urban, Z. Meng, K. J. Pagadala Damadaram, A. J. Myrick, M. F. Cooperband and M. J. Domingue. 2019. Progression of seasonal activities of adults of the spotted lanternfly, Lycorma delicatula, during the 2017 season of mass flight dispersal behavior in eastern Pennsylvania. Journal of Asia-Pacific Entomology 22(3), 705–713. https://doi.org/10.1016/j.aspen.2019.05.006

Baker, T.C., A.J. Myrick, M.S. Wolfin, and Y. Wang. 2021. Visual responses of flight-dispersing spotted lanternflies, Lycorma delicatula toward a tall vertical silhouette in a vineyard. Journal of Insect Behavior 34: 49-60. https://doi.org/10.1007/s10905-021-09766-0.

BAO, Ke-Xin, Ying-Qiao DANG, Xiao-Yi WANG, and Jian-Zhou CUI. 2022. Influencing factors for natural populations of Lycorma delicatula in different habitats. The Journal of Applied Ecology 33(1): 248-254. http://dx.doi.org/10.13287/j.1001-9332.202201.037. [Chinese]

BAO, Ke-Xin, Ying-Qiao DANG, Xiao-Yi WANG, and Wenyu CHEN. 2022. Diurnal rhythm of adult behavior of Anastatus orientalis under different photoperiod-temperature conditions. Chinese Journal of Biological Control (2022): (PrePrint, 9 pp.). https://doi.org/10.16409/j.cnki.2095-039x.2022.01.009. [Lycorma delicatula] [Chinese]

BAO, Ke-xin, Xiao-yi WANG, Liang-ming CAO, Bei XIN, Hannah J. BROADLEY, and Juli R. GOULD. 2023. Effects of transgenerational photoperiod experience on the reproduction and development of Anastatus orientalis, an egg parasitoid of the spotted lanternfly. Frontiers in Insect Science 3: 1153723. https://doi.org/10.3389/finsc.2023.1153723.

Barringer, L. E. 2014. Pest Alert. Spotted lanternfly Lycorma delicatula (Hemiptera: Fulgoridae). Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania. 2 pp. pdf.

Barringer, L. and C. M. Ciafré. 2020. Worldwide feeding host plants of spotted lanternfly, with significant additions from North America. Environmental Entomology 49(5): 999–1011. https://doi.org/10.1093/ee/nvaa093

Barringer, L. E. and C. R. Bartlett. 2018. Pennsylvania planthoppers (Hemiptera: Auchenorrhyncha: Fulgoroidea): relative abundance and incidental catch using novel trapping methods. Insecta Mundi 0661: 1–31. pdf

Barringer, L. E. and E. Smyers. 2016. Predation of the spotted lanternfly, Lycorma delicatula (White) (Hemiptera: Fulgoridae) by two native Hemiptera. Entomological News 126(1): 71–73.

Barringer, L. E., L. R. Donovall, S.-E. Spichiger, D. Lynch and D. Henry. 2015. The first New World record of Lycorma delicatula (Insecta: Hemiptera: Fulgoridae). Entomological News 125(1): 20-23.

Belouard, N. and J.E. Behm. 2023. Spotted! Computer-aided individual photo-identification allows for mark-recapture of invasive spotted lanternfly (Lycorma delicatula). Frontiers in Insect Science 3: 1112551. https://doi.org/10.3389/finsc.2023.1112551.

Belouard, N., and J.E. Behm. 2023. Multiple paternity in the invasive spotted lanternfly (Hemiptera: Fulgoridae). Environmental Entomology (2023): (PrePrint, 7 pp.). https://doi.org/10.1093/ee/nvad083.

Bielski, J. and J. Steffel. 2019. Field observations on spotted lanternfly behavior and host suitability. Pp. 31-36. In: M.W. Choi (ed.). Proceedings of the Cumberland-Shenandoah Fruit Workers Conference 95th Annual Meeting, December 5th-6th, 2019, Winchester, Virginia. pdf.

Bien, T., B. H. Alexander, E. White, S. T. Hsieh, and S. A. Kane. 2023. Sizing up spotted lanternfly nymphs for instar determination and growth allometry. Plos one 18(2): e0265707; 1-13. https://doi.org/10.1371/journal.pone.0265707.

Bien, T., B.H. Alexander, C. Li, N. Goeler-Slough, S.T. Hsieh, and S. Amador Kane. 2023. How spotted lanternflies get back on their feet: 3D mechanical modeling explains terrestrial self-righting strategies. bioRxiv: 2023-02. https://doi.org/10.1101/2023.02.06.527347. [not listed online]

Blake, C. 2014. New invasive pest could threaten stone fruit, grapes. Western Farm Press (12/6/2014) 36(23): 1-8 (2 pp).  http://www.westernfarmpress.com/grapes/spotted-lanternfly-new-threat-grapes-stone-fruit (txt)

Blanchard, E. 1847. Descriptions of various genera of Homoptera with fixation. In: D’Orbigny A. C. V. D. 1847. Dictionnaire universel d’histoire naturelle, 10. p. 612 [642]

Broadley, H.J., J.R. Gould, L.T. Sullivan, X.Y. Wang, K.A. Hoelmer, M.L. Hickin, and J.S. Elkinton. 2020. Life history and rearing of Anastatus orientalis (Hymenoptera: Eupelmidae), an egg parasitoid of the spotted lanternfly (Hemiptera: Fulgoridae). Environmental Entomology 50(1): 28–35. https://doi.org/10.1093/ee/nvaa124.

Broadley, H. J., S. J. Sipolski, D. B. Pitt, K. A. Hoelmer, X. Y. Wang, L. M. Cao, L. A. Tewksbury, T. J. Hagerty, C. R. Bartlett, A. D. Russell, Y. Wu, S. C. Davis, J. M. Kaser, J. S. Elkinton, and J. R. Gould. 2023. Assessing the host range of Anastatus orientalis, an egg parasitoid of spotted lanternfly (Lycorma delicatula) using Eastern US non-target species. Frontiers in Insect Science 3: 21; 1-14. https://doi.org/10.3389/finsc.2023.1154697.

Brooks, R. K., A. Toland, A. C. Dechaine, T. McAvoy and S. Salom. 2020. The inability of spotted lanternfly (Lycorma delicatula) to vector a plant pathogen between its preferred host, Ailanthus altissima, in a laboratory setting. Insects 11 (ePub article 515): 10 pp. https://doi.org/10.3390/insects11080515

Burrows, M., A. Ghosh, G. P. Sutton, H. M. Yeshwanth, S. M. Rogers, and S. P. Sane. 2021. Jumping in lantern bugs (Hemiptera, Fulgoridae). Journal of Experimental Biology 224: jeb243361; 1-14. https://doi.org/10.1242/jeb.243361.

Byeon, Dae-hyeon, Jae-Min Jung, Sunghoon Jung and Wang-Hee Lee. 2020. Effect of types of meteorological data on species distribution predicted by the CLIMEX model using an example of Lycorma delicatula (Hemiptera: Fulgoridae). Journal of Asia-Pacific Biodiversity 13(1): 1-6. https://doi.org/10.1016/j.japb.2019.11.010

Cai, Hui-Xian and Xiao-Zhong Wu. 2013. Occurrence and control of Lycorma delicatula in Ailanthus altissima in Jiaozuo, China [Abstract]. In: The Proceedings of Chinese Society of Plant Protection in 2013, Shandong, China, 22-25 October, 2013. Chinese Academy of Agricultural Sciences, China. pp. 146-149.

Calvin, D. D., J. Keller, J. Rost, B. Walsh, D. Biddinger, K. Hoover, B. Treichler, A. Johnson, and R. T. Roush. 2021. Spotted Lanternfly (Hemiptera: Fulgoridae) nymphal dispersion patterns and their influence on field experiments. Environmental Entomology 50(6): 1490-1504. https://doi.org/10.1093/ee/nvab104.

Calvin, D. D., J. Rost, J. Keller, S. Crawford, B. Walsh, M. Bosold, and J. Urban. 2023. Seasonal activity of spotted lanternfly (Hemiptera: Fulgoridae), in Southeast Pennsylvania. Environmental Entomology (2023): (PrePrint, 18 pp.). https://doi.org/10.1093/ee/nvad093.

Cantu, A. 2023. Evaluating volatile organic compounds from the spotted lanternfly (Lycorma delicatula) using headspace odor sampling methods. Doctoral dissertation. Texas Tech University, Lubbock, Texas.

Chase, K.D., and S.E. Wright. 2023. Preemergent control of Spotted Lanternfly, 2020. Arthropod Management Tests 48(1): tsad001; 1-2. https://doi.org/10.1093/amt/tsad001.

CHEN, X. X. and J. H. HE. 2006. Lycorma delicatula Ashmead. In: X.X. Chen and J.H. He. 2006. Parasitoids and Predators of Forest Pests in China. China Forestry Publishing House. Beijing, China. P. 128.

Cho, S. R., J. E. Lee, J. W. Jeong, J. O. Yang, C. M. Yoon and G. H. Kim. 2011. Comparison of cuticular hydrocarbons of different developmental stages of the spot clothing wax cicada, Lycorma delicatula (Hemiptera: Fulgoridae). Korean Journal of Applied Entomology 50(3): 185-194.

Choi, Man-Youn, Zhong-Qi Yang, Xiao-Yi Wang, Yan Long Tang, Zhen Rong Hou, Jeong Hwan Kim and Young Woong Byeon. 2014. Parasitism rate of egg parasitoid Anastatus orientalis (Hymenoptera: Eupelmidae) on Lycorma delicatula (Hemiptera: Fulgoridae) in China.  Korean Journal of Applied Entomology 53(2) 135-139. (link should open to PDF)

Choi, D. H., K .H. Kim and Y. K. Jang. 2011. Agonistic interactions between nymphs of Lycorma delicatula (Hemiptera: Fulgoridae). Journal of Asia-Pacific Entomology 14(1): 21-25.

Choi, D. S., D. I. Kim, S. J. Ko, B. R. Kang, J. D., Park, S. G. Kim and K. J. Choi. 2012. Environmentally-friendly control methods and forecasting the hatching time Lycorma delicatula (Hemiptera: Fulgoridae) in Jeonnam Province. Korean Journal of Applied Entomology 51(4): 371-376. (link – the DOI doesn’t work)

CHOI, ManYoung, YANG ZhongQi, WANG XiaoYi, TANG YanLong, HOU ZhenRong, KIM JeongHwan, and BYEON YoungWoong. 2014. Parasitism rate of egg parasitoid Anastatus orientalis (Hymenoptera: Eupelmidae) on Lycorma delicatula (Hemiptera: Fulgoridae) in China.  Korean Journal of Applied Entomology 53(2) 135–139.  (here)

Chou. I. A. 1946. A study on Lycorma delicatula White. Entomologia et Ars 1(2/3/4): 31-54.

Chu, H. J. 1931. Notes of the Life-History of Lycorma delicatula White in Nanking. Peking Natural History Bulletin 5(2): 33-35.

Clifton, E.H. and A.E. Hajek. 2022. Efficacy of Beauveria bassiana and Cordyceps javanica mycoinsecticides against spotted lanternflies, Lycorma delicatula, in laboratory bioassays. Biocontrol Science and Technology 32(7): 824-836. https://doi.org/10.1080/09583157.2022.2052804.

Clifton, E. H., L. A. Castrillo, A. Gryganskyi and A. E. Hajek. 2019. A pair of native fungal pathogens drives decline of a new invasive herbivore. Proceedings of the National Academy of Sciences (PNAS) article 201903579 (ePub): 3 pp. https://doi.org/10.1073/pnas.1903579116

Clifton, E.H., A.E. Hajek, N.E. Jenkins, R.T. Roush, J.P. Rost and D.J. Biddinger. 2020. Applications of Beauveria bassiana (Hypocreales: Cordycipitaceae) to Control Populations of Spotted Lanternfly (Hemiptera: Fulgoridae), in Semi-Natural Landscapes and on Grapevines, Environmental Entomology 49(4, ePub article nvaa064): 854–864. https://doi-org.udel.idm.oclc.org/10.1093/ee/nvaa064

Clifton, E. H., L. A. Castrillo, and A. E. Hajek. 2021. Discovery of two hypocrealean fungi infecting spotted lanternflies, Lycorma delicatula: Metarhizium pemphigi and a novel species, Ophiocordyceps delicatula. Journal of invertebrate pathology 186: 107689. https://doi.org/10.1016/j.jip.2021.107689

Clifton, E.H., L.A. Castrillo, S.T. Jaronski, and A.E. Hajek. 2023. Cryptic diversity and virulence of Beauveria bassiana recovered from Lycorma delicatula (spotted lanternfly) in eastern Pennsylvania. Frontiers in Insect Science 3: 1127682. https://doi.org/10.3389/finsc.2023.1127682.

Contreras-Orendain, L. 2021. Limitations of Genomic Analysis on Novel Species. Haverford  College, Haverford PA. Senior thesis. http://hdl.handle.net/10066/23531

Cook, R.T. 2021. Exotic invasive species dynamics and impacts in forest ecosystems. Purdue University, West Lafayette, IN. Directorate dissertation. https://doi.org/10.25394/PGS.14501778.v1 (abstract).

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DING, Zhanlai, Jing YANG, Peiliu LI, Yan XING, Shousheng TANG, and Lei WANG. 2020. An interlayer of multiple microscale hollow channels enhances the durability of surface topographies. ChemNanoMat 2020: 6; 1-6 (pre-print ePub, article #6). https://doi.org/10.1002/cnma.201900726 [Lycorma delicatula]

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Hajek, A.E., E.H. Clifton, S.E. Stefanik, and D.C. Harris. 2022. Batkoa major infecting the invasive planthopper Lycorma delicatula. Journal of Invertebrate Pathology (2022): (PrePrint, 27 pp.). http://dx.doi.org/10.2139/ssrn.4132010.

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JUNG, Jong-Kook, Mannyeon KIM, Cha Young LEE, Beom-Jun JANG, Dongsoo KIM, Hae Yeon KWON, and Yunmi PARK. 2021. Comparison of insect pest communities on 30 cultivars of Hibiscus syriacus. Journal of Korean Society of Forest Science 110(1): 116-127. https://doi.org/10.14578/jkfs.2021.110.1.116. [Korean, with English summary] [Lycorma delicatula, Metcalfa pruinosa]

Jung, Jae-Min, S. Jung, D.-h. Byeon and W.-H. Lee. 2017 Model-based prediction of potential distribution of the invasive insect pest, spotted lanternfly Lycorma delicatula (Hemiptera: Fulgoridae), by using CLIMEX. Journal of Asia-Pacific Biodiversity 10(4): 532-538. https://doi.org/10.1016/j.japb.2017.07.001.  [PDF]

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JUNG, Minhyung, Jung-Wook KHO, Do-Hun GOOK, Young Su LEE, and Doo-Hyung LEE. 2022. Dispersal and oviposition patterns of Lycorma delicatula (Hemiptera: Fulgoridae) during oviposition period in Ailanthus altissima (Simaroubaceae). Research Square preprint 17 pp. https://doi.org/10.21203/rs.3.rs-1201537/v1

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Kane, S.A., T. Bien, L. Contreras-Orendain, M.F. Ochs, and S.T. Hsieh. 2021. Many ways to stick the landing: Novel righting strategies allow spotted lanternfly nymphs to land on diverse substrates. bioRxiv (2021): (PrePrint, 27 pp.). https://doi.org/10.1101/2021.04.12.439561. {not peered reviewed}

Kane, S.A., Theodore B., L. Contreras-Orendain, M. Ochs, and T. Hsieh. 2021. Spotted lanternfly nymphs use multiple self-righting behaviors during landing. Bulletin of the American Physical Society (2021) [Abstract]. https://meetings.aps.org/Meeting/MAR21/Session/P14.1.

Kane, S.A., E.O. Aviles-Rosa, and N.J. Hall. 2023. Development and assessment of Spotted Lanternfly (Lycorma delicatula) training aids for detection canines. Applied Animal Behaviour Science 266: p106011. https://doi.org/10.1016/j.applanim.2023.106011.

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Keena, M.A., and A.L. Nielsen. 2021. Comparison of the hatch of newly laid Lycorma delicatula (Hemiptera: Fulgoridae) eggs from the United States after exposure to different temperatures and durations of low temperature. Environmental Entomology (2021): (PrePrint, 8 pp.). https://doi.org/10.1093/ee/nvaa177.

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Park, M., K. S. Kim and J. H. Lee. 2012. Isolation and characterization of eight microsatellite loci from Lycorma delicatula (White) (Hemiptera: Fulgoridae) for population genetic analysis in Korea. Molecular Biology Reports 39(5): 5637-5641.

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Ramirez, V. A., S. De Bona, M. R. Helmus, and J. E. Behm. 2022. Multiscale assessment of oviposition habitat associations and implications for management in the spotted lanternfly (Lycorma delicatula), an emerging invasive pest. bioRxiv. https://doi.org/10.1101/2022.09.08.507131.

Ramirez, V.A., S. De Bona, M.R. Helmus, and J.E. Behm. 2022. Multiscale assessment of oviposition habitat associations and implications for management in the spotted lanternfly (Lycorma delicatula), an emerging invasive pest. Journal of Applied Ecology (2022): (PrePrint, 10 pp.). https://doi.org/10.1111/1365-2664.14365.

REKHA, Deepika GOSWAMI, Deeksha ARYA and B.R. KAUSHAL. 2022. Insects diversity in an agroecosystem of Bhabar region of Uttrakhand. Indian Journal of Entomology e21216; 1-9. https://doi.org/10.55446/IJE.2021.382. [Lycorma delicatula]

Riggins, D.J. [no date] Spotted lanternfly. Mississippi Forestry Commission Technical Bulletin Forest Health Notes #76. 6 pp. pdf

Rowe, L.M., P.J. Higman, and H.D. Enander. 2020. Screening the Michigan forest inventory and Midwest invasive species network databases to locate host plants of Lycorma delicatula (Spotted Lanternfly) in Michigan. Michigan Natural Features Inventory, Report No. 2020-11, Lansing, Mi. 19 pp.

Rohde, B.B., M.F. Cooperband, I. Canlas, and R.W. Mankin. 2022. Evidence of receptivity to vibroacoustic stimuli in the Spotted Lanternfly Lycorma delicatula (Hemiptera: Fulgoridae). Journal of Economic Entomology 115(6): 2116-2120. https://doi.org/10.1093/jee/toac167.

Schaum, H. R. 1850. Fulgorellae. Erster Section A-G. In: I. S. Ersch  and I. G. Gruber. Allgemeine Encyklopädie der Wissenschaften und Kunste in alnhaberischen folge von Genannten Schriftstellern bearbeitet und herausgegeben, 51. p. 58-73.

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Shim, Jae-Kyoung and Kyeong-Yeoll Lee. 2015. Molecular characterization of heat shock protein 70 cognate cDNA and its upregulation after diapause termination in Lycorma delicatula eggs. Journal of Asia-Pacific Entomology. https://doi.org/10.1016/j.aspen.2015.08.005  [pdf of preprint]

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Yang, Zhong-qi, Man-Young Choi, Hwi-Jong Lee and Young-Woong Byeon. 2015. Asynchronous Emergence Mechanisms of Anatatus orientalis (Hymenoptera: Eupelmidae), an Egg Parasitoid of Lantern-fly Lycorma delicatula (Hemiptera: Fulgoridae). Presentation P054 at the Biannual meeting of the Korean Society of Applied Entomology (www.entomology.or.kr). April 23-25, 2015. [abstract available]

Yoon, C. M., S. R. Moon, J. W. Jeong, Y. H. Shin, S. R. Cho, K. S. Ahn, J. O. Yang and G. H. Kim. 2011. Repellency of lavender oil and linalool against spot clothing wax cicada, Lycorma delicatula (Hemiptera: Fulgoridae) and their electrophysiological responses. Journal of Asia-Pacific Entomology 14(4): 411-416.

Yun, Ji-Hyun, Seong Woon Roh, Tae Woong Whon, Mi-Ja Jung, Min-Soo Kim, Doo-Sang Park, Changmann Yoon, Young-Do Nam, Yun-Ji Kim, Jung-Hye Choi, Joon-Yong Kim, Na-Ri Shin, Sung-Hee Kim, Won-Jae Lee and Jin-Woo Bae. 2014. Insect Gut Bacterial Diversity Determined by Environmental Habitat, Diet, Developmental Stage, and Phylogeny of Host. Applied and Environmental Microbiology  80 (17): 5254–5264.  DOI: 10.1128/AEM.01226-14 [Lycorma noted]

Zhang, Jian-Hui, Jian-Wei Chen and Xiang Li. 2009. Research progress on medicinal resources of Mylabris and close origin species. China Journal of Chinese Materia Medica 34 (6) 647-650. [In Chines see pubmed for abstract] [abstract Research progress on the medicinal resources of Mylabris spp. and related species is reviewed. Besides the 45 species in 7 genera within the Meloidae which produce cantharidin, 9 other species in 7 genera of other insect families also contain cantharidin: Zanna, Fulgora and Lycorma within the Fulgoridae (Homoptera), and Oxocopis, Heliocis, Xanthochroa and Oedemera within the Oedemeridae (Coleoptera). The biosynthesis and function of cantharidin are discussed, and attention is given to new methods of utilizing Mylabris resources.]

 ZHANG, Li, Wenhui ZHAO Fuping WANG, and Daozheng QIN. 2019. Genetic Diversity and Population Structure of Natural Lycorma delicatula (White) (Hemiptera: Fulgoridea) Populations in China as Revealed by Microsatellite and Mitochondrial Markers. Insects 10(10): 312; 1–13. https://doi.org/10.3390/insects10100312.

ZHANG, Yanlong, Kexin BAO, Bei XIN, Liangming CAO, Ke WEI, Yingqiao DANG, Zhongqi YANG, Zhaozhi LV, and Xiaoyi WANG. 2023. The biology and management of the invasive pest spotted lanternfly, Lycorma delicatula White (Hemiptera: Fulgoridae). Journal of Plant Diseases and Protection (2023): 1-20. https://doi.org/10.1007/s41348-023-00794-w.

Zheng J., J. Du, Q. Liu, S. Wang, Y. Li and J. Wu. 2009. Investigation on insect pest and plant diseases damaging apple trees and their dynamics in Tianshui, Gansu. Acta Agriculturae Boreali-Sinica 18(2): 293-298. [record from China North-Central (Gansu) ]

Zuo, J. H., S. M. Ding, X. B. Qin, Z. R. Wang and L. L. Bai. 2008. [Dryinus browni Ashmead biological characteristics observational study]. Journal of Shandong Forestry Science and Technology 2724(5): 16-18 (pdf in Chinese).

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