Abstract
The taxonomy of the deltocephaline leafhopper tribe Faltalini (13 genera, 63 species) is revised. A key to all genera and keys to species for each genus are provided. All genera are described and at least one species of each genus is illustrated including images of the habitus, male and female genitalia, and scanning electron microscope (SEM) images of the head, face, and other characters. The new genera Dietrichana n.gen. [type species D. pampas n. sp.] and Paraclorindaia n.gen. [type species P. pinguis n. sp.] are described and the following 36 new species are described: Ackbaria jujuyensis n. sp. [Argentina], A. ojosverdes n. sp. [Argentina], Aequcephalus laplata n. sp. [Chile], Bonamus jussaral n. sp. [Brazil], Clorindaia obrienorum n. sp. [Uruguay], C. otamendi n. sp. [Argentina], Dietrichana pampas n. sp. [Argentina], Faltala catalanoae n. sp. [Argentina, Uruguay], F. paradellae n. sp. [Argentina], F. viscacha n. sp. [Argentina, Uruguay], Hecullus mexicanus n. sp. [Mexico], Kramerana saltensis n. sp. [Argentina], Paraclorindaia pinguis n. sp. [Argentina, Uruguay], P. vesperis n. sp. [Argentina], Tenucephalus admirabilis n. sp. [Argentina], T. bartletti n. sp. [Peru], T. brevilamina n. sp. [Argentina], T. cerrado n. sp. [Brazil], T. ecuadorensis n. sp. [Ecuador], T. flagellifer n. sp. [Peru], T. heppneri n. sp. [Peru], T. iguazu n. sp. [Argentina], T. interstinctus n. sp. [Peru], T. johnsoni n. sp. [Brazil], T. lingulatus n. sp. [Bolivia, Peru], T. manuensis n. sp. [Peru], T. misiones n. sp. [Argentina], T. musettiae n. sp. [Brazil], T. nielsoni n. sp. [Brazil], T. novafriburgo n. sp. [Brazil], T. parana n. sp. [Argentina], T. spectabilis n. sp. [Peru], T. takiyae n. sp. [Brazil, Peru], T. umbrinus n. sp. [Brazil], T. viperinus n. sp. [Argentina], and Virganana danasensis n. sp. [Ecuador]. The new combinations Paraclorindaia brasileira (Zahniser) n.comb. [transferred from Clorindaia] and Tenucephalus amabilis (Linnavuori & Heller) n.comb. [transferred from Bonamus] are recognized. A phylogenetic analysis based on 51 morphological characters scored for 42 taxa including two outgroup taxa was performed. The resulting phylogeny resolved Faltalini as monophyletic, but the tribe remains difficult to diagnose because it encompasses a large amount of morphological diversity in characters traditionally used to define tribes of Deltocephalinae including head morphology, wing length and venation, and male genitalia. Despite this diversity, members of the tribe share the following characters: 1) phragma of male genital capsule partly developed into sclerotized plates or setose bulbous lobes (exception: Acrolithus); 2) first valvula shape relatively straight; 3) first valvula dorsal sculpturing separated from dorsal margin by unsculptured band; 4) first valvula dorsal sculpturing pattern maculose, with maculae separate or only slightly overlapping, or granulose; 5) first valvula with distinctly delimited ventral sculptured area; and 6) second valvula without dorsal teeth. Two major clades within the tribe were resolved and strongly supported, consisting of (Bonamus, Tenucephalus) and (Hecalocorica, (Hecullus, (Acrolithus, (Ackbaria, (Aequcephalus, Dietrichana, Virganana, Clorindaia, Faltala (Kramerana, Paraclorindaia)))))). A stepwise evolution of brachyptery in the latter clade is suggested by the phylogeny, beginning with evolution of subbrachypterous females in Hecullus and Acrolithus and culminating in completely brachypterous males and females in the monophyletic “Faltala group” of eight genera. It is hypothesized that a transition from an ancestral forested habitat to an open or grassland habitat facilitated the evolution of brachyptery. The hindwings and associated thoracic structures of eight species of Faltalini and three other Deltocephalinae were imaged with SEM. Brachyptery in Cicadellidae and other insects are briefly reviewed and discussed considering the hypothesis proposed here.
References
Ball, E.D. (1901) New Jassidae from the Rocky Mountain and Pacific region. Canadian Entomologist, 33, 4–11.
https://doi.org/10.4039/Ent334-1
Beamer, R.H. (1937) Five new species of leafhoppers (Homoptera – Cicadellidae). Journal of the Kansas Entomological Society, 10 (1), 10–13.
Blocker, H.D. & Fang, Q. (1992) A synopsis of the Faltala group (Homoptera: Cicadellidae) with descriptions of new taxa. Journal of the Kansas Entomological Society, 65, 341–346.
Bourgoin, T., Rong-Rong, W., Asche, M., Hoch, H., Soulier-Perkins, A., Stroiński, Yap, S. & Szwedo, J. (2015) From micropterism to hyperpterism: recognition strategy and standardized homology-driven terminology of the forewing venation patterns in planthoppers (Hemiptera: Fulgoromorpha). Zoomorphology, 134, 63–77.
https://doi.org/10.1007/s00435-014-0243-6
Campodonico, J.F. & Zahniser, J.N. (2017) A new genus and species of grass specialist short-winged leafhopper from Chile and Argentina (Hemiptera: Cicadellidae: Deltocephalinae: Faltalini). Acta Entomologica Musei Nationalis Pragae, 57 (2), 381–390.
https://doi.org/10.1515/aemnp-2017-0082
Castro-Huertas, V., Forero, D. & Grazia, J. (2021) Evolution of wing polymorphism and genital asymmetry in the thread-legged bugs of the tribe Metapterini Stål (Hemiptera, Reduviidae, Emesinae) based on morphological characters. Systematic Entomology, 46, 28–43.
https://doi.org/10.1111/syen.12445
Cheng, Y.J. (1980) New leafhopper taxa (Homoptera: Cicadellidae: Deltocephalinae) from Paraguay. Journal of the Kansas Entomological Society, 53, 61–111.
Crumb, S.E. (1911) The Jassoidae of Kansas. Transactions of the Kansas Academy of Science, 24, 232–238.
https://doi.org/10.2307/3624590
Cwikla, P.S. & Blocker, H.D. (1981) Neotropical genera of Deltocephalinae not included in Linnavuori’s 1959 key. Bulletin of the Entomological Society of America, 27 (3), 170–178.
https://doi.org/10.1093/besa/27.3.170
DeLong, D.M. (1944) A new genus (Tenucephalus) and species of Mexican leafhopper related to Parabolocratus. Ohio Journal of Science, 44, 236–237.
DeLong, D.M. & Knull, D.J. (1946) Check list of the Cicadellidae (Homoptera) of America, north of Mexico. Ohio State University Graduate School Studies, Biological Science Series, 1, 1–102.
DeLong, D.M. & Thambimuttu, C.C. (1973) Three closely related new genera and five new species of shortwinged Chilean leafhoppers (Homoptera: Cicadellidae). Florida Entomologist, 56, 165–172.
https://doi.org/10.2307/3493121
Denno, R.F., Olmstead, K.L. & McCloud, E.S. (1989) Reproductive cost of flight capability: a comparison of life history traits in wing dimorphic planthoppers. Ecological Entomology, 14, 31–44.
https://doi.org/10.1111/j.1365-2311.1989.tb00751.x
Dietrich, C.H. (2005) Keys to the families of Cicadomorpha and subfamilies and tribes of Cicadellidae (Hemiptera: Auchenorrhyncha). Florida Entomologist, 88 (4), 502–517.
https://doi.org/10.1653/0015-4040(2005)88[502:KTTFOC]2.0.CO;2
Dietrich, C.H. (2017) South American leafhoppers of the tribe Hecalini (Hemiptera: Auchenorrhyncha: Cicadellidae: Deltocephalinae). Entomologica Americana, 122 (3), 398–404.
https://doi.org/10.1664/1947-5144-122.3.398
Dietrich, C.H., Allen, J.M., Lemmon, A.R., Lemmon, E.M., Takiya, D.M., Evangelista, O., Walden, K.K.O., Grady, P.G.S. & Johnson, K.P. (2017) Anchored hybrid enrichment-based phylogenomics of leafhoppers and treehoppers (Hemiptera: Cicadomorpha: Membracoidea). Insect Systematics and Diversity, 1 (1), 57–72.
https://doi.org/10.1093/isd/ixx003
Dietrich, C.H. & Goncalves, A.C. (2014) New Baltic amber leafhoppers representing the oldest Aphrodinae and Megophthalminae (Hemiptera, Cicadellidae). European Journal of Taxonomy, 74, 1–13.
https://doi.org/10.5852/ejt.2014.74
Dietrich, C.H. & Perkovsky, E.E. (2020) A new genus and species representing the first leafhopper (Hemiptera: Cicadellidae) from Eocene Rovno amber. Palaeoentomology, 3 (2), 180–187.
https://doi.org/10.11646/palaeoentomology.3.2.7
Dudley, R. (2000) The biomechanics of insect flight: form, function, evolution. Princeton University Press, Princeton, New Jersey, 476 pp.
https://doi.org/10.1515/9780691186344
Evans, J.W. (1947) A natural classification of leaf-hoppers (Jassoidea,Homoptera). Part 3: Jassidae. Transactions of the Royal Entomological Society of London, 98, 105–271.
https://doi.org/10.1111/j.1365-2311.1947.tb01054.x
Freytag, P.H. & Gaiani, M. (2017) Neotropical Leafhoppers. Available from: http://neotropicalcicadellidae.myspecies.info/ (accessed 25 January 2020)
Freytag, P.H. & Ma, N. (1988) An unusual new genus and species of hecaline leafhopper from Venezuela (Homoptera: Cicadellidae). Entomological News, 99 (3), 153–156.
Giesbrecht, R.M. (2015) Estações Ferroviárias, Available from: http://www.estacoesferroviarias.com.br/rmv_tronco/jussaral.htm (accessed 14 March 2020)
Hamilton K.G.A. (1975) Review of the tribal classification of the leafhopper subfamily Aphrodinae (Deltocephalinae of authors) of the Holarctic region (Rhynchota: Homoptera: Cicadellidae). Canadian Entomologist, 107, 477–498.
https://doi.org/10.4039/Ent107477-5
Hamilton, K.G.A. (1994) Evolution of Limotettix Sahlberg (Homoptera-Cicadellidae) in peatlands, with descriptions of new taxa. Memoirs of the Entomological Society of Canada, 169, 111–133.
https://doi.org/10.4039/entm126169111-1
Hamilton, K.G.A. (2000) Five genera of New-World “Shovel-Headed” and “Spoon-Bill” leafhoppers (Hemiptera: Cicadellidae: Dorycephalini and Hecalini). Canadian Entomologist, 132 (4), 429–503.
https://doi.org/10.4039/Ent132429-4
Hamilton, K.G.A. & Zack, R.S. (1999) Systematics and range fragmentation of the Nearctic genus Errhomus (Rhynchota: Homoptera: Cicadellidae). Annals of the Entomological Society of America, 92, 312–354.
https://doi.org/10.1093/aesa/92.3.312
Ikeda, H., Nishikawa, M. & Sota, T. (2012) Loss of flight promotes beetle diversification. Nature Communications, 3 (648), 1–7.
https://doi.org/10.1038/ncomms1659
Knight, W.J. & Webb, M.D. (1986) The Pacific Iassine cicadellid genus Thalattoscopus (Homoptera). Journal of Natural History, 20, 1051–1082.
https://doi.org/10.1080/00222938600770761
Kramer, S. (1950) The morphology and phylogeny of Auchenorrhynchous Homoptera (Insecta). Illinois Biological Monographs, 20, 1–111.
https://doi.org/10.5962/bhl.title.50340
Lawson, P.B. (1920) The Cicadellidae of Kansas. Kansas University Science Bulletin, 12, 5–306, pls. II–XVII.
Linnavuori, R. (1957) The neotropical Hecalinae (Hom. Cicadellidae). Suomen Hyonteistieteellinen Aikakauskirja, 23 (3), 133–143.
Linnavuori, R. (1959) Revision of the Neotropical Deltocephalinae and some related subfamilies (Homoptera). Annales Zoologici Societatis Zoologicae-Botanicae Fennicae ‘Vanamo’, 20, 1–370.
Linnavuori, R. (1975) Studies on Neotropical Deltocephalinae (Homoptera, Cicadellidae). Notulae Entomologicae, 45, 49–52.
Linnavuori, R.E. & DeLong, D.M. (1976) New Neotropical leafhoppers from Peru and Bolivia (Homoptera: Cicadellidae). Revista Peruana de Entomologia, 19 (1), 29–38.
Linnavuori, R. & DeLong, D.M. (1977) The leafhoppers (Homoptera: Cicadellidae) known from Chile. Brenesia, 12–13, 163–267.
Linnavuori, R.E. & DeLong, D.M. (1977b) New deltocephaline leafhoppers from Central America (Homoptera: Cicadellidae) and illustrations of some Osborn Bolivian species. Journal of the Kansas Entomological Society, 50 (4), 558–568.
Linnavuori, R.E. & DeLong, D.M. (1978) Seventeen new species and three new genera of Central and South American Deltocephalini (Homoptera: Cicadellidae). Brenesia, 14 & 15, 195–226.
Linnavuori, R. & Heller, F. (1962) Beitrag zur Cicadelliden-Fauna von Peru. Entomologische Ergebnisse der Stuttgarter Anden-Expedition 1957-Nr. 1. Stuttgarter Beitraege zur Naturkunde aus dem Staatlichen Museum fur Naturkunde in Stuttgart, 67, 1–14.
Medeiros, M.J. & Gillespie, R.G. (2011) Biogeography and the evolution of flightlessness in a radiation of Hawaiian moths (Xyloryctidae: Thyrocopa). Journal of Biogeography, 38, 101–111.
https://doi.org/10.1111/j.1365-2699.2010.02402.x
Mejdalani, G. (1998) Morfologia externa dos Cicadellinae (Homoptera, Cicadellidae): comparação entre Versigonalia ruficauda (Walker) (Cicadellini) e Tretogonia cribrata Melichar (Proconiini), com notas sobre outras espécies e análise da terminologia. Revista Brasileira de Zoologia, 15, 451–544.
https://doi.org/10.1590/S0101-81751998000200015
Metcalf, Z.P. (1963) General Catalogue of the Homoptera. Fascicle VI. Cicdelloidea. Part 9. Hecalidae. U.S. Department of Agriculture, Agricultural Research Service, Washington, D.C., 123pp.
Metcalf, Z.P. (1967) General Catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Part 10. Euscelidae. Section II. U.S. Department of Agriculture, Agricultural Research Service, Washington, D.C., pp. 1078–2074.
Mitterboeck, T.F. & Adamowicz, S.J. (2013) Flight loss linked to faster molecular evolution in insects. Proceedings of the Royal Society B: Biological Sciences, 280 (1767), 20131128, 1–8.
https://doi.org/10.1098/rspb.2013.1128
Nielson, M.W. (1996) A New Genus, Hecalocorica, and a New Species of Hecaline leafhopper from Costa Rica (Homoptera: Cicadellidae). Entomological News, 107 (3), 125–127.
Nova Coelho, L.B. & Ribeiro DaSilva, E. (2001) New records for leafhoppers (Homoptera: Cicadellidae) from Brazil. Revista de Biologia Tropical, 49 (3–4), 1282–1283. Available from: http://www.scielo.sa.cr/scielo.php?script=sci_arttext&pid=S0034-77442001000300070&lng=en&tlng=en (accessed 21 September 2019)
Novotny, V. (1994) Relation between temporal persistence of host plants and wing length in leafhoppers (Hemiptera: Auchenorrhyncha). Ecological Entomology, 19, 168–174.
https://doi.org/10.1111/j.1365-2311.1994.tb00407.x
Oman, P.W. (1938) A generic revision of American Bythoscopinae and South American Jassinae. University of Kansas Science Bulletin, 37, 343–420.
Oman, P.W. (1949) The Nearctic leafhoppers. A generic classification and check list. Memoirs of the Entomological Society of Washington, 3, 1–253.
Oman, P.W., Knight, W.J. & Nielson, M.W. (1990) Leafhoppers (Cicadellidae): a Bibliography, Generic Check-list, and Index to the World Literature 1956–1985. CAB International Institute of Entomology, Wallingford, 368 pp.
Oman, P.W. & Musgrave, C.A. (1975) The Nearctic genera of Errhomenini (Homoptera: Cicadellidae). Melanderia, 21, 1–14.
Paradell, S. & Remes Lenicov, A.M.M. (2005) Redescription of Clorindaia hecaloides Linnavuori, 1975, and distribution in the Argentina Republic (Hemiptera, Cicadellidae). Bulletin de la Société entomologique de France, 110, 133–136.
Paradell, S. & Zahniser, J.N. (2018) Designation of a neotype for Clorindaia hecaloides Linnavuori (Hemiptera: Cicadellidae: Deltocephalinae: Faltalini). Zootaxa, 4532 (2), 293–295.
https://doi.org/10.11646/zootaxa.4532.2.9
Qin, D. & Zhang, Y. (2009) A revision of Malaxella Ding & Hu (Hemiptera: Delphacidae) with description of a new species. Zootaxa, 2208 (1), 44–50.
https://doi.org/10.11646/zootaxa.2208.1.3
Rakitov, R.A. (1998) On differentiation of cicadellid leg chaetotaxy (Homoptera, Auchenorrhyncha, Membracoidea). Russian Entomological Journal, 6, 7–27.
Rakitov, R.A. (2009) Brochosomal coatings of the integument of leafhoppers (Hemiptera, Cicadellidae). In: Gorb, S.N. (Ed.), Functional surfaces in biology, vol. 1. Springer, Dordrecht, pp. 113–137.
https://doi.org/10.1007/978-1-4020-6697-9_8
Remes Lenicov, A.M. & Paradell, S.L. (2009) Argentinean species of the Faltala leafhopper group (Hemiptera: Cicadellidae: Deltocephalinae). Revista de la Sociedad Entomológica Argentina, 68, 263–269.
Ren, F., Huang, Y., Zheng, L. & Qin, D. (2015) Review of the genus Malaxella (Hemiptera: Fulgoroidea: Delphacidae) endemic in China, with a description of a new species. Florida Entomologist, 98 (1), 104–113.
https://doi.org/10.1653/024.098.0117
Rios, N., Bart Jr., H.L., Sturgill, C., Park, S., Abibou, D., Ranipeta, A., Belame, S., Huan, S., Boutte, J., Tseng, A. & Spears, N. (2020) GEOLocate, Available from: https://www.geo-locate.org/ (accessed 19 June 2020)
Roff, D.A. (1986) The evolution of wing dimorphism in insects. Evolution, 40, 1009–1020.
https://doi.org/10.1111/j.1558-5646.1986.tb00568.x
Roff, D.A. (1990) The evolution of flightlessness in insects. Ecological Monographs, 60, 389–421.
https://doi.org/10.2307/1943013
Sattler, K. (1991) A review of wing reduction in Lepidoptera. Bulletin of the British Museum of Natural History, Entomology, 60, 243–288.
Scherbakov, D.A. (2020) The earliest fully brachypterous auchenorrhynchan from Cretaceous Burmese amber (Homoptera: Fulgoroidea: Jubisentidae). Russian Entomological Journal, 29 (1), 6–11.
https://doi.org/10.15298/rusentj.29.1.02
Scholtz, C.H. (2000) Evolution of flightlessness in Scarabaeoidea (Insecta, Coleoptera). Deutsche Entomologische Zeitschrift, 47 (1), 5–28.
https://doi.org/10.1002/mmnd.4800470102
Schuh, R.T. & Slater, J.A. (1995) True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, New York, 336 pp.
Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: https://www.simplemappr.net (accessed 18 June 2020)
Snäll, N., Tammaru, T., Wahlberg, N., Viidalepp, J., Ruohomaki, K., Savontaus, M.L. & Huoponen, K. (2007) Phylogenetic relationships of the tribe Operophterini (Lepidoptera, Geometridae): a case study of the evolution of female flightlessness. Biological Journal of the Linnean Society, 92, 241–252.
https://doi.org/10.1111/j.1095-8312.2007.00834.x
Southwood, T.R.E. (1962) Migration of terrestrial arthropods in relation to habitat. Biological Reviews, 37, 171–214.
https://doi.org/10.1111/j.1469-185X.1962.tb01609.x
Strömberg, C.A.E. (2011) Evolution of grasses and grassland ecosystems. Annual Review of Earth and Planetary Sciences, 39, 517–544.
https://doi.org/10.1146/annurev-earth-040809-152402
Swofford, D.L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]
Van Duzee, E.P. (1916) Check List of Hemiptera (excepting the Aphididae, Aleurodidae, and Coccidae) of America North of Mexico. New York Entomological Society, New York, New Yorks, xi + 111 pp.
https://doi.org/10.5962/bhl.title.7967
Waloff, N. (1983) Absence of wing polymorphism in the arboreal, phytophagous species of some taxa of temperate Hemiptera: an hypothesis. Ecological Entomology, 8, 229–232.
https://doi.org/10.1111/j.1365-2311.1983.tb00502.x
Wagner, D.L. & Liebherr, J.K. (1992) Flightlessness in insects. Trends in Ecology and Evolution, 7 (7), 216–220.
https://doi.org/10.1016/0169-5347(92)90047-F
Wahlberg, N., Snäll, N., Viidalepp, J., Ruohomäki, K. & Tammaru, T. (2010) The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae). Molecular Phylogenetics and Evolution, 55, 929–938.
https://doi.org/10.1016/j.ympev.2010.01.025
Wei, C., Webb, M.D. & Zhang, Y. (2010) A review of the morphologically diverse leafhopper subfamily Stegelytrinae (Hemiptera: Cicadellidae) with description of new taxa. Systematic Entomology, 35 (1), 19–58.
https://doi.org/10.1111/j.1365-3113.2009.00480.x
Zahniser, J.N. (2007) An online interactive key and searchable database of Deltocephalinae (Hemiptera: Cicadellidae). Available from: http://zahniser.speciesfile.org/ (accessed 20 June 2020)
Zahniser, J.N. & Dietrich, C.H. (2010) Phylogeny of the leafhopper subfamily Deltocephalinae (Hemiptera: Cicadellidae) based on molecular and morphological data with a revised family-group classification. Systematic Entomology, 35 (3), 489–511.
https://doi.org/10.1111/j.1365-3113.2010.00522.x
Zahniser, J.N. & Dietrich, C.H. (2013) A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae). European Journal of Taxonomy, 45, 1–211.
https://doi.org/10.5852/ejt.2013.45
Zahniser, J.N. & Dietrich, C.H. (2015) Phylogeny, evolution, and historical biogeography of the grassland leafhopper tribe Chiasmini (Hemiptera: Cicadellidae: Deltocephalinae). Zoological Journal of the Linnean Society, 175, 473–495.
https://doi.org/10.1111/zoj.12292
Zahniser, J.N. & Webb, M.D. (2004) Placement of the Faltala Oman leafhopper group (Hemiptera: Cicadellidae: Deltocephalinae) with descriptions of three new species. Annals of the Entomological Society of America, 97, 667–674.
https://doi.org/10.1603/0013-8746(2004)097[0667:POTFOL]2.0.CO;2
Zanol, K.M.R. (2006) Catalogue of the neotropical (including north of Mexico) Deltocephalinae (Hemiptera, Cicadellidae). Part I—Athysanini and Deltocephalini excluded Acta Biológica Paranaense, Curitiba, 35 (3–4), 89–161.
https://doi.org/10.5380/abpr.v35i0.6869
Zanol, K.M.R. (2008) Catalogue of the Neotropical Deltocephalinae (Hemiptera: Cicadellidae). Part III – Tribe Athysanini. Acta Biológica Paranaense, Curitiba, 37 (1–2), 1–104.
Zeng, Y., O’Malley, C., Singhal, S., Rahim, F., Park, S., Chen, X. & Dudley, R. (2020) A tale of winglets: evolution of flight morphology in stick insects. Frontiers in Ecology and Evolution, 8 (121), 1–15.
https://doi.org/10.3389/fevo.2020.00121
Zhang, C.-X., Brisson, J.A. & Xu, H.-J. (2019) Molecular mechanisms of wing polymorphism in insects. Annual Review of Entomology, 64, 297–314.
https://doi.org/10.1146/annurev-ento-011118-112448