Membracidae

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Paraneoptera
Superordo: Condylognatha
Ordo: Hemiptera
Subordo: Auchenorrhyncha
Infraordo: Cicadomorpha
Superfamilia: Membracoidea

Familia: Membracidae
Subfamiliae (9): Centronodinae – Centrotinae – Darninae – Endoiastinae – Heteronotinae – Membracinae – Nicomiinae – Smiliinae – Stegaspidinae Centrotinae - Heteronotinae - Membracinae - Nicomiinae - Smiliinae - Stegaspidinae

Genera Incertae sedis (5): Brachytalis – Problematode – Smergotomia – Smilidarnis – Togotolania
Overview of genera
A

Abelus – Acanthicoides – Acanthophyes – Acanthucalis – Acanthuchus – Achantomedes – Aconophora – Acutalis – Adippe – Afraceronotus – Alchisme – Alcmeone – Aleptocentrus – Alobia – Alocanthella – Alocebes – Alosextius – Amastris – Amblycentrus – Amblyophallus – Amphilobocentrus – Ananthasubramanianum – Anchistrotus – Anchon – Anchonastes – Anchonobelus – Anchonomonoides – Anisostylus – Anobilia – Antialcidas – Antianthe – Antillotolania – Antonae – Anzac – Aphetea – Archasia – Arcuatocornum – Arimanes – Ashmeadea – Aspasiana – Aspona – Atymna – Atymnina – Atypa – Aurinotus – Awania
B

Bajulata – Barsumas – Barsumoides – Bathoutha – Beaufortiana – Bilimekia – Bleccia – Bocchar – Bocydium – Bolbonota – Bolbonotodes – Boocerus – Bordonia – Brachybelus – Brachycentrotus – Brachytalis – Bryantopsis – Bubalopa – Bucktoniella – Bulbauchenia – Bunyella – Butragulus
C

Callicentrus – Calloconophora – Camelocentrus – Campylenchia – Campylocentrus – Capeneralus – Capeneriana – Carynota – Cebes – Centriculus – Centrobelus – Centrochares – Centrodontus – Centrolobus – Centronodus – Centrotus – Centrotusoides – Centrotypus – Centruchoides – Centruchus – Ceraon – Ceresa – Chelyoidea – Choucentrus – Cladonota – Clepsydrius – Coccosterphus – Colisicostata – Cornutalis – Cornutobelus – Creonus – Crito – Cryptaspidia – Cryptoparma – Cymbomorpha – Cyphonia – Cyphotes – Cyrtolobus
D

Dacaratha – Daconotus – Dagonotus – Daimon – Darnis – Darnoides – Daymfus – Dectonura – Deiroderes – Demanga – Dingkana – Dioclophara – Distantobelus – Dograna – Dukeobelus – Dysyncritus
E

Ebhul – Elaphiceps – Emphusis – Enchenopa – Enchophyllum – Endoiastus – Ennya – Entylia – Erechtia – Erecticornia – Erosne – Eualthe – Euceropsila – Eucoccosterphus – Eucyphonia – Eudonica – Eufairmairia – Eufairmairiella – Eufrenchia – Eumela – Eumocentrulus – Eumonocentrus – Eunusa – Euritea – Eutryonia – Euwalkeria – Evanchon – Evansiana
F

Fairmairiana – Farcicaudia – Flatyperphyma – Flexanotus – Flexocentrus – Flynnia – Foliatrotus – Folicarina – Funkhouserella
G

Gargara – Gargarina – Gelastogonia – Germariana – Gigantorhabdus – Glischrocentrus – Glossonotus – Goddefroyinella – Godingia – Goniolomus – Gonoconophora – Grandolobus – Guayaquila
H

Hadrophallus – Hamma – Harmonides – Havilandia – Hebetica – Hebeticoides – Heliria – Helonica – Hemicardiacus – Hemicentrus – Hemikyptha – Hemiptycha – Heranice – Heteronotus – Holdgatiella – Horiola – Hybanda – Hybandoides – Hygris – Hypheodana – Hyphinoe – Hypsauchenia – Hypsolyrium – Hypsoprora – Hypsoprorachis
I

Idioderma – Ilithucia – Imporcitor – Indicopleustes – Insitor – Insitoroides – Iria – Ischnocentrus
J

Jacobiana – Jibarita – Jingkara – Joveriana
K

Kallicrates – Kanada – Kronides
L

Lallemandia – Lanceonotus – Lecythifera – Leioscyta – Leprechaunus – Leptobelus – Leptocentrus – Leptoceps – Leptosticta – Lewdeitzia – Lirania – Lobocentrus – Lubra – Lycoderes
M

Maarbarus – Mabokiana – Machaerotypus – Madlinus – Maguva – Marshallella – Matonotus – Matumuia – Maturnaria – Maurya – Megaloschema – Melusinella – Membracidoidea – Membracis – Mendicea – Menthogonus – Mesocentrina – Metcalfiella – Metheisa – Micreune – Microcentrus – Microschema – Micrutalis – Mitranotus – Monanchon – Monobeloides – Monobelus – Monocentrus – Multareis – Multareoides
N

Nassunia – Negus – Neocanthuchus – Neocentrus – Neomachaerotypus – Neoproterpia – Neosextius – Nessorhinus – Nicomia – Nilautama – Nodonica – Nondenticentrus – Notocera – Notogonioides
O

Occator – Ochrolomia – Ochropepla – Oeda – Omolon – Ophicentrus – Ophiderma – Orekthophora – Orthobelus – Otinotoides – Otinotus – Oxyrhachis
P

Palonica – Pantaleon – Paracentronodus – Paraceresa – Paradarnoides – Paragara – Parantonae – Parapogon – Paraxiphopoeus – Parayasa – Peltosticta – Peltzerella – Penichrophorus – Periaman – Philya – Phormophora – Phyllotropis – Pieltainellus – Platybelus – Platycentrus – Platycotis – Pogon – Pogonella – Pogonotus – Pogonotypellus – Pogontypus – Polonius – Polyglypta – Polyglyptodes – Poppea – Postanomus – Potnia – Potnioides – Problematode – Procyrta – Promitor – Protinotus – Proxolonia – Psilocentrus – Publilia – Pyrgauchenia – Pyrgonota
Q

Quadrinarea
R

Rachinotus – Ramedia – Ramosella – Rentzia – Rexicornia – Rhexia – Rigula
S

Sakakibarella – Sarantus – Saudaraba – Scalmophorus – Scytodepsa – Sertorius – Sextius – Sinodemanga – Sipylus – Smerdalea – Smergotomia – Smilia – Smilidarnis – Smiliorachis – Spalirises – Spathenotus – Spathocentrus – Spinodarnoides – Spissistilus – Stalobelus – Stalomia – Stalotypa – Stegaspis – Stictocephala – Stictodepsa – Stictolobus – Stictopelta – Stilbophora – Stirpis – Streonus – Sturmella – Stylocentrus – Subrincator – Sundarion
T

Takliwa – Tapinolobus – Taunaya – Telamona – Telamonanthe – Telingana – Telonaca – Terentius – Thelia – Thelicentrus – Thrasymedes – Thuris – Tiberianus – Todea – Togotolania – Tolania – Tomogonia – Tortistilus – Trachytalis – Tragopa – Tribulocentrus – Tricentroides – Tricentrus – Trichaetipyga – Tricoceps – Trinarea – Trioxiphus – Tritropidia – Tropidarnis – Tropidolomia – Truncatocornum – Tsunozemia – Tumecauda – Turrialbia – Tylocentrus – Tylopelta – Tynelia – Tyrannotus
U

Umbelligerus – Umbonia – Umfilianus – Undarella – Uroxiphus
V

Vanduzea – Vecranotus – Vestistilus
X

Xantholobus – Xanthosticta – Xiphopoeus
Y

Yaponotus – Yasa
Z

Zanzia – Zigzagicentrus

All genera above are taken from Catalogue of Life: 29th November 2018 edition with new additional valid genera from Treehoppers Insect Museum.org
Name

Membracidae Rafinesque, 1815
References

Camacho, L.F., Flórez-V, C. & Evangelista, O. 2017. Notes on the genus Sakakibarella Creão-Duarte with description of three new species (Membracidae: Membracinae: Hoplophorionini). Pp 90—107 in: Evangelista, O., Takiya, D.M. & Dietrich, C.H. (eds.) 2017. The art and science of describing nature’s surrealists: Festschrift Celebrating the Career of Albino Morimasa Sakakibara. Zootaxa 4281(1): 1–290. DOI: 10.11646/zootaxa.4281.1.1. title page (PDF) Reference page. . Reference page.
Creão-Duarte, A.J., Cabral, V.A. & Laurenco, A.P. 2017. A new species of Lycoderes Sakakibara (Hemiptera, Membracidae, Stegaspidinae) from Brazil. Pp 58—62 In: Evangelista, O., Takiya, D.M. & Dietrich, C.H. (eds.) 2017. The art and science of describing nature’s surrealists: Festschrift Celebrating the Career of Albino Morimasa Sakakibara. Zootaxa 4281(1): 1–290. DOI: 10.11646/zootaxa.4281.1.1. title page (PDF) Reference page. . Reference page.
Creão-Duarte, A.J., Rothéa, R.R.A.D., Lourenço, A., Cabral, V.A. & Evangelista, O. 2017. Two remakable new species of Notocera Amyot and Serville, 1843 (Hemiptera, Membracidae, Hypsoprorini) from the Brazilian Caatinga. Pp 77—89 in: Evangelista, O., Takiya, D.M. & Dietrich, C.H. (eds.) 2017. The art and science of describing nature’s surrealists: Festschrift Celebrating the Career of Albino Morimasa Sakakibara. Zootaxa 4281(1): 1–290. DOI: 10.11646/zootaxa.4281.1.1. title page (PDF) Reference page. . Reference page.
Day, M.F. 1999. The genera of Australian Membracidae (Hemiptera: Auchenorrhyncha). Invertebrate Taxonomy 13(4): 629–747. DOI: 10.1071/IT98010
Dmitriev, D.A. & Deitz, L.L. 2017. Nomenclatural changes in the family Membracidae (Hemiptera, Auchenorrhyncha, Membracoidea). Zootaxa 4365(1): 94–98. DOI: 10.11646/zootaxa.4365.1.9. Full article (PDF) Reference page.
Deitz, L.L.; Wallace, M.S. 2012: Richness of the Nearctic treehopper fauna (Hemiptera: Aetalionidae and Membracidae). Zootaxa 3423: 1–26. Preview PDF Reference page.
Flórez-V, C. 2017. A new species of Cornutalis Sakakibara (Membracidae: Smiliinae: Acutalini) from Colombia. Zootaxa 4272(3): 439–445. DOI: 10.11646/zootaxa.4272.3.9. Reference page.
Gaiani, M.A. 2017. Problematode: an enigmatic new genus of Membracidae (Hemiptera) with two new species from Venezuela. Zootaxa 4281(1): 70–76. DOI: 10.11646/zootaxa.4281.1.7 Reference page.
Gonzalez-Mozo, L., McKamey, S., Ware, J.L. & Hamilton, G. 2017. Two new species of Darnini (Hemiptera: Membracidae) from Colombia and Peru. Pp 108—114 in: Evangelista, O., Takiya, D.M. & Dietrich, C.H. (eds.) 2017. The art and science of describing nature’s surrealists: Festschrift Celebrating the Career of Albino Morimasa Sakakibara. Zootaxa 4281(1): 1–290. DOI: 10.11646/zootaxa.4281.1.1. title page (PDF) Reference page. . Reference page.
Haviland, M. D. 1926. "The Membracidae of Kartabo, Bartica District, British Guiana: With Descriptions of New Species and Bionomical Notes"' Zoologica 6(3): 229-290 (BHL) Reference page.
Flynn, D.J. 2012: Checklist of treehoppers of Panama (Hemiptera: Membracidae) with a list of checklists and keys to the Nearctic and Neotropical fauna. Zootaxa 3405: 35–63. Preview PDF Reference page.
Marino De Remes Lenicov, A.M., Mariani, R., Scelsio, N. & Gutierrez, A. 2010. Membracidae types (Hemiptera-Membracoidea) housed at the Museo de La Plata entomological collection (Argentina). Zootaxa 2566: 21–38. Preview (PDF).
McKamey, S.H. 2017. The identity of three South American “smiliine” treehoppers (Hemiptera, Membracidae) and related taxonomic changes, including description of a new genus in Thuridini. Zookeys 678: 65—72. DOI: 10.3897/zookeys.678.10340. Reference page.
McKamey, S.H.; Brodbeck, B.V. 2013: Immature stages and hosts of two plesiomorphic, Antillean genera of Membracidae (Hemiptera) and a new species of Antillotolania from Puerto Rico. ZooKeys 301: 1–12. DOI: 10.3897/zookeys.301.4234 Reference page.
Özdikmen, H. & Demir, E. 2010. Daymfus nom. nov., a substitute name for the preoccupied genus Strzeleckia Day, 1999 (Hemiptera: Membracidae). Munis Entomology & Zoology 5(1): 303. PDF.

Links

Membracidae - Taxon details in World Auchenorrhyncha Database

Vernacular names
English: Treehoppers
日本語: ツノゼミ科
中文: 角蟬科

Treehoppers (more precisely typical treehoppers to distinguish them from the Aetalionidae) and thorn bugs are members of the family Membracidae, a group of insects related to the cicadas and the leafhoppers. About 3,200 species of treehoppers in over 400 genera are known.[1] They are found on all continents except Antarctica; only five species are known from Europe. Individual treehoppers usually live for only a few months.

Morphology
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A treehopper of the species Heteronotus nodosus investigated by Micro-CT, revealing the interior morphology. The interior of the extended pronotum, in this species mimicking a wasp's body, is empty.

Treehoppers, due to their unusual appearance, have long interested naturalists. They are best known for their enlarged and ornate pronotum, expanded into often fantastic shapes that enhance their camouflage or mimicry, often resembling plant thorns (thus the commonly used name of "thorn bugs" for a number of treehopper species). The specialised pronotum (or helmet) may not be simply an expansion of the prothoracic sclerite, but a fused pair of dorsal appendages of the first thoracic segment. The treehopper has specialized muscles in its hind femora that unfurl to generate sufficient force to jump.[2]

These may be serial homologues of insect wings, which are dorsal appendages of the second and/or third thoracic segments, although this interpretation has been seriously challenged.[3] Evidence for this theory included the development of the helmet, which arises as a pair of appendages attached to each side of the dorsal prothorax by an articulation with muscles and a flexible membrane that allow it to be mobile (also, similar genes are involved in development of the helmet and the wings[4]).

Distinguishing males from females is accomplished only by looking at the genitalia.
Ecology

Treehoppers pierce plant stems with their beaks and feed upon sap. The young can frequently be found on herbaceous shrubs and grasses, while the adults more often frequent hardwood tree species. Excess sap becomes concentrated as honeydew, which often attracts ants. Some species have a well-developed ant mutualism, and these species are normally gregarious as well, which attracts more ants. The ants provide protection from predators. Treehoppers mimic thorns to prevent predators from spotting them.

Others have formed mutualisms with wasps, such as Parachartergus apicalis.[5] Even geckos form mutualistic relations with treehoppers, with whom they communicate by small vibrations of the abdomen.[6]

Eggs are laid by the female with her saw-like ovipositor in slits cut into the cambium or live tissue of stems, though some species lay eggs on top of leaves or stems. The eggs may be parasitised by wasps, such as the tiny fairyflies (Mymaridae) and Trichogrammatidae. The females of some membracid species sit over their eggs to protect them from predators and parasites, and may buzz their wings at intruders. The females of some gregarious species work together to protect each other's eggs. In at least one species, Publilia modesta, mothers serve to attract ants when nymphs are too small to produce much honeydew. Some other species make feeding slits for the nymphs.[7]

Like the adults, the nymphs also feed upon sap, and unlike adults, have an extensible anal tube that appears designed to deposit honeydew away from their bodies. The tube appears to be longer in solitary species rarely attended by ants. It is important for sap-feeding bugs to dispose of honeydew, as otherwise it can become infected with sooty moulds. Indeed, one of the evident benefits of ants for Publilia concava nymphs is that the ants remove the honeydew and reduce such fungal growth.

Most species are innocuous to humans, although a few are considered minor pests, such as Umbonia crassicornis (a thorn bug), the three-cornered alfalfa hopper (Spissistilus festinus), and the buffalo treehopper (Stictocephala bisonia), which has been introduced to Europe. The cowbug Oxyrachis tarandus has been recorded as a pest of Withania somnifera in India.
Systematics

The diversity of treehoppers has been little researched, and their systematic arrangement is tentative. It seems three main lineages can be distinguished; the Endoiastinae are the most ancient treehoppers, still somewhat resembling cicadas. Centrotinae form the second group; they are somewhat more advanced but the pronotum still does not cover the scutellum in almost all of these. The Darninae, Heteronotinae, Membracinae and Smiliinae contain the most apomorphic treehoppers.

Several proposed subfamilies seem to be paraphyletic. Centronodinae and Nicomiinae might need to be merged into the Centrotinae to result in a monophyletic group.

References

Treehoppers. Dr. Metcalf. NCSU Libraries. North Carolina State University.
Burrows, M. (2013) Jumping mechanisms of treehopper insects (Hemiptera, Auchenorrhyncha, Membracidae). https://doi.org/10.1242/jeb.078741
Yoshizawa, K. (2012) The treehopper’s helmet is not homologous with wings (Hemiptera: Membracidae) Systematic Entomology. 37, 2–6.
B. Prud'homme et coll (2011). "Body plan innovation in treehoppers through the evolution of an extra wing-like appendage". Nature. 473 (7345): 83–86. Bibcode:2011Natur.473...83P. doi:10.1038/nature09977. PMID 21544145. S2CID 4401372.
Wetterer, J. K.; et al. (2000). "Diel shifts in treehopper-tending by ants and wasps in Costa Rica (Hymenoptera)" (PDF). Sociobiology. 36: 1.
BBC documentary Life in Cold Blood, episode Dragons of the Dry, minute 22

Lin, C.; et al. (2004). "Molecular phylogenetics and evolution of maternal care in membracine treehoppers" (PDF). Systematic Biology. 53 (3): 400–421. doi:10.1080/10635150490445869. PMID 15503670.