Athysanini Van Duzee, 1892
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publication ID |
https://doi.org/10.5852/ejt.2013.45 |
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publication LSID |
lsid:zoobank.org:pub:41B10E4D-7DAB-40CA-A8FE-4ECA078E04A3 |
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DOI |
https://doi.org/10.5281/zenodo.3844611 |
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persistent identifier |
https://treatment.plazi.org/id/6903BC00-A36A-FFCD-AC71-E02C2AA6F9E5 |
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treatment provided by |
Carolina (2020-05-12 20:03:55, last updated 2024-11-26 00:28:06) |
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scientific name |
Athysanini Van Duzee, 1892 |
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Athysanini Van Duzee, 1892 View in CoL View at ENA
Fig. 11 View Fig
Type genus: Athysanus Burmeister, 1838 View in CoL .
= Phrynomorphini Kirkaldy, 1907.
= Thamnotettigini Distant, 1908.
= Anoterostemmini Haupt, 1929 syn. nov.
= Bobacellini Kusnezov, 1929.
= Platymetopiini Haupt, 1929 View in CoL syn. nov.
= Colladonini Bliven, 1955.
= Cerrillini Linnavuori, 1975.
= Allygidiina Dmitriev, 2006 syn. nov.
Diagnosis
Because this is such a large tribe, it is difficult or impossible to provide a set of characters that will easily diagnose it. There is substantial morphological diversity in the group, but most members have a Y -shaped connective and lack the distinctive features of other tribes.
Description
HEAD. Head subequal to or wider than pronotum or distinctly narrower than pronotum. Discal portion of crown glabrous, radially or longitudinally striate, or shagreen. Anterior margin of head shagreen, glabrous, irregularly textured, with numerous transverse striations, foliaceous, with 2 or 3 parallel carinae, or with numerous carinae. Frontoclypeus not tumid or tumid; texture shagreen or rugose. Clypellus parallel-sided or widening apically; apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus near base or distinctly narrower than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head or long, 1.5 x width of head or longer. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge absent or weakly developed (carinate or weakly carinate). Ocelli present; close to eyes or distant from eyes; on anterior margin of head.
THORAX. Pronotum lateral margin carinate or not; lateral margin shorter than basal width of eye.
WINGS. Forewing brachypterous to macropterous; appendix absent or reduced (in submacropterous or brachypterous forms) or restricted to anal margin; with 3 anteapical cells; veins not raised; with or without reflexed costal veins; A1-A2 crossvein absent or present; apical venation usually not strongly reticulate, rarely reticulate.
LEGS. Profemur with AM1 absent or reduced, with AM1 seta only, or with AM1 and with one or more additional proximal setae; intercalary row with one row of five or more fine setae; row AV with thin, hair-like setae, without setae, with short, stout setae, or with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1 or 2+2+1 with additional proximal macrosetae. Metatarsomere I not expanded apically or expanded apically; plantar setae usually simple, tapered.
MALE GENITALIA. Valve articulated with pygofer or fused to subgenital plates; lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous cleft present; macrosetae absent or reduced (≤ two rows) or macrosetae well differentiated into several rows. Subgenital plates free from each other; articulated with or rarely fused to valve; macrosetae absent or present, scattered and irregularly arranged, uniseriate laterally, with two lateral rows, with some irregularly arranged macrosetae near lateral margin, or uniseriate and distant from lateral margin. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced or present, connected or articulated to connective or near base of aedeagus. Aedeagus with single shaft and gonopore. Connective anterior arms somewhat divergent, Y - or U -shaped, or rarely anterior arms closely appressed anteriorly; articulated with or rarely fused to aedeagus.
FEMALE GENITALIA. Pygofer with numerous macrosetae.Ovipositor not protruding far beyond or protruding far beyond pygofer apex. First valvula convex or not strongly convex; dorsal sculpturing pattern strigate, concatenate, reticulate, or imbricate (with overlapping scales); sculpturing reaching dorsal margin or submarginal; with indistinctly delimited ventroapical sculpturing or rarely with ventroapical sculpturing distinctly delimited. Second valvula abruptly broadened medially or subapically or broad, gradually tapered; with or without dorsal median tooth; dorsal teeth on apical 1/3 or more; teeth large, regularly shaped or small, regularly or irregularly shaped.
Geography and ecology
Distribution: cosmopolitan. Athysanini can be found in nearly all terrestrial ecosystems. Athysanini feed on a wide variety of eudicots and some species occasionally feed on grasses or sedges. This group contains numerous vectors of crop diseases. Some of the most well-known include: Colladonus clitellarius (Say, 1830) , C. geminatus (Van Duzee, 1890) and C. montanus (Van Duzee, 1892) , which transmit Eastern and Western X-diseases of peach, aster yellows, yellow leaf roll stain, and Euscelis incisus (Kirschbaum, 1858) , a vector of numerous diseases of clover.
Remarks
This is the largest tribe of Deltocephalinae , including 228 genera and 1123 species. Habitus images are available for most genera of the tribe through the online database. Athysanini constitute a polyphyletic assemblage of genera which have historically been placed in the tribe mostly because they retain the most common deltocephaline external and genitalic characters but lack the distinctive features that define other tribes. More detailed studies of the phylogeny of Deltocephalinae including many more representatives of Athysanini are needed to more fully understand the relationships of these genera and the evolution of all of Deltocephalinae . Such studies may also help to determine morphological characters that define smaller, more particular lineages, particularly those belonging to clades that received strong branch support in our phylogenetic analyses. For now, the revised classification presented here including the new tribes Bahitini and Phlepsiini and the revised interpretation of Scaphoideini separates some groups of genera which were previously placed in Athysanini or were previously unplaced to tribe. Revisions to Athysanini here include the addition of many genera that were previously unplaced in Deltocephalinae , thus resulting in a classification of the subfamily that has no genera unplaced to tribe. This arrangement seems preferable to an alternative option of restricting Athysanini to the interpretation of the Athysanus genus group, as defined here, and considering all other genera once included in Athysanini to be unplaced to tribe in Deltocephalinae . The latter option would likely create more confusion, as did the previous classification which contained both unplaced genera and a large number of genera in Athysanini without any clear distinction between the two.
Other changes made here include considering Allygidiina, Anoterostemmina, and Platymetopiina synonyms of Athysanini . Emeljanov (1999) and Dmitriev (2006b) recognized up to 4 subtribes of Athysanini (Allygidiina, Athysanina, Cicadulina , and Platymetopiina) although their generic classifications partly conflict. Both authors included the genera treated here as Cicadulini in Athysanini (Emeljanov in Athysanina, and Dmitriev as the subtribe Cicadulina ). The phylogenetic analyses here included representatives of 5 of the 14 genera included in Platymetopiina by Dmitriev (2006b): Anoplotettix , Colladonus , Phlepsius , Platymetopius , and Thamnotettix . Except for a relationship between Colladonus and Platymetopius , the analyses strongly suggest that these genera are not closely related and that the morphological characters previously used to define the group are not informative of phylogenetic relationships. Therefore, Platymetopiina is not recognized here. Further investigation may allow for a revised concept of this group. The genera Eusama and Twiningia were found with strong branch support to be related to Colladonus and Platymetopius , which could in part inform future recognition of such a taxon. The phylogenetic analyses grouped the included representatives of Athysanina and Allygidiina in the same area of the tree and they share some morphological features. They are therefore recognized together in the informal Athysanus genus group, and some other genera that were not within the geographical ranges of the previous taxonomic studies are included in the group.
The morphologically unique Neotropical genus Chimaerotettix was included in our analyses, but its placement with respect to other Athysanini was poorly resolved.
Justification for the inclusion of other genera not covered in the previous discussion is given below.
Egenus was previously included in Faltalini by Zahniser & Dietrich (2010) who had not examined female specimens. It is transferred to Athysanini here because female specimens became available for study which showed that they do not possess the synapomorphies of the first and second valvulae that in part define Faltalini. Preliminary phylogenetic work on the tribe Hecalini (Catanach, personal communication) suggests that Egenus may be related to Arrugada .
Loralia is transferred to Athysanini from Deltocephalini . It clearly does not belong in Deltocephalini and was found here to be closely related to Occinirvanini. However, we were unable to identify morphological features uniting Loralia and Occinirvana , so more study of the Australian fauna will be needed to determine whether including Loralia or other genera in Occinirvanini is justifiable.
Nesothamnus is transferred from Scaphytopiini . It is not closely related to Scaphytopius but is potentially more closely related to some Neotropical Athysanini . In several analyses, it was resolved as sister to or near Idioceromimus with which it shares a relatively short, broad body form and bright orange/red coloration.
Phycotettix was listed in Fieberiellini by Oman et al. (1990). It does not possess the characters that are diagnostic for Fieberiellini and its placement there was perhaps a mistake due to the similarity of the name to the fieberielline genus Placotettix . Phycotettix was not included in Fieberiellini in a comprehensive revision of the tribe (Meyer-Arndt & Remane 1992a, b).
Selected references
Oman (1949), Linnavuori (1959), Linnavuori & DeLong (1978b), Cwikla & Blocker (1981), Anufriev & Emeljanov (1988), Emeljanov (1999), Dmitriev (2006b).
Included genera
Acunasus DeLong, 1945 placement nov. (previously unplaced in Deltocephalinae )
Aeternus Distant, 1918 placement nov. (previously unplaced in Deltocephalinae )
Alladanus DeLong & Harlan, 1968
Allotapes Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Anaemotettix Korolevskaya, 1980 placement nov. (previously placed in Platymetopiina)
Ancudana DeLong & Martinson, 1974
Aplanusiella Nielson & Haws, 1992
Asthenotettix Korolevskaya, 1980 placement nov. (previously placed in Platymetopiina)
Awasha Heller & Linnavuori, 1968
Bardana DeLong, 1980 placement nov. (previously unplaced in Deltocephalinae )
Bicoloratum Dai & Li, 2011
Chimaerotettix Dietrich & Rakitov, 2002 placement nov. (previously unplaced in Deltocephalinae )
Cocrassana Blocker & Larsen, 1991
Colladonus Ball, 1936 placement nov. (previously placed in Platymetopiina)
Comayagua Linnavuori & DeLong, 1978
Consepusa Linnavuori & DeLong, 1977
Cozadanus DeLong & Harlan, 1968
Crassana DeLong & Hershberger, 1947
Cyanidius Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Dampfiana DeLong & Hershberger, 1948
Danbara Oman, 1949 placement nov. (previously unplaced in Deltocephalinae )
Desertana DeLong & Martinson, 1973
Egenus Oman, 1938 placement nov. (transferred from Faltalini)
Ephelodes Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Euleimonios Kirkaldy, 1906 placement nov. (previously unplaced in Deltocephalinae )
Eusama Oman, 1949 placement nov. (previously placed in Platymetopiina)
Eusceloidia Osborn, 1923 placement nov. (previously unplaced in Deltocephalinae )
Fistulatus Zhang, 1997 placement nov. (previously unplaced in Deltocephalinae )
Gunawardenea Fletcher & Moir, 2008
Hamulotettix Dai & Zhang, 2011
Hardya Edwards, 1922 placement nov. (previously placed in Platymetopiina)
Idioceromimus Dietrich & Rakitov, 2002 placement nov. (previously unplaced in Deltocephalinae )
Idiodonus Ball, 1936 placement nov. (previously placed in Platymetopiina)
Inghamia Evans, 1966 placement nov. (previously placed in Platymetopiina)
Jakarellus Webb, 1980 placement nov. (previously placed in Platymetopiina)
Krameraxus Maldonado-Capriles, 1968 placement nov. (previously unplaced in Deltocephalinae )
Lamprotettix Ribaut, 1952 placement nov. (previously placed in Platymetopiina)
Lascumbresa Linnavuori & DeLong, 1979
Latinocesa Kocak & Kemal, 2010 placement nov. (previously unplaced in Deltocephalinae )
Lojanus Linnavuori, 1959 placement nov. (previously unplaced in Deltocephalinae )
Loralia Evans, 1966 placement nov. (transferred from Deltocephalini )
Malasiella Evans, 1954 placement nov. (previously placed in Platymetopiina)
Nakaharanus Ishihara, 1953 placement nov. (previously placed in Platymetopiina)
Napo Linnavuori & DeLong, 1976
Neodonus DeLong & Hershberger, 1948 placement nov. (previously placed in Platymetopiina)
Neohegira Linnavuori & DeLong, 1978
Nesothamnus Linnavuori, 1959 placement nov. (transferred from Scaphytopiini )
Norvellina Ball, 1931 placement nov. (previously placed in Platymetopiina)
Nurenus Oman, 1949 placement nov. (previously placed in Platymetopiina)
Ophionotum Emeljanov, 1964 placement nov. (previously placed in Allygidiina)
Paralaca Lozada, 1998 placement nov. (previously unplaced in Deltocephalinae )
Paramacrosteles Dai, Li & Chen, 2006
Perundanus Zanol, 1993 placement nov. (previously unplaced in Deltocephalinae )
Phlepsobahita Linnavuori, 1959
Phycotettix Haupt, 1929 placement nov. (transferred from Fieberiellini )
Platymetopius Burmeister, 1838 placement nov. (previously placed in Platymetopiina)
Poliona Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Pseudalaca Linnavuori, 1959 placement nov. (previously unplaced in Deltocephalinae )
Pseudaligia Kramer & DeLong, 1968
Pseudocephalelus Linnavuori, 1969
Rinconada Linnavuori & DeLong, 1977
Saudallygus Dlabola, 1979 placement nov. (previously unplaced in Deltocephalinae )
Scaphytoceps Dlabola, 1957 placement nov. (transferred from Scaphytopiini )
Scenergates Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Sincholata DeLong, 1982 placement nov. (previously unplaced in Deltocephalinae )
Sobara Oman, 1949 placement nov. (previously unplaced in Deltocephalinae )
Stymphalella Evans, 1954 placement nov. (previously placed in Platymetopiina)
Tapetia Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Thanomahia Dlabola, 1987 placement nov. (previously unplaced in Deltocephalinae )
Tingolix Linnavuori & DeLong, 1978
Twiningia Ball, 1931 placement nov. (previously placed in Platymetopiina)
Anufriev G. A. & Emeljanov A. F. 1988. Suborder Cicadinea - (Auchenorrhyncha). In: Lehr P. A. (ed.) Keys to the Insects of the Far East of the USSR. Volume II: 1 - 10. Nauka Publishing House, Leningrad.
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.
Dai W., Viraktamath C. A., Zhang Y. & Webb M. D. 2009. A review of the leafhopper genus Scaphotettix Matsumura (Hemiptera: Cicadellidae: Deltocephalinae), with description of a new genus. Zoological Science 26: 656 - 663. http: // dx. doi. org / 10.2108 / zsj. 26.656
Dai W., Dietrich C. H. & Zhang Y. 2011. Review of the Oriental leafhopper genus Lampridius (Hemiptera: Cicadellidae: Deltocephalinae), with description of a related new genus. Zoological Science 28: 380 - 387. http: // dx. doi. org / 10.2108 / zsj. 28.380
DeLong D. M. 1938. The genus Phlepsius. A study of the North American species with special reference to the characters of the male genitalia. Lloydia 1: 232 - 252.
DeLong D. M. 1945. The genus Chlorotettix (Homoptera-Cicadellidae) in Mexico. Lloydia 8: 1 - 30.
Dietrich C. H. & Rakitov R. A. 2002. Some remarkable new deltocephaline leafhoppers (Hemiptera: Cicadellidae: Deltocephalinae) from the Amazonian rainforest canopy. Journal of the New York Entomological Society 110 (1): 1 - 48.
Distant W. L. 1908. Rhynchota - Homoptera. In: Bingham C. T. (ed.) The Fauna of British India, including Ceylon and Burma. Volume 4. Taylor and Francis, London.
Distant W. L. 1918. Rhynchota. Homoptera: Appendix. Heteroptera: Addenda. In: Shipley A. E. & Marshall G. A. K. (eds) The Fauna of British India, including Ceylon and Burma. Volume 7. Taylor and Francis, London.
Dmitriev D. A. 2006 a. 3 I, a new program for creating Internet-accessible interactive keys and taxonomic databases and its application for taxonomy of Cicadina (Homoptera). Russian Entomological Journal 15 (3): 263 - 268.
Dmitriev V. D. [sic] 2006 b. Larvae of leafhoppers of the subfamily Deltocephalinae (Homoptera, Cicadellidae) of European Russia and adjacent territories: IV. Tribe Athysanini. Entomological Review 86 (1): 11 - 48. http: // dx. doi. org / 10.1134 / S 0013873806010027
Emeljanov A. F. 1962. New tribes of the leafhopper subfamily Euscelinae (Auchenorrhyncha, Cicadellidae). Entomologicheskoe Obozrenie 41: 388 - 397. [In Russian with English summary, translation in Entomological Review 41 (2): 236 - 240]
Emeljanov A. F. 1999. A key to genera of the subfamily Deltocephalinae s. l. (Homoptera, Cicadellidae) from Kazakhstan, Middle Asia, and Mongolia, with description of new genera and subgenera. Entomological Review 79: 547 - 562.
Evans J. W. 1954. Les Cicadellidae de Madagascar (Homopteres). Memoires de l'Institut Scientifique de Madagascar (Serie E) 4: 87 - 137.
Evans J. W. 1966. The leafhoppers and froghoppers of Australia and New Zealand (Homoptera: Cicadelloidea and Cercopoidea). Memoirs of the Australian Museum 12: 1 - 347.
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. 1961. Hemiptera (Homoptera): Cicadellidae. In: Hanstrom B., Brinck P. & Rudebeck G. (eds) South African Animal Life 8: 452 - 486.
Linnavuori R. 1969. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries XCIII. Hemipteres Hylicidae et Cicadellidae. Bulletin de l'Institut Fondamental d'Afrique Noire (Series A) 31 (4): 1129 - 1185.
Linnavuori R. 1975. Revision of the Cicadellidae (Homoptera) of the Ethiopian region 3. Deltocephalinae, Hecalini. Acta Zoologica Fennica 143: 1 - 37.
Linnavuori R. & DeLong D. M. 1977 a. The leafhoppers (Homoptera: Cicadellidae) known from Chile. Brenesia 12 - 13: 163 - 267.
Linnavuori R. & DeLong D. M. 1978 b. Neotropical leafhoppers of the Bahita group (Homoptera: Cicadellidae: Deltocephalinae). A contribution to the taxonomy. Brenesia 14 - 15: 109 - 169.
Linnavuori R. & DeLong D. M. 1978 c. Genera and species of Hecalini (Homoptera, Cicadellidae, Deltocephalinae) known to occur in Mexico. Annales Entomologici Fennici 44: 48 - 53.
McKamey S. H. 2003 a. Review of the New World leafhopper genus Excultanus Oman (Hemiptera: Cicadellidae: Deltocephalinae). Proceedings of the Entomological Society of Washington 105 (2): 292 - 299.
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.
Meyer-Arndt S. & Remane R. 1992 a. Phylogenie und Speziation der Fieberiellini Wagner, 1951 (Homoptera: Auchenorrhyncha: Cicadellidae). Teil 1. Marburger Entomologische Publikationen 2 (7): 1 - 387.
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. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.2010.00522. x
Zhang Y. & Webb M. D. 1996. A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae). Bulletin of the Natural History Museum (Entomology) 65 (1): 1 - 103.
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
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Class |
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Order |
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Family |
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SubFamily |
Deltocephalinae |
Athysanini Van Duzee, 1892
| Zahniser, James N. & Dietrich, Chris H. 2013 |
| Phrynomorphini Kirkaldy, 1907 |
| Thamnotettigini Distant, 1908 |
Euscelini
| Euscelini Van Duzee, 1917 |
Anoterostemmini
| Anoterostemmini Haupt, 1929 |
| Bobacellini Kusnezov, 1929 |
Platymetopiini
| Platymetopiini Haupt, 1929 |
| Colladonini Bliven, 1955 |
| Cerrillini Linnavuori, 1975 |
| Allygidiina Dmitriev, 2006 |
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