Athysanini Van Duzee, 1892

Zahniser, James N. & Dietrich, Chris H., 2013, A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae), European Journal of Taxonomy 45, pp. 1-211: 37-44

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Athysanini Van Duzee, 1892


Athysanini Van Duzee, 1892  

Fig. 11 View Fig

Type genus: Athysanus Burmeister, 1838   .

= Phrynomorphini Kirkaldy, 1907.

= Thamnotettigini Distant, 1908.

= Euscelini Van Duzee, 1917   .

= Anoterostemmini Haupt, 1929   syn. nov.

= Bobacellini Kusnezov, 1929.

= Platymetopiini Haupt, 1929   syn. nov.

= Colladonini Bliven, 1955.

= Cerrillini Linnavuori, 1975.

= Allygidiina Dmitriev, 2006 syn. nov.


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.


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; A 1- A 2 crossvein absent or present; apical venation usually not strongly reticulate, rarely reticulate.

LEGS. Profemur with AM 1 absent or reduced, with AM 1 seta only, or with AM 1 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.


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

Abrus Dai & Zhang, 2002  

Acacimenus Dlabola, 1979  

Acunasus DeLong, 1945   placement nov. (previously unplaced in Deltocephalinae   )

Aeternus Distant, 1918   placement nov. (previously unplaced in Deltocephalinae   )

Aindrahamia Linnavuori, 1965  

Aligia Ball, 1907  

Alladanus DeLong & Harlan, 1968  

Allophleps Bergroth, 1920  

Allotapes Emeljanov, 1964   placement nov. (previously placed in Platymetopiina)

Amblytelinus Lindberg, 1954  

Anaemotettix Korolevskaya, 1980   placement nov. (previously placed in Platymetopiina)

Ancudana DeLong & Martinson, 1974  

Andanus Linnavuori, 1959  

Angubahita DeLong, 1982  

Angulanus DeLong, 1946  

Antoniellus Linnavuori, 1959  

Aplanatus Cheng, 1980  

Aplanus Oman, 1949  

Aplanusiella Nielson & Haws, 1992  

Arahura Knight, 1975  

Arawa Knight, 1975  

Argyrilla Emeljanov, 1972  

Aricanus Linnavuori, 1959  

Artucephalus DeLong, 1943  

Asiotoxum Emeljanov, 1964  

Asthenotettix Korolevskaya, 1980   placement nov. (previously placed in Platymetopiina)

Atanus Oman, 1938  

Awasha Heller & Linnavuori, 1968  

Bambusana Anufriev, 1969  

Bambusananus Li & Xing, 2011  

Bandara Ball, 1931  

Bardana DeLong, 1980   placement nov. (previously unplaced in Deltocephalinae   )

Baroma Oman, 1938  

Benibahita Linnavuori, 1959  

Bergolix Linnavuori, 1959  

Bicoloratum Dai & Li, 2011

Biluscelis Dlabola, 1980  

Bolotheta Kramer, 1963  

Bonneyana Oman, 1949  

Brachylope Emeljanov, 1962  

Brachypterona Lindberg, 1954  

Branchana Li, 2011  

Brasilanus Linnavuori, 1959  

Brazosa Oman, 1938  

Cahya Linnavuori, 1959  

Caladonus Oman, 1949  

Caranavia Linnavuori, 1959  

Carelmapu Linnavuori, 1959  

Cenedaeus Distant, 1908  

Cerrillus Oman, 1938  

Cetexa Oman, 1949  

Chaparea Linnavuori, 1959  

Chimaerotettix Dietrich & Rakitov, 2002   placement nov. (previously unplaced in Deltocephalinae   ) Chroocacus Emeljanov, 1962  

Cocrassana Blocker & Larsen, 1991  

Colladonus Ball, 1936   placement nov. (previously placed in Platymetopiina)

Comayagua Linnavuori & DeLong, 1978  

Conoscelis Abdul-Nour, 1988  

Consepusa Linnavuori & DeLong, 1977  

Conversana DeLong, 1967  

Costamia DeLong, 1946  

Cozadanus DeLong & Harlan, 1968  

Crassana DeLong & Hershberger, 1947  

Cyanidius Emeljanov, 1964   placement nov. (previously placed in Platymetopiina)

Cyclevertex Li & Xing, 2011  

Dagama Distant, 1910  

Dampfiana DeLong & Hershberger, 1948  

Danbara Oman, 1949   placement nov. (previously unplaced in Deltocephalinae   )

Deltorhynchus DeLong, 1943  

Desertana DeLong & Martinson, 1973  

Devolana DeLong, 1967  

Didymotettix Yang, 1996  

Divitiacus Distant, 1918  

Dlabolaracus Remane, 1961  

Doleranus Ball, 1936  

Dolyobius Linnavuori, 1959  

Egenus Oman, 1938   placement nov. (transferred from Faltalini)

Elphnesopius Nast, 1984  

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   )

Eusora Oman, 1949  

Eutettix Van Duzee, 1892  

Excavanus DeLong, 1946  

Fistulatus Zhang, 1997   placement nov. (previously unplaced in Deltocephalinae   )

Fitchana Oman, 1949  

Floridonus Oman, 1949  

Fridonus Oman, 1949  

Garapita Oman, 1938  

Gunawardenea Fletcher & Moir, 2008  

Hamulotettix Dai & Zhang, 2011  

Hardya Edwards, 1922   placement nov. (previously placed in Platymetopiina)

Hegira Oman, 1938  

Houtbayana Linnavuori, 1961  

Huancabamba Linnavuori, 1959  

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)

Jaacunga Nielson, 1988  

Jakarellus Webb, 1980   placement nov. (previously placed in Platymetopiina)

Krameraxus Maldonado-Capriles, 1968   placement nov. (previously unplaced in Deltocephalinae   ) Lajolla Linnavuori, 1959  

Laminacutus Abdul-Nour, 2002  

Lamprotettix Ribaut, 1952   placement nov. (previously placed in Platymetopiina)

Lascumbresa Linnavuori & DeLong, 1979  

Latinocesa Kocak & Kemal, 2010   placement nov. (previously unplaced in Deltocephalinae   )

Laylatina Abdul-Nour, 1988  

Lineana Li & Xing, 2011  

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)

Melillaia Linnavuori, 1971  

Mesadorus Linnavuori, 1955  

Mesamia Ball, 1907  

Mexicananus DeLong, 1944  

Mimodorus Linnavuori, 1959  

Mocolinna McKamey, 2003  

Moskgha Deeming & Webb, 1982  

Nakaharanus Ishihara, 1953   placement nov. (previously placed in Platymetopiina)

Napo Linnavuori & DeLong, 1976  

Neocrassana Linnavuori, 1959  

Neodonus DeLong & Hershberger, 1948   placement nov. (previously placed in Platymetopiina) Neohegira Linnavuori & DeLong, 1978  

Neomacednus Xing & Li, 2011  

Neomesus Linnavuori, 1959  

Neoreticulum Dai, 2009  

Nesophryne Kirkaldy, 1907  

Nesothamnus Linnavuori, 1959   placement nov. (transferred from Scaphytopiini   )

Neurotettix Matsumura, 1914  

Nigridonus Oman, 1949  

Norvellina Ball, 1931   placement nov. (previously placed in Platymetopiina)

Nurenus Oman, 1949   placement nov. (previously placed in Platymetopiina)

Occiplanocephalus Evans, 1941  

Okaundua Linnavuori, 1969  

Ollarianus Ball, 1936  

Ophionotum Emeljanov, 1964   placement nov. (previously placed in Allygidiina)

Orientus DeLong, 1938  

Oxytettigella Metcalf, 1952  

Pachytettix Linnavuori, 1959  

Paracolladonus Nielson, 1988  

Paracrassana Nielson, 1988  

Paraganus Linnavuori, 1955  

Paralaca Lozada, 1998   placement nov. (previously unplaced in Deltocephalinae   )

Paramacrosteles Dai, Li & Chen, 2006  

Paranurenus Nielson, 1988  

Paratanus Young, 1957  

Perundanus Zanol, 1993   placement nov. (previously unplaced in Deltocephalinae   )

Phlepsanus Oman, 1949  

Phlepsobahita Linnavuori, 1959  

Phycotettix Haupt, 1929   placement nov. (transferred from Fieberiellini   )

Pingellus Evans, 1966  

Platymetopius Burmeister, 1838   placement nov. (previously placed in Platymetopiina)

Poliona Emeljanov, 1972   placement nov. (previously placed in Platymetopiina)

Protensus Zhang & Dai, 2001  

Pseudalaca Linnavuori, 1959   placement nov. (previously unplaced in Deltocephalinae   )

Pseudaligia Kramer & DeLong, 1968  

Pseudaraldus Bonfils, 1981  

Pseudocephalelus Linnavuori, 1969  

Pseudosubhimalus Ghauri, 1974  

Pseutettix DeLong, 1967  

Pygotettix Matsumura, 1940  

Renonus DeLong, 1959  

Retusanus DeLong, 1945  

Rinconada Linnavuori & DeLong, 1977  

Sanuca DeLong, 1980  

Sardius Ribaut, 1946  

Saudallygus Dlabola, 1979   placement nov. (previously unplaced in Deltocephalinae   )

Scaphetus Evans, 1966  

Scaphomonoides Li, 2011  

Scaphytoceps Dlabola, 1957   placement nov. (transferred from Scaphytopiini   )

Scenergates Emeljanov, 1972   placement nov. (previously placed in Platymetopiina)

Selachina Emeljanov, 1962  

Serratus Linnavuori, 1959  

Sincholata DeLong, 1982   placement nov. (previously unplaced in Deltocephalinae   )

Sobara Oman, 1949   placement nov. (previously unplaced in Deltocephalinae   )

Spinulana DeLong, 1967  

Stenometohardya Dlabola, 1981  

Stenomisella Evans, 1954  

Stoneana DeLong, 1943  

Streptopyx Linnavuori, 1958  

Stymphalella Evans, 1954   placement nov. (previously placed in Platymetopiina)

Tambocerus Zhang & Webb, 1996  

Tapetia Emeljanov, 1964   placement nov. (previously placed in Platymetopiina)

Tenuisanus DeLong, 1944  

Thanomahia Dlabola, 1987   placement nov. (previously unplaced in Deltocephalinae   )

Tingolix Linnavuori & DeLong, 1978  

Twiningia Ball, 1931   placement nov. (previously placed in Platymetopiina)

Usanus DeLong, 1947  

Watanabella Vilbaste, 1969  

Yungasia Linnavuori, 1959  

Zabrosa Oman, 1949  

Zercanus Dlabola, 1965  

Zilkaria de Menezes, 1974  


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Athysanini Van Duzee, 1892

Zahniser, James N. & Dietrich, Chris H. 2013

Phrynomorphini Kirkaldy, 1907

Thamnotettigini Distant, 1908


Euscelini Van Duzee, 1917


Anoterostemmini Haupt, 1929

Bobacellini Kusnezov, 1929


Platymetopiini Haupt, 1929

Colladonini Bliven, 1955

Cerrillini Linnavuori, 1975

Allygidiina Dmitriev, 2006