Athysanini Van Duzee, 1892
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|Athysanini Van Duzee, 1892|
Fig. 11 View Fig
Type genus: Athysanus Burmeister, 1838 .
= Phrynomorphini Kirkaldy, 1907.
= Thamnotettigini Distant, 1908.
= 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.
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).
Oman (1949), Linnavuori (1959), Linnavuori & DeLong (1978b), Cwikla & Blocker (1981), Anufriev & Emeljanov (1988), Emeljanov (1999), Dmitriev (2006b).
Allotapes Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Anaemotettix Korolevskaya, 1980 placement nov. (previously placed in Platymetopiina)
Asthenotettix Korolevskaya, 1980 placement nov. (previously placed in Platymetopiina)
Bicoloratum Dai & Li, 2011
Colladonus Ball, 1936 placement nov. (previously placed in Platymetopiina)
Cyanidius Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Egenus Oman, 1938 placement nov. (transferred from Faltalini)
Ephelodes Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Eusama Oman, 1949 placement nov. (previously placed in Platymetopiina)
Hardya Edwards, 1922 placement nov. (previously placed in Platymetopiina)
Inghamia Evans, 1966 placement nov. (previously placed in Platymetopiina)
Jakarellus Webb, 1980 placement nov. (previously placed in Platymetopiina)
Lamprotettix Ribaut, 1952 placement nov. (previously placed in Platymetopiina)
Malasiella Evans, 1954 placement nov. (previously placed in Platymetopiina)
Nakaharanus Ishihara, 1953 placement nov. (previously placed in Platymetopiina)
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)
Platymetopius Burmeister, 1838 placement nov. (previously placed in Platymetopiina)
Poliona Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Scenergates Emeljanov, 1972 placement nov. (previously placed in Platymetopiina)
Stymphalella Evans, 1954 placement nov. (previously placed in Platymetopiina)
Tapetia Emeljanov, 1964 placement nov. (previously placed in Platymetopiina)
Twiningia Ball, 1931 placement nov. (previously placed in Platymetopiina)
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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|