A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae) Author Zahniser, James N. C44D6E44-FA1C-4B29-B7BB-FAF5940CD225 Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816 S. Oak St., Champaign, IL 61820, United States of America & urn: lsid: zoobank. org: author: C 44 D 6 E 44 - FA 1 C- 4 B 29 - B 7 BB-FAF 5940 CD 225 & corresponding author e-mail: zahniser @ illinois. edu Author Dietrich, Chris H. 82FCB86C-54B4-456A-AE5E-D7847D271CB9 Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816 S. Oak St., Champaign, IL 61820, United States of America & urn: lsid: zoobank. org: author: 82 FCB 86 C- 54 B 4 - 456 A-AE 5 E-D 7847 D 271 CB 9 text European Journal of Taxonomy 2013 2013-05-29 45 1 211 journal article 22221 http://dx.doi.org/10.5852/ejt.2013.45 a66592b8-63a8-4c2d-9471-e58ddb2c0559 2118-9773 3822710 41B10E4D-7DAB-40CA-A8FE-4ECA078E04A3 Deltocephalinae Dallas, 1870 Note Because of the size and morphological diversity of this subfamily, nearly all of the characters listed below, which are the most common states in Deltocephalinae , occur in different states in some members of the subfamily. Diagnosis Small to large, usually wedge-shaped leafhoppers. Head with ocelli on the anterior margin close to eyes; frontoclypeus not inflated, without carinae; lateral frontal sutures extending to ocelli; antennal ledges reduced or absent; gena with a fine erect seta laterad of lateral frontal suture; gena large, mostly concealing proepisternum. Forewing macropterous to brachypterous; if macropterous, with apices overlapping at rest; with 3 anteapical cells; often with 1 or more crossveins between A1 and claval suture; inner apical cell tapered distally, not extended to wing apex. Profemur AM 1 seta distinct; row AV with short stout setae extending from base to 1/2 to 2/3 length of femur; intercalary row with numerous thin setae arranged in one row. Mesotrochanter with apical posteroventral stout seta. Metafemur macrosetal formula usually 2+2+1 with penultimate pair close-set. Metatibia usually anteriorly-posteriorly compressed, ventral side with median ridge. Male pygofer with basolateral oblique membranous cleft. Valve produced posteriorly, lateral margins short, articulated with pygofer laterally. Subgenital plates articulated with each other and with valve; triangular, more or less depressed or flattened; with dorsal slot or fold articulating with style. Connective Y -shaped or ‘linear’ (with anterior arms closely appressed), rarely T -shaped (with arms widely divergent); without anteromedial lobe or process. Style broadly bilobed basally; apophysis not elongate. First valvula convex to relatively straight; dorsal sculpturing pattern reaching or not reaching dorsal margin; sculpturing pattern strigate, concatenate, reticulate, imbricate, maculose, or granulose. Second valvula with basal fused section as long as or longer than distal paired blades; with or without median dorsal tooth; usually with small to large, regularly or irregularly shaped dorsoapical teeth on apical 1/3 or more; teeth sometimes restricted to apical 1/4, or absent. Included Tribes and Subtribes: Acinopterini Oman , 1943 Acostemmini Evans, 1972 Arrugadini Linnavuori, 1965 Athysanini Van Duzee, 1892 Bahitini Zahniser & Dietrich, 2013 Bonaspeiini Zahniser & Dietrich, 2013 Chiasmini Distant, 1908 Cicadulini Van Duzee, 1892 Cochlorhinini Oman , 1943 Deltocephalini Dallas, 1870 Dorycephalini Oman , 1943 Drabescini Ishihara, 1953 Drabescina Ishihara, 1953 Paraboloponina Ishihara, 1953 Drakensbergenini Linnavuori, 1979 Eupelicini Sahlberg, 1871 Eupelicina Sahlberg, 1871 Paradorydiina Evans, 1936 Faltalini Zahniser & Dietrich, 2010 Fieberiellini Wagner, 1951 Goniagnathini Wagner, 1951 Hecalini Distant, 1908 Glossocratina Dmitriev, 2002 Hecalina Distant, 1908 Hypacostemmini Linnavuori & Al-Ne’amy, 1983 Koebeliini Baker, 1897 Grypotina Haupt, 1929 Koebeliina Baker, 1897 Limotettigini Baker, 1915 Luheriini Linnavuori, 1959 Macrostelini Kirkaldy, 1906 Magnentiini Linnavuori, 1978 Mukariini Distant, 1908 Occinirvanini Evans, 1966 Opsiini Emeljanov, 1962 Achaeticina Emeljanov, 1962 Circuliferina Emeljanov, 1962 Eremophlepsiina Dmitriev, 2006 Opsiina Emeljanov, 1962 Paralimnini Distant, 1908 Aglenina Dmitriev, 2004 Paralimnina Distant, 1908 Pendarini Dmitriev, 2009 Penthimiini Kirschbaum, 1868 Phlepsiini Zahniser & Dietrich, 2013 Scaphoideini Oman , 1943 Scaphytopiini Oman, 1943 Selenocephalini Fieber, 1872 Adamina Linnavuori & Al-Ne’amy, 1983 Dwightlina McKamey, 2003 Ianeirina Linnavuori, 1978 Selenocephalina Fieber, 1872 Stegelytrini Baker, 1915 Stenometopiini Baker, 1923 Tetartostylini Wagner, 1951 Vartini Zahniser & Dietrich, 2013 Key to Tribes 1. Crown sharply produced anteriorly ( Fig. 48A ); head narrower than pronotum; gena not incised laterally below eye ( Fig. 48C ), visible behind eye in dorsal view ( Fig. 48A ); frontoclypeus elongate ( Fig. 48C ); forewing with numerous reflexed costal veins ( Fig. 48B ); connective a transverse bar with two lateral arms directed obliquely laterad, stem absent ( Fig. 48G ); usually with pair of long processes articulated with connective( Fig.48G,J ); Nearctic,Neotropical ...................... Scaphytopiini – Without the above combination of characters; crown variable; head width variable; gena incised laterally below eye (exceptions: some Acostemmini, Opsiini , Stegelytrini , Scaphoideini, and Vartini); forewing with or without reflexed costal veins; connective not as above; usually without long processes articulated with connective (exceptions: Adamina, some Scaphoideini).................2 2. Connective T -shaped, bar-shaped, or platelike or, if Y-shaped, with arms forming broadly obtuse angle (e.g., Figs 8G , 25G , 33G , 53G ); style more or less linear, not broadly bilobed at base (e.g., Figs 8G , 25G , 33G , 53G )..................................................................................................................3 – Connective with anterior arms somewhat or not at all divergent, connective Y - or U -shaped (e.g., Figs 11G , 52G ) or ‘linear’ with anterior arms closely appressed (e.g., Fig. 17G ); style broadly bilobed at base (e.g., Fig. 11G ).........................................................................................................................7 3. Head much narrower than pronotum; profemur row AM with numerous preapical macrosetae; metatibia dorsal surface with supernumerary macrosetae between rows PD and AD, especially basally; pro- and mesotibia row PD and often row AD with numerous (> 6) macrosetae; forewing appendix often large, extending around apex;Palearctic, Oriental ..................................... Stegelytrini – Head slightly narrower to wider than pronotum; profemur row AM with not more than 2 preapical macrosetae; metatibia dorsal surface without supernumerary macrosetae; pro- and mesotibia with <6 macrosetae; forewing appendix reduced or absent...........................................4 4. Anterior margin of head usually with a single distinct transverse carina between ocelli; epistomal suture keeled ( Fig. 9C ) or evident from change in texture on frontoclypeus; subgenital plates completely or partially fused to each other and/or to valve (if completely articulated then pronotum produced dorsally, with longitudinal crest, Ikelibeloha ); Madagascar ..................Acostemmini (part) – Anterior margin of head shagreen, glabrous, or with several irregular striae or carinae, without single distinct carina between ocelli; epistomal suture not keeled; subgenital plates not fused, articulated with valve..........................................................................................................................................5 5. Forewing acuminate apically ( Fig. 8B ); head narrower than pronotum ( Fig. 8A ) (exception: Cariancha ); connective bar-shaped or plate-like, without well-defined stem ( Fig. 8G ); Nearctic, Neotropical ...................................................................................................................Acinopterini – Forewing rounded apically; head as wide as or wider than pronotum; connective with welldeveloped anterior arms posterior stem...........................................................................................6 6. Connective with median, darkly sclerotized Y - or T -shaped ridge following stem and arms of connective ( Fig. 25G ); forewing central anteapical cell not strongly constricted medially; female pregenital sternite VIII not concealed and folded beneath sternite VII,movably articulated with sternite VII( Fig.25J );Palearctic, Nearctic ( Fieberiella florii introduced in NorthAmerica).......... Fieberiellini – Connective evenly sclerotized throughout ( Fig. 33G ); forewing central anteapical cell constricted medially ( Fig. 33B ); female pregenital sternite VIII folded beneath and concealed by sternite VII, sternites not articulated with each other; Neotropical ............................................... Luheriini 7. Subgenital plates fused to each other ( Figs 9F , 26F ); valve straplike, fused to pygofer laterally ( Fig. 9D, F ) OR not readily apparent (fused to plates); connective fused with aedeagus ( Figs 9H , 26H ).....................................................................................................................................................8 – Subgenital plates usually free; valve present, articulated with pygofer laterally and usually articulated with subgenital plates; connective articulated with or fused to aedeagus; rarely with plates fused and connective fused with aedeagus, and if so then body slender or relatively small in size, clypellus tapering apically ( Deltocephalini : Miradeltaphus , Yuanamia )............................9 8. Anterior margin of head with a distinct transverse carina between ocelli; forewings tectiform, not overlapping at rest ( Fig. 9A ); forewing appendix reduced or absent (9B); valve strap-like, fused to pygofer laterally ( Fig. 9D, F ); Afrotropical (especially diverse in Madagascar ), Oriental ( India ) ..................................................................................................................Acostemmini (part) – Anterior margin of head without transverse carina; forewing, if macropterous, with apices overlapping at rest ( Fig. 26A ) and appendix extending around wing apex ( Fig. 25A ); valve not readily apparent, fused to subgenital plates; Palearctic, Afrotropical, Oriental , and Australian .......................................................................................................................... Goniagnathini 9. Head strongly produced, elongate or spatulate ( Figs 18A , 21A , 22A , 23A ); gena acutely incised laterally below eye ( Figs 18C , 21C , 22C , 23C ); profemur intercalary row absent or with few scattered setae; metafemur macrosetal formula 2+0 or 2+1; metatarsomere I shorter than metatarsomeres II and III combined, expanded apically; metatibia straight throughout length and distinctly bent distally in dorsal view...................................................10 – Without above combination of characters; head produced or not; gena obtusely incised (e.g., Fig. 11C ) or not incised laterally below eye; profemur intercalary row more or less welldelimited and with several to many long fine setae; metafemur macrosetal formula usually 2+2+1; metatarsomere I longer than metatarsomeres II and III combined, usually not expanded apically; metatibia slightly arcuate throughout its length in dorsal view...................................15 10. Aedeagus movably hinged between base and shaft ( Fig. 14H ); connective with anterior arms closely appressed ( Fig. 14G ); first valvula dorsal sculpturing maculose to granulose, submarginal; second valvula teeth obtusely triangular, serrate, uniformly shaped; Old World .... ............................................................................... Chiasmini (part, Listrophora , some Chiasmus ) – Aedeagus without hinge; connective with anterior arms separate ( Y -shaped); first valvula dorsal sculpturing pattern variable; second valvula teeth variable.............................................11 11. Ocelli on crown.............................................................................................................................12 – Ocelli on anterior margin of head..................................................................................................13 12. Macropterous; crown encroaching onto eyes ( Fig. 22 A-C); frontoclypeus with median longitudinal carina ( Fig. 22C ); Palearctic ........................................Eupelicini (part, Eupelicina) – Brachypterous ( Fig. 21 A-B); crown not encroaching onto eyes; frontoclypeus and clypellus tumid, without longitudinal carina ( Fig. 21 B-C); Afrotropical ( South Africa , Lesotho ) ............... ...........................................................................................................................Drakensbergenini 13. Macropterous; hindwing submarginal vein evanescent apically; valve fused to pygofer ( Fig. 23D ); male pygofer without basolateral cleft ( Fig. 23D ); style base connected or articulated only laterally with apical part ( Fig. 23J ); Old World ....................Eupelicini (part, Paradorydiina) – Brachypterous or macropterous, hindwing submarginal vein complete; valve articulated to pygofer; male pygofer with basolateral cleft; style base broadly fused with apical part (e.g., Fig. 27G )........14 14. Aedeagus with terminal processes (e.g., Fig. 27 H-I); first and second valvulae humpbacked dorsally; first valvula with distinctly delimited ventroapical sculptured patch; second valvula without teeth; Nearctic .................................................. Hecalini (part, Attenuipyga , Neoslossonia ) – Aedeagus without terminal processes ( Fig. 18 H-I); first and second valvulae not humpbacked dorsally; first valvula without distinctly delimited ventroapical sculptured patch; second valvula with distinct teeth; Palearctic .......................................................................................Dorycephalini 15. Anterior margin of pronotum exceeding eyes in dorsal view ( Fig. 35A ); integument black, shiny, punctate ( Fig. 35 A-B); Afrotropical, Oriental .................................................................Magnentiini – Anterior margin of pronotum not exceeding eyes in dorsal view; integument rarely completely black and punctate....................................................................................................................................16 16. Antennal ledge strong, forming distinct ledge, sometimes seemingly continuous with anterior margin of head ( Figs 19C , 45C ); protibia dorsal side flat, margins carinate; forewing appendix large, extending around wing apex................................................................................................17 – Antennal ledge absent or represented by weak carina not forming ledge; protibia rounded dorsally; forewing appendix usually smaller, not extending around wing apex...........................................18 17. Large, robust leafhoppers, not squat or dorsally rounded or ventrally flattened ( Fig. 19 A-B); ocelli on anterior margin of head (19C); frontoclypeus texture longitudinally striate or rugose ( Fig. 19C ); Palearctic, Ethiopian, Oriental , Australian .....................Drabescini (part, Drabescina) – Robust and squat, dorsally rounded, ventrally flattened leafhoppers ( Fig. 45 A-B); ocelli on crown distant from eyes ( Fig. 45B ); frontoclypeus texture glabrous or finely shagreen; Cosmopolitan .. ...............................................................................................................................Penthimiini 18. Crown and face strongly rugose ( Fig. 10 A-C); body greater than 7 mm in length; Neotropical ............. ............................................................................................................................................Arrugadini – Crown and face shagreen, glabrous, or striate, not rugose, or if rugose then body less than 7 mm in length..............................................................................................................................................19 19. Aedeagus bifurcate, with 2 gonopores............................................................................................20 – Aedeagus with one shaft and gonopore............................................................................................21 20. Face ( Fig. 36B ) strongly depressed, nearly horizontal, sometimes concave posteroventrally in profile, sometimes inflated anterodorsally; anterior margin of head often with one or more carinae or sharp ridges; antennal bases near anterodorsal (upper) corners of eyes ( Fig. 36C ); stem of connective short; Afrotropical, Oriental ....................................................Mukariini (part) – Face oblique, not strongly depressed, not concave in profile; anterior margin of head without carinae (foliaceous in Chlidochrus ); antennal bases near middle or posteroventral (lower) corners of eyes; stem of connective longer; Cosmopolitan .................................................. Opsiini 21. Head often somewhat produced ( Fig. 36A ); face flattened or frontoclypeus inflated medially or anteriorly, lying in horizontal plane in lateral view ( Fig. 36 B-C); antennal bases near anterodorsal (upper) corners of eyes ( Fig. 36C ); eyes large ( Fig. 36C ); antennae often long, longer than width of head; profemur row AV setae absent; outer anteapical cell often confluent with or nearly touching costal vein ( Fig. 36B ); Oriental , feeding on bamboo ..........................Mukariini (part) – Without above combination of characters; head produced or not; face flattened or not, not distinctly inflated anteriorly, lying in horizontal plane in lateral view or not; size of eyes and antennae variable; profemur row AV setae present or absent; outer anteapical cell usually not touching or confluent with costal vein......................................................................................22 22. Connective Y -shaped and fused to aedeagus ( Fig. 16 G-H); male pygofer with one or more pairs of teeth, spines, or processes ( Fig. 16 D-E); male pygofer side excavated medially, scoop-like ( Fig. 16E ); male segment X long, sclerotized laterally, sometimes sclerotized ventrally and/or dorsally ( Fig. 16 D-E); second valvulae with a median dorsal tooth; Nearctic, Neotropical ( Cochlorhinus pluto introduced in Chile ) .........................Cochlorhinini – Connective shape variable, but not Y -shaped and fused to aedeagus; male pygofer with or without processes; male pygofer side variable; male segment X variable; second valvulae with or without dorsal tooth on shaft......................................................................................................23 23. Metatarsomere I with platellae on plantar surface; ocelli distant from eyes, at least 4 x own diameter from adjacent eye (e.g., Fig. 30C ).............................................................................24 – Metatarsomere I without platellae on plantar surface; ocelli usually close to eyes (exceptions: some Athysanini , Chiasmini, and Faltalini)..................................................................................26 24. Head not produced anteriorly ( Fig. 30A ); crown rounded to face ( Fig. 30B ) or anterior margin of head carinate, not foliaceous; Palearctic, Oriental , Nearctic ( Grypotes puncticollis introduced to northeastern North America) ..............................Koebeliini, in part (Grypotina) – Head strongly produced, foliaceous anteriorly ( Figs 31 A-C, 37A-C).....................................25 25. Ocelli on face ( Fig. 31C ); antennae not very long; forewing veins pustulate, appendix absent ( Fig. 31 A-B); Nearctic (western North America) ..........................................Koebeliini (part, Koebeliina) – Ocelli on crown ( Fig. 36A 1-A 2 ); antennae very long; forewing veins not pustulate, appendix present; Australian ( Western Australia ) .......................................................................Occinirvanini 26. Color often greenish or bluish with red or orange longitudinal stripes ( Fig. 56 A-B); crown produced, pointed ( Fig. 56A ); gena not incised laterally below eye, visible behind the eye in dorsal view ( Fig. 56 B-C); frontoclypeus elongate ( Fig. 56C ); profemur intercalary row setae thick and extending to or basad of middle of profemur; forewing macropterous, truncate apically (56B); apodemes of male sternite II long, subrectangular, flared apically, directed posterolaterally ( Fig. 56J ); connective with anterior arms closely appressed ( Fig. 56G ); male segment X tube-like, protruding from pygofer, often well-sclerotized ( Fig. 56 D-E, K); Afrotropical, Oriental ....................... Vartini – Color variable; crown produced or not; gena usually incised laterally below eye, not visible behind eye in dorsal view (exception: some Scaphoideini); frontoclypeus variable, usually not very elongate; profemur intercalary row setae fine;if macropterous, forewing usually not truncate apically; apodemes of male sternite II not as above; connective variable; male segment X variable.......................27 27. Clypellus parallel-sided or tapering apically (e.g., Fig. 17C ); connective linear, triangular, or racquetlike in shape, with anterior arms closely appressed anteriorly (e.g., Figs 17G , 42G )......................28 – Clypellus variable; connective usually Y -shaped (e.g., 54G) or U -shaped (e.g., Fig. 52G ), with anterior arms separated (exception: some Athysanini , Scaphoideini)...............................................32 28. Subgenital plates fused with valve and apex of plate with sharply pointed process ( Fig. 55F ); connective articulated posteriorly with a pair of long paraphyses ( Fig. 55G ); female sternite VII ovate, sometimes very long; Palearctic, Afrotropical .......................... Tetartostylini – Subgenital plates usually articulated with valve, apices without sharply pointed processes; paraphyses present or absent; female sternite VII not ovate, not very long................................29 29. Connective fused with aedeagus ( Fig. 17G )................................................................................30 – Connective articulated with aedeagus ( Fig. 42G )......................................................................31 30. Body elongate, somewhat depressed; anterior margin of head transversely striate or carinate; first valvula dorsal sculpturing pattern maculose to granulose and distinctly submarginal, with distinctly delimited ventroapical sculptured area; second valvula without dorsal teeth; ovipositor protruding well beyond pygofer apex; Nearctic ( Mexico ), Neotropical ..............Faltalini (part, Tenucephalus ) – Body not elongate or depressed; crown rounded to face, anterior margin shagreen; first valvula sculpturing pattern imbricate (scale-like) and reaching dorsal margin, without distinct apicoventral sculptured area; second valvula with irregular teeth; ovipositor not protruding far beyond pygofer; Cosmopolitan ............................................................................................... Deltocephalini 31. Aedeagus movably hinged between base and shaft ( Fig. 14H ) (exceptions: Baileyus , some Gurawa , Paraphrodes ); ovipositor protruding beyond pygofer apex (exceptions: Gurawa , Baileyus , Omaranus , Paraphrodes ); first valvula dorsal sculpturing pattern maculose to granulose, submarginal; second valvula teeth uniformly shaped, obtusely triangular, serrated; Cosmopolitan ........................................................................................................... Chiasmini (part) – Aedeagus entire, not movably hinged; ovipositor not protruding far beyond pygofer; first valvula dorsal sculpturing pattern usually imbricate, reaching dorsal margin; second valvula teeth not uniformly obtuse and triangular, serrated or not; Cosmopolitan ................................ Paralimnini 32. Macropterous, forewing with 2 anteapical cells ( Fig. 34B ); usually small, slender leafhoppers ( Fig. 34 A-B); Cosmopolitan .......................................................................................... Macrostelini – Macropterous to brachypterous, if macropterous then usually with 3 anteapical cells; if with 2 anteapical cells, then larger, more robust leafhoppers...........................................33 33. Small to medium sized, slender, elongate leafhoppers ( Fig. 15A ); usually yellow, ochraceous, or pale green, often with paired black or brown markings or spots on head ( Fig. 15 A-C); subgenital plate often with row of macrosetae distant from lateral margin, with numerous long fine setae laterally ( Fig. 15F ); male pygofer incised dorsally nearly to base ( Fig. 15 D-E); male segment X elongate and sclerotized dorsally ( Fig. 15 D-E) (not elongate in Knullana DeLong ); Palearctic, Nearctic, Neotropical ....................................................................................... Cicadulini – Size and color variable; subgenital plate macrosetae variable, with or without long fine setae; male pygofer dorsal incision variable; male segment X usually not elongate and strongly sclerotized.......................................................................................................................................34 34. Macropterous to brachypterous, if macropterous then with appendix large, extending around forewing apex; aedeagus movably hinged between base and shaft ( Fig. 14H ) (exceptions: Leofa , Picchusteles ); ovipositor extending well beyond pygofer apex (exception: Paraphrodes ); first valvula dorsal sculpturing pattern granulose to maculose, submarginal; second valvula teeth uniformly shaped, obtusely triangular and serrated; Cosmopolitan ................. Chiasmini (part) – Macropterous to brachypterous, if macropterous, with appendix not large or extending around forewing apex; aedeagus not movably hinged; ovipositor length variable; first valvula dorsal sculpturing variable; second valvula teeth not obtusely triangular and serrated................................35 35. Crown narrow, width between eyes less or equal to median width of eye ( Fig. 54A ); crown completely shagreen to base; pronotum lateral margin not carinate; male pygofer declivous, directed posteroventrally, with few macrosetae, basolateral cleft absent or not membranous ( Fig. 54D ); ovipositor extending far beyond pygofer; first valvula with distinctly delimited ventroapical sculptured area; second valvula without teeth; Cosmopolitan ..................Stenometopiini – Crown not very narrow; discal portion of crown texture usually glabrous or striate (rarely shagreen in some Athysanini [e.g., Anoterostemma ], some Pendarini [ Chlorotettix and Copididonus ], and Limotettigini); pronotum lateral margin usually carinate; male pygofer shape variable, basolateral cleft present, membranous; ovipositor not protruding far beyond pygofer apex; first valvula without distinctly delimited ventroapical sculpturing; second valvula with teeth...................................36 36. Dorsoventrally depressed or flattened leafhoppers (e.g., Fig. 24B ); head strongly or sometimes weakly produced; crown broad, more or less parabolically shaped ( Figs 24A , 28A ); crown to face transition acutely angled; anterior margin sometimes foliaceous ( Figs 24B , 28B ); face lying slightly obliquely to horizontal plane ( Figs 24B , 28B ); lateral margin of pronotum long, nearly as long or longer than least width of eye ( Figs 24A , 28A ); first valvula dorsal sculpturing maculose to granulose, distinctly submarginal; second valvula without dorsal teeth (exception: Glossocratus )...37 – Not particularly dorsoventrally depressed or flattened; head not strongly produced; crown broad or not, rarely parabolically shaped; crown to face transition less acute; face usually lying in a more upright plane; lateral margin of pronotum usually shorter than least width of eye; first valvula dorsal sculpturing variable; second valvulae with teeth.............................38 37. Ocelli close to eyes, closer to eyes than lateral frontal sutures ( Fig. 28C ); male apodemes of abdominal sternite I long, narrow, as long as apodemes of sternite II ( Fig. 28J ); male apodemes of abdominal sternite II large, spatulate ( Fig. 28J ); first and second valvulae humpbacked dorsally; second valvula concave ventrally; Cosmopolitan .................................. Hecalini (part) – Ocelli relatively distant from eyes, usually 3-4 x their diameter from ocellus or more (exception: some Faltala , Bonamus ), sometimes absent or obscure; male apodemes of abdominal sternite I short or undeveloped; male apodemes of abdominal sternite II short or undeveloped; first and second valvulae not strongly humpbacked dorsally; second valvula not concave ventrally; Nearctic, Neotropical .......................................Faltalini (part) 38. Body usually broad and robust ( Fig. 13 A-B); forewings brachypterous to submacropterous ( Fig. 13 A-B) (exception: Renosteria ); hindwings when present narrowed anteriorly-posteriorly when outstretched or otherwise reduced in size ( Fig. 13 J-K) (exception: Renosteria ); connective U - or sometimes Y - shaped with stem often very short or absent ( Fig. 13G ); male segment X broad and sclerotized; ovipositor usually distinctly protruding beyond pygofer apex ( Fig. 13B ); Afrotropical ( South Africa ) ...........................................................Bonaspeiini – Body shape variable; forewings usually macropterous; hindwings not narrowed anteriorlyposteriorly when outstretched; connective variable; male segment X variable; ovipositor usually not protruding far beyond pygofer apex.........................................................................................39 39. Anterior margin of head with 2 or more distinct carinae or with several distinct transverse striations (e.g., Fig. 50C ); antennal ledge with oblique carina (e.g., Fig. 49C ); medium to large-sized leafhoppers......................................................................................................................................40 – Anterior margin of head without carinae or distinct transverse striations (rarely with single transverse carina), texture shagreen or glabrous, crown rounded to face; antennal ledge absent or represented by weak carina; size variable...............................................................44 40. Antenna long, at least as long as half length of body; antennal pits near upper (anterodorsal) corners of eyes, large and encroaching onto frontoclypeus ( Fig. 20C ) (exception: Nirvanguina ); profemur row AV setae reduced or absent, usually with fine setae; Afrotropical, Oriental , Australian ...................................................................................Drabescini (part, Paraboloponina) – Antenna not very long; antennal pits near lower corners of eyes (exception: Selenocephalini [Dwightlina]), not very large or encroaching onto frontoclypeus); profemur row AV with distinct stout setae.......................................................................................................................................41 41. Profemur row AM often with 1 or more extra macrosetae basad of AM 1; pygofer usually with macrosetae reduced in number or absent (e.g., Fig. 52D ) (exception: Adamina); connective stem usually very short or absent, connective U -shaped ( Fig. 52G ); male segment X usually broad and well-sclerotized dorsally; Afrotropical, Palearctic, Oriental .......................... Selenocephalini – Profemur row AM usually without extra macrosetae basad of AM 1; connective stem present, relatively long, connective Y -shaped; male segment X variable, usually not very broad or large.............42 42. Head distinctly narrower than pronotum ( Fig. 46A ); antennal ledges carinate ( Fig. 46C ); lorum long, relatively large ( Fig. 46C ); clypellus long, widening apically ( Fig. 46C ); pronotum lateral margin long, longer than half basal width of eye; forewing often with brown irrorate markings; profemur row AM sometimes with one or more setae basad of AM 1; nymph with pygofer short, with 6 rows of abdominal macrosetae ............................Phlepsiini (part) – Head usually as wide as or wider than pronotum; antennal ledges variable; lorum not very large; clypellus usually not particularly long; pronotum lateral margin short, usually less than half basal width of eye; forewing coloration variable; profemur row AM only AM 1 seta; nymph variable...........................................................................................................................43 43. Medium to large sized, robust leafhoppers ( Fig. 12 A-B); often predominantly brown in color ( Fig. 12 A-B); crown usually short, broad ( Fig. 12A ); often with anterior margin of head distinctly upturned, raised, or with several transverse carinae ( Fig. 12 B-C); clypellus widening apically, parallelmargined, or tapering apically ( Fig. 12C ), sometimes distinctly tumid ( Fig. 12C ); forewing often with strongly pigmented reflexed costal veins ( Fig. 12B ); pygofer often with processes ( Fig. 12D ); aedeagus often with apical processes ( Fig. 12 H-I); Nearctic, Neotropical ..................Bahitini (part) – Without the above combination of characters; Cosmopolitan ......................... Athysanini (part) 44. Aedeagus with single, unpaired sclerotized apophysis (= ‘dorsal connective’) projecting dorsally from atrium, membranously connected to segment X ( Fig. 32 H-I); male pygofer dorsal margin thickened, declivous, and often ending in ventrally directed curved hook or spine ( Fig. 32D ); Palearctic, Oriental , Australian, Nearctic, Neotropical ............................................Limotettigini – Without the above combination of characters..........................................................................45 45. Large, yellow to greenish species ( Fig. 29 A-B); forewing veins greenish ( Fig. 29 A-B); forewing venation highly reticulated apically ( Fig. 29B ); Afrotropical ( South Africa ) ........Hypacostemmini – Color variable; forewing venation rarely highly reticulate, but if so, reticulate portion not restricted to forewing apex............................................................................................................46 46. Ocelli on face below anterior margin of head ( Fig. 51C ), distant from eyes;Afrotropical ................... ............................................................................................. Selenocephalini (part, Ianeirini: Ianeira ) – Ocelli on anterior margin of head, close to eyes......................................................................47 47. Head distinctly narrower than pronotum ( Fig. 46A ); antennal ledge carinate ( Fig. 46C ); lorum long, relatively large ( Fig. 46C ); clypellus long, widening apically ( Fig. 46C ); pronotum lateral margin longer than half basal width of eye ( Fig. 46A ); forewing often with brown irrorate markings ( Fig. 46 A-B); profemur row AM sometimes with one or more setae basad of AM 1; nymph with pygofer short, with 6 rows of abdominal macrosetae ............................Phlepsiini (part) – Head width variable; antennal ledge variable; lorum not very large; clypellus usually not particularly long; pronotum lateral margin usually less than half basal width of eye; forewing coloration variable; profemur row AM without one or more macrosetae basad of AM 1; nymph variable.......48 48. Medium to large sized, robust leafhoppers (12A-B); often predominantly brown in color (12A- B); crown usually short, broad (12A); often with anterior margin of head distinctly upturned, raised (12B-C); clypellus widening apically, parallel-margined, or tapering apically (12C), sometimes distinctly tumid (12C); forewing often with strongly pigmented reflexed costal veins (13B); pygofer often with processes (12D); aedeagus often with apical processes (12H- I); Nearctic, Neotropical ....................................................................................Bahitini (part) – Size variable; color variable; anterior margin of head usually not upturned, raised; clypellus usually widening apically or parallel-sided, not tumid; forewing with or without reflexed costal veins; pygofer with or without processes; aedeagus with or without apical processes.....................49 49. (Note: the following characters are not present in all included genera, but some combination of most of these characters is present in all): Head narrower than pronotum, produced anteriorly ( Fig. 47A ); frontoclypeus long, narrow ( Fig. 47C ); antennae long, longer than width of head; body slender ( Fig. 47A ); head and wings often with brown, orange, ochraceous, or ivory markings ( Fig. 47 A-B); forewing with one or more darkly pigmented reflexed costal veins ( Fig. 47B ); profemur row AV setae absent or reduced (not stout); metatibia row PD macrosetae long, as long as or longer than 0.5x length of protibia; male or female pygofer with dense tufts of long fine or regular setae ( Fig. 47D ); subgenital plate apex membranous or long, digitate, somewhat membranous or weakly sclerotized; subgenital plates with long fine setae laterally ( Fig. 47F ); basal processes of aedeagus or connective often present, connected or articulated to base of aedeagus or apex of connective stem ( Fig. 47J, M ); Cosmopolitan ................Scaphoideini – Without some combination of above characters.............................................................................50 50. Ground color yellowish or greenish OR forewing with numerous brown, tawny, or reddishbrown ramose or reticulate pigment lines, or connected dots forming such lines ( Fig. 44 A-B); head as wide as or wider than pronotum; male pygofer often with a spine or process arising from dorsal side; nymph crown-face transition carinate; face of nymph with longitudinal carina in upper part; nymph pygofer elongate; Nearctic, Neotropical .....................Pendarini – Without the above combination of characters; Cosmopolitan ...................... Athysanini (part)