Penthimiini Kirschbaum, 1868 Fig. 45 Type genus: Penthimia Germar, 1821. = Thaumatoscopini Baker, 1923. Diagnosis Penthimiini are medium sized to large, squat, robust, often black or brown leafhoppers; often with ventral part of face and/or entire ventral side flattened and dorsal side convex. They can be identified by the ocelli on the crown and often distant from the eyes, strong antennal ledge, dorsally flattened and carinate protibia, and forewing with appendix large and extending around the wing apex. Description HEAD. Head subequal to, wider than, or distinctly narrower pronotum. Discal portion of crown glabrous with radial or longitudinal striae. Anterior margin of head with numerous transverse striations or carinae, or foliaceous. Head and pronotum usually sloping anteriorly. Face short, broad. Frontoclypeus tumid or not; texture striate or glabrous. Clypellus widening apically; apex following or slightly surpassing normal curve of gena. Lorum subequal to or wider than clypellus near base. Antennal bases near upper or anterodorsal corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledges strongly developed (with a distinct ledge); sometimes forming lateral part of anterior margin of head. Ocelli present; distant from eyes; on crown. THORAX. Pronotum lateral margin carinate; lateral margin shorter or longer than basal width of eye. WINGS. Forewing macropterous or submacropterous; sometimes coriaceous, setose, punctate, or tuberculate; appendix large and extending around wing apex or rarely restricted to anal margin; with 3 anteapical cells; veins not raised; without reflexed costal veins; A1-A2 crossvein absent or present; apical venation not highly reticulate. LEGS. Profemur 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 usually with thin, hair-like setae or without setae, more rarely with short, stout setae or with relatively long macrosetae. Protibia dorsal surface flat, AD and PD margins sharply carinate or at ~90º (not rounded). Metafemur apex macrosetae 2+2+1. Metatarsomere I not expanded apically, plantar setae simple, tapered. MALE GENITALIA. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows. Subgenital plates free from each other; articulated with valve; without macrosetae or macrosetae scattered, irregularly arranged. Style linear, median anterior lobe not pronounced or 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; articulated with aedeagus. FEMALE GENITALIA. Pygofer with numerous macrosetae. Ovipositor not protruding or rarely protruding (e.g., Penthimidia) far beyond pygofer apex. First valvula convex; dorsal sculpturing pattern strigate, concatenate, or reticulate; sculpturing reaching dorsal margin; without distinctly delimited ventroapical sculpturing. Second valvula abruptly broadened medially or subapically or broad, gradually tapered; with or without dorsal median tooth; teeth on apical 1/3 or more; teeth large, regularly shaped or small, regularly or irregularly shaped. Geography and ecology Distribution: cosmopolitan, poorly represented in North America. Penthimiini are collected on trees, shrubs, and herbaceous vegetation. Remarks Penthimiini contains46 genera and 203species.It is a relatively early diverging lineage of Deltocephalinae but has the typical derived deltocephaline characters of the male genitalia (pygofer with basolateral membranous cleft, Y -shaped connective, and style broadly bilobed basally). The phylogenetic analyses included 4 exemplars of the tribe as circumscribed here (Penthimia, Penthimidia, Penthimiola, and Jafar included in analyses; Citorus Stål, 1866 is removed from the tribe based on these analyses). Parsimony analyses resolved the former 3 exemplars as monophyletic and sister to Magnentius Singh-Pruthi, 1930, while ML and Bayesian analyses resolved Penthimidia and Penthimiola together and Penthimia in an uncertain position elsewhere in the tree. It is unclear why Penthimia does not always group with other penthimiines in phylogenetic analyses, but based on the morphological evidence supporting Penthimiini as a taxon, we suspect that the phylogenetic placement of Penthimia in the ML and Bayesian analyses is artifactual. Jafar was also included for the first time and grouped with one or two undescribed African genera, somewhat distant from other Penthimiini but with no branch support. Because its phylogenetic position is uncertain, and because it possesses several characters diagnostic for the tribe (large and robust body, ocelli on crown, inflated frontoclypeus, strong antennal ledges, and dorsally flattened and bicarinate protibiae), it is retained in the tribe pending further study. Citorus, which was previously included in Penthimiini based on its body shape and large forewing appendix, was found with strong branch support to be more closely related to Selenocephalini (Dwightla and Selenocephalus). Based on the phylogenetic results and on some morphological characters that differ from Penthimiini (anterior margin of the head with distinct sharp carinae, ocelli on anterior margin of head and close to eyes, antennal ledge not very strong or ledge-like) it is transferred to Selenocephalina here. The squat, robust body form and large appendix are similar to Penthimiini but these are apparently convergent characters in this case. Selected references Evans (1966, 1972), Linnavuori (1977). Included genera Alopenthimia Evans, 1972 Amberbakia Distant, 1912 Chanohirata Hayashi & Machida, 1996 Chinaella Evans, 1935 Ectopiocephalus Kirkaldy, 1906 Eovulturnops Evans, 1947 Eupenthimia Evans, 1972 Eusallya Evans, 1972 Foroa Linnavuori, 1977 Gressittella Evans, 1972 Haranga Distant, 1908 Irenaella Linnavuori, 1977 Jafar Kirkaldy, 1903 Kronos Distant, 1917 Ledroides Dammerman, 1910 Macutella Evans, 1972 Malichus Distant, 1918 Megalopenthimia Evans, 1954 Musosa Linnavuori, 1977 Neodartellus Evans, 1955 Neodartus Melichar, 1903 Neopenthimia Evans, 1972 Neovulturnus Evans, 1937 Nielsoniella Linnavuori, 1977 Nortoides Evans, 1972 Nubelella Evans, 1972 Nubelloides Evans, 1972 Osella Evans, 1972 Penthimia Germar, 1821 Penthimidia Haglund, 1899 Penthimiella Evans, 1972 Penthimiola Linnavuori, 1959 Penthimiopsis Evans, 1972 Pentria Evans, 1972 Piorella Evans, 1972 Platyscopus Evans, 1941 Reticuluma Cheng & Li, 2005 Sidelloides Evans, 1972 Tambila Distant, 1908 Thaumatopoides Evans, 1947 Thaumatoscopus Kirkaldy, 1906 Tolasella Evans, 1972 Tomaloides Evans, 1972 Uzelina Melichar, 1903 Vertigella Evans, 1972 Vulturnus Kirkaldy, 1906

Published as part of Zahniser, James N. & Dietrich, Chris H., 2013, A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae), pp. 1-211 in European Journal of Taxonomy 45 on pages 142-145, DOI: 10.5852/ejt.2013.45, http://zenodo.org/record/3822710