ZOOLOGIA 30 (5): 519–568, October, 2013 http://dx.doi.org/10.1590/S1984-46702013000500008 South American leafhoppers of the tribe Typhlocybini (Hemiptera: Cicadellidae: Typhlocybinae) Christopher H. Dietrich Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 S. Oak St., Champaign, IL 61820, USA. E-mail: [email protected] ABSTRACT. The fauna of Typhlocybini (sensu stricto, excluding Empoascini) endemic to South America is reviewed and comprises seven closely related genera, five described herein as new, and 55 species, 52 here described as new. The genera and species are described and keys and illustrations are provided to aid in their identification. Columbonirvana Linnavuori comprises 17 species, 16 new. Eualebra Baker comprises 19 species, 17 new. Euzyginella gen. nov., comprises four new species. Neozyginella gen. nov., comprises six new species. Pseudhadina gen. nov., comprises one new spe- cies. Pseudozyginella gen. nov., comprises three new species. Tahurella gen. nov., comprises five new species. One new synonym is recognized: Eualebra smithii Baker, 1899 equals Dikraneura (Hyloidea) reticulata Osborn, 1928, syn. nov. One previously described species placed in Eualebra belongs in tribe Dikraneurini; thus, a new combination is proposed: Alconeura (Hyloidea) rubra (Van Duzee), comb. nov. Most of the specimens available for this study were from Malaise trap and canopy fogging samples obtained at very few localities in Bolivia, Brazil, Colombia, Ecuador, and Peru, sug- gesting that further sampling in South America, particularly in the Amazonian rainforest and eastern foothills of the Andes Mountains, will reveal large numbers of additional species. KEY WORDS. Distribution; Homoptera; morphology; Neotropical; taxonomy. The leafhopper subfamily Typhlocybinae comprises ~6,000 also includes Columbonirvana Linnavuori, a genus previously described species of mostly small, delicate leafhoppers that feed placed in Nirvaninae (LINNAVUORI 1959), but transferred to preferentially on the contents of leaf parenchyma cells of their Typhlocybinae by DIETRICH (2004) based on phylogenetic analy- host plants. Based on described species, this is currently the sec- sis of morphological characters. These two genera include a ond largest leafhopper subfamily (after Deltocephalinae, ZAHNISER total of three previously described species (plus one new syn- & DIETRICH 2010), but recent sampling of tropical faunas indi- onym, designated below). Recent sampling indicates that the cates that the extant, but mostly undescribed, fauna of South American fauna of Typhlocybini is much richer in both Typhlocybinae is far larger than that of any other leafhopper species and genera. In this paper, I review the endemic South subfamily (DIETRICH & WALLNER 2002). The Neotropical fauna, in American genera and species of Typhlocybini and discuss their particular, is highly diverse but appears to remain largely un- possible relationships to the fauna of this tribe in other parts documented (DIETRICH & DMITRIEV 2006, 2008). of the world. Considering that the 52 new species of Study of canopy fogging samples from lowland rainforests Typhlocybini described herein were collected from only eight in eastern Peru and Ecuador (DIETRICH & WALLNER 2002) indi- localities in Bolivia, Brazil, Colombia, Ecuador and Peru, fur- cates the presence of an enormously diverse typhlocybine fauna. ther sampling in South America, especially in cloud forests of Composition of this fauna contrasts strikingly with that of the the eastern Andean foothills, will likely reveal the presence of somewhat better known, but still poorly documented faunas of a much more speciose assemblage belonging to this tribe. the Old World tropics. Tribes Dikraneurini, Empoascini (= Jorumini) and Alebrini, in decreasing order of species rich- MATERIAL AND METHODS ness, are well represented in the samples from Amazonia; but Erythroneurini and Typhlocybini, so abundant in north tem- Morphological terminology follows that of YOUNG (1952) perate forests and in the Old World tropics, are relatively rare. as modified by DIETRICH (2005) except wing veins are labeled Malaise trap samples from the eastern Andean foothills according to the simplified system illustrated in figures 3i, j. above 500 m elevation differ from those obtained from the Drawings of genitalia were made by tracing over photographs canopy of lowland rainforest (unpublished data) in that they taken using a digital camera mounted on a compound micro- include larger numbers of Typhlocybini (sensu stricto). These scope. In drawings of the male genital capsule, the entire anal species belong to a group of apparently closely related, endemic complex (segments X-XI) is shown in lateral view but only the South American genera that includes Eualebra Baker. The group well sclerotized and pigmented parts of segment X are shown © 2013 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | All rights reserved. 520 C.H. Dietrich in dorsal view. Material examined is deposited in the follow- 2’. Hind wing submarginal vein incomplete, either extended ing institutions: Humboldt Institute, Villa de Leyva, Colombia along costal margin but not connecting vein apices along (HIC); Carnegie Museum of Natural History (ICCM); Illinois apical margin, or connecting vein apices along apical Natural History Survey, Champaign (INHS); Universidad de San margin but not extended along costal margin (terminated Marcos, Lima (USML), United States National Museum (USNM). at apex of vein RP or R+M)............................................... 3 3. Forewing with inner apical cell parallel-sided, hind wing TAXONOMY submarginal vein absent along apical margin, vannal vein unbranched ................................................ Erythroneurini Typhlocybinae Kirschbaum, 1868 3’. Forewing with inner apical cell usually distinctly tapered The tribal classification of Typhlocybinae has long been through most of length (often short and oblique), hind unstable, with some authors (e.g., YOUNG 1952) having recog- wing submarginal vein, if present, not extended along costal nized as few as four tribes and others (e.g., RUPPEL 1987) as many margin basad of RP or R+M, vannal vein branched ....... 4 as ten. In his review of Western Hemisphere Typhlocybinae, 4. Ocelli absent or vestigial, hind wing submarginal vein absent YOUNG (1952) employed a relatively broad concept of between apices of veins MA or RP+MA and MP (Fig. 78) or, Typhlocybini, including as synonyms the tribes Eupterygini if present (Fig. 70), body strongly depressed with face nearly Kirkaldy, 1906, Jorumini McAtee, 1926, and Empoascini Dis- horizontal ...................................................... Typhlocybini tant, 1908, despite differences in the hind wing venation of 4’. Ocelli well developed (except in Paulomanus), hind wing these groups. Dworakowska, the principal worker on the Old submarginal vein present between apices of veins MA or World typhlocybine fauna for the past four decades, recognized RP+MA and MP, body not strongly depressed, face in profile tribes Typhlocybini, with synonym Eupterygini; Empoascini, oblique, not nearly horizontal ........................ Empoascini with synonym Jorumini; and Zyginellini (DWORAKOWSKA 1979). The latter tribe was erected to comprise species having the hind Typhlocybini Kirschbaum wing submarginal vein apparently connected directly to CuA Typhlocybini Kirschbaum, 1868: 16. rather than being joined by a crossvein (alternatively inter- Eupteryginae Kirkaldy, 1906: 296. preted as the absence of the segment of vein CuA distad of its Zyginellini Dworakowska, 1979: 299. junction with the submarginal vein). Despite the unique hind Diagnosis. Species of Typhlocybini (sensu AHMED 1983) wing venation, Zyginellini, as defined by DWORAKOWSKA (1979), differ from other Typhlocybinae in having the following com- is a rather heterogeneous assemblage. Most genera placed in bination of morphological features: ocelli absent or vestigial; this tribe strongly resemble many Typhlocybini (sensu stricto), forewing appendix absent, inner apical cell short and oblique, particularly in the structure of the male genital capsule (e.g., not extended to apical wing margin; hind wing vannal vein presence of a single macroseta near the base of the plate) and branched, submarginal vein present or absent at wing apex; aedeagus (e.g., presence of elongate paired apical processes). male pygofer with few or no macrosetae (except Eualebra). Moreover, some of the Neotropical species described herein Remarks. The endemic South American taxa treated vary intraspecifically (occasionally intra-individually) for the herein fit AHMED’s (1983) concept of Typhlocybini, but form a features used to distinguish Zyginellini from Typhlocybini. distinctive group within this tribe. Members of this group are AHMED (1983) considered Zyginellini to be an artificial group readily distinguished by their depressed form, produced head, and treated it as a synonym of Typhlocybini. Until phyloge- and presence of two or more macrosetae on the male subgenital netic analyses can clarify the status and relationships of the plate (most Old World and Nearctic Typhlocybini have 0-1). various tribes proposed for groups of typhlocybine genera, it Some of the genera included here have hind wing vena- seems reasonable to adopt the classification of AHMED (1983), tional patterns not known to occur among Typhlocybini in which recognizes five tribes in the subfamily: Alebrini, other parts of the world.