James N. Zahniser

Phylogeny of the leafhopper subfamily Deltocephalinae (Hemiptera: Cicadellidae) based on molecular and morphological data with a revised family-group classification.

Deltocephalinae, a highly diverse and economically important subfamily of leafhoppers, contains over 6200 species and 36 tribes distributed worldwide in habitats ranging from xeric grasslands and shrublands to tropical rainforests. Recent morphological and molecular phylogenetic analyses of Cicadellidae and a morphology‐based analysis of Deltocephalinae and related subfamilies indicated that several previously recognized cicadellid subfamilies are closely related to or derived from within Deltocephalinae, but these analyses did not provide a comprehensive or well‐supported hypothesis of the phylogeny of Deltocephalinae s.l. due to either low taxon sampling or low branch support. Here, taxon sampling was increased to include members of most family‐group taxa of Deltocephalinae and molecular data (∼2800 bp 28S rDNA and ∼350 bp histone H3) were added to improve the phylogenetic estimate. Five putative outgroup taxa were included, and parsimony and Bayesian analyses of the combined molecular and morphological (119 characters) data and maximum likelihood analyses of the 28S data showed strong support for the monophyly of Deltocephalinae as defined here. Branches near the base of the tree and towards the tips were longer and better supported than many of the shorter internal branches. Similar to a previous morphological phylogenetic analysis of Deltocephalinae, all of the grass‐ and sedge‐specializing tribes were recovered in one common clade, with a few apparent reversals to nongrass feeding. Although support for this clade was low and requires further testing, the results suggest that grass/sedge specialization is a phylogenetically conservative trait within Deltocephalinae. The history of the classification of Deltocephalinae and related subfamilies is reviewed, and based on the results of the phylogenetic analyses presented here, a revised family‐group taxonomic classification is proposed. In addition to subfamilies that were recently included in Deltocephalinae, the following are considered junior synonyms of Deltocephalinae: Acostemminae syn.n., Arrugadinae syn.n., Drakensbergeninae syn.n., Mukariinae syn.n. and Stegelytrinae syn.n. The morphological characters supporting this interpretation of Deltocephalinae are provided and discussed, and a description of the subfamily is provided. A new tribe, Faltalini tribe n. (11 genera, 31 species) is described, and Magnentiini placement n. and Paraphrodini placement n. are transferred to Deltocephalinae from Nioniinae and Aphrodinae, respectively. New placements of genera include: Twiningia Ball and Eusama Oman (Athysanini: Platymetopiina), placement n.; Cerrillus Oman (Athysanini), placement n.; Scaphotettix Matsumura and Agrica Strand (Mukariini), placement n.; Loralia Evans and Phlogotettix Ribaut (Deltocephalinae, unplaced to tribe), placement n. The recognition of Scaphoideini Oman 1943 as a nomen nudum results in the following placements: Acunasus DeLong, Cantura Oman, Danbara Oman, Osbornellus Ball, Prescottia Ball, Scaphodhara Viraktamath & Mohan, Scaphoideus Uhler, Scaphoidophyes Kirkaldy, Sincholata DeLong, Sobara Oman and Soleatus DeLong (Deltocephalinae, unplaced to tribe), placement n.

Phylogeny of the leafhopper subfamily Deltocephalinae (Insecta: Auchenorrhyncha: Cicadellidae) and related subfamilies based on morphology

Abstract Previous phylogenetic analyses of the leafhopper family Cicadellidae based on morphological and molecular data suggested that Deltocephalinae, as traditionally defined, is polyphyletic. These analyses recovered a large clade comprising Deltocephalinae and several other subfamilies. To further elucidate relationships within this large clade of deltocephaline‐like leafhoppers, an expanded dataset of 119 morphological characters and 68 taxa was compiled and analysed using cladistic methods. The taxon sample included one or more representatives of nearly all previously recognised tribes of Deltocephalinae, representatives of the non‐deltocephaline family‐group taxa which grouped with Deltocephalinae in previous analyses, and six putative outgroup taxa drawn from other cicadellid subfamilies. The resulting most parsimonious trees consistently recovered the putative ingroup as monophyletic. However, in agreement with previous analyses, most subfamilies and tribes represented by multiple exemplars were not resolved as monophyletic groups. Based on these results, subfamilies derived from within the deltocephaline lineage include Acostemminae, Arrugadinae, Drakensbergeninae, Eupelicinae, Koebeliinae, Mukariinae, Paraboloponinae, Penthimiinae, Selenocephalinae and Stegelytrinae. The phylogeny indicates that grass specialisation is much more phylogenetically conservative than implied by the current higher classification of leafhoppers. A list of synapomorphies on the strict consensus tree and an illustrated key to all family‐group taxa of deltocephaline‐like leafhoppers are provided.

First Reports of the Invasive Grass-Feeding Leafhopper Balclutha rubrostriata (Melichar) (Hemiptera: Cicadellidae) in the United States

ABSTRACT: The invasive leafhopper Balclutha rubrostriata (Melichar) was found in abundance in Bexar County, Texas. An arthropod diversity study along highway right-of-way in September and October, 2008 identified B. rubrostriata as the single most abundant species. This is the first peerreviewed published record of this invasive species in the mainland United States. Other records of this species in the U.S. are discussed. Diagnostic characters of the adults are provided and illustrated, and the nymph is described and illustrated for the first time. Genomic DNA was extracted from one adult and one nymph specimen, and the mitochondrial COI gene was sequenced for each and yielded identical sequences, thus confirming the association of the nymph and adult. Sequences are deposited in GenBank for future use in diagnostic or other studies.

Placement of the Faltala Oman Leafhopper Group (Hemiptera: Cicadellidae: Deltocephalinae) with Descriptions of Three New Species

Abstract Three new species of the Faltala Oman leafhopper group, Kramerana junina n. sp. and K. adusta n. sp. from the Andes Mountains of Peru and Clorindaia brasileira n. sp. from the Serra da Mesa of Brazil, are described. The tribal placement of the group and the high incidence of brachyptery and reduced (or absent) ocelli in the group are discussed. A revised key to the Faltala group genera and additional characters for Kramerana are provided.

A New Genus of Deltocephalinae (Hemiptera: Cicadellidae) from Ecuador and a New Synonym of Luheria constricta Osborn with Illustrations of the Female

Abstract The new deltocephaline leafhopper genus Oxycephalotettix is described and illustrated based one species, Oxycephalotettix tiputini sp. nov., from Ecuador, and is placed in Athysanini. The genus bears a superficial resemblance to Luheria Osborn (Luheriini), especially in the coloration of the forewing. Based on the examination of type material, Luheria constricta Osborn is herein considered a senior synonym of Tenucephalus hamatus DeLong, new synonymy. The female genitalia of L. constricta are illustrated for the first time, and the known range of this species is expanded to include Bolivia, northern Argentina, and northeastern Brazil.