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

Abstract

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 Uhl...

A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae)

“…Also, taxonomic concepts of many family-group taxa have often not been clearly stated or universally agreed upon, or even if they have, their circumscriptions have often not been tested by examination of multiple character sets or in phylogenetic analyses. These deficiencies have been partly addressed by taxonomic revisions (e.g., Linnavuori & Al-Ne'amy 1983, Knight & Webb 1993, Zhang & Webb 1996, Dietrich & Dmitriev 2003 and recent phylogenetic studies based on morphological (Dietrich 1999, Knight & Webb 1993, Zahniser & Dietrich 2008) and molecular data (Dietrich et al 2001, Zahniser & Dietrich 2010. These studies provided strong evidence European Journal of Taxonomy 45: 1-211 (2013) that Deltocephalinae sensu Oman et al (1990) was not monophyletic and that several other leafhopper subfamilies had their closest relatives within the deltocephaline lineage.…”

“…The taxon sample used by Zahniser & Dietrich (2010) was expanded here to include 37 additional exemplars of Deltocephalinae. All analyses except one supplementary analysis included 152 total taxa, 5 of which were putative outgroup taxa.…”

“…The difficulty in the classification of these genera appears to be due to a rapid radiation of deltocephaline leafhoppers that left very little phylogenetic signal in morphological and molecular characters (Zahniser & Dietrich 2010) among a morphologically diverse group of deltocephalines. they possess the typical Y-shaped connective of the male genitalia but do not possess any of the characters that define other tribes, and are consequently a polyphyletic assemblage of genera that is difficult to make sense of as a whole.…”

“…These studies provided strong evidence European Journal of Taxonomy 45: 1-211 (2013) that Deltocephalinae sensu Oman et al (1990) was not monophyletic and that several other leafhopper subfamilies had their closest relatives within the deltocephaline lineage. Based on the accumulated data and understanding of the relationships of Deltocephalinae and other leafhoppers, Zahniser & Dietrich (2010) adopted a revised, broader concept of the subfamily and provided a detailed history of the classification of the subfamily.…”

“…*Identification updated from previous study (Zahniser & Dietrich 2010). **28S data taken from previous study (Dietrich et al 2001;Rakitov, unpublished) …”