Morphological phylogeny of Dictyopharidae analysis for the first time

Morphological phylogeny of Dictyopharidae (Hemiptera: Fulgoromorpha)

To explore the phylogenetic basis of the current classification of Dictyopharidae, especially the relationships among the tribes of Dictyopharinae, we present the first cladistic analysis of this family based on 146 morphological characters of adults. Our analysis includes 104 of 125 recognized genera and subgenera within 12 extant tribes of Dictyopharinae, plus nine genera representing all four tribes of Orgeriinae.

The results of this study support Dictyopharidae as a monophyletic group with Aluntiini sister to the remaining Dictyopharidae, but do not support Orgeriinae as sister to Dictyopharinae. Seven major lineages – Aluntiini, Arjunini, Hastini, (Taosini+Lappidini)+Nersiini, a Xenochasma+ complex (including ‘Orgeriinae’), Orthopagini, and Dictyopharini – are recovered in Dictyopharidae. The Xenochasma+ complex is proposed here and includes Xenochasma clade + (Scoloptini +((Fernandea clade+Phylloscelini)+(Rancodini+[Capenini+Orgeriinae]))). Within this complex, some genera are of uncertain tribal placement, and the Orgeriinae are retained as a subfamily until a molecular phylogeny can confirm the results of this paper. The implied weighting analysis supports the monophyly of most tribes of Dictyopharinae (except Taosini), the sister-group relationships of (Taosini+Lappidini) with Nersiini, and Orthopagini+Dictyopharini, and the current tribal classification for Dictyopharinae. Most Dictyopharinae genera fit their respective tribal affiliation, but some proposed changes are that Pharodictyon, Paramisia, Dictyopharoides s.s., Chondrophana, Sicoris, Chondrodire, and Tupala are provisionally placed in Hastini; Pukuakanga is moved into Nersiini; Sinodictya and Raphiophora are transferred into Orthopagini; and Chiltana, Litocras, and Viridophara are placed in Dictyopharini.

Although most current tribes are recovered, most tribal and generic concepts lack solid autapomorphic support. In connection with molecular approaches yet to be undertaken, future investigations for the morphological side should combine a new set of characters. Female internal genitalia (Bourgoin, 1993; Soulier-Perkins, 2001; Gnezdilov, 2003; Weirauch, 2008), nymphs, fossil and even outgroup selection to lessen the impact of homoplasy need to be taken properly into consideration. Female accessory genitalic glands, in particular (Song & Bourgoin, in preparation), but also nymphal morphological characters, especially pattern and structure of the sensory and glandular units (wax-plate areas) on the tergites, may also display important new sets of characters . 

Our results offer a framework for further explorations into the systematics of the family.We are confident that future systematic investigation will soon lead to an updated classification (including identification keys). Well-tested phylogenetic hypotheses will also allow investigations into the biogeographic history and the evolution of multitrophic ecological interactions in this conspicuous group of herbivores.

The work was supported by grants from the National Natural Science Foundation of China (no. 31572297, to ZSS) and Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (no. Y4ZK111B01), and partially by grants from the National Natural Science Foundation of China (nos 31561163003 and 31572298, to APL). And published as "Morphological phylogeny of Dictyopharidae (Hemiptera: Fulgoromorpha) "on Systematic Entomology.

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A, Orthopagus sp.; B, Saigona fuscoclypeata; C, Tenguna medogensis, A–C photographed by Z.-S. Song; D, Centromeria triangulata, photographed by Paul Bertner; E, Centromeria speilinea, photographed by Petr Kočárek.

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Phylogenetic Tree Based on Morphological Characteristics of Dictyopharidae

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Partial morphological features of Dictyopharidae