Morphology‐based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

Abstract

Summary A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.

Systematics and phylogeny of the Neotropical treehopper subfamily Nicomiinae (Hemiptera, Membracidae)

“…Antattended and gregarious species are common in both Aetalionidae and Membracidae and this seems to be the plesiomorphic condition for these two families (DIETRICH et al 2001, CRYAN et al 2004. Nicomiines are collected rarely, usually only as single individuals (WOOD 1984), suggesting that these species, unlike many membracids, are solitary as adults.…”

“…The treehopper family Membracidae Rafinesque, 1815 (Hemiptera, Cicadomorpha), is a group of approximately 3,100 species of sap-sucking insects classified into nine subfamilies (DIETRICH et al 2001, WALLACE & DEITZ 2004). The family is distributed worldwide but is most diverse in the New World tropics.…”

“…The family is distributed worldwide but is most diverse in the New World tropics. Recent phylogenetic analyses (DIETRICH et al 2001, CRYAN et al 2004) indicated that some subfamilies, as currently defined, are para-or polyphyletic and that more detailed analyses of certain lineages are needed. One such lineage includes the subfamily Nicomiinae, a little known group of Neotropical treehoppers previously comprising two genera, Tolania Stål and Nicomia Stål (DIETRICH et al 2001).…”

“…Endoiastinae, the sister group to the remaining membracids, apparently arose prior to the initial acquisition of a posterior pronotal process. Morphological evidence , DIETRICH et al 2001 indicates that Nicomiinae are derived from treehoppers with the posterior process well developed. In particular, some nicomiine species retain the notch or groove in the scutellum in which the posterior process of other membracids rests.…”

“…HAMILTON (1971) considered Euwalkeria to be closely related to the cicadellid genus Coloborrhis (Ulopinae) based on characters of the head and forewing, while Euwalkeria, Holdgatiella, and three additional genera (Antillotolania Ramos, Deiroderes Ramos, and Togotolania CRYAN & DEITZ) appear to be related to Nicomiinae because they lack a posterior pronotal process (the process is present or absent in Euwalkeria) and have the width of the head greater than the distance across the pronotal humeri. The morphology-based phylogeny of DIETRICH et al (2001) grouped these genera (Togotolania Cryan & Deitz listed as "New Genus T") in a clade with Nicomiinae, Stegaspidinae, Centronodinae, and Centrodontinae, but relationships among these taxa were poorly resolved. Subsequent, more detailed morphology-based analyses of Stegaspidinae placed Antillotolania + Deiroderes as sister to a clade comprising two undescribed Dominican amber species and the extant members of Stegaspidini and Microcentrini, but the position of Togotolania remained unresolved (CRYAN et al 2003).…”