Paraphyly of Homoptera and Auchenorrhyncha inferred from 18S rDNA nucleotide sequences


Western Regional Research Center, USDA-ARS, 800 Buchanan St., Albany, CA94710–1100, U.S.A.


Evolutionary affiliations of eighteen families of Hemiptera (s.l.) are inferred using molecular phylogenetic analysis of nucleotide (nt) sequences of 18S rDNAs. Exemplar taxa include: Archaeorrhyncha (=Fulgoromorpha): flatid, issid, dictyopharid, cixiid and delphacid; Prosorrhyncha (=Heteropterodea): Peloridiomorpha (=Coleorhyncha) -peloridiid, Heteroptera gerrid, lygaeid and mirid; Clypeorrhyncha [=extant (monophyletic) cicadomorphs]: cicadid, cercopoids (cercopid, aphrophorid), membracid and cicadellids (deltocephaline and cicadelline); and Sternorrhyncha: psyllid, aleyrodid, diaspidid and aphid. Analysed sequences encompass a region beginning ≅550 nucleotides (nts) from the 5'-end to ≅200 nts upstream from the 3'-end of the gene [≅1150 base pairs (bp) in euhemipteran to >1400 bp in sternorrhynchan taxa]. Maximum parsimony and bootstrap analyses (PAUP) identify four principal hemipteran clades, Stenorrhyncha, Clypeorrhyncha, Archaeorrhyncha and Prosorrhyncha. These lineages are identified by synapomorphies distributed throughout the gene. Sternorrhyncha is a sister group to all other Hemiptera (i.e. Euhemiptera sensu Zrzavy), rendering Homoptera paraphyletic. Within Euhemiptera, clades Clypeorrhyncha, Archaeorrhyncha, Prosorrhyncha and Heteroptera are supported by one, three, two and three synapomorphic sites, respectively. There is equitable parsimonious inference for Archaeorrhyncha as the sister group to Prosorrhyncha (Neoherriiptera sensu Sorensen et al.) or Clypeorrhyncha, in either case rendering Auchenorrhyncha paraphyletic. Neohemiptera is supported by one synapomorphy. Within Clypeorrhyncha, clade cicada + cercopoids is the sister group of the clade cicadellids + membracid (Membracoidea sensu Dietrich & Deitz). Among archaeorrhynchans, clade delphacid + cixiid is the sister group of the clade dictyopharid + flatid + issid. Within Prosorrhyncha, the peloridiid is sister to the Heteroptera. Within Heteroptera, gerrid is the sister group of the clade mirid + lygaeid (Panheteroptera sensu Schuh). Based on secondary structure of synonymous 18S rRNA, two synapomorphies each of Sternorrhyncha, Prosorrhyncha and Heteroptera are compensatory substitutions on stem substructures. All other synapomorphies identifying major lineages of Hemiptera are noncompensatory substitutions on either bulges or stems. Short basal internodal distances suggest radiation of hemipteran lineages at the suborder level occurred rapidly. Morphological, palaeoentomological and eco-evolutionary factors supporting the 18S rDNA-based phylogenetic tree are discussed.