A subfamily‐level classification of mealybugs (Hemiptera: Pseudococcidae) based on integrated molecular and morphological data

Abstract

The mealybugs (Hemiptera: Coccoidea: Pseudococcidae) are a speciose and ubiquitous group of sap‐sucking plant parasites, many of which are very serious agricultural pests. There has been much work on the alpha‐level taxonomy, amounting to the description of more than 2000 species, but suprageneric relationships remain poorly known. Downie & Gullan reviewed proposed schemes for a mealybug subfamily‐level classification and used DNA sequence data from three nuclear genes to infer the mealybug phylogeny. They recognized three subfamilies: the Rhizoecinae, the Phenacoccinae, and the Pseudococcinae; and within the Pseudococcinae, recognized the tribes Pseudococcini, Trabutinini, and Planococcini. Excepting the Trabutinini, none of these groupings was well‐supported. We improve our estimation of the mealybug phylogeny by: (i) increasing the taxon sampling by 50%; (ii) adding a morphological character matrix; and (iii) performing mixed model Bayesian and maximum likelihood inference procedures. We recover two primary clades within the Pseudococcidae, to which we apply the subfamily names Phenacoccinae and Pseudococcinae. Within the Phenacoccinae, we recover support for the monophyly of the hypogaeic mealybugs (Rhizoecini), and within the Pseudococcinae, we find support for the tribes Pseudococcini, Trabutinini, and Planococcini. Our results suggest a clear sequence of (i) β‐Proteobacteria primary endosymbionts infecting the mycetome of the most recent common ancestor of the Pseudococcinae, followed by (ii) γ‐Proteobacteria secondary endosymbionts infecting the primary β‐Proteobacteria endosymbionts. For each subfamily, we provide a morphological diagnosis and a comprehensive list of included genera. We resurrect the genus Ceroputo Šulc, currently under synonymy with the genus Puto Signoret (Putoidae), and place it in the mealybug subfamily Phenacoccinae. Greenoripersia kaiseri Bodenheimer is transferred from the Pseudococcidae to the Eriococcidae.

Ovaries of Puto superbus and Ceroputo pilosellae (Hemiptera: Coccoidea): Morphology, ultrastructure, phylogenetic and taxonomic implications

“…The genus Ceroputo that was synonymized with the genus Puto (Ferris, 1918) was resurrected by Hardy at al. (2008) and is now considered to be one of the genera of the Pseudococcidae placed in the subfamily Phenacoccinae.…”

Molecular phylogenetic analysis of a scale insect (Drosicha mangiferae; Hemiptera: Monophlebidae) infesting mango orchards in Pakistan

“…Because the use of standard taxonomic characters for identification has proven difficult (Danzing, 1997;Gullan & Cook, 2007), various molecular markers have been used to differentiate species and to ascertain evolutionary relationships (Beuning et al, 1999;Mowry & Barbour, 2004;Gadagkar et al, 2005;Gariepy et al, 2007). In some cases, molecular phylogenetic analyses have been combined with morphological characters to assess taxonomic relationships (Hardy et al, 2008). The most commonly used genes for the differentiation of insect species have been ribosomal RNA (18S, 28S, internal transcribed spacers) and mitochondrial cytochrome c oxidase I (COI) (Simon et al, 1994;Li et al, 2005).…”

“…In fact, sequence diversity in the 5' region of the COI gene has been established as the standard for species identification across the animal kingdom (Hebert et al, 2003;Miller, 2007;Linares et al, 2009). Several prior studies (Hardy et al, 2008;Rung et al, 2008;Ashfaq et al, 2010) have examined mealybug classification based on sequence variation in ribosomal and mitochondrial DNA. Similarly, sequence data of nuclear, mitochondrial and endosymbiotic genes have been used to estimate the phylogenies of scale insects (Cook et al, 2002;Morse & Normark, 2006;Gullan & Cook, 2007).…”

A new species of mealybug (Hemiptera: Pseudococcidae) from critically endangered Banksia montana in Western Australia

“…Pseudococcus has more than 150 named species worldwide (Ben-Dov 2102) but molecular data suggest that the genus is not monophyletic (Hardy et al 2008). However the relationships of sufficient numbers of species of Pseudococcus have not been studied for anyone to attempt to change the current functional classification of species in this and related genera.…”