Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing the re‐interpretation of morphology and a revised generic classification

Abstract

The scale insect tribe Iceryini (Coccoidea: Monophlebidae) is a group of relatively large and polyphagous insects found worldwide. Currently, the tribe contains about 80 named species placed in seven genera, which are diagnosed largely on features associated with egg protection. We reconstruct the phylogeny of the Iceryini on the basis of nucleotide sequence data from nuclear ribosomal (18S and D2, D3 and D10 regions of 28S) and protein‐coding (histone H3) gene regions of 40 iceryine species representing six of the seven genera and seven outgroup taxa, mostly from two other tribes of Monophlebidae. Bayesian and maximum parsimony analyses recover a monophyletic tribe and clades that correspond more to geography than to the existing morphology‐based classification. Gueriniella Fernald is sister to the rest of the Iceryini and the genera Crypticerya Cockerell, Icerya Signoret and Steatococcus Ferris are not monophyletic. Our data imply that the distinctive iceryine reproductive strategies, such as protecting eggs in a waxy ovisac or inside a marsupium, are poor indicators of relationships. On the basis of molecular relationships and the re‐examination of morphological characters, we recognize only five genera of Iceryini –Crypticerya, Echinicerya Morrison, Gigantococcus Pesson & Bielenin, Gueriniella and Icerya – and substantially revise the generic concepts of Crypticerya, Gigantococcus and Icerya. We provide a key to the genera based on adult females. We redescribe and illustrate the adult female and first‐instar nymph of the type species Crypticerya rosae (Riley & Howard), Echinicerya anomala Morrison, Gigantococcus maximus (Newstead) (adult female only), Gueriniella serratulae (Fabricius) and Icerya seychellarum (Westwood). We recognize Auloicerya Morrison as a junior synonym (syn.n.) of Icerya, and transfer the two Auloicerya species to Icerya as I. acaciae (Morrison & Morrison) comb.n. and I. australis Maskell comb.rev. We recognize Steatococcus and Proticerya Cockerell as junior synonyms (syn.n.) of Crypticerya. From Steatococcus, we transfer five species to Crypticerya [C. mexicana Cockerell & Parrott comb.rev., C. morrilli (Cockerell) comb.n., C. tabernicola (Ferris) comb.n., C. townsendi Cockerell comb.rev., C. tuberculata (Morrison) comb.n.], four species to Gigantococcus [Gi. euphorbiae (Brain) comb.n., Gi. gowdeyi (Newstead) comb.n., Gi. madagascariensis (Mamet) comb.n., Gi. theobromae (Newstead) comb.n.] and three species to Icerya [I. assamensis (Rao) comb.n., I nudata Maskell comb.rev., I. samaraia (Morrison) comb.n.]. From Icerya, we transfer 14 species to Crypticerya [C. brasiliensis (Hempel) comb.n., C. colimensis (Cockerell) comb.n., C. flava (Hempel) comb.n., C. flocculosa (Hempel) comb.n., C. genistae (Hempel) comb.n., C. littoralis (Cockerell) comb.n., C. luederwaldti (Hempel) comb.n., C. minima (Morrison) comb.n., C. montserratensis (Riley & Howard) comb.n., C. palmeri (Riley & Howard) comb.n., C. rileyi (Cockerell) comb.n., C. similis (Morrison) comb.n., C. subandina (Leonardi) comb...

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

“…A noticeable difference between northern and southern region of Pakistan is the high altitude and hilly terrain of north versus the lowlands of the south, factors that may have played a role in explaining the regional divergence we detected in D. mangiferae. Other members of the Monophlebidae are typically polyphagous (Unruh & Gullan, 2008) so the presence of D. mangiferae on multiple host plants is not unusual. We conclude that D. mangiferae and D. stebbingi (Latif, 1949) are simply host races of the same species.…”

Scanning electron microscopy of six selected mealybug (Hemiptera: Pseudococcidae) species of Sri Lanka

“…Such differences have been used to revise the generic placement of some scale insect species into families, such as Coccidae, Diaspididae and Monophlebidae (Takagi, 1990;Ülgentürk and Wilhem, 2001;Unruh, 2008;Unruh and Gullan, 2008). Each quinquelocular pores consist of a pentagonal, raised area of cuticle containing a circle of five openings or loculi.…”

A new species of Crypticerya Cockerell (Hemiptera: Monophlebidae) from Colombia, with a key to species of the tribe iceryini found in South America

“…5) and was found to be closely related to C. zeteki, C. brasiliensis and C. montserratensis (Unruh & Gullan 2008a). As a result, it was included in the informal Crypticerya montserratensis group described in Unruh & Gullan (2008b) as it shares several key features to other species in the group that are not seen in any other iceryine species.…”

“…The following abbreviations are for institutions or collections where material were deposited: BME, The Unruh & Gullan (2008a)] and stored at BME.…”

“…Although many South American Crypticerya species resemble one another in life, especially species in the montserratensis group (Fig 1), see images of C. montserratensis (Riley & Howard), C. brasiliensis (Hempel), C. zeteki (Cockerell) and C. multicicatrices Kondo & Unruh sp. n., they are genetically distinct (Unruh & Gullan 2008a). Several Crypticerya species are well-documented pests in the Neotropics.…”