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

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 (historic 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 L acaciae (Morrison & Morrison) comb.n. and L 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 (Cockerel]) 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.n., C. zeteki (Cockerell) comb.n.] and nine species to Gigantococcus [Gi. alboluteus (Cockerell) comb.n., Gi. bimaculatus (De Lotto) comb.n., Gi. brachystegiae (Hall) comb.n., Gi. longisetosus (Newstead) comb.n., Gi. nigroareolatus (Newstead) comb.n., Gi. pattersoni (Newstead) comb.n., Gi. schoutedeni (Vayssiere) comb.n., Gi. splendidus (Lindinger) comb.n., Gi. sulfureus (Lindinger) comb.n.]. From Crypticerya, we transfer seven species to Icerya [I. clauseni (Rao) comb.n., I. jacobsoni Green eomb.rev., I. jaihind (Rao) comb.n., I. kumri (Rao) comb.n., I. mangiferae (Tang & Hao) comb.n., I. natalensis (Douglas) comb.rev., L nuda Green comb.rev.] and five species to Gigantococcus [Gi. bicolor (Newstead) comb.n., Gi. cqjani (Newstead) comb.n., Gi. caudatus (Newstead) comb.n., Gi. ewarti (Newstead) comb.n., Gi. rodriguesi (Castel-Branco) comb.n.]. Both I. hyperici (Froggatt) and Palaeococcus dymocki (Froggatt) are syn.n. of I. nudata (all previously placed in Steatococcus). We recognize I. maynei Vayssiere as a syn.n. of Gi. nigroareolatus, I. tremae Vayssiere as a syn.n. of Gi. schoutedeni and L townsendi plucheae Cockerell as a syn.n. of C. townsendi. We revalidate the species name I. crocea Green stat.reval. In addition, we transfer I. taunayi Hempel to Laurencella Foldi (Monophlebidae: Llaveiini) as I. taunavi (Hempel) comb.n. Four species, Coccus hirticornis Boyer de Fonscolombe, I. chilensis Hempel, L insulans Hempel and I. paulista Hempel, are considered incertae sedis. We designate lectotypes for C. rosae, E. anomala and I. candida (a junior synonym of I. seychellarum). Following this revision, we recognize 74 species of Iceryini, distributed as follows: 22 in Crypticerya, one in Echinicerya, 19 in Gigantococcus, two in Gueriniella and 30 in Icerya.