Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing the re‐interpretation of morphology and a revised generic classification
Authors: UNRUH C. M. , GULLAN P. J. 
Published: November 2007

AbstractThe 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.GueriniellaFernald is sister to the rest of the Iceryini and the generaCrypticeryaCockerell,IceryaSignoret andSteatococcusFerris 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,EchiniceryaMorrison,GigantococcusPesson & Bielenin,GueriniellaandIcerya– and substantially revise the generic concepts ofCrypticerya,GigantococcusandIcerya.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 speciesCrypticerya rosae(Riley & Howard),Echinicerya anomalaMorrison,Gigantococcus maximus(Newstead) (adult female only),Gueriniella serratulae(Fabricius) andIcerya seychellarum(Westwood). We recognizeAuloiceryaMorrison as a junior synonym (syn.n.) ofIcerya, and transfer the twoAuloiceryaspecies toIceryaasI. acaciae(Morrison & Morrison)comb.n.andI. australisMaskellcomb.rev.We recognizeSteatococcusandProticeryaCockerell as junior synonyms (syn.n.) ofCrypticerya. FromSteatococcus, we transfer five species toCrypticerya[C. mexicanaCockerell & Parrottcomb.rev.,C. morrilli(Cockerell)comb.n.,C. tabernicola(Ferris)comb.n.,C. townsendiCockerellcomb.rev.,C. tuberculata(Morrison)comb.n.], four species toGigantococcus[Gi. euphorbiae(Brain)comb.n.,Gi. gowdeyi(Newstead)comb.n.,Gi. madagascariensis(Mamet)comb.n.,Gi. theobromae(Newstead)comb.n.] and three species toIcerya[I. assamensis(Rao)comb.n.,I nudataMaskellcomb.rev.,I. samaraia(Morrison)comb.n.]. FromIcerya, we transfer 14 species toCrypticerya[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.n.,C. zeteki(Cockerell)comb.n.] and nine species toGigantococcus[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(Vayssière)comb.n.,Gi. splendidus(Lindinger)comb.n.,Gi. sulfureus(Lindinger)comb.n.]. FromCrypticerya, we transfer seven species toIcerya[I. clauseni(Rao)comb.n.,I. jacobsoniGreencomb.rev.,I. jaihind(Rao)comb.n.,I. kumari(Rao)comb.n.,I. mangiferae(Tang & Hao)comb.n.,I. natalensis(Douglas)comb.rev.,I. nudaGreencomb.rev.] and five species toGigantococcus[Gi. bicolor(Newstead)comb.n.,Gi. cajani(Newstead)comb.n.,Gi. caudatus(Newstead)comb.n.,Gi. ewarti(Newstead)comb.n.,Gi. rodriguesi(Castel‐Branco)comb.n.]. BothI. hyperici(Froggatt) andPalaeococcus dymocki(Froggatt) aresyn.n.ofI. nudata(all previously placed inSteatococcus). We recognizeI. mayneiVayssière as asyn.n.ofGi. nigroareolatus,I. tremaeVayssière as asyn.n.ofGi. schoutedeniandI. townsendi plucheaeCockerell as asyn.n.ofC. townsendi. We revalidate the species nameI. croceaGreenstat.reval.In addition, we transferI. taunayiHempel toLaurencellaFoldi (Monophlebidae: Llaveiini) asL. taunayi(Hempel)comb.n.Four species,Coccus hirticornisBoyer de Fonscolombe,I. chilensisHempel,I. insulansHempel andI. paulistaHempel, are consideredincertae sedis. We designate lectotypes forC. rosae,E. anomalaandI. candida(a junior synonym ofI. seychellarum). Following this revision, we recognize 74 species of Iceryini, distributed as follows: 22 inCrypticerya, one inEchinicerya, 19 inGigantococcus, two inGueriniellaand 30 inIcerya.