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Supplementum 2, 2018, pp. 1-260. Published 31 May 2018

Managing editor: D.A. Gapon

I.A. Gavrilov-Zimin. 2018. Ontogenesis, morphology and higher classification of archaeococcids (Homoptera: Coccinea: Orthezioidea). Zoosystematica Rossica, Supplementum 2: 1-260.


The monograph summarizes original research data and published literature data on reproduction, life cycles, individual development and morphology of scale insects of the superfamily Orthezioidea (archaeococcids). The superfamily system is accepted mainly in its traditional concept, i.e. with four well-defined families: Margarodidae s. l., Ortheziidae, Carayonemidae, and Phenacoleachiidae. The tribe Matsucoccini (Margarodidae s. l.: Xylococcinae s. l.) is considered as a most archaic group of scale insects according to morphological, reproductive and ontogenetic characters. A complicated ontogenesis with an alternation of movable/immovable instars and with arostrate imago of both sexes (as in Matsucoccus Cockerell, 1909 and many other Margarodidae s. l.) is presumed to be initial in scale insect evolution and such ontogenesis is supposed to be an apomorphy of suborder Coccinea. Distribution of different variants of ovoviviparity/viviparity amongst scale insect families is overviewed. It is demonstrated that the evolution of scale insects shows multiple cyclic conversions of oviparous reproduction pattern to ovoviviparous/viviparous ones with the appearance of new and new peculiar adaptations to eggs protection; the most ancient scale insects (Matsucoccini and their ancestor) were probably facultatively ovoviviparous, whereas the origin of the whole neococcid phylogenetic line (Coccoidea s. s.) was probably connected with obligate complete ovoviviparity, which also appeared in some “derived” archaeococcids of the tribe Iceryini (Margarodidae s. l.), in the families Phenacoleachiidae and Carayonemidae. New taxonomic additions and changes in generic composition of some tribes are provided for the family Margarodidae s. l., in its subfamilies Monophlebinae and Callipappinae s. l. The tribe Labioproctini tr. nov. (Monophlebinae) is erected for six genera possessing peculiar quadrilocular wax pores: Aspidoproctus Newstead, 1901, Hemaspidoproctus Morrison, 1927, Labioproctus Green, 1922, Lecaniodrosicha Takahashi, 1930, Misracoccus Rao, 1950, and Walkeriana Signoret, 1876. The presence of quadrilocular pores are considered as a synapomorphic character of the Labioproctini tr. nov. and Ortheziidae. Disputable taxonomic position of Xenococcidae Tang, 1992 is discussed and this family is also placed in Orthezioidea. New genera and species are described and illustrated, based mainly on material collected in the Oriental region: Eremostoma klugei gen. et sp. nov., Crambostoma largecicatricosum gen. et sp. nov. (both in Callipappinae s. l.: Coelostomidiini s. l.), Buchnericoccus reynei sp. nov., Monophlebus neglectus sp. nov. (both in Monophlebinae: Monophlebini), Crypticerya ovivivipara sp. nov., Icerya oculicicatricata sp. nov., I. siamensis sp. nov. (all three in Monophlebinae: Iceryini).

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