Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae)

Authors: NOMARK, BENJAMIN - UNIVERSITY OF MASSACHUSETTS; OKUSU, ARIKO - UNIVERSITY OF MASSACHUSETTS; MORSE, GEOFFREY - UNIVERSITY OF SAN DIEGO; PETERSON, DANIEL - UNIVERSITY OF MASSACHUSETTS; ITIOKA, TAKAO - KYOTO UNIVERSITY; Schneider, Scott

Submitted to: Zootaxa
Publication Type: Peer Reviewed Journal
Publication Acceptance Date: 2/3/2019
Publication Date: 6/17/2019
Citation: Nomark, B.B., Okusu, A., Morse, G.E., Peterson, D.A., Itioka, T., Schneider, S.A. 2019. Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae). Zootaxa. 4616:1-98. https://doi.org/10.11646/zootaxa.4616.1.1.
DOI: https://doi.org/10.11646/zootaxa.4616.1.1

Interpretive Summary: Armored scale insects, or the family Diaspididae, are an important group of common agricultural pests that cause billions of dollars in global economic losses each year. They are among the most invasive pests in the USA and other countries. This article reports on a phylogeny – a family tree – of armored scale insects and methods for curating specimens to produce morphological and molecular vouchers. This is the most comprehensive study of relationships among species of armored scale insects. We provide information about the classification of groups within the family Diaspididae. We also introduce updated identification keys to revised groups. This information is useful to port identifiers and to scientists studying scale insects. Armored scale insects also serve as model systems for a host of basic evolutionary questions. This work will contribute to both applied and basic fields of scientific research.

Technical Abstract: Armored scale insects are major economic pests and are among the world's most invasive species. We describe a system of specimen and ID management that establishes a basis for well-vouchered molecular identification. We also present an expanded Bayesian phylogenetic analysis based on concatenated fragments of four genetic loci: the large ribosomal subunit (28S), elongation factor 1-alpha, cytochrome oxidase I and II, and the small ribosomal subunit (16S) of the primary endosymbiont, Uzinura diaspidicola. Our sample includes 1389 individuals, representing 11 outgroup species and at least 311 described and 61 undescribed diaspidid species. The results broadly support Takagi's 2002 classification but indicate that some revisions are needed. We propose a revised classification recognizing four subfamilies: Ancepaspidinae Borchsenius, new rank, Furcaspidinae Balachowsky, new rank, Diaspidinae Targioni Tozzetti, and Aspidiotinae Westwood. Within Diaspidinae, in addition to the large tribes Diaspidini Targioni Tozzetti and Lepidosaphidini Shimer, we recognize 5 new tribes for phylogenetically isolated lineages: Aotearoaspidini Normark (for Aotearoaspis Normark, n. gen.), Umzantsiaspidini (for Umzantsiaspis Normark, n. gen.), Prodiaspidini Normark, Furchaspidini Normark, and Ischnaspidini Normark. Within Aspidiotinae, in addition to the existing tribes Aspidiotini Westwood, Parlatoriini Leonardi, Odonaspidini Ferris, Leucaspidini Atkinson, and Smilacicolini Takagi, we recognize as tribes Gymnaspidini Balachowsky, new rank, and Aonidiini Balachowsky, new rank. Within Diaspidini we recognize three subtribes: Diaspidina Targioni Tozzetti, Fioriniina Leonardi, and Chionaspidina Brues and Melander. We provide keys to these taxa. We also offer many additional minor modifications to diaspidid taxonomy, including many new combinations. Several taxa heretofore supposed to be primitive are shown to be relatively recently derived from more "advanced" taxa, often involving losses of structures, and we sink these taxa accordingly. We regard Kuwanaspidina Borchsenius as a junior synonym of Fioriniina, Thysanaspidini Takagi as a junior synonym of Leucaspidini, and Protodiaspidina Takagi and Ulucoccinae Takagi as junior synonyms of Chionaspidina. Indeed the morphologies that characterize a large proportion of sampled genera are derived from within a few highly paraphyletic genera, notably Pseudaulacaspis, Chionaspis, Aspidiotus, Pseudaonidia, and Parlatoria. We argue that each of these genera includes anciently polyphagous, abundant species -- today regarded as pests -- that have served as a kind of phylogenetic meristem, spewing out many more-specialized, range-restricted, and morphologically-reduced species in a process of nonadaptive radiation.