The phylogeny of the superfamily Coccoidea (Hemiptera: Sternorrhyncha) based on the morphology of extant and extinct macropterous males

Abstract

Abstract. Currently, 49 families of scale insects are recognised, 33 of which are extant. Despite more than a decade of DNA sequence-based phylogenetic studies of scales insects, little is known with confidence about relationships among scale insects families. Multiple lines of evidence support the monophyly of a group of 18 scale insect families informally referred to as the neococcoids. Among neococcoid families, published DNA sequence-based estimates have supported Eriococcidae paraphyly with respect to Beesoniidae, Dactylopiidae, and Stictococcidae. No other neococcoid interfamily relationship has been strongly supported in a published study that includes exemplars of more than ten families. Likewise, no well-supported relationships among the 15 extant scale insect families that are not neococcoids (usually referred to as 'archaeococcoids') have been published. We use a Bayesian approach to estimate the scale insect phylogeny from 162 adult male morphological characters, scored from 269 extant and 29 fossil species representing 43/49 families. The result is the most taxonomically comprehensive, most resolved and best supported estimate of phylogenetic relationships among scale insect families to date. Notable results include strong support for (i) Ortheziidae sister to Matsucoccidae, (ii) a clade comprising all scale insects except for Margarodidae s.s., Ortheziidae and Matsucoccidae, (iii) Coelostomidiidae paraphyletic with respect to Monophlebidae, (iv) Eriococcidae paraphyletic with respect to Stictococcidae and Beesoniidae, and (v) Aclerdidae sister to Coccidae. We recover strong support for a clade comprising Phenacoleachiidae, Pityococcidae, Putoidae, Steingeliidae and the neococcoids, along with a sister relationship between this clade and Coelostomidiidae + Monophlebidae. In addition, we recover strong support for Pityococcidae + Steingeliidae as sister to the neococcoids. Data from fossils were incomplete, and the inclusion of extinct taxa in the data matrix reduced support and phylogenetic structure. Nonetheless, these fossil data will be invaluable in DNA sequence-based and total evidence estimates of phylogenetic divergence times.

Morphology of the Males of Seven Species of Ortheziidae (Hemiptera: Coccoidea)

“…Differences in the genital segments were also noted by Koteja (1987a) who therefore assigned the fossil to a separate genus Palaeonewsteadia. Despite the assumption that those two genera are closely related, Hodgson and Hardy (2013) retrieved Palae-onewsteadia as sister group to all recent Ortheziidae. Further analyses involving the male Ortheziidae described here is necessary to reassess their relationships.…”

“…In general, only the adult females (and sometimes late-instar nymphs) are currently identifiable because the original descriptions were based solely on the former, along with their associated nymphs. However, our understanding of the morphology of adult male scale insects is gradually improving, with the recognition that phylogenetic analyses based on male morphology have given us a much improved understanding of scale insect phylogeny (Hodgson and Hardy, 2013) compared with the structure of the larviform adult females. additionally, specimens of fossil adult males of many scale insect families are now known and are commonly preserved in amber around the world (almost always without their adult female counterpart).…”

Diverse New Scale Insects (Hemiptera: Coccoidea) in Amber from the Cretaceous and Eocene with a Phylogenetic Framework for Fossil Coccoidea

“…113. Type of glandular pouch (adapted from Hodgson and Hardy [2013]): 0 = group of pores but not pouchlike and flat (e.g., Phenacoleachia); 1 = shallow pouch (e.g., Puto, Steingelia); 2 = deep pouch (e.g., Pseudococcidae, Eriococcidae, Coccidae); 3 = absent. 114.…”

“…nevertheless, sufficient comparative information exists among families and genera of some families, which potentially allows phylogenetic placement of fossilized males (Hodgson and Hardy, 2013).…”

New fossil taxa of Monophlebidae (Sternorrhyncha: Coccoidea) from Baltic amber

“…Morrison considered that it comprised 5 tribes: Drosichini, Iceryini, Llaveiini, Monophlebini and Monophlebulini. Now all subfamilies and most of the tribes sensu Morrison (1928) have been given family status (Koteja, 1974;Foldi, 2005;Hodgson & Foldi, 2006;Gullan & Cook, 2007;Hodgson & Hardy, 2013).…”