A molecular phylogeny of the cicadas (Hemiptera: Cicadidae) with a review of tribe and subfamily classification

Abstract

A molecular phylogeny and a review of family-group classification are presented for 137 species (ca. 125 genera) of the insect family Cicadidae, the true cicadas, plus two species of hairy cicadas (Tettigarctidae) and two outgroup species from Cercopidae. Five genes, two of them mitochondrial, comprise the 4992 base-pair molecular dataset. Maximum-likelihood and Bayesian phylogenetic results are shown, including analyses to address potential base composition bias. Tettigarcta is confirmed as the sister-clade of the Cicadidae and support is found for three subfamilies identified in an earlier morphological cladistic analysis. A set of paraphyletic deep-level clades formed by African genera are together named as Tettigomyiinae n. stat. Taxonomic reassignments of genera and tribes are made where morphological examination confirms incorrect placements suggested by the molecular tree, and 11 new tribes are defined (Arenopsaltriini n. tribe, Durangonini n. tribe, Katoini n. tribe, Lacetasini n. tribe, Macrotristriini n. tribe, Malagasiini n. tribe, Nelcyndanini n. tribe, Pagiphorini n. tribe, Pictilini n. tribe, Psaltodini n. tribe, and Selymbriini n. tribe). Tribe Tacuini n. syn. is synonymized with Cryptotympanini, and Tryellina n. syn. is synonymized with an expanded Tribe Lamotialnini. Tribe Hyantiini n. syn. is synonymized with Fidicinini. Tribe Sinosenini is transferred to Cicadinae from Cicadettinae, Cicadatrini is moved to Cicadettinae from Cicadinae, and Ydiellini and Tettigomyiini are transferred to Tettigomyiinae n. stat from Cicadettinae. While the subfamily Cicadinae, historically defined by the presence of timbal covers, is weakly supported in the molecular tree, high taxonomic rank is not supported for several earlier clades based on unique morphology associated with sound production.

Evolutionary relationships amongMassosporaspp. (Entomophthorales), obligate pathogens of cicadas

“…Two Massospora species treated in this study, Mt and Ml / Mp have cicada hosts both belonging to the subfamily Tibicinae, while the hosts of Md and Mc belong to two other subfamilies, Cicadinae and Cicadettinae, respectively (Sanborn 2013, Łukasik et al 2018, Marshall et al 2018 The results of the morphological study presented here indicate that spore measurements may not be useful for species level identifications. Unfortunately, the numbers of isolates sampled for many of these species were insufficient to confidently conclude whether differences truly exist.…”

“…Conidia are passively disseminated during mating attempts or flights, or possibly in crowded settings where high densities of cicadas promote close contact (Soper 1963, Cooley et al 2018. Cicadas infected by conidia develop secondary infections (Soper et al 1976b, Cooley et al 2018, resulting in the production of resting spores inside cicada hosts. These resting spores are incorporated back into the soil to infect new cohorts of cicadas as they emerge in later years (Soper et al 1976a).…”

“…AHT behavior in Massospora is thought to be chemically induced (Boyce et al 2019), and includes hypersexual behavior where infected male cicadas mimic female-specific behaviors to attract copulation attempts from other males (Cooley et al 2018). Massospora and Strongwellsea, a fly pathogen, are the only two genera where all species are known to induce AHT behavior in their hosts, although AHT has also been reported in select species of Entomophthora (E. erupta and E. thripidum) and Entomophaga (E. kansana) (Roy et al 2006).…”

Checklist, endemism, English vernacular names and identification of the cicadas (Insecta, Hemiptera, Cicadidae) of KwaZulu-Natal, South Africa

“…If a robust and stable phylogenetic classification of a group has been established (e.g. Marshall et al 2018), the opportunity to coin vernacular names is also an opportunity to co-ordinate the folk taxonomy with the hierarchy of relationships that is inherent in the scientific classification.…”

“…Based on the species list, relevant taxonomic literature (e.g. Distant 1906;Villet 1987Villet , 1989Villet , 1994Villet , 1997Villet and Reavell 1989;Marshall et al 2018) and experience, a taxonomic key to adult males of the subfamilies, genera and species of cicadas occurring in KwaZulu-Natal was drawn up. The identification of many species of cicadas relies on characteristics of the males' sound-producing organs and genitalia.…”