Molecular phylogenetics, diversification, and systematics of Tibicen Latreille 1825 and allied cicadas of the tribe Cryptotympanini, with three new genera and emphasis on species from the USA and Canada<br />(Hemiptera: Auchenorrhyncha: Cicadidae)

Abstract

North America has a diverse cicada fauna with multiple genera from all three Cicadidae subfamilies, yet molecular phylogenetic analyses have been completed only for the well-studied periodical cicadas (Magicicada Davis). The genus Tibicen Latreille, a large group of charismatic species, is in need of such work because morphological patterns suggest multiple groups with complicated relationships to other genera in the tribe Cryptotympanini. In this paper we present a molecular phylogenetic analysis, based on mitochondrial and nuclear DNA, of 35 of the 38 extant USA species and subspecies of the genus Tibicen together with their North American tribal allies (Cornuplura Davis, Cacama Davis), selected Tibicen species from Eurasia, and representatives of other Eurasian and Pacific cryptotympanine genera. This tree shows that Tibicen contains several well-supported clades, one predominating in eastern and central North America and related to Cryptotympana Stål and Raiateana Boulard, another in western North America related to Cacama and Cornuplura, and at least two clades in Eurasia. We also present a morphological cladistic analysis of Tibicen and its close allies based on 27 characters. Character states identified in the cladistic analysis define three new genera, two for North American taxa (Hadoa gen. n. and Neotibicen gen. n.) including several Mexican species, and one for Asian species (Subsolanus gen. n.). Using relaxed molecular clocks and literature-derived mtDNA rate estimates, we estimate the timeframe of diversification of Tibicen clades and find that intergeneric divergence has occurred since the late Eocene, with most extant species within the former Tibicen originating after the mid-Miocene. We review patterns of ecology, behavior, and geography among Tibicen clades in light of the phylogenetic results and note that the study of these insects is still in its early stages. Some Mexican species formerly placed in Tibicen are here transferred to Diceroprocta, following refinement of the definition of that genus.

A new Neotibicen cicada subspecies (Hemiptera: Cicadidae) from the southeastern USA forms hybrid zones with a widespread relative despite a divergent male calling song

“…1 and Results). Although the "narrowly V-shaped" eighth sternite described by Lee is more acutely angled than in other Neotibicen, it is close to that found in N. davisi harnedi (Davis) and N. davisi davisi (Smith and Grossbeck) (see also Hill et al 2015). Sanborn and Heath (2017) have noted as well that intraspecific variation in the shape of the eighth sternite causes problems for generic definitions based on single specimens.…”

“…Other than the highly distinctive uncus, which was noted by earlier authors (Davis 1922;Hill et al 2015;Smith & Grossbeck 1907), the genitalic characters identified by Lee are subtle and unlikely to be applied consistently (see pygofer images in Supplementary Fig. 1).…”

“…Very soon after the publication of Paratibicen, Sanborn and Heath (2017) returned similaris to Neotibicen, noting that the molecular dataset of Hill et al (2015) does not consistently support similaris as the sister lineage to the remainder of Neotibicen. In addition, Hill et al (2015) include a cladistic morphological tree showing similaris in a derived position within Neotibicen, although this is inconclusive because not all of the characters referenced by Lee were included.…”

“…In addition, Hill et al (2015) include a cladistic morphological tree showing similaris in a derived position within Neotibicen, although this is inconclusive because not all of the characters referenced by Lee were included.…”

Taxonomy as a hypothesis: testing the status of the Bermuda buckeye butterfly Junonia coenia bergi (Lepidoptera: Nymphalidae)

“…These data suggest that J. coenia bergi has a North American geographic origin and is a descendant of mainland populations of J. coenia coenia . In this regard, J. coenia bergi is similar to many other endemic species, such as the Bermuda skink ( Plestiodon longirostris ) (Brandley et al , ), Bermuda cicada ( Tibicen bermudiana ) (Hill et al , ), Bermuda cedar ( Juniperus bermudiana ) (Guo et al , ), and Bermuda palmetto ( Sabal bermudana ) (Heyduk et al , ), all of which appear to have colonized Bermuda from the North American mainland. That being said, at least one endemic species, the Bermuda petrel ( Pterodroma cahow ), appears to be more closely related to species from the Caribbean and the islands of the eastern Atlantic than to species from the North American mainland (Welch et al , ).…”