Off-target capture data, endosymbiont genes and morphology reveal a relict lineage that is sister to all other singing cicadas

Phylogenetic asymmetry is common throughout the tree of life and results from contrasting patterns of speciation and extinction in the paired descendant lineages of ancestral nodes. On the depauperate side of a node, we find extant ´relict´ taxa that sit atop long, unbranched lineages. Here, we show that a tiny, pale green, inconspicuous and poorly known cicada in the genus Derotettix, endemic to degraded salt-plain habitats in arid regions of central Argentina, is a relict lineage that is sister to all other modern cicadas. Nuclear and mitochondrial phylogenies of cicadas inferred from probe-based genomic hybrid capture data of both target and non-target loci and a morphological cladogram support this hypothesis. We strengthen this conclusion with genomic data from one of the cicada nutritional bacterial endosymbionts, Sulcia, an ancient and obligate endosymbiont of the larger plant-sucking bugs (Auchenorrhyncha) and an important source of maternally inherited phylogenetic data. We establish Derotettiginae subfam. nov. as a new, monogeneric, fifth cicada subfamily, and compile existing and new data on the distribution, ecology and diet of Derotettix. Our consideration of the palaeoenvironmental literature and host-plant phylogenetics allows us to predict what might have led to the relict status of Derotettix over 100 Myr of habitat change in South America.Fil: Simon, Chris. University of Connecticut; Estados UnidosFil: Gordon, Eric R. L.. University of Connecticut; Estados UnidosFil: Moulds, M.S.. Australian Museum Research Institute; AustraliaFil: Cole, Jeffrey A.. Pasadena City College; Estados UnidosFil: Haji, Diler. University of Connecticut; Estados UnidosFil: Lemmon, Alan R.. Florida State University; Estados UnidosFil: Lemmon, Emily Moriarty. Florida State University; Estados UnidosFil: Kortyna, Michelle. Florida State University; Estados UnidosFil: Nazario, Katherine. University of Connecticut; Estados UnidosFil: Wade, Elizabeth J.. Curry College. Department of Natural Sciences and Mathematics; Estados Unidos. University of Connecticut; Estados UnidosFil: Meister, Russell C.. University of Connecticut; Estados UnidosFil: Goemans, Geert. University of Connecticut; Estados UnidosFil: Chiswell, Stephen M.. National Institute of Water and Atmospheric Research; Nueva ZelandaFil: Pessacq, Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Centro de Investigación Esquel de Montaña y Estepa Patagónica. Universidad Nacional de la Patagonia "San Juan Bosco". Centro de Investigación Esquel de Montaña y Estepa Patagónica; ArgentinaFil: Veloso, Claudio. Universidad de Chile; ChileFil: McCutcheon, John P.. University of Montana; Estados UnidosFil: Lukasik, Piotr. University of Montana; Estados Unidos. Swedish Museum of Natural History. Department of Bioinformatics and Genetics; Sueci

Figs 1-2 in Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Figs 1-2. Euthemopsaltria laeta sp. n.: (1) live male, lateral view; (2) live female, lateral view. Photos Stan and Kaisa Breeden.Published as part of <i>Moulds, M.S., 2014, Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland, pp. 177-190 in Australian Entomologist 41 (4)</i> on page 179, DOI: <a href="http://zenodo.org/record/8290212">10.5281/zenodo.8290212</a&gt

Fig. 11 in Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Fig. 11. Map of northeastern Queensland centred around Cairns, showing distribution of Euthemopsaltria laeta sp. n.; black dots indicate known localities.Published as part of <i>Moulds, M.S., 2014, Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland, pp. 177-190 in Australian Entomologist 41 (4)</i> on page 183, DOI: <a href="http://zenodo.org/record/8290212">10.5281/zenodo.8290212</a&gt

A revision of the Australian cicada genus Punia Moulds, 2012 (Cicadidae Cicadettinae: Cicadettini) with the description of four new species

Moulds, M.S. (2020): A revision of the Australian cicada genus Punia Moulds, 2012 (Cicadidae Cicadettinae: Cicadettini) with the description of four new species. Zootaxa 4860 (1): 101-115, DOI: https://doi.org/10.11646/zootaxa.4860.1.

Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Moulds, M.S. (2014): Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland. Australian Entomologist 41 (4): 177-190, DOI: 10.5281/zenodo.829021

Figs 3-8 in Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Figs 3-8. Euthemopsaltria laeta sp. n., male: (3) abdomen, ventral view; (4) forewing; (5) hind wing; (6) timbal; (7) aedeagus, lateral view; (8) basal plate, dorsal view, apex at bottom. op operculum; ti timbal; ty tympanum; I-VII numbered sternites.Published as part of <i>Moulds, M.S., 2014, Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland, pp. 177-190 in Australian Entomologist 41 (4)</i> on page 181, DOI: <a href="http://zenodo.org/record/8290212">10.5281/zenodo.8290212</a&gt

Fig. 12 in Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Fig. 12. One of two equally parsimonious trees from a cladistic analysis derived using the procedure described in the text above (length 42, CI 73, RI 78) from an analysis employing Lembeja vitticollis as outgroup, with all characters unordered and equally weighted. Numbers at nodes are bootstrap values greater than 50% from 1,000 replications. Character transformations are shown at each node: black bars = nonhomoplasious forward change; grey bars = homoplasious forward change; white bars = reversal (whether homoplasious or not).Published as part of <i>Moulds, M.S., 2014, Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland, pp. 177-190 in Australian Entomologist 41 (4)</i> on page 186, DOI: <a href="http://zenodo.org/record/8290212">10.5281/zenodo.8290212</a&gt

Figs 9-10 in Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland

Figs 9-10. Euthemopsaltria laeta sp. n., male genitalia: (9) lateral view; (10) ventral view with claspers spread apart. cl clasper; pyg pygofer; th theca of aedeagus.Published as part of <i>Moulds, M.S., 2014, Euthemopsaltria laeta, a remarkable new genus and species of cicada (Homoptera: Cicadidae: Cicadettinae: Chlorocystini) from Queensland, pp. 177-190 in Australian Entomologist 41 (4)</i> on page 182, DOI: <a href="http://zenodo.org/record/8290212">10.5281/zenodo.8290212</a&gt