TY - JOUR
AU - Łukasik,, Piotr
AB - Abstract 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. Amaranthaceae, anchored hybrid enrichment, Argentina, Derotettiginae, Derotettix, hybrid capture bycatch, palaeobiology, phylogenomics, Sulcia, South America INTRODUCTION Phylogenetic tree asymmetry is a phenomenon that has captivated evolutionary biologists for a number of reasons. Some biologists focus on the expectation of asymmetrical trees in phylogenetic tree construction (Raup et al., 1973; Farris, 1976; Kirkpatrick & Slatkin, 1993; Mooers & Heard, 1997; Blum & François, 2006); others search for ‘key innovations’ that might have resulted in descendant lineages that differ dramatically in species richness (Sanderson & Donoghue, 1994; Ree, 2005; Rabosky et al., 2007; Nicholson et al., 2014; Simões et al., 2016; Branstetter et al., 2017), and still others focus on the long unbranched lineages, or relict taxa [e.g. horseshoe crabs (Lamsdell, 2013, 2016), coelacanths (Takezaki & Nishihara, 2016), tuataras (Jones et al., 2009), ginkgoes (Wang et al., 2017) and spotted wren babblers (Alström et al., 2014)]. Long branches in asymmetrical phylogenies can result from trivial but common causes, such as lack of taxon sampling (Hedtke et al., 2006), or from evolutionary processes, such as extinction of species (Crisp & Cook, 2005), different rates of cladogenesis in different lineages (e.g. Ellis & Oakley, 2016; Janicke et al., 2018), long periods of time when cladogenesis does not happen (Vaux et al., 2016) or combinations of the above. By studying depauperate lineages, we may learn as much about the ability of species to adapt to changing climates and landscapes as by studying more species-rich lineages (Rabosky, 2017). Cases of phylogenetically asymmetrical living diversity are found in many well-studied taxonomic groups; for example, caecilians (212 species), which split from the remaining extant amphibians > 350 Mya and make up only 3% of current amphibian diversity (Amphibiaweb, 2019; Roelants et al., 2007). Among insects, ancient depauperate relict lineages are also common, e.g. the myxophagan and archostematan beetles (McKenna et al., 2015), myerslopiid leaf hoppers (Hamilton, 1999; Dietrich et al., 2017) and Gondwanan moss bugs (Coleorrhyncha; Yoshizawa et al., 2017). Examples from cicadas include the two extant species of hairy cicadas (Tettigarctidae) that are sister to ~3000 living species of singing cicadas (Cicadidae) (Moulds, 2018; Kaulfuss & Moulds, 2015) and, in the cicada subfamily Cicadettinae, the now depauperate tribe Pictilini (four known species), which at ~60 Mya split from the tribe Cicadettini (500 species; Marshall et al., 2016). Asymmetry is not restricted to deep time. A more recent example is two single species of New Zealand shade-singing cicada lineages in the genus Kikihia (Cicadettinae), which ~6–7 Mya split successively from the remaining 28 Kikihia species and subspecies (Marshall et al., 2008, 2011; Ellis et al., 2015; Banker et al., 2017). The same asymmetry can be found in hundreds of other clades of animals. The study organism Cicadas (Box 1) belong to the superfamily Cicadoidea that, along with the spittle bug superfamily Cercopoidea, make up the sap-feeding bug infraorder Cicadomorpha in the suborder Auchenorrhyncha (large plant-sucking bugs) of the order Hemiptera. The age of Auchenorrhyncha can be traced by fossils to 250 Mya and by molecular dating to > 300 Mya (Misof et al., 2014; Johnson et al., 2018). Cicadoidea comprise two families: the largely extinct hairy cicadas (Tettigarctidae, one extant genus) and the modern, singing cicadas (Cicadidae, ~450 genera) (Marshall et al., 2018). Fossils of hairy cicadas are rare in the Cenozoic geological record but relatively abundant in the Mesozoic and date back to 200 Mya (Shcherbakov, 2009; Moulds, 2018; Lambkin, 2019). The fossil record of the family Cicadidae places modern cicadas with some doubt in the Cretaceous (~99 Mya) and with certainty in the Palaeocene (~59.2–56 Mya; Moulds, 2018). BOX 1. Cicadas occupy a broad range of habitats and are distributed on all continents except Antarctica (Marshall et al., 2018). Cicadas are unique among non-diapausing Hemiptera in having a typical life cycle (egg to adult) that, with few exceptions, spans 3 years or more (Table S3 in Campbell et al. [2015]). Life cycles longer than 1 year allow the development of synchronized episodic life cycles (Duffels, 1988; Heliövaara et al., 1994; Lehmann-Ziebarth et al., 2005; Hajong & Yaakop, 2013; Sota et al., 2013; Chatfield-Taylor & Cole, 2017; Cooley et al., 2018). Despite the difficulty of captive rearing, cicadas offer useful study systems by virtue of their acoustic sexual signals, ease of collection and widespread distribution. Songs of cicadas are highly species specific and facilitate rapid gathering of distributional data and identification of cryptic species (e.g. Marshall & Cooley, 2000; Puissant & Sueur, 2001; Marshall et al., 2011; Hertach et al., 2016). Short-lived adults are known for low dispersal rates (e.g. Duffels, 1988; de Boer & Duffels, 1996; Duffels & Turner, 2002) and high levels of phylogeographical structure within species (e.g. Hill et al., 2009; Marshall et al., 2009; Ellis et al., 2015; Hertach et al., 2016; Liu et al., 2018). Although dispersal rates are generally low, over the span of tens of millions of years, occasional long-distance dispersal has resulted in colonization of distant islands and continents worldwide (Arensburger et al., 2004; Marshall et al., 2016). Low dispersal also enhances the utility of geological events as meaningful calibrations for molecular clocks (e.g. Buckley & Simon, 2007; Marshall et al., 2016; Owen et al., 2017). In addition, drought and cold tolerance (e.g. Toolson, 1987; Sanborn et al., 1995) has equipped cicadas to persist through challenging environmental shifts (Buckley & Simon, 2007; Marshall et al., 2009, 2012; Owen et al., 2017). An aspect of cicada biology that has recently attracted attention is their interaction with heritable nutritional endosymbiotic microorganisms, which provides independent insights into the phylogenetic relationships among the cicada hosts and also offers a unique window into the genomic evolutionary processes related to symbiosis (Van Leuven et al., 2014; Campbell et al., 2015, 2017; Łukasik et al., 2018; Matsuura et al., 2018). Cicadas have been studied taxonomically since the time of Linnaeus, and their subfamily structure has been a subject of continuous debate, having gone through at least seven substantial revisions (Fig. 1) since Distant’s (1906) original classification scheme. Many of the early classification schemes were based on convergent characters associated with sound-producing structures, including covered or uncovered ribbed timbal membranes, resonating chambers of various morphologies, stridulatory organs and wings designed for snapping. Marshall et al. (2018) produced the first scheme based on both molecular and morphological phylogenetic data, but despite their sampling 46 of the 53 tribes, there are still gaps in our understanding of the evolutionary history of cicadas. Figure 1. Open in new tabDownload slide Historical shifts in the number of families and subfamilies in Cicadidae classification, excluding Tettigarctidae (updated from Moulds, 2005; Goemans 2016; Marshall et al., 2018). *Plautilla (Plautillinae) is strongly supported as a member of the Cicadinae (Goemans, 2016; Marshall et al., 2018). Figure 1. Open in new tabDownload slide Historical shifts in the number of families and subfamilies in Cicadidae classification, excluding Tettigarctidae (updated from Moulds, 2005; Goemans 2016; Marshall et al., 2018). *Plautilla (Plautillinae) is strongly supported as a member of the Cicadinae (Goemans, 2016; Marshall et al., 2018). Here, we broaden the taxonomic and environmental scope of our worldwide survey of cicadas with the addition of a tiny, pale green, inconspicuous cicada, Derotettix mendosensis Berg, 1882, which we found living in degraded salt-plain habitats in the ‘Monte de Llanuras y Mesettas’ (plateaus/plains) and the ‘Dry Chaco’ regions of central Argentina (Fig. 2) (Pometti et al., 2012). A recent mitochondrial phylogeny of > 100 members of the family Cicadidae (Łukasik et al., 2019) suggested that Derotettix (represented by two living Argentine species) might be the only surviving genus in a lineage that is the sister group to all other subfamilies in the family Cicadidae. By exploiting reduced representation genome sequencing, we were able to generate nuclear, mitochondrial and symbiont phylogenies to test this hypothesis. Along with a morphological phylogeny, also presented here, these data support and strengthen this sister-group relationship and support the monophy of all other subfamilies sensuMarshall et al. (2018). We create a new, fifth, cicada subfamily, Derotettiginae subfam. nov., which we propose split from the rest of Cicadidae ~100–60 Mya in the transition between the Mesozoic and Cenozoic eras. We discuss the factors that might have led to the relict status of Derotettix over 100 Myr of habitat change in South America. Figure 2. Open in new tabDownload slide Provinces and ecoregions of Argentina redrawn from Pometti et al. (2012), with Derotettix localities coloured by collection year and collectors (see key). Those new to the present study are detailed in Table 1. Sanborn & Heath’s (2014 records include data from five personal expeditions and examination of 11 major relevant museum collections, including three major museums in Argentina and the National History Museum, London. Two additional localities are taken from Torres (1945) but are not exact because he lists only the names of cities or villages. All records are Derotettix mendosensis unless noted as Derotettix wagneri in the key. Figure 2. Open in new tabDownload slide Provinces and ecoregions of Argentina redrawn from Pometti et al. (2012), with Derotettix localities coloured by collection year and collectors (see key). Those new to the present study are detailed in Table 1. Sanborn & Heath’s (2014 records include data from five personal expeditions and examination of 11 major relevant museum collections, including three major museums in Argentina and the National History Museum, London. Two additional localities are taken from Torres (1945) but are not exact because he lists only the names of cities or villages. All records are Derotettix mendosensis unless noted as Derotettix wagneri in the key. MATERIAL AND METHODS Taxon sampling, song recording and analysis From 7 to 23 December 2015, we surveyed Derotettix localities and documented individual cicadas with photographs (14 individuals photographed from San Juan, Rio Negro and Neuquén provinces, Argentina; Fig. 2; Table 1; Supporting Information, Figs S1–S5). We recorded songs from two populations (See Box 2). From 6 to 10 January 2018, we collected D. mendosensis specimens from two localities in Mendoza province, Argentina (Fig. 2; Table 1; Supporting Information, Figs S6–S8) by locating individual males from their songs and capturing them in nets or by hand. We captured females opportunistically, often near males. Specimens collected in 2018 were preserved in 95% ethanol or RNAlater and stored on wet ice for 2 weeks before laboratory storage at −20 °C. We deposited vouchers of specimens collected in 2018 in the collections of M. S. Moulds (Kuranda, Queensland, Australia) and the Department of Ecology and Evolutionary Biology, University of Connecticut Biodiversity Collections (Storrs, CT, USA). In total, we collected and exported ten Derotettix specimens (eight males and two females) from Mendoza province, Argentina. We took data for other cicada species used in our phylogenetic studies from Marshall et al. (2018) and from bycatch from an anchored hybrid enrichment study of the family Cicadidae, in progress (see ‘Sample preparation, sequencing and data handling’, below). Table 1. Derotettix mendosensis localities new to this paper (mapped in Fig. 2). Sample label Date Site name Latitude Longitude Elevation (m) 18.AR.MZ.CLG 10 January 2018 Calle Lugones* -33.513802 -69.065634 910 18.AR.MZ.EVT 6 January 2018 East of Villa Tulimaya† -32.726388 -68.564623 600 PL767 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL769 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL754 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL755 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL756 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL757 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL758 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL954 (song) 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL955 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL752 22 December 2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL753 22 December /2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL618 (song) 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL623 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL624 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 Sample label Date Site name Latitude Longitude Elevation (m) 18.AR.MZ.CLG 10 January 2018 Calle Lugones* -33.513802 -69.065634 910 18.AR.MZ.EVT 6 January 2018 East of Villa Tulimaya† -32.726388 -68.564623 600 PL767 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL769 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL754 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL755 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL756 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL757 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL758 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL954 (song) 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL955 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL752 22 December 2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL753 22 December /2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL618 (song) 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL623 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL624 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 *Just off of Ruta Provincial 96, Mendoza, AR, corner of Calle Lugones. †Highway 34, East of Villa Tulumaya, Mendoza, AR. ‡Side of Ruta Provincial l 6, 23 km NW from Rincón de Los Sauces, Neuquén, AR. §Side of Ruta Nacional 151, 2 km north of Sgto. Vidal, Río Negro, AR. ‖Side of Ruta Nacional 250, 33 km south of the village of Pomona, Río Negro, AR. ¶Side of Ruta Nacional 22 between Choele Choel & Gral Roca, Río Negro, AR. **Road 141 between Bermejo and Marayes, San Juan, AR. Open in new tab Table 1. Derotettix mendosensis localities new to this paper (mapped in Fig. 2). Sample label Date Site name Latitude Longitude Elevation (m) 18.AR.MZ.CLG 10 January 2018 Calle Lugones* -33.513802 -69.065634 910 18.AR.MZ.EVT 6 January 2018 East of Villa Tulimaya† -32.726388 -68.564623 600 PL767 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL769 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL754 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL755 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL756 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL757 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL758 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL954 (song) 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL955 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL752 22 December 2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL753 22 December /2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL618 (song) 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL623 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL624 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 Sample label Date Site name Latitude Longitude Elevation (m) 18.AR.MZ.CLG 10 January 2018 Calle Lugones* -33.513802 -69.065634 910 18.AR.MZ.EVT 6 January 2018 East of Villa Tulimaya† -32.726388 -68.564623 600 PL767 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL769 29 December 2015 NW de Rincón de Los Sauces‡ -37.26473333 -69.07856667 664 PL754 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL755 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL756 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL757 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL758 28 December 2015 Sgto. Vidal§ -38.65226667 -68.13903333 322 PL954 (song) 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL955 23 December 2015 Ruta de Pomona‖ -39.665793 -65.482162 265 PL752 22 December 2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL753 22 December /2015 Ruta a Choele Choel¶ -39.08821667 -66.38621667 355 PL618 (song) 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL623 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 PL624 7 December 2015 San Juan ruta 141** -31.55763333 -67.43775 554 *Just off of Ruta Provincial 96, Mendoza, AR, corner of Calle Lugones. †Highway 34, East of Villa Tulumaya, Mendoza, AR. ‡Side of Ruta Provincial l 6, 23 km NW from Rincón de Los Sauces, Neuquén, AR. §Side of Ruta Nacional 151, 2 km north of Sgto. Vidal, Río Negro, AR. ‖Side of Ruta Nacional 250, 33 km south of the village of Pomona, Río Negro, AR. ¶Side of Ruta Nacional 22 between Choele Choel & Gral Roca, Río Negro, AR. **Road 141 between Bermejo and Marayes, San Juan, AR. Open in new tab BOX 2. Taxonomic description of Derotettiginae subfam. nov. and Derotettigini tribe nov. Subfamily Derotettiginae Moulds subfam. nov. Type genus: Derotettix Berg, 1882 (type species Derotettix mendosensis Berg, 1882). Included tribes:Derotettigini Moulds tribe nov. Diagnosis: Metanotum partially exposed at dorsal midline (Supporting Information, Fig. S12). Forewing veins CuP and 1A unfused, adjacent for two-thirds of length but widely diverging in distal third. Hindwing veins RP and M unfused at their bases (Fig. 6). Male opercula rounded, reduced, enclosing tympanal cavity but not meeting. Abdominal timbal cavity lacking timbal covers. Pygofer with distal shoulder undeveloped; pygofer upper lobe absent. Claspers absent. Aedeagus restrained by tubular encapsulation below uncus, with ventrobasal pocket present; basal plate reduced on more than its basal half to form a pair of long, slender lateral arms attached to theca by sinuation (Fig. 7). The unique morphology of the male genitalia might be a feature of this subfamily rather than a feature of the tribe Derotettigini or Derotettix. The theca is loosely hinged with the basal plate at the extremities of the lateral projections of both structures, as in Tettigarcta (Tettigarctidae). The endotheca enters the somewhat flattened theca beneath a short dorsal overhang at its proximal end. At its distal end, the theca attaches to the thickened membranous vesica in an area weakly membranous, meaning that there is some flexibility between the two. At rest, the nearly straight distal half of the vesica is held within a groove along the ventral surface of the uncus. Distinguishing features: With the following combination of features: forewing veins CuP and 1A and hindwing veins RP and M unfused (Fig. 6); aedeagus with a ventrobasal pocket present and a basal plate deeply divided basally and attached to the theca by sinuation (compare Fig. 7A–C with F–H). Distribution: Neotropics: Argentina. Dry Chaco and Monte de Llanuras y Mesettas ecoregions. Comments: The shape of the basal plate and its attachment to the theca by sinuation are unique among Cicadidae but are features also found in the Tettigarctidae. All other cicadas have the basal plate undivided, as illustrated by Tibicina Kolenati, 1857 (Fig. 7I–L), subfamily Tibicininae. Tribe Derotettigini Moulds tribe nov. Type genus: Derotettix Berg, 1882 (type species Derotettix mendosensis Berg, 1882). Included genera: Derotettix Berg, 1882. Diagnosis: Head including eyes wider than lateral margins of pronotum, but with supra-antennal plates much wider than distance between supra-antennal plate and eye. Postclypeus shape in transverse cross-section rounded; postclypeal ridges lacking transverse grooves towards distal ends. Pronotal collar narrow, with lateral margins confluent with adjoining sclerites and no lateral tooth. Mesonotum lacking auxiliary sound-producing structures. Forewing pterostigma absent; veins C and R+Sc close together; vein RA1 aligned closely with subcosta (Sc) for its length; vein CuA1 divided by crossvein so that distal portion is longest. Hindwing with anal lobe broad and vein 3A straight, very long and widely separated from wing margin. Foreleg femoral primary spine small and prostrate, lacking auxiliary spines. Hind-coxae lacking a large inner protuberance. Meracanthus broadly rounded. Male opercula not completely encapsulating meracanthus; completely covering tympanal cavity but not meeting. Male abdominal tergites with sides convex in cross-section; tergite 2 larger than tergites 3–7; epipleurites reflexed to ventral surface, without an inward V-shaped kink. Timbals extended below level of wing bases; timbal cavity with a rounded rim. Pygofer with basal lobe moderately developed; dorsal beak absent. Uncus undivided, not retractable within pygofer. Aedeagus with theca broad, almost flat but concave ventrally, lacking appendages; vesica much longer than theca, not retractable; basally with a small sclerotized plate either side; conjunctival claws and pseudoparameres absent; ventral rib of basal plate ill defined, short, fused with surface of basal plate (Figs 6, 7 A–E; Supporting Information, Figs S12–S17). Distinguishing features: The Derotettigini tribe nov. differs from all other tribes in having, in combination, the foreleg femoral primary spine small and prostrate and no auxiliary spines, a male uncus that is not retractable within the pygofer, and an aedeagus that has a very broad, almost flat theca. Seven (rather than eight) apical cells in the forewing might be unique to this genus and tribe, but we cannot be certain. Likewise, there are other unusual features of the wings that would normally be considered generic attributes but might not be relevant here at tribal rank, in particular the very elongate basal cell, thickened costal veins (C, Sc + R), the enlarged sixth apical cell in the forewings and the very wide space between hindwing vein 3A and the wing margin. The Supporting Information (Table S3), updated from Marshall et al. (2018) to include Derotettiginae subfam. nov., compares distinguishing features of all five subfamilies of cicadas. Figure 1, also updated from Marshall et al. (2018), traces the historical shifts in the number of subfamilies of Cicadidae. Genus Derotettix Berg, 1882 Included species: Derotettix mendosensis Berg, 1882; Derotettix wagneri Distant, 1905 (Supporting Information, Figs S13–S17). Ecology: Derotettix mendosensis is found largely in patches of dry, salty soils in the Dry Chaco, Monte de Llanuras y Mesettas ecoregions of Argentina (high plains and plateaus; as defined by Pometti et al., 2012), with one or two specimens located nearby in the Estepa Patagónica and Espinal ecoregions (Fig. 2). The only other species in the genus, D. wagneri, is known from several localities in the Dry Chaco of Santiago de Estero province, Argentina (Fig. 2; (Torres, 1945; Sanborn et al., 2004; Sanborn & Heath, 2014). Derotettix species are found on plants in the Amaranthaceae (Allenrolfea and Heterostachys) and Chenopodiaceae (Atriplex, salt bush; Supporting Information, Fig S1A) typical of alkaline salty soils. Heterostachys is a new host record. These habitats can seemingly be degraded by human activity and still support populations of Derotettix (Supporting Information, Figs S2–S4, S6, S8). Derotettix are cryptically coloured to match their host plants (Fig. 3 insets; Supporting Information, Figs S1–S8). Other cicadas have been found on related fleshy halophytic plants. For example, in the desert southwest of North America two species of the genus Okanagodes are cicadas of a similar colour and found on Atriplex, but they belong to the subfamily Tibicininae; another tibicinine cicada, Babras sonorivox, is found on Allenrolfea in Argentina but is not pale green (Torres, 1945; Sanborn et al., 2004; Sanborn & Heath, 2014). Derotettix has one of the highest known thermal tolerances of any cicada species (Sanborn et al., 2004). Derotettix mendosensis Calling song: Songs were recorded from two populations in Mendoza province during the 2018 field season (N = 6 from site 18.AR.MZ.EVT and N = 4 from site 18.AR.MZ.CLG) between 12.30 and 13.30 h at temperatures that ranged from 30.6 to 36.7 °C. Male Derotettix produced a monotonous buzz in long bouts (~30–45 s). Males were wary and ceased singing upon disturbance but were reluctant to fly, instead relying on crypsis to avoid detection. After disturbance, calling resumed as intermittent bouts of short duration (~6–8 s). No interpopulation differences were found in pulse rate or peak frequency (Welch’s two-sample t-tests, P = 0.643 and P = 0.812, respectively). Two singletons from San Juan and Río Negro provinces recorded during the 2015 field season were compared with the song character distributions estimated from Mendoza province. Single-comparison t-tests (Sokal & Rohlf, 1995: pp. 227–228) could not reject the null hypothesis that the song characters of these specimens were drawn from the same distributions. The following descriptive statistics thus include all recordings (N = 12 from three Argentine provinces). Male D. mendosensis calling songs have a pulse rate of 210.4 ± 9.3 (range 194.8–224.7) s−1 (Supporting Information, Fig. S18a) and a peak frequency of 9.5 ± 0.6 (range 8.6–10.4) kHz (Supporting Information, Fig. S18). Neither pulse rate (generalized linear model, N = 9, P = 0.93) nor peak frequency (P = 0.57) depended on ambient temperature over the range at which our recordings were made. The fact that pulse rate was independent of temperature suggests that males thermoregulate their acoustic behaviour. Genetic data: See GenBank numbers in Results and Discussion, above. Endosymbionts: Derotettix mendosensis, like many but not all cicadas (Matsuura et al., 2018), harbours two obligate endosymbionts: Hodgkinia cicadicola and Sulcia muelleri. The 235 kb genome of Sulcia is one of the smaller genomes observed for cicadas, but the family has not been characterized fully. Similar to many other cicadas (Łukasik et al., 2019), the Hodgkinia of Derotettix comprise cytologically and genetically distinct but complementary lineages: one with the expected genome size of ~144 kb and high coding density; the other at much lower abundance, substantially smaller and not yet assembled fully. Our data suggest that after the split, the resulting Hodgkinia lineages degenerated more asymmetrically than in the previously characterized case of the Hodgkinia of Tettigades undata (Van Leuven et al., 2014), but the mechanisms underlying this asymmetry are unclear. During the 2018 field season, we recorded male calling songs in the field using a digital linear pulse code modulation recorder (model PCM-D50; Sony Corp.) with an integral condenser microphone pair. Set at bit depth 16, a 96 kHz sampling rate and a low cut-off frequency of 75 Hz, this equipment recorded a frequency range of 75 Hz to 40 kHz. In the 2015 field season, we recorded songs with a device (model H4n; Zoom Corp.) set to similar specifications. We analysed recordings in Audacity v.2.1.0 (available at www.audacity.sourceforge.net/) and visualized them with RavenLite v.2.0 (available at www.ravensoundsoftware.com). Before analysis, we used a high-pass filter set to a 1 kHz cut-off frequency and 6 dB roll-off to remove wind and other ambient noises that were not already reduced by the low cut-off frequency setting used during recording. For each recording, we calculated pulse rate manually from a 0.5 s oscillogram window. We measured the peak frequency (i.e. frequency at maximal amplitude) from three randomly selected pulses with a fast Fourier transformation (Hanning window, size 512). We performed statistical analysis using R v.3.5.2 (R Core Team, 2018). Sample preparation, sequencing and data handling We extracted DNA from muscle tissue of two legs of a D. mendosensis specimen (specimen code 18.AR.MZ.EVT.01) with a QIAGEN DNeasy Blood & Tissue kit (QIAGEN, Valencia, CA, USA), augmenting the included protocol with an overnight incubation at 56 °C as described by Marshall et al. (2018). We conducted PCR with EmeraldAmp GT PCR Master Mix (Takara, Shiga Japan) for the genes cox1, cox2, EF1a and the 18S rRNA using the primers and annealing temperatures described by Marshall et al. (2018). The PCR products were electrophoresed on a 1% agarose gel, and excess nucleotides and primers were digested with ExoSAP-IT (USB Corp., Cleveland, OH, USA) before submission to Eurofins Genomics (Louisville, KY, USA) for Sanger sequencing with forward and reverse primers. We visualized, quality trimmed and assembled chromatographs and confirmed accurate protein translation for relevant loci in Geneious (Biomatters Ltd, Auckland, New Zealand). We removed intronic sequences from EF1a. For anchored hybrid enrichment, we dissected bacteriomes under a stereomicroscope from specimens preserved in ethanol and stored at −20 °C. We removed the abdominal sternites from each specimen and excised all the bacteriomes found (minus as much excess cicada tissue as possible) and placed them directly in lysis buffer for DNA extraction. We performed separate DNA extractions of bacteriomes and muscle tissue from one leg for each cicada using the QIAGEN DNeasy Blood & Tissue kit. We used the manufacturer’s instructions but added an overnight incubation at 56 °C and removed RNA with RNase A (QIAGEN). We assessed the quality and quantity of DNA using the Qubit fluorometer v.2.0 (Invitrogen, Carlsbad, CA, USA) and agarose gel electrophoresis. We trialled three different pooling methods and demonstrated that we could sequence endosymbiont (extracted from dissected cicada bacteriome tissue) and cicada DNA simultaneously using a mixture of 0.1% cicada bacteriome DNA and 99.9% cicada leg DNA. Average coverage was 146× for cicada anchored hybrid enrichment loci and 864× for endosymbiont loci. High coverage is needed to compensate for the high variance in cicada-to-endosymbiont DNA ratio (owing to variation in the size of bacteriomes among samples). We prepared Illumina libraries from DNA extracts at the Center for Anchored Phylogenomics (www.anchoredphylogeny.com), following Lemmon et al., (2012) and Prum et al. (2015). More specifically, we sonicated DNA using a Covaris ultrasonicator to a fragment size of 175–325 bp. We then used a Beckman-Coulter FXp liquid handling robot to add universal Illumina adapters with 8 bp indexes. After pooling, we enriched libraries using the anchored hybrid enrichment approach (Lemmon et al., 2012). The targets for enrichment were developed for Paraneoptera by Dietrich et al. (2017), who produced a probe set containing probes representing cicadas, among other lineages. This target set was derived from 941 core loci that were determined previously to be orthologous across Diptera (Young et al., 2016), Holometabola (Niehuis et al., 2012), Arthropoda (Misof et al., 2014) or Neuropteroidea (McKenna & Farrell, 2010; Beutel & McKenna, 2016). Dietrich et al. (2017) scanned for these core loci in 17 genomes and 46 transcriptomes of Paraneoptera. After the sequences obtained were aligned and filtered for taxon presence, we designed probes from 514 target loci (total target size = 151 944 bp), including ten genes (dnaE, dnaK, fusA, groL, mnmA, prfA, rpoA, rpoB, rpoC and tufA) of the obligate heritable cicada endosymbiont Candidatus Sulcia muelleri (hereafter, Sulcia) to assess whether the phylogeny of this bacterium mirrored that of the host genome. Agilent Technologies produced the probe kit, which included 55 700 probes. We captured a total of 515 anchored cicada loci, which ranged from 342 to 969 bp in length. The anchored loci themselves will be used in a future publication that includes increased taxon sampling. To achieve the most complete possible phylogenetic dataset incorporating global cicada diversity, we supplemented existing Sanger-sequenced data for the 28S, 18S, EF1a, ARD1, cox1 and cox2 genes from Marshall et al. (2018) by mining the capture assemblies of the same or closely related individuals for loci that were previously missing and were likely to have been a part of capture bycatch owing to a high genomic copy number (18S rRNA, cox1 and cox2). In addition, we supplemented this dataset with the 28S rRNA, which, although off target, was also frequently recoverable in capture assemblies. We deduplicated both merged and unpaired reads from the capture library sequencing using clumpify in the BBMap suite (Bushnell, 2014) and trimmed them of TruSeq adaptor and low-quality (Quality Score < 20) sequences with Trimmomatic (Bolger et al., 2014). We assembled the resulting trimmed reads using SPAdes v.3.12.0 (Nurk et al., 2013). For host genes, we queried capture assemblies with blastn (18S and 28S rRNA) or tblastn (cox1 and cox2) using Magicicada references on GenBank (MG953107.1 and KM000130.1) or the 28S rRNA of an unidentified cicada (JQ309936.1) as the query sequence. We aligned matching contigs back to the query sequence in Geneious v.10.1.3 and stitched them together if they consisted of two or more contigs. We processed captured loci for Sulcia and the 28S rRNA in a similar manner, then implemented additional processing using iterative read mapping with MIRA v.4.0.2 (Chevreux et al., 1999). We then used MITObim v.1.9.1 (Hahn et al., 2013), an additional read mapper that produced slightly better results, on the MIRA-corrected bait sequences, which we edited by manual trimming of apparent misassembled or duplicated segments. We aligned Sulcia and host 28S rRNA loci using the MAFFT v.7 E-INS-i algorithm (Katoh et al., 2017), and we trimmed alignments of apparently misassembled or duplicated segments further. We constructed individual unpartitioned Sulcia gene trees using RAxML v.8 (Stamatakis, 2014) on the CIPRES web server (Miller et al., 2010) to check and remove sequence data that showed evidence of cross-contamination based on similarity to sequence data from distantly related taxa. The methods used to acquire and assemble the metagenome of a D. mendosensis specimen, PL623x1, are described by Łukasik et al. (2018, 2019). Briefly, we extracted DNA from the dissected bacteriome after fragmenting it using Covaris, prepped following a modified protocol by Meyer & Kircher (2010) and sequenced on the Illumina HiSeq 4000 platform. We assembled the reads using Spades v.3.7.1. We obtained the cicada ARD1 and 28S rRNA sequence in addition to all Sulcia loci for Derotettix by querying this assembly with blastn or by using a query sequence from a relative as a seed for MITObim on the trimmed paired end reads. Phylogenetic analysis We aligned cicada and Sulcia loci using the MAFFT v.7 E-INS-i algorithm (Katoh et al., 2017) and inspected and trimmed them in Geneious v.10.1.3 based on amino acid translations for protein-encoding loci. We used SequenceMatrix (Vaidya et al., 2011) to concatenate loci. We created partitioning schemes based on codon positions of each protein-encoding gene and separate partitions for the two rRNA genes and the 5′ untranslated region of ARD1 and analysed them using PartitionFinder v.1.0.1 (Lanfear et al., 2012) with the greedy search algorithm and the best combination of possible partitions chosen by the Bayesian information criterion. We generated a maximum likelihood tree using this partitioning scheme with RAxML v.8 (Stamatakis, 2014) on the CIPRES web server (Miller et al., 2010), with 1000 rapid bootstrap replicates. We visualized trees in FigTree v.1.4.0 (Rambaut & Drummond, 2012) and edited with ggtree (Yu et al., 2017). Morphological cladistic analysis For the morphological cladistic analysis, we used the same 117-character dataset as Moulds (2005). We scored D. mendosensis for these characters and added them to the dataset along with scores for Tettigomyia vespiformis Amyot & Serville, 1843 to ensure that all subfamilies were represented. See Moulds (2005) for a full description of these characters and character states. We analysed the data using the heuristic search parsimony algorithms in PAUP* v.4.0b2 (Swofford, 1998). We used the tree bisection–reconnection algorithm for tree searches and conducted 1000 random additional searches starting from random trees; we left other settings at their default values. We weighted all characters equally and treated all multistate characters as unordered. We found the most resolved trees by filtering the set of shortest trees using the Filter Trees option. We prepared the chosen tree using CLADOS v.1.2 (Nixon, 1992) with DELTRAN optimization. We dissected male genitalia needed for study and illustration from relaxed adults by cutting the intersegmental membrane holding the pygofer (often also along with sternite 8); we then cleared the genitalia in 10% KOH at room temperature for ~4–8 h, with the length of time depending on the degree of sclerotization of the genitalia. After removing the genitalia from the KOH, we washed them thoroughly in water. Using a stereomicroscope, with the genitalia submerged in a Petri dish of water, we removed excess intersegmental membrane from the pygofer hind-margin and then removed any internal undissolved muscle tissue (dark matter). When closer examination of the aedeagus was required, we separated it from the pygofer by cutting the translucent membrane surrounding the theca and pulling the aedeagus backwards. Descriptions of new subfamily and new tribe The terminology for morphological features follows that of Moulds (2005, 2012). The relevance of characters in defining higher taxa follows the cladistic analysis of Moulds (2005). A discussion of the song, endosymbionts, ecology and biogeography of Derotettix is included in the formal description in the Results and Discussion (Box 2). RESULTS AND DISCUSSION Exploitation of bycatch in sequence capture data enables integration with existing datasets The data acquired via sequence capture experiments typically allow for robust phylogenomic analyses based on hundreds of preselected loci (Bi et al., 2013; Blaimer et al., 2016; McCormack et al., 2016). The success of enrichment of these selected loci varies depending on the phylogenetic distance of sampled species to those for which probes are designed (Bragg et al., 2016; Kieran et al., 2019) and many other factors, such as the amount and integrity of target DNA in individual samples, probe tiling depth and whether probes are synthesized as RNA or DNA (Gasc et al., 2016). For studies in which successful captures of species within clades dating to 100–200 Mya have been performed, the ranges of on-target reads have been reported to be anywhere from ~10 to ~60% on average across the entire dataset, with rates of only up to 80% on-target reads for species from which the probes were designed (Schott et al., 2017; Knyshov et al., 2019). Owing to the imperfect nature of hybridization of targeted DNA, additional loci may also be recoverable from naïve assemblies of reads from capture experiments given adequate sequencing depth. In particular, we found that high-copy number genes, including those on the mitochondrion and those found as part of the rRNA operon, were frequently recoverable from non-target reads. Given that these multi-copy genes happen to be ones that were first selected as commonly used phylogenetic markers because they allowed relatively easy PCR amplification, we can integrate newly sampled taxa meaningfully with datasets collected previously that encompass much wider sampling. Such non-target bycatch data have begun to be exploited for systematic studies only in recent times (Guo et al., 2012; Gasc et al., 2016; Lyra et al., 2017; Barrow et al., 2017; Caparroz et al., 2018; Matsuura et al., 2018; Taucce et al., 2018; Percy et al., 2018; Łukasik et al., 2019), but continued use of this valuable, albeit hidden, resource will help to resolve the tree of life by allowing more complete sampling in phylogenies. Note that we were particularly successful at obtaining these data from the bycatch because of the relatively low enrichment efficiency of anchored hybrid enrichment loci (between 1 and 6% of reads map to target anchored hybrid enrichment loci), which is attributable to the large size of the cicada genome (Hanrahan & Johnston, 2011) and its diversity. Mitochondrial DNA genomes can be more difficult to obtain from bycatch for systems in which the size of the genome is small and/or the probes are designed for taxa with less variation (i.e. when > 50% of reads map to target loci). The complete mitochondrial genome of D. mendosensis (minus the control region) was sequenced as genomic bycatch from exon capture (Łukasik et al., 2019; GenBank no. MG737807.1). Nuclear and mitochondrial metadata and gene segments used in the present study (28S, 18S, Ef1a and ARD1; cox1 and cox2) can be found at GenBank numbers MN241535-MN241813). Sulcia gene sequences can be found at GenBank numbers MN219733-MN219984. Details of each Genbank submission by species and gene are given in Tables S1 and S2. Multifaceted evidence for a monogeneric subfamily Before our work, there were four Cicadidae subfamilies. Three of these, Cicadettinae (worldwide), Cicadinae (worldwide) and Tettigomyiinae (Africa + Madagascar), appear to have split from each other close together in time (Marshall et al., 2018). The fourth subfamily (Tibicininae) is the sister group to the other three, as shown in our trees (Figs 3–5; Supporting Information, Figs S9–S11) and by Marshall et al. (2018). Our results demonstrate that four datasets [nuclear gene (Supporting Information, Fig. S9), mitochondrial genome (Łukasik et al., 2019), Sulcia endosymbiont genes (Fig. 4) and morphological data (Fig. 5)] all strongly support the hypothesis that the genus Derotettix is sister to all these subfamilies. Derotettix is also strongly supported as sister to the rest of Cicadidae in our nuclear plus mitochondrial DNA phylogeny (Fig. 3), in the phylogeny built with 28S data alone (Supporting Information, Fig. S10) and in a tree made with all genetic data combined (Supporting Information, Fig. S11). All trees also strongly support Tibicininae as sister to the remaining three subfamilies: Tettigomyiinae, Cicadettinae and Cicadinae. Maximum likelihood bootstrap support is strong (98–100%) for the monophyly of each subfamily except Cicadinae, for which bootstrap support varied from unresolved in 28S alone to 69–89% in the other dataset combinations. Figure 3. Open in new tabDownload slide A, RAxML phylogeny, RNA + codon partitioned, nuclear (28S, 18S, EF1a and ARD1) plus mitochondrial DNA data. Of 8819 total characters, 2123 are parsimony informative; for the ingroup only, 1787 characters are parsimony informative. B, Derotettix mendosensis (PL954), Ruta de Pomona, Provincia Rio Negro, Argentina with fingertip for scale (photograph: P.Ł.). C, D, D. mendosensis green and yellow colour morphs, respectively, both from site 18.AR.MZ.EVT, on Heterostachys (see Table 1) (photographs: J.A.C.). Figure 3. Open in new tabDownload slide A, RAxML phylogeny, RNA + codon partitioned, nuclear (28S, 18S, EF1a and ARD1) plus mitochondrial DNA data. Of 8819 total characters, 2123 are parsimony informative; for the ingroup only, 1787 characters are parsimony informative. B, Derotettix mendosensis (PL954), Ruta de Pomona, Provincia Rio Negro, Argentina with fingertip for scale (photograph: P.Ł.). C, D, D. mendosensis green and yellow colour morphs, respectively, both from site 18.AR.MZ.EVT, on Heterostachys (see Table 1) (photographs: J.A.C.). Figure 4. Open in new tabDownload slide RAxML phylogeny, codon partitioned, Sulcia endosymbiont genes (dnaE, dnaK, fusA, groL, mnmA, prfA, rpoA, rpoB, rpoC and tufA). Of 22 804 sites, 1400 are parsimony informative, 899 for the ingroup. Inset photograph borders match the colour of subfamily tree branches. A, Tettigarcta tomentosa (Tettigarctidae), Tasmania (photograph: Simon Grove, Tasmanian Museum & Art Gallery). B, Derotettix mendosensis (Derotettiginae subfam. nov.), Argentina (photograph: C.S.). C, Platypedia sp. (Tibicininae), Arizona (photograph: David Marshall). D, Stagira sp. (Tettigomyiinae), Uganda (photograph: Nick Dean). E, Kikihia ochrina (Cicadettinae), New Zealand (photograph: C.S.). F, Gaeana maculata (Cicadinae), Hong Kong (photograph: Ray Li). Figure 4. Open in new tabDownload slide RAxML phylogeny, codon partitioned, Sulcia endosymbiont genes (dnaE, dnaK, fusA, groL, mnmA, prfA, rpoA, rpoB, rpoC and tufA). Of 22 804 sites, 1400 are parsimony informative, 899 for the ingroup. Inset photograph borders match the colour of subfamily tree branches. A, Tettigarcta tomentosa (Tettigarctidae), Tasmania (photograph: Simon Grove, Tasmanian Museum & Art Gallery). B, Derotettix mendosensis (Derotettiginae subfam. nov.), Argentina (photograph: C.S.). C, Platypedia sp. (Tibicininae), Arizona (photograph: David Marshall). D, Stagira sp. (Tettigomyiinae), Uganda (photograph: Nick Dean). E, Kikihia ochrina (Cicadettinae), New Zealand (photograph: C.S.). F, Gaeana maculata (Cicadinae), Hong Kong (photograph: Ray Li). Figure 5. Open in new tabDownload slide One of 716 most parsimonious trees (length 313, consistency index [CI] 55, rescaled consistency index [RI] 87) derived using the morphological dataset from Moulds (2005), with Derotettix and Tettigomyia included; hence, all subfamilies are represented. Differences between the 716 trees were confined to terminal taxa within Cicadinae and Cicadettinae, meaning that character transformations for more basal nodes shown on the tree were identical for all trees. Character transformations on branches are represented as follows: black bars, non-homoplasious forward change (unique); grey bars, homoplasious forward change (a shared state); white bars, reversal (whether homoplasious or not). Bootstrap values are shown for branches supporting subfamilies. Figure 5. Open in new tabDownload slide One of 716 most parsimonious trees (length 313, consistency index [CI] 55, rescaled consistency index [RI] 87) derived using the morphological dataset from Moulds (2005), with Derotettix and Tettigomyia included; hence, all subfamilies are represented. Differences between the 716 trees were confined to terminal taxa within Cicadinae and Cicadettinae, meaning that character transformations for more basal nodes shown on the tree were identical for all trees. Character transformations on branches are represented as follows: black bars, non-homoplasious forward change (unique); grey bars, homoplasious forward change (a shared state); white bars, reversal (whether homoplasious or not). Bootstrap values are shown for branches supporting subfamilies. Figure 6. Open in new tabDownload slide Derotettix mendosensis left fore- and hindwing, with veins labelled. Abbreviations: a, apical cells; A, anal; bc, basal cell; C, costa; CuA, cubitus anterior; CuP, cubitus posterior; M, median; R, radius; Ra, radius anterior; rc, radial cell; RP, radius posterior; Sc, subcostal. Derotettix mendosensis wing photograph: E.R.L.G. (using Macropod camera, Macroscopic Solutions). Figure 6. Open in new tabDownload slide Derotettix mendosensis left fore- and hindwing, with veins labelled. Abbreviations: a, apical cells; A, anal; bc, basal cell; C, costa; CuA, cubitus anterior; CuP, cubitus posterior; M, median; R, radius; Ra, radius anterior; rc, radial cell; RP, radius posterior; Sc, subcostal. Derotettix mendosensis wing photograph: E.R.L.G. (using Macropod camera, Macroscopic Solutions). Figure 7. Open in new tabDownload slide Distinguishing morphological features of Derotettiginae subfam. nov. and comparisons of genital characters of Derotettiginae with Tettigarctidae and Tibicininae. Clockwise from upper right: A–E, Derotettix mendosensis pygofer (genital capsule), lateral view (A); pygofer, ventral view (B); aedeagus (dissected) (C); front leg (D); timbal (E); F–H, Tettigarcta crinita Distant pygofer, lateral view (F); pygofer, ventral view (G); aedeagus (H); I–L, Tibicina haematodes Scopoli pygofer, lateral view (I); pygofer, ventral view (J); aedeagus (K); basal plate (L). C, H, K, views of aedeagi of the three representative species. Figure 7. Open in new tabDownload slide Distinguishing morphological features of Derotettiginae subfam. nov. and comparisons of genital characters of Derotettiginae with Tettigarctidae and Tibicininae. Clockwise from upper right: A–E, Derotettix mendosensis pygofer (genital capsule), lateral view (A); pygofer, ventral view (B); aedeagus (dissected) (C); front leg (D); timbal (E); F–H, Tettigarcta crinita Distant pygofer, lateral view (F); pygofer, ventral view (G); aedeagus (H); I–L, Tibicina haematodes Scopoli pygofer, lateral view (I); pygofer, ventral view (J); aedeagus (K); basal plate (L). C, H, K, views of aedeagi of the three representative species. The nuclear gene tree (Supporting Information, Fig. S9) includes four gene segments: EF1a, ARD1, 28S and 18S. The 28S gene was not included in the study by Marshall et al. (2018) but proved to be informative in our phylogeny at the deeper nodes (Supporting Information, Fig. S10). This locus, consisting of a total of 4622 sites (629 informative within Cicadidae), is able to resolve the relevant relationships of Derotettix with respect to the rest of cicadas and the relationship of the Tibicinae as sister to the remaining three subfamilies with 100% bootstrap support but loses resolution shallower in the tree (Supporting Information, Fig. S10). Nuclear anchored hybrid enrichment genomic data analyses (C. Owen, D. Marshall, E. J. Wade, R. C. Meister, G. Goemans, K. B. R. Hill, A. R. Lemmon, E. M. Lemmon, M. Kortyna, M. S. Moulds, V. Sarkar, K. Marathe, K. Kunte, C. Simon, unpublished observations) are predicted to strengthen support for the monophyly of the subfamily Cicadinae and to resolve shallower nodes in the tree. Auchenorrhyncha were ancestrally associated with one or more obligate bacterial endosymbionts that produced essential amino acids and vitamins and were transmitted faithfully through the female reproductive system to subsequent generations (Moran et al., 2005). One of them, Sulcia, has been retained by the majority of Auchenorrhyncha lineages, including all cicadas characterized to date. Unlike the second ancestral endosymbiont of cicadas, Candidatus Hodgkinia cicadicola (hereafter, Hodgkinia) (McCutcheon et al., 2009a, b), Sulcia evolves in a relatively slow manner and is easy to align across Cicadidae (Campbell et al., 2015), making it useful for phylogenetic reconstructions (e.g. Matsuura et al., 2018).Our maximum likelihood phylogeny of ten conserved Sulcia genes supports and strengthens the conclusions of the nuclear DNA, mitochondrial DNA and morphological data, but with Cicadinae, Cicadettinae and Tettigomyiinae represented as a trichotomy. The three subfamilies that were represented by multiple taxa were all clearly monophyletic. Our morphological tree for 82 taxa (plus one outgroup; Fig. 5) adds Derotettix and Tettigomyia to the 117-character dataset used by Moulds (2005) (Supporting Information, Table S4) and also supports the hypothesis that Derotettix is sister to the other cicada subfamilies and should be placed in a new subfamily, Derotettiginae. Although clade support for Derotettiginae on the morphological tree was not strong, owing to its single non-homoplasious synapomorphy being outweighed by five homoplasious (shared) synapomorphies, the clade support for the remaining subfamilies of Cicadidae to the exclusion of Derotettiginae was strong (82% bootstrap). Despite Derotettiginae having only one non-homoplasious synapomorphy, it does have six shared attributes in unique combination, a situation not unusual for more basal nodes in large morphological analyses of groups with reasonably conservative morphology (e.g. Cicadinae has only one non-homoplasious synapomorphy and five shared attributes in unique combination; see Supporting Information, Table S4). Hiding in plain sight Derotettix was described in 1882 but has never been singled out as unusual. Thus, given the long history of work on higher-level cicada taxonomy (Fig. 1), the resolution of D. mendosensis as sister to all other species in the family Cicadidae (Łukasik et al., 2019) was unexpected. The tribe Parnisini, to which Derotettix had been assigned, includes 23 genera, of which 13 genera are restricted to the Ethiopian biogeographical realm (four of those are endemic to Madagascar); two genera are found in both the Ethiopian and Palaearctic realms; one genus is restricted to the Palaearctic; and finally, five genera (including Derotettix) are found only in the Neotropical realm. Marshall et al. (2018) reviewed the tribes and subfamilies of the family Cicadidae, including four genera previously classified as Parnisini. The Neotropical genera Parnisa Stål, 1862 and Calyria Stål, 1862 were retained in this tribe, but Quintilia Stål, 1866 was moved into the new African subfamily Tettigomyiinae; Arcystasia Distant, 1882 was found to belong to Cicadettini in an earlier work (Marshall et al., 2016) but not formally reassigned. Marshall et al. (2018) were not able to review the other 19 parnisine genera and noted that the tribe Parnisini needs further revision. Our present study does not include parnisines outside of the Neotropics; therefore, a complete evaluation of the members of this tribe awaits further sampling and future genomic studies. Marshall et al. (2018) questioned the make-up of tribes that, like Parnisini, have deep, seemingly global distributions (e.g. Chlorocystini, Cryptotympanini and Taphurini) and removed some taxa; future studies might remove more. However, global tribes do exist. The subfamily Cicadettinae contains two well-sampled tribes with Northern, Southern, Eastern and Western Hemisphere components: Cicadettini (Marshall et al., 2016) and Lamotialnini (Marshall et al., 2018). Parnisini (minus Derotettix) might turn out to be another widely distributed cicadettine tribe. It is not impossible that future taxon sampling will turn up other lineages that branch deep in the cicada tree. Such candidates could come from genera in two poorly characterized tribes that have been found to be polyphyletic, i.e. Parnisini or Taphurini, or from as yet unsampled genera. For example, a new South American genus of cicada, Gibbocicada Ruschel, 2018 (Tibicinini, Tibicininae), was described recently from museum material and is the only member of its tribe found in the Southern Hemisphere (Ruschel, 2018). Long branch, little cicada: palaeoclimatic and landscape changes We propose that species extinction is a more likely cause for the lack of other species in Derotettiginae than lack of speciation over millions or tens of millions of years. This hypothesis parallels the situation we see in the Tettigarctidae, the sister lineage to modern singing cicadas. Tettigarctids were diverse throughout the Mesozoic and into the Eocene (from 250 to ~40 Mya; Moulds 2018), but today are represented by only two relatively closely related species that are cold adapted and live in remote mountainous regions of Tasmania and southern New South Wales (Kaulfuss & Moulds, 2015). The derotettigine lineage dates back to the late Cretaceous or earliest Palaeocene. We suggest that former species in this lineage went extinct as a result of landscape and climatic changes. The Cenozoic was a time of extensive habitat modification caused by continental movements and changes in sea level and ocean currents that had a profound effect on the global distribution and diversification of plants and animals [e.g. in Australia (Byrne et al., 2018), Africa (Linder & Bouchenak-Khelladi, 2015) and South America (Ortiz-Jaureguizar & Cladera, 2006)]. A general cooling and drying of the Southern Hemisphere starting in the Late Eocene/early Oligocene was triggered by many factors, including the establishment of the Antarctic Circumpolar Current and a reduction in global carbon dioxide (~41–33 Mya; (DeConto & Pollard, 2003; Speelman et al., 2009). Various taxa invaded these newly arid domains around the world (e.g. Rabosky et al., 2007; Kadereit et al., 2012; Woodburne et al., 2014; Owen et al., 2017; Byrne et al., 2018), with plants being aided by the rise of C4 photosynthesis independently in many lineages (Edwards & Smith, 2010; Morando et al., 2014; Zucol et al., 2018). Roig et al. (2009) reviewed the biogeography of Argentina and concluded that the Monte and Chaco ecoregions were savanna at the beginning of the Cenozoic (65 Mya). Despite the wet climate of the Eocene (50–33 Mya), pollen fossils suggest that dry conditions persisted in parts of central western Argentina and increased in extent as the Andes uplifted. Retreats of extensive epicontinental seaways (Supporting Information, Fig. S19) left salt deposits in many parts of South America (Benavides, 1968), most recently in the mid Miocene, when most of the current range of both Derotettix species was under the Paranense Sea (Hernández et al., 2005). The influence of this incursion has been seen in the population structures of many central and northern Argentinian animal species (e.g. Delsuc et al., 2012; Morando et al., 2014; Brusquetti et al., 2019). From the middle Miocene to the present in Argentina, rain-shadow aridity has increased owing to the rise of the Andes over the last 15 Myr, with accelerated uplift in the central Andes ~5 Mya (Farías et al., 2008; Folguera et al., 2011). The resulting climatic changes influenced all groups of flora and fauna (e.g. Ortiz-Jaureguizar & Cladera, 2006; Roig et al., 2009; Ruzzante & Rabassa, 2011; Turchetto-Zolet et al., 2013; Wallis et al., 2016). In the last 2.6 Myr, the region has been affected by a series of no fewer than eight glaciation events that caused major fluctuations in the climate (Rabassa, 2008; Rabassa et al., 2011; Elderfield et al., 2012). Although the derotettigine lineage stretches back >60 Myr, the current host plants of Derotettix are thought to have arrived much later. Allenrolfea and Heterostachys are predicted to have arrived in South America from Eurasia some time in the Miocene, 19 Mya at the earliest (Piirainen et al., 2017). Atriplex arrived in North America in the mid Miocene (14 Mya or later) and moved into South America from there (Kadereit et al., 2010). Salt tolerance, succulence and the evolution of C4 photosynthesis might have preadapted these plant taxa to invade the steppes and deserts of the interior of South America. Derotettix might have arisen via a host shift in the mid Miocene after their host plants arrived (Piirainen et al., 2017). Host shifts in insects are common and often lead to speciation (Forbes et al., 2017). Examples are known from cicadas, including changes in gene expression likely to be associated with a shift in host plant in Subpsaltria yangi from an angiosperm to a gnetophyte (Hou & Wei, 2019). The ability of Derotettix to adapt to saline environments might have been a key innovation that facilitated their survival. The Monte regions of Argentina currently lie in the transition between the tropical biota to the north and the Patagonian biota to the south. During our fieldwork, we observed that the cicadas of this region are a mixture of these northern (tribe Fidicinini-dominant) and southern (tribe Tettigadini-dominant) elements. The rise of cicadas in the subfamily Tibicininae, whose members in the tribe Tettigadini now dominate the temperate habitats of southern South America (Sanborn & Heath, 2014), probably also contributed to the decline of Derotettiginae. The key innovations that are lacking in Derotettiginae but present in Tibicininae are unknown but would be a fruitful area for future research. Age of Cicadidae and a possible South American origin The fossil record (Shcherbakov, 2009; Moulds, 2018) suggests that the modern cicadas (Cicadidae) arose during the late Cretaceous or early Palaeozoic at the latest. This would place the origin of the family Cicadidae no later than 99–60 Mya, during the time of the main Angiosperm radiation. The most comprehensive dating analyses of a major clade of cicadas conducted so far (Marshall et al., 2016; Owen et al., 2017) suggests that the tribe Cicadettini most probably originated around the time of the greenhouse–ice house transition at the end of the Eocene (~41–33 Mya). This tribe is contained within the subfamily Cicadettinae. The subfamily Tibicininae, known from a Palaeocene fossil 59.2–56 Myr old (Moulds 2018), is sister to Cicadettinae plus Cicadinae plus Tettigomyiinae (Figs 3–5). Thus, the Derotettiginae must have split from the rest of Cicadidae before the deposition of this tibicinine late Palaeocene fossil. Formal molecular dating studies using fossilized birth–death methods (Heath et al., 2014), with much larger taxonomic and genomic sampling, are in progress. Given that Derotettiginae is known only from South America and Tibicininae is heavily represented in South America, we hypothesize that the family Cicadidae had a South American origin. Tibicininae is largely New World, with at least two independent amphitropical Northern Hemisphere–Southern Hemisphere clade splits. Tibicininae includes an additional five (out of 23) genera that are endemic to the Palaearctic (Marshall et al., 2018) and closely related to the North American tibicinines (Sueur et al., 2007). Genomic sampling of additional South American and world taxa (in progress) will allow us to test the South American origin hypothesis put forth here for the first time. SUPPORTING INFORMATION Additional supporting information may be found in the online version of this article at the publisher’s website: Table S1. Additional details for cicada genes used in this study. Table S2. Additional details for Sulcia endosymbiont genes used in this study. Table S3. Characters for the five subfamilies of the family Cicadidae, updated from Marshall et al. (2018). Autoapomorphies are highlighted in grey. Note that the Tettigomyiinae and Derotettiginae subfam. nov. each lack an autapomorphy and are diagnosable only by a combination of attributes. Table S4. Character matrix (83 taxa × 117 characters) used in the maximum parsimony morphological character analysis. Missing data and character states not relevant to a taxon are scored as ‘?’. Figure S1. A, Derotettix mendosensis male (PL754) on saltbush, Atriplex sp., Río Negro province, Argentina. B, male, dorsal view. C, male, side view. D, male, ventral view. Photographs: P.Ł. See Table 1 and Figure 2 for exact locality. Figure S2.Derotettix mendosensis male (PL618), Ruta 141, San Juan province, Argentina. Clockwise from upper left: A, side view; B, dorsal view; C, side view; D, head. Photographs: P.Ł. See Table 1 and Figure 2 for exact locality. Figure S3.Derotettix mendosensis (PL755 and PL756), Río Negro province, Argentina. A, female, green morph, dorsal. B, female, green morph, ventral. C, male, yellow morph. D, male, yellow morph, dorsal. E, male, yellow morph, side view. Photographs: P.Ł. See Table 1 and Figure 2 for exact locality. Figure S4.Derotettix mendosensis (PL767), NW de Rincón de Los Sauces, Neuquén province, Argentina. A, male, side view. B, male, ventral view. C, male, dorsal view. Photographs: P.Ł. See Table 1 and Figure 2 for exact locality. Figure S5.Derotettix mendosensis (PL954), Ruta de Pomona, Provincia Rio Negro, Argentina. A, male, dorsal view. B, male, side view. C, male, ventral view. Photographs: P.Ł. See Table 1 and Figure 2 for exact locality. Figure S6.Derotettix mendosensis site (18.AR.MZ.CLG), highway 34, east of Villa Tulumaya, Mendoza province, Argentina. Three views of the habitat of D. mendosensis, with obvious human disturbance. Photographs: C.S. See Table 1 and Figure 2 for exact locality. Figure S7.Derotettix mendosensis, east of Villa Tulumaya, Mendoza province, Argentina (site 18.AR.MZ.EVT). A, female, ventral view. B, female, side view. C, male, side view. D, male, ventral view. Photographs: C.S. See Table 1 and Figure 2 for exact locality. Figure S8.Derotettix mendosensis habitat (18.AR.MZ.CLG), Calle Lugones, just off Provincial Road 96, Mendoza province, Argentina. Habitat with obvious human disturbance. Photographs: C.S. See Table 1 and Figure 2 for exact locality. Figure S9. RAxML phylogeny for nuclear genes only, RNA + codon partitioned (28S, 18S, EF1a and ARD1). Of 6652 total characters, 1034 are parsimony informative; for the Cicadidae ingroup only, 714 are parsimony informative. Figure S10. RAxML phylogeny, 28S gene only. Of 4622 total sites, 721 are parsimony informative, 629 within Cicadidae. Note 100% support on deepest nodes, including Cicadidae, Derotettiginae subfam. nov., Tibicininae and the three remaining subfamilies as a trichotomy (Cicadinae unresolved). Figure S11. RAxML phylogeny, all genetic data combined (nuclear NDA, mitochondrial DNA and Sulcia, RNA + codon partitioned). Of 31 623 total sites, 3523 are parsimony informative, 2686 within Cicadidae. All subfamilies were resolved as monophyletic. Figure S12. Illustration of the subfamily character ‘metanotum at dorsal midline’ with states ‘partially visible’ and ‘completely hidden’. Clockwise from upper left: A, Cicadettinae, Amphipsalta zelandica (photograph: C.S.); B, Derotettiginae subfam. nov., Derotettix mendosensis (photograph: E.R.L.G.); C, Tibicininae, Alarcta micromacula (photograph: C.S.); D, Cicadinae Neotibicen pronotalis (photograph: David C. Marshall). Figure S13.Derotettix mendosensis, 18.AR.MZ.EVT (Table 1). Colour faded by ethanol. A, dorsal view. B, head. C, lateral view. Macropod photographs: E.R.L.G. Figure S14.Derotettix mendosensis male holotype, La Plata. A, dorsal view. B, ventral view. Macropod photographs: K.N. Figure S15.Derotettix mendosensis female allotype, La Plata. A, dorsal view. B, ventral view. Macropod photographs: K.N. Figure S16.Derotettix wagneri (= Derotettix proseni) male holotype, La Plata. A, dorsal view. B, ventral view. Macropod photographs: K.N. Figure S17. A, Derotettix wagneri (= Derotettix proseni) male holotype, lateral view, La Plata. B, Derotettix mendosensis male holotype, lateral view, La Plata. Macropod photographs: K.N. Figure S18. A, Derotettix mendosensis song structure, recording 080110-05, 35.4 °C, 18.AR.MZ.CLG. A, 0.5 s oscillogram (above). B, spectrogram. Figure S19. (A) Paleomap reconstructions redrawn/excerpted as permitted from Scotese, C. R., 2001. Atlas of Earth History, Volume 1, Paleogeography, PALEOMAP Project, Arlington, Texas, 52 pp. Note extensive South American inland sea incursions 80 Mya and again 20 Mya.; (B) Paranense Sea transgression 13-15 Mya, with all Derotettix localities superimposed. Map redrawn from Hernandez et al. 2005, Figure 2. Inset below is Figure 2 from the present paper for comparison. ACKNOWLEDGEMENTS Thanks to Ana Maria Marino de Remes Lenicov and her museum staff at Museo de La Plata for facilitating our examination of type specimens in their collections. Thanks to Argentine permitting authorities and Raúl Adolfo Pessacq for help with permits and to Iris Peralta of the National University of Cuyo, Mendoza, Argentina for identification of Heterostachys. Thanks to Ivan Nozaic, Kyra Kopestonsky, Sally Beech and Virge Kask for assistance with illustrations. We thank Sean Holland and Kirby Birch at the Center for Anchored Phylogenomics for assistance with data collection and analysis. Thanks to David Marshall, Bert Orr and Heinrich Fliedner for advice on nomenclature and to David Marshall and Kathy Hill and various cicada colleagues around the world (listed by Marshall et al., 2018) for help with collection of non-Derotettix taxa. Thanks to Allen Sanborn for information on D. wagneri localities. David Marshall provided valuable discussion on the systematics and ecology of cicadas, a careful reading of the paper and many excellent suggestions. One anonymous reviewer, Tatiana Ruschel, Masami Hayashi and John A. Allen made helpful suggestions that improved the manuscript. Thanks to the Biodiversity Research Collections, University of Connecticut for housing vouchers of specimens used in this publication. Collection, transportation and export of Derotettix specimens was permitted by the province of Mendoza and the Argentine Ministerio de Ambiente Y Desarrollo Sustentable, Dirección de Fauna Silvestre DSN no. 52460852/18. This research was supported by the National Science Foundation (DEB 1655891 and IOS 1553529), the National Geographic Society grant 9760-15, an Ernst Mayr Travel Grant from the Museum of Comparative Zoology (Harvard University) and several grants from the University of Connecticut. REFERENCES Alström P , Hooper DM , Liu Y , Olsson U , Mohan D , Gelang M , Le Manh H , Zhao J , Lei F , Price TD . 2014 . Discovery of a relict lineage and monotypic family of passerine birds . Biology Letters 10 : 20131067 . Google Scholar Crossref Search ADS PubMed WorldCat Amphibiaweb . 2019 . University of California, Berkeley, CA, USA. Available at: https://amphibiaweb.org. Accessed 30 Jul 2019. Arensburger P , Buckley TR , Simon C , Moulds M , Holsinger KE . 2004 . Biogeography and phylogeny of the New Zealand cicada genera (Hemiptera: Cicadidae) based on nuclear and mitochondrial DNA data . Journal of Biogeography 31 : 557 – 569 . Google Scholar Crossref Search ADS WorldCat Banker SE , Wade EJ , Simon C . 2017 . The confounding effects of hybridization on phylogenetic estimation in the New Zealand cicada genus Kikihia . Molecular Phylogenetics and Evolution 116 : 172 – 181 . Google Scholar Crossref Search ADS PubMed WorldCat Barrow LN , Soto-Centeno JA , Warwick AR , Lemmon AR , Lemmon EM . 2017 . Evaluating hypotheses of expansion from refugia through comparative phylogeography of south-eastern Coastal Plain amphibians . Journal of Biogeography 44 : 2692 – 2705 . Google Scholar Crossref Search ADS WorldCat Benavides V . 1968 . Saline deposits of South America. In: Mattox RB , Holser WT , eds. Saline deposits: a symposium based on papers from the International Conference on Saline Deposits, Houston, Texas, 1962 . New York: Geological Society of America , 249 – 290 . Google Preview WorldCat COPAC Beutel RG , McKenna DD . 2016 . Systematic position, basal branching pattern and early evolution. In: Beutel RG , Leschen RAB , eds. Coleoptera, beetles. Morphology and systematics . Berlin: Walter de Gruyter, 1–12 . Google Preview WorldCat COPAC Bi K , Linderoth T , Vanderpool D , Good JM , Nielsen R , Moritz C . 2013 . Unlocking the vault: next-generation museum population genomics . Molecular Ecology 22 : 6018 – 6032 . Google Scholar Crossref Search ADS PubMed WorldCat Blaimer BB , Lloyd MW , Guillory WX , Brady SG . 2016 . Sequence capture and phylogenetic utility of genomic ultraconserved elements obtained from pinned insect specimens . PLoS One 11 : e0161531 . Google Scholar Crossref Search ADS PubMed WorldCat Blum MGB , François O . 2006 . Which random processes describe the tree of life? A large-scale study of phylogenetic tree imbalance . Systematic Biology 55 : 685 – 691 . Google Scholar Crossref Search ADS PubMed WorldCat de Boer AJ , Duffels JP . 1996 . Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific: a geotectonic explanation . Palaeogeography, Palaeoclimatology, Palaeoecology 124 : 153 – 177 . Google Scholar Crossref Search ADS WorldCat Bolger AM , Lohse M , Usadel B . 2014 . Trimmomatic: a flexible trimmer for Illumina sequence data . Bioinformatics 30 : 2114 – 2120 . Google Scholar Crossref Search ADS PubMed WorldCat Boulard M . 1976 . Un type nouveau d’appareil stridulant accessoire pour les Cicadoidea: révision de la classification supérieure de la superfamille [Hom.] . Journal of Natural History 10 : 399 – 407 . Google Scholar Crossref Search ADS WorldCat Bragg JG , Potter S , Bi K , Moritz C . 2016 . Exon capture phylogenomics: efficacy across scales of divergence . Molecular Ecology Resources 16 : 1059 – 1068 . Google Scholar Crossref Search ADS PubMed WorldCat Branstetter MG , Danforth BN , Pitts JP , Faircloth BC , Ward PS , Buffington ML , Gates MW , Kula RR , Brady SG . 2017 . Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees . Current Biology 27 : 1019 – 1025 . Google Scholar Crossref Search ADS PubMed WorldCat Brusquetti F , Netto F , Baldo D , Haddad CFB . 2019 . The influence of Pleistocene glaciations on Chacoan fauna: genetic structure and historical demography of an endemic frog of the South American Gran Chaco . Biological Journal of the Linnean Society 126 : 404 – 416 . Google Scholar Crossref Search ADS WorldCat Buckley TR , Simon C . 2007 . Evolutionary radiation of the cicada genus Maoricicada Dugdale (Hemiptera: Cicadoidea) and the origins of the New Zealand alpine biota . Biological Journal of the Linnean Society 91 : 419 – 435 . Google Scholar Crossref Search ADS WorldCat Bushnell B . 2014 . BBMap: a fast, accurate, splice-aware aligner . In: Lawrence Berkeley National Laboratory . CA: Ernest Orlando Lawrence Berkeley National Laboratory in Berkeley. LBNL-7065E. Retrieved from https://escholarship.org/uc/item/1h3515gn. COPAC Byrne M , Joseph L , Yeates DK , Roberts JD , Edwards D . 2018 . Evolutionary history. In: Lambers H , ed. On the ecology of Australia’s arid zone . Cham : Springer International Publishing , 45 – 75 . Google Preview WorldCat COPAC Campbell MA , Łukasik P , Simon C , McCutcheon JP . 2017 . Idiosyncratic genome degradation in a bacterial endosymbiont of periodical cicadas . Current Biology 27 : 3568 – 3575 .e3. Google Scholar Crossref Search ADS PubMed WorldCat Campbell MA , Van Leuven JT , Meister RC , Carey KM , Simon C , McCutcheon JP . 2015 . Genome expansion via lineage splitting and genome reduction in the cicada endosymbiont Hodgkinia . Proceedings of the National Academy of Sciences of the United States of America 112 : 10192 – 10199 . Google Scholar Crossref Search ADS PubMed WorldCat Caparroz R , Rocha AV , Cabanne GS , Tubaro P , Aleixo A , Lemmon EM , Lemmon AR . 2018 . Mitogenomes of two neotropical bird species and the multiple independent origin of mitochondrial gene orders in Passeriformes . Molecular Biology Reports 45 : 279 – 285 . Google Scholar Crossref Search ADS PubMed WorldCat Chatfield-Taylor W , Cole JA . 2017 . Living rain gauges: cumulative precipitation explains the emergence schedules of California protoperiodical cicadas . Ecology 98 : 2521 – 2527 . Google Scholar Crossref Search ADS PubMed WorldCat Chevreux B , Wetter T , Suhai S . 1999 . Genome sequence assembly using trace signals and additional sequence information . German Conference on Bioinformatics 99 : 45 – 56 . WorldCat Cooley JR , Arguedas N , Bonaros E , Bunker G , Chiswell SM , DeGiovine A , Edwards M , Hassanieh D , Haji D , Knox J , Kritsky G , Mills C , Mozgai D , Troutman R , Zyla J , Hasegawa H , Sota T , Yoshimura J , Simon C . 2018 . The periodical cicada four-year acceleration hypothesis revisited and the polyphyletic nature of Brood V, including an updated crowd-source enhanced map (Hemiptera: Cicadidae: Magicicada) . PeerJ 6 : e5282 . Google Scholar Crossref Search ADS PubMed WorldCat Crisp MD , Cook LG . 2005 . Do early branching lineages signify ancestral traits? Trends in Ecology & Evolution 20 : 122 – 128 . Google Scholar Crossref Search ADS PubMed WorldCat DeConto RM , Pollard D . 2003 . Rapid Cenozoic glaciation of Antarctica induced by declining atmospheric CO2 . Nature 421 : 245 – 249 . Google Scholar Crossref Search ADS PubMed WorldCat Delsuc F , Superina M , Tilak MK , Douzery EJ , Hassanin A . 2012 . Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos . Molecular Phylogenetics and Evolution 62 : 673 – 680 . Google Scholar Crossref Search ADS PubMed WorldCat Dietrich CH , Allen JM , Lemmon AR , Lemmon EM , Takiya DM , Evangelista O , Walden KKO , Grady PGS , Johnson KP . 2017 . Anchored hybrid enrichment-based phylogenomics of leafhoppers and treehoppers (Hemiptera: Cicadomorpha: Membracoidea) . Insect Systematics and Diversity 1 : 57 – 72 . Google Scholar Crossref Search ADS WorldCat Distant WL . 1906 . A synonymic catalogue of Homoptera: Pt. 1. Cicadidae . London : British Museum . Google Preview WorldCat COPAC Duffels JP . 1988 . The cicadas of the Fiji, Samoa and Tonga Islands: their taxonomy and biogeography (Homoptera, Cicadoidea) with a chapter on the geological history of the area by A Ewart. Entomonograph 10: 1–108 . Google Preview WorldCat COPAC Duffels JP , Turner H . 2002 . Cladistic analysis and biogeography of the cicadas of the Indo‐Pacific subtribe Cosmopsaltriina (Hemiptera: Cicadoidea: Cicadidae) . Systematic Entomology 27 : 235 – 261 . Google Scholar Crossref Search ADS WorldCat Edwards EJ , Smith SA . 2010 . Phylogenetic analyses reveal the shady history of C4 grasses . Proceedings of the National Academy of Sciences of the United States of America 107 : 2532 – 2537 . Google Scholar Crossref Search ADS PubMed WorldCat Elderfield H , Ferretti P , Greaves M , Crowhurst S , McCave IN , Hodell D , Piotrowski AM . 2012 . Evolution of ocean temperature and ice volume through the Mid-Pleistocene climate transition . Science 337 : 704 – 709 . Google Scholar Crossref Search ADS PubMed WorldCat Ellis EA , Marshall DC , Hill KBR , Owen CL , Kamp PJJ , Simon C . 2015 . Phylogeography of six codistributed New Zealand cicadas and their relationship to multiple biogeographical boundaries suggest a re-evaluation of the Taupo Line . Journal of Biogeography 42 : 1761 – 1775 . Google Scholar Crossref Search ADS WorldCat Ellis EA , Oakley TH . 2016 . High rates of species accumulation in animals with bioluminescent courtship displays . Current Biology: CB 26 : 1916 – 1921 . Google Scholar Crossref Search ADS PubMed WorldCat Farías M , Charrier R , Carretier S , Martinod J , Fock A , Campbell D , Cáceres J , Comte D . 2008 . Late Miocene high and rapid surface uplift and its erosional response in the Andes of central Chile (33°–35°S) . Tectonics 27 : 1 – 22 . Google Scholar Crossref Search ADS WorldCat Farris JS . 1976 . Expected asymmetry of phylogenetic trees . Systematic Zoology 25 : 196 – 198 . Google Scholar Crossref Search ADS WorldCat Folguera A , Orts D , Spagnuolo M , Vera ER , Litvak V , Sagripanti L , Ramos ME , Ramos VA . 2011 . A review of Late Cretaceous to Quaternary palaeogeography of the southern Andes . Biological Journal of the Linnean Society 103 : 250 – 268 . Google Scholar Crossref Search ADS WorldCat Forbes AA , Devine SN , Hippee AC , Tvedte ES , Ward AKG , Widmayer HA , Wilson CJ . 2017 . Revisiting the particular role of host shifts in initiating insect speciation . Evolution 71 : 1126 – 1137 . Google Scholar Crossref Search ADS PubMed WorldCat Gasc C , Peyretaillade E , Peyret P . 2016 . Sequence capture by hybridization to explore modern and ancient genomic diversity in model and nonmodel organisms . Nucleic Acids Research 44 : 4504 – 4518 . Google Scholar Crossref Search ADS PubMed WorldCat Goemans G . 2016 . The classification and phylogeny of the Neotropical Cicada Tribe Zammarini (Hemiptera, Cicadidae) and a revision of its type genus Zammara Amyot & Audinet Serville, 1843 and its sister genus Zammaralna Boulard & Sueur, 1996 . Unpublished D. Phil. Thesis, University of Connecticut . Google Preview WorldCat COPAC Guo Y , Long J , He J , Li CI , Cai Q , Shu XO , Zheng W , Li C . 2012 . Exome sequencing generates high quality data in non-target regions . BMC Genomics 13 : 194 . Google Scholar Crossref Search ADS PubMed WorldCat Hahn C , Bachmann L , Chevreux B . 2013 . Reconstructing mitochondrial genomes directly from genomic next-generation sequencing reads—a baiting and iterative mapping approach . Nucleic Acids Research 41 : e129 . Google Scholar Crossref Search ADS PubMed WorldCat Hajong SR , Yaakop S . 2013 . Chremistica ribhoi sp. n. (Hemiptera: Cicadidae) from North-East India and its mass emergence . Zootaxa 3702 : 493 – 500 . Google Scholar Crossref Search ADS PubMed WorldCat Hamilton KGA . 1999 . The ground-dwelling leafhoppers Sagmatiini and Myerslopiidae (Rhynchota: Homoptera: Membracoidea) . Invertebrate Taxonomy 13 : 207 – 235 . Google Scholar Crossref Search ADS WorldCat Hanrahan SJ , Johnston JS . 2011 . New genome size estimates of 134 species of arthropods . Chromosome Research 19 : 809 – 823 . Google Scholar Crossref Search ADS PubMed WorldCat Hayashi M . 1984 . A review of the Japanese Cicadidae . Cicada 5 : 25 – 76 . WorldCat Heath TA , Huelsenbeck JP , Stadler T . 2014 . The fossilized birth–death process for coherent calibration of divergence-time estimates . Proceedings of the National Academy of Sciences of the United States of America 111 : E2957 – E2966 . Google Scholar Crossref Search ADS PubMed WorldCat Hedtke SM , Townsend TM , Hillis DM . 2006 . Resolution of phylogenetic conflict in large data sets by increased taxon sampling . Systematic Biology 55 : 522 – 529 . Google Scholar Crossref Search ADS PubMed WorldCat Heliövaara K , Väisänen R , Simon C . 1994 . Evolutionary ecology of periodical insects . Trends in Ecology & Evolution 9 : 475 – 480 . Google Scholar Crossref Search ADS PubMed WorldCat Hernández RM , Jordan TE , Dalenz Farjat A , Echavarría L , Idleman BD , Reynolds JH . 2005 . Age, distribution, tectonics, and eustatic controls of the Paranense and Caribbean marine transgressions in southern Bolivia and Argentina . Journal of South American Earth Sciences 19 : 495 – 512 . Google Scholar Crossref Search ADS WorldCat Hertach T , Puissant S , Gogala M , Trilar T , Hagmann R , Baur H , Kunz G , Wade EJ , Loader SP , Simon C , Nagel P . 2016 . Complex within a complex: integrative taxonomy reveals hidden diversity in Cicadetta brevipennis (Hemiptera: Cicadidae) and unexpected relationships with a song divergent relative . PLoS One 11 : e0165562 . Google Scholar Crossref Search ADS PubMed WorldCat Hill KBR , Simon C , Marshall DC , Chambers GK . 2009 . Surviving glacial ages within the Biotic Gap: phylogeography of the New Zealand cicada Maoricicada campbelli. Journal of Biogeography 36 : 675 – 692 . Google Scholar Crossref Search ADS WorldCat Hou Z , Wei C . 2019 . De novo comparative transcriptome analysis of a rare cicada, with identification of candidate genes related to adaptation to a novel host plant and drier habitats . BMC Genomics 20 : 182 . Google Scholar Crossref Search ADS PubMed WorldCat Janicke T , Ritchie MG , Morrow EH , Marie-Orleach L . 2018 . Sexual selection predicts species richness across the animal kingdom . Proceedings of the Royal Society B: Biological Sciences 285 : 20180173 . Google Scholar Crossref Search ADS WorldCat Johnson KP , Dietrich CH , Friedrich F , Beutel RG , Wipfler B , Peters RS , Allen JM , Petersen M , Donath A , Walden KKO , Kozlov AM , Podsiadlowski L , Mayer C , Meusemann K , Vasilikopoulos A , Waterhouse RM , Cameron SL , Weirauch C , Swanson DR , Percy DM , Hardy NB , Terry I , Liu S , Zhou X , Misof B , Robertson HM & Yoshizawa K . 2018 . Phylogenomics and the evolution of hemipteroid insects . Proceedings of the National Academy of Sciences of the United States of America 115 : 12775 – 12780 . Google Scholar Crossref Search ADS PubMed WorldCat Jones MEH , Tennyson AJD , Worthy JP , Evans SE , Worthy TH . 2009 . A sphenodontine (Rhynchocephalia) from the Miocene of New Zealand and palaeobiogeography of the tuatara (Sphenodon) . Proceedings of the Royal Society B: Biological Sciences 276 : 1385 – 1390 . Google Scholar Crossref Search ADS WorldCat Kadereit G , Ackerly D , Pirie MD . 2012 . A broader model for C4 photosynthesis evolution in plants inferred from the goosefoot family (Chenopodiaceae s.s.) . Proceedings of the Royal Society B: Biological Sciences 279 : 3304 – 3311 . Google Scholar Crossref Search ADS WorldCat Kadereit G , Mavrodiev EV , Zacharias EH , Sukhorukov AP . 2010 . Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis . American Journal of Botany 97 : 1664 – 1687 . Google Scholar Crossref Search ADS PubMed WorldCat Kato M . 1954 . On the classification of Cicadoidea (Homoptera: Auchenorrhyncha) . Kontyû 21 : 97 – 100 . WorldCat Katoh K , Rozewicki J , Yamada KD . 2017 . MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization . Briefings in Bioinformatics 1–7. doi:10.1093/bib/bbx108 . WorldCat Kaulfuss U , Moulds MS . 2015 . A new genus and species of tettigarctid cicada from the early Miocene of New Zealand: Paratettigarcta zealandica (Hemiptera, Auchenorrhyncha, Tettigarctidae) . ZooKeys 484 : 83 – 94 . Google Scholar Crossref Search ADS WorldCat Kieran TJ , Gordon ERL , Forthman M , Hoey-Chamberlain R , Kimball RT , Faircloth BC , Weirauch C , Glenn TC . 2019 . Insight from an ultraconserved element bait set designed for hemipteran phylogenetics integrated with genomic resources . Molecular Phylogenetics and Evolution 130 : 297 – 303 . Google Scholar Crossref Search ADS PubMed WorldCat Kirxpatrick M , Slatkin M . 1993 . Searching for evolutionary patterns in the shape of a phylogenetic tree . Evolution 47 : 1171 – 1181 . Google Scholar Crossref Search ADS PubMed WorldCat Knyshov A , Gordon ERL , Weirauch C . 2019 . Cost-efficient high throughput capture of museum arthropod specimen DNA using PCR-generated baits . Methods in Ecology and Evolution 10 : 841 – 852 . Google Scholar Crossref Search ADS WorldCat Lambkin KJ . 2019 . Mesodiphthera Tillyard, 1919, from the Late Triassic of Queensland, the oldest cicada (Hemiptera: Cicadomorpha: Cicadoidea: Tettigarctidae) . Zootaxa 4567 : 358 – 366 . Google Scholar Crossref Search ADS WorldCat Lamsdell JC . 2013 . Revised systematics of Palaeozoic ‘horseshoe crabs’ and the myth of monophyletic Xiphosura: re-evaluating the Monophyly of Xiphosura . Zoological Journal of the Linnean Society 167 : 1 – 27 . Google Scholar Crossref Search ADS WorldCat Lamsdell JC . 2016 . Horseshoe crab phylogeny and independent colonizations of fresh water: ecological invasion as a driver for morphological innovation . Palaeontology 59 : 181 – 194 . Google Scholar Crossref Search ADS WorldCat Lanfear R , Calcott B , Ho SY , Guindon S . 2012 . PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses . Molecular Biology and Evolution 29 : 1695 – 1701 . Google Scholar Crossref Search ADS PubMed WorldCat Lehmann-Ziebarth N , Heideman PP , Shapiro RA , Stoddart SL , Hsaio CCL , Stephenson GR , Milewski PA , Ives AR . 2005 . Evolution of periodicity in periodical cicadas . Ecology 86 : 3200 – 3211 . Google Scholar Crossref Search ADS WorldCat Lemmon AR , Emme SA , Lemmon EM . 2012 . Anchored hybrid enrichment for massively high-throughput phylogenomics . Systematic Biology 61 : 727 – 744 . Google Scholar Crossref Search ADS PubMed WorldCat Linder HP , Bouchenak-Khelladi Y . 2015 . The causes of southern African spatial patterns in species richness: speciation, extinction and dispersal in the Danthonioideae (Poaceae) . Journal of Biogeography 42 : 914 – 924 . Google Scholar Crossref Search ADS WorldCat Liu Y , Qiu Y , Wang XU , Yang H , Hayashi M , Wei C . 2018 . Morphological variation, genetic differentiation and phylogeography of the East Asia cicada Hyalessa maculaticollis (Hemiptera: Cicadidae) . Systematic Entomology 43 : 308 – 329 . Google Scholar Crossref Search ADS WorldCat Łukasik P , Chong RA , Nazario K , Matsuura Y , Bublitz AC , Campbell MA , Meyer MC , Van Leuven JT , Pessacq P , Veloso C , Simon C , McCutcheon JP . 2019 . One hundred mitochondrial genomes of cicadas . The Journal of Heredity 110 : 247 – 256 . Google Scholar Crossref Search ADS PubMed WorldCat Łukasik P , Nazario K , Van Leuven JT , Campbell MA , Meyer M , Michalik A , Pessacq P , Simon C , Veloso C , McCutcheon JP . 2018 . Multiple origins of interdependent endosymbiotic complexes in a genus of cicadas . Proceedings of the National Academy of Sciences of the United States of America 115 : E226 – E235 . Google Scholar Crossref Search ADS PubMed WorldCat Lyra ML , Joger U , Schulte U , Slimani T , El Mouden EH , Bouazza A , Künzel S , Lemmon AR , Lemmon EM , Vences M . 2017 . The mitochondrial genomes of Atlas Geckos (Quedenfeldtia): mitogenome assembly from transcriptomes and anchored hybrid enrichment datasets . Mitochondrial DNA Part B 2 : 356 – 358 . Google Scholar Crossref Search ADS WorldCat Marshall DC , Cooley JR . 2000 . Reproductive character displacement and speciation in periodical cicadas, with description of new species, 13-year Magicicada neotredecem . Evolution 54 : 1313 – 1325 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Hill KB , Cooley JR , Simon C . 2011 . Hybridization, mitochondrial DNA phylogeography, and prediction of the early stages of reproductive isolation: lessons from New Zealand cicadas (genus Kikihia) . Systematic Biology 60 : 482 – 502 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Hill KB , Fontaine KM , Buckley TR , Simon C . 2009 . Glacial refugia in a maritime temperate climate: cicada (Kikihia subalpina) mtDNA phylogeography in New Zealand . Molecular Ecology 18 : 1995 – 2009 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Hill KB , Marske KA , Chambers C , Buckley TR , Simon C . 2012 . Limited, episodic diversification and contrasting phylogeography in a New Zealand cicada radiation . BMC Evolutionary Biology 12 : 177 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Hill KBR , Moulds M , Vanderpool D , Cooley JR , Mohagan AB , Simon C . 2016 . Inflation of molecular clock rates and dates: molecular phylogenetics, biogeography, and diversification of a global cicada radiation from Australasia (Hemiptera: Cicadidae: Cicadettini) . Systematic Biology 65 : 16 – 34 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Moulds M , Hill KBR , Price BW , Wade EJ , Owen CL , Goemans G , Marathe K , Sarkar V , Cooley JR , Sanborn AF , Kunte K , Villet MH , Simon C . 2018 . A molecular phylogeny of the cicadas (Hemiptera: Cicadidae) with a review of tribe and subfamily classification . Zootaxa 4424 : 1 – 64 . Google Scholar Crossref Search ADS PubMed WorldCat Marshall DC , Slon K , Cooley JR , Hill KB , Simon C . 2008 . Steady Plio-Pleistocene diversification and a 2-million-year sympatry threshold in a New Zealand cicada radiation . Molecular Phylogenetics and Evolution 48 : 1054 – 1066 . Google Scholar Crossref Search ADS PubMed WorldCat Matsuura Y , Moriyama M , Łukasik P , Vanderpool D , Tanahashi M , Meng XY , McCutcheon JP , Fukatsu T . 2018 . Recurrent symbiont recruitment from fungal parasites in cicadas . Proceedings of the National Academy of Sciences of the United States of America 115 : E5970 – E5979 . Google Scholar Crossref Search ADS PubMed WorldCat McCormack JE , Tsai WL , Faircloth BC . 2016 . Sequence capture of ultraconserved elements from bird museum specimens . Molecular Ecology Resources 16 : 1189 – 1203 . Google Scholar Crossref Search ADS PubMed WorldCat McCutcheon JP , McDonald BR , Moran NA . 2009a . Convergent evolution of metabolic roles in bacterial co-symbionts of insects . Proceedings of the National Academy of Sciences of the United States of America 106 : 15394 – 15399 . Google Scholar Crossref Search ADS WorldCat McCutcheon JP , McDonald BR , Moran NA . 2009b . Origin of an alternative genetic code in the extremely small and GC-rich genome of a bacterial symbiont . PLoS Genetics 5 : e1000565 . Google Scholar Crossref Search ADS WorldCat McKenna DD , Farrell BD . 2010 . 9-genes reinforce the phylogeny of holometabola and yield alternate views on the phylogenetic placement of Strepsiptera . PLoS One 5 : e11887 . Google Scholar Crossref Search ADS PubMed WorldCat McKenna DD , Wild AL , Kanda K , Bellamy CL , Beutel RG , Caterino MS , Farnum CW , Hawks DC , Ivie MA , Jameson ML , Leschen RAB , Marvaldi AE , McHugh JV , Newton AF , Robertson JA , Thayer MK , Whiting MF , Lawrence JF , Slipinski A , Maddison DR , Farrell BD . 2015 . The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution . Systematic Entomology 40 : 835 – 880 . Google Scholar Crossref Search ADS WorldCat Metcalf ZP . 1963 . General catalogue of the Homoptera. Fascicle VIII Cicadoidea. Part 1 Cicadidae . Raleigh : University of North Carolina State College . Google Preview WorldCat COPAC Meyer M , Kircher M . 2010 . Illumina sequencing library preparation for highly multiplexed target capture and sequencing . Cold Spring Harbor Protocols 2010 : pdb.prot5448 . Google Scholar Crossref Search ADS PubMed WorldCat Miller MA , Pfeiffer W , Schwartz T . 2010 . Creating the CIPRES Science gateway for inference of large phylogenetic trees . Gateway Computing environments workshop (GCE) . New Orleans, LA: IEEE. Misof B , Liu S , Meusemann K , Peters RS , Donath A , Mayer C , Frandsen PB , Ware J , Flouri T , Beutel RG , Niehuis O , Petersen M , Izquierdo-Carrasco F , Wappler T , Rust J , Aberer AJ , Aspöck U , Aspöck H , Bartel D , Blanke A , Berger S , Böhm A , Buckley TR , Calcott B , Chen J , Friedrich F , Fukui M , Fujita M , Greve C , Grobe P , Gu S , Huang Y , Jermiin LS , Kawahara AY , Krogmann L , Kubiak M , Lanfear R , Letsch H , Li Y , Li Z , Li J , Lu H , Machida R , Mashimo Y , Kapli P , McKenna DD , Meng G , Nakagaki Y , Navarrete-Heredia JL , Ott M , Ou Y , Pass G , Podsiadlowski L , Pohl H , von Reumont BM , Schütte K , Sekiya K , Shimizu S , Slipinski A , Stamatakis A , Song W , Su X , Szucsich NU , Tan M , Tan X , Tang M , Tang J , Timelthaler G , Tomizuka S , Trautwein M , Tong X , Uchifune T , Walzl MG , Wiegmann BM , Wilbrandt J , Wipfler B , Wong TK , Wu Q , Wu G , Xie Y , Yang S , Yang Q , Yeates DK , Yoshizawa K , Zhang Q , Zhang R , Zhang W , Zhang Y , Zhao J , Zhou C , Zhou L , Ziesmann T , Zou S , Li Y , Xu X , Zhang Y , Yang H , Wang J , Wang J , Kjer KM , Zhou X . 2014 . Phylogenomics resolves the timing and pattern of insect evolution . Science 346 : 763 – 767 . Google Scholar Crossref Search ADS PubMed WorldCat Mooers AO , Heard SB . 1997 . Inferring evolutionary process from phylogenetic tree shape . The Quarterly Review of Biology 72 : 31 – 54 . Google Scholar Crossref Search ADS WorldCat Moran NA , Tran P , Gerardo NM . 2005 . Symbiosis and insect diversification: an ancient symbiont of sap-feeding insects from the bacterial phylum Bacteroidetes . Applied and Environmental Microbiology 71 : 8802 – 8810 . Google Scholar Crossref Search ADS PubMed WorldCat Morando M , Medina CD , Avila LJ , Perez CHF , Buxton A , Sites JW Jr . 2014 . Molecular phylogeny of the New World gecko genus Homonota (Squamata: Phyllodactylidae) . Zoologica Scripta 43 : 249 – 260 . Google Scholar Crossref Search ADS WorldCat Moulds MS . 2005 . An appraisal of the higher classification of cicadas (Hemiptera: Cicadoidea) with special reference to the Australian fauna . Records of the Australian Museum 57 : 375 – 446 . Google Scholar Crossref Search ADS WorldCat Moulds MS . 2012 . A review of the genera of Australian cicadas (Hemiptera: Cicadoidea) . Zootaxa 3287 : 1 – 262 . Google Scholar Crossref Search ADS WorldCat Moulds MS . 2018 . Cicada fossils (Cicadoidea: Tettigarctidae and Cicadidae) with a review of the named fossilised Cicadidae . Zootaxa 4438 : 443 – 470 . Google Scholar Crossref Search ADS PubMed WorldCat Myers JG . 1929 . Insect singers: a natural history of the cicadas . London : George Routledge and Sons . Google Preview WorldCat COPAC Nicholson DB , Ross AJ , Mayhew PJ . 2014 . Fossil evidence for key innovations in the evolution of insect diversity . Proceedings of the Royal Society B: Biological Sciences 281 : 20141823 . Google Scholar Crossref Search ADS WorldCat Niehuis O , Hartig G , Grath S , Pohl H , Lehmann J , Tafer H , Donath A , Krauss V , Eisenhardt C , Hertel J , Petersen M , Mayer C , Meusemann K , Peters RS , Stadler PF , Beutel RG , Bornberg-Bauer E , McKenna DD , Misof B . 2012 . Genomic and morphological evidence converge to resolve the enigma of Strepsiptera . Current Biology: CB 22 : 1309 – 1313 . Google Scholar Crossref Search ADS PubMed WorldCat Nixon KC . 1992 . Clados, version 1.2 . Ithaca : L.H. Bailey Hortorium, Cornell University . Google Preview WorldCat COPAC Nurk S , Bankevich A , Antipov D , Gurevich AA , Korobeynikov A , Lapidus A , Prjibelski AD , Pyshkin A , Sirotkin A , Sirotkin Y , Stepanauskas R , Clingenpeel SR , Woyke T , McLean JS , Lasken R , Tesler G , Alekseyev MA , Pevzner PA . 2013 . Assembling single-cell genomes and mini-metagenomes from highly chimeric reads. In: Research in computational molecular biology . Berlin and Heidelberg : Springer , 158 – 170 . Google Preview WorldCat COPAC Ortiz-Jaureguizar E , Cladera GA . 2006 . Paleoenvironmental evolution of southern South America during the Cenozoic . Journal of Arid Environments 66 : 498 – 532 . Google Scholar Crossref Search ADS WorldCat Owen CL , Marshall DC , Hill KBR , Simon C . 2017 . How the aridification of Australia structured the biogeography and influenced the diversification of a large lineage of Australian cicadas . Systematic Biology 66 : 569 – 589 . Google Scholar PubMed WorldCat Percy DM , Crampton-Platt A , Sveinsson S , Lemmon AR , Lemmon EM , Ouvrard D , Burckhardt D . 2018 . Resolving the psyllid tree of life: phylogenomic analyses of the superfamily Psylloidea (Hemiptera ). Systematic Entomology 43 : 762 – 776 . Google Scholar Crossref Search ADS WorldCat Piirainen M , Liebisch O , Kadereit G . 2017 . Phylogeny, biogeography, systematics and taxonomy of Salicornioideae (Amaranthaceae/Chenopodiaceae) – a cosmopolitan, highly specialized hygrohalophyte lineage dating back to the Oligocene . Taxon 66 : 109 – 132 . Google Scholar Crossref Search ADS WorldCat Pometti CL , Bessega CF , Vilardi JC , Saidman BO . 2012 . Landscape genetic structure of natural populations of Acacia caven in Argentina . Tree Genetics & Genomes 8 : 911 – 924 . Google Scholar Crossref Search ADS WorldCat Prum RO , Berv JS , Dornburg A , Field DJ , Townsend JP , Lemmon EM , Lemmon AR . 2015 . A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing . Nature 526 : 569 – 573 . Google Scholar Crossref Search ADS PubMed WorldCat Puissant S , Sueur J . 2001 . Contribution à l étude des cigales de Corse (Hemiptera, Cicadidae) . Bulletin de La Societe Entomologique de France 106 : 429 – 436 . WorldCat Rabassa J . 2008 . Late Cenozoic glaciations in Patagonia and Tierra del Fuego . Developments in Quaternary Sciences 11 : 151 – 204 . Google Scholar Crossref Search ADS WorldCat Rabassa J , Coronato A , Martinez O . 2011 . Late Cenozoic glaciations in Patagonia and Tierra del Fuego: an updated review . Biological Journal of the Linnean Society 103 : 316 – 335 . Google Scholar Crossref Search ADS WorldCat Rabosky DL . 2017 . Phylogenetic tests for evolutionary innovation: the problematic link between key innovations and exceptional diversification . Philosophical Transactions of the Royal Society B: Biological Sciences 372 : 20160417 . Google Scholar Crossref Search ADS WorldCat Rabosky DL , Donnellan SC , Talaba AL , Lovette IJ . 2007 . Exceptional among-lineage variation in diversification rates during the radiation of Australia’s most diverse vertebrate clade . Proceedings of the Royal Society B: Biological Sciences 274 : 2915 – 2923 . Google Scholar Crossref Search ADS WorldCat Rambaut A , Drummond A . 2012 . FigTree version 1.4 . Available at: http://tree.bio.ed.ac.uk/software/figtree/ Google Preview WorldCat COPAC Raup DM , Gould SJ , Schopf TJM , Simberloff DS . 1973 . Stochastic models of phylogeny and the evolution of diversity . The Journal of Geology 81 : 525 – 542 . Google Scholar Crossref Search ADS WorldCat R Core Team . 2018 . R: a language and environment for statistical computing. 3.4.0 ed. Vienna: R Foundation for Statistical Computing . Available at: https://www.R-project.org/. WorldCat Ree RH . 2005 . Detecting the historical signature of key innovations using stochastic models of character evolution and cladogenesis . Evolution; international journal of organic evolution 59 : 257 – 265 . Google Scholar Crossref Search ADS PubMed WorldCat Roelants K , Gower DJ , Wilkinson M , Loader SP , Biju SD , Guillaume K , Moriau L , Bossuyt F . 2007 . Global patterns of diversification in the history of modern amphibians . Proceedings of the National Academy of Sciences of the United States of America 104 : 887 – 892 . Google Scholar Crossref Search ADS PubMed WorldCat Roig FA , Roig-Juñent S , Corbalań V . 2009 . Biogeography of the Monte desert . Journal of Arid Environments 73 : 164 – 172 . Google Scholar Crossref Search ADS WorldCat Ruschel TP . 2018 . Gibbocicada brasiliana, new genus and new species from Brazil and a key for the genera of Tibicinini (Hemiptera: Auchenorrhyncha: Cicadidae) . Acta Entomologica 58 : 559 – 566 . WorldCat Ruzzante DE , Rabassa J . 2011 . Palaeogeography and palaeoclimatology of Patagonia: effects on biodiversity . Biological Journal of the Linnean Society 103 : 221 – 228 . Google Scholar Crossref Search ADS WorldCat Sanborn AF . 1997 . Body temperature and the acoustic behavior of the cicada Tibicen winnemanna (Homoptera: Cicadidae) . Journal of Insect Behavior 10 : 257 – 264 . Google Scholar Crossref Search ADS WorldCat Sanborn AF , Heath JE , Heath MS , Noriega FG . 1995 . Thermoregulation by endogenous heat production in two South American grass dwelling cicadas (Homoptera: Cicadidae: Proarna) . The Florida Entomologist 78 : 319 – 328 . Google Scholar Crossref Search ADS WorldCat Sanborn AF , Heath MS . 2014 . The cicadas of Argentina with new records, a new genus and fifteen new species (Hemiptera: Cicadoidea: Cicadidae) . Zootaxa 3883 : 1 – 94 . Google Scholar Crossref Search ADS PubMed WorldCat Sanborn AF , Heath MS , Heath JE , Noriega FG , Phillips PK . 2004 . Convergence and parallelism among cicadas of Argentina and the southwestern United States (Hemiptera: Cicadoidea) . Biological Journal of the Linnean Society 83 : 281 – 288 . Google Scholar Crossref Search ADS WorldCat Sanderson MJ , Donoghue MJ . 1994 . Shifts in diversification rate with the origin of angiosperms . Science 264 : 1590 – 1593 . Google Scholar Crossref Search ADS PubMed WorldCat Schott RK , Panesar B , Card DC , Preston M , Castoe TA , Chang BS . 2017 . Targeted capture of complete coding regions across divergent species . Genome Biology and Evolution 9 : 398 – 414 . Google Scholar PubMed WorldCat Shcherbakov DE . 2009 . Review of the fossil and extant genera of the cicada family Tettigarctidae (Hemiptera: Cicadoidea) . Russian Entomological Journal 17 : 343 – 348 . WorldCat Simões M , Breitkreuz L , Alvarado M , Baca S , Cooper JC , Heins L , Herzog K , Lieberman BS . 2016 . The evolving theory of evolutionary radiations . Trends in Ecology & Evolution 31 : 27 – 34 . Google Scholar Crossref Search ADS PubMed WorldCat Sokal RR , Rohlf FJ . 1995 . Biometry, the principles and practice of statistics in biological research . New York : H. Freeman and Company . Google Preview WorldCat COPAC Sota T , Yamamoto S , Cooley JR , Hill KBR , Simon C , Yoshimura J . 2013 . Independent divergence of 13- and 17-y life cycles among three periodical cicada lineages . Proceedings of the National Academy of Sciences of the United States of America 110 : 6919 – 6924 . Google Scholar Crossref Search ADS PubMed WorldCat Speelman EN , Van Kempen MM , Barke J , Brinkhuis H , Reichart GJ , Smolders AJ , Roelofs JG , Sangiorgi F , de Leeuw JW , Lotter AF , Sinninghe Damsté JS . 2009 . The Eocene Arctic Azolla bloom: environmental conditions, productivity and carbon drawdown . Geobiology 7 : 155 – 170 . Google Scholar Crossref Search ADS PubMed WorldCat Stamatakis A . 2014 . RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies . Bioinformatics 30 : 1312 – 1313 . Google Scholar Crossref Search ADS PubMed WorldCat Sueur J , Vanderpool D , Simon C , Ouvrard D , Bourgoin T . 2007 . Molecular phylogeny of the genus Tibicina (Hemiptera, Cicadidae): rapid radiation and acoustic behaviour . Biological Journal of the Linnean Society 91 : 611 – 626 . Google Scholar Crossref Search ADS WorldCat Swofford DL . 1998 . Phylogenetic analysis using parsimony (PAUP), version 4 . Sunderland : Sinauer Associates . Google Preview WorldCat COPAC Takezaki N , Nishihara H . 2016 . Resolving the phylogenetic position of Coelacanth: the closest relative is not always the most appropriate outgroup . Genome Biology and Evolution 4 : 1208 – 1221 . Google Scholar Crossref Search ADS WorldCat Taucce PPG , Canedo C , Haddad CFB , Lemmon AR , Lemmon EM , Vences M , Lyra M . 2018 . The mitochondrial genomes of five frog species of the Neotropical genus Ischnocnema (Anura: Brachycephaloidea: Brachycephalidae) . Mitochondrial DNA Part B 3 : 915 – 917 . Google Scholar Crossref Search ADS WorldCat Toolson EC . 1987 . Water profligacy as an adaptation to hot deserts: water loss rates and evaporative cooling in the Sonoran Desert cicada, Diceroprocta apache (Homoptera: Cicadidae) . Physiological Zoology 60 : 379 – 385 . Google Scholar Crossref Search ADS WorldCat Torres BA . 1945 . Revisión de los géneros Chonosia Dist. Mendozana Dist. y Derotettix Berg. y algunas interesantes notas cicadidologicas (Homoptera-Cicadidae) . Notas Del Museo de La Plata 10 : 55 – 82 . WorldCat Turchetto-Zolet AC , Pinheiro F , Salgueiro F , Palma-Silva C . 2013 . Phylogeographical patterns shed light on evolutionary process in South America . Molecular Ecology 22 : 1193 – 1213 . Google Scholar Crossref Search ADS PubMed WorldCat Vaidya G , Lohman DJ , Meier R . 2011 . SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information . Cladistics 27 : 171 – 180 . Google Scholar Crossref Search ADS WorldCat Van Leuven JT , Meister RC , Simon C , McCutcheon JP . 2014 . Sympatric speciation in a bacterial endosymbiont results in two genomes with the functionality of one . Cell 158 : 1270 – 1280 . Google Scholar Crossref Search ADS PubMed WorldCat Vaux F , Trewick SA , Morgan-Richards M . 2016 . Lineages, splits and divergence challenge whether the terms anagenesis and cladogenesis are necessary . Biological Journal of the Linnean Society 117 : 165 – 176 . Google Scholar Crossref Search ADS WorldCat Wallis GP , Waters JM , Upton P , Craw D . 2016 . Transverse Alpine speciation driven by glaciation . Trends in Ecology & Evolution 31 : 916 – 926 . Google Scholar Crossref Search ADS PubMed WorldCat Wang Z , Sun F , Jin P , Chen Y , Chen J , Deng P , Yang G , Sun B . 2017 . A new species of Ginkgo with male cones and pollen grains in situ from the Middle Jurassic of eastern Xinjiang, China . Acta Geologica Sinica/Dizhi Xuebao 91 : 9 – 21 . Google Scholar Crossref Search ADS WorldCat Woodburne MO , Goin FJ , Raigemborn MS , Heizler M , Gelfo JN , Oliveira EV . 2014 . Revised timing of the South American early Paleogene land mammal ages . Journal of South American Earth Sciences 54 : 109 – 119 . Google Scholar Crossref Search ADS WorldCat Yoshizawa K , Ogawa N , Dietrich CH . 2017 . Wing base structure supports Coleorrhyncha + Auchenorrhyncha (Insecta: Hemiptera) . Journal of Zoological Systematics and Evolutionary Research 55 : 199 – 207 . Google Scholar Crossref Search ADS WorldCat Young AD , Lemmon AR , Skevington JH , Mengual X , Ståhls G , Reemer M , Jordaens K , Kelso S , Lemmon EM , Hauser M , De Meyer M , Misof B , Wiegmann BM . 2016 . Anchored enrichment dataset for true flies (order Diptera) reveals insights into the phylogeny of flower flies (family Syrphidae) . BMC Evolutionary Biology 16 : 143 . Google Scholar Crossref Search ADS PubMed WorldCat Yu G , Smith DK , Zhu H , Guan Y , Lam TTY . 2017 . GGTREE: an R package for visualization and annotation of phylogenetic trees with their covariates and other associated data . Methods in Ecology and Evolution 8 : 28 – 36 . Google Scholar Crossref Search ADS WorldCat Zucol AF , Krause JM , Brea M , Raigemborn MS , Matheos SD . 2018 . Emergence of grassy habitats during the greenhouse–icehouse systems transition in the Middle Eocene of southern South America . Ameghiniana 55 : 451 – 482 . Google Scholar Crossref Search ADS WorldCat © 2019 The Linnean Society of London, Biological Journal of the Linnean Society This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model)
TI - Off-target capture data, endosymbiont genes and morphology reveal a relict lineage that is sister to all other singing cicadas
JF - Biological Journal of the Linnean Society
DO - 10.1093/biolinnean/blz120
DA - 2019-12-06
UR - https://www.deepdyve.com/lp/oxford-university-press/off-target-capture-data-endosymbiont-genes-and-morphology-reveal-a-j9b6wfO3uv
SP - 1
VL - Advance Article
IS - 
DP - DeepDyve
ER -