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The Decticini and Ctenodecticini (Orth., Tettigoniidae) Are Not Monophyletic: a Total Evidence Approach

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The Decticini and Ctenodecticini (Orth., Tettigoniidae) Are Not Monophyletic: a Total Evidence Approach Archive of SID ÎÕÕ ÎÐîéŹƺƿźƸƃìîƱřźƿřƾſŚƴƃƵźƄůƾƬƬưƫřƲ ǀŝƵźĮƴƧƲǀƫƹř The Decticini and Ctenodecticini (Orth., Tettigoniidae) are not monophyletic: a total evidence approach M. Mofidi-Neyestanak Insect Taxonomy Research Department (ITRD) & Hayk Mirzayans Insect Museum (HMIM), Iranian Research Institute of Plant Protection (IRIPP), Agricultural Research, Education and Extension Organization (AREEO), Tehran, Iran, [email protected] Abstract The first phylogenetic hypothesis for the Decticini and Ctenodecticini is presented based on the total evidence of morphological and molecular datasets of 37 representatives of the two tribes and close taxa. The simultaneous dataset includes 1779 characters, of which 211 are morphological (discrete and continuous) and the rest are molecular data (mitochondrial cytochrome oxidase-b, nuclear 28S rDNA, and mitochondrial 16S rDNA). Parsimony analyses were carried out on each of the datasets, singly and simultaneously. The phylogenetic trees of each analysis were compared with each other and matched up with the results of previous works. As a result, monophyly of the Decticini and Ctenodecticini is rejected and both are suggested as junior synonyms of Platycleidini. Keywords: Continuous, discrete, morphology, rDNA, mtDNA, phylogeny, Tettigoniidae, Decticini, Ctenodecticini, total evidence, synonym Orth., Tettigoniidae Ctenodecticini Decticini ƵŶǀƨģ ŚƷƵ ŵřŵƖǀưŬţƁƹŹŶƴŤƀǀƳŚǀƳƦ ţ ŶƴƬŝƦ ųŚƃƽŚƷŲ ƬƯŻř ƹ ŠƬǀŞƣƹŵ ƦƳŚŤƀǀƳƽŶǀƠƯƲƀŰƯ Ctenodecticini Decticini ŹŵŚŝƹƾƫƺƨƫƺƯƹƾſŚƴƃƪ ƨƃƽŚƷƵŵřŵƖǀưŬţŽŚſřźŝ ƹ ŠƬǀŞƣƹŵƦǀŤƀƿŵLjƧƾſŚƴƃŹŚŞţŠǀƋźƟƲǀƫƹř çææƶƧŢſřźŤƧřŹŚƧæììîƪƯŚƃƾƿŚƸƳƾŞǀƧźţžƿźţŚƯŢſřƵŶƃƶŗřŹřƦƿŵżƳƽŚƷƵƹźĭƹƶƬǀŞƣƹŵƱŚĭŶƴƿŚưƳƱřƺƴƗƶŝƶƳƺĭèìƲŤƟźĭźƔƳ ƾŝŻřŶǀƀƧřƭƹźƧƺŤǀſƾƿŚƿŹŶƴƧƺŤǀƯƽŚƷƱ ĥ ƾƫƺƨƫƺƯƽŚƷƵŵřŵƶŝƍƺŝźƯƶǀƤŝƹ ƶŤƀƀĭƹƶŤſƺǀě ƾſŚƴƃƪ ĪƃƽŚƷƾ ĭĦƿƹƶŝƍƺŝźƯŵŶƗ 16S 28S ƹƾƿŚţƹŵƽŚƷŜ ǀƧźţƾƯŚưţżǀƳƹŚƷž ƿźţŚƯƦƿŚƨƿƽƹŹƾƳƺưǀſŹŚěƪǀƬŰţŢſř ƾƯƹŻƺŞƿŹ ƾƿŚƿŹŶƴƧƺŤǀƯƹƾƯƹŻƺŞƿŹƽřƶŤƀƷ Ctenodecticini Decticini ƱřƺƴƗƶŝƶƬǀŞƣƹŵźƷƹŶƃŵŹ ƹ ŠƬǀŞƣƹŵƱŵƺŝŚǀƳƦ ţƶŬǀŤƳŹŵŶƃƭŚŬƳřŚƷƵŵřŵƪƧŢƿŚƸƳŹŵƹŚƷƵŵřŵƾƿŚţŶƴģ Platycleidini ŶƳŶƃƭLjƗř ŵƺųƾƬŞƣƭŚƴưƷ Introduction Although Tettigoniinae is the largest subfamily of Tettigoniidae (Orthoptera: Ensifera), there have been little molecular phylogenetic studies within the subfamily. Based on morphological information, Harz (1969) presented an arbitrary key comprising 8 previously described tribes (Arytropteridini, Drymadusini, Glyphonotini, Nedubiini, Onconotini, Plagiostirini, Platycleidini, and Tettigoniini) within the subfamily. Rentz & Colless (1990), based on a numerical taxonomic analysis, reduced this number to 5 (Glyphonotini, Nedubiini, Onconotini, Platycleidini, and Tettigoniini). In the most recent work, Eades et al. (2015) presented a list of 12 tribes in their database, by adding Bergioini (Bergiola Shchelkanovtsev, Bienkoxenus Èejchan, Eulithoxenus Bei-Bienko, and Uvarovina Ramme), Ctenodecticini (Ctenodecticus Bolivar and Miramiola Uvarov), Decticini (Bicolorana Zeuner, Chizuella Furukawa, Decticus Serville, Eobiana Bei-Bienko, Hypsopedes Bei-Bienko, Metrioptera, Montana Zeuner, Sphagniana Zeuner, and Tessellana Zeuner), and Gampsocleidini (Gampsocleis Fieber and Uvarovites Tarbinsky). After raising a separate group of decticine bushcrickets as an available family (Herman, 1874), the longstanding genus of Decticus Audinet-Serville was classified by the coming orthopterists at different ranks (see Kirby, 1906). Afterwards, the validity of the group as a subfamily and then a tribe within the Tettigoniidae was discussed based on morphological characters and followed by most recent authors (Zeuner, 1941; Kaltenbach, 1971; Rentz, 1979, 1985; McE.Kevan, 1982; Storozhenko, 2004; Ünal, 2012; Eades et al., 2015). Similarly, the closely related taxon, the tribe Ctenodecticini that was raised by Uvarov (1939) and followed by the others (Kaltenbach, 1971; Storozhenko, 2004; Eades et al., 2015) is a heterogeographical group; the species of the Ctenodecticus Bolívar are distributed in the North of West Africa and the South of West Europe, while the only species within the other genus (Miramiola Uvarov) is distributed in East Europe and the South of Russia (see Eades et al., 2015; Roskov et al., 2015). The 28S rDNA gene has been used in phylogenetic analyses within the Orthoptera at higher taxonomic levels due to its relatively slow substitution rate. In this gene segment, the unambiguously aligned regions are relatively conserved; making st 188 1 Iranian International Congress of Entomology, 29–31 August 2015 www.SID.ir Archive of SID ÎÕÖ ÎÐîéŹƺƿźƸƃìîƱřźƿřƾſŚƴƃƵźƄůƾƬƬưƫřƲ ǀŝƵźĮƴƧƲǀƫƹř the gene especially useful for recovering deeper divergence levels (Belshaw & Quicke, 1997; Belshaw et al., 2001). On the other hand, mitochondrial fragments of cytb and 16S rDNA have been widely employed within the Orthoptera to reconstruct phylogenetic relationships at various taxonomic levels (Chapco et al., 2001; Jermiin & Crozier, 1994; Lunt et al., 1996; Maekawa et al., 1999; Filipenko & Timofeeva, 2000; Ren et al., 2004; Allegrucci et al., 2005; Hui-Meng & Yuan, 2006; Jost & Shaw, 2006; Robillard & Desutter-Grandcolas, 2006). Theses genes, because of high substitution rate, are normally most informative at interspecific level. Since mitochondrial and nuclear genes evolve at different rates, combining of them will probably increase the phylogenetic estimation and resolution of cladograms (Caterino et al., 2000). These three gene fragments were chosen because of their different levels and types of variations in the hope that they will complement each other. Here a combined molecular and morphological approach is taken to provide a more objective and robust phylogenetic hypothesis of the group, focusing on the composition of Decticinae sensu lato and also on the composition of Decticini and Ctenodecticini. Materials and methods A total of 200 fresh, alcohol-preserved, or pinned specimens (total of 37 species belonging to 19 genera/subgenera) representative of major lineages of the Decticinae and Ctenodecticini were studied using molecular and morphological techniques. Most borrowed material had initially been identified by the senders or by the museums from which it had been borrowed. Other specimens and most of the borrowed material was then determined or their identifications confirmed by reference to various taxonomic keys. When necessary, specimens were compared with type material in the Natural History Museum London or the Muséum National d'Histoire Naturelle France for further confirmation. Five datasets of continuous and discrete morphological characters, 28S nuclear rDNA, mitochondrial 16S rDNA and mitochondrial cytochrome oxidase b (cytb) were analysed singly and simultaneously. The final dataset comprises total of 1779 characters, of which 38 are continuous morphological, 173 discrete morphological characters and 1568 DNA characters (including gaps), of which there were 327 parsimony-informative characters (approximately 21%). The number of parsimony informative characters for each dataset is given in Table 2. In a few cases the samples included in the morphological study were not available for molecular work (and vice versa). Total genomic DNA of alcohol-preserved material was individually extracted from hind femoral muscle of bushcrickets (Bensasson et al., 2000). Sequences were amplified using the primers given in Table 1. All the sequence electropherograms were checked manually and edited with Sequence Navigator software (Applied Biosystems Inc). Ambiguities and gaps were treated as missing data and the primer ends were removed. The mitochondrial protein-coding sequences were initially aligned by eye and then inspected and edited by exporting into MacClade 4.0 (Maddison & Maddison 2000) using amino acid translation graphic facilities. The ribosomal gene sequences were firstly aligned visually to locate the most conserved parts. Then, to determine what global alignment strategy would preserve best the previously indicated conserved parts, they were aligned using the Clustal-W programme (Chenna et al., 2003) with a range of Gap Opening to Gap Extension (GO:GE) penalties. As a result, for the 16S, 10:2.5 and for 28S, 2:1 GO:GE penalties were selected. The Clustal alignments were not manually adjusted. The included taxa, their provenances, morphological data and GenBank accession numbers are available on demand. The molecular data matrix can be downloaded from the TreeBase webpage (submission accession no. SN3593-16279). Terminology followed Maran (1953), Ragge (1955), Rentz & Birchim (1968), Harz (1969), and Mofidi-Neyestanak & Quicke (2007). st 189 1 Iranian International Congress of Entomology, 29–31 August 2015 www.SID.ir Archive of SID ÎÖÍ ÎÐîéŹƺƿźƸƃìîƱřźƿřƾſŚƴƃƵźƄůƾƬƬưƫřƲ ǀŝƵźĮƴƧƲǀƫƹř Phylogenetic analyses The data were analysed without a priori weighting. Dataset attributes were calculated and statistical analyses carried out using PAUP* version 4.0b10 (Swofford, 2001) and TNT version 1.1 (Goloboff et al., 2003). Multistate characters were coded as polymorphisms and the gaps in the molecular alignments were treated as the ‘fifth base’. The entire characters, except for the continuous morphological data, were treated as non-additive. In analyses that were carried out in TNT, all the criteria (Sectorial search, Ratchet, Drift, and Tree fusing) of the New Technology Search were selected and the parameters set as deafult; however, careful consideration was made to understand how datasets behave to changes in parameters. As a result, some minor alterations were applied in some analyses. In the maximum parsimony analyses using PAUP*,
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