Archive of SID

ÎÕÕ ÎÐîéŹƺƿźƸƃìîƱřźƿřƾſŚƴƃƵźƄůƾƬƬưƫřƲ ǀŝƵźĮƴƧƲǀƫƹř  The Decticini and Ctenodecticini (Orth., ) are not monophyletic: a total evidence approach

M. Mofidi-Neyestanak 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 ţƶŬǀŤƳŹŵŶƃƭŚŬƳřŚƷƵŵřŵƪƧŢƿŚƸƳŹŵƹŚƷƵŵřŵƾƿŚţŶƴģ Platycleidini ŶƳŶƃƭLjƗř ŵƺųƾƬŞƣƭŚƴưƷ Introduction Although is the largest subfamily of Tettigoniidae (: ), 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

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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).

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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*, a total of 10,000 random additions with TBR branch swapping holding only one tree, was applied. In total 15 single and combined analyses were carried out. Based on their presence in each dataset, Ephippiger ephippigera (Fiebig, 1784) (Ephippigerinae) or Tettigonia viridissima (Linnaeus, 1758) (Tettigoniinae) were selected as outgroup. In order to assess the robustness of the generated phylogenetic trees, nodal support for the monophylies of clades was assessed using bootstrapping and jackknifing for the whole cladograms. The standard bootstrapping values were calculated in TNT using 1000 replicates with 10 random additions per replicate. The jackknifing nodal support values were calculated with a deletion of 50% removal probability (Felsenstein, 2004). To examine the difference in phylogenetic signal between the datasets, incongruence length differences (ILDs) were calculated in PAUP* (Farris et al., 1994) by executing 1000 random partitions of the data for 100 replications. All uninformative characters were removed and the only taxa common to the tested partitions included in the analysis (Cunningham, 1997).

Results Two datasets of discrete and continuous data were analysed singly and simultaneously. The continuous and the combined datasets were analysed using TNT as described above, while the discrete morphological data was analysed using heuristic search (in PAUP*). In the cladogram resulted from the simultaneous analyses of morphological data, near the base of the tree comprises Tettigonia (the outgroup), Decticus and Medecticus. In this grade the two representative of Tettigonia are not recovered together (as are also found in the simultaneous analysis, Fig. 1). Results of the ILD tests revealed incongruence between most combinations of datasets. Among the molecular dataset combinations, only 28S and cytb were found to be congruent (p= 0.12). The phylogenetic estimate based on 28S was not resolved well. The parsimony analysis of this dataset yielded 855 MPTs and the strict consensus of these was extremely poorly resolved. Less than one third of the characters were parsimony informative (see Table 2). The 50% majority rule consensus tree of 28S has few resolved clades, though, a well-supported group of (Ctenodecticus granatensis (Tessellana orina (Anonconotus baracunensis (Metaballus mucronatus (Sporadiana sporadarum (Pachytrachis striolatus+Squamiana sp) is recovered. This clade also appears in the single MPT from the 28S+discrete data analysis, comprises 7 morphologically divergent genera and suggests large amount of homoplasy in their morphology. In the strict consensus tree of 2 MPTs derived from analysis of cytb the Decticus, Platycleis, Tessellana, and Eupholidoptera Groups were resolved and the latter group is placed as sister group to Pholidoptera. Among the three molecular datasets, 28S and cytb were found to be congruent (ILD p value= 1.00). The combined analysis of these datasets yielded a consensus tree includes monophyletic Platycleis and Tessellana Groups as sister. The Eupholidoptera and Decticus Groups are grouped as a grade and placed as sister group to Pholidoptera and Bolua turkiyae. However, this clustering is not recovered in the cladogram resulting from analysis of the 3 molecular datasets combined.

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For total evidence analyses, the ILD tests results revealed that among the combined matrices of molecular and morphological datasets, the 28S+ discrete is congruent (p= 1.00). However, the entire eight combinations of the 2 morphological and 3 molecular datasets were analysed. In most of the combined analyses, the clades were often more robust, almost certainly because of complementarities of the data; nevertheless, some clades of the simultaneous analyses were less supported than particular single analyses. The most resolved cladogram of the simultaneous analyses of morphological and molecular data was 16S+discrete. In this tree, the Platycleis, Parnassiana, Anterastes, Pholidoptera, Eupholidoptera and Koroglus Groups are formed. The Decticus Group (excluding Montana barretii) is sister group to Pholidoptera Group+Bolua. Also, Koroglus is placed as sister group to Anterastes. All the included species of Tessellana are grouped together except for T. incerta. In the strict consensus tree resulted from the simultaneous parsimony analysis of all the molecular and the discrete morphological data the Decticus Group is recovered. Parnassiana is placed as sister group to the Tessellana Group and the Decticus Group is paraphyletic to (Bolua (Pholidoptera (Eupholidoptera))). In the single MP tree yielded from analysis of the total evidence (Fig. 1) the Decticus Group is partially recovered, M. barretii falls outside of it and the species of Tettigonia and Decticus are mixed up within the group. A clade comprising Bolua, Pholidoptera, the Eupholidoptera group, and the Decticus Group is formed in the cladograms of cytb, 28S+cytb, and in the 3 molecular datasets combined. However, this clade is not recovered in any of the morphological analyses and the above mentioned genera are spread all over the 16S and the total evidence trees.

Table 1. Primers used in this study (5′—3′).

locus Primer Reference

Forward ′TATGTTTTACCATGAGGACAAATATC′ Jermiin & Crozier (1994) cytb Reverse ′TATTTCTTTCTTATGTTTTCAAAAC′ Jermiin & Crozier (1994) Forward 1 ′CGCCTGTTTAACAAAAACAT′ Simon et al. (1994) Reverse 1 ′TTTAATCCAACATCGAGG′ Cognato & Vogler (2001) 16S Forward 2 ′CACCTGTTTATCAAAAACAT′ Costa et al. (2003) Reverse 2a ′CGTCGATTTGAACTCAAATC′ Costa et al. (2003) Reverse 2b ′CTTATTCAACATCGAGGTC′ Whitfield (1997) Forward 1 ′AAGAGAGAGTTCAAGAGTACGTG′ Belshaw & Quicke (1997)

Reverse 1 ′TAGTTCACCATCTTTCGGGTCCC′ Mardulyn & Whitfield (1999) 28S Forward 2 ′CCACAGCGCCAGTTCTGCTTAC′ Muraji & Tachikawa (2000)

Reverse 2 ′CCCGTCTTGAAACACGGACCAA′ Muraji & Tachikawa (2000)

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Table 2. Statistical comparisons of the maximum parsimony trees from singly and simultaneous analyses of the five data partitions.

ILD Total Informativ MPT Tree P- character e CI RI s length valu s characters e 0.22 Discrete 173 125 1 1011 0.53 - Morphological 7 datasets Continuous 38 38 5 1846.36 0.81 0.77 - Discrete+continuous 211 193 5 3076.545 0.3 0.67 - 0.31 0.62 16S 499 174 35 1299 - 3 9 0.85 0.91 28S 546 161 855 256 - 9 6 0.34 0.58 cytb 523 200 2 996 - 5 6 0.46 0.61 0.01 Molecular datasets 16S+28S 1045 293 2 1776 6 8 * 0.39 0.57 0.01 16S+cytb 1022 319 21 1575 5 8 * 0.35 0.58 28S+cytb 1069 209 2 1015 1.00 3 8 0.37 0.57 0.01 16S+28S+cytb 1568 327 11 1536 5 9 * 0.26 0.54 0.01 16S+discrete 672 294 12 2306 6 9 * 0.34 0.57 28S+discrete 719 284 1 1279 0.12 9 7 0.38 0.56 0.01 cytb+discrete 696 284 3 1304 Combined 3 7 * molecular and 0.32 0.54 0.01 16S+28S+discrete 1218 412 6 2491 morphological 7 8 * datasets 0.41 0.57 0.01 16S+cytb+discrete 1195 395 2 1772 9 5 * 0.38 0.56 0.01 28S+cytb+discrete 1242 292 3 1327 7 6 * 0.42 0.57 0.01 16S+28S+cytb+discrete 1741 402 2 1794 1 5 * 16S+28S+cytb+discrete+continuo 0.32 0.59 Total evidence 1779 969 1 6414.291 - us 1 2

* Significant incongruence between datasets is indicated with an asterisk

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Table 3. Recovered clades resulted from phylogenetic analyses of single and combined molecular and morphological data 

PLAss TES PAR ANT DEC PHO EUP KOR (PHO+EUP) (Bolua+ANT) Continuous Discrete Continuous+discrete 16S 28S cytb - - - 16S+28S 16S+cytb - - - - 28S+cytb - - - - 16S+28S+cytb - - - - 16S+discrete 28S+discrete cytb+discrete - - - - 16S+28S+discrete 16S+cytb+discrete - - - - - 28S+cytb+discrete - - - - 16S+28S+cytb+discrete - - - - Total evidence

PLA ss= Platycleis sensu stricto, examined species of the subgenus Platycleis (sensu Eades et al., 2015); TES= examined species of the genus Tessellana; PAR= examined species of the subgenus Parnassiana (sensu Eades et al., 2015); ANT= Antaxius+Yersinella+Rhacocleis; DEC= Decticus+Medecticus+Uvarovistia+Montana barretii; PHO= examined species of the genus Pholidoptera; EUP= examined species of the genus Eupholidoptera excluding E. krueperi and E. mirzayani; KOR= Koroglus+Anterastes. Gray= recovered clade; Dark grid= not recovered clade; (-)= no species of the given clade was available in the analysis.

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Fig. 1. Phylogenetic estimate of the tribes Decticini and Ctenodecticini (Orth., Tettigoniidae) based on simultaneous parsimony analysis of molecular (cytb, 28S, 16S) and morphological (continuous and discrete) datasets. Part of the single MPT is shown with bootstrap and jackknifing support values (>50%) indicated above and below branches respectively.

Discussion From Herman (1874) to recent workers, different subfamilial systems are proposed for the Tettigoniinae, the largest subfamily of Tettigoniidae. All the suggested hypotheses have been based on morphological information and there has not been any agreement on its tribes or genera. The earliest attempts were largely based on identifying the actual tettigoniines from the others. Originally, Krauss (1890) and after him, Brunner von Wattenwyl (1893) used the term decticine for the latter group and consequently Kirby (1906) and Caudell (1907, 1908) used it as a subfamily (Decticinae). Zeuner (1941) defined Platycleidini to include a small number of decticines and workers after him added more taxa within Platycleidini and made it much larger (see Ramme, 1951; Harz, 1969; Rentz & Colless, 1990). In addition to Platycleidini, several other tribes have been defined (Harz, 1969; Rentz & Colless, 1990; Storozhenko, 1994) and in total, twelve tribes are proposed within the Tettigoniinae: Arytropteridini, Bergiolini, Ctenodecticini, Decticini, Drymadusini, Gampsocleidini, Glyphonotini, Nedubini, Onconotini, Plagiostirini, Platycleidini, and Tettigoniini (Roskov et al., 2015; Eades et al., 2015). Based on the yielded results of the present work, several taxonomic implications are suggested as follows: Decticini: Rentz (1979) was the first worker who used Decticini as a tribe, based on an incorrect spelling of a family group (Dectici) used by Brunner von Wattenwyl (1893). The genera that Rentz (1979, 1985) included in the Decticini were:

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Bicolorana Zeuner, Chizuella Furukawa, Decticus Serville, Eobiana Bei-Bienko, Hypsopedes Bei-Bienko, Metrioptera Zeuner, Montana Zeuner, Sphagniana Zeuner, and Tessellana Zeuner. This was followed by Eades et al. (2015) and Roskov et al. (2015). In the majority of the cladograms Decticus, Medecticus, and Uvarovistia are grouped together and their clade falls within the other genera of the Platycleidini. In addition, Eobiana, Metrioptera, Montana, and Tessellana are not clustered with Decticus. The Medecticus and Decticus are traditionally placed in Decticinae (Harz 1969) or Tettigoniini (Rentz & Colless, 1990; Eades et al., 2015). The Uvarovistia is a member of the Platycleidini (Eades et al., 2015). The monophyly of Decticini sensu Rentz (1979) is not supported in any of the analyses and its genera are spread among the genera of Platycleidini. Thus, the Decticini is considered to be a junior synonym of the Platycleidini. Ctenodecticini: This tribe was originally erected by Uvarov (1939) for two genera of Ctenodecticus Bolivar and Miramiola Uvarov. The latter genus could not be obtained for the present study. In analyses of combined morphological data Ctenodecticus is placed in a well-supported clade as (Ctenodecticus+Koroglus). In the 16S+28S it is formed as (Bucephaloptera (Rhacocleis+Ctenodecticus+). Similarly, in the consensus tree obtained from analysis of 16S+28S+discrete it is placed as sister group to the Antaxius Group. In the cladogram obtained from the total evidence analysis (Fig. 1) Ctenodecticus is placed as sister group to Yersinella and their clade is paraphyletic to Bolua. In all the obtained trees Ctenodecticus falls within the tribe Platycleidini. Thus Ctenodecticini becomes a junior synonym of Platycleidini.

Acknowledgements I never forget the attentive support and kind encouragement of the late Hayk Mirzayans, my beloved teacher in Orthoptera.Thanks to collegues, friends and contributors in the Hayk Mirzayans Insect Museum, Iran; the Imperial College London; and the Natural History Museum, London, UK; the Muséum National d'Histoire Naturelle; and the Université Louis Pasteur, France; Abant Izzet Baysal Üniversitesi; and the Akdeniz University, Turkey; Jordan University; Tokyo Metropolitan University, Japan; Universität Osnabrück, Germany; Zoological Institute Academy of Sciences St. Petersburg, Russia. Special appreciation to Prof. Dr. D.L.J. Quicke, the great emeritus of Imperial College London, and the Natural History Museum, London, UK for providng me support and fascilities to work in his lab after my PhD in University of London.

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st 196 1 Iranian International Congress of Entomology, 29–31 August 2015

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st 197 1 Iranian International Congress of Entomology, 29–31 August 2015

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st 198 1 Iranian International Congress of Entomology, 29–31 August 2015

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